A Cultural History of Heredity

M A X - PLA N C K - I N S T I T U T F Ü R W I S S E N S C H A FT S G E S C H I C H T E

Max Planck Institu te for the History of Science

2002

PREPRINT 222

Conference

A Cultural History of Heredity I: 17th and 18th Centuries

ISSN 0948-9444

Table of Contents

Introduction

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Cabbage, Tulips, Ethiopians – “Experiments” in Early Modern Heredity Staffan Müller-Wille

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Speculation and Experiment in Enlightenment Life Sciences Mary Terrall

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Duchesne’s Strawberries: Between Grower’s Practices and Academic Knowledge Marc J. Ratcliff

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Natural Things and Non-natural Things. The Boundaries of the Hereditary in the 18th Century Carlos López-Beltrán

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List of Authors

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Introduction This volume assembles some of the contributions to the workshop “A Cultural History of Heredity I: Seventeenth and Eighteenth Centuries” which took place at the Max-Planck-Institute for the History of Science MAY 24-26, 2001. The workshop was the first in a series of workshops dedicated to the cultural history of heredity. There are a number of histories of genetics written from the perspective of history of ideas. François Jacob’s La logique du vivant (1970; engl. transl. 1973) and Robert Olby’s Origins of Mendelism (1966, 2nd ed. 1985) have certainly set lasting standards in this field. There are also some sophisticated, far from whiggish histories written from a disciplinary perspective, like Hans Stubbes Kurze Geschichte der Genetik (1963; engl. transl. 1972), Leslie Clarence Dunn’s A short history of genetics (1965), and Elof Axel Carlson’s The gene: a critical history (1966), to name just a few. What is missing, however, is a comprehensive study that embraces the cultural history of heredity by presenting the knowledge of heredity in its broader practical and historical contexts. In our project we wish to focus on the scientific and technological procedures, in which the knowledge of heredity was materially anchored and by which it affected other cultural domains. Such a project will be content neither with conventional history of ideas nor with mere social history. It will rather explore the various practices, standards, and architectures of hereditary knowledge and the “spaces” which they formed by their respective historical conjunctions. “Heredity”, under this perspective, is more than the scientific discipline “genetics”. The project is less about the history of a science than about the history of a broader knowledge regime, in which a naturalistic conception of heredity developed historically that today affects all domains of society. This knowledge regime dates back to the social illusions and illuminations of the Enlightenment. What does it mean that nature determines history so that it appears as if history could be controlled by nature? And: are we today, with the advent of gene technologies, witnessing the end of a deterministic world view, or are we confronted with its definite restoration? A project like this is vitally dependent on the participation of experts from a broad range of disciplines, covering cultural history in its various subdomains of science, technology, medicine, politics, economy, law, literature, and art. It will be pursued in a series of workshops, each focussing on a loosely defined “epoch” characterised by a certain development in hereditary knowledge. The first workshop, organized by Hans-Jörg Rheinberger, Peter McLaughlin and Staffan MüllerWille, concentrated on the late seventeenth and the eighteenth century and assembled historians of science, medicine, politics and literature from the United States, Mexiko, Germany, Switzerland, and Italy. Four poapers presented at this workshop make up this volume. Discussion during the workshop turned around two main questions: 1), if a concept of heredity existed at all in this period; and 2), in how far 18th century theories of generation were guided by empirical experience. In regard to the first question, several contributions could show that there was no general concept of heredity underlying the discourse of the life sciences (Fantini, Terrall). However, there did exist some isolated, well-defined and sometimes, especially in breeding and medicine, highly localised fields structured by the recognition of hereditary transmission of differential characters

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in the 18th century – the definition of specific difference in natural history (Müller-Wille), the explanation of hereditary diseases in pathology (Lopez-Beltran), political organisation of colonial societies according to racial characteristics (Mazzolini), and the application of hybridisation in plant (Ratcliff). In regard to the second question, the workshop disclosed a rich spectrum of theoretical approaches to generation in the 18th century and made clear that this diversity is only insufficiently captured by the conventional dichotomy of preformation vs. epigenesis (Fantini, Terrall). This spectrum, however, was rather determined by different positions in regard to the politics and poetics of production, both experimental and social, than by a secured and welldefined domain of empirical data (Müller-Sievers, Roe, Terrall). Future workshops in the series follow a very rough chronological order. Their thematic structure will be developed from the results reached in the preceding workshop. The folllowing list of planned workshops, therefore, indicates in a very preliminary way how the project will proceed: Eighteenth to Nineteenth Century: Heredity Becomes Central (January 10-12, 2003) This workshop will focus on the period demarcated by the publication of Kant’s Von den verschiedenen Rassen der Menschen 1775 and Darwin’s The Variation of Animals and Plants under Domestication 1868. This “epoch” in the cultural history of heredity was characterised by a decisive development at the end of which stood the establishment of “inheritance” as one of the central problems of the life sciences. Parallel to this shift one can observe conceptual displacements: Heredity began to be conceptualised as a relation between parental and filial dispositions, rather than between over-all constitutions. Accordingly, theory formation began to revolve around the production and combination of traits within a species rather than around its over-all morphology. Finally, adaptation (or degeneration) under changing conditions, rather than a pre-established balance of nature, became its general framework. Four fields have preliminarily been identified in which these displacements took place: natural history, breeding research, medicine (including psychiatry), and anthropology. Nineteenth to Twentieth Century: Heredity Becomes Exact This workshop will cover the period from Galton’s Hereditary Genius to the formulation of a thoroughly mathematical populations genetics in the 1930s. Population thinking and statistics replace the taxonomic regime of race and character, that previously provided the conceptual framework for hereditary knowledge. The historical background for this development can be seen in the social transformations triggered by industrialisation and concurrent eugenic visions of regulating and controlling populations, including their notorious 20th century versions. Early Twentieth Century: Heredity Becomes Molecular Molecular genetics did not immediately result from the developments characterised in the previous workshop, but from a plurality of methodological achievements in other biological domains, few of which had to do with genetics. The result of these conjunctions, however, was a further naturalisation of hereditary thought with the “cracking” of the genetic code. Heredity was

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brought down from the structural level of populations and genetic traits to a material level of molecules capable of being “read”, “copied”, “translated”, and “transmitted”, be it in human bodies, be it in laboratories. The question of genetic determination and control thus gained renewed prominence after the demise of eugenics. Late Twentieth Century: Heredity Becomes Technological The field of heredity was again thoroughly transformed with the advent of “gene technologies”. Potentially all areas of social life are affected by these practices, and their application in medical and industrial contexts open up perspectives whose limits are typically not foreseeable. Gene patenting, DNA-fingerprinting, gene therapy, and cloning are areas hotly debated in this regard. For the discipline, moreover, the question arises how future research will be oriented after the sequencing of large genomes has become routine work. With the longue durée historical perspective provided by the previous workshops we hope to gain innovative, even surprising outlooks on these problems of the day. Hans-Jörg Rheinberger Peter McLaughlin Staffan Müller-Wille

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Cabbage, Tulips, Ethiopians – “Experiments” in Early Modern Heredity Staffan Müller-Wille

0. Introduction Genealogy is the oldest logic. Not only do the key concepts of ancient logic – genos and eidos – have genealogical connotations, but even the way in which these concepts were seen to relate deductively was modelled on genealogy: eidos and genos relate in the same way “as Agamemnon is an Atride, a Pelopide, a Tantalide, and finally of Zeus”, as it says in Porphyry’s Isagoge to Aristotle’s Categoria.1 Some identity persists in descending genealogies, and thus things can be inferred for individuals by retracing their origin (arch or principium, again a term with a genealogical connotation) through the chain of their antecedents. 2 And yet, the old world, as is well known, was a world full of monstrous births, strange transmutations and unnatural copulations. Though Aristotle, e. g., repeatedly maintained that “a man is generated by a man”,3 he equally conceded the possibility that “two animals different in species produce offspring which differs in species; for instance a dog differs in species from a lion, and the offspring of a male dog and a female lion is different in species”. 4 And still in 1690 we see no one less than John Locke resuming in his Essay concerning human understanding that he “once saw a Creature that was the Issue of a Cat and a Rat, and had the plain marks of both about it.” 5 As fantastic as this promiscuity of nature – in which every combination seemed as plausible as possible – may seem to modern eyes, it rested on a rational ground: The foundation for the similarity among parents and offspring was provided by the recurrence of similar physiological and climatic conditions during procreation and development, which, inversely, meant that any deviation from this ordinary course of things – e. g. mesalliances as the ones referred to by Aristotle and Locke – would produce as deviant results. “All things are governed by law” is the conventional translation for the opening sentence of a Hippocratic tract De genitura.6 Yet, it is worthwhile to consult its Renaissance Latin translation: “Lex quidem omnia corroborat” – “Law strenghtens” – the original indeed has “kratunei”, which signifies both strengthen and govern – “everything”, with “law (nomos)” – as the commentator Girolamo Mercuriale carefully noted – 1

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Porphyrius (1887), 2b1-4, p. 6. The full text in its Latin translation by Boethius (ibid, p. 31) is: “ea vero quae sunt ante specialissima usque ad generalissimum ascendentia et genera dicuntur et species et subalterna genera, ut Agamemnon Atrides et Pelopides et Tantalides et ultimum Jovis.” Cf. Porphyrius (1998), I 1-3, II 9. Cf. Heinrich (1981), pp. 98-100, also Gayon and Wunenburger (1995), p. 8: “[...] la valorisation des filiations historiques, des ascendances et descendances d’un être, a longtemps servi de vecteur quasi unique d’intellegibilité canonique”. E. g. Aristotle De gen. anim. 735a20. Cf. Lesky (1950), p. 139, for further references and discussion. Aristotle De gen. anim. 747b33-36. Aristotle refers to this as an “abstract argument” in the discussion of mules, which are infertile, yet, as he himself states (ibid 748a16), he gave a wealth of examples for such “hybrids” in both De generatione and Historia animalium. Cf. Zirkle (1935), pp. 15-17. (Locke ([1690] 1975), p. 451, §23. For a rich account on such “hybrids” through history see Zirkle (1935), for iconographical evidence also Daston and Park (1998). E.g. Lloyd (1978), p. 317. Cf. the French translation – “La loi gouverne tout” – in Hippocrates (1851), p. 471.

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meaning “customs, pasture, region, class (instituta, pascua, regionem, classem)” in the dialect of the Hippocratic texts.7 Genealogy was seen to consist less in relations resulting from the lawful transmission and redistribution of some hereditary material but rather in the local persistence of economical, political and social configurations, in the persistence of a “fabric”– as Claude LéviStraus has formulated it – “in which warp and filling yarn correspond to localities and tribes”. 8 In this context, heredity was at once something trivial and precarious: It was stabilised and reinforced by the persistence of its own, municipal bounds (local rules of marriage and residence), and yet remained infinitely open to disturbances by illicit transgressions of these bounds. 9 It is, I presume, behind this background that we must see the achievements of the 18th century in regard to the formation of a modern concept of heredity. Peter Bowler has rightly argued, I think, that one important condition had to be fulfilled, before one could speak of such a concept: a clear distinction had to be drawn “between the transmission of characters from one generation to the next and the process by which the characters are produced in the growing organism”. 10 As an indication of how difficult it is, indeed, to see this, I take it that one 18th century theory of organic reproduction, in which this precondition seems to have been fulfilled – if only to a certain degree, which is the very topic of this paper – has consistently been underrated, if not simply overlooked, by historians of 18th century theories of organic reproduction. 11 I speak of the theory of organic reproduction of Carolus Linnaeus. 1. Laws of generation One of the reasons for the neglect of this theory by modern historians may be the place and the form in which it first appeared: Linnaeus laid out his theory not – as one might expect – in a coherent tract on the physiology of generation, but in a set of short aphorisms introducing the first editions of his famous taxonomic works, the Systema naturae of 1735 and the Genera plantarum of 1736.12 Moreover, it did not enter the scene as an independent account on the mechanisms of propagation, but was contained in the definition of a central category of Linnaean taxonomy: the species. Let us first have a close look at this definition:

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Hippocrates (1588), p. 10 & 15. Cf. Stubbe (1965), pp. 18-21. I thank Volkmar Schüller and Friedrich Steinle from the Max-Planck-Institute for the History of Science, Berlin, for discussing this passage with me. Lévi-Strauss ([1962] 1969), p. 97. Cfr. ch. 2 “La logique des classifications totémiques” in Lévi-Strauss (1962), pp. 48-99, who explicitly subsumes “the naturalists and hermetics of antiquity and the middle ages: Galen, Plinius, Hermes Trismegistos, Albert the Great” (ibid, p.57) under his analysis. Bowler (1989), p. 6. Lippmann (1933), pp. 49, 63 & 78, Cole (1930), pp. 18 & 137 and Roger ([1963] 1993), p. 322 mention Linnaeus only in passing. Ritterbush (1964), pp.101-103, though discussing Linnaeus’ theory of (plant) reproduction at length, tries to side him in the debate between “ovists” and “pollenists”, an opposition Linnaeus discarded, as we shall see. Delaporte (1983), pp. 124-127 discusses the anthropomorphism of Linnaeus’ theory of reproduction, but otherwise misses its idiosyncrasies. Interestingly, and rightly so, as we shall see, Linnaeus is compeletely missing in François Duchesneau voluminous study of 18th century physiology. Far from reducing the impact of Linnaeus’ theory of reproduction, this place rather had the effect that it “swept all of Europe and North America”. Cf. Farley (1982), p. 5.

Cabbage, Tulips, Ethiopians – “Experiments” in Early Modern Heredity

There are as many Species as different forms produced by the Infinite Being in the beginning. Which forms afterwards produce more, but always similar forms according to inherent laws of generation; so that there are not more Species now than came into being in the beginning. Hence, there are as many Species as different forms or structures of Plants occurring today, those rejected which place or accident exhibits to be less different (varieties).13

This short, and in its brevity so concise, passage has been endlessly quoted as indicating Linnaeus’s adherence to a “typological” species concept.14 And yet, as a close reading reveals, it does not so much tell us, what species are, but rather how many there are on the assumption of certain “laws of generation” . If we analyse Linnaeus’s species definition with regard to these laws, three peculiar absences come to the fore, which on the one hand set off this theory from contemporary theories of generation and on the other hand indicate its propensity towards the modern concept of heredity: 1. The subjects of Linnaeus’s “laws of generation” are not substances – seminal principles, moules interieures or the like – but “forms or structures”. “Structura” – in contrast to the much more wider term “forma”, which the definition uses synonymously – means something highly specific in Linnaeus’s terminology: It designates the totality of characters differentiating the members of a species from all other species according to “all its parts [...] in four dimensions: number, form, proportion, position”.15 The similarity relation posited among parents and offspring “according to inherent laws of generation” is one of mere structural analogy, not of substantial contiguity. There are two peculiar consequences to this: First, that the traditional “three tier” set-up of the problem of heredity – heredity of generic, sexual and individual traits – does not play, as far as I can see, the least role in Linnaeus’s writings on organic reproduction. 16 Sexual and individual differences – ontogenetic as well as intraspecific differences, e. g. in the number of certain organs – simply collapse with generic differences as they become differences within one and the same structural whole.17 And second – as is also clear from the stress put on the number, rather than the essence, of species –, that the order reigning between species is not a contiguous “scale of being” but a simple juxtaposition of discrete entities, without overlaps or intermediates. 18 2. The second absence within Linnaeus’s theory of reproduction is the absence of “physiology”. If one follows the “laws of generation” referred to by the definition, up to their full explication in the 13

14 15 16 17

Linné (1737), Ratio operis §5, [p.2]: “Species tot sunt, quot diversas formas ab initio produxit Infinitum Ens; quæ deinde formae secundum generationis inditas leges produxere plures, at sibi semper similes, ut Species nunc nobis non sint plures, quam quæ fuere ab initio. Ergo Species tot sunt, quot diversæ formæ seu structuræ Plantarum, rejectis istis, quas locus vel casus parum differentes (Varietates) exhibuit, hodienum occurrunt.” For the locus classicus of this interpretation see Mayr (1957). Linnaeus (1736), §92, p.11. Cfr. ibid, § 326, p.32. Olby (1966), p. 1, describes this set-up, which, e. g., was still shared by Buffon, as one of the main obstacles on the way to the modern (Mendelina) concept of heredity. Thus Linnaeus designated sexual dimorphism as a “natural variety (varietas naturalis)”; Cf. Linnaeus (1736), § 308, p. 30. A particular good example for the collapse of generic, sexual, and individual differences is provided by the description of the “structure” of the genus Urtica in the Genera plantarum, where one line says: “Female flowers, either in one or two different plant individuals”. (Linnaeus 1737), p. 283. This collapse is why Linnaeus was prepared to accept “constant varieties (varietates constantes)” – i. e. distinct and constantly reproducing forms within one and the same species –, in later writings on the reproduction of plants (see Linnaeus ([1755] 1788), pp. 380-383). I will come back to that in the section on “Ethiopians”.

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very first two paragraphs of the Systema naturae of 1735, it becomes evident, that the substrate of the structural similarity posited between ancestors and descendants is made up of purely genealogical relations. All in all, these two paragraphs formulate three basic laws: a) that “individual living beings are propagated by the egg (viventia singula ex ovo propagari)”; b) that “each egg produces offspring similar to the parents (omne ovum producere sobolem parenti simillimam)”; and c) that “individuals are multiplied by generation (ex generatione multiplicantur individua)”.19 Translated into negative statements, the first two “laws” reject any possibility for spontaneous generation or transmutation: No generation can occur outside the context of genealogical relations and no generation can transgress the bounds of this context. The only change that does occur is one in the number of individuals. That “ ovum”, the fertilised egg, does not represent a physiological unit (in the sense, e. g., in which it does in William Harvey’s work), but simply that entity which mediates generations of parents and offspring, can be inferred from another paragraph of the introduction to the Systema naturae, where Linnaeus states that “natural bodies are made up of elements, but in a way inexplicable except for creation and laws of generation.”20 Consequently, Linnaeus did not side himself in the debate between animalculists/ pollenists and ovists, but rejected both versions of preformation with the (basically genealogical) argument, that both parents, male and female, enter into the procreative act to leave traces in the offspring (which, of course, was known to preformationists) and that otherwise “it remains an obscure matter, now and once, how generation or fertilisation happen”. 21 3. The third absence in Linnaeus’s theory of organic reproduction is explicit in his formulation of a species concept: “[T]hose [structures are to be] rejected which place or accident exhibit to be less different (varieties).” Though formulated in a rather awkward way, the message is clear: Linnaeus draws a distinction between structural differences distinguishing organisms on a generic (species) level and obeying the laws of generation, and structural differences distinguishing organisms on the level of individuals and being due to local, accidental causes – “climate, soil, heat, wind”, as a later formulation of this distinction in the Philosophia botanica (1751) has it.22 In modern words: Linnaeus drew a distinction between nature and nurture, the latter’s effects being excluded from the realm of “laws of generation”. Accordingly, and in stark contrast to contemporary theories of generation, Linnaeus’s theory of reproduction, even in its later, more elaborated versions, does not show the least trace of the age old theories of pangenesis and “inheritance of acquired characters”.23 I take all three abstractions – the absence of substance, the absence of physiology, and the absence of environment – to be hallmarks of the distinction of transmission and development 18

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That there are no intermediates between species and, by extension, genera is frequently stressed in Linnaeus’s Critica botanica; see, e. g. Linnaeus (1737a), §224, p. 29. As is well known, Linnaeus rejected the “scala” in favour of the map as representing relations among generic entities (cf. Linnaeus (1751), §77, p. 27). For a discussion see Rheinberger (1986) and Müller-Wille (1999), pp. 89-97. Barsanti (1995), p. 35, considers the map, instead of the scale, to be the “image plus apte à acceullir une logique de filiation”. Linnaeus (1735), §§1-2, [p. 1]. Ibid, §7, [p. 1]: “Naturalia illa ex elementis constructa, licet modo, praeter creationem & leges generationis, inexplicabili.” For Harveys concept of “ovum” as a fundamental physiological unit, which, e. g., also could effect spontaneous generation of living beings, see Jacob (1970), pp. 63/64. Linnaeus ([1746] 1749), p. 347: Quomodo fiat generatio, vel fecundatio, innumerae sententiae Physiologorum fuerunt; sed aeque ac olim obscura res est.” Cfr. ibid, p. 349. Linnaeus (1751), § 158, p. 100.

Cabbage, Tulips, Ethiopians – “Experiments” in Early Modern Heredity

which according to Bowler was vital for the formation of the modern (Mendelian) concept of heredity, with the important limitation, however, that in Linnaeus’s theory of organic reproduction “transmission” did not consist in the redistribution of independent characters, but in the universal persistence of a totality of characters, i. e. of “structure” (I will come back to this in the end of my paper). To be sure, also, Linnaeus later supplemented his theory with a physiological model of generation, which, interestingly, however, hypostatised the abstractions inherent to the earlier formulation of “laws of generation”: According to this model, all organisms consist of two, antagonistic substances: the inner “pith (medulla)”, which is propagated via the maternal line, and has a “power of infinite multiplication” (in animals, this substance corresponds to the nervous system). And the outer “bark (cortex)”, which is propagated via the paternal line and has the power to attract and conduct nutriments, thus nourishing and protecting the medulla, and thus controlling its growth. The medulla, to put it shortly, is a merely propagative, the cortex a merely distributive substance, and it is their balanced, yet inherently antagonistic interaction through which life is maintained and develops.24 One of the earliest formulations of this physiological model in the Philosophia botanica 1751 shows, how closely it was tied to Linnaeus’s peculiar theory of generation: The root [containing the pith] extends infinitely, until the integuments [i. e. the bark] break up at the top to form the flower, and the seeds develop as the continuation and the utmost end of vegetation. This seed falls down, sprouts, and sort of continues the plant at a different place. Thus similar offspring is produced as the tree produces the branch, the branch the bud, and the bud the plant; therefore the generation of plants is a continuation.25

As much as some of the assumptions within Linnaeus’s theory of reproduction seem to be evident in the context of modern biology – as the rejection of spontaneous generation and transmutation, or the distinction of environmentally induced variation and “genetic” determination – at their time they were very strong claims. The strength of these claims, however, stood in a peculiar contrast to their empirical foundation. In the Fundamenta botanica of 1736, e. g., the sentence “Omne vivum ex ovo” is just said to be “repeatedly asserted by reason and experience and confirmed by the cotyledons”.26 If this already seems to be a rather weak, arbitrary, and ad hoc substantiation of the claim in question, even more so do Linnaeus’s references to “experiences (experimenta)” in his first published essay on the reproduction of organisms, the 23

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See Zirkle (1946) for rich material of the occurrence of these beliefs in the eighteenth century. Linnaeus did speculate sometimes, that certain plant species had resulted from the prolonged exposition of other species to a different climate or to intense cultivation (cfr. Ramsbottom 1938). This was also the basis for his attempts to “acculturate” exotic plants like tea to the climate of Sweden (cf. Körner 1994). Linnaeus remained undecided, however, if these changes were not reversible, and thus just special cases of “varieties” (e. g. Linnaeus (1737a), §316, p. 255). See, e. g., Linnaeus ([1759] 1763). For a detailed discussion of this theory see Stevens and Cullen (1990). Linnaeus (1751), § 157, p. 99: “Radix extenditur in herbam inque infinitum, usque dum apice rumpantur integumenta in florem, formantque semen contiguum, ultimum terminum vegetationis; Hoc semen cadit, prognascitur, & in diverso loco quasi plantam continuat; hinc simillimam sobolem producit, uti Arbor ramum, Ramus gemmam, Gemma herbam; ergo Continuatio est generatio plantarum.” Ibid, §79, p. 37, identifies the extending root with the medullar, and the integuments with the cortical substance. Linnaeus (1736), §135, p. 16: “Omne vegetabile ex ovo (134) provenire dictitat ratio & experientia, confirmant cotyledones.”

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Sponsalia plantarum of 1746: To modern eyes they just appear as a potpourri of chance observations, poorly designed experiments, just-so stories and circular reasonings. 27 Yet, a closer look at these empirical references is necessary to identify a historical space for the formation of Linnaeus’s “laws of generation” and to reach a better understanding of their position in the cultural history of heredity. 2. Ethiopians The bulk of the Sponsalia plantarum tries to identify the sexual organs of plants in analogy to animals, and employs morphological arguments for this aim. Among this, however, we find a few, which refer to what we today would call hereditary phenomena. One of them is especially intriguing, as it appeals to an experiential background, that was shared by the learned since antiquity: the “blended” character of children resulting from interracial “crossings” among humans of African and European origin.28 Yet, Linnaeus did not leave it at a general reference to this experiential background, but quoted a particular example from literature in the following words: “A common Ethiopian detained in the Copenhagen prison and kindled with love for a maid, secretly slept with her. Pregnant from that, and after due time of bearing, was brought forth a child of male sex, which resembled the mother in the whiteness of skin all over the body, only the darker penis showing the kind of its father.” Barthol. cent. 4. obs. 5. Which all evinces, that the beginnings of the coming fetus by no means lie hidden in one sex only.29

That Linnaeus chose to quote this instance, rather than to report a general experience, has certainly to do with the “empirical weight” gained by the authority of its source: The quote is from the Historiarum anatomicarum rariorum centuria which Thomas Bartholin (1606-1680), influential professor of medicine and theology at Copenhagen university, published in four volumes 1656 and 1657, each volume containing a centuria (a hundred) of “histories”. This format, also used by Bartholin in his Epistolarum medicinalium à doctis vel ad doctos scriptarum 1663-1667, appears rather strange to modern eyes: Completely unrelated observations of various medical interest – dissection reports, observations of unusual phenomena, cures and receipts, clinical cases – follow each other without any attempt at a systematisation. The headings of the first ten histories – including the one quoted by Linnaeus – of the fourth centuria may illustrate this:

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Cf. Sachs (1875), p. 104-106, whose reading of the Sponsalia plantarum has long influenced the picture of Linnaeus as a poor empirist. That such “crossings” were common in the Mediterranean of Antiquity is clear, and thus we find frequent references to them in the writings of ancient natural philosophy (e. g. in Aristotle, cf. Stubbe (1965), p. 21). How continuous these contacts remained even in Central Europe of the Middle Ages is discussed at length in Martin (1993). Linnaeus ([1746] 1749), p. 349: “‚Æthiops cerdo in ergastulo Hafniensi detentus, amore puellæ servæ accensus, clanculum illam compressit. Gravida inde, legitimo partus tempore enixa est prolem virilis sexus, quæ matrem universo corpore cutis candore referebat, solus vero penis paternum genus nigrore commonstravit. Barthol. cent. 4. obs. 5.‘ Quæ omnia, rudimentum futuri fœtus neutiquam in uno tantum sexu delitescere, evincunt.”

Cabbage, Tulips, Ethiopians – “Experiments” in Early Modern Heredity

I

Anatome Civettae (Dissection of a civet cat)

II

Prolapsus uteri cum urinae incontinentia (Prolapsus with incontinence)

III

Convulsiones paralyticae (Paralytic convulsions)

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Unicornu Groenlandicum (Greenlandian unicorn)

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Ex Aethiope natus (Birth from an Ethiopian)

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Anatome Hominis sanis (Dissection of a healthy man)

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Epilepsia ex vermibus (Epilepsy from worms)

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Calculus ex scroto suppuratu (Stone gathered from a scrotum)

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Emplastra magna (Large plasters)

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Anatomi Monachi (Dissection of a monk)

This “aphoristic” make-up was not used in deficiency of a system or theory, as Lorraine Daston has argued, but rather to avoid the pitfalls of speculation, to immunise the gathering of observations from rash conjecture and system building to enhance their “facticity”. 30 Yet this very format entailed a difficulty in regard to the empirical foundation of Linnaeus’s theory of generation, which comes to the fore, if we compare Linnaeus’s quote with the original text of Bartholin: A common Ethiopian detained in the Copenhagen prison and kindled with love for a maid, secretly slept with her. Pregnant from that, and after due time of bearing, was brought forth a child of male sex, which resembled the mother in the whiteness of skin all over the body, only the darker penis showing the kind of its father, which several people eye-witnessed and wondered at. I assign this wholly to the imagination of the mother, which seizing the desired part with a fixed and vigorous mind, impressed its colour on the offspring. Of mixed colour are children otherwise brought forth from an Ethiopian and a white, which often shows us, how both sexes have their separate commands over generation.31

What Linnaeus leaves out in his quote is what is interesting to Bartholin, while what Linnaeus tries to prove by his quote is what Bartholin takes for granted, as something known anyway. Bartholin concentrates on those features of the case reported – the fact that the colours do not blend in the child, as is usual, but remain seperated – which single it out as a singular case. And consequently, he is looking for singular circumstances in this case (the passion of the mixed couple, which, as it seems, was looked upon as something unusual by Bartholin) in search for an explanation. Only by leaving out this argument, and by reformulating the general conclusion, could Linnaeus adapt Bartholin’s observation to his theory of generation in which a combination of structural 30 31

Cf. Daston (1998). Bartholinus (1657), pp. 220-221: “Æthiops Cerdo in Ergastulo Hafniensi detentus, amore puellæ servæ accensus, clanculum illam compressit. Gravida inde legitimo partus tempore enixa est prolem virilis sexus, qvæ matrem universo corpore cutis candore referebat, solis verò penis paternum genus nigrore prodebat, qvod oculati testes plures & viderunt & mirati sunt. Imaginationi matris id universum assigno, qvæ partem vehementis desideratam animo fixo comprehendens ejusdem colorem fœtui impressit. Mixti |221| aliàs coloris solent fœetus ex Æthiope & alba procreari, qvod sæpe nobis visum, si qvidem uterq; sexus divisum in generatione imperium habet.”

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differences in the offspring rather was to be expected than posed a problem, and in which such developmental causes as “maternal imagination” did notably not play a role. Quite in line with this, the problem that Linnaeus saw in the well known case of “Ethiopians” was not to explain that and how their skin colour persisted, but rather that they possessed a distinctive character – black colour – that remained constant even under varying climatic and geographic conditions, while they nevertheless doubtlessly belonged to the same species as other humans: “Who would deny that the Ethiopian is of the same species as we humans”, as it says in a paragraph discussing difficulties in distinguishing varieties from species in the Critica botanica (1737), “and yet the Ethiopian brings forth black children in our countries.”32 What this shows, is that something more than isolated observations of hereditary phenomena had to enter the scene before Linnaeus’ could formulate his “laws of generation”. These laws, as we saw, transcended the bounds of individual parentage, and only in hindsight could the numerous isolated instances of inheritance reported in medicine and natural history appear to Linnaeus as approving his laws. What is missing is an empirical context in which the hereditary relations constitutive of Linnaeus’ theory of organic reproduction were actively implemented to reach beyond the exceptional and the ordinary course of things. The problem of the historical formation of this theory is one of synthesis, of how to knit together hereditary phenomena to form a network of relations rather than a number of individual cases.33 3. Tulips For understandable reasons, this problem of synthesis could not be solved in anthropology. A much better candidate for that would have been plant and animal breeding, and the Sponsalia plantarum indeed abound with examples taken from this ancient realm of technology. A particular interesting example, as it expressly referred to an “experiment”, is the following: TULIP. Delightful is this horticultural experiment: If someone perchance rejoices in completely red tulips and tears out the anthers from some flower before the pollen is scattered, and afterwards takes a tulip with a white flower and sprinkles the other, red one’s stigma with its anthers; and finally puts the ripe seeds in their own bed, he will obtain in this bed some red, some white, and for the greatest part two coloured flowers, no less than variously coloured offspring is produced from two animals of different colour.34 32

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Linnaeus (1737a), §316, p. 255: “Quis neget æethiopes esse ejusdem speciei ac nos homines, tamen æthiops nigros procreat infantes, in nostra terra”. This problem of “constant varieties” became more and more pressing for Linnaeus, resulting in the development of a theory of hybridization to account for the rise of new species and varieties from 1751 on. Cf. Müller-Wille (1998). For Linnaeus’s classification of human races according to skin colour, which he already presented in his Systema naturae of 1735, see Sloan (1995). The “aphoristic” style of Bartholin resembles that of the “consilia”, a collection of court cases which was posthumously added to Paolo Zacchias’ Quaestiones medico-legales 1621-1650. This latter publication contained, among other topics, a systematic exploration of ancient and early modern theories about “the similarity and dissimilarity of children (De similitudine & dissimilitudine natorum)”, reaching the conclusion that both male and female have “equal potentials” in the “process of generation” (see Bajada (1988), pp. 23-60, who calls the consilia a collection of “experiments”; ibid. p. 31). This forensic context, in which Bartholin was active too (see Dansk Biografisk Leksikon, Copenhagen 1979, vol. 1, pp. 475-6) and which may very well have been a source of “synthesis” of individual cases, could not be explored in this paper.

Cabbage, Tulips, Ethiopians – “Experiments” in Early Modern Heredity

The “experiment” Linnaeus recounts here came from a well known historical background: Not only had the trade in tulip varieties – after their introduction to Europe by the Turks in the early sixteenth century, Matthias L’Obel already could include illustrations of 20 different sorts in his Kruydtboeck of 1581 – been responsible for the first emergence of widespread, public option trading and ensuing market crash in the Dutch “tulip craze” 1633-1637; 35 this trade also sparked off an intense activity of breeding new varieties of tulips which survived the “craze”. But far from praising this activity as giving insights into the “laws of generation”, we see Linnaeus complaining about it elsewhere, namely in his Critica botanica, curiously enough invoking just the example of tulips: The prime reason why [our] precursors came out with wrong species names only consisted in that they refused to distinguish natural characters and parts, or certain ones from sportive ones. As they accepted all characters, accidental and natural ones alike, they erected new species from the most insignificant character, from whence so much confusion, such a barbarity of names, such an accumulation of wrong species, that it were easier to clean the stable of Augias, than that of Botany. [...] Certainly, if each character would equally constitute new species, there would be no wiser and accurate Botanists among mortals than those FLOWER-LOVERS, who each year in tulips, primroses, anemones, daffodils and hyacinths alone present to the curious thousands [of plants] unknown to the Botanists, and hence new species. [However:] The Omnipotent Builder of Creation stood off from work on the seventh day, so that there is no new creation each day, but a continuous multiplication of things once created. He created one human, as the Holy Scripture teaches: but if the smallest character were enough, there would stand out thousands of human species today; there stand out namely [those] with white, red, black, doggish [?] hair; with white, pink, brown, black face; with erect, short, curved, snub, aquiline nose; there stand out giants and pygmies, fat and thin, straight and bowed, leprosic [?] and lame people etc. etc. Yet who would ever lightheadedly [?] call them different species? You see, therefore we assume certain characters, and look for the deceptive ones, which lead astray and do not change the thing. [... e. g.], Tournefort counts 93 Tulips (where there is only one) and 63 hyacinths (were there are only two), and no less do others sport in others.36

But not only do such “flower-lovers (anthophili)” as Tournefort – who in fact was counted among the greatest botanists in Linnaeus’s time – burden botany with trivial distinctions and wrong species; Linnaeus even believed that they were to count as no botanists at all: Flower-lovers and Botanists have the same objects in varieties, however, with the difference, that the Flower-lover enters the scene, where the Botanist leaves it. The latter, sort of weary of it, sets an end to the work; the former, vigorous, begins to build that he may reach the stars.37

34

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Linnaeus ([1746] 1749), p. 370: “TULIPA. Jucundum est horticulturæ experimentum: si forsan rubris tantum gaudeat Tulipis, in flore aliquo antheras omnes decerpat ante pollinis dispersionem, assumat deinde Tulipam flore albo hujusque antheris stigma alterius rubræ aspergat6; maturis deinde seminibus, eadem in areolam propriam projiciat, & in hac areola flores reportabit, alios rubros, alios albos, bicolores plerosque ceteros, haud secus ac ex duobus animalibus diversi coloris, fœtus variis decoratus coloribus producitur.” Examples from animal breeding – mules, hens, dogs and sheep – are listed on p. 349. Cf. Jessen (1864), pp. 256-257, Kulischer ([1929] 1988), pp. 319-320, Zirkle (1935), p. 88.

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And, as a comment to his own, consequent dictum, that the “botanist does not care for those fortuitious monstrosities and varieties (casuales monstroisitates varietatesque [...] non curat Botanicus)”, Linnaeus added in obvious allusion to his species concept: The number of species of Botanists remains the same, now or in the future, as when they poured out of the hand of the Omnipotent Creator. Of the Flower-lovers, however, new and different ones are produced each day from the species (as the Botanists call them), and descended from these they finally ruin them. To the former, therefore, [i. e. the species of Botanists], there have been set certain limits by nature, which cannot be transgressed; in the latter, however, there is an infinite sport of nature without end; the former’s species come from the all-wise hand of the Omnipotent, the latter’s varieties from the sport of nature, especially under the auspices of the gardeners. From hence the greatest difference between Botanists and Flower-lovers.38

I have quoted Linnaeus here at length, because it is not easy to see, why Linnaeus so polemically criticised the “Anthophili”, who, after all, seem to have provided him with a “delightful experiment” to back his theory of organic reproduction. It is not a difference in “scientificity”, as the mentioning of Tournefort as one of the proponents of the “Anthophili” proves.39 It is equally not a difference in the realm of objects studied by Botanists and flower-lovers respectively: Varieties are said to be as much a topic for botanists as they are for Flower-lovers. The best way to describe the divide raised between botanists and flower-lovers by Linnaeus is to say that it is a difference of aspect: While the botanist is interested in limits inherent to the “continuous multiplication of things once created ”, the flower-lover concentrates on quite the opposite, namely change and variety – the “ infinite sport of nature without end” – and the means to effect these, notably by gardening technologies.40 That this is not just nit-picking reasoning to reach some social distinction for botanists, but has important consequences on the level of theorising can be confirmed by a glance at the content of seventeenth and early eighteenth century horticultural literature: Far away from expounding something like “laws of generation” it indulges in the multifarious means of effecting the 36

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Linnaeus (1737a), § 259, p. 152-155: “Primaria causa, cur nomina specifica Antecessorum fallacia evaserint, sola in eo consistit, quod partes & notas naturales, ac certas, a ludicris distinguere recusarint. Cum autem assumserint omnes notas, accidentales & naturales indifferenter, indeque constituerint ob minimum notam, novam speciem, orta fuit tanta confusio, tanta nominum barbaries, tanta specierum falsarum accumulatio, ut facilius stabulum Augias purgare, quam Botanicen. [...]. Certe si omnis nota indifferens novam constituat speciem, nulli mortalium Botanici Sapientiores & acutiores ANTHOPHILIS, qui in solis Tulipis, Primulis, Anemonibus, Narcissis, Hyacinthis omni anno aliquot millia Botanicis ignotas, novas proinde, species ostendunt curiosis. Desistebat ab opere Creationis Omnipotens Conditor die septima, nec nova creatio omni die, sed continuata multiplicatio creatorum. Hominem creavit unicum, dictiante S. Scriptura: at si minima nota sufficiat, vel mille hominum species hodie prostant; prostant enim capillis albis, rubris, nigris, canis; facie alba, rosea, fusca, nigra; naso erecto, brevi, inflexo, simo, aquilino; prostanbt gigantes, pygmæi, obesi, macilenti, erecti, incurvi, tophosi, claudi &c. &c. sed quis leviter ianus hos distinctas diceret species. En itaque assumamus notas certas, & inquiramus notas fallaces, quae seducunt, nec variant rem, [...]. Tournefortius Tulipas 93. (ubi una est) & Hyacinthis 63. (ubi duo sint) numerat, nec minus sæpe in aliis alii luxuriarunt.” Ibid, § 306, p. 238: “Anthophilorum & Botanicorum in Varietatibus objecta eadem sunt, ea tamen cum differentia, ut Anthophilus incipiat scenam, ubi Botanicus definit; hic, dum lassus quasi, finem operi imponit; ille, vegetus struere incipit, ut astra petat.”

Cabbage, Tulips, Ethiopians – “Experiments” in Early Modern Heredity

“transmutation” of plants, like soil preparation, watering, application of heat and manure, grafting, transplantation etc. Thus we find Laurembergius setting out the following general principle in the first book of his Horticultura 1631: [...] for the prosperity and flourishing of gardens; for the growth and augmentation of fruits, flowers, and vegetables, we wish for two things: [...]: These are a benign sky and a fecund earth. The sky is the father of everything sown; the earth the mother.41

This is as far away as can be from the principles expounded in Linnaeus’s species definition. And it is clear, that, with such principles, the transmutation, or degeneration, and the spontaneous generation of plants would make perfect sense. And even though doubts in transmutation should grow towards the turn to the eighteenth century;42 and even though artificial pollination should more and more raise the interest of authors on gardening, the situation in horticultural literature seems to have remained basically the same up to the time of Linnaeus, i. e. artificial pollination should remain one among many means to meliorate plants, and it should basically be thought of in analogy to technological processes.43 And still in 1773, we see that Joh. Chr. Fabricius, one of the more famous of Linnaeus’s students, was able to announce similar principles as his teacher in a chapter on “Gartenbau”:44 Die unzähligen Abänderungen der Gewächse beobachtet der Gärtner. Wir verlangen die besten unter denselben, und man muß mehr auf die Cultur der Abänderung, als die Art selbst sehen. [...]. Die Ursachen dieser vielen Abänderungen untersucht der Gärtner nach dem Boden, dem Clima, der Cultur und vielleicht der Zeugung (generatio hybrida).

The distinction of horticulture as a basically technological discipline does of course not mean that the knowledge accumulated on the basis of horticultural practices was of no significance for Linnaeus’s theory of (plant) reproduction. Quite the contrary is true: In the Philosophia botanica Linnaeus explicitely stated that “culture is the mother of so many varieties, and thus also the best mean of examining varieties.”45 After all, he was, as director of the Uppsala university garden, also 38

39 40 41

42

Ibid, § 310, p. 245-246: “Species omnes Botanicorum eodem numero, quo hoc vel futuro tempore existent, ab Omnipotentis Creatoris manu profluxerunt: Anthophilorum autem a Speciebus (Botanicis dictis) omni die novæ & diversæ prognascuntur, & prognatae in priores tandem ruunt. Illis itaque impositi sunt limites a natura certi, ultra quos progredi nequeat; in his vero lusus infiniti naturae absque fine; Illorum species Sapientissima a Manu Omnipotentis, horum Varietates a Ludente natura, sub auspiciis præsertim Hortulanorum, prodiere. Hinc differentia inter Botanicum & Anthophilum maxima.” In his Genera plantarum Linnaeus even declared his dependance on Tournefort, by stating that he understood “no one but Tournefort and his school” (Linnaeus (1737b), § 11, [p. 6]). Cf. Linnaeus (1739), which is discussed in detail in Müller-Wille (1999), pp. 151-155. Laurembergius ([1631] o.J.), lib. I, cap. i, § 7, p. 38: “[...] ad hortorum prosperitatem, florentemq; constitutionem; ad fructum, florum, olerum felicem proventum & incrementa, duo adesse optamus [...]. Ea sunt Coelum benignum, & terra foecunda. Coelum satorum omnium Pater est. Terra mater.” In the foreword it says in ragard to wine “tot genera, tot species, tot deliciae, quot regiones, quot oppida & urbes”, something any wine afficiando would wholeheartedly agree to even today. See, e. g., Sharrock (1660), pp. 28-32, who, however, concludes at one instance that “it was reason we should believe the report [on a transmutation] of good artists in matters of their own faculty”. In Rudbeck jun. (1686), who was one of the teachers of Linnaeus, we find Laurembergius principles still expounded (p. 5) and spontaneous generation admitted (p. 8ff.). Rich material on the belief in “degeneration” till the end of the seventeenth century see Zirkle (1935), pp. 61-88.

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very much himself involved in gardening.46 Yet it shows, again, that something else had to enter the scene apart from the mere production of variety for the formation of his theory.47 4. Cabbage CABBAGE. “Baal (Rich.), gardener from Brainford, had collected an enormous amount of seeds from flowered cabbage [cauliflower?] in his garden and sold it to most of the gardeners living in the suburbs of London. But these, after having sown these seeds with care in fat earth, produced common, long-leafed cabbage, wherefore they lamented to have been betrayed and summoned the aforesaid Baal before the Westminster court, who was condemned by the decision of the judge not only to repay the price to them, but also to restore the waste of time and loss of land use.” Raj. hist. I. p. 42. The deed is not to be ascribed to that gardener Baal, but to an impregnation of his better cabbage by the common cabbage. Therefore, if someone possesses that better cabbage, he should not let it flower on the same bed with another one, so that the better one is not fecundated with pollen from the lower and the lower one is generated from the seeds.48

This third example for empirical references in Linnaeus’s Sponsalia plantarum also stems from horticulture, yet with a decisive, additional shift in comparison with the case of the tulips: The “criminal case” reported does not only encompass the local production of varieties (in which Baal seems to have been quite successful) but also their circulation. Its geographical localisation – in suburbanis Londini – is revealing in this respect: It was near this large city and – some decades earlier – near the cities of Holland that from the 1650ies onwards and due to population growth in the urban centers crop production changed from subsistence to commercial production, involving a separation of production and consumption and a consequent interposition of trade mechanisms.49 For this trade – either in the products themselves, or in seeds as an important means of production – it was vital, of course, that the products exchanged did not change due to their transport from one locality to the other. Baals failure to guarantee this for his seed variety – he did notably not fail in the production of this variety as such – was responsible for his trial and – most probably – subsequent ruin.50 There is something else noteworthy about the reference to Baal’s fate: As in the case of the Copenhagen Ethiopian discussed before, Linnaeus quotes from literature and adds an explanation 43

44 45 46 47

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Thus we see Richard Bradley, one of the “fathers” of plant sexuality and probably the source for Linnaeus’s report on the “delightful experiment” with tulips (cf. Roberts ([1929] 1965), pp. 65/66), speak of fertilisation as a “mixing of qualities” and a possible source of “adulteration”. Well into the 17th century, plant sexuality was generally denied, and even the fig, on which articial pollination had been used since Babylonian times, was regarded either as exceptional or as only analogically sexual (thus Laurembergius; cf. Prest (1988), pp. 81-84). Fabricius (1773), p. 45. Linnaeus (1751), § 316, p. 247: “Cultura tot Varietatum mater, optima quoque Varietatum examinatrix est.” Accordingly, he knew the horticultural literature very well; see Stearn (1976). A close study of seventeenth and early eigteenth century literature on gardening and animal husbandry is, I believe, the great lacunae in the history of biology, which I, for the purposes of this paper, could of course not fill. The two most important publications on this topic are Zirkle (1935) and Roberts ([1929] 1965), which, however, because of their interest in the prehistory of Mendelism, exclusively focus on early accounts of “hybridisation”. Henrey (1975), though voluminous and highly informative, rather serves bibliographical puposes.

Cabbage, Tulips, Ethiopians – “Experiments” in Early Modern Heredity

to the case in question that rests on (and thus supposedly validates) his own theory of (plant) reproduction. And again, we can observe that this addition is contrary to the context of the original source: The quote is from a book John Ray published in 1686 and which bore the monstrous title Historia plantarum Species hactenus editas aliasque insuper multas noviter inventas & descriptas complectens : in qua agitur primò De Plantis in genere, Earúmque Partibus, Accidentibus & Differentiis; Deinde Genera omnia tum summa tum subalterna ad species usque infimas, Notis suis certis & characteristicis definita, Methodo Naturae vestigiis insistente disponuntur, Species singulae accurate describuntur, obscura illustrantur, omissa supplentur, superflua resecantur, Synonyma necessaria adjiciuntur; Vires denique & usus recepti compendiò traduntur (not surprisingly Linnaeus should refer to this work as one of the few “universal” plant histories in his Bibliotheca botanica51). This title already indicates, that Brassica florida belonged to a context much more systematic than that of the Copenhagen Ethiopian, and it occupied a central position within this context: Other than Linnaeus, Ray added the case of Brassica florida under the heading “About the transmutation of species among plants (De Specierum in Plantis transmutatione)” as a problematic case to a chapter, which discusses the “so called specific differences of plants (De specifica (ut vocant) Plantarum differentia)”, thus revealing the translation of plants as a relevant field of empirical evidence for his discussion of “specific differences”. Interestingly, a central passage of this discussion reveals some close resemblances to Linnaeus species definition: As, namely, the difference in sex in animals is not enough to argue for a difference in species, because both sexes originate from the same kind of seed and not rarely from the same parents, although they differ from each other in many and insignificant properties; [...]: so, also, there is no surer sign for a specific conformity than that plants originate from the same seed, be it individually or specifically. For what differs in species perpetually serves its species and not does this [species] originate from that seed [of another species] or vice versa. Therefore I propose not to hold for different species of plants those that differ by the colour [...] of the flower alone. [...]. [Because these varieties] can be brought about by art and display, not less than by repeated translation from one place to the other and by irrigation with water tinged with some colour. [...] Laurembergius, a worthy and truthful man, wrote in his Horticultura that he experienced [this] often in pinks [...].52 48

49

50 51

Linnaeus ([1746] 1749), p. 370: “BRASSICA. Baal (Rich.) hortulanus Brainfordensis, ingentem copiam seminis Brassicæ floridæ in horto suo collectam, hortulanis quam plurimis in suburbanis Londini degentibus vendidit; At hi cum summa cura eadem semina terræ pingui commiserunt, brassicas longifolias vulgares ipsis produxere, quare se fraudatos queruntur, & prædicto Baal litem intendunt in foro Vestmonasteriensi, qui ex sententia judicum condemnatus est, non solum ut ipsis pecunias restitueret, sed jacturam temporis, & amissum terræ usum fructum resarciret. Raj. hist. I. p. 42. Facinus hoc hortulano Baal non adscribendum est, sed impræagnationi Brassicæ ejus optimæ a Brassica vulgari factæ. Quare si quis Brassicam possideat optimam, eandem cum alia in eadem areola florescere non sinat, ne præstantior vilioris polline fecundutur, & ex seminibus vilior generetur.” Slicher van Bath (1963), pp. 14/15, Grigg (1982), pp. 102ff. Grigg also notes a steep rise in the number of books published on agriculture in the sebventeenth century (ibid., p. 159). The distinction of the spheres of production/consumption and circulation is reflected in Sharrock (1660), p.3: “The end of the Artist is to Propagate and Improve”. On the emergence of seed trade see Webber (1968). Cf. Linnaeus (1751), § 12, p. 10.

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In contrast to Linnaeus’s axiomatic species definition this formulation of a criterion for “specific conformity (convenientia specifica)” reveals its experiential basis and a peculiar shift in regard to it: While Ray’s formulation already foreshadows the three “absences” we observed in Linnaeus – absence of substance, physiology and environment53– Laurembergius used the experiences Ray drew upon in quite the opposite direction: The diverse regions and parts of the earth are imbued with diverse liquids of diverse properties, with which the plant adorns itself, and as it draws new food who wonders that it also acquires a new nature because of the subordination and mutual proportion between that which nourishes and that which is nourished? Thus we see white flowers change into red, yellow, blue on dousing with liquid tinged with these colours, which attracted by the roots bring a similar shape to the flowers.54

Behind this peculiar shift lay two decades of botanical activity, in which Ray concentrated on two projects: cataloguing plant species both indigenous and exotic to England - with the Catalogus plantarum circa Cantabrigiam nascentium (1660) and the Methodus plantarum nova (1682) demarkating this project – and experimental studies into plant embryology and physiology. The aim of the first project was announced in the 1660 Catalogus as asking “by which similarity and by which characters [any unknown plant] coincides with its congeners (ex similitudine, & notis quibus cum congeneribus conveniret)”55. The experiments, carried out in collaboration with Francis Willoughby and Martin Lister, regarded the “motion of sap in trees” and operated by “bleeding” various trees to determine the flow directions of the “sap” under various conditions. 56 Parallel to that, Ray studied the structure of the seed plant, reaching the distinction of mono- and dicotyledons.57 Both cataloguing and experimenting on plants depended on a highly specific locality: the botanic garden, in which on the one hand plants from all over the world were collected and 52

53

54

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Ray (1686), lib. I, cap. xx, p. 40: “Sicut enim in Animalibus sexuum distnctio non sufficit ad speciei diviersitatem arguendam, quia sexusa uterque ex eodfem speciei semine, eisdémque non rarò parentibus orìtur, quamvis multis & insignibus Accidentibus inter se differant; [...]: sic pariter in plantis convenientiæ specificæ non aliud certius indicium est quàm ex semine ajusdem plantæ seu in specie seu in individuo oriri. nam quæ specie differunt speciem suam perpetu’ servant, neque hæc ab illius semine oritur, aut vice versa. Hinc pro distinctis plantarum seciebus non habendas censeo, 1. Quæ solo floris colore [...] differunt. [...] Denique arte & mangonio induci possunt, nimirum translatione iterata de loco in locum, & irrigitaione aquà colore aliquo imbutà. Nam P. Laurembergius, vir fide dignus, Horticult. cap. 28. Sect. 3. se in Caryophyllis sæpius expertum scribit [...].” Ray ended his unpublished “discourse on the specific differences of plants” with the following words, which clearly foreshadowed Linnaeus’s species definition: “By this way of sowing [“in rich soil”] may new varieties of flowers and fruits be still produced ad infinitum, which affords me with another argument to prove them not specifically distinct; the number of species being in nature certain and determinate, as is generally acknowledged by philosophers and might be proved also by divine authority, God having finished his works of creation, that is, consummated the number of species in six days” (Ray 1674), p. 173. Laurembergius ([1631] o.J.), cap. XIII, § vi, p. 77: “Diversae autem regiones & telluris partes diversis sunt imbutae humoribus, diversis proporietatibus, & novo cibo ali assuscit, quid mirum si & novam acquirat naturam, propter illam quae est inter alitum & alens subordinatium & proportionem mutuam? Ita videmus flosculos albos permutari in rubros, flavos, ceruleos, affusione humoris his coloribus imbuti; qui per radices attractus similem florem ideam conciliat.” Ray (1660), p. 5.

Cabbage, Tulips, Ethiopians – “Experiments” in Early Modern Heredity

compared (Ray mainly acquired seeds for his garden commercially from London), 58 and in which, on the other hand, plant specimens were produced under controlled and – for reason of experimental studies – manipulable conditions.59 Though Ray, in contrast to Linnaeus, remained inconclusive and tentative about his findings in both projects,60 it is here that we can identify a space in which hereditary relations could be observed to transcend local conditions: the botanic garden – with its peculiar double nature of a site for collection and experimentation– installed the same separation of production/consumption and circulation that emerged in commercial horticulture in the course of the 17th century,61 and it was thus the site that could provide the material space for the abstractions expressed in Linnaeus’s “laws of generation”. And thus we can see Ray expressing the same “structuralist” approach to species as Linnaeus: 62 It is to be observed, however, that the distinct propagation by seed is not by itself that which constitutes the essential or specific difference, or that in which it [i. e. the essential or specific difference] consists, but its sign or indication alone.

5. Conclusion: Linnaeus’s concept of heredity By looking at three empirical references adduced by Linnaeus to back his theory of organic reproduction and by retracing some of their background in 17th century medicine, horticulture and natural history, I hope to have provided some evidence that observations of hereditary phenomena occupied a much more systematic space than is usually acknowledged by the history of science,63 and that the organisation of this space – in the case of plants – hinged upon a separation of plant reproduction and circulation instituted by botanical gardens. With this background in mind it is worthwhile to return to Linnaeus concept of “heredity” as expressed in his species definition. It is striking that Linnaeus – at least to my knowledge – never 56 57 58

59

60

Willoughby and Ray ([1669] 1928); cf. Ray (1928), pp. 45-47. Ray (1674). See Raven (1942), p. 109. In respect to this collecting practice it is interesting to see, that Ray quoted – apart from Laurambergius – another author of a gardening book, Johann Baptista Ferrarius, who directed his book not to gardeners, but to “rei florae scriptoribus” (see Ferrarius (1638), p. 13) and included a “law of sowing flowers (lex floris serendi)” and a “law of propagating flowers (lex floris propagandi” in this book. The first “law” consisted, among other things, in prescriptions of how to produce a “paper garden (hortus papyraceus)” representing the garden and guranteeing the identification of the plants sown out over time. In regard to the second “law”, Ferrarius stated, that “two things are necessary in the affairs of the propagator, namely that he augments the domestic [plants] and that he acquired foreign ones (duplex in propagatore industria requiritur, ut flores domesticos augeat, externosque conquirat)” (ibid. p. 294), and that for this “trade in flowers (florum commercia) was necessary. The importance of the garden was emphasised by Ray in a letter to James Petiver in 1701 when complaining of the loss of access to gardens: “Since I came to prosecute the work [i. e. the second edition of Rays Methodus, which appeared in 1703] in good earnest, I have been in no case to travel to visit gardens, and to see plants growing, flowering, and seeding. Dried specimens, figures, descriptions, and names of plants, is all I have had to work by, so that I must needs be liable to commit a thousand mistakes.” Ray called the report he sent to the Royal Society in 1674 on plant embryology and physiology “inchoate & imperfect” (see Ray (1928), p. 68). Likewise, he warned the reader of his Methodus, not to “believe that it [i.e. his “universal method”] extends so far as to signify all plant genera over the whole world (Quod Methodum nostram generalem apello, cave credas me vocare hanc extendere ad omnia per totum terrarum orbem nascentia plantarum genera significanda.)” (Ray 1682), pp. 7/8).

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used this or similar concepts in the context of organic reproduction. I think there was a good reason for this: If the “structure” characterising organisms as belonging to certain species consists in nothing but properties which identify and differentiate organisms “universally”, i. e. irrespective of any physiological and environmental contiguity, there is no need to postulate any mechanism of heredity. The different “structures” postulated in Linnaeus species concept just persist through time, or rather time-lessly. In a way Linnaeus stood halfway between the traditional and the modern concept of heredity, by maintaining persistence as the mechanism responsible for the similarity among parents and offspring and at the same time reducing that which persists to universal relations of formal identity and difference. 64 For a historical moment therefore we see heredity vanishing as a “biological” concept. And yet, this moment inadvertently provided a decisive precondition for the emergence of modern heredity: The taxonomically defined species of Linnaeus was to serve as the substrate for the “transmission of characters from one generation to the next” characterising the Mendelian concept of heredity (thus Mendel spoke of plant species as the “Träger” of his experiments). 65 The role of taxonomy for the history of heredity still awaits its analysis. 66

61 62

63 64 65

22

Cf. Stearn (1961) and Wijnands (1988) for the development of exchange relations among European botanical gardens during the seventeenth century. Ray (1686), p. 42: “Notandum tamen distinctam propagationem ex semine non esse illud ipsum quod constituit differentiam essentialem seu specificam, sive in quo illa consistit, sed ejus signum seu indicium tantúm.” Sloan (1972) has connected John Rays well known scepticism about knowing the “essences” of plants with Lockean philosophy. Though Ray and Locke probably knew each other (they became fellows of the Royal Society the same day; see Ray (1928), p. 44), I doubt this interpretation. Rays scepticism concerned the possibility to get a knowledge of taxonomic criteria through physiological studies, not the possibility to determine taxonomic criteria as such. Locke, on the contrary, explicitely rejected the criterion of “constant” reproduction in his Essay concerning human understanding (see above, n. 5). Mary Slaughter has also situated Rays scientific projects in a “philosophical” context, namely the search for an “universal language” by John Wilkins, a context, to which Ray – in hindsight – referred himself (Ray 1703, p. 5). While I think, that, beyond this, Rays attempts have to be seen in the much more specific context of physiology and natural history, I share Slaughters general conclusion: “The isolation of these variables [i. e. structure, sign, essence] or the decomposition of the organism into these elements permits organic form to be reconstituted or retranslated into a linear language, into a series of successively ordered elements which constitute a taxonomy” (Slaughter 1982, pp. 9/10). Cf., e. g., Ritterbush (1964), p. vii. In this respect, Linnaeus’ theory of organic reproduction showed striking resemblances to the 18th century concept of the “genealogical state”, see Paulson Eigen (2000) and Müller-Wille (2000). Mendel (1866), p. 5.

Cabbage, Tulips, Ethiopians – “Experiments” in Early Modern Heredity

References Bajada, Joseph. 1988. Sexual impotence. The contribution of Paolo Zacchia (1584 - 1659), Analecta Gregoriana 252. Rom: Ed. Pontificia Univ. Gregoriana. Barsanti, Giulio. 1995. Schémas biologiques de la descendance (XVIIIe - XIXe siècles). In Le paradigme de la filiation, edited by J. Gayon and J.-J. Wunenburger. Paris: L’Harmattan. Bartholinus, Thomas. 1657. Historiarum anatomicarum rariorum centuria III et IV ejusdem cura accessere observationes anatomicae Cl. Viri Petri Pawi. Hafniae: Haubold. Bowler, Peter J. 1989. Mendelian Revolution: the emergence of hereditarian concepts in modern science and society. Baltimore: John Hopkins Univ. Pr. Cole, F. J. 1930. Early theories of sexual generation. Oxford: Clarendon Pr. Daston, Lorraine. 1998. The Language of Strange Facts in Early Modern Science. In Inscribing Science. Scientific texts and the materiality of Communication, edited by T. Lenoir. Stanford: Stanford Univ. Pr. Daston, Lorraine, and Katharine Park. 1998. Wonders and the order of nature : 1150 - 1750. New York: Zone Books. Delaporte, François. 1983. Das zweite Naturreich. Über die Fragen des Vegetabilischen im XVIII. Jahrhundert. Frankfurt/M. etc.: Ullstein. Fabricius, Johann Cristian. 1773. Anfangs-Gründe der oeconbomischen Wissenschaften zum Gebrauch academischer Vorlesungen. Flensburg: Serringhausen. Farley, John. 1982. Gametes & Spores. Ideas about Sexual Reproduction. 1750-1914. Baltimore - London: John Hopkins Univ. Pr. Ferrarius, Johann Baptista. 1638. Flora s. de Florum cultura. Amsterdam. Gayon, Jean. 1995. Entre force et structure: genèse du concept naturaliste de l’hérédité. In Le paradigme de la filiation, edited by J. Gayon and J.-J. Wunenburger. Paris: L’Harmattan. Gayon, Jean, and Jean-Jacques Wunenburger. 1995. Présentation. In Le paradigme de la filiation, edited by J. Gayon and J.-J. Wunenberger. Paris: L’Harmattan. Grigg, D. 1982. Dynamics of Agricultural Change. Heinrich, Klaus. 1981. Tertium datur : eine religionsphilosophische Einf. in d. Logik. Dahlemer Vorlesungen. Band 1. Basel – Frankfurt/M.: Stroemfeld/Roter Stern. Henrey, Blanche. 1975. British Botanical and Horticultural Literature before 1800. Comprising a history and bibliography of botanical and horticultural books printed in England, Scotland, and Ireland from the earliest times until 1800. 2 vols. London: Oxford Univ. Pr. Hippocrates. 1588. De genitura. In Opera Hippocratis Coi qvae Graece et latine extant. Venetiis: Apud iuntas. ———. 1851. De la génération (Peri gones). In Oevres complètes d’Hippocrate, edited by É. Littré. Paris: J.B. Baillière. Jacob, François. 1970. La logique du vivant. Paris: Gallimard. Jessen, Karl F.W. 1864. Botanik der Gegenwart und Vorzeit in culturhistorischer Entwicklung. Ein Beitrag zur Geschichte der abendländischen Völker. Leipzig: Brockhaus. Körner, Lisbet. 1994. Nature and nation in Linnaean travel, Dissertation Abstracts International, Dissertation at Harvard Univ., 1993. Adviser: Simon Schama. Univ. Microfilms order no. 94 – 12358. 480 pp. Kulischer, Josef. [1929] 1988. Allgemeine Wirtschaftsgeschichte des Mittelalters und der Neuzeit. Bd. 2: Die Neuzeit. München und Berlin. Laurembergius, Petrus. [1631] o.J. Horticultura, Libris II. comprehensa; huic nostro coelo & solo accomodata; Regulis Observationibus, Experimentis, & Figuris novis instructa: in qua quicquid ad hortum proficue colendum, et eleganter instruendum facit, explicatur. Francofurti ad Moenam: Merian. Lesky, E. 1950. Die Zeugungs- und Vererbungslehren der Antike und ihr Nachwirken. Abhandlungen der Geistes- und Sozialwissenschaftlichen Klasse der Akademie der Wissenschaften und der Literatur in Mainz, Jahrgang 1950, Nr. 19. Wiesbaden: Franz Steiner. Lévi-Strauss, Claude. 1962. La pensée sauvage. Paris: Plon. ———. [1962] 1969. Das Ende des Totemismus. Vol. 128, edition suhrkamp. Frankfurt/M.: Suhrkamp. Linnaeus, Carl von. 1735. Systema Naturae, sive Regna Tria Naturae systematice proposita per classes, ordines, genera, & species. Lugduni Batavorum: de Groot. 66

Contributions in that direction are Barsanti (1995), p. 35, who maintained “une priorité de classification sur la pense généalogique”, and Gayon (1995), who described the transition from a concept of heredity as “force” to a concept of heredity as “structure” as the main development in the formation of 20th century heredity.

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———. 1736. Fundamenta Botanica quae Majorum Operum Prodromi instar Theoriam Scientiae Botanices per breves Aphorismos tradunt. Amstelodami: Solmon Schouten. ———. 1737a. Critica botanica in quo nomina plantarum generica, specifica, & variantia examini subjicuntur, selectiora confirmantur, indigna rejicintur; simulque doctrina circa denominationem plantarum traditur. Seu Fundamentorum Botanicorum pars 4. Lugduni Batavorum: Wishoff. ———. 1737b. Genera plantarum Eorumque characters naturales Secundum numerum, figuram, situm, proportionem Omnium fructificationis Partium. Lugduni Batavorum: Wishoff. ———. 1739. Rön om växters plantering grundat på Naturen. Kungliga Svenska Vetenskaps-Akademiens Handlingar 1:5-24. ———. [1746] 1749. Sponsalia plantarum [Diss., Resp. J. G. Wahlbom]. In Caroli Linnaei Ammoenitates academicae, seu Dissertationes variae Physicae, Medicae, Botanicae antehac seorsim editae. Holmiae et Lipsiae: Godofredus Kiesewetter. ———. 1751. Philosophia botanica in qua explicantur Fundamenta Botanica cum definitionibus partium, exemplis terminorum, observationibus rariorum [...]. Stockholmiae: Kiesewetter. ———. [1755] 1788. Metamorphoses plantarum [Diss., Resp. N. E. Dahlberg]. In Caroli Linnaei Amoenitates academicae, seu Dissertationes variae Physicae, Medicae, Botanicae antehac seorsim editae. Erlangae: Jo. Jacobus Palm. ———. [1759] 1763. Generatio ambigena [Diss., Resp. C. L. Ramström]. In Caroli Linnaei Ammoenitates academicae, seu Dissertationes variae Physicae, Medicae, Botanicae antehac seorsim editae. Holmiae: Laurentius Salvius. Lippmann, Edmund O. 1933. Urzeugung und Lebenskraft. Zur Geschichte dieser Probleme von den ältesten Zeiten an bis zu den Anfängen des 20. Jahrhunderts. Berlin: Julius Springer. Lloyd, G. E. R., ed. 1978. Hippocratic Writings. Pelican classics. Harmondsworth: Penguin Books. Locke, John. [1690] 1975. An Essay Concerning Human Understanding. Oxford: Clarendon Pr. Martin, Peter. 1993. Schwarze Teufel, edle Mohren. Afrikaner in Bewußtsein und Geschichte der Deutschen. Hamburg: Junius. Mayr, Ernst. 1957. Species Concepts and Definitions. In The Species Problem, edited by E. Mayr. Washington D.C.: American Association for the Advancement of Science. Mendel, Gregor. 1866. Versuche über Pflanzen-Hybriden. Verhandlungen des Naturforschenden Vereins zu Brünn 4:3-47. Müller-Wille, Staffan. 1998. ‘Varietäten auf ihre Arten zurückführen’ - Zu Carl von Linnés Stellung in der Vorgeschichte der Genetik. Theory in Biosciences 117:346-376. ———. 1999. Botanik und weltweiter Handel. Zur Begründung eines Natürlichen Systems der Pflanzen durch Carl von Linné (1707-1778). Edited by O. Breidbach and M. Weingarten, Studien zur Theorie der Biologie Band 3. Berlin: Verlag für Wissenschaft und Bildung. ———. 2000. Genealogie, Naturgeschichte und Naturgesetz bei Linné und Buffon. In Genealogie als Denkform in Mittelalter und Früher Neuzeit, edited by K. Heck and B. Jahn. Tübingen: Niemeyer. Olby, Robert C. 1966. Origins of Mendelism. London: Constable. Paulson Eigen, Sara. 2000. A Mother’s Love, a Father’s Line: Law, Medicine and the 18th-Century Fictions of Patrilineal Genealogy. In Genealogie als Denkform in Mittelalter und Früher Neuzeit, edited by K. Heck and B. Jahn. Tübingen: Niemeyer. Porphyrius. 1887. Isagoge et in Aristotelis Categoria commentarium. Übers. von Boethius. Hrsg. von Adolf Busse. Vol. 4.1, Commentaria in Aristotelem Graeca. Berolini: Reimer. Prest, John. 1988. The Garden of Eden: the botanic garden and the re-creation of paradise. New Haven: Yale Univ. Pr. Ramsbottom, J. 1938. Linnaeus and the Species Concept. Proceedings of the Linnean Society London 150:192219. Raven, Charles E. 1942. John Ray, naturalist. His life and works. Cambridge: Cambridge Univ. Pr. Ray, John. 1660. Catalogus plantarum circa Cantabrigiam nascentium : in quo exhibentur quotquot hactenus inventæ sunt, quæ vel sponte proveniunt, vel in agris seruntur : unà cum synonymis selectioribus, locis natalibus & observationibus quibusdam oppidò raris : adjiciuntur in gratiam tryonum, index anglicolatinus, index locorum, etymologia nominum, & explicatio quorundam terminorum Index plantarum agri Cantabrigiensis. Cantabrigiæ: Excudebat J. Field. ———. 1674. A discourse on the seeds of plants. In The History of the Royal Society of London, edited by T. Birch. London: Millar. ———. 1674. A discourse on the specific differences of plants. In The History of the Royal Society of London, edited by T. Birch. London: Millar. ———. 1682. Methodus plantarum nova, brevitatis & perspicuitatis causa synoptice in tabulis exhibita : cum notis generum tum summorum tum subalternorum characteristicis, observationibus nonnullis de feminibus

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plantarum & indice copioso. Londini: Faitborne & Kersey. ———. 1686. Historia plantarum Species hactenus editas aliasque insuper multas noviter inventas & descriptas complectens : in qua agitur primò De Plantis in genere, Earúmque Partibus, Accidentibus & Differentiis; Deinde Genera omnia tum summa tum subalterna ad species usque infimas, Notis suis certis & characteristicis definita, Methodo Naturae vestigiis insistente disponuntur, Species singulae accurate describuntur, obscura illustrantur, omissa supplentur, superflua resecantur, Synonyma necessaria adjiciuntur; Vires denique & usus recepti compendiò traduntur. 3 vols. Londini: Clark & Faithorne. ———. 1703. Methodus plantarum emendata et aucta. Londini: Smith & Walford. ———. 1928. Further Correspondence of John Ray. London: Ray Society. Rheinberger, Hans-Jörg. 1986. Aspekte des Bedeutungswandels im Begriff organismischer Ähnlichkeit vom 18. zum 19. Jahrhundert. History and Philosophy of the Life Sciences 8:237-250. Ritterbush, Philip C. 1964. Overtures to Biology. The Speculations of Eighteenth Century Naturalists. New Haven & London: Yale Univ. Pr. Roberts, H. F. [1929] 1965. Plant Hybridization before Mendel. New York - London: Hafner. Roger, Jaques. [1963] 1993. Les sciences de la vie dans la pensée française du XVIIIe siècle. Paris: Albin Michel: Biblioth`que de synthèse historique. Rudbeck jun., Olaus. 1686. De propagatio plantarum botanico-physicae [Diss.]. Upsaliae. Sachs, Julius. 1875. Geschichte der Botanik vom 16. Jahrhundert bis 1860. Edited by H. C. b. d. K. A. d. Wissenschaften. Vol. 15, Geschichte der Wissenschaften in Deutschland, Neuere Zeit, Bd 15. München: Oldenbourg. Sharrock, Robert. 1660. The History of the Propagation and Improvement of vegetables. By the concurrence of Art & Nature: Showing the several ways for the cultivation of Plants usually cultivated in England [...]; with the effect of Nature, and her manner of working upon the several Endeavours and Operations of the Artist. Oxford: Robinson. Slaughter, M.M. 1982. Universal languages and scientific taxonomy in the seventeenth century. Cambridge etc.: Cambridge Univ. Pr. Slicher van Bath, and Bernard Hendrik. 1963. The agrarian history of western Europe. London: Arnold. Sloan, Phillip Reid. 1972. John Locke, John Ray, and the Problem of the Natural System. Journal of the History of Biology 5 (1):1-55. ———. 1995. The Gaze of Natural History. In Inventing Human Science. Eighteenth-Century Domains, edited by C. Fox, R. Porter and R. Wokler. Berkeley etc.: Univ. of California Pr. Stearn, William T. 1961. Botanical Gardens and Botanical Literature in the Eighteenth Century. In Catalogue of Botanical Books in the collection of Rachel McMasters Miller Hunt., edited by J. Quinby and A. Stevenson. Pittsburgh/PA. ———. 1976. Carl Linnaeus and the Theory and Practice of Horticulture. Taxon 25 (1):21-31. Stevens, P. F, and S. P Cullen. 1990. Linnaeus, the cortex-medulla theory, and the key to his understanding of plant form and natural relationships. Journal of the Arnold Arboretum 71:179-220. Stubbe, Hans. 1965. Kurze Geschichte der Genetik bis zur Wiederentdeckung der Vererbungsregeln Gregor Mendels. Edited by H. Stubbe. 2 ed. Vol. Beitrag 1, Genetik. Grundlagen, Ergebnisse und probleme in Einzeldarstellungen. Jena: VEB Fischer. Webber, Ronald. 1968. The Early Horticulturalists. Newton Abbot: David and Charles. Wijnands, D. Onno. 1988. Hortus auriaci: the gardens of Orange and their place in late 17-th century botany and horticulture. Journal of Garden History 8:61-86, 271-304. Willoughby, Francis, and John Ray. [1669] 1928. Experiments concerningh the motion of the Sap in Trees, made this spring. In Further Correspondence of John Ray, edited by R. W. T. G. Gunther. London: Ray Society. Zirkle, Conway. 1935. The Beginnings of Plant Hybridization. Philadelphia: Univ. of Pennsylvania Pr. ———. 1946. The Early History of the Idea of the Inheritance of Aquired Characters and of Pangenesis. Transactions of the American Philosophical Society. New Series 35 (2):91-151.

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Speculation and Experiment in Enlightenment Life Sciences Mary Terrall

Although, as Carlos López-Beltrán has argued convincingly, heredity was not a coherently theorized concept until the 1830s, the inheritance of particular traits entered into quite a range of discussions about the nature and organization of living matter in the Enlightenment. Since the scientific community was not yet parsed strictly by discipline or profession, it is not surprising that these discussions easily crossed lines that later would become more impermeable. In addition to the writings of physicians on the cause (or even existence) of hereditary diseases, hereditary phenomena from animal breeding and from observations of human families made their way into in works of various genres that addressed the vexed question of generation. 1 Precisely because biology was not yet constituted as a discipline, this topic was open to interpretation, speculation and experimentation by a range of writers including philosophically-minded materialists (or crypto-materialists) as well as practicing physicians, anatomists, and observers of animal behavior. Inheritance of traits, whether abnormal or not, was one kind of evidence mustered in the discourse about generation and the origin of organization. Much of this discourse remained speculative, but some writers drew on empirical evidence to buttress their conjectures about unseen forces and hidden mysteries. Ideas and experiments related to the passing of traits from parent to offspring thus belong in a cultural context that includes a shifting and fluid discourse about the nature of life. The obscurity of the principles governing generation and organization meant that the means for uncovering them were as fraught as the elusive principles themselves. Turning to the Encyclopédie article on generation, for example, we find first a description of the mechanics of copulation, and then the reflection that the really interesting part of the process is not mechanical, but “physical.” The physical lies at a more fundamental level than the mechanical, apparently, and “nature employs the most secret means, the least available to the senses, to put fertilization into operation.” Ironically, the most sensual of phenomena masks a process beyond the reach of empirical investigation. This mystery has always excited the curiosity of physiciens and has led them to conduct so many investigations in order to penetrate it, so many experiments in an effort to take nature in the act; ... they have imagined so many different systems, which have destroyed each other successively, ... without resulting in more light on the subject. On the contrary it seems that the veil behind which nature hides herself is essentially impenetrable to the eyes of the subtlest mind, and that the cause of the formation of animals must be ranked among first causes, like that of motion and gravity, of which we will never be able to know anything but the effects.2

Diderot, too, in his article “Animal” referred to the ordered succession of generations by which species preserve themselves as “the greatest marvel”. He went on to rhapsodize, “The machine is 1 2

On the medical discourse about heredity López-Beltrán (1995); see also López-Beltrán (1994). Aumont (1757), p. 568.

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finished, and the hours strike under the eye of the clockmaker. But among the sequences of the mechanism, we must admit that this faculty of animals and vegetables to produce their kind, ... this procreative power that operates perpetually ... is for us ... a mystery whose depths it seems we will not be allowed to sound.”3 Here Diderot articulates the tension between predictable clockwork mechanism and the “procreative power” essential to life. The sense of mystery, and the recalcitrance of the problem, pervaded discussion of generation and hereditary phenomena in this period. If we are to understand the efforts of 18th-century thinkers to explore this mystery, we need to keep this in view. Otherwise we risk reading these writers as simply groping towards 19 thcentury biology, an approach that ignores the complexity of their efforts and the meanings attached to them by their readers. The suspicion that the depths of the mystery might remain beyond the reach of human understanding did not keep people from trying to sound them. By the mid-18 th century, whenever the question of how to understand the generation of living forms came up, it brought along related philosophical, theological and methodological freight. Empirical evidence came from anatomical observations going back to Harvey, Malpighi, Leeuwenhoek and others working in the previous century. By the time the volumes of the Encyclopédie started to appear, in 1751, the ramifications of the problem of generation extended across several disciplines, and involved overlapping sets of medical men, academic anatomists, experimenters, natural historians and philosophers. It was not a problem that found a ready solution and no consensus was reached. Some were willing, even eager, to peer beyond the visible (behind the veil, as it were) to speculate about fundamental forces and principles; others recoiled with something like repugnance. The interplay between eager speculation and prudent restraint characterized this discursive terrain, and gave an edge of danger, a whiff of suspicion, to theories of generation that went beyond the mechanical. I look here at competing theories and interpretations in the context of related debates about method, and especially about how speculation (or “conjecture,” in the terminology of the day) was interconnected with experiment and observation of organisms. Claims about method – what was allowable or legitimate – played into hotly contested questions that stretched the physical toward the metaphysical. Defining a science of life – setting parameters for how to go about investigating organic phenomena like generation – was inseparable from the question of the nature of life itself, which inevitably shaded toward claims with moral or religious implications. The investigation of inherited traits also entailed reflection on the properties of matter, not to mention the role of God in the universe. In the spectrum of discourse about generation, and the boundary between living and inert, we find conflict not just about content of theories, or interpretations of experiments, but about what is thinkable, what is dangerous, what is threatening or liberating. The disjuncture between the mechanical and the living inspired attention to generation and inheritance, and especially the arguments against the pre-existence of germs, whose “development” was only a

3

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“La machine est faite, & les heures se marquent sous l’oeil de l’horloger. Mais entre les suites du mécanisme, il faut convenir que cette faculté de produire son semblable qui réside dans les animaux and dans les végétaux, ... cette vertu procréatrice qui s’excerce perpétuellement sans se détruire jamais, est pour nous ... un mystere dont il semble qu’il ne nous est pas permis de sonder la profondeur.” (Diderot 1751).

Speculation and Experiment in Enlightenment Life Sciences

mechanical unfolding of forms already organized. Getting behind the mechanical, for some at least, meant going beyond the visible and rehabilitating conjecture, or speculation. Theories of generation Debates about generation revolved around two sets of oppositions: female eggs and male sperm (what is contribution of each sex?), and pre-existence and epigenesis. 4 Pre-existence theories, the main contenders in the aftermath of microscopical/anatomical work at the end of the 17 th century, located a fully organized creature in the germ, whether in egg or sperm. The organism was assumed to be contained in sub-microscopic particles, only just beyond the reach of microscope-enhanced vision. The origin of organization was, then, referred back to God, and therefore not open to scientific investigation. Any alternative to this way of thinking had to assign a cause (whether physical or metaphysical) responsible for getting unorganized material particles organized into a functioning animal. Epigenesis, for example, assigned responsibility for the process of organization to properties or forces associated with elements of organic matter. Organisms formed from unorganized matter, rather than unfolding or expanding according to pre-ordained plan. The problem was how to get at the non-mechanical aspect of the process without raising other sorts of philosophical obscurity. Suffice it to say that recent developments in physics (gravity and electricity) and chemistry (selective affinities driving reactions) made it conceivable to associate forces with matter, in very un-Cartesian ways. In spite of the obvious conceptual difficulties (all future organisms had to be present either in Eve or in Adam; or else seeds have to be created directly by God in each generation) pre-existence was compelling because it avoided the problem of self-acting matter. This was not a question that could be settled definitively by a crucial experiment, and there was a remarkable lack of consensus about how to explain even the most mundane phenomena of inheritance, like the resemblance of children and parents. Nevertheless, observational and experimental evidence was continually brought into the discussion, and certain phenomena became touchstones for the debate. One of these was the freshwater polyp. This tiny organism’s capacity not just to propagate by budding, but to regenerate whole organisms from pieces of itself, or to grow several heads where one had been, captured the imagination of naturalists, philosophers and their readers alike. It made an appearance in virtually every text that touched on the origin of life, the relation of soul to body, or generation. The polyp became emblematic of the mysterious capacities of organized matter, and by virtue of its stranger-than-fiction quality, inspired some people to speculate about what it might mean for a science of life. Abraham Trembley, though, who first noticed these “little organized bodies,” undertook to show that, however unusual they might appear, they followed normal patterns just like any other species. He identified their feeding habits, their normal mode of reproduction by budding and their response to being cut in all conceivable ways. 5 His focused, almost obsessive, effort was an attempt to contain the startling nature of his discovery, and to avoid possible implications for thinking about the properties of matter, and the relation between body and soul. Others were not so timid. Buffon argued, for example, that in polyps “each part contains a whole,” and he went on to argue that these organisms imply the existence of a pervasive 4 5

The standard source is Roger (1971). Trembley (1744). See also Dawson (1987), Vartanian (1950).

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multitude of organic particles from which organisms form and develop. It takes thousands or millions of them to form a single germ. In polyps, according to Buffon, organization means a simple repetition of the same “form;” more complex organisms more complex combinations of organized parts.6 Maupertuis described the polyp as a “Hydra more marvelous than that of the fable,” with its ability to regenerate a tail from a severed head and vice versa. “What are we to think of this strange kind of generation? of this principle of life spread throughout each part of the animal? Are these animals anything other than collections of embryos just ready to develop, as soon as they are allowed? Or do they reproduce by unknown means all that the mutilated parts are missing? Would Nature, which in all other animals attached pleasure to the act by which they multiply, cause these [creatures] to feel some kind of sensual delight when they are cut into pieces?” 7 For Maupertuis, this “principle of life spread throughout each part” was the organizing force driving the process of generation. In higher organisms, he speculated, parts of the body send organic elements to the reproductive organs, where they collect and combine by virtue of selective organizing forces, analogous to chemical affinities. The mixing of male and female fluids is essential, as evidenced by the resemblance of children to both parents; neither sex is privileged. At the elemental level, on this view, matter must be self-organizing; whatever the forces are, they must be intimately associated with material elements. Looking at how different people responded to strange phenomena like polyps, we see uncertainty, not only around how to interpret unexpected observations, but about whether they are even open to interpretation. Where were the limits of intelligibility? And just how far could speculation – based on the visible, but extending beyond it– safely go? Diderot, to take one example, drifted easily (and quite consciously) from the evidence of natural history to a vision of nature’s activity going well beyond empirically verifiable sights. “It seems that nature varies the same mechanism in an infinity of different ways. ... When we consider the animal kingdom, ... wouldn’t we willingly believe that there never has been more than one original animal, prototype of all animals, in which nature only lengthened, shortened, transformed, multiplied, or obliterated certain organs?”8 The prototype is more than an ideal – it’s a presumed ancestor, dating back to a moment before the proliferation of natural forms. Diderot calls his suggestion a “philosophical conjecture,” bringing the imagination to bear on solid evidence from natural history. From the indefinite past, and the malleable form of the original prototype, he ventured into the submicroscopic realm of organic molecules and proposed “muffled sensibility” as a property of all matter, the property that made all those variations in form possible. This force drives molecules to seek situations of stable equilibrium, by an “automatic restlessness [ inquiétude automate].”9 Diderot’s primary interest was not in a theory of generation as such, but a theory of matter, which was in turn tied to a theory of life. He embedded the theory in reflections on method, arguing for a combination of exploration and synthesis, experiment and interpretation. The philosopher seeks new knowledge in the same way that molecules seek their places, by a kind of 6 7 8 9

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Buffon (1954), p. 239. “Il y a dans la Nature une infinité de parties organiques actuellement existantes, vivantes, et dont la substance est la même que celle des êtres organisés.” Maupertuis (1980), pp. 107/108. Diderot ([1754] 1981), Pensée XII, pp. 36/37. Ibid., Pensée LI, pp. 84/85.

Speculation and Experiment in Enlightenment Life Sciences

restless touching and retouching, trial and error, a “tatonnement” like that of a blind man’s stick. The ideal method involves moving back and forth from sense impressions to reflection, from experiment to theory, in a kind of oscillatory exploration: “C’est le travail de l’abeille.” 10 It’s productive, useful, like making honey, but it’s also synthetic, and physical, or empirical. Any position, whether material or philosophical, is always either in flux or in a state of dynamic equilibrium, and potentially unstable. Diderot’s Pensées sur l’interpretation de la nature (1754) is a defense of “conjecture” as a productive method; the interpreter of nature starts where the senses leave off. “He draws, from the order of things, abstract and general conclusions that are for him just as evident as sensible and particular truths.”11 Diderot developed his radical defense of “philosophical conjecture” or “ esprit de divination” in dialogue with the books and experiments of Buffon and Maupertuis. Toward the beginning of his investigation of reproduction, Buffon declared that “the living and animate, instead of being a metaphysical aspect [un degré métaphysique] of creatures, is a physical property of matter.”12 Organic molecules are not yet organisms – they cannot reproduce themselves – but they are the material of life, the matter from which organisms build themselves. In collaboration with the English microscopist John Turberville Needham, who had earlier found teeming animalcules in the milt of squid, Buffon examined fluids taken from the reproductive organs of dogs and other animals. They found “moving bodies,” many with tails, in both male and female fluids, and decided that these apparently ubiquitous entities must be organic elements that combined to produce “germs” that in turn combined to generate new organisms. From the modern point of view, these experiments involved considerable confusion about what was actually seen. 13 But for Buffon and Needham, their observations of seminal fluids implied first of all that mammalian eggs did not exist, which they used as evidence against the preexistence of organized germs, and secondly that male and female functioned symmetrically in reproduction. After further experiments with infusions of seeds and meat, in which moving particles appeared spontaneously, they concluded that organic particles could be found throughout nature, and not just in reproductive organs. They claimed to have actually seen the building blocks of living organisms through their microscope. The next question was how to understand the organization of these elements into functioning, living organisms. Buffon’s organic molecules come together through the action of “penetrating forces” that guide them into “internal molds” where they take on the appropriate structure and form of body parts. “In the same way that we can make molds by which we give to the exterior of bodies whatever shape we please, let us suppose that Nature can make molds by which she gives not only the external shape, but also the internal form, would this not be a means by which reproduction could be effected?”14 This notoriously obscure formulation was ridiculed and misunderstood from its inception, though on careful consideration it seems that Buffon was selfconsciously searching for an analytic tool to give him a way of talking about “hidden mysteries” 10 11 12 13

Ibid., p. 34. Pensée LVI, p. 88. Buffon, Histoire Naturelle, vol. ii, In Buffon (1954), p. 238. On Needham, see Roger (1971), pp. 424/520; Roe (1983). For the experiments on seminal fluids and the microscope they used, see Roger (1997), pp. 140/145 and Sloane (1992). Sloane concludes that Buffon was very likely seeing bacteria and cell fragments in Brownian motion.

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without retreating into agnosticism. I want to emphasize the interpretive power of analogy for Buffon. He did not claim that physical, three-dimensional molds hover in the sex organs, waiting to receive matter exactly as metal molds do. “Nature can have these internal molds, which we will never have, just as she has the qualities of gravity, which in effect penetrate to the interior; the supposition of these molds is therefore founded on good analogies.” 15 If our senses were not limited to the surfaces of things, we might be able to conceive of internal molds more immediately. As it is, Buffon tells us, we have no reason to assume that Nature cannot employ means that are beyond our direct intuition. In fact, this may be the only way to know about natural processes, and Buffon invoked the authority of Newton to support the analogy between gravity and internal molds: I have admitted in my explanation of development and reproduction first the accepted principles of mechanics, then that of the penetrating force of gravity that we must accept, and by analogy I thought I could say that there were other penetrating forces that act on organic bodies, as experience assures us. I have proved by facts that matter tends to organize itself, and that there are an infinite number of organic particles. I have thus done nothing but generalize from observations, without having advanced anything contrary to mechanical principles.16

The internal molds are conceptually slippery because they sometimes seem to operate as forces, by analogy to gravity, and sometimes as constraining structures, by analogy to the sculptor’s molds. Buffon wrestled with how to combine a quasi-mechanical explanation of forces, shapes and motions with a notion of active organic matter that resisted this kind of explanation. He tried to escape the limitations of the human senses, while simultaneously drawing on empirical evidence. Experiment and observation were crucial components of his theory of life – but he insisted that the limitations of the human senses made it necessary to bring into play other means, if complex phenomena were to be understood. Opponents attacked him at precisely this point, unwilling to make the analogical leap; his most outspoken defender, Charles Joseph Panckoucke, remarked that we would need a sixth or even a seventh sense, in order to be able to witness or understand internal molds more immediately.17 This attitude threatened deists like Réaumur and Pluche, who explicitly pursued a descriptive approach restricted to visible surfaces, and avoided mysteries and metaphysics. Following his collaboration with Buffon, Needham continued his microscopical work. His experiments with infusions of animal and plant material, as well as observations of seminal fluids 14

15

16

17

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De la même façon que nous pouvons faire des moules par lesquels nous donnons à l’extérieure des corps telle figure qu’il nous plaît, supposons que la Nature puisse faire des moules par lesquels elle donne non seulement la figure extérieure, mais aussi la forme intérieure, ne seroit-ce pas un moyen par lequel la reproduction pourroit être opérée?” (Buffon 1954, p. 243). “Ces moules intérieures, que nous n’aurons jamais, la Nature peut les avoir, comme elle a les qualités de la pesanteur, qui en effet pénètrent à l’intérieur; la supposition de ces moules est donc fondée sur de bonnes analogies.” (Ibid., p. 244). “J’ai admis dans mon explication du développement et de la reproduction, d’abord les principes mécaniques reçus, ensuite celui de la force pénétrante de la pesanteur qu’on est obligé de recevoir, et par analogie j’ai cru pouvoir dire qu’il y avoit d’autres forces pénétrantes qui s’exercoient dans les corps organisez, comme l’expérience nous assure. J’ai prouvé par des faits que la matiere tend à s’organiser, et qu’il existe un nombre infini de parties organiques, je n’ai donc fait que généraliser des observations, sans avoir rien avancé de contraire aux principes mécaniques.” (Ibid., p. 254). Panckoucke (1761).

Speculation and Experiment in Enlightenment Life Sciences

of animals, led him to insist upon the existence of “a new Class of Beings,” creatures he saw come to life in his infusions. These “microscopical Animals ... furnish a key to the Generation of all others.” Infusions of seeds and meat gravy produced microscopic “moving Globules” after sitting for a few days. Based on many observations, he decided that all organic substances, animal and plant alike, had “vegetative Powers.”18 I won’t stop to elaborate on Needham’s interpretation of what he saw; he appears here simply to show the centrality of his surprising experiments for those of his contemporaries worried about making a science of life. Maupertuis was not convinced by Needham’s talk of epigenesis through vegetation, but the array of new observations of infusoria inspired his own conjectures. Writing to Buffon, Maupertuis commented, “I have just been reading Needham’s book. What disorder, what a reasoner! But it also contains plenty of marvels. It’s too bad that such a man wants to make systems, and that a maker of bricks wants to be an architect.”19 In the same vein, he wrote to La Condamine: “Have you read Needham’s book? What are we to think? What a new universe! What a shame that a man who observes so well reasons so poorly! After reading his book, my mind was so dizzy [étourdi] from all the ideas it presented to me that I had to go to bed like an invalid, and I have not yet completely recovered from the upheaval that this reading put me in. I hope when this tumult calms down a bit to take up again the thread of some meditations that I have begun some time since on this subject, and see if it is possible to pull out something reasonable from it.” 20 This passage captures the contemporary sense, experienced bodily by the reader in this case, of the inexplicability and astounding novelty of Needham’s observations, not to mention the dissatisfaction with his theoretical excursions. The exciting prospect of a “new world” was tempered by frustration at how to make sense of it. The ultimate question remained unsolved, according to Maupertuis, regardless of the status of Buffon’s and Needham’s microscopic bodies. “Even if material organic parts of the bodies of animals had been found, it [still] would not have fundamentally explained generation: because the formation of these organic particles would [also] need to be explained. It is too bad that those who do the experiments hardly attempt these speculations, and those who do the speculating are devoid of experiments.” 21 In his writings on generation (none of them in the form of a systematic treatise), he attempted to bridge this gap between speculation and experiment, by opening up the boundaries of both categories. Maupertuis used evidence from microscopy, animal breeding and inheritance of human traits to argue that matter must have more properties than impenetrability, extension and inertia. While gravity and chemical affinities could account for many kinds of phenomena, organization required properties “of another order than those we call physical. ... We must have recourse to some principle of intelligence, to something similar to what we call desire, aversion, memory.” 22 18 19 20 21

22

Needham (1748). Maupertuis to Buffon, 1 September 1750, Saint-Malo Municipal Archives. Maupertuis to La Condamine, 24 August 1750, Saint-Malo Municipal Archives. Quand on n’auroit trouvé des parties organiques materiaux des corps des animaux ce ne seroit pas avoir expliqué primordialement la generation: car il faudroit expliquer la formation de ces parties organiques elles memes. C’est domage que ceux qui font des experiences ne s’elevent queres à ces speculations, et que ceux qui font ces speculations soyent denués des experiences. (Ibid.). “Il faut avoir recours à quelque principe d’intelligence, à quelque chose semblable à ce que nous appellons désir, aversion, memoire.” (Maupertuis, Systême de la nature, In Maupertuis (1756), vol. ii, p. 147).

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This “principle of intelligence” resides in matter, down to its smallest parts. In effect, the elements of matter have special organic affinities analogous to psychic entities: desires, aversions, perceptions, habits and memories. As for generation, both male and female contribute material to the embryo, which forms from a mixture of the seminal fluids. Desire and aversion direct the organic elements to their places, and memory provides a link (albeit immaterial) between elements and comparable particles in the parent organism. Each one “retains a kind of memory (souvenir) of its previous situation and will resume it whenever it can, in order to form the same part in the fetus.”23 The same process produces individual variations and congenital defects. The original cause of excesses or deficiencies could be strictly accidental, but their effects may then be perpetuated through normal generation. Once a trait was established, “the particles become accustomed to their locations, which makes them place themselves similarly [in succeeding generations].”24 Once again, we see a mix of speculation and empirical evidence – a theory of selfdirecting matter required more than observation, because it could not be seen directly. Responses to Epigenesis The works of Buffon, Maupertuis, and Needham were widely read, cited, admired and challenged. A number of writers regarded these theories as dangerous, morally and theologically, as well as problematic from the anatomical point of view (especially on the question of whether male and female fluids actually mix in the uterus). Haller, for example, couched his objections to Buffon in the language of anatomical expertise. He argued, point by point, that Buffon just was not a competent anatomist, and that his theoretical conclusions were not well grounded in observation. “I have opened, without preconception or prejudice, the bodies of hundreds and hundreds of women, old and young,” he boasted, but rarely encountered the glandular bodies where Buffon located the female seminal fluid. Internal molds (being invisible) were even more problematic for Haller; he argued against them not on the basis of his fundamental incredulity about the process, but as an anatomist who knew from experience that no two individuals really resemble each other. “It is anatomy that has taught me the troublesome truth” that no two individuals have exactly the same layout of arteries, nerves, veins, muscles and bones. Therefore, since no two individuals truly resemble each other, to build a theory of heredity on resemblance is to start from false premisses. 25 This may seem a perverse argument, but it is built on Haller’s sense of himself as the consummate anatomist and also reflects the divide separating medical practitioners from more philosophical writers like Buffon.26 Réaumur also objected to giving matter the capacity to order itself through the action of attractive forces, and he likewise objected to speculating about unseen causes and forces. Even supposing that male and female fluids mix to form the germ, how can we imagine that order will emerge from the chaos of the mixture? “Who is the agent which is to disentangle and clear this 23

24 25 26

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“Mais chacun extrait de la partie semblable à celle qu’il doit former, conserve une espece de souvenir de son ancienne situation; et l’ira reprendre toutes les fois qu’il le pourra, pour former dans le foetus la même partie.” (Ibid., p. 158). “L’habitude de la situation des parties dans le premier individu les fait se replacer de la même manière.” (Ibid., pp. 160/61). Haller (1751), quotations on pp. 48, 32. See López-Beltrán (1995).

Speculation and Experiment in Enlightenment Life Sciences

chaos, to sort the several parts which are to come together, to construct organs with them, ... in short, to finish that germ...? We must not expect ... that the bare action of a gentle heat can ever be capable of producing such a work, a work infinitely more complicated than any repeating watch can possibly be.”27 To speculate, to turn the imagination loose, was tantamount to irreligion for Réaumur. Design and knowledge could not reside in matter, however fashionable attractive forces might be. “In order to arrive at the formation of so complicated a piece of work, it is not enough to have multiplied and varied the laws of attraction at pleasure; one must besides attribute the most complete stock of knowledge to that attraction.”28 To avoid this sort of trouble, Réaumur was content to refrain from theorizing altogether. Réaumur’s protégé Lelarge de Lignac, a Jesuit amateur of natural history, went further and accused the epigenesists of materialism. Buffon, he charged, relied on chance as an explanation: “animals make themselves from what he calls living elements, equally appropriate for entering into the formation of animals and vegetables.”29 Lignac was incensed by every aspect of Buffon’s book, from its content to its style, pointing to its subversion of morals along with what he considered its absurd use of attractive forces. He denounced the excesses of Buffon’s style (“worthy of a modern novel”) used to reinforce heterodox accounts of the origin of human life and the order of the solar system. These critiques share a squeamishness about active matter, chance, sexual license, modern style: all compounded together. Natural history is safe; argument from design is useful; epigenesis is materialist, and therefore scary. Anxieties about self-directing matter were not limited to priests like Lignac; Voltaire also found pre-existence (in the sperm) the only possible explanation for generation, even though he recognized that the implied infinite regress is counter-intuitive and must remain a “mystery.” He preferred the mystery to the threatening implications of active matter. Voltaire stubbornly misunderstood Maupertuis’s and Buffon’s analogical use of forces, in order to more easily caricature epigenesis. Maupertuis, he complained, “has supposed that, in the fertile principles of man and woman mixed together, the left leg of the fetus unerringly attracts the right leg; that an eye attracts an eye leaving the nose between them, that a lung is attracted by the other lobe, etc.” 30 Voltaire here found himself on the same side of the fence as Lignac, the Cartesian Jesuit, since they shared a conviction that God alone could understand the first principles behind phenomena. What was left of divine design if matter could organize itself? Voltaire and Lignac agreed that epigenesis implied a disturbing lack of orderly causality; “a man could be born from a clod of earth just like an eel from a bit of flour paste. Besides, this ridiculous system would lead obviously to atheism. ... Needham’s microscope appears to be the laboratory of atheists.” 31 27 28 29

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Réaumur ([1749] 1751), p. 457. Ibid., p. 463. “Dans son ouvrage tout s’opere fortuitement; les animaux meme se composent d’élémens qu’il appelle vivans et egalement propre à entrer dans la construction des animaux et des végétaux. Il est vrai qu’il met l’efficace de l’attraction à la place du hazard d’Epicure...” (Lelarge de Lignac 1751). “Il [l’auteur] a pretendu que, dans les principes feconds de l’homme et de la femme meles ensemble, la jambe gauche du foetus attire la jambe droite sans se méprendre; qu’un oeil attire un oeil en laissant le nez entre deux, qu’un poumon est attiré par l’autre lobe, etc.” (Voltaire 1752, p. 446). Si des animaux naissaient sans germe, il n’y aurait plus de cause de la generation: un homme pourrait naitre d’une motte de terre tout aussi bien qu’une anguille d’un morceau de pate. Ce systeme ridicule menerait d’ailleurs visiblement a l’atheisme. ... Le microscope de Needham passa pour être le laboratoire des athées.” Voltaire pamphlet against Needham, 1765; cited Mazzolini and Roe (1986), p. 82.

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Animal Breeding I now turn to another kind of experiment that contributed to the debates about life and how to investigate the origin of organization, namely breeding experiments. Domesticated animals had been bred for selected characteristics for centuries, by farmers, by pet fanciers, and by dealers in animals for the luxury market. These breeders didn’t worry about what caused certain traits to be inherited with a certain frequency. But in the 18th century hybrids and domestic animals entered the discourse about generation alongside Needham’s microscopic animals, and the inheritance of visible traits became evidence for the mixing of contributions from male and female parents. In breeding practices, and in thought experiments about hypothetical crosses, we can see how the specialist discourse of life science (coming out of anatomy and microscopy) intersected with polite culture, specifically in the cultivation of pets and other domestic or farm animals, like horses and fowl, and also in literature written for this same genteel readership. Maupertuis himself kept quite a menagerie of pets (as did Buffon and Réaumur), some of which he bred systematically, looking for hereditary patterns. When he came across a rare combination of colors in the coat of a female dog, for example, he attempted to perpetuate the trait; after four litters a male puppy with identical markings was born. 32 This dog eventually went on to father yet another with the distinctive coloring, showing that the trait was not sex linked. Another example came from a human family in Berlin, some of whom were born with extra fingers: he recorded the genealogy of this family for three generations with names of individuals, their spouses, and their children, and the occurrence of extra digits among them. The trait occurred frequently enough to show that it was transmitted bilaterally, and that marriage to a fivefingered spouse affected the frequency of its appearance in the offspring. These examples seem prosaic, and not particularly startling; but they opened up questions about the meaning of individual variations for a science of life – especially the analytic value of examining populations over time – and about the relation of individuals to their species (or races). To show that the sixth finger was inherited rather than accidental, Maupertuis estimated its probability of appearing at random in consecutive generations of one family. The probability decreases by the same factor for each generation, to the point where the chances of three generations of the same family producing such individuals at random would be impossibly high; “numbers so large that the certainty of the best demonstrated things in Physics does not approach these probabilities.” 33 Breeding also raised questions about agency: could manipulation by human breeders produce new species? How malleable were living forms? And how could human efforts to breed new animals illuminate natural processes? Maupertuis understood the perpetuation of naturally occurring variations, like albinism, by analogy to selective breeding practices: “Whether one takes this whiteness [of the albino’s skin] for an illness or accident, it will never be anything but a 32

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Formey recalled making his way fearfully through Maupertuis’ s menagerie when he went to visit. “Il étoit dangereux quelquefois de passer a travers le plupart de ces animaux, par lesquels on étoit attaqués Je craignois surtout beaucoup les chiens Islandois.” These were the dogs used in Maupertuis’s breeding experiments. (Formey 1789, vol. i, pp. 218/219). Many letters attest to Maupertuis’s trading in dogs with his correspondents. The dog-breeding experiments are described in Maupertuis, “Sur la génération des animaux,” Lettres de M. de Maupertuis, in Maupertuis (1756) vol. ii, pp. 310/311. See also Hoffheimer (1982). Ibid., p. 310.

Speculation and Experiment in Enlightenment Life Sciences

hereditary variation that reinforces itself or erases itself over the course of generations.” 34 Similarly for polydactyly: “By such repeated marriages, it would probably die out, and it would perpetuate itself through marriages where the trait was common to both sexes.” 35 Varieties occurring by chance could be perpetuated through cultivation, just as particular traits are preserved in animals by systematically selecting for them. Over time, cultivation could cause varieties to solidify into a separate species that would be stable enough to perpetuate itself. In his most speculative mode, Maupertuis suggested that naturalists might use the collections of animals in menageries to explore heredity through hybridization, even to the point of crossing animals that would never mate in nature. “The efforts of a hardworking and enlightened naturalist would result in plenty of curiosities of this type, by causing animals of different species to lose their natural repugnance for each other, through education, habit and need.” And for animals that couldn’t be pressured to mate, he imagined the enlightened naturalist developing techniques for artificial insemination to create new “marvels”: “We might see from these unions plenty of monsters, of new animals, perhaps even entire species that nature has not yet produced.” 36 These suggestions were meant to push beyond the limits of current knowledge into the unknown, the possible, the marvelous, not just by looking at nature, but by manipulating it. Maupertuis implied that much of what we don’t know is only mysterious because it hasn’t yet been carefully (or imaginatively) investigated. Reflecting on the regenerative capacity of polyps and lizards (who regrow their tails), he asked, “Is it probable that this marvelous property belongs only to the small number of animals we know about? ... perhaps it only depends on the method for separating the parts of other animals to see them reproduce themselves [in the same way].” 37 Maupertuis’s projects along these lines remained in the realm of the hypothetical, of course. I would point out, though, that interspecies crossing carried a definite flavor of the fashionably transgressive and risqué. An extreme example, that undoubtedly draws on contemporary discussions about breeding, is found in Diderot’s D’Alembert’s Dream, where the characters imagine the production of a hybrid race of “goat-footed men” to work as servants: they would be “vigoureuse, intelligente, infatigable et véloce.”38 Mlle de l’Espinasse only realizes belatedly that such a breed would require people to mate with goats; this causes her to withdraw her approval for the scheme. This fantasy, intended to shock as well as to amuse, is an extreme example of the resonance of epigenetic discourse with social concerns, cast in the enlightened medical framework articulated by Diderot’s version of the vitalist doctor Bordeu. Even Réaumur, a denizen of salon culture though not a freethinker by any means, wrote about the “philosophical amusements of the barnyard,” giving detailed voyeuristic accounts of the amorous adventures of a duck and a rooster and, more engaging still, of a rabbit and a hen. These animals, which Réaumur had kept under observation in his house for a time, had entertained “the whole of Paris,” and prompted, naturally enough, speculation about the outcome of the interspecies union: “It was the general wish, as well as my own, that it might have procured us chickens covered with hair, or rabbits clothed with feathers.”39 34 35 36 37 38

Maupertuis (1980), p. 138. Maupertuis (1756), vol. ii, p. 308. Maupertuis, Lettre sur le progrès des sciences (Berlin 1752), in Maupertuis (1756), vol. ii, pp. 420/421. Ibid., pp. 421-422. Diderot, Le Rêve de d’Alembert, in Diderot (1964), p. 383.

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A furry chicken or a feathered rabbit would have called into question Réaumur’s commitment to preformed germs, so it’s perhaps just as well that the hen’s eggs never hatched. Réaumur did note in his work on poultry that chickens could provide clues to understanding inheritance, through selective breeding for readily observable variations like extra claws. In fact, he described breeding programs to enlist the efforts of “those who love to find in their poultry-yards amusements conducive to the progress of natural knowledge.”40 He claimed to have done such systematic crosses with his own chickens, but coyly refused to disclose the results. His readers continued to send him material, some of which made it into the second edition of the book. Réaumur’s impulse to collect stories and specimens of hybrids, as well as to proceed with his own breeding experiments, brought him up against recalcitrant evidence for bilateral inheritance. He managed to avoid the troubling conclusion, however, by focussing on the “curiosities” themselves – including wingless chickens, a North African beast of burden resulting from crossing an ass with a cow, and even six-fingered people. This last example, which corroborated Maupertuis’s account of the Berlin family, was provided by an informant in Malta with a genealogy over several generations. Réaumur quoted the report in full with no comment other than “it does not seem favorable to the preexistence of germs.”41 Buffon also devoted substantial effort to breeding across species (wolves and dogs, for example), not so much to show how traits were passed from generation to generation, but to map out relations among species. His programs for breeding agricultural animals (within species) were intended to counter what he saw as the natural degeneration of animal forms if left to themselves. “In order to have beautiful horses, good dogs, etc., it is necessary to give foreign males to the native females, and reciprocally to the native males, foreign females; failing that, animals will degenerate ... In mixing the races, and above all in renewing them constantly with foreign races, the form seems to perfect itself, and Nature seems to revive herself.”42 Nature is flexible, with the potential to be molded by environmental factors, including human intervention. Buffon argued that people can use the forces of nature to shape animals to their purposes. Breeding for specific traits (as in pet dogs, for example) required constant attention, however, to avoid reversion to the original forms. The Paris physician Charles Augustin Vandermonde incorporated Buffon’s theory of organic molecules into a system for improving the human species through application of principles of nutrition, education, hygiene – and breeding. Vandermonde played on contemporary concerns about declining population and degeneration by prescribing healthy practices for everything from the choice of mate to engendering vigorous offspring to the care of pregnant women and the feeding and education of children. Since, following Buffon, both male and female contribute equally to the offspring, they must be well matched to complement each other. Just as animal stock is strengthened by crossing with natives of other regions, he says, humans benefit from mating with people from different climates, who will necessarily have different constitutions. When people move from the provinces to the city and intermarry with urban dwellers, the result is 39 40 41 42

38

Réaumur ([1749] 1751), p. 457. Ibid., p. 467. This discussion about crosses in chickens was added to the second edition of Réaumur’s book. (Réaumur, Art de faire éclorre ..., 2nd ed. (Paris 1751), p. 376.) Quoted from Spary (2000), pp. 113/114.

Speculation and Experiment in Enlightenment Life Sciences

improvement in strength and health of the city stock. Further, he implies that human agency could easily be put to work improving its own species, by analogy to animal breeders who “create new races of dogs, cats, horses.” “We can easily see that one could perfect animals by varying them in different ways. Why should we not work also on the human species? It would be just as possible, ... in combining all our rules, to embellish men, as it is routine for an able sculptor to cause a model of beautiful nature to emerge from a block of marble.”43 Here the breeder appears as an artist, molding active matter to do his bidding. Vandermonde’s book reads as a handbook for healthy living; he assumes his readers will take an interst in shaping their own progeny. While he does not indulge in fantasies about cross-species mating, his rules for choosing suitable partners from other climates dictate a kind of subtle hybridization, or at least mixing. The program rests on a mechanics of heredity inspired by Buffon’s organic molecules, whereby specific traits materialize in oscillating spiral-shaped particles that wind together selectively to make up the germ of the offspring. These operations at the submicroscopic level, well beyond the reach of direct observation, locate macroscopic properties firmly in the stuff of organic matter with its resilience, activity, vibrations and self-propulsion. Vandermonde adapted Buffon’s version of active matter to his own more medical vision of prescriptions for human betterment where heredity is key. In the realm of satire, boundaries can be tested with even greater abandon. An anonymous pamphlet published in Paris in 1772 tells the “true and marvelous story of a “lynx-girl” who could see into solid objects. “I take the thing from an Englishman, and those gentlemen do not pass as very gullible, nor are they prone to immediately crying ‘Miracle!’” The author discusses the odd trait of the lynx-girl by analogy to hydroscopes, people with the ability to find water underground by seeing through the earth. He presumed that the trait was heritable, and therefore could be perpetuated by breeding. Inspired by Maupertuis’s speculations about cross-breeding, he goes on to suggest that the Royal Society and the Paris Academy of Sciences really ought to preside over (and pay the expenses of) the marriage and breeding of this girl, in order to produce more such gifted lynx-people. “There is no need to mention what advantages would result from a lynx race, for the good of humanity; what light [lumières], what vision, what insight, these living telescopes, born in the sanctuary and under the auspices of physics, could communicate to savants, the authors and the cause of their existence!”44 He went on to calculate how long it would take for the trait to multiply in subsequent generations, and how useful these people, bred in academies would be for police work, for uncovering court intrigues, and so on. Conclusion I have tried to lay out the landscape of discourse about heredity and generation to highlight resonances with other claims – about active matter, about the origin of organization, about progress and change, and also about method. We are not surprised to see that experiments played an essential role, often as provocation for conjectural interpretation and rarely as conclusive or indisputable. While Haller, who has only come into my story as a critic of Buffon, implied that everything could be sorted out by multiplying careful and professional observations of embryos, a number of French writers at mid-century addressed the question of how to accommodate 43 44

Vandermonde (1756), p. 155. Histoire véritable et merveilleuse d’une jeune angloise... (1772) p. 58.

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method to mystery, and to the limits of human senses. If we are to understand their attempts to lay the foundations of a science of life, we should consider the role of hypothetical experiments, as well as conjectures and queries, and how all these theoretical modes interact with actual experiments and observation. When Réaumur, for example, imagines crosses designed to determine the location of the germ, but then refrains from reporting his results, does this count as experiment? How is this different from Diderot’s goat-men or Maupertuis’s hypothetical hybrids between exotic African jungle animals? Or Vandermonde’s perfected humans? If we examine these speculative experiments alongside the breeding programs that traced abnormalities through generations of dogs or birds, we can read them as part of a program to get below the surface to general laws of nature and of life. Maupertuis articulated this desideratum in his critique of natural history (and here he had Réaumur in mind as the contemporary exemplar): “All our treatises on animals, even the most methodical, form nothing but paintings pleasing to the eye. To make natural history into a true science [véritable science] we would have to apply ourselves to researches that would allow us to know, not the particular figure of such and such an animal, but the general processes of nature in her production and preservation.” 45 This comment is partly about designing experiments to ask the right questions, but it also points to a willingness to interpret them. Even d’Alembert addressed the question of the role of conjecture in his discussion of experiment in the Encyclopédie, where he says, “When I proscribe from physics the mania for explanations, I am very far from proscribing that spirit of conjecture, which at once timid and enlightened sometimes leads to discoveries: ... that spirit of analogy, whose wise strength pierces beyond that which nature seems to want to show, and foresees facts before having seen them.” 46 For many thinkers at mid-century seeking access to what Buffon called “the hidden means that nature might be employing for the generation of creatures,” the complementary use of conjecture and experiment was the only viable method.

Department of History, University of California, Los Angeles

45

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“Tous ces traités des animaux que nous avons, les plus méthodiques meme, ne forment que des Tableaux agréables à la vue: pour faire de l’histoire naturelle une véritable science, il faudroit qu’on s’appliquat à des recherches qui nous fissent connoitre, non la figure particulière de tel ou tel animal, mais les procédés généraux de la nature dans sa production & sa conservation.” Maupertuis, Lettre sur le progrès des sciences, p. 418. “...quand je proscris de la Physique la manie des explications, je suis bien éloigné d’en proscrire cet esprit de conjecture, qui tout-a la-fois timide & éclairé conduit quelquefois à des découvertes, pourvu qu’il se donne pour ce qu’il est, jusqu’à ce qu’il soit arrivé à la découverte réelle: cet esprit d’analogie, dont la sage hardiesse perce au dela de ce que la nature semble vouloir montrer, & prévoit les faits, avant que de les avoir vus.” D’Alembert, “Experimental,” in Diderot and Alembert (1751-1765).

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Reference List Aumont, Arnulfe d’. 1757. “Génération (physiologie)”. In Encyclopédie, edited by Diderot and d’Alembert. vol. vii, pp. 559–574. Paris: Briasson. Buffon, Georges-Louis Leclerc de. 1954. Oeuvres philosophiques, edited by Jean Piveteau. Paris: Presses Universitaires de France. Dawson, Virginia P. 1987. Nature’s enigma: the problem of the polyp in the letters of Bonnet, Trembley, and Reaumur. Philadelphia: American Philosophical Society. Diderot, Denis. 1751. “Animal,” In Encyclopédie, edited by Diderot and d’Alambert. vol. i, pp. 468–474. Paris: Briasson. ———. 1964. Oeuvres philosophiques, edited by Paul Vernière. Paris: Garnier. ———. [1754] 1981. Pensées sur l’interprétation de la nature. In Diderot. Oeuvres complètes, vol. ix, edited by Jean Varloot. Paris: Hermann. Diderot, Denis and Jean Le Rond d’Alembert. 1751–1765. Encyclopédie; ou Dictionnaire raisonné des sciences, des arts et des metiers. 17 vols. Paris: Briasson. Formey, Jean-Henri-Samuel. 1789. Souvenirs d’un citoyen, 2 vols. Berlin: Lagarde. Haller, Albrecht von. 1751. Réflexions sur le systême de la génération de M. de Buffon. Geneve: Chez Barrillot. Hoffheimer, Michael H. 1982. “Maupertuis and the Eighteenth-Century Critique of Preexistence.” Journal of the History of Biology 15(1):119–144. Lelarge de Lignac, Joseph Adrien. 1751. Lettres à un Amériquain sur l’Histoire naturelle, générale et particuliere de Monsieur de Buffon. 2 vols. Hambourg. López-Beltrán, Carlos. 1994. “Forging Heredity: From Metaphor to Cause, a Reification Story.” Studies in the History and Philosophy of Science 25:211–235. ———. 1995. “‘Les maladies héréditaires’: Eighteenth-century disputes in France.” Revue d’histoire des sciences 48:307–350. Maupertuis, P.-L. M. de. 1756. Oeuvres. 4 vols. Lyons: Bruyset. ———. 1980. Vénus physique. edited by Patrick Tort. Paris: Aubier Montaigne. Mazzolini, Renato and Shirley Roe. 1986. Science against the unbelievers: the correspondence of Bonnet and Needham, 1760-1780. Oxford: The Voltaire Foundation. Needham, John Turberville. 1748. “A Summary of some late Observations upon the Generation, Composition, and Decomposition of Animal and Vegetable Substances.” Philosophical Transactions 45:615–666. Panckoucke, Charles Joseph. 1761. De l’homme, et de la reproduction des différens individus. Ouvrage qui peut servir d’introduction & de défense à l’Histoire naturelle des animaux par m. de Buffon. Paris. Réaumur, René-Antoine Ferchault de. [1749] 1751. On the Hatching and Breeding of Domestick Fowls. London. translation of L’art de faire éclorre ... des oiseaux domestiques. Roe, Shirley. 1983. “John Turberville Needham and the generation of living organisms.” Isis 74:159–184. Roger, Jacques. 1971. Les sciences de la vie dans la pensée française du XVIIIième siècle. 2nd ed. Paris: Armand Colin. ———. 1997. Buffon: A life in natural history. Sarah Bonnefoi, trans. Ithaca, N.Y.: Cornell University Press. Sloane, Philip. 1992. “Organic molecules revisited.” Buffon 88: actes du Colloque international pour le bicentenaire de la mort de Buffon. Paris & Lyon: J. Vrin. Spary, E. C. 1999. “Codes of passion: Natural history specimens as a polite language in late eighteenthcentury France.” In Wissenschaft als kulturelle Praxis, 1750–1900, edited by H. E. Bödeker, P. H. Reill and J. Schlumbohm. Göttingen: Vanderhoek and Ruprecht. pp. 105–135. ———. 2000. Utopia’s Garden: French Natural History from Old Regime to Revolution. Chicago: University of Chicago Press. Trembley, Abraham. 1744. Mémoires pour servir a l’histoire d’un genre de polypes d’eau douce. Leiden. Vandermonde, Charles-Augustin. 1756. Essai sur la manière de perfectionner l’espèce humain. 2 vols. Paris: Vincent. Vartanian, Aram. 1950. “Trembley’s Polyp, La Mettrie, and Eighteenth-Century French Materialism.” Journal of the History of Ideas 11:259–286. Voltaire, François Marie Arouet de. [1752] 1877–1885. “Les Oeuvres de M. de Maupertuis,” Bibliothèque raisonnée 49: 158–172, In Voltaire Oeuvres, vol. 39. 52 vols., edited by Beuchot. Pari:, Garnier freres.

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Duchesne’s Strawberries: Between Grower’s Practices and Academic Knowledge Marc J. Ratcliff

INTRODUCTION This paper studies the history of eighteenth century hybridization and genealogy while looking at the interaction of growers’ practices and theoretical botany. I discuss a case-study based on the discovery of a new vegetable species during the second part of the eighteenth-century. Antoine Duchesne, a Paris gardener and botanist, discovered an undescribed form of strawberry in 1766. While a certain work was done by James L. Larson on Koelreuter and Linnaeus’ experiments on hybridization, the secondary literature did not pay a lot of attention to Duchesne although the story displays a much revealing example of the strategy a scholar elaborated to deal with new species. Indeed, the interest of this discovery lies at least in the fact that Duchesne attempted to understand this new production of “nature and art”, the new sort of strawberry, thanks to a model labeled genealogical tree. Thus through Duchesne’s work, hybridization crosses genealogy. The main epistemological and historical points are to understand what does genealogy mean, and to what extent is this a sign of a modern issue of genealogical tree as defined later by Darwin, Haeckel and de Vries and, eventually, what are the influences that shaped this story. After a brief description of the discovery reported by Duchesne, I focus on what did historiography with this story. Nineteenth-century biologists and twentieth-century historians made various uses of Duchesne’s strawberries. When using Duchesne’s species, nineteenthcentury evolutionists show outspoken ideas yet different from that of the historians of biology and classification. The historians of classification and evolutionism have slightly discussed Duchesne, who was neglected by the tradition of the history of the biological thought as outlined by Jacques Roger, Elizabeth Gasking or John Farley. Yet, the recent comments on Duchesne placed contradictory interpretations on the meaning of the genealogy. My paper aims partly at bringing more data on what does “genealogy” and genealogical classification mean for a breeder and botanist in the second part of the eighteenth-century. I examine the several factors that determine this story in order to explain where does such a conception of genealogy come from. Putting Duchesne’s ideas in context thus means to investigate on his working milieu, the continent of practitioners: breeder, gardeners, growers, florists, a world close to that of cattle farmers. Two problems may need investigation. First, what are the current and shared practices of hybridization in the agricultural and “domestic economy” milieu which could have influenced Duchesne’s experiments and conception? Second, what are the differences – of object, practices and conceptions – between this milieu and the botanical academic milieu which discusses similar questions. One shall examine the methods of hybridization used by growers and florists, before attempting to define the borders between botanists and breeder-growers-florists. This shall bring us to more precise definitions of the terms species and varieties according to the “professional” contexts using these concepts. Indeed, it seems that the definition varied according to the use each community made of it, though certain concepts were shared by botanists and practitioners.

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Another problem relates to fixity, history and genealogy. First, in relation to species and classifications, the status of time was at stake in the debates on varieties and species. A fixed species is conceived as being out of time, while an organism suddenly appearing raises the question of time and of a constancy of species. This issue was much discussed by the classical botanists such as John Ray, Robert Morison, Jean Marchant and Linnaeus, and Duchesne did not evade it. Second, time is also related to history, and the use Linnaeus, and Duchesne made of the history of botanical species was new in contrast to the descriptive tradition of botany. Including time in the history of botany shows a new way of reading history, which relates to the changes from a narrative to a temporal history. Third the use of genealogy in Duchesne seems to have been influenced by human genealogy and I shall use context clues consistent with this interpretation. All this should enhance our view of the origin and function of the genealogical tree and classification in Duchesne, while putting greater emphasis on a synthetic moment that made the field of practitioners and botanists crossing each other. 1. CONSTRUCTING THE DISCOVERY In 1763, a young Paris grower and botanist, Antoine-Nicolas Duchesne (1747-1827) discovers an unknown kind of strawberries in his Versailles garden. Unknown with regard to its morphology, this plant had not been described by previous botanists. The main morphological characteristic of the plant was its monophylla leaves, with only one lobe, while leaves of common strawberries contain three lobes (fig. 1).

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Duchesne’s Strawberries: Between Grower’s Practices and Academic Knowledge

Fig. 1 Fragaria Monophylla by S. Edwards. From The Botanical Magazine 2 (1788).

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At the age of seventeen, in July 1764, Duchesne who had published a small handbook of botany, “personally presented King Louis XV with a pot of strawberries.” 1 Thanks to a particular hybridization, the result was spectacular and the King decided to patronise him. “He authorized Duchesne to raise more F. chiloensis in the royal kitchen garden at Versailles and to collect all varieties of strawberries known in Europe for the Trianon garden.” 2 Such a high patron opened him many doors. Tutored by Bernard de Jussieu, who had helped him from the beginning, he wrote to Haller, to Linnaeus, to other French and European naturalists to ask for seeds and specimens. After he made the plant reproducing from budding, Duchesne collected the seeds obtained which he sowed in 1764. Two weeks after, the first strawberries appeared and he was much astonished to find that they had reproduced from the seeds. This was indeed one of the features of a stable species, constant in time, and the plant was yet undescribed. Duchesne dispatched the seeds obtained and asked the scholars to sow them in order to repeat the experiment. Sowed in other places the seeds stubbornly reproduced the same plant. On this series of fact, Duchesne launched a two years research that led him to publish his 1766 Histoire naturelle des Fraisiers. Still non extant for the botanist community, the plant had to be at least baptised. It was named, the year after Duchesne’s book, Fragaria monophylla, the latter term being not used by Duchesne. Linnaeus was, at this time, at the apex of his career, for he had been recently elected a foreign member of the Paris Académie des Sciences.3 In Philosophia botanica, Linnaeus had prescribed strict rules that enabled only “orthodox Botanists” to give the “right name” to a plant. 4 This secured him and a few other privileged botanists against amateurs, while himself and other emergent professionals shared, among other, the power of naming plants. In the 1767 twelfth edition of Systema Naturae,5 Linnaeus thus named the new plant F. monophylla, a name later currently used, for instance by Abbé Rozier in his entry “strawberry” of his 1782 Cours Complet d’agriculture. The only plate in Duchesne’s book illustrates a “genealogy of strawberries” (fig. 2) in which the author classifies ten kinds of strawberries thanks to a genealogical key. 6

1 2 3 4 5 6

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Darrow (1966), ch. 5, section “Duchesne and his work.” (I quote Darrow from an Internet version without pagination). Darrow (1966), ch. 5, section “Duchesne and his work.” Larson (1994), p. 17. Linnaeus (1751), §211. Orthodox Botanists were those “who established their method on a real base, such as fructification” (Ibid, § 26), i.e. a few botanists among whom was Linnaeus. Linnaeus, Systema Naturae (1767) II, 349. Duchesne (1766a), p. 228, pl. I.

Duchesne’s Strawberries: Between Grower’s Practices and Academic Knowledge

Fig. 2 Genealogy of the strawberries according to Duchesne. 2. DUCHESNE’S STRAWBERRIES IN THE HISTORIOGRAPHY OF SCIENCE On the basis of the genealogical plate, scientists and historians have regarded Duchesne as an early representative of mutationism and evolutionism, while other historians have rejected the presence of a concept of genealogy in the book. The relationship between the discovery of new species in the eighteenth-century and the framework of heredity and generation was poorly discussed by historians. Jacques Roger, Peter Bowler, John Farley, Walter Bernardi, Roselyne Rey who have studied the interplay between generation, species, and sometimes, heredity, have mainly looked for a theoretical debate of ideas

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that included Maupertuis, Diderot, Buffon and Bonnet’s speculations. 7 Notably much work has been dedicated to the debates on teratological cases that illustrate both imaginary and detailed conceptions of heredity, for instance in the works of Jean-Louis Fischer and of Javier Moscoso. But, the theory-oriented approach was much interested in animal and human field, and it has notably set apart the influence practitioners had on the debate over heredity and new species. On the other hand, the historians of eighteenth-century botany and natural history have focused more on Linnaeus and Jussieu’s works. Yet the question of new species has been mainly discussed through the example of plant hybridization for Linnaeus on the Peloria, Adanson’s conceptions, Koelreuter’s hybridization, and Lamarck’s view.8 One has to look to another field – the history of classification, evolutionism, and breeding – to find a place for Duchesne’s strawberries in the history and historiography of science. Historians of breeding have no problems in declaring that Duchesne’s work represents the starting point of the modern strawberry.9 Certain scientists have discussed the relevance of the species itself Fragaria monophylla to the issue of mutation and new species, while historians have emphasised Duchesne’s conception of species, fixism or mutationism, grounded on his genealogical classification. Nineteenth-century scientists in particular have constructed a historical narrative that includes Duchesne’s study into a whiggish history. Discovering Fragaria and the why and how of new species was less examined than the so-called anticipation of modern evolutionism based on the genealogical tree. Alphonse de Candolle (1806-1893), a Swiss botanist who wrote one of the first sociological studies of science, is among the first to quote Duchesne as a forerunner of Darwin’s genealogical definition of species.10 On the other hand, Hugo de Vries (1848-1935), who defended a mutationist version of evolutionism in his 1900 work on the mutation of species and varieties, had included Fragaria monophylla among various examples of mutations remarked by previous botanists. He noticed that a similar example of F. monophylla was represented in a painting by Holbein (1495-1573). De Vries’ was an argument showing that the same mutation had occurred in various places and times.11 He used it as a rationale useful to his scientific programme, assigning the value of a proof to a historical example. This use seems to be different from that of Candolle who regarded Duchesne’s model as an anticipation of evolution, for De Vries did not comment on Duchesne’s genealogical tree. Still, not much time later, his French translator, L. Blaringhem, published a book on mutationism in which he developed the right same example. However he did not content himself to discuss the presence of F. monophylla as a mutation vouched for historically by several authors, but considered Duchesne to have been a supporter of mutable species. To Blaringhem, Duchesne’s questions “show that he had, at least since 1766, the very precise notion of the sudden transformation of forms (...) he refused to believe in the fixity of species”.12 This was closer to Candolle than to De Vries’s thought.

7

8 9 10 11 12

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Roger (1963), Bowler (1973), Farley (1977), Bernardi (1986) and Rey (1989) have usually not much discussed botanical works, and have almost neglected topical authors for the question of the new species such as Marchant, Linnaeus, Adanson, Gleditsch, Koelreuter and Duchesne. Stafleu (1971), pp. 261/62, 305/06; on Koelreuter, see Larson (1994). Darrow (1966), ch. 6, section “Breeding in France 1770 to 1900.” De Candolle (1881). De Vries (1909), p. 378. Blaringhem (1911), p. 8.

Duchesne’s Strawberries: Between Grower’s Practices and Academic Knowledge

Later this interpretation of Duchesne became topical in Emile Guyénot’s 1941 standard French handbook of history of biology. Guyénot, himself a biologist, followed Blaringhem’s reconstruction, but was disturbed by the use Duchesne had made of the term race, and added in a footnote: “the author uses frequently the term race with the meaning of species, and the term species with that of genus”.13 And Guyénot quoted Duchesne, who wrote “thus there are new [species]”.14 Eventually, through recent literature other options brought up concerning not Duchesne’s conception of species, but his ideas on genealogy. Pascal Tassy notably devoted a book to the issue of the historical construction and to the function of the genealogical tree in modern biology. He considered Duchesne was the first to make use of a temporal genealogy for a classification of living organisms.15 On the other hand, Giulio Barsanti’s study of the images and representations of classification up until the time of Darwin highlights another conception of Duchesne’s genealogical tree. Barsanti regarded Duchesne’s use of a genealogical tree as a misconception. For Barsanti, Duchesne’s is not a genealogical tree, it has no bough, no bifurcation and intermediary species, and is furthermore turned over.16 According to Barsanti, Duchesne’s use of a genealogy stems from misunderstanding this notion, treated in a obscure and “misterioso” way.17 It appears thus that secondary literature offers several opinions on the particular meaning of the “new species” for Duchesne. In front of so opposed ideas from historians of cladism and of classification, one shall look at the sources. If the secondary literature is not outdated, there is nevertheless no study on Duchesne that discusses its apparition in the second half of the 18th century – what means the publication of a whole book on a new strawberries in 1766? More particularly, I could not find studies on the origins, contexts, social network, sources of inspiration and continuation of this enterprise. Such a work is a preliminary historical investigation before explaining Duchesne’s conceptions, and their influence on the realm he addressed. In order to understand Duchesne’s enterprise, I shall follow several lines and hypothesis: a. Duchesne’s work stems from a hybridization of two particular “traditions”: the grower and breeder’s practices, and the academic botany. I thus examine both traditions, show their relationships, in order to understand what were their respective contributions to Duchesne’s enterprise. b. Once identified the origin of certain practices and ideas, it will be more easy to analyse and display Duchesne’s conception of genealogy, species, race, heredity, etc., in order to characterise his own contribution. c. I shall address the problem of the impact this enterprise had on the second half of the eighteenth-century practices and knowledge of natural history, notably on two points: what became the new strawberries, and what was Duchesne’s impact on heredity and new species.

13 14 15 16 17

Guyénot (1941), p. 376n. Guyénot (1941), p. 376; Duchesne (1766), p. 13. Tassy (1991), p. 26. “avec la publication d’une véritable généalogie d’êtres vivants qui se veut comme telle, à la manière des fraisiers de Duchesne, apparaît la notion du temps (...).” Barsanti (1992), p. 85. Barsanti (1992), p. 84. Ibid., pp. 84/85.

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3. BOTANY AND PRACTITIONERS IN THE MID EIGHTEENTH-CENTURY 3.1 Grower and breeder’s practices: morphological changes of plants G. M. Darrow has given a comprehensive account of the history and breeding of strawberries in England. Among the wide variety of growers and breeders’ practices, I shall characterise those affecting morphological changes of plant during the years of Duchesne’s enterprise. These practices established a common background for growers, florists, breeders – and cattle farmers – from the seventeenth-century onwards; they had been progressively improved along several treatises on husbandry, gardening and agriculture. At the time of Duchesne, the leading publication providing information on empirical practices in France was Journal économique. A monthly journal created in 1751 by a group of people favoring practical economics and agricultural topic, it was partly controlled by the Académie des Sciences through the well-disposed censorship of Jean-Etienne Guettard.18 The Journal économique excluded non-applied scientific research, and served as a link between applied knowledge and civil society, for subjects such as natural history, domestic economy, law, public health, navy, geography, technology, and, of course gardening, growing and breeding. The growers of the mid eighteenth-century distinguished several methods that caused morphological changes in a plant or an animal. One of the concrete problems was, for the florists, to transform a specimen with simple flowers into a specimen with double flowers, or in other words, to double or even triple the number of petals. The simple flowered and non cultivated specimens had been described by botanists with the minimum number of petals. However, sustained by adequate cultivation, they could be converted into double flowered specimens, with a great increase in value for the rare ones. This was not just a joke invented by a few gardeners. In the seventeenth-century and early eighteenth-century, the price of certain rare tulips was so high that the Dutch government published a law regulating the market, to avoid certain people losing everything for a couple of flowers... In the mid eighteenth-century the German and Dutch florists and growers distributed catalogues of their best specimens of carnations and other flowers all over Europe. Several methods were used by these practitioners to transform a flower from simple to double. Soil transplantation was one of these methods, and especially frequent transplantation from a soil to another without blooming was regarded as being much successful. Such method echoed that of the cattle farmers. When a sheep flock was seen as “degenerating”, being too meagre and small – “like rabbits”! – farmers imported a more than three-years old strong ram from another region to cover the sheep. An experiment was said to have been carried out by a farmer in Limousin who used the latter method, and, in about three years, “multiplied by three the income of the wool”. 19 While the growers and florist had in mind to transform the shape of flowers, the cattle farmers experimented on the selection of domestic animals with the aim of “improving the species.” 20 18

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In the history of the relationship between Paris Académie des Sciences and French societies directed to utilitarian practices, the first Société des Arts created in the 1720s in Paris failed apparently because of rivalries with the Académies des Sciences (see Hahn (1971), pp. 109/10). However this situation was not reproduced with the Journal économique and Guettard seems to have been well-disposed toward the group of people that led it. Puismarais (1754), p. 74

Duchesne’s Strawberries: Between Grower’s Practices and Academic Knowledge

Certain horticulturists wondered if it was possible to “treat the young trees in the same manner as the flowers, in order to change the structure of their organs, and create, so to speak, double species”.21 Notably, there was a list of “good species” in which transformations were easier to carry out: anemone, auricula, carnation, crocuses, daisy, larspur, hepatica, primrose, tulip, violet and wallflower. It was also currently admitted that if a double flower ceased to be cared for, it returned to the state of simple flower. The second example of changes in plants was well-known in the eighteenth-century, and concerns the transmission of colors in tulip and other flowers. Indeed, “when cutting all the stamens of a red tulip before the emission of dust, and when powdering the stigma of this same tulip with the dust of the stamen of a white tulip”, the results was the following: “the seeds obtained were red for a part, white for another part and red and white for the last part”. However no quantification of the result was observed or reported by eighteenth-century scholars. This example was reported by Adanson who presented it as the result of a basic hybridization of varieties of the same species.22 Perhaps because it was a too much known example – which is what Adanson says – careful experiments on hybridization are not frequently reported in Journal économique, contrary to their place in academic Mémoires, like those by Koelreuter and Spallanzani. However, hybridization was certainly used by growers and breeders. Yet, to a large extent based on tacit knowledge, heredity was embodied in various practices. Except the word degeneration, I could not find other particular names (such as “atavism”, “reversion”, “heredity” used in the nineteenth-century) referring to the return to the natural condition of the specimen. The language for hybridization is not particularly technical. The positive impact of human art on nature was a common model by which florists, growers and cattle farmers understood their own practices of transforming the sorts and species. 23 As shown by historians of gardening, a shared project of the practitioners was to push nature out of its limits, including morphology and calendar rhythms. In the case of plants, not only the morphological properties of the organisms were affected, but even their blooming season. Indeed certain methods displayed in Journal économique made particular flowers (hyacinth and others) blooming at a precise date, to serve for private parties and Christian festivals. 24 However, such positive impact of art on nature was limited by teratological considerations, notably when produced by man hands. Grafting was a technique used to change either the morphology or color of certain species, for instance an anonymous was able to produce “green or yellow roses”, thanks to a graft.25 In several papers, authors reported various grafting “experiments,” through which a horned cock was obtained, or a pear-apple, or seedless cherries, etc. 26 For some of these productions, the authors discussed the question of constancy, yet they usually regarded them as monstrous productions with no possibility of sexual reproduction.

20 21 22 23 24 25 26

Anonymous E (1758). Anonymous I (1761), p. 172. Adanson (1763), p. cxiii. Anonymous F (1758), p. 500. Anonymous G (1760), p. 335. Anonymous E (1758), p. 170. Anonymous D (1757); Anonymous H (1761); Anonymous I (1761).

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To understand Duchesne’s enterprise, it is necessary to remind that he knew a great deal of the breeder, florist and grower’s practices. Notably he quoted several authors of this “tradition” and his hundred page of domestic economy section shows that he was himself a grower. This will help us to establish the contribution of the grower’s tradition to his own enterprise. For instance, Duchesne knew well a town particularly renowned in Journal économique as an important essay and market place for breeders and florists: Montreuil. The Montreuil community of growers was famous for its products supported by horticultural and agricultural experiments. This small town close to Paris was especially known for its practices of selection of new races of fruits, notably peaches and strawberries, which dated back to the first part of the seventeenth-century. 27 Duchesne had taken much of his domestic economy section from “the practices of the grower traders of the town of the Wood [from Boulogne?] and of Montreuil”. 28 As a synthesis of the grower and breeder’s practices, it is worth saying that one could hardly identify a single tradition of gardening and breeding in the Ancien Régime. The know-how depended on the country, and on the type of plant or animal breeding implemented. Silkworm was present in Italy and France since the Renaissance, while breeding flowers occupied much place in the Dutch Republic and in certain areas of Germany. Moreover, new trends were developed for gardening, for instance the expansion of the English home gardening between 1750 and 1850. 29 Thus it is perhaps better to speak of the diversity of grower and breeders’ applied practices, opposed to the monolithic system of the botanical tradition. Yet, the diversity among practitioners had nevertheless a unity which was confined geographically to particular places. Indeed Paris was, during the eighteenth-century and up until the middle of the nineteenth-century, a city surrounded by many small towns which produced goods and especially vegetables. They fed the capital bringing the goods to the Halles. There are scenes of the Parisian life of les Halles described by Balzac that show the wide circulation of vegetable goods in Paris during the first half of the nineteenth century. From the seventeenth to the eighteenth-century, many of these surrounding villages were material to the training of growers, breeders and practitioners as well. Rozier’s Cours complet d’agriculture reports that young apprentice florists complied with masters from many of these towns which had a specialised production of vegetables. Montreuil was specialised in producing fruits and particularly peaches, while other such as Montmorency, Bagnolet, Vincennes, Charonne, etc. cultivated specialty fruits. This environment gave its bedrock to a localised tradition of practitioners as opposed to the international tradition of botany. The regional tradition was destroyed before the end of the nineteenth century with the growth of Paris and the railways that facilitated transport of goods from other parts of France and developed other markets. 3.2 Varieties and new species between botanists and practitioners Even criticised by Linnaeus since the 1730s as a tower of Babel, the botanical tradition was certainly much more united than were the florists and growers, who were usually organised in local guilds. Botanists worked within an international network. Indeed the methodological rules 27 28 29

52

Schabot (1758), pp. 75/76; Rozier (1787), p. 56. Duchesne (1766a), p. x. See Lustig (2000).

Duchesne’s Strawberries: Between Grower’s Practices and Academic Knowledge

and skills of communication, notably the international use of Latin, enabled a network of scholars stretched over Europe and the world to collect plants and information, to give plants a place – in any classification or in the Systema naturae – and to spread botanical information. As well, an important difference between botanists and growers is that the botanical tradition kept its memory alive, listed hundred of names, while among the growers and practitioners, such a relation to memory was much less vivid. They wrote papers in journals, treatises, dictionaries of gardening and Traité d’Agriculture, but except for a few heroes such as Thomas Fairchild (16671729) or Olivier de Serres (1539-1619), the vast majority remained actually anonymous until the second half of the eighteenth-century. The new species question was also an affair of controlling geographical areas. However it was a type of control different from the strategies and know-how of the practitioners. When Peloria was reported to Linnaeus, when Fragaria Monophylla appeared in Versailles, both were considered undescribed species, which is not saying they had a recent origin. Adanson and other travelers described hundreds of new species during their travels out of Europe, but they did not consider them as new species. There are actually two ways of conceiving the novelty of a species, which depended much on the extant and previous control of an area by botanists. In an area already checked by a centuries-long exploration, like Europe, any undescribed species should appear as a new one, which was of course not the case for species abroad. The new species problem reveals us something of the geographical control by botanists, a territory into which worked nevertheless also the practitioners. It is known that Linnaeus’ specific project was to unify his “army of botanists” thanks to institutional means as well as guideline books such as Critica botanica and Philosophia botanica.30 In these books he notably formalised the botanical rules, for nomenclature and the morphological language, addressed to the community of the botanists. Franz Stafleu, Jean-Marc Drouin and Pascal Duris have stressed both the sociological model supplied by Linnaeus for botanists and his social impact on the formation of a new type of scientific society at the end of the eighteenthcentury. Moreover, Linnaeus was also very attentive in working economically, reducing the number of species while eliminating varieties from the botanist’s realm. Indeed, while, in 1623, Bauhin had described about 6000 species, Ray 18’500, Linnaeus reduced the number to 7000. The barrier towards varieties he raised pushed dramatically the study of variations back to nature and put it in the hands of the florists and growers, a fracture that strengthened the distinction between the two professions.31 Furthermore this impetus helped to forge the botanical memory and identity for it extended what John Ray said half a century before, that botanists should not be interested in varieties. The economic way of treating botany by Linnaeus entailed the elimination of sub-species, but, when discarding varieties, Ray and Linnaeus had touched a level other than organising the botanist’s duties. They actually blocked any access to “sources of variation” and avoided bringing confusion between things that vary (varieties) and things that, as a rule, do not vary: species and genera.32 Rejecting the varieties out of botany was a sort of political and theological act that eliminated variation from the botanist’s cabinet, whose role was to lock up the myth of the fixity of the species – or genus. Thus, the world of growers and breeders, whose role 30 31

Stafleu (1971). Linnaeus (1751).

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was to play with varieties, sorts, hybrids, etc., was clearly set apart from Systema Naturae. Four events appeared in the story that disturbed this decision of Linnaeus. First, the definition of species by Ray himself was a sort of genealogical definition. Second, the fixist system was seriously challenged by several plants that did not obey to Linnaeus’ rules, such as Mercurialis from Marchant, Peloria from Rudbeck and Linnaeus himself, and Delphinium from Gmelin. Third, certain botanists, such as Michel Adanson, considered that varieties shall not be neglected by botanists, a position that appealed for a massive work of re-classification Adanson was not able to carry out alone. It nevertheless gave birth to the latter “evolutionary” theory. 33 Fourth, Duchesne’s strawberries are a kind of return of the repressed voice of the florists, breeders and growers, who acquire their meaning, partly from being placed at the exact cross border which Linnaeus wanted to remain a no man’s land. Duchesne pushed further the exploration of this no man’s land. He had published, at the age of seventeen, his 1764 Manuel de botanique, that used both French and Latin Linnaean nomenclature, and dealt only with the utility of plants for feeding, medicine, arts and decoration.34 In Histoire naturelle des fraisiers, the true border was the question of the constancy: Botanist worked with species and rejected varieties which were not constant. This was clearly stated by a botanist such as Chrétien Guillaume Lamoignon de Malesherbes in the early 1750s: “All these alleged species [pears, peaches, carnation, tulips] degenerate when one wants to multiply them from seeds; consequently they are only varieties for botanists”.35 It is a question whether the disciplinary boundaries of botany were strong enough to make someone considering himself a botanist even working on varieties, such as Adanson or Duchesne. Especially, many voices gathered at this time to agree with Linnaeus, but what about the botanists who defied the “taboo”? As noticed, Duchesne was a grower, as his discussion on Montreuil and other clues testify, but he was also a botanist, son of a grower and botanist. One shall stress Duchesne’s botanical skills, close to those of Linnaeus: use of Latin name, morphological description of species, use of classification, historical knowledge, quotation of botanists, correction of “bad description”, etc. Duchesne followed a similar method to that of the botanical tradition: a systematical and historical survey, for a particular species, of its name, description, morphological characters, differences, reproduction, cultivation, synonymy, authors, geography, to which remarks were added. Such a schema is as old as the Renaissance botany and natural history. Other works show that he was deeply rooted in natural history tradition. The strawberries 32

33

34 35

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It is not clear to me what is the main fixed unity in the eighteenth-century botanical tradition, either the genus, the species or both. Such that the relationship of genus and species to nature was considered to be not arbitrary, while classes and orders could vary according to the system and methods of classification used. The diversity of systems of classification, that touched on orders, classes, section and other subdivisions, should not, as a rule, touch on the genera. In a text written in 1749 as a general criticism toward Buffon’s first two volumes of Histoire Naturelle, Malesherbes (sur l'histoire naturelle, I, 43) defined well the role of the various systems, “which are not made in order to establish the genera, but only to order [ranger] the genera already established” (Malesherbes ([1798] 1971), I, pp. 115/116). Adanson claimed his evolutionary conception in his 1763 Familles des Plantes He indeed admitted thus the possibility of creation of new species: “one could perhaps apply these examples to a number of insects, shellfish and worms, which would demonstrate the possibility of the mutations [mutations] or of the creations of new species in animals, as it seems proved that there are new species in plants showing to be not immutable” (Adanson (1763), p. clxii). See also Atran (1993), p. 306. Duchesne (1764), p. xv. Malesherbes ([1798] 1971), I, p. 31.

Duchesne’s Strawberries: Between Grower’s Practices and Academic Knowledge

particularly lead him to correspond with Linnaeus, Monti in Bologna, Allioni in Turin, Haller in Bern, other European botanists and collectors, and he acknowledged Bernard de Jussieu for the classification used in Histoire naturelle des fraisiers.36 The classification of the strawberries in the Rosacea family was used in Jardin de Trianon and Jussieu had authorized Duchesne to publish an abridgment of this family.37 The management of variety and variation by the botanical tradition and by grower-breeders reveals much of the tensions among botanists. Adanson remarked that “modern botanists do no agree with these changes, which are, to speak properly, only varieties more characterised”. 38 This led him to criticise the definition of species proposed by Buffon: defining the species as the product of a sexual generation worked for superior animals and vegetables. But, what about the other species that reproduced through parthenogenesis, budding, section, and hermaphroditism? “What about those whose individual produce varieties that change at each generation, or that will be fixed during several generation?”39 In term of Linnean botany, such a question did not have a lot of meaning, or perhaps better, it was cast out by the Linnean rules. Indeed, since varieties should be the object of practitioners such as gardeners and florists, they were excluded from the botanical discipline, as written in Philosophia botanica.40 Still, Linnaeus cast doubts on the constancy of the species during a decade after he had described the Peloria. Of course, the practitioners had not waited the Philosophia botanica to work on varieties and on the various methods of generation. The local tradition of the practitioners probably was enough unified to work with certain routines on varieties and races. Still, much probably, there was nothing such as a systematical search for new varieties appearing through the techniques of hybridization, although several example vouch for the existence of the latter practice. Duchesne was to treat much of these problems in several remarks of Histoire naturelle des fraisiers. 4. DUCHESNE’S ENTERPRISE: CULTURE, HISTORY, AND GENEALOGY. In his book, Duchesne first gives a historical survey on strawberries, including the changes due to the climate and other causes. Then he describes the various species of strawberries, placed within their family (Rosacea) and compared with other genera of the family. Duchesne thus begins to study each particular race, including morphological differences and history, “notifying at the same time what race seems the older, and concerning the new ones I indicate those among which they came from”.41 This is followed by a summary of the characters particular to each race together with their genealogy. The section on domestic economy of strawberries includes reflections on art, ornamental, gardening, growers’ practices, harmful insects, recipes, remedies, etc. 42 Histoire naturelle des fraisiers contains a second part, distinguished by a separate pagination. Duchesne 36 37 38 39 40 41 42

Duchesne (1766a), pp. vii-viii. Antoine-Laurent de Jussieu who worked with his uncle Bernard published his major work Genera plantarum (1789), a foundation for the natural method, see Stevens (1994). Adanson (1763), p. clxii. “les botanistes modernes, ne conviennent pas de ces changements qui cependant ne sont, à proprement parler, que des variétés seulement plus marquées.” Adanson (1763), p. clxiii. Linnaeus (1751), § 306: “Botanists have nothing to sort out about varieties.” Duchesne (1766a), p. ix: “avertissant en même temps quelles races paroissent les plus anciennes, et indiquant pour les nouvelles celles dont elles ont pris naissance”. Duchesne’s plan takes place at pages viii-x.

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added there a hundred pages containing five “particular remarks”, intended to develop “certain interesting points, too much weighted with details [which] I could not discuss as much as they deserved”.43 The five remarks deal respectively with the question of the constancy of the race, the definition of varieties, species and genus, the existence of hermaphrodite strawberries, the comparison between hybrids in plants and animals, and the synonymy. 44 4.1 Of species, races and constancy The question of the limits between the species, the variety and the race is topical both to understand the relationship between botany and practitioners, as well as Duchesne’s enterprise. In his first remark, he addresses the influence of the method of generation (sexual or budding) on the constancy of a race. Commenting on a passage from Miller’s Gardener’s Dictionary, Duchesne emphasised that only the sexual reproduction allowed certain races and varieties to appear and to become constant, while budding and even grafting never give birth to a species, because the plant obtained was an extension of the same specimen. Such that it could not change. For these plants, the second generation – thus the new specimen – does not usually produce seeds. 45 This distinction, used by practitioners, could be useful to understand the limits between species and varieties. On these limits, the second remark focused stressing the status of the new strawberries: “is it a species? Then there are new species. Is it a variety? Thus how many varieties are there in other genera, considered to be species?”.46 Through these questions, Duchesne opened the door to a redefinition of the species that expected to bring the fixed in harmony with the variable. As we saw above, several historians have considered that, for Duchesne, the race was a species, or that he had changed one term for the other. Nothing is less sure. Actually Duchesne adopted the botanical definitions of his time and was sharply clear on the distinction between species and race, the first being fix and the second mutable.47 Duchesne himself criticised the usual confusion between race and species, taking the example of the human being.48 Once established the necessity to eliminate the confusion, yet a second problem emerged, i.e. to distinguish the meaning of constancy for a species and for a variety. Indeed, constancy was a property of the species, and how could one deal with constancy for a race? The botanists had carefully erased the problem of the constancy of the race from their field of vision. It was the discovery of F. Monophylla which obliged Duchesne to rethink the story as a whole and led him write that “cultivation and other accidental causes do not produce new species, but they cause, in certain individuals, some changes which, being persistent through their posterity, make new races”.49 Once admitted that new races could originate from stables species, the questions were two: first, how to secure the distinction between the race and the species; second, 43 44 45 46 47

48 49

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Duchesne (1766b), p. 1. Duchesne (1766a), p. xi-xii. Duchesne (1766b), pp. 5/6, 8. Ibid., 13. Duchesne (1766a), p. 135; Idem (1766b), p. 14. The second remark comments on the “distinction one must establish of the fixed and invariable characters of the species, from the slight and changing differences of the Races; on the Constancy of the one and the Mutability of the others.” Duchesne (1766b), pp. 18/19. Ibid., p. 21.

Duchesne’s Strawberries: Between Grower’s Practices and Academic Knowledge

what is the relationship between the parent species and the race offspring? For the first problem, cross-breeding of various species allowed to decide which were the species, but it was a too long method – and it could raise particular difficulties – such that Duchesne reverted to consideration on an adequate morphological description. But a new plant – such as the Peloria – and new races such as F. Monophylla raised the unknown problem, and helped to cross the borders of the botanical world to penetrate into that of the practitioners, which made the system of the fixed species overturning. The cause of such a “mess”, Duchesne saw it in the fact that many specific characters were not the true one. Thus, what had been built on the Peloria – making hybridization a model to understand the origin of many new species by Linnaeus – was made “in small” with F. monophylla, raising attention to the conceptions and practices of growers and breeders. It was actually the practitioners, growers and cattle farmers, who employed the category of “constant variety”, usually termed race. Buffon had used it in Histoire Naturelle des animaux, and, for Duchesne, this term needed “to be introduced into that of the vegetables”.50 The field in-between the classical botany and the grower’s practices could be defined through this notion, because botanists maintained the fatal confusion where practitioners had eliminated it: “following Ray’s axiom, since one can not label variety a constant race, they are named species”. 51 Being both a botanist and a grower, Duchesne could easily admit constancy for certain types of strawberries and thus land on his feet, considering “all the strawberries as a species distinct from all the others, and each strawberry, taken particularly, as a race or a variety. [The reasoning] led me to look for their genealogy”.52 4.2 A new reading of history? What kind of influence did the botanical tradition have on Duchesne’s enterprise? What were the routines and mental habits of a botanist in the second half of the eighteenth-century? Although there was a tendency to use vernacular languages for publication, botanists still knew Latin, used it for morphology and nomenclature of plants, as well as classification. They also made reference to a two to three centuries year old tradition. Which means that an eighteenth-century botanist had a particular relation to memory and, going back to the Renaissance, referred to names and morphological descriptions of plants recorded by previous botanists. Such relationship to history in the botanical tradition was much distinct from that of another important tradition, the experimental tradition, which takes its roots in the second half of the seventeenth-century. 53 And indeed, when one looks after the historical perception demonstrated by eighteenth-century experimentalists, they usually – with a few exceptions54– do not refer to works before the 16601680s. The line that separates the “Ancients” from the “Moderns” is roughly situated at the end or 50 51 52 53 54

Ibid., p. 18. Ibid., p. 26. Ibid., p. 14. Many historians have studied the social, rhetorical and cognitive novelty both for physical and natural sciences; see Dear (1985), Shapin and Schaffer (1985), Bernardi (1986) and Licoppe (1994).” For instance certain Italian scholars such as Neapolitan (Bovi 1769, Cavolini 1785) still quote Aristotle, Galen or Pliny the Elder. But it is not the case of many experimentalists in the Center and North of Italy, such as Felice Fontana, Lazzaro Spallanzani, Bonaventura Corti, Maurizio Roffredi, Scarpa, who worked during the second half of the eighteenth-century.

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the second half of the seventeenth-century, corresponding with the birth of Academies and the famous querelle des anciens et des modernes. Such a line of separation did not exist in the botanical and natural history tradition. To speak only of the botanical tradition, although there are controversies of historians on the boundaries between the herbalist tradition and the botanical tradition,55 at least, eighteenth-century botanists has in mind to consider everything of the tradition. Indeed Linnaeus’ classification of botanists, his Bibliotheca botanica, and Haller’s Bibliotheca botanica left place for Renaissance authors. Similarly when Adanson speaks of the botanical iconography, he begun his inventory with Corbichon and Cuba in 1482, and Leonicenus in 1491.56 Of course, the plan for systematically surveying the ancient literature was both critical and cumulative. Critical in order to identify and separate copyists from creators; for instance among the 70’000 engravings of plants Adanson had identified, he regarded only 10’000 as original species, 60’000 were copies of the originals, and only 2000 to 3000 were “good” according to him.57 The aim was also to establish a cumulative selection of data to avoid loosing information on authors, plants, figures, names, descriptions, classifications, etc. While the reading of history was a duty in the botanical tradition, it was probably seldom carried out in the way Duchesne established in his book. An important point of rupture in Duchesne’s thought was his temporal interpretation of a historical material. Linnaeus, who did it for the Peloria, probably influenced him.58 Duchesne conferred a historical meaning to the observations on strawberries made by the botanists since Antiquity. An example of his method of reading the botanical history will make clear what I mean by temporal interpretation of history: “In 1665, Colbert (...) asked for the catalogue of the King’s garden (..) They included the double strawberry. At around the same time, Morison also describes it (...) Father Barrelier, who died in 1673, drew it (..) it has been engraved since, and published in 1714. (...). Zanoni, at the same time, drew also it (...); it is in his botanical history published in 1675 (..). It appears in Furetière’s dictionary, from 1690 (...) M. Haller quotes a description of it (..) in the memoir of Breslau, July 1722”.59 Much of Duchesne’s reading of the history of strawberries is characterised by a focus on context, dates and places, and by a frequent use of temporal connectors. This temporal history supplied Duchesne’s with the historical ground for his genealogical conception, as showed in the following example: “From Plymouth where our strawberry seems to be born, it arrived in Leyde, then in Paris, and eventually in Bologna, where Zanoni saw it in 1675. It also stayed in England, its native country, for Ray said in 1686, he had cultivated it during several years in Cambridge. But since that time no one saw it”.60 Everything thus pushed Duchesne to reshape the relationships among races into a classification that would include the multiple pressures of time. The genealogical tree was the main and probably the most efficient model available for the job.

55 56 57 58 59 60

58

See Atran (1993). Adanson (1763), pp. lxxxi, cxlii. Ibid., p. cxlii. I’m indebted to Staffan Müller-Wille for this information. Duchesne (1766a), pp. 77/79. My italics. Ibid., p. 104.

Duchesne’s Strawberries: Between Grower’s Practices and Academic Knowledge

4.3 From genealogy to experiment on new races Duchesne’s use of genealogy was probably enriched also by a socio-cultural influence, notably the use of the nobiliary conception and language. For instance even the term race seldom appears in the papers of the Journal économique,61 which usually prefers to use “sort”. “Souche” i.e. stock, stump, is used by Marquis de Puismarais to qualify sheep; he also employs the term alliance. 62 Other terms used by Duchesne point clearly at their origin in the language of nobility and human heredity. “Head of the race” was used by Buffon; “Offspring” (postérité) is frequent in Duchesne; “Head of the tree”, “branch of the same house”, “genealogical tree”, “genealogy”, “family”, belong to a vocabulary borrowed to the nobiliary conception of heredity.63 Duchesne had been among the first, after Adanson, to generalise the term family in his Manuel de Botanique. Another interesting clue for the relationship between hybridization and noble genealogy, is that many specialty fruits, and especially strawberries, received noble names: Duke of Kent’s Scarlet, Comte de Paris, Princesse Royale, Duc de Malakoff were, among others, names given to new varieties of strawberries during the second part of the nineteenth century.64 I see three rationale that could account for that. First, the awareness of a true genealogical origin was a shared knowledge in the breeding milieu, yet it did not necessarily vouch for the presence of an evolutionary issue. Second, the mixture supposed by the process of hybridization could be exorcised through a noble name. Third, a patron could be thanked through this habits. Still the genealogical tree of Duchesne (fig. 2) owns its proper logic, which makes it a particular model for understanding the descent from a race of strawberries to another. It is organised according to certain rules and contains symbols that obey to them. The first distinction Duchesne made was to separate races that were constant through the seeds (bigger rectangles), from varieties or monstrous specimens resulting from these constant races (smaller and dotted line rectangles). Such that the genealogical descent was stopped with the varieties but could theoretically last with a possible constant race. The second rule was to place the head of all races on the top of the tree, quite similarly to the way used for a human being genealogical tree. Consequently, the younger races are on the bottom. The third rule relates to the proximity relation among several races. The three races 3, 4 and 5, close to each other “have several common characters” which make possible reducing them to a common root. The places of the other strawberries were organised according to morphological and ecological similarities, and of course, genealogical descent. The 7th comes from the 6th, the 10th from the 2d, and the 6th from the first. The kind of lines used in the tree had also a signification, the normal line designates a descent without hybridization, and the wavy line a hybridization: the 9th, F. Ananassa (fig. 3) was suspected to come from a crossbreeding of 8 and 10,65 and a new race was expected to come from the crossbreeding of 7 and 8. 61 62 63

64 65

Anonymous H (1761), p. 122. Puismarais (1754), p. 70. “Souche”: Duchesne (1766a), p. 224. “Chef de race”: idem (1766b), pp. 23, 47, 63. “Tête de l’arbre”, “branches de la même maison,” “Arbre généalogique”: idem (1766a), pp. 214, 221. “Famille”, a botanical term much used by Adanson and Jussieu is also represented in Duchesne’s work. Darrow (1966), ch. 6, “Breeding in France 1770 to 1900.” Indeed, modern botanists such as Darrow considers that Duchesne “must be credited as the first to identify F. chiloensis x F. virginiana as the origin of the modern strawberry” (Darrow (1966), ch. 6). In the plate, F. chiloensis is frutiller (8) F. virginiana is fraisier écarlate (10) and the modern strawberry (F. ananassa) is fraisier-ananas (9).

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Fig. 3 Fragaria Ananassa from Rozier’s Cours d’agriculture. However, if it established a temporal relationship from the origin (top) to the last races (bottom), the time in the genealogical tree was not quantified precisely, even if Duchesne had proposed many hints in his historical reconstruction of strawberries that could help to do so. He was able to date the appearance of certain strawberries, and which were the older to have been described by the ancients. This helped him to establish Fragaria semperflorens as the head of the line. In addressing the questions of genealogy and new constant races, Duchesne proved also to be consistent with his main methodology and his numerous appeals for naturalists to use empirical means. Indeed, his genealogical investigation had an important counterpart in his experimentation. Duchesne actually carried out himself experiments of hybridization, with the scope of testing the constant race issue. Such that Histoire naturelle des fraisiers discusses two new races and not only one. The first race was accidental and named strawberry from Versailles (F. monophylla) by Duchesne, who managed to reproduce it through seeds. But the second race was

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the result of an experimentation. Having received a female Frutiller, without a male of the race, Duchesne placed it in the house of the male Capiton, to which it resembled. A week after, the Frutillier gave a fruit. Duchesnes tried to repeat the experiment, and, thanks to the help of people from the Jardin du Roi, they obtained four Frutiller pollinated by the Capiton. The seeds were given to several botanists, such as Jussieu and Richard, who sowed them at the time the printing press was running on Duchesne’s book. Other scholars, such as Le Monnier, and Le Normand had tried to cross the Frutiller with F. Silvestris, but with no success. On this point, Histoire naturelle des fraisiers closes on a disappointing end, and the fate of the current hybridization is not reported. However, the practitioners conserved its memory. Thirty years later, Abbé Rozier and a society of people’s Cours complet d’agriculture solved the question when showing, at the entry “Strawberries”, the genealogical tree of Duchesne, with the new race which was now undoubtedly constant (fig. 4) and identified to the Scarlet strawberry from Bath. 66 Indeed the genealogy contains the new name for the race number 11, and another race was added by the author (number 12). Duchesne’s experiment seems to have left, at least in the late eighteenth-century, some offspring.

66

Rozier (1787), p. 50, pl. II.

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Fig. 4 Genealogy of the strawberries in Rozier’s Cours d’agriculture. Yet, in the 1780s, after Duchesne’s discovery was diffused and adapt to the knowledge of practitioners and botanists, the difference were still important between them, notably on the concept of species. Rozier remarked that the botanists distinguished only three species of fragaria, (semperflorens, muricata et sterilis). While, on the other hand, practitioners gave to the definition of species a larger extension, because they considered the constant varieties to be species, and

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labeled “variety” the sorts that derive from these “species”. It seems thus that two models for the species co-existed in France at the end of the eighteenth century, which served the rather different interests of two distinct communities, the practitioners and the botanists. Yet it was, during this time, in the restricted context of the practitioners that the genealogical model emerged. CONCLUSION Various influences brought thus Duchesne to use the genealogical tree. In addition to the empirical approach to hybridization used by Marchant, Réaumur, Buffon, Sprengel, Koelreuter, Adanson and Gmelin, Duchesne developed a proper methodology. In order to explore the no man’s land of the “constant races” in-between botany and the practitioners, he systematically examined the history of botany-gardening, looking for the historical meaning of what means describing a plant, in term of human activity and history. In this respect, Duchesne cultivated a temporal history as opposed to a narrative history, that circulated, for instance through Buffon’s Histoire naturelle. The account of nature according to a temporal scheme was nevertheless the method used later by Darwin, Haeckel and De Vries. Duchesne’s attractive attempts of explanation such as the relationships between race, species and genealogy, his genealogical tree and use of a historical methodology, explain perhaps why certain historians of science tried to acclimatise his strawberries into their sometimes heroic history of science without practitioners. This story thus tells us the entry of a skilled practitioner into the no man’s land left unexplored in-between the two traditions, botany and breeding, and their hybridization that created a new field which crossed each border. Practices of heredity took a shape through secret know-how and hybridization cultivated by breeders, who demonstrate no particular quest for the understanding of these influences of art on nature. The double education of Duchesne drew him to look for a theoretical and schematic account of novelty according both to the current language and concepts used by breeders and to that of botany. Relevant to his problem, a nobiliary custom of illustrating genealogy was available by which he designed the first genealogical account of species in transformation. The tensions between tacit and explicit knowledge, morphological and genealogical definitions of the species, and each tradition, allowed to bring into focus new questions and are perhaps symptoms of an emergent discourse about heredity.

Institut d’histoire de la médecine et de la santé Université de Genève

Acknowledgements This paper is a improved version of a paper presented at a workshop on the cultural history of heredity at the MPIWG, Berlin. I want to thank Hans-Jörg Rheinberger, Peter McLaughlin, Staffan Müller-Wille and the participants to the MPI workshop for the discussion of this text.

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REFERENCES Primary sources Adanson, Michel. 1763. Familles des plantes, Paris. Anonymous A. 1752. ‘Pour faire venir sur un même pied des fleurs de la même espèce et de différentes couleurs’. Journal économique, juin 1752: 25–28. Anonymous B. 1756. ‘Méthode facile pour avoir des pommes et des poires dont les quartiers soient de différentes espèces’. Journal économique, avril 1756: 75–76. Anonymous C. 1756. ‘Observations sur la manière de greffer par M***’. Journal économique, avril 1756: 83– 95. Anonymous D. 1757. ‘Manière d’élever les oies et les canards, et de se procurer des canards mulets’. Journal économique, décembre 1757: 44–57. Anonymous E .1758. ‘Manière de faire venir des roses vertes et des jaunes’. Journal économique, avril 1758: 170. Anonymous F. 1758. ‘Observations oeconomiques sur les brebis, et la manière de les élever’. Journal économique, novembre 1758: 498–505. Anonymous G. 1760. ‘Méthode pour avoir en hiver des fleurs naturelles nouvelles épanouies le jour que l’on veut’. Journal économique, juillet 1760: 335. Anonymous H. 1761. ‘Des coqs cornus, et de la manière de leur faire venir des cornes sur la tête’. Journal économique, Mars 1761: 120–122. Anonymous I. 1761. ‘Moyen d’avoir des cerises sans noyau’. Journal économique, Avril 1761: 171–172. Bonnet, Charles. [1764] 1781-1782. La contemplation de la nature. Neuchâtel: Fauche. Dhermont. 1756. ‘Observation Botanique’. Journal économique, mars 1756: 112–115. Duchesne, Antoine-Nicolas. 1764. Manuel de botanique. Paris: Didot. Duchesne, Antoine-Nicolas. 1766a. Histoire naturelle des Fraisiers. Paris. Duchesne, Antoine-Nicolas. 1766b. Remarques particulières. [second and separated section in Histoire naturelle des Fraisiers. Paris]. Krachenninikow. 1755. ‘Nouvel espèce d’érable’. Journal économique, février 1755: 173–174. Lardilion. 1756. ‘Dissertation sur la monstruosité des prunes’. Journal économique, mai 1756: 35–58. Le Camus. 1754. ’sur le mémoire de Jourdan de Pelerin’. Journal économique, décembre 1754: 121–124. Linnaeus, Carl von. 1751. Philosophia botanica. Stockholm. Linnaeus, Carl von. 1767. Systema naturae (1766-68). Editio duodecima. Stockholm: Laurentius Salvius. Malesherbes, Chrétien-Guillaume de Lamoignon de. [1798] 1971. Observations sur l'histoire naturelle, générale et particulière de Buffon et Daubenton. Paris, Genève: Slatkine reprints. Miller. The Gardener’s and Florist’s dictionary, or a complete system of Horticulture. London. Puismarais, Marquis de. 1754. ‘Mémoire sur les moyens de bonnifier les laines dans les provinces du royaume’. Journal économique, août 1754: 68–77. Roger, Abbé. 1755. ’sur les village de Montreuil, Bagnolet, Vincennes, Charonne et village adjacents à deux lieus ou environ de Paris, au sujet de la culture des végétaux, avec une idée de leur méthode de traiter les arbres, surtout les pêchers’. Journal économique, février 1755: 44–79. Rozier. 1787. “De la culture des fraisiers,” Cours complet d’agriculture, Vol. 5. Paris: Serpente, 56–59. Schabot, Roger Abbé. 1758. ‘De la culture des fraisiers’. Journal économique, avril 1758: 248–253. Secondary literature Atran, Scott. 1993. Cognitive foundations of natural history. Cambridge: Cambridge University Press. Barsanti, Giulio. 1992. La Scala, la Mappa, l’Albero Immagini e classificazioni della nature fra Sei et Ottocento. Firenze: Sansoni. Bernardi, Walter. 1986. Le metafisiche dell’embrione, scienza della vita e filosofia da Malpighi a Spallanzani, 1672-1793. Firenze: Olschki. Blaringhem, L. 1911. Les transformations brusques des êtres vivants. Paris: Flammarion. Bowler, Peter J. 1973. ‘Bonnet and Buffon, Theories of Generation and the Problem of Species’. JHB 6: 259– 281. Darrow, G.M. 1966. The Strawberry: History, Breeding and Physiology. New York: Holt, Rinehart and Winston. De Candolle, Alphonse. 1881. ‘Darwin considéré au point de vue des causes de son succès et de l’importance de ses travaux’. Arch Sc. Genève: VII.

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De Vries Hugo. 1909. Espèces et variété. Paris: Alcan. Dear, Peter. 1985. ‘Totius in Verba: Rhetoric and Authority in the Early Royal Society’. Isis 76: 145–161. Farley, John. 1977. The Spontaneous Generation Controversy from Descartes to Oparin. Baltimore and London: The Johns Hopkins University Press. Guyénot, Emile. 1941. Les sciences de la vie au XVII et XVIIIe siècles, l’idée d’évolution. Paris: Albin Michel. Hahn, Roger. 1971. The anatomy of a scientific institution: the Paris Academy of Sciences, 1666-1803. Berkeley, Los Angeles: University of California Press. Hylander, N. 1945. ‘Linné, Duchesne och smultronen’. Svenska Linné-Sällskapets årsskrift 28: 17–40. Larson, James L. 1994. Interpreting Nature The Science of Living Forms from Linnaeus to Kant. Baltimore: The Johns Hopkins Press. Licoppe, Christian. 1994. ‘The Crystallization of a New Narrative Form in Experimental Reports (16601690) The Experimental Evidence as a Transaction between Philosophical Knowledge and Aristocratic Power’. SIC 7(2): 205–244. Lustig, A. J. 2000. ‘Cultivating knowledge in nineteenth-century English gardens’. Science in Context 13: 155–181. Rey, Roselyne. 1989. ‘Génération et hérédité au XVIIIe siècle’. In L’ordre des caractères, aspects de l’hérédité dans l’histoire des sciences de l’homme, edited by C. Bénichou. Paris: Science en Situation, pp. 7–48. Roger, Jacques. 1963. Les sciences de la vie au XVIIème et XVIIIème siècle. Paris: Colin. Shapin, Steven & Simon Schaffer. 1985. Leviathan and the Air-Pump, Hobbes, Boyle, and the Experimental Life. Princeton: Princeton University Press. Stafleu, Franz. 1971. Linnaeus and the Linnaeans. The spreading of their ideas in systematic botany, 1735-1789. Utrecht: International Association for Plant Taxonomy. Stevens, P. F. 1994. The development of biological systematics. New York: Columbia University Press. Tassy, Pascal. 1991. L’arbre à remonter le temps, s.l. Christian Bourgois Editeur.

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Natural Things and Non-natural Things. The Boundaries of the Hereditary in the 18th Century Carlos López-Beltrán

1. Hereditary transmission of bodily (physical) and behavioral (moral) features from parents to offspring became an independent subject of scientific theorizing only in the mid-decades of the nineteenth century. Only then did it become clear that the questions around the stability of species and the question about the contingent similarities responsible for family resemblance were tightly intermingled, and that both could be explained in a unified way. The problem of how transmission is effected became central issue for biologists.1 Until then the issue of the conservation of type within a genealogical line was considered a major biological problem, whereas the phenomenology of hereditary communication of accidental traits within the lineages was normally seen as of secondary importance, except in a few local contexts, such as among animal and plant breeders and particularly among some physiologists and medical men, whose interests guided them towards considering the possibility of a regularity in the transmission of traits from ancestors to offspring within a genealogical line. Elsewhere I defended the view that the appearance of the word Heredity, after the decade of the 1830’s, first in French and then in other European languages, signals the turning point, as the shift from an adjectival use (in which there always was a peculiarity that was called hereditary) to the nominal use (Hérédité) reveals a growing awareness of the existence of a causal pathway responsible for observed regularities, and irregularities.2 I have also contended that before that period there existed among medical men and naturalists a set of accepted phenomena associated with the use of the adjective hereditary which were explained (or explained away) in different fashions during the different epochs. A rather modest tradition of paying attention to how hereditary accidents are communicated through lineages can be followed from Antiquity to modern times. One can find within it a growing number of descriptions, and of attempted explanations of how the phenomenom occurs and why it can be responsible for similarities of physical and moral characters within such lineages. Some well known examples are Aristotle’s mixed (cultural-hereditarian) explanation of the “Longheads”,3 Aristotle’s complex physiological (dynamic) explanation of resemblance of the offspring to both parents.4 Hippocratic and Galenic appeals to dual-seminal models of reproduction in order to account for resemblances in general.5 Medieval astrological accounts of children’s peculiarities of feature. Paracelsian influential dualistic theory, in which the

1 2 3 4 5

López-Beltrán (1992), Idem. (1994), Churchill (1987). López-Beltrán (1992), Idem. (1994). See Glacken (1967). See Coles (1995). See Boylan (1984); Idem. (1986); Jacob (1970); Lloyd (1983); Coles (1995).

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imagination could induce both resemblances and monstrosities 6. The medical attempts (after the Renaissance) to account for hereditary transmission of disease within family lines under different theoretical frameworks.7 In 1775 I. Kant proposed an analytical distinction that can be of use in our descriptions. He suggested that hereditary variations should be called “resemblances” if “they agree with their derivations” (the offspring “takes after” one or both parents, or some ancestor), and be called degenerations (or “expeciations” according to one translator) if they moved away from the norm in such a way “that the original stem-formation cannot be restored”. 8 By the mid-eighteenth century a more or less stable set of phenomena were usually included in this peculiar category of the hereditary. Be it resemblances or degenerations, the fact that inessential peculiarities of feature not shared by all the members of a species or lineage, managed to be re-produced with some fidelity in the descendants, was considered a stumbling block for descriptive and explanatory systems in both Natural History and Physiology. It can be said that a special relationship was forged during the 17th and 18th century between such set of hereditary phenomena and the theoretical models of generation that were vehemently discussed. Resemblance to both parents, the mixed peculiarities of mules, the origins and transmission of monstrosity, etc. became a probing, evidential ground for the dispute. What I call the hereditary constituted thus, from the mid decades of the eighteenth century on, a set of heterogeneous phenomena that anyone had to “save” in order to consolidate his view on generation. Chamber´s Dictionary (1738) for instance, in his entry for “Generation” mentions that Sir John Floyer “starts a difficulty which seems to press equally against each system (ovism and animalculism), taken singly”... the fact that mules partake of the characteristics of both horse and ass, and that the defenders of both systems capriciously choose the characters that favor their view as the important ones for the determination of the origin of the foetus, having the characters conveyed by the opposing sex as secondary.9 When Diderot was preparing in the 1750´s his Élemens de Physiologie, he assigned a special weight the hereditary facts for the evaluation of the several systems of generation he intended to describe. The difficulty that preformationist views had in dealing with “maladies héréditaires; resemblance des parens; mules et mulets qui engendrent” was particularly highlighted by him in those notes.10 At the beginning of his Considérations sur les Corps Organisés, Charles Bonnet posed some of the challenges his preformationist stance had to face: Si les germes sont contenus originairement dans les ovaires de la femelle, et si la matiere séminale n’est qu’une espece de fluide nourricier, destiné à devenir le principe du développement, d’où viennent les divers trait de resemblance des enfans avec ceux qui leur ont donné le jour? Pourquoi les Monstres? Comment se forment les Mulets?11 6 7 8

9 10 11

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See for instance Glacken (1967) and Radl (1930). See López-Beltrán (1992). See Kant (1775) In Chukwade (1997). Chukwade uses the term “expeciation” to translate Kant’s expression which covers similar semantical grounds as Buffon’s “dégénération”. Though Kant’s appeal to the original stem-formation shows, I believe, an attempt to a generality influenced by Bonnet and Blumenbach that Buffon´s concepts lacked, I will keep “degeneration” as the adequate translation. See “Generation” in Chamber’s Dictionary, vol. I, 2nd. edition (1738). See “Génération”, chapter XXIV. in Diderot ([1875] 1964), pp. 182–185. C.Bonnet ([1778] 1985), p. 31.

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In his Primae Linae Physiologiae, written in the period when he still sided with epigenesis, Haller outlined the facts that pushed him necessarily in that direction. That organisms, especially hybrids, resemble both parents, simply “rules out” he wrote, any possibility that the new being is preformed in one or the other parent.12 Diderot, in his adjudicator’s stance, knew well that even if the dual seminal (pangenetical) 13 models could account with more ease for the hereditary transmission of features by both parents, they had serious problems of their own when facing actual anatomical observations, and detailed physiological questioning. In his Élemens de Physiologie he writes “Dans ce sisteme (pangenesis) placenta, et envelopes impossibles a expliquer”. This kind of criticism was of course made forcefully by Haller and Bonnet against Buffon.14 An important difference to point out concerns the character of the empirical facts that posed problems for each competing approach to generation. While detailed observation of the organs of generation, and of the development of the embryo, backed strongly preformationist (specially ovist) positions, dual seminal (pangenetic) accounts were favored by what may be called “genealogical” observations (or pedigree following); that is the following of patterns of similitude and difference within lineages. While the first set of facts depend on a focus on single individual development, the second set imply a higher level, comparative perspective, that needs the observation of many individuals belonging to several generations. The kind of features that, due to curiosity or a special interest, were followed through the generations varied widely. From very vague family resemblances to precise weird characters like an extra digit, a spectacular mole or a snub nose, or on the pathological side from general tendencies to ill health to precise ailments that develop in exactly the same manner at a particular age. The “genealogical” approach to evidence and observation, I believe, opens up the possibility for setting external limits to physiological speculation,15 in contrast with the interior limits set by dissection and microscopy. The gathering of convincing cases of hereditary transmission of a wide range of different features, the progressive closure of different causal avenues for dealing with them,16 was one main theme of 18th century debates around generation. Important aspects of Bonnet’s increasingly sophisticated ovism (in which he strategically adopts several explanatory resources from the rival dual seminal theories) were no doubt a consequence of the strains put on preformationism by hereditary facts. It was not however an all powerful set of facts. The hereditary was an unstable domain, plagued with irregularities and exceptions. Not everybody agreed to grant it some reality. The fact of hereditary transmission itself could be put into question. When reviewing Buffon’s theory of generation, Haller considered that the hereditary posed a challenge important enough, to go to the 12

13

14 15 16

Haller collected a series of hereditary facts in his miscelaneous collection Similitude Parentum. In this period he was convinced that only a new formation could acccount for them. See Roe (1981), p. 25 and Guyenot (1957), p. 295. Whereas when discussing generation theories the main opposition between systems seems to be between preformation and succesion (say, epigenesis), in the case of hereditary transmission a crucial issue is the origin of the “information” concerning details (snub noses), and the opposition preformation vs. dual semen (or seed) is more relevant. See Haller (1752). By considering for instance the changing sets of conditions under which herediatary transmission occurs, when and with what regularity some features are preserved within a lineage. For example explaining them away by ascribing them to chance ocurrences or calling them irrelevant.

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extreme of denying its reality. “I prefer simply to deny to Mr. Buffon that offspring resemble their parents... the offspring are no longer images of their parents” (“et le reste de l’édifice tomberá de lui même”). Haller bases his belief in the greater number of exceptions than positive cases, and especially on the fact that internally (anatomically) there is never a shared pattern of nerves or veins between parents and offspring.17 When commenting this extreme denial by Haller, Duchesneau writes that it illustrates with particular clarity the “individual” character of generation, in the eyes of the savants of the first half of the 18th century. 18 I believe this statement deserves some clarification. Rather than “individual” the act of generation was considered the result of natural (or divine) laws that produced each individual separately. This made the consideration of genealogical links rather irrelevant for all the fundamental issues. What however remained under dispute was if generation was to be given responsibility for all the features, both general and paticular, of the individual, or only for the former. The latter option left the door open for the all things individual, accidental, to be affected by influences from both external and ancestral (hereditary) influences, somehow independently from how generation occurred. A denial similar to Haller’s was made in 1747 by the French physician Antoine Louis in a notorious discussion on hereditary disease.19 What Louis was keen in establishing there is the impossibility that any anatomical (“solid”) feature of the parents can serve as the origin of a similar feature in the offspring. Any similitude has to be due to common, external causes. What is singular, individual, is the acquisition of peculiarities by each new being. Generation is entirely another question. About the possibility of hereditary disease Louis writes: les desordres de l’oeconomie animale doivent s’acquerir particuliérement par chaque homme: toutes les maladies seront individuelles puisqu’elles doivent être postérieures a la formation des germes qui n’ont reçu aucune alteration dans leur principe.20

Causal boundaries of the hereditary were drawn in a very different manner than the one we have become habituated after the instauration of our nature-nurture distinction. The explanation of individual bodily peculiarities was still closely linked to complex and open-ended medical notion such as constitution, temperament, etc. where an interaction between external and internal elements was responsible for idiosyncratic features of form and (dis)function. The restrictive view that Haller and Louis (in their different projects) put forward for the notion of hereditary transmission of individual peculiarities somehow strangles (squeezes) the possibility of such transmission by closing the gap between two causally independent domains. The internal of the generation (or reproduction) of the germ (the first formation), and the external, circumstantial, influences, on the body. Among other things, what was being blocked by their arguments is the possibility that external influences (climate, nutrition, etc.) became somehow integrated into the lineages and eventually adopted in a non-accidental manner. For such authors, there is no conceivable way in which accidental variations of any kind could be 17

18 19 20

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Haller (1752), p. 32. The English translation by Phillip Sloan ( “Reflections on the theory of generation of Mr. Buffon”, p. 318) insists that the resemblances whose existence Haller is denying are “exact” replications, but that modification is not to be found in the French. Duchesneau (1982), footnote 132, p. 539. For a discussion of this dissertation see López-Beltrán (1994). Louis (1749), p. 35.

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inherited from parents to offspring with any regularity. Even if for instance family resemblances were caused by a transmission of peculiar features from parents to offspring, they would be unimportant. For them the hereditary belonged to the domain of anecdote. 2. A particularly suitable and surprising approximation to the conceptual frame under which hereditary matters were conceived within the medical tradition in the 18th century is given by the Galenic distinction between natural things and non-natural things. The boundary between body and environment, between physiology and milieu, for 18th century physicians can be neatly outlined with these concepts. Arnulfe D’Aumont writing for the Encyclopédie defined the (six) non-natural things| on appelle donc choses non-naturelles (d’aprés Galien21) celles qui en composent pas notre nature ou notre etre, mais dont l’économie animale éprouve des grands effects, des grands changémens, des grandes altérations.22

The list of the six non-naturals is somehow surprising for a modern eye: “l’Air, les Alimens, le Travail et le Repos, le Somneil et la Veille, les Excrétions retenues ou évacuées, et les Passions de l’Ame”.23 Galenists contrast these external factors with those called the (seven) natural things, and which are essential part by nature of the individual’s bodily constitution: les élemens, les tempéramens, les parties, les humeurs, les esprits, les facultés et les actions: ce sont celles qui concurrent à former le physique de notre être.

The coherence, logical and explanatory power of this conceptual framework has been studied and discussed in several papers by William Coleman. He writes for instance that in the 18th century “the doctrine of the non-naturals provided a concise, flexible, and widely accepted framework for articulating the primary demands imposed by the conditions of existence upon men and women who sought seriously to preserve their physical well-being”. (...)“The non-naturals became an integral part of a new and largely secular moral order.”24 I believe we can also deploy such conceptual frame to focus on the way the boundary between internal and external determinations of bodily features and constitution were delineated in the period, in order to raise the question of the permeability, and , in some sense, the fluidity, between the internal (natural) and the external (non-natural) actions. The ancient notion of the body as both a product of a mixture of humors ( crasis, temperament) and a constant subject to multiple humoral influences played an important role in this kind of issues. Questions linked to the individuality of temperament, and to the possible explanatory role 21

22 23 24

Galen inspired the concept of the six non–naturals, but did not really coin it himself. From a few suggestions he gave about which external influences were important to check in order to promote a healthy life, his followers ended up establishing a narrow list of six determinant factors. For historical studies of the concept see Nyebil (1971), Rather (1968), D’Aumont (1765), Nutton (1971). D’Aumont (1765). See Louis (1747), p. 18; D’Aumont (1765). Coleman (1974), p. 406. See also Idem. (1984).

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that geography (“airs, waters, and places”) and genealogy (family, tribe, nation) could play on some of its aspects. Taking Antoine Louis’ analysis again as an example of a radical view, he insisted in the extreme individuality of the body: Le tempérament des enfans qui naissent d’un même pere, et d’une même mere est presque tojours différent; les un son bilieux, les autres sanguins; les uns son guais, les autres sérieux, pésans: ces différences d’humeur, de caractere et d’inclination dans les freres et soeurs, sont des suites de la différence des tempéramens; et elle depend peut-être moins de la constitution primitive ou radicale, qui paroît devoir être la même dans tous les enfans; que d’une diposition acquise par la combinaison infiniment variée de toutes les choses extérieures.25

Among the external influences Louis mentions are the weather at birth, the suffering during birth, the amount of blood in vessels at birth, the quality of the nurse’s milk, the thickness of the air that was breathed during the first hours, etc. (“on ne finiroit à faire l’enumeration”). The earlier external influences have the more lasting effects on the individual’s temperament. Future illnesses (or dispositions to them) are often acquired at very early stages. Louis was adamant that this acquisitions (pathological or not) were not however transmitted to the next generation (si la) diversité des tempéramens n’est point héréditaire, comment les maladies qui ont les suites pourroient-elles se transmettre par les parens26 (...) les variations décident donc rien en faveur de la question des maladies héréditaires, puis quelles en vienent d’un principe interne et des dispositions inhérentes et immuables; mais qu’elles dependant uniquement des choses non-naturelles qui sont toutes extérieures27

As I have shown elsewhere,28 most of Louis’ critics, a few decades later, focused their attack on his “unbelievable” argument against the reality of hereditary transmission on what they saw as the false assumption that temperaments are all in all individual, secondary and accidental. A considerable proportion of medics believed that the possibility of transmitting bodily peculiarities through the family line was just impossible to deny. Contrary to Haller’s and Louis’ attitude, they claimed, facts should be given precedence over theory. To understand the differences on this issue among 18th century physicians an important “theoretical” division should be considered (one that to my knowledge has not been sufficiently discussed). The one which separated solidists from humoralits. The latter is of course the older tradition, and is responsible for the main conceptual scheme for Hippocratic-Galenic medicine. 29 Its rationale for a unity and diversity of human bodily and spiritual propensities turns around the balance of its fluid (humoral) constituents and their relation to the environment. 30 As Vivian Nutton wrote “the advantages of this logical scheme can best be seen in the development within humoralism of the theory of the six non-naturals”.31.Solidism is a medical outgrowth of modern 25 26 27

28 29

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Louis (1747), p. 35. Ibid., p. 37. Ibid., pp. 74/75. Further on he writes: “Les hommes sont soumis a cette regle generale comme les plantes et les animaux, leur caractère et leur tempérament dependent d’une infinie des choses extérieures qui peuvent être variées a l’infini: c’est une verité recinue en médecine.” López-Beltrán (1994). About the history of these medical doctrines and its links to hereditary disease see Portal (1808), and Adams (1814).

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mechanicism. It is tied to the search for mechanical, structural causes within the body. Theoretical stances like iatromechanics and the physiology of fibers are expressions of its main contention: disease and/or normality should be found in the physical properties of organization. Lesions (not a bad mixture of humors) are the causes of diseases. Over the issue of the permeability of the boundary body and environment these two theoretical positions worked under different presuppositions. If under a mechanical stance the solid parts (fibers, tissues, organs) are given causal (etiological) primacy over the fluid portions in the body,32 then it becomes (as Louis argues) much less likely that external presences (air, efluvia, waters, food, climate in general) could irreversibly affect the body’s properties; specially if the origin of the main features of the solids (organization) is thought of as preformed. In other words, the solid parts of each new being , ultimate bearers of all functional responsibility, will not receive any kind of regular, permanent influence, physiologically normal, from the equivalent solid parts of its ancestors; nothing acts at the first formation of the new being that deserves the name of heredity. On the other hand, if primacy is given to humoral mixtures (and if solids are thought of as product of a solidification or condensation of humors after fecundation), then the possibility arises that at the first formation of the new being its organizational plan acquires “original” alterations due to the peculiarities (the qualities) of the materials contributed by the parental seeds. It also becomes possible that those alterations acquire some “permanence” within the lineage, so that any causal influence that can find its way, through air, water, food, etc., and that dramatically changes a person’s humoral balance will alter his physical constitution. This will happen because such alteration in turn will affect the next generation’s reproductive humors like semen, blood, milk.33 In the context of hereditary traits (resemblances, degenerations) then, I believe that the distinction between naturals and non-naturals, and the conception of their interactions, can be illuminating. A traditional explanatory resource for variation within biological species, especially in the human case, is the appeal to external influences. The climatic (geographic) explanation is ancient. 34 During the 18th century the action of the sun, the air, the water, etc. was again given a central role in the explanation of diversity of bodily constitution (and of moeurs) by important theorists as 30

31 32

33

34

As Vivian Nutton defines it: “Humoralism is a system of medicine that considers illness to be the result of some disturbance in the natural balance of the humours, within the body as a whole or within one particular part. It stresses the unity of the body and the strong interaction between mental and physical processes. It is at one and the same time highly individualistic, for each person and each bodily part has their own natural humoral composition (also known as krasis, mixture, or temperament), and universal, for the range of variation is limited and the same patterns of illness (diseases) can be seen to occur in many individuals. The natural balance of health is always pecarious, for it is constantly subject to potentially harmful influences from one’s diet, life-style and environment.” “Humoralism” in Porter and Bynum (1993), p. 281. Nutton (1993), p. 288. Louis writes “l’action des fibres plus ou moins forte et vigoureuse, façonne et modifie différemment les humeurs de notre corps; ces huneurs agissent suivant leur quantité sur les solides dans lequels elles sont contenues, et elles en determinent diversement les actions: de-là viennent les complexions particulieres qui mettent tant de différence entre les hommes, tant par rapport aux dispositions du corps qu’aux caracteres de l’esprit.” Louis (1747), pp. 37/38. The capacity that one is willing to concede to a humor was a point in dispute. Haller was very critical of the way humoralists were willing to accept that amorfous fluids could by themselves produce any sensible order. Its locus classicus is the Hippocratical Airs, Waters, Places.

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Arbuthnot, Falconer, Montesquieu, Buffon, etc.35 This external origin of difference was often combined with discussions concerning resemblances and degenerations. What could be called the “staying power” of externally induced alterations was often a point of both empirical and theoretical discussion. Competing systems (or models) of generation had to adapt themselves to collected facts and consensus opinion around this issue. It will be of interest to notice that conceptions of hereditary transmission associated with French vitalism tended to side with a more permeable view of the link between body and environment. This open position, that saw in (complex) equilibria between internal and external elements of the body the only plausible realistic approach to health (and to natural history) allowed for the eventual conception of a Hygienic approach to both health and the physical betterment of humanity. A crucial difference between French and British hereditarianism in the course of the 19th century was due exactly to this permeability allowed by the French. As Coleman clearly showed, the notion of the non-naturals was the antecedent to the hygienic programs in the post-Napoleonic years. Elizabeth Williams has followed brilliantly the influence of vitalism throughout the 19th century approaches to the link between body and environment by French physicians.36 Humoralists on the whole were prone to what we could call a holistic view of the links between the body and the environment, and found it conceivable that a kind of a flux (or flow) of externally induced characters could travel (trough physiological means) within the family lines and conform a peculiar bodily heritage for families, tribes, races. On the other hand, solidists (à la Louis), given that no possible route of influence can be imagined that connect the parent’s initial, solid bodily frame, with that of its infant’s, and that every single case of variation and resemblance can be explained (away) using external factors, hereditary attributions become just a “façon de parler”. What I want to discuss then is the question of the permeability of the (possible) hereditary causal routes to the external factors during the 18th century. The task should be to show how the two levels of analysis, the influence of external factors on bodily constitution, and the hereditary influence of the parents bodily constitution on the bodily constitution of the child could be (and were) coordinated by different authors with different viewpoints. The specific discussion around the hereditary transmission of disease in the frame of eighteenth century views of physiology and generation provides a privileged access to the evasive conceptualization of the possible routes through which the environment could affect, in dramatic and more or less permanent fashion, the constitution of individuals and lineages. 3. It seems to me that a useful way to frame the discussion of heredity in the 18th century is to be found in Peter McLaughlin’s proposal37 to make a “distinction between the two types of inheritance – the law-like transmission of species form and the contingent disturbance or supplementation of this transmission by individual traits.” I believe however that such distinction should be made in another fashion, and more care should be taken in the choice of words. For the 35 36 37

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See relevant chapters in Glacken (1967). See Coleman (1984), Williams (1994). McLaughlin (2000).

Natural Things and Non-natural Things. The Boundaries of the Hereditary in the 18th Century

most, during the eighteenth century, what McLaughlin refers to as the first type of inheritance was considered outside the domain of the hereditary. Generally speaking, all theoretical positions about generation did not refer to inheritance when dealing with a lawful transmission of form; that is a notion that was adopted well into the 19th century. Authors in the 18th century, both preformationists and succesionists,38 thought that the transmission of form from generation to generation was due to a constant, invariable cause (or fact) that gave the species its fixity and stability, and which had nothing to do with the contingencies of genealogy. Both the idea of the preexistence of the germ, and that of a lawful (successive) production of the new being driven by a mould, or by a generative force, or some other epigenetic principle, shared the notion of a basic common structure for each species, over which the singular, accidental characters of the ancestors had no permanent influence. A shared attitude towards hereditary characteristics (the peculiarities of individual, family, or major lineages) was thus to confine them to the domain of the accidental. While accepted and discussed in both Hippocratic treatises and Aristotle’s works, the facts of hereditary transmission of physical and moral resemblance, deformation and disease, were seen both as undeniable (though bothersome) factual givens that had to be explained away. Perhaps only in the medical tradition, with its preoccupation with the singularity of disease and patients had more room for thinking about the peculiarities of individual temperaments and their transmission through the family line. For that period the distinction that MacLaughlin is aiming at should in my view be refered to as that between type vs. accident, and only the latter should be seen as linked to the hereditary. The type-accident distinction allows for a careful following of the place that the hereditary received in different theoretical frameworks. For instance, Hippocratic-Galenic medics were content with the fact that dual seminal, pangenetic, solidification of fluids, model could among other explanatory virtues, account for the facts of hereditary recurrences. At the same time, the more theoretically sophisticated model of generation devised by Aristotle was seen to generate some puzzles difficult to sort out in regards to this kind of facts.39 There is in fact an ongoing discussion among Aristotelian scholars about how to understand his proposal of explaining (away) in his scheme, the hereditary fact of resemblance to both parents, and sometimes to previous ancestors. As is well known, in Aristotle’s scheme of generation male seed was responsible for form and female matter for individual peculiarities.40 In an illuminating recent piece, Andrew Coles provides a complex physiological analysis of the origin and function of the male seed (semen) in Aristotle, so that the main difficulties that arise with hereditary facts are overcome. Contrary to widespread belief, according to Coles it is by rescuing the Cnidian (Hippocratic) notion of pangenetic origin of the semen that he manages to do the trick: “It is in Aristotle’s conception of 38 39

40

Succesionist, as I said, is a label that includes both epigenetists and the partisans af dual seminal rapid formation of the germ or first rudiment. At least since Empedocles, for anybody in the business of giving an account of human, (or animal) generation, the paradoxes of the hereditary were a serious stumbling block: Aristotle’s view of the male seed as the only causal contributor to the form (shape) of the body of the offspring had to by-pass the empirical evidence of female transmitted characteristics (resemblances, mules). The most convincing account of of the irregular mixtures of resemblances to both parents was given by dual seminal theories. The question then (as Sharples poses it) is, if it is supposed that the father imparts form and nothing else, why snub-nosed fathers tend to have snub-nosed children. See Sharples (1985).

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the physiological origins of semen, and more particularly of its hereditary properties, that the links between his biology and that of the Cnidian’s are the closest”.41 Apparently, the view that the (male) semen is the product of a special process of separation from the blood, and that in its passing through all the different parts of the father’s body, it acquires particular kinds of secondary movements (dunamis) which can be responsible for carrying on the offspring hereditary resemblances. The result from the (part by part) struggle of these paternal dunamis with similar movements that oppose them and that come in the mother’s generative matter defines the side that the offspring’s different parts are going to take after.42 Aristotle’s acute perception of the thorny causal problems that hereditary resemblances posited was progressively diluted by his interpreters, so the aspects of his system specifically designed for coping with them seem to have been lost. Galen for instance was convinced that there was no way in which Aristotle could account for female transmitted resemblances, and for that reason made a forceful argument in favor of the existence of a female seed (semen), with equivalent physiological powers to transmit hereditary peculiarities. 43 Harvey believed he was being Aristotelian when he limited the influence of material physiology to the action of male spirits in the production (conception) of form in the new being, and pushed out the hereditary influence (resemblances, etc.) to the Paracelsian domain of the action of the mother’s imagination.44 As McLaughlin points out, the conflict between pangenetic (dual seminal) models of generation (with their hereditary support) and the epistemological (and theological) demands that promoted preformationist views an the 17th century produced different attempts at “mixing” the virtues of both accounts, as Aristotle had apparently done. At the beginning of the 18th century Bourguet proposed one such synthetic model. A preformed germ affected after fecundation by a pangenetic, dual seminal influence, responsible for all things hereditary. As has been said, Haller first used the hereditary as a weapon for epigenesis, and later changed sides, denying emphatically its reality when criticizing Buffon. Bonnet eventually adopted and deepened Bourguet’s strategy separating clearly the origin of form (in the germ) from the external origins of resemblances 45 which were due to the incorporation (intussusception) of accidents in the nutritional growth-development process where (pangenetically gathered) particles from both origins could play a role. Although succesionist authors like Maupertuis and Buffon insisted that the manner of the (new) production of the germ in each fecundation is important, several authors (including McLaughlin) have argued that there is a common structure between their hypothesis and elaborate preformationism such as Bonnet’s. More than a hundred years ago, in an insightful historical essay called “Evolution in Biology”(1878), T.H. Huxley arrived at the conclusion that if we set aside around the time and 41 42 43 44 45

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Coles (1995), p. 50. See Coles (1995), pp. 70/76. Coles (1995), p. 76, Boylan (1984). See Harvey ([1651] 1981). Though in his later versions of his theory of generation Bonnet came closer and closer to a quasiepigenetical scheme, in which organised structure is not actually present in the preformed germ but somehow “preprogrammed”. See Huxley ([1878] 1896), and Bonnet ([1769] 1985).

Natural Things and Non-natural Things. The Boundaries of the Hereditary in the 18th Century

manner of the production of the germ (Harvey’s rudiment), most 18th century theorists of generation, preformationists or succesionists, supported some form of the doctrine of evolution or development, which considers a dual phase in the production of the new being’s body; first the creation or reproduction of the basic structure in the germ, and second the growth and development of this rudiment.46 Huxley sees in Bonnet’s description of such process the common “exemplar” for the period. For the case of the hen’s egg, Bonnet in his Considérations ... states that: fecundation and incubation simply cause the germ to absorb nutritious matters, which are deposited in the interstices of the elementary structures of which the miniature chick, or germ, is made up. The consequence of this intussusceptive growth is the “development” or “evolution” of the germ into the visible bird. Thus an organized individual is a composite body consisting of the original, or elementary parts and of the matters which have been associated with them by the aid of nutrition; so that if these matters could be extracted from the individual, it would, so to speak, become concentrated in a point, and would thus be restored to its primitive condition of a germ.47

Further modifications to his notion of germ made Bonnet’s position even more general and inclusive.48 Ce mot (germe) en designéra pas seulement un corps organisé reduit en petit, il désignera encore toute espèce de préformation originelle dont un tout organique peut résulter comme de son principe immédiat49

Huxley’s conlusion is eloquent: But thus defined, the germ is neither more nor less than the “particule genitalis” of Aristotle, or the “primodium vegetale” or “ovum” of Harvey; and the “evolution” of such germ would not be distinguishable from “epigenesis”.50

So even something as obtuse as Buffon’s “moule interieur” could somehow become equivalent to Bonnet’s generalized “germ”. Notwithstanding important subtleties that stand in the way of Huxley’s coarse integration of these concepts, he is no doubt making an important point. Most 18th century theorists shared the idea of a deep conceptual hiatus between the explanation of organization (taxonomic similarities), and that of accidental individual peculiarities (variations, resemblances). For our purposes, the similarity Huxley sees between the generation models of the kind Bonnet-Haller, and of the kind Buffon-Maupertuis, turns around the fact that both deliver a germ that suffers a process of growth and transformation under the influence of a fluid milieu that

46 47 48

49 50

Huxley added that even Cuvier, in following century, adopted a very similar scheme. Huxley ([1878] 1896), p. 190. Ibid., p. 191. Huxley writes: “Bonnet ... in his later writings and at length ... admits that the germ need not be the actual miniature of the organism, but that it may be merely a “original preformation” capable of producing the latter.” (Huxley [1878] 1896, p. 193. he quotes from Bonnet, 1769, X., ch. ii). Bonnet (1769), X, ch. ii. quoted by Huxley ([1878] 1896), p. 193. Huxley ([1878] 1896), p. 193.

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provides the elements that become incorporated. This milieu is responsible for hereditary influence. The origin, qualities and manner of action becomes the question for hereditary analysis. The typological, essential, permanent features of organization are identified with the solid frame, whereas accidental, ephemeral features come into play through humoral (fluid) influences. The dialectics of solid-humoral interaction have a crucial role in the split between structural stability and individual deviations. The solid organization (given for instance by the germ or driven by a solidifying force) receives constant influences, and matter from the fluid environment. These alter and shape the bodily frame. Aggregation or intussuseption of particles of different origins (paternal, maternal, external) are processes indifferent with regards to the hereditary route. Both a succesionist and a preformationist account of generation can equally explain (away) hereditary transmission through the incorporation of elements during growth or development. There is an important difference to keep in mind about the consequences of the acceptance of a preformationist account and that of a succesionist account of the origin of the germ (or the “first formation”). Although some hereditary facts can in both cases be accounted for through external supplements51 to the first formation during “evolution”, or growth, the preservation of a clear-cut split between the (causal) origin of form and the (causal) origin of resemblances or degenerations becomes more problematic for the succesionist. In a Maupertuis-Buffon kind of model, for instance, some of the more dramatic, important hereditary variations (degenerations) could up to a point be incorporated into the essential genealogical sequence. Six-digitism, or other degenerations, become susceptible of being transmitted or “copied” by the generation process 52 responsible for the first formation in a similar fashion as the more basic characteristics of the species. In fact, for some defendants of the preformation, like Haller, this vagueness about the limits of the individual (resemblances) and the formal (type) became one of the main reasons for their opposition to succesionist schemes. At the same time the failure of preformation to make sense (by itself) of preexistence within genealogical lines of accidental features made the subsequent appeal to a supplementary dual pangenetic influence methodologically suspicious. 53 Again, the permeability or impermeability of the specific form to external, accidental influences is one of the issues at stake within these discussions. Though it must be remembered that hereditary facts could, during the 18th century, still have alternative non-physiological explanations (v.gr. a constancy or repetition of some external climatic influences or an appeal to the action of imagination or other “mind over matter” interventions) the search for a regular, stable physiological source was increasingly seen as the only sensible explanatory strategy. For such a strategy one important question turned around the origin, kind, and qualities of the material particles that were incorporated (through generation, nutrition, development and growth) into the different parts of the body. The notion of a strict ontological boundary between organic and inorganic particles, as proposed for instance by 51 52

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Dual seminal, plus maternal blood, plus other sources of nourishment. For example through the modified moule interieur, or Maupertuis’ material memory. There seems to be evidence both for a possibility of modification of the moule interieur, and for its inalterability in Buffon’s writings. See Aréchiga (1996). In his 1878 work Huxley concludes that it was only with the careful observation of development of the chick by C.F. Wolff that the speculative impasse ended and embriology could take a progressive route.

Natural Things and Non-natural Things. The Boundaries of the Hereditary in the 18th Century

Buffon, provided another boundary to take into account. The access of external influences to the peculiar bodily features of each individual could clearly be limited by such considerations. Climatic, nutritional and other sources of bodily variation, captured eloquently by the label of the non-naturals can have very different capacities to alter the constitution according to the continuity or discontinuity postulated between bodily constituents and external matter (air, water, places, etc.). The medical frame of natural and non-natural, and of solid vs. humoral influences was the main available resource for coping with these questions. It was undoubtedly at the root of Buffon’s speculations concerning the influence of climate and food in the production of degenerations (degenerations), which begin as individual variations and are progressively generalized within a lineage as they are hereditarily transmitted (“like diseases are communicated from fathers or mothers to the children”).54 A further question, not often formulated clearly is the “staying power” (which makes them more or less permanent or ephemeral) of the peculiarities incorporated by environmental and nutritional routes, within the genealogical succesions. This question can become crucial in a succesionist account, if there is a dilution of the difference between individual and specific organization.55 The question presented itself to Maupertuis and Buffon, and also to Blumenbach. They all allowed for a semi–permanent transformation of the type due to the conservation of accidental variation within the lineage.56 It is probably in the extensive discussions of Buffon on the notion of degeneration that the intricacies of this issue began to be sorted out. The belief in a strict and widespread correlation between climate and bodily temperament in human groups was heavily questioned in the course of the 18th century. The tenacity or the feebleness (i.e. differences in “staying power”) of accidental racial features had both evidence and testimonies in their favor.57The issue of establishing how and when external influence on the physical characteristics became “rooted”58 in a lineage acquired a progressive importance after Buffon’s work. 59 This French author always maintained that a limit existed to the amount and kind of variation accepted by the interior mould.60 Kant for instance believed in the feebleness of accidental acquisitions. He writes: Gradually and at last the constitution of the soil (moisture or drought), and food, also, induce a hereditary difference or strain among animals of one and the same stock and race, especially in stature, proportion of limbs, and also in the temperament; which later hybridizes when mixed with another kind: but on another soil and in the presence of other food (even without alteration of the climate) disappears but in a few generations. 61 54 55

56 57

58 59 60

See Buffon ([1749] 1971). See also Aréchiga (1996), p. 74. The process of using the complete bodily constitution or temperament of a parent as an original pattern or mould for the production of the new being (at the moment its first formation) has even induced a paradoxical talk of the conservation of an individual type(!): See Duchesneau (1982), p. 539, and LópezBeltrán (1992), chapter V, and Borie (1985). John Hunter arrived at a similar view, influenced by Blumenbach. See López-Beltrán (1992), chapter IV. “such is the difference of this effects – writes Blumenbach – (some) are preserved unimpaired by a sort of tenacious constancy through long series of generations, or by some power of change withdraw themselves again in a short space of time”. (Blumenbach [1795] 1865). He metaphor of depth, of rootedness, taken I believe from iatrochemical speculations of the seed or material cause of disease, was frequently invoqued in this context. See Roger (1989), Aréchiga (1996), Glacken (1967). Blumenbach himself wavered in front of the evidence. Compare Blumenbach ([1775 and 1795] 1865). See also López-Beltrán (1992), chapter IV.

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Blumenbach wavered in front of the evidence. After first having also dismissed the view that what Buffon called degenerations could become a stable hereditary part of an animal’s (or a person’s) constitution, he came to accept what he called “hereditary peculiarities of animals from diseased temperaments”: An hereditary disposition to disease would seem at first sight rather to belong to pathology than to natural history of animals. But when the matter is more carefully looked into, it is plain that in more ways than one it has something to do with those causes of degeneration we are concerned with.62

Blumenbach later writes that when some of these (constitutional diseases or disorders)... are propagated by hereditary causes for a long series of generations it shades sensibly away into a sort of second nature and in some species of animals gives rise to peculiar and constant varieties.63

4. Blumenbach’s analogy between hereditary disease and hereditary variation, and his blurring of the distinction is to my mind revealing, in that within medical tradition (as I said) there existed a frame that allowed a way of dealing with the issues of the link between the body (temperament, constitution) and the environment. A complex, and mutually dependent set of causal factors could be considered as working over the same product (the body) in such a way that the final result could not be attributed wholly to one or another particular influence. The notions of crasis (mixture), fluidity, individuality, allowed the imagination to both conceive a given outcome as determined and caused, and as complex product of an un-analyzable and variable set. On the other hand the distinction between the natural things (bodily frame, temperament) and the nonnatural things (that affect and change the former) allowed for a separation of causal spaces, that is however very different from the modern nature-nurture split we have become habituated to. The fact for instance that passions, dreams, and other psychological elements were placed on the same grounds as the air, or food, or water, reveals a very different conception of the boundaries, and of the causal dependencies that are implied. Organisms (bodies) were not seen diachronically as we see them, that is as being in a dialectical sequence of expression of a hereditary information within succesive environments, but were rather seen as embedded in, open to and intimately dependent on its physical surroundings. The complex web ob influences that acted over each individual (human) body was a main preoccupation of the medical tradition, and the physical and moral inheritance that was passed on from parents to offspring through generation was only a small portion of such web. The natural things (humors, elements, temperament) can be equaled for the sake of our story with the bodily elements that are put in place by typical generation. That is to say, those which constituted the germ, and/or the preordered set of elements that get together to form the

61 62 63

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Kant (1775), in Chukwudi (1997). Blumenbach ([1795] 1865), p. 202. Ibid., p. 259.

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organized new individual are product of the natural things. The non-naturals (air, food, dreams, passions, etc.) are the set of “external” influences that somehow come into contact with, and/or determinantly, affect the body as it grows, lives and decays. From a superficial analysis the two sets of causes are disconnetced. They were treated for instance in two different Hippocratic treatises.64 But of course the link between them is powerful, and as Arnulfe D’Aumont writes, it is when these interactions (natural/non-natural) goes wrong that the praeter-natural (diseases) appear.65 As we saw in Antoine Louis’ argument above, the six non-naturals can readily play a part in the discussion concerning external causes of degeneration, and the “staying power” they acquire according to their origin and moment of influence. 66 The possibility that external influences could become more or less permanent through generation within a genealogical line was seen as more or less open by differently oriented physicians. A privileged access to how these matters were dealt with, and into the details of physiology and etiology, during the18th century is gained when we look at discussions of hereditary disease. 67 Through them we can clarify the different consequences for the phenomenology of the hereditary that medical men could see if they were to accept certain openness (permeability) or closure to external, accidental influences at the moment of the first formation. According to different physiological views, different boundaries could be drawn. The question can again be posed: are accidental hereditary bodily features induced by the six non-naturals(?) and are these transformations of such an order that the family lineage is affected in a permanent way? I will attempt in the following paragraphs a short summary of several positions that can be found among physicians during the second half of the 18th century. A strict solidist position can readily drive to an anti-hereditary stance. We have seen that with Louis.68 The impossibility of the incorporation (at the required depth) of anything external or accidental into the body’s frame (i.e. the original germ). This is Louis’ main argument. If diseases have a solidist root, any lesion would have to be local, isolated, and its effects would end with the death of the bearer. In contrast a humoralist view incorporates the hereditary to a choreography of internal and external influences. Within them there is no clear-cut separation between external and internal humors. The hereditary cause does not have a special ontological status. It is more a consequence (a “side effect”) of the flux of succesive generations irregularly acquiring (not only through generation) physical (and moral) peculiarities and transmitting them (through generation) to their offspring. At the most, the adjective “hereditary” refers to a possible route of transmission of influences that may also operate through other routes. An important theoretical development occurs among medical men when committed solidists, uncomfortable with humoral, proteic explanations of hereditary disease, nevertheless accept the 64 65 66

67 68

“Airs, Waters, Places” and “The Nature of Man”. D’Aumont (1765). The division we described between humoralists and solidist becomes relevant. In an humoralist view the balance among humors is crucial, and there is a continuous flow of materials, mixtures, in and out of the body, carried by air and liquids and food. The solid parts of the body are always changing according to these interactions. In a solidist view the root of the disease is in a lesion or malformation at the level of the organs or solid parts. It is the solids and their properties who master a control the humors. I have made a thorough study of the disputes around this issue that happened in France between 1745 and 1810. See López-Beltrán (1996). Louis himself locates the thrust of his antihereditary argument in the older work of Luis Mercado, where apparently all solid to solid influence is also denied. See Louis (1749), footnote, p. 14.

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reality of the fact of hereditary transmission, on the base of the accumulated evidence provided by the accumulation of cases were features, traits and diseases were seen to travel through lineages. A new kind of speculation about the possibility of solid to solid hereditary influence is then developed, and as a consequence a distinction between properly hereditary transmission and parallel congenital influences is proposed.69. We could also summarize the different “models” of hereditary disease transmission according to which notion of generation is presupposed by physicians. Under preformation: the germ is invaded by the “seed” of disease,70 and its humoral balance is disturbed. Or it is damaged by an external influence (v.gr. a fluid) during its development and the alteration can then be somehow transmitted to the following generation through parental semen (pangenetically for instance). The transmission then is not necessary and is avoidable. In a clear-cut succesionist account something analogous can happen. After the “first formation”, heritable alterations can be suffered. Humors are responsible for idiosyncracies. There is not an important etiological distinction between receiving the evil action for the first time in the lineage (by an environmental cause) or receiving it through the liquids of the parents during growth and development, or even during nursing. Towards the end of the 18th century a very different notion of hereditary transmission is adumbrated by solidists that is only possible in a succesionist scheme. A form or other of a copying mechanism is proposed as responsible for producing, each generation, the solid parts of the new being.71 The latter can be framed (as for Buffon and Maupertuis) under the analogy of crystallization, or not. The (solid) bodies of the parents are taken to be the source (or mould) that is used as a base for the new production of organization in the offspring. Both essential and nonessential features can be thus candidates for copying. The hereditary act (transmission) occurs always at the moment of the “first formation” of solid parts (the production of the germ). Bodily variations due to external influence can become hereditary only if they originate at the moment of the “first formation”. Besides, not every feature is transmitted identically. There is rather a different tendency among them to be transmitted according to their properties. Essential features are transmitted with more constancy, and accidental features have less chance to be copied. Similarities have the way more open than degenerations. The older an accidental feature has been running in the family the stronger will its propensity to be communicated due to a sort of rootedness it has developed. What we have been calling the “staying power” of a variation is thus linked to its propensity to be carried along to posterior generations by the “copying” mechanism. Variation within the offspring is thus explained. The bodily organization of the new being is a version of the parent’s organization, in which both essential and accidental features are copied from them. Even deformities can be transmitted if their origin is an accident at the moment of the first formation. What can be called a “frozen accident” explanation of hereditary transmission of accidental variation due to external causes. This kind of thinking can be traced to the model of Maupertuis’ generation theory and was used by John Hunter and his followers. The clear-cut distinction it aims at establishing between hereditary and non-hereditary transmission within lineages was used by French medics, and also by Hunter and his followers, to establish a difference 69 70 71

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For a detailed discussion of this conceptual innovation, and its consequences in both French and British discussions of the hereditary see López-Beltrán (1992), idem. (1996). In a iatrochemical wording. See Pujol (1802), Prichard (1813) and idem. (1824).

Natural Things and Non-natural Things. The Boundaries of the Hereditary in the 18th Century

between congenital and connate transmission of disease.72 What is often discussed is, I repeat, if the depth of the incorporation of the hereditary variations into the normal flux driven by the reproduction mechanism. Usually, among medics, the question attended to is how, if in any way, can hereditary disease be eliminated. The answer to such question reflects the view held by the physician.73 5. (Conclusions) We can talk of the existence during the 18th century of two kinds of limits or boundaries to hereditary transmission. The “channelization” of hereditary influences through the physiological “line of production” of the new beings can be seen, during that period as being limited by an internal and an external boundary. The internal boundary is given by the separation between the re-production (generation after generation) of the specific form, and the acquisition and transmission of peculiar accidents of constitution first by individuals and later by lineages. Both preformationist and epigenist views of generation accepted the challenge of explaining (away) the facts of hereditary links between parents and offspring. And in both their positions a tight boundary was placed in order to preserve the integrity and stability of the form. The accidents could not be incorporated definitively into the essential organizational features or form. Each new being when first constituted as a germ was free from accidental variations produced by external influences, including hereditary ones. The external boundary is given by the physiological facts of growth, development, nutrition. The amount of isolation they were thought to have with respect to the physical milieu made the outcome of growth and development (the mature individual’s temperament) more or less variable, and the depth of the changes could also vary. The belief that only those modifications of temperament by the non-naturals that could find their way to the flux of pangenetic chain of transmission was shared by many. The mystery of course was how this could actually take place. What exactly had to be modified by external influences? The mould or the particles? The bodily features of the parent and the particles that integrated them. Buffon, Blumenbach and other 18th century thinkers conceived mechanisms by which physical modifications induced by the environment could be preserved hereditarily and give rise to varieties within species. The reversibility by external influences was always an option, as was the idea of betterment through hygienic measures.74 The vitalist program in France preserved this attitude throughout the 19th century, allowing the notion of hereditary determination of physical and moral characters to be balanced by a hygienist counterpart. Though the notion of hereditary transmission was restricted and had a minor importance in the 18th century. The breaching of what I just called of the internal boundary is responsible for the hardening of heredity, and its progressive transformation during the 19th century into a central cause in physiological thought. The permanence and transcendence of “accidental” variations needed a new conceptualization of the hereditary. Towards the turn of the century (18th to 19th), as can be seen in Blumenbach’s changing views, and more prominently in 72 73 74

See Pagès (1798), Pujol (1802), and Adams (1814). See López-Beltrán (1996). Changing the climate, the food, the way of life.

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Lamarck’s use of hereditary (and in a sense “accidental”) modifications as the source of species transformation, the narrow boundaries between which hereditary transmission could be conceived to occur, became progressively breached, and blurred. The outcome of this opening was that external climatic and psychological causes could be now seen to affect and modify in the long run the essential form of organisms, and of their descendants. Deeply embedded in the organism’s constitution, some degenerations with considerable staying power could be conceived as witnesses to the power of both environment and heredity to shape bodies (and minds). Resemblances, degenerations and the transmission of form through generation could be now seen as having similar causal dependencies.

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References Adams, Joseph. 1814. A Treatise on the Hereditary Properties of Diseases. London: J. Callow. (a facsimile reproduction of this work was published in Charles Rosenberg’s series History of Hereditarian Thought, 1984, Garland Publishing). The second edition of this work, in 1815, with only a few additions as appendixes received a much more ambitious title: A philosophical dissertation on the hereditary peculiarities of the human constitution. A German translation appeared in the Neu Sammlung auserlesener Abhandlungen (tome II, p. 503). Amoreux, Pierre-Joseph. 1790. “Des Maladies Héréditaires”. handwritten essay at the archives from the Societé Royale de Paris (10; 120-3-1). This piece was given the accesit. An outstanding, modified and extended version of the 1788 document. Amoreux shows abundantly the historical depth of the theme, and displays an impressive knowledge of the bibliography. Several of the comments on 17th and 18th century texts above are taken from this important manuscript. This text seems to have been close to winning the contest, and the judges wrote of it that: “L’Auteur de ce Mémoire a développé une érudition très-étendue. La Société auroit desiré de trouver plus de détails dans les traitemens prophylactique & curatif.(Hist. Soc. Roy.,vol.9,p. xi) Anderson, James. 1799. “An inquiry into the nature of that department of Natural History which is called VARIETIES”. Recreations in Agriculture, Natural-History, Arts, and Miscellaneous Literature 1:49–100. (6 vols. 1799-1806) London: T. Bensley. Arbuthnot. 1733. Essay Concerning the Effect of Air on Human Bodies. London. Aréchiga, Violeta. 1996. El concepto de Degeneración en Buffon. tesis de maestría. UNAM-Iztapalapa. Bénichou, Claude, ed. 1989. L’ordre des caractères -Aspects de l’hérédité dans l’histoire des sciences de l’homme. Paris: Vrin. Bernardi, Walter. 1986. Le Metafisiche dell’Embrione; Scienze della vita e filosofia da Malpighi a Spallanzani (1672-1793). Firenze: Leo S. Olschki. Bichat, M. F. X. 1800 (an VIII). Recherches Physiologiques sur la Vie et la Mortt. Paris: Brosson & Gabon. Blumenbach, J.F. [1775 and 1795] 1865. The Anthropological Treatises. London: Anthropological Society. (translations by T. Bendysche of his works on On the Natural Variety of Mankind, of 1775 and 1795). Böhme, Gernot and Hartmut Böhme. 1998. Fuego, Agua, Tierra, Aire, una historia cultural de los elementos. Madrid: Herder (original de 1996). Bonnet, Charles. [1779] 1985. Considérations sur les Corps Organisés. vol. 3 of Oeuvres d’Histoire Naturelle et de Philosophie, 8 vols., Neuchatel. Hay una reproducción (1985) de la edición de 1778 en la colección Corpus des Oeuvres de Philosophie an Langue Francaise, de Fayard. Bonnet, Charles. 1769. La paligénésie philosophique. Geneva. Borie, Jean. 1981. Les Mythologies de l’Hérédité au XIXe siècle. Paris: Editions Galilee. Bowler, P. J. 1971. “Preformation and Pre-existence in the 17th Century”. J. Hist. Biol. 4(2):221–244. ———.1973. “Bonnet & Buffon -theories of generation and the problem of Species”. J. Hist. Biol. 6:259– 281. Bowler, Peter. 1989. The Mendelian Revolution. London: Athlone Press. Boylan, Michael. 1984. “The Galenic and Hippocratic Challenges to Aristotle’s Conception Theory”. Jour. Hist. Biol. 17:83–112. ———. 1986. “Galen’s Conception Theory”. Jour. Hist. Biol. 19:47–77. Buffon. [1749] 1971. “Histoire Naturelle de l’Homme”. In De l’Homme. Paris. Burns, Chester R. 1976 “The Nonnaturals : A Paradox in the Western Concept of Health”.The Journal of Medicine and Philosophy 1(3):202–211. Bynum W.F. and Roy Porter. 1993. Companion Encyclopedia of the History of Medicine. London: Routledge. Cabanis, P.J.G. [1802] 1956. (an X). Rapports du Physique et du moral de l’Homme. In Oeuvres Philosophiques de Cabanis, I:126, edited by Claude Lehec and Jean Cazeneuve. 2 vols. Paris: Presses universitaires de France. (in English: On the Relations between the Physical and Moral Aspects of Men. 1981. John Hopkins U. P. , 2 vols.). Canguilhem, Georges, G.Lapassade, J.P. Piquemal and J. Ulmann. 1962. Du Devéloppement a l’Évolution au XIXe siecle. Paris: Presses Universitaires de France. (1985, 2a. ed). Canguilhem, George. 1977. La formation du concept de reflex aux XVIIe et XVIII siècle. Paris: Vrin. ———. 1983. Etudes d’Histoire et de Philosophie des Sciences. Paris: Vrin. Chambers Dictionary -Cyclopedia or an Universal Dictionary of Arts & Sciences. 1738. 2 vols., edited by E. Chambers. 2nd. edition. London.

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Chukwudi Eze, Emmanuel, ed. 1997. Race and the Enlightenment : a Reader. Cambridge, Massachusetts: Blackwell. Churchill, Frederick.1987. “From Heredity to Vererbung, The transmission problem 1850-1915”. Isis 78:337–364. Coleman, William. 1965. “Cell, Nucleus and Inheritance -a historical study”. Proc. Amer. Phil. Soc. 109(3):124–158. ———. 1974. “Health and Higiene in the Encyclopédie: A Medical Doctrine for the Bourgeosie”. Journal of the History of Medicine 29:399–421. ———. 1984. “Inventig Demography, Montyon on hygiene and the state”. In Transformation and Tradition in the Sciences, edited by E. Mendelsohn. Cambridge: Cambridge University Press. Coles, Andrew. 1995. “Biomedical models of Reproduction in the Fifth Century BC and Aristotle’s Generation of animals”. Phronesis XL(1). Corsi, Pietro. 1988. The Age of Lamarck -Evolutionay Theories in France 1790-1830. traducción de la edición edición italiana (1983) by J. Mandelbaum. Univerity of California Press D’Aumont, Arnulfe. 1765. “Non-naturelles, choses”. In L’Encyclopédie, edited by Diderot and D’Alambert, vol. XI, 217–224. Diderot, Denis and Jean D’Alembert. 1750–1775. L’Encyclopédie, ou Dictionnaire des Sciences, Arts et Métiers. ———. [1876] 1964. Élemens de Physiologie. edition critique by J. Mayer. Paris: Marcel Didié. Duchesneau, Francois. 1982. La physiologie des Lumières; empirisme, modèles, théories. La Haye: Martin Nijhoff. Fourke, Daniel C. 1989. “Mechanical and “organical” models in seventeenth-century explanations of biological reproduction”. Science in Context 3(2):365–381. Fox, Christopher, Roy Porter and Robert Wokler, eds. 1995. Inventing Human Science, 18th-century domain. Los Angeles: University of California Press. Galton, Francis. 1869. Hereditary Genius. London: Macmillan. ———. 1872. “on Blood-Relashionship”. Proceedings of the Royal Society. 20:394–402. ———. 1889. Natural Inheritance. London: Macmillan. Gayon, Jean. 1998. Darwinism’s struggle for survival : heredity and the hypothesis of natural selection. Cambridge, New York: Cambridge University Press. (traducción del francés Darwin et l’Aprés Darwin , Paris: Kimé, 1994, de Matthew Cobb). Ghiselin, Michael T. 1975. “The Rationale of Pangenesis”. Genetics 79:47–57. Glacken, Clarence J. 1967. Traces on the Rhodian shore : nature and culture in Western thought from ancient times to the end of the eighteenth century. Berkeley: University of California Press. Guyenot, Emile. 1957. Les Sciences de la Vie au XVII et XVIII siecles. Paris: Albin Michel. Haller, Albert v. 1752. Reflexions sur le Système de la Génération de M.de Buffon. Geneva: Barrilot et Fils. (translation from the preface to the 2nd. volume of the 1st. German edition of Buffon’s Oeuvre), Paris. An English translation by P. S. Sloan, “Reflections on the Theory of Generation of Mr. Buffon”, in Lyon & Sloan (eds.) 1981. ———. 1772. La génération ou exposition des Phénomenes relatives a cettte function naturelle, 2 tomes, Paris: Chez de la Dove. Hannaway, Caroline. 1993. “Environment and Miasmata”. In Companion Encyclopedia of the History of Medicine. edited by W. F. Bynum and R. Porter. London: Routhledge. pp.292–307. Harvey, William. [1651| 1981. Disputations touching The Generation of Animals. Oxford: Blackwell Scientific Publications. Héritier-Augé, Françoise. 1985. “Le Sperme el le sang, de quelques théories anciennes sur leur genèse et leurs rapports”. Nouvelle Revue de Psychanalyse 32:111–122. (English version: “Semen and Blood: Some ancient theories concerning their genesis and relationship” in Zone : Fragments for a History of the Human Body . 3, edited by M. Feher (1989). New York. pp. 159– 175). Hilts, Victor. 1984. “Enlightenment views on the genetic perfectibility of man”. In Transformation and tradition in the Sciences, edited by E. Mendelsohn. Cambridge U.K.: Cambridge University Press. Hippocrates. 1923. “Airs, Waters, Places”, “The Nature of Man”. In Works. London: Heinemann, Loeb Classical Library. Hoffheimer, M. H. 1982. “Maupertuis and the 18th-Century Critique of Preexistence”. J. Hist. Biol. 15(1):119–144. Hunter, John. 1786. Observations on certain parts of the animal oeconomy. London: Castle Street. ———. 1861. Essays and Observations on Natural History , edited by R. Owen, 2 vols. London. Huxley, Thomas H. [1878] 1896. “Evolution in Biology”. In Darwiniana; Essays. New York: Appleton and Co.

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Jacob, François. 1970. La logique du vivant, une histoire de l’hérédité. Paris: Gallimard. Jarcho, Saul. 1970. “Galen’s Six non-Naturals”. Bulletin of the History of Medicine 44:372–377. Kant, Immanuel. [1775] 1997. “On the Different Races of Man”. In Chukwudi. 1997. pp.38–48. Lloyd, G.E.R. 1983. Science, Folklore and Ideology, Studies in the life Sciences of Ancient Greece. Cambridge: Cambridge University Press. Lindeboom, G. A. 1970. “Boerhaave’s concept of the basic structure of the body”. Clio Medica 5: 203–208. López-Beltrán, Carlos. 1992. Human Heredity 1750-1870, The Construction of a Domain. PhD Thesis. London: King’s College. ———. 1994. “Forging Heredity, from metaphor to cause: a reification story”. Studies in the History and Philosophy of Science 25. ———. 1995. “«Les maladies héréditaires» 18th-century disputes in France”. Revue d’Histoire des Sciences XLVIII(3):307–350. ———. 1997. “ Perfectionner le Corps: Des défauts héréditaires à l’hérédité fatale”, trabajo presentado en la reunión l’Eugenisme, 1945 et aprés, Universidad de Dijon, Francia, Septiembre 1997. Aparecerá en Jean Gayon y Daniel Jacobi (eds.). in press. ———. 2000. “Por una crítica de la noción de Raza”, en Revista Ciencias, UNAM, no. 60-61, Enero. Lonie, Ian M. 1981. The Hippocratic Treatises (translation and commentary of “On Generation”, “On the Nature of the Child” and “Diseases IV”). Berlin, New York: Walter Gruyter. Louis, Antoine. 1749. Dissertation envoyée á l’Academie des Sciences de Dijon, pour le prix de l’année 1748, sur la question ...Comment se fait la transmission des maladies héréditaires? Paris, Delaguete, 12o. Lyon, John and P. R. Sloan, eds. 1981. From Natural History to the History of Nature. Readings from Buffon and his critics. Notre Dame: University of Notre Dame Press. Maupertuis, P.L.M. 1744. Dissertation Physique a l’ocassion du Nègre Blanche. Paris: Leyde. ———. 1745. Vénus Physique. La Haye. McLaughlin, Peter. 2000. “Preliminary Notes on biological notions of Heredity in the Enlightenment”. presented in the meeting on Cultural History of Heredity, Max Planck Institute for the History of Science, May, 2000. Miller, Genevieve. 1962. “ “Airs, Waters and Places” in History”. Journal of the History of Medicine 17:129– 140. Mazzolini, Renato G. and Shirley A. Roe. 1986. Science against the unbelievers : the correspondence of Bonnet and Needham, 1760-1780. Oxford: Voltaire Foundation at the Taylor Institution. (Studies on Voltaire and the eighteenth century 243). Morsink, J. 1979. “Was Aristotle’s Biology Sexist?”. Jour. Hist. Biol. 12:83–112. Niebyl P.H. 1971. “The Non-Naturals”. Bull. Hist. Med. 45:486–492. Nutton, Vivian. 1993. “Humoralism”. In Companion Encyclopedia of the History of Medicine, edited by W. F. Bynum and R. Porter. London: Routhledge. (vol. 1, pp. 281–290). Odom, Herbert. 1970. Groundwork for Darwin, Theories of Heredity and Variation, Great Britain 1790-1820. PhD Thesis. Cambridge Mass: Harvard. Pagès, Jean-François. 1798. “Héréditaires (maladies)”. In Dictionnaire de Médecine, part of the Encyclopedie Methodique, vol. VII, an VI de la République, chez Agasse. pp. 160–176. This entry is essentially the piece that was honoured in 1790 by the Royal Society of Medicine. It is an excellent defense of solid to solid hereditary transmission, and as a Dictionary piece must have been widely read, preceding Petit’s essay (1817) in breaking the ground for a broader view of heredity. It was chosen for the Dictionary by Vicq D’Azyr among the many very competent dissertations for the competition. Portal, Antoine. 1808. Considerations sur la nature et le traitement de quelques maladies héréditaires, ou de famille. Mem. Inst. Nat. de France, T.8, Semestre 2, p. 156. (Read at the Institute the 25 January 1808). Later published as book in 1808. Paris, 4o. And in a revised and extended version in 1814. 3eme edition, Paris. Extracts of this work were also published in Graperon, Bull. des Sci. Méd., Tome 2, p. 328. Almost certainly this work prompted Joseph Adams’ famous treatise on the subject, in which he tried to claim for John Hunter, and himself, the “modernization” of the concept of hereditary transmission of disease (see below). The English (edited) version of the piece was published in two installments in Adam’s London Medical and Physical Journal, volume 21, in Dec. 1808 and June 1809, pp. 229-239, and 281-296. Porter, Roy. 1995. “Medical Science and Human Science in the Enlightenment”. In Inventing Human Science, edited by C. Fox, R. Porter and R. Wolkler. University of California Press. pp.53–87. Preus, A. 1975. Science and Philosophy in Aristotle’s Biological Works. New York: Olms. Prichard, James Cowles. 1813. Researches into the Physical History of Man, 1st. edition. London: J & A Arch.

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(Based on his 1808 thesis, De generis humani varietate, Abernethy & Walker, Edinburgh; a facsimile edition was edited by G. W. Stocking, Chicago University Press, 1973). ———. 1826. Researches into the Physical History of Mankind, 2nd. edition, revised and augmented, 2 vols. London: J & A Arch. ———. 1833. “Temperament”. Cyclopedia of Practical Medicine 4:159–174. Pujol, Alexis. 1802. “Essai sur les Maladies Héréditaires”. In Oeuvres Médicales, 4 vol., Castres. (Second edition edited by F. G. Boisseau, with notes, a biography and additions, as Oeuvres de Médecine Practique, 2 vols., Paris, Bailliere & Bechet, 1823. This essay was written for the Royal Society of Medicine competition in 1788-90). Rádl, Emanuel. 1930. The History of Biological Theories, translated from the 1909 German text by E. J. Hatfield. Oxford University Press (specially chapter XXII: Human Heredity). Spanish translation by Revista de Occidente, Madrid, 1931. Rather, L-J. 1968. “The six non-naturals: a note on the origins and fate of a doctrine and phrase”. Clio Med. 3:337–347. Rey, Roselyne. 1989. “Génération et Hérédité au 18e siècle”. In L’ordre des caractères, edited by C. Bénichou. Paris: Vrin. (pp. 7–48). ———. 2000. Naissance et développement du vitalisme en France de la deuxieme moitié du 18e siecle a la fin du Pemier Empire. Oxford: Voltaire Foundation. Richards, Robert J. 1992. The Meaning of Evolution, The Morphological Construction and Ideological Reconstruction of Darwin’s theory. Chicago: Chicago: The University of Chicago Press. Ritvo, Harriet. 1997. The Platypus and the Mermaid, and other figments of the Classifying Imagination. Boston: Harvard University Press. Robinson, Gloria. 1979. A Prelude to Genetics, Theories of a material substance of Heredity: Darwin to Weismann, Lawrence. Kansas: Coronado Press. Roe, Shirley. 1981. Matter, Life and Generation, 18th century embriology and the Haller-Wolff debate. Cambridge U.K.: Cambridge University Press. ———., ed. 1981. The Natural Philosophy of Albert von Haller. New York: Arno Press. Roger. Jacques.1963. Les Sciences de la Vie dans la Pesée Française du XVIIIe siècle. Paris: Armand Colin. ———. 1989. Buffon, un Philosophe au Jardin de Roi. Paris: Fayard. Russell, Nicholas. 1986. Like Engend’ring Like, Heredity and Animal Breeding in Early Modern England. Cambridge U.K.: Cambridge University Press. Schiller, J. 1974. “Queries, Answers and unsolved problems in 18th Century Biology”. History of Science 12:191–192. ———. 1978. La notion d’Organistion dans l’Histoire de la Biologie. Paris: Maloin. Schopfer, W.H. 1946. “L’Histoire des theories relatives a la generation aux 18eme et 19eme siecles”. Gesnerus. Sharples, R.W. 1985. “Species, Form and Inheritance: Aristotle and after”. In Aristotle on Nature and Living Things, edited by A. Gotthelf. Bristol: Bristol Classical Press. (pp.117–128). Staum, Martin S. 1980. Cabanis -Enlightenment and Medical Philosophy in the French Revolution. Princeton University Press. Williams, Elizabeth Ann. 1994. The physical and the moral : Anthropology, physiology, and philosophical medicine in France, 1750-1850. Cambridge U.K.: Cambridge University Press. Zirkle, Conway. 1946. “The early history of the idea of of the Inheritance of Acquired characters and of Pangenesis”.Trans. Amer. Phil. Soc. 38:91–151. ———. 1951. “The Knowledge of Heredity before 1900”. In Genetics in the 2oth centuy. Essays on the Progress of Genetics during its first 50 years, edited by L.C. Dunn. New York: Macmillan Co.

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List of Authors

Dr. Staffan Müller-Wille Max Planck Institute for the History of Science Wilhemstraße 44 10117 Berlin, Germany [email protected]

Prof. Mary Terrall Harvard University Department of the History of Science Science Center 235 Cambridge, MA 02138, USA [email protected]

Dr. Marc J. Ratcliff Université de Genève Faculté des Sciences - Faculté des Lettres Historie et Philosophie des Sciences 10, avenue Jules Crosnier CH-1206 Genève [email protected] Prof. Carlos López Beltran Instituto de Investigaciones Filosóficas Universidad Nacional Autónoma de México Ciudad Universitaria Coyoacan D.F. 04500, México [email protected]

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A Cultural History of Heredity II: 18th and 19th Centuries

Table of Contents

Introduction Hans-Jörg Rheinberger & Staffan Müller-Wille

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Heredity old and new; French physicians and l’hérédité naturelle in early 19th century. Carlos López-Beltrán

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The Sheep Breeders’ View of Heredity (1723-1843) Roger J. Wood

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Characters written with invisible ink. Elements of Hybridism 1751-1875 Staffan Müller-Wille

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Comments on the papers given by Roger Wood and Staffan Müller-Wille Raphael Falk

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Acquired character: the (pre-genetic) material of the ‘self-made man’ Paul White

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Kant on Heredity and Adaptation Peter McLaughlin

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Inheritance of Acquired Characters in Lamarck’s and Geoffroy Saint Hilaire’s Zoology Wolfgang Lefèvre

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Erasmus Darwin on Hereditary Disease: Conceptualizing Heredity in Enlightenment English Medical Writings Philip K. Wilson

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Pathological Heredity as a Bid for Greater Recognition of Medical Authority in France, 1800-1830 Laure Cartron

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Poor Old Ancestors: The Popularity of Medical Hereditarianism, 1770-1870 John C. Waller

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Comments on the papers given by Phillip Wilson, John C. Waller, and Laure Cartron Gianna Pomata

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Heredity, Milieu and Sin: the works of Bénédict Augustin Morel (1809-1873) Jean-Christophe Coffin

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George Combe’s law of hereditary descent John van Wyhe

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Majorat: Literature and the Law of Succession in the 19th Century Ulrike Vedder

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“Victor, l’enfant de la forêt” – Experiments on the Heredity of Human Nature in Savage Children Nicolas Pethes

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Program

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Introduction

The contributions to this volume were prepared for the second of a series of workshops dedicated to the cultural history of heredity. Concentrating in turn on a succession of time periods in chronological order, this series attempts to uncover and relate to each other the agricultural, technical, juridical, medical, and scientific practices in which knowledge of inheritance was materially anchored and in which it gradually revealed its effects. While the first workshop concentrated on the late seventeenth and eighteenth centuries, the second dealt with a time period demarcated by two classical publications: Immanuel Kant’s Von den verschiedenen Rassen der Menschen (1775) and Charles Darwin’s The Variation of Plants and Animals under Domestication (1868).1 One of the major results of the first workshop was corroborated by the contributions to the second, namely that no general concept of heredity was underlying the discourse on life (including medicine, anthropology and the moral sciences) in the eighteenth century and that such a concept was only slowly emerging in the first half of the nineteenth century. 2 Carlos López Beltrán illustrates this in his contribution to this volume by directing our attention to a decisive linguistic shift: while the use of the adjective hereditary can be dated back to antiquity in the context of nosography (maladies héréditaires), a transition to a nominal use (hérédité) took place only from the 1830s onwards, first among French physiologists, then in other European scientific communities. This shift indicates a reification of the concept, or, in López Beltrán’s words, the establishment of a “structured set of meanings that outlined and unified an emerging biological conceptual space […] produc[ing] the first appearance of our modern concept of biological heredity.” For the fields of natural history and breeding one can recognise a similar shift from the use of adjectives like ‘constant’ and ‘true’ to refer to characters that remain unchanged in the course of generations, to the recognition of ‘heredity’ or ‘inheritance’ as one of the central life forces.3 Alongside this development, one can observe the erosion of a set of very ancient distinctions in regard to observed similarities between parents and offspring: the distinction of specific vs. individual, paternal vs. maternal, normal vs. pathological similarities all gave way gradually to a generalised notion of heredity capturing relations among traits independent of the particular life forms they were part of.4 This can be viewed as the main outcome of the first two workshops on the cultural history of heredity, to which all contributions, in one way or the other, attest. At the same time, however, this result provides the central, historiographical problem for our project. How is it that a phenomenon that, from a contemporary perspective, appears to be of central importance, and in 1 2 3 4

For more details on the project and the workshops see http://www.mpiwg-berlin.mpg.de/Heredity/. The original program of the second workshop is reproduced at the end of this volume. Results of the first workshop have been published in: A Cultural History of Heredity I: 17th and 18th Centuries, Max-Planck-Institute for the History of Science Preprint 222, Berlin 2002. See Mclaughlin, Müller-Wille, and Wood, this volume. See López Beltrán, McLaughlin, Müller-Wille, and Pomata in this volume; cf. Coffin, this volume, on the work of Bénédict Augustin Morel who, in the mid nineteenth-century, still upheld the distinction of normal vs. pathological heredity.

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its effects appears to be so tangible in everyday life, was subjected to conceptualisation so late? This seems to be the most curious feature of the cultural history of heredity: while ‘heredity’ today belongs to the most fundamental concepts of the life sciences, it entered the scenes of inquiry into the phenomena of life only very late in history. The late emergence of heredity coincides, moreover, with the important transformation that the life sciences underwent in general around 1800 and that Michel Foucault and François Jacob have described so succinctly. 5 Though both of these authors focussed on the concept of organisation in studying this transformation, it seems highly probable that the emergence of heredity represents an aspect of utmost significance for fully understanding this transformation.6 Most of what the essays assembled in this volume have to say on this point, leads to a solution of the historiographical problem just outlined that may come as a surprise. It is not that the concept of heredity emerged from a growing attention to regularities, a sort of fixation of the scientific mind on the laws of nature at the expense of the contingenices and complexities of real life. It seems, rather, that the emergence of heredity occurred within an epistemic space that unfolded while people, objects, and their relationships were set into motion. 7 This means that it was a condition for distinguishing between inherited and environmentally induced traits in organisms, for example, that organisms were actually removed from their natural and (agri-) cultural habitats. Only then could an environmental difference express itself in a difference in traits, and only then could heritable traits manifest their steadiness against environmental changes. Breeding new varieties for specific marketable traits, the exchange of specimens among botanical and zoological gardens, experiments in fertilisation and hybridisation, the dislocation of Europeans and Africans that accompanied colonialism, and the appearance of new social strata in the context of industrialisation and urbanisation, all these processes interlocked in mobilising cultural and natural ties and thus provided, as will be explained in more detail in the following, the material substrate for the emerging concept of heredity. It is true, of course, that the principle of ‘like engenders like’ had been around since the earliest times of Greek poetry and philosophy, as an expression for what ought to happen as a rule. 8 This ‘law’ remained unanalysed, however, so that it lacked the kind of inner structure that could have provoked the application of a metaphor that in its proper context, that of legal regulations of property transmission, possessed such complex semantics as ‘heredity’ did. And this, as Wolfgang Lefèvre demonstrates for the cases of Lamarck and Geoffroy St. Hilaire in this volume, remained valid for the preformationist and epigenetic theories of evolution up to the early nineteenth century. In a sense, even, both preformation and epigenesis – and both conceptions have a wellknown, ancient legacy – excluded inheritance: according to preformation nothing is transmitted in generation because everything has been there from the beginning; according to epigenesis nothing is transmitted in generation because in each instant everything is built up from scratch. 5 6 7 8

4

Michel Foucault, Les mots et les choses, (Paris, 1966); François Jacob, La logique du vivant, (Paris, 1970). Cf. Michel Foucault, Histoire de la sexualité, vol. 1: La volonté de savoir, (Paris, 1976). This is emphasised in the comments by Raphael Falk and Gianna Pomata included in this volume. For a different perspective see Waller and van Wyhe in this volume. See Erna Lesky, Die Zeugungs- und Vererbungslehren der Antike und ihr Nachwirken, (Wiesbaden, 1950); Hans Stubbe, Kurze Geschichte der Genetik bis zur Wiederentdeckung der Vererbungsregeln Gregor Mendels, (Cambridge, Mass., 1965; engl. transl. 1972).

Introduction

As Peter McLaughlin points out in his contribution, it is in Immanuel Kant that we encounter a theory of propagation which is neither preformationist nor epigenetic – and in which, at the same time, conceptions of heredity began to unfold a manifold of specific meanings. “Anerben”, “ererben”, “vererben”, “forterben” are all terms that Kant used in this context to distinguish, as McLaughlin puts it, “various aspects, permutations and combinations of hereditary phenomena.” The phenomenon that gave rise to this proliferation of terms was not simply the similarity that offspring exhibited with regard to their parents. It was rather a narrowly circumscribed, highly specific phenomenon, the existence of distinct races in the human species distinguished by traits that were invariably transmitted to offspring even if environmental conditions should change. Empirically this peculiar behaviour was exhibited to Kant by Portuguese colonists in Africa (whose children remained white, despite dislocation) and black Africans in Europe (who likewise continued to produce black children there). Such a phenomenon undercut the ancient distinction of specific forms and individual peculiarities: characterising classes at a subspecific level, racial characters belonged to the individual peculiarities that interfered with the universality of species form; yet being infallibly reproduced generation by generation they seemed to be subject to the same regularities that governed species form. To account for this, Kant brought together natural law and contingent (family) history in his concept of Vererbung: the potential or Anlagen for hereditary traits were included in the original organisation of ancestors, but once they had been expressed in reaction to a change in environment, they were permanently and irrevocably transmitted. The way in which Kant set up the problem and the way in which he advanced a solution by exploring the complex semantics of Vererbung can be regarded as prototypical for the emergence of heredity. The problem was not the constancy of species form but the patterns of variety that structure life at a sub-specific level. As long as such patterns coincided with the distribution of organisms over locally circumscribed environments, however, such patterns were readily explainable by the permanence of ties between organisms and their “natural places”. In these cases, it is, in a sense, the place that “inherits” its inhabitants and impresses its character upon them. It is only when this tie is dissolved to open up a variety of correlations between forms, places, and modes of transmission, that a need arises for a complex metaphor like heredity to be inserted in order to account for the proliferating phenomena. From this perspective, also, Kant’s preoccupation with human races seems less eccentric: it is, after all, first and foremost through human activities that people and, along with them, things are actually seen to be mobilised. Kant’s famous dictum, that better progress in knowledge may be achieved by letting the spectator revolve around objects quite consequentially put anthropology in the centre of thought. This motivation to apply and explore the concept of heredity in face of a mobilisation of social and natural ties was observed in all the cultural sub-fields that were explored at the workshop. Inheritance regulations that imposed restrictions to succession for future generations as well as the extent to which cousin marriages should be outlawed came under intense discussion in the aftermath of the French revolution. They were perceived as guaranteeing the stability of possessions and privileges on the one hand, and, on the other, as an obstacle to social strategies aimed at coping with the progressive dominance of mobile vis-à-vis landed property. 9 The interest of physicians in hereditary diseases turned from “noble” maladies like gout, seen to be “softly”

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inherited by over-consumption, to diseases like phthisis (tuberculosis) and madness that were ascribed to the rapidly growing class of the landless and poor urban populations and perceived as “degenerative” diseases based in a permanent defect of hereditary dispositions. 10 Breeders were progressively engaging in attempts to transplant strains from one country to the other, and to “mould” their creatures for specific traits in the context of industrialisation, and refining, alongside with this, the image of the “self-made man” which was reflecting concerns among the middle classes about inborn or acquired character in the wider, moral sense. 11 Natural historians and evolutionists came to recognise patterns of heredity in the series of transplantations and hybridisations that they set off in their experiments, experiments which were carried out not rarely despite the abhorrence experimenters sensed in face of their monstrous productions. 12 Anthropologists, finally, began to concentrate on racial differences as exhibited in the dislocation of European and African people or on the traits of “savage children” who had been uprooted and raised without education.13 These cultural subfields did not cohere immediately, after the model of mutual influences, but they were connected by a kind of domino effect that mobilisation in one field had on others. The import of plants for collection purposes of natural history inspired attempts at their acclimatisation for economic purposes (and vice versa). The successes of breeders in establishing marketable strains provided the model for the “self-made man”. The growth of a class that depended on mobile property evoked a culture of leisure collecting and breeding. Thus, the results of the first two workshops do not so much point to a unitary “culture of heredity” that suddenly emerged around 1800, but rather to a piecemeal relaxation of social and natural ties in several, highly specific cultural sub-fields that opened up the epistemic space which would eventually come to be occupied by the concept of heredity. We would like to thank the government of Liechtenstein for financial support.

Hans-Jörg Rheinberger

9

10

11 12 13

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See Ulrike Vedder’s contribution on the majorate. The topic of cousin marriage regulations was explored by David Sabean in the comment he gave at the workshop (unfortunately, he was prevented from preparing them for publication in this volume); see his Kinship in Neckarhausen, 1700-1870, (Cambridge 1998). These aspects are covered in great detail in the contributions to this volume by Laure Cartron, Phillip Wilson, John C. Waller, Jean-Christophe Coffin, John van Wyhe, and Paul White. Gianna Pomata discusses the first three of these contributions. See Roger Wood and Paul White in this volume. See Staffan Müller-Wille on the experimental tradition of “hybridism” in this volume. The former aspect is discussed by Peter McLaughlin, the latter by Nicolas Pethes in their contributions to this volume.

Heredity old and new; French physicians and l’hérédité naturelle in early 19thcentury. Carlos López-Beltrán

The coinage of hérédité At some point in the early decades of the 19th century, French medical men and physiologists adopted the noun “hérédité” as the carrier of a structured set of meanings that outlined and unified an emerging biological conceptual space. The elements of this domain had previously been loosely connected by the metaphorical mirroring between physical resemblance between parents and offspring and the passing on of property and titles through the generations and projected over the medical, zoological, agricultural, and ethnological fields. The development of such a conceptual space, and in those domains, during the first half of the 19 th century in several European countries, produced the first appearance of our modern concept of biological heredity. 1 Le Trésor de la Langue Francaise gives a 1821 quote as the first registered occurrence of the noun “hérédité” with a biological meaning.2 The work cited is Joseph de Maistre’s Les Soirées de Saint-Pétersbourg, where he talks of “cette triste hérédité” referring to the physical ailments bequeathed to the infants due to the sins of their elders from several generations. I have located several previous instances linked also to the transmission of disease, where the notion of a noxious bequeathal (facheuse hérédité) dropping down the generations is dominant.3 The link between the Christian (Augustinian) notion that hereditary physical ailments stem from a divine punishment linked to the original sin and the substantialization of hereditary diseases into a reified entity hérédité seems to me undeniable.4 The early 19th century French medical community played a major role in the articulation of our modern concept of heredity, only comparable with the role played by animal and plant breeders. The aim of this presentation is to outline that process. I begin with an empirical fact: after c.1830 “hérédité” became an increasingly popular noun amongst French medical men. In the preceding years the traditional medical formula maladies héréditaires (a derivation from the usage of morbi hereditarii at least since Avicenna) was being transformed into phrases like “hérédité des maladies”, “hérédité morbeuse”, “héredité pathologique”, transferring the stress of the fact of transmission to a noun, that opens the space for a more general notion, and somehow eroding the 1

2

3 4

For heredity during the 18th century see Rey (1989); A Cultural History of Heredity I (2002). For recent views on the history of the concept of heredity see Olby (1992); López-Beltrán (1994); Gayon (1999); Wood and Orel (2001); Waller (2001b). The metaphorical, adjectival use in several European languages derive from morbis haereditarii, and was well in place amongst physicians by the 16th century; we thus have “maladies héréditaires”, “Erbkrankheit”, “hereditary disease” and several variants scattered in medical treatises, with an increasing rate during the 18th century. See Appendix 1 in López-Beltrán (1992). See for instance Fodéré (1813), 5: p. 365. Le Grand Robert gives the same occurrence as the first one. De Maistre’s book is really from 1822. Two contests set by the Royal Society of Medicine in the 18th century where instrumental for the development of this analysis (López-Beltrán 1994). For the English world John Hunter did the most surprising and clarifying discussion; see Hunter ([1786] 1835-1837), 1: pp. 353-359.

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metaphorical cushion. The presence of the noun suggests the existence of a “thing” (a force, a law, a mechanism), the nature and reach of which was then progressively shaped by French physicians. After its adoption in French medical literature hérédité frequently was qualified by different adjectives that established important oppositions. Common among these was the contrast between “hérédité physiologique” and “hérédité pathologique”. But after a few decades the most influential opposition was that between “hérédité physique” and “hérédité moral”. The first pair was used to emphasize the growing perception that there was a natural aspect of hereditary phenomena, which was free from the common noxious connotations. 5 Such opposition served, I believe, to define the axis through which the structure of the medical concept of hereditary transmission was transported into a more general, biological frame, and was “de-pathologized” for the consumption by a wider spectrum of savants. I speak of the “structure of the concept” because I want to emphasize the existence in that period of a cluster of classificatory and explanatory elements that the medical analysis and disputes around the notion of hereditary transmission produced. In previous works I have shown how medics were the first theorists to propose and develop important features of biological heredity, like the distinction between innate and congenital characters, or the latent and pre-dispositional nature of hereditary causes; these and other criteria were used as pointers towards the necessary separation between accidental and hereditary variation.6 Once heredity was in place different physiological schools struggled to take over the increasingly powerful new domain. Several generations of French physicians, alienists, physiologists and naturalists contributed with alternative accounts of how heredity was acting in the shaping of individual, familiar, and even national constitutions. 7

Hérédité and Constitution The traditional medical concepts of temperament, complexion and constitution, which had been adapted in the different periods according to dominant physiological creeds had been for centuries the depositaries of underlying, general potencies and dispositions that could account for both typical and idiosyncratic physical responses of individual organisms under different surroundings. In contrast, the moral peculiarities of human beings were alternatively linked to, or separated from the physical (constitution, temperament) in accordance to theological and metaphysical positions.8 Late 18th century medics, when the pendulum was moving in some places towards materialistic approaches, found once more in the medical concepts of temperament and constitution a good framing device for their attempt at grounding the moral on the physiological, and on using that as a launching base for hygienist programs of physical and moral improvement 5

6 7 8

8

John Hunter in 1786 maintained that the hereditary principle “[…] may be divided into two kinds: the transmission of natural properties, and the transmission of diseased, or what I shall call acquired or accidental properties” (Hunter 1835-1837, 1: pp. 353-354). See López-Beltrán (1992). See also Olby (1992); Gayon (1999); Waller (2001a). Aliéniste was the name given in France to the early psychiatrists as they dealt with mental alienation, or insanity. The word for insane was aliéné. Roger (1963) remains the outstanding investigation of this for the French context from the 16th to the 18th centuries.

Heredity old and new; French physicians and l’hérédité naturelle in early 19th century

of humanity. The influential work of P. J. G. Cabanis set the tone for a repositioning among French medical men with regards to the plasticity or permanence of the inborn temperamental features with respect to the influence of the environment. Temperament could be modified, improved, up to a certain point. A crucial consideration in the hygienists project was that […] les habitudes de la constitution se transmettent des pères et mères aux enfants; qu’elles se conservent comme une marque ineffaceable, au milieu des circonstances les plus diverses de l’éducation, du climat, des travaux, du régime.9

The notion that some aspects of the bodily features could become entrenched in some lineages through this hereditary transmission was one that was progressively made explicit by physicians in the period. The transmission of the set of empirical facts that I have elsewhere called the hereditary (resemblance of offspring to parents, atavism, recurrence of disease or of striking peculiarities within families or groups...)10 provided a link between parents’ temperaments (or constitutions) and those of their children; that connection, extended over time to whole genealogies, justified the talk of family, or even national characters. Cabanis emphasized this, and insisted that it could be the basis for a program (similar to those followed by breeders) that eliminated the undesirable and consolidated the desirable traits. This of course was one of the influences that shaped Lamarck’s notions of the inheritance of bodily adaptations. 11 But it was disease that had driven the medics to pay such close attention to hereditary transmission in the first place. Physicians had found useful the notion of actual physical transmission (by both parents) of some kind of casual influence, through physiological means at the moment of conception, as a useful theoretical resource for the explanation of some diseases. Especially those with familial patterns, and mainly chronic ailments, which were also generically known as “constitutional.” Among them insanity, epilepsy and other mental abnormalities were sometimes included, but not particularly stressed.12 It is important not to mistake the presence of the notion of hereditary transmission with the presence of the concept of heredity itself. My story turns around this distinction. Lamarck is a good example of what I mean. Although posterior developments situated the name and the work of Lamarck in the center of the debates about heredity of acquired characters, neither him nor Cabanis (nor anyone from their generation) could pay attention to heredity itself, as it wasn’t there yet. As André Pichot has recently phrased it: […] même si l’hérédité est bien le centre de la théorie de Lamarck, ce centre n’est pas theorisé. Par la suite, quand on parlera de la théorie de l’hérédité lamarckienne, on va projeter sur la théorie de Lamarck la notion d’hérédité elaborée après sa théorie.13 9 10 11 12

Cabanis, ([1802] 1824), 3: p. 431. See Staum (1980); and Williams (1994), for Cabanis’ influence on the French medical world in early 19th century. López-Beltrán, (2002). See Cabanis (1824), 3: p. 434. See also Carol (1995), p. 24. For the influence of Cabanis and other ideologues on Lamarck see Corsi (1998). The hereditary character of mental diseases had been a characteristic observed and discussed since ancient times. In the Hippocratic corpus it is in discussing epilepsy, the sacred disease (morbus sacer) that some of the clearer passages concerning the hereditary and its relation to generation, is to be found. See for this Lonie (1981) and Boylan (1984).

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This elaboration required, I believe, that medical men integrated the notion of hereditary transmission within the semantic field of the powerful and prestigious concepts of temperament and constitution. There have been several successful studies of the expansion of hereditarianism into different areas of the medical practice and of natural history after the 1840s. 14 However, the previous, necessary, and rather dramatic shift in emphasis, during the first few decades of the century, from the hereditary as an important but secondary (predisposing) component of many physical and mental conditions, to heredity as a main (if not the main) cause responsible for all the natural bodily (and thus, for some, moral) endowments of individuals has not been adequately described. William Coleman has persuasively shown how, with the turn of the century, French physicians abandoned the Galenic old language of the “six non-naturals” when alluding to the external influences on health (like nutrition, climate, etc.) and began to speak of hygiene. I believe that a parallel move occurred with the complementary concept of the “naturals” (i.e. the temperament), which was eventually substituted by heredity, in order to reproduce the previous dialectic between body and external milieu.15 The social and political developments during the French Revolution, and afterwards during the Napoleonic reforms, opened the space in which medical practitioners found the opportunity to promote these two new powerful conceptual weapons that could help promote their gremial aspirations for a major role in the reorganization of civil life. Hereditary transmission had never before been looked at as a subject of special analytical and theoretical interest. Then, in a few years, it was all over the place.16 There was a notorious change of emphasis, and what had been kept aside on footnotes or short discussions, became the subject of chapters in books, and an increasing number of dissertations. Particularly curious is the adoption after 1810 of hereditary explanations by some medical authors who had written several works before and had not resorted to them. I can mention Emmanuel Fodéré, Antoine Portal, and Philippe Pinel; all of which played leading parts in the post-revolutionary medical reforms. They had of course acknowledged the hereditary influences in their early works, but only at this period they gave them a central role. 17 After the Revolution the hereditarian wave gathered momentum, and with the exception of isolated cases of skepticism,18 the French medical community seems to have been finally overtaken by it by the beginning of the 1830s. The process by which the conversion of heredity from a marginal metaphoric use into an important explanatory tool within the biological disciplines occurred and the momentum it 13 14 15 16

17 18

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Pichot (2002). Dowbiggin, (1991); Gayon (1999); Orel (1996); Waller (2001a). Coleman (1974); (1984); Williams (1994); and López-Beltrán (2002). For the appeal to the dialectics of hygiene and heredity see Williams (1994). Cabanis managed to defend simultaneously that the breeds of human beings ought to be improved to attain a basic equality, and that diversity is a basic human value that ought not to be jeopardized (Cabanis 1824, pp. 435-436). For instance Fodéré (1792); Portal (1781), (1800); and Pinel (1785). Wide perspectives can be had in Poilroux (1821); Caillot (1818). See also Ackerknecht (1967); Williams (1994). An important skeptical argument was made by Sersiron (1836). He maintained that true hereditary transmission of disease ought to be as “fatal” and deterministic as the hereditary transmission of specific characters, like the shape of the bones or the form of the eyes. Any accidental character can disappear from the genealogical line after a few generations, so it cannot properly be claimed to be affected by the same hereditary cause that maintains the unity of the species. Their transmission is therefore also accidental and not lawfully governed.

Heredity old and new; French physicians and l’hérédité naturelle in early 19th century

gathered in these decades remains to be better understood. Among the several factors influencing the process a few more can be mentioned, besides the post-revolutionary reforming zeal: The rise and fall of the phrenological movement, and the disputes around vitalist physiology and medicine.19 The outbreak of hard hereditarian, degenerationist and racist thinking in the French scene after the 1850s is well documented. Its brewing phase among the alienists in the decade of the 40s has been also described. What has not been seen clearly is that this in its turn was only possible due to the great amount of groundwork that the previous generation of French medical men had been accomplishing by shaping and structuring a working notion of hérédité.

Hérédité in the physiological battleground The French milieu transformed its marginal 18th century medical disputes around les maladies héréditaires into a widespread theoretical and ideological 19th century preoccupation with the general workings of l’hérédité. The last decade of the 18th century, and especially the early years of the 19th, had witnessed the publication in France of a number of treatises, essays, articles, and dictionary entries on “hereditary diseases”. François Pagès, Alexis Pujol, and Joseph Claude Rougemont published essays on hereditary disease written for a Royal Society of Medicine competition (1788-90).20 These were soon joined by a succession of very authoritative works on the subject. Antoine Portal, Antoine Petit, Emmanuel Fodéré wrote extensively on the subject. 21 Hereditary causation was with increasing frequency emphasized in general pathologies and in treatises concerning the main chronic diseases.22 With some reluctance, but unequivocally, the famous alienist Philippe Pinel had acknowledged the importance of hereditary predisposition in the onset of mental illness, and very soon his followers enthusiastically took to exploring the theme. Among them Esquirol and Fodéré, who promoted the hereditary influence from the “back row” of secondary influences on insanity to the forefront of one of its main predisposing physical causes.23 The understanding of hereditary transmission of disease had of course a close connection to the idea that constitution and/or temperament somehow “run in the families” as well as in the wider genealogical groups. Both the way these influenced or predisposed the individual’s body to react in given manners, and the fashion in which the physical state of both parents could actually influence that of the new being through their seminal fluids were main considerations. The complex transition from old humoralism to solidist and vitalist physiologies, during the 17 th and 18th centuries, had the accounting for hereditary diseases as one of its battlegrounds. 24 One can find representatives of all these approaches in early 19th century France. This created some tension when French medics sought to unify their views under the common cause of general (pathological) hérédité. The outcome was that the general term physiological heredity became accepted as referring to the normal mechanism by which bodily resemblances are transmitted through the generations (whatever their actual instantiation). Pathological heredity was then to be 19 20 21 22 23 24

See Pick (1989); Chamberlin (1985); Borie (1981); Carol (1995); Dowbiggin (1991). See López-Beltrán (1994) for the complete story about that competition. See Fodéré (1813) “maladies hereditaires”. See Fodéré (1809); Baumes (1805); Portal (1808). Pinel (1812); Fodéré (1817); and Esquirol (1820). The major source for this story is Rey (2000).

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seen as based on the same principles, but having as object the transmission of deviant particularities that predisposed to disease.25 Probably by 1820 most French medical men, and some physiologists and naturalists, considered biological heredity an important issue. For them, hereditary transmission of a whole range of characters occurred without doubt and what really remained to be understood was the reach, the power and the limitations this phenomenon had in both humans and other species. Particularly crucial for different reasons was to know 1) if some socially damaging diseases, specially mental insanity, were indefinitely preserved within genealogical lines (in this case families), 2) if the racial, national and other group differences between humans could be entirely ascribed to the preservation within genealogical lines of hereditary variations (or degenerations), and 3) if characters that affected the specific type of the living organisms could also be preserved within genealogical lines in such a way as to challenge the age old belief in the immutability of species. It was increasingly believed that these relatively different questions (concerning medical men and alienists, anthropologists and naturalists) could be confronted with a unified analysis of the phenomena: a general theory of hereditary transmission. This was the idea that occurred to several medical authors during the second decade of the 19th century. A very convincing register of this development can be found in the 60 volumes of the Dictionnaire des Sciénces Médicales, which from the period of 1812 to 1820 captured, in its different entries, this progressive generalization of the hereditary (in the loose metaphorical use) into a nomological approach to biological heredity. There is a sense then in which this dictionary can be described as a kind of forum where the positions of different influential physicians were being rehearsed and criticized successively. This dictionary opens a window with which we can look at the concept of heredity being shaped. As late as 1812, Philippe Pinel, the grandfather of French alienism did not consider the hereditary cause important enough to deserve a mention in his inaugural paper on “aliénation”, preoccupied as he was in giving his “moral causes” the leading role. The editors managed to give heredity a greater role by commissioning a further article on the overlapping subject of the “aliéné”, to a disciple of Pinel’s, Marc, who in his piece stressed the importance of hereditary predisposition to insanity. “Elle établit -he wrote- une des plus fortes présomptions en faveur de la realité de l’aliénation mentale.”26 But it was Esquirol, the crown prince among Pinel’s followers (and teacher to the openly hereditarian generation that followed) who gave heredity the leading role as an influence for mental disease in his 13 articles for the Dictionnaire.27 In both “folie” and “manie” he gave advances of what was to become his classic book on Maladies Mentales (1838).28 As is well known, Esquirol was the first to organize in statistical tables the cases of mental insanity, with the intention of sorting out the importance of each causal influence. Heredity he found to be a major “physical” cause, and in certain circumstances a dominant one. 29

25 26 27 28

12

Physiologists like Burdach (1837) and Flourens(1863) were among the most influential to propose a general physiological account of heredity. Marc (1812). Like Georget, Moreau de Tours, Baillarger, Morel. For accounts of Esquirol and his school’s work see Ackerknecht (1959), chap.6, pp. 37-51. See also Semelaigne (1894). Esquirol (1816), (1818).

Heredity old and new; French physicians and l’hérédité naturelle in early 19th century

For the entry on “maladie héréditaire” a recently published essay by Antoine Petit (1817) was included in the Dictionnaire. 30 This piece remained the most influential analysis published on the subject until the 1840s. It was a clear and convincing attack on humoralist heredity. Echoes of Petit’s precisely worded piece can be found in works written sixty or seventy years later. Petit summarizes what he considers to be the main achievements that medics had attained in the definition of the hereditary cause. Heredity, he asserts, has to be based upon particular states of the bodily constitution communicated to children by parents. These states give an “organic disposition” to re-produce a given effect, for instance a particular disease. He adds that they can be both localized states, or states of the whole économie, but he denies that some kind of general qualities of the constitution (like weakness) that establish in the body vague and indefinite tendencies (to disease) are to be seen as similarly hereditary. In heredity a specific, one to one connection must be shown to exist. Petit praises insightfully the ancient distinction between predisposing and efficient causes as the main analytic resource to deal with the hereditary.31 He summarizes, with more clarity than any previous author, the determinant features of heredity. Latency, homochrony (to use Haeckel’s later term), atavism, all can be accounted for with a proper causal analysis. He upholds the importance of separating clearly congenital and connate influences, and accepts that only through the process of generation can real hereditary influence be transmitted. He however joins previous authors in condemning attempts to solve the mystery of heredity by an even deeper and more unsolvable mystery of generation. Hypothetical theories of generation (“systèmes”) only confuse the issue. It is far more likely he adds that the proper observation of the patterns and nature of hereditary disease will illuminate the theorizing in generation, than the other way round. 32 Although he is skeptical about the feasibility of any success, Petit leaves it to other specialists to decide over the real (intimate) nature of the inherited dispositions. The good observer however can on occasions find visible, exterior characters that are linked to the disposition, before its effects are noticeable. Generally, however, this is not the case, and though there is an organic base to hereditary causes, they usually remain hidden (latent) until the time, in the life pattern, comes for their expression. This theme of the hidden cause that exposes itself at a given time was to be retaken by different authors of the Dictionnaire of both medical and physiological orientation. After Petit’s solid defense of heredity, the articles of the Dictionnaire on all constitutional, chronic diseases gave a preeminent role to heredity. The entries on “scrophules” and “phthisie”, for instance, join vigorously the attack on humoralistic, taint dependent explanations of hereditary transmission,33 favoring without reserve the view that heredity is to be ascribed to inborn constitutional (organic) peculiarities, that predispose for certain conditions. There is a wish in several authors of the Dictionnaire to make it clear that there is nothing particularly 29

30 31 32 33

Esquirol held heredity as an influential cause of insanity among the wealthy. His views were close to the solidist tradition; he spoke of it as a physical, predisposing cause, and believed that homochrony and latency were particular signs of the presence of an hereditary cause. Like Pinel, he was sure of a physical base for human mental states, but was not a fatalist and gave more importance to efficient, moral causes. Petit (1817). “Distinction lumineuse […] qui repose tout entière sur les faits, sera toujours une des sources fécondes où le médecin habile puisera les notions plus positives” (Petit 1817, p. 59). Petit (1817), p. 63. Fournier-Pescay and Begin (1820), pp. 278-386; Maygrier (1820), pp. 15-168.

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pathological with the route (or mechanism) through which the structural anomalies are communicated from parents to children. Normal physiological processes were responsible for that. Once the constitution acquires a flaw, the natural trend would be to transmit it through generation, as are transmitted all other constitutional features and qualities responsible for general and particular resemblance between parents and offspring. The open end of the discussion (which Petit shied away from) was the question of how to understand constitution,34 and how to describe its causal influence in the life of the organism. Where some medics saw constitution as a synthetic (cluster) term referring to the sum of the organic parts of the body (organs, tissues, etc,) and their organization, others saw the term as linked to functional qualities, non-reducible to general or particular dispositions. The different attitudes had a root in the tension between material and functional explanations. Between anatomy and physiology. And within physiology itself, between purely materialistic, and vitalist ontologies. “Constitution” was traditionally a term with enough breath to encompass different, and relatively incompatible conceptions of the body, of its organization and function. Heredity, as a concept that was being integrated into the same semantic niche, acquired the same quality. “Constitution”, with its relative synonyms “temperament” and “complexion”, defined a general space whose details, whose actual goings on, had still to be fought over by the proponents of different physiological theories. De Montenegre, in his Dictionnaire piece on “maladie constitutionelle” provides a striking illustration of this view of the body as a battlefield: […] le corps animal peut être considerée comme formé de plusieurs êtres independens, jusqu’à un certain point les uns des les autres, par leur manière d’agir; mais concourans tous à former un résultat général qui est la vie. Il doit nécessairement exister entre ces différens un sorte d’equilibre d’action, […]. C’est ainsi que l’on peut concevoir ces dispositions individuelles qui s’étendent au moral comme au physique, et établissent entre tous les hommes une variété infinie.35

A constitution could be ascribed general states, or forms of being, that in turn would be responsible for reactions to stimuli, for dispositions, etc. Or it could be ascribed particular states or forms of organization responsible for localized reactions, in a given organ or part. The peculiarity of the constitutional variation could be material, and observable in principle, or it could be only a potentiality rooted in some emergent quality (like irritability) or a vital or dynamique state. Coincidence focused around the existence of a basically fixed set of physical dispositions that characterize each individual. An important turning point seems to have been the adoption by French physiologists, influenced heavily by the work of Burdach, of a set of analogies from the physical sciences that aimed at framing difficult causal issues in dynamical terms. The idea that some dispositions in living beings were not to be explained solely in terms of their basic structural physical properties but also depended on particular sets of “dynamic” properties was used to reframe the ancient, and threatened, medical concepts of constitution and temperament, in order to give them a new 34

35

14

Fournier (1820) tried to make the differences between these concepts explicit. “Il existe -he wrote- entre les mots tempérament, constitution et complexion, quelle que soit leur synonymie, dans certains cas, des nuances qui permettent pas les employer indistinctement les uns pour les autres.” Montenègre (1820), p. 246.

Heredity old and new; French physicians and l’hérédité naturelle in early 19th century

respectability.36 Both concepts were aimed at capturing the fixed set of physical dispositions (structural and dynamic) that each individual possessed through his life. After Cabanis, few authors doubted that there was a link (a rapport) between the parents’ constitution and that of the new beings they gave rise to. “Hérédité”, after 1830, was ideal as a basic conceptual tool that could be deployed to highlight and investigate such relationship. As Michel Lévy summed it up: […] á l’étude de la Constitution se rattache essentiellement celle de l’hérédité dans la santé et dans les maladies […]. L’hérédité éclate chez l’homme dans sa forme génerale et dans la proportion relative des ses parties; elle se manisfeste par les propriétés intimes de la fibre organique.37

This close explanatory link with ‘constitution’ gave the notion of ‘hérédité’ a wide appeal both for the medical men and naturalists, and eventually transformed it into an common frame accepted by most, whose contents where being debated and defined en route. Resemblances, in form and function, in health and illness, in body and mind, could with its help be rooted in a historical causal chain. Eventually, naturalists and anthropologists that adopted hérédité participated in the struggle to fashion it into a broader rational explanatory scheme, for which the boundaries and ways of actions needed to be clearly defined. A part of that work was directed towards having clarity about the kinds of characters heredity would affect. A clearly defined set of types of characters in a hierarchical scheme was seen by many as crucial. From specific, through racial, to individual, features (on one axis); and from physical to moral characters (on the other axis), authors would debate the reality and relative importance of an increasing number of claims of hereditary transmission of traits. The main problem that most theoretical generalizations faced was, again, irregularity; the proliferation of exceptions. Besides, given the bad track record of general hypothetical systems, medics had strong feelings about not letting those outsiders impose definitions of constitutional dispositions and heredity without giving enough weight to their (medical) accumulated experience.38 For them “l’hérédité pathologique” should inform “l’hérédité physiologique” before the latter could reciprocate. And so the definition of heredity itself should be based on medical men’s assessments of what generally is the case. To which extent, for instance, were only general, non-localized constitutional dispositions (or characters) inherited; or were also very localized and particular ones (like moles, or bladder stones) transmitted. Such a question was better answered, some medics believed, observing the patterns of disease communication, given that a disease (or a malformation) was a much clearer sign than other normal resemblances, just like in the case of moral phenomena, it is easier to follow, in a family, a pattern of a distinctive set of symptoms, as those of insanity, than it is to follow vaguer, positive qualities, like honesty or strength of will. It is a sign of the effectiveness with which contributors to the Dictionnaire made their cases that many of the matters they discussed around hereditary transmission were considered a given by most French medical men after them. Elaborations of their main tenets followed. The schools of 36 37 38

Some authors made complex analytical distinctions between these two concepts wheras others saw them as equivalent. Lévy ([1844] 1869) 1: pp. 114, 186. For this position of the medics see particularly Petit (1817); Lereboullet (1834); and Piorry (1840).

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medicine of Paris, Montpellier, and Strasbourg were regularly producing theses, both by students and professors, dealing with different aspects of hereditary transmission of diseases. Works in which with increasing frequency a nomological attitude towards l’hérédité was assumed. D.A. Lereboullet (1834), M. Lévy (1844), C. Béclere (1845), and specially P.A. Piorry (1840), produced good expositions of how heredity worked in normal features and in the communication of disease. But by the time these medics wrote, the field was ceasing to be a purely medical and pathological one. Slowly but constantly heredity was becoming a hotly debated social and scientific issue. What medical men and some physiologists had for decades been arguing about the hereditary base of human nature, finally captured the attention of broader sections of mid-19 th century French intelligentsia, who saw the potential power of their ideas for accounting for the unaccountable: the human soul in its collective and individual dimensions, and its dependence upon the body’s constitutional make-up, or organization. D.A. Lereboullet summarized the general importance of heredity, outside the purely medical realm. He stressed the uniqueness of the human case, of which medics had the privilege of having more experience. Among humans, he wrote “l’organisation nous présente des différences individuelles” based on the innumerable combinations that different constitutions, temperaments, and idiosyncrasies can produce. Together with the many modifications that external factors (climatic, passions, education) can make they can “rendre raison des nuances infinies que nous observons entre les hommes […] sous le rapport de leurs charactères physiques et moraux.” But, he added, these subtleties can be further analyzed […] si nous appliquons à l’étude physiologique de l’espèce humaine la méthode des naturalistes, nous pourrons encore distinguer, au milieu de ces nombreuses différences, certains caractères communs, certains types originaux dont plusieurs auront persisté à travers une longue series de siècles. Les points de ressemblance seront plus nombreux entre les individus d’une nation isolée et qui n’aura pas contracté d’alliances étrangères. Enfin, si nous portons nos regards sur les membres d’une même famille, nous trouverons entre les enfans et les parens une conformité des plus évidentes: traits du visage, taille, son de la voix, couleur de la peau, constitution, tempérament, habitudes, caractère, moeurs, penchans, tout se ressemble. C’est sous l’influence de cette loi immuable, en vertu de laquelle l’homme donne le jour à des êtres semblables à lui, que l’on voit aussi quelquefois des vices de conformation se transmettre de génération en génération. Ainsi nous héritons de la constitution et du tempérament de nos parens; nous héritons de leurs caractères physiques et moraux; nous héritons de leurs vices de conformation.39

The possibility, described by many 18th and 19th century authors,40 of having stable, hereditarily and genealogically based, natural human groups under the level of the species (races, varieties), could easily be extended to other “socially useful” categories, like the family, and the nation. Genealogy as the basis for classification, with heredity as the main explanatory concept was profiling itself as a promising approach outside the medical realm. At the same time the vagueness

39 40

16

Lereboullet (1834). Among them Maupertuis (1745); Hunter (1786); Joseph Adams (1814); Prichard (1813); Pujol (1802).

Heredity old and new; French physicians and l’hérédité naturelle in early 19th century

of key working concepts, like latency and predisposition, was an open invitation for all sorts of analogical reasoning among imaginative theoreticians. After the 1840s, increasing attention was being paid to the ‘moral’, or later on ‘psychological’ aspect of heredity in humans. Heredity had ceased to be a pathological term with analogical links with normal resemblances. It ceased to be a simple concept and began to embody more than just a small sector of the medical community’s view of the human body’s original make and dispositions. Its empirical basis was expanded by its closer linkage to biological phenomena (like the origin of varieties), and its theoretical structure was also thoroughly expanded. By 1834 D. A. Lereboullet, then a candidate for medical chair at Strasbourg, could confidently assert that the majority of authors understood Hérédité as the transmission of particular (bodily) dispositions that tend to re-produce in children the same characteristics their parents had (resemblances, diseases), at the same age, or in the presence of the same exciting cause. More or less at the same period Charles Darwin began his lifelong effort to clarify the complexities of hereditary transmission within lineages. As Mary Bartley and others have shown, the features of the notion of heredity and hereditariness that he considered more useful to begin his explorations were basically the same that French and British medical men had been deploying for several decades: latency, homochrony, reversion.41 A cursory look at Darwin’s marginalia on his copies of medical works, like those of Lucas, Piorry or Henry Holland will eliminate any doubt about a robust connection.42

References Ackerknecht, Erwin H. 1959. A Short History of Psychiatry. New York: Hafner. _____. 1967. Medicine at the Paris Hospital 1794-1848. Baltimore: The John Hopkins University Press. _____. 1982, “Diathesis: the word and the concept in medical history.” Bulletin for the History of Medicine 56: 317-325. A Cultural History of Heredity I: Seventeenth and Eighteenth Centuries. 2002. Preprint 222. Berlin: Max Planck Institut für Wissenschaftsgeschichte. Adams, Joseph. 1817. Memoir of Life and Doctrine of the late John Hunter. London: Callow. Bartley, Mary M. 1992. “Darwin and Domestication.” Journal of the History of Biology 25 (2): 307-333. Baumes, Jean B. T. 1805. Traité de la Phthisie pulmonaire. Paris. Béclère, Claude. 1845. De l’Hérédité dans les Maladies, Doctorat Thesis, Paris: Faculté de Médecine. Rignoux imp., 22 pp. Bleynie, Francis. 1865. Considerations Génerales sur l’Hérédité Physique et l’Hérédité Morale. Paris. Borie, Jean. 1981. Les Mythologies de l’Hérédité aux XIXe siècle. Paris: Galilee. Boylan, Michael. 1984. “The Galenic and Hippocratic Challenges to Aristotle’s Conception Theory.” Journal of the History of Biology 17 (spring): 83-112. Burdach, Karl F. 1837. Traité de Physiologie. 13 volumes. Translated by A.L. Jourdan. Paris. Burton, Robert. [1651] 1932. The Anatomy of Melancholy. London: Dent and Sons. Everyman’s Library. Cabanis, Pierre J.G. 1802. Rapports du physique et du moral de l’homme. 2nd edition. Paris. _____. 1824. Oeuvres Complètes. Paris: Bossange-Firmin Didot. Caillot. 1819. Traité de Pathologie Generale. Paris. 41 42

See Bartley (1992) and Winther (2000). See Di Gregorio (1990) and (1986).

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Carol, Anne. 1995. Histoire de L’Eugénisme en France: les médecins et la procreation XIXe-XXe siècle. Paris: Seuil. Chamberlin, J. Edward, ed. 1985. Degeneration: The dark side of progress. New York: Columbia University Press. Coleman, William. 1974. “Health and Hygiene in the Encyclopédie: A Medical Doctrine for the Bourgeoisie.” Journal of the History of Medicine 29: 399-421. _____. 1984. “Inventing Demography, Montyon on hygiene and the state.” In Transformation and Tradition in the Sciences, edited by E. Mendelsohn. Cambridge: Cambridge University Press. Corsi, Pietro. 1988. The Age of Lamarck: Evolutionary Theories in France 1790-1830. Berkeley: University of California Press. Delage, Yves. 1903. L’Hérédité et les grandes problèmes de la Biologie Générale. 2nd edition. Paris: Reinwald. Di Gregorio, Mario. 1986. “Unveiling Darwin’s Rotos.” Archives of Natural History 13 (3): 313-324. _____. 1990. Charles Darwin’s Marginalia. Volume 1. London: Garland. Dowbiggin, Ian R. 1991. Inheriting Madness: Professionalization and Psychiatric Knowledge in 19th-Century France. Berkeley: University of California Press. Esquirol, J. Etienne D. 1816. “Folie.” In Dictionnaire des Sciences Médicales. Volume 16. 118. _____. 1818. “Manie.” Dictionnaire des Sciences Médicales. Volume 30. 37-472. _____. 1838. Des Maladies Mentales. Paris: Baillère. Flourens, Pierre. 1861. De la raison du genie et de la folie. Paris: Garnier. _____. 1863. Ontologie Naturelle, ou Etudes Philosophiques des Etres. Paris: Garnier. Fodéré, François Emmanuele. [1792] 1800. Traité de Goître et du Crétinisme. Paris. _____. 1813. Traité de Médecine Legale et d’Hygiene Publique. Paris: Croullebois. _____. 1817. Traité du Délire. Paris: Croullebois. _____. 1821. “Vie.” Dictionnaire des Sciences Médicales. Volume 57. 434-603. Fournier-Pescay. 1812. “Constitution.” Dictionnaire des Sciences Médicales. Volume 4. 158. Fournier-Pescay and Begin. 1820. “Scrophules.” Dictionnaire des Sciences Médicales. Volume 50. 278-386. Gayon, Jean. 1999. Darwinism’s struggle for survival: Heredity and the Hypothesis of Natural Selection. Cambridge: Cambridge University Press. Hamlin, Christopher. 1992. “Predisposing Causes and Public Health in Early Nineteenth-Century Medical Thought.” Social History of Medicine 5 (1): 43-70. Holland, Henry. 1839. “On Hereditary Disease.” Medical Notes and Reflections. 1st edition. London. 9-37. (3rd edition, 1855, 16-53.) Hunter, John. 1786. Observations of certain parts of the Animal Oeconomie. London: Castle-Street. _____. 1835-1837. The Works of John Hunter. 3 volumes. London: Longman Rees. Larousse, Pierre. 1873. Grand Dictionnaire du XIXe siècle. Paris. Lereboullet, D.A. 1834. De l’hérédité dans les maladies. Strasbourg: Silbermann. Lévy, Michel. [1844] 1869. Traité d’Hygiène Publique et Privée. Paris: Baillière. Littré, Emile. 1863. Dictionnaire de la Langue Française. Paris. Lonie, Ian M. 1981. The Hippocratic Treatises. Berlin and New York: Walter deGruyter. López-Beltrán, Carlos. 1992. Human Heredity 1750-1870: The construction of a domain. Ph.D. diss., London: King’s College. _____. 1994. “Forging Heredity: from metaphor to cause, a reification story.” Studies in the History and Philosophy of Science 25: 211-235. _____. 1995. “‘Les maladies héréditaires’ 18th century disputes in France.” Revue d’Histoire des Sciences. 47 (3): 307-350. _____. 1999. “Statistics and Human hereditary talent.” Ludus Vitalis. 7 (11): 11-27. _____. 2002. “Natural Things and Non-Natural Things: The Boundaries of the Hereditary in the 18th century.” In A Cultural History of Heredity. Preprint 222. Berlin: Max Planck Institut für Wissenschaftsgeschichte. 67-88. Lucas, Prosper. 1847-1850. Traité Philosophique et Physiologique de l’Hérédité Naturelle dans les état de santé et de maladie du système nerveux. 2 volumes. Paris: Mason. Maistre, Joseph de. 1822. Les Soirées de Saint-Petersbourg. Paris: Rusand. Marc, J. 1812. “Aliéné.” Dictionnaire des Sciences Médicales. Volume 1. 311-329. Maupertuis, Pierre Louis Moreau. 1745. Vénus Physique. La Haye. Maygrier, J.P. 1820. “Phthisie.” Dictionnaire des Sciences Médicales. Volume 42. 15-168. Montenègre, Alphonse de. 1820. “Maladie Constitutionelle.” In Dictionnaire des Sciences Médicales. Volume 6. 246.

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Olby, Robert. 1992. “Constitutional and Hereditary Disorders.” In Companion Encyclopedia to the History of Medicine, edited by W. Bynum and R. Porter. London: Routledge. Pagés, Jean François. 1798. “Héréditaires (maladies).” Dictionnaire de Médecine: Encyclopedie Méthodique. Volume 7. Paris: Chez Agasse. Pariset, J. and Villeneuve, A. 1812-1822. “Diathèse.” Dictionnaire des Science Médicales. Volume 10. 248-250. Petit, Antoine. 1817. Essai sur les Maladies Héréditaires. Paris: Gabon. Reprinted in Dictionnaire des Sciences Médicales.Volume 17. 58-86. Petit, Antoine, ed. 1812-1822. Dictionnaire des Sciences Médicales. 60 volumes. Paris: Panckoucke. Pichot, André. 2002. “Génetique Humaine: rêve ou cauchemar?” www.ogmdangers.org/action/cr_conference/Pichot_Benichou.htm Pick, Daniel. 1989. Faces of Degeneration: A European disorder c.1848-c.1918. Cambridge: Cambridge University Press. Pinel, Philippe. [1801] 1806. Traité Médico-philosophique sur l’Alienation Mental, ou la Manie. Paris: Brosson. _____. 1812. “Aliénation.” Dictionnaire des Sciences Médicales. Volume 1. Piorry, Pierre-Adolphe. 1840. De l’Hérédité dans les maladies. Paris: Bury. Poilroux, J. 1821. Nouvelles Recherches sur les Maladies Chroniques. Paris. Portal, Antoine. 1781. Observations sur la Phthisie de naissance. Paris: Memoires. _____. 1808. “Considérations sur la nature et le traitement de quelques maladies héréditaires, ou de famille.” Memoires Institute Nationale de France 8 (Semestre 2): 156-180. Prichard, James Cowles. 1813. Researches into the Physical History of Man. 1st. edition. London: J & A Arch. Pujol (de Castres), Alexis. [1802] 1823. Oeuvres Médicales. 4 volumes. Reissued by Boisseau as Oeuvres de Médecine Practique. 2 volumes. Paris: Bailliere & Bechet. Rey, Alain, ed. 1992-1999. Dictionnaire Historique de la Langue Française. Paris: Le Robert. Rey, Roselyne. 1989. “Génération et Hérédité au 18e siècle.” In L’ordre des caractères, edited by Bénichou Claude. Paris: Vrin. 7-48. _____. 2000. Naissance et développement du vitalisme en France de la deuxieme moitié du 18e siècle a la Fin du Premier Empire. Oxford: Voltaire Foundation. Roger, Jacques. 1963. Les Sciences de la vie dans la Pensée Française de XVIII siecle. Paris: Armand Colin. Russell, Nicholas. 1986. Like Engenderin’ Like: Animal Breeding in Early Modern England. Cambridge: Cambridge University Press. Semelaigne. 1894. Les Grandes Aliénistes Françaises. 2 volumes. Paris: G. Steihel. Schiller, Joseph. 1978. La notion d’organisation dans l’histoire de la biologie. Paris: Maloine. Sersiron. Patrick. 1836. De l’hérédité dans les maladies. Thèse de doctorat. no. 339, Paris: Faculté de Médecine. Staum, Martin S. 1980. Cabanis: Enlightenment and Medical Philosophy an the French Revolution. Princeton: Princeton University Press. Virey, Jean Joseph. 1808. L’Art de Perfectionner l’Homme. Paris: Deterville. _____. 1812-1822, “Monstres.” “Physionomie.” “Variation.” Dictionnaire des Sciences Médicales. 60 volumes. Paris: Panckoucke. Waller, John C. 2001a. The Social and Intellectual Origins of Sir Francis Galton’s Ideas on Heredity and Eugenics. Ph.D. diss., London: University College. _____. 2001b. “Ideas of Heredity, reproduction and Eugenics in Britain, 1800-1875.” Studies in History and Philosophy of Biology and the Biomedical Sciences 32 (3): 457-489. Williams, Elizabeth. 1994. The Physical and the Moral: Anthropology, Physiology, and Philosophical Medicine in France, 1750-1850. Cambridge: Cambridge University Press. Winther, Rasmus G. 2000. “Darwin on Variation and Heredity.” Journal of the History of Biology 33: 425455. Wood, Roger and Vítezslav Orel. 2001. Genetic Prehistory in Selective Breeding: A Prelude to Mendel. Oxford: Oxford University Press.

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The Sheep Breeders’ View of Heredity (1723-1843) Roger J. Wood

What can be revealed from horse breeding within a century can be achieved in sheep breeding within a decade. (A. von Weckherlin 1846, original in German)

Breeding sheep for desired characteristics has a long history. The ingenuity of generations of farmers ensured that the best of breeds, and the most ‘noble’ animals among them, would become valued objects of trade, war booty and gifts between monarchs. In eighteenth century Europe something new and radical occurred when advances in selective breeding technology became associated with improved rearing conditions and carefully designed feeding experiments. It was then that artists began to bring the evidence of breed variability and transformation to a fascinated public. Meanwhile an expanding army of agricultural writers was attempting to explain how the various breeding successes had come about. By the mid-nineteenth century, principles of breeding had been established that would serve the farmer for years to come. Attempts made to discover patterns of relationship between generations, to predict the results of breeding, met with gratifying success in skilled hands although never to the point of providing a functional explanation of biological inheritance. Theory was restricted to practice-based axioms and breeding rules, and it evolved with technological advance. It is these technology-driven changes in the concept of heredity1 that I shall be reviewing during the period under consideration (1723-1843). The purpose is to observe the relationship between experience and theory, as the latter becomes progressively modified as an aid to further practical advance. Three significant dates will serve to guide us through the period: •

•

•

1

1723 when Jonas Alströmer (1685-1761) imported the first Merino sheep from the warm dry atmosphere of Spain to the cold dampness of Sweden, and thereby set a trend in fine wool production throughout Europe and beyond, even finally to the Antipodes; 1783 when Robert Bakewell (1725-1795) and his friends in the Midland counties of England formed the Dishley Society, the first breed association, set up to regularise the letting and exchanging of breeding stock of the ‘New Leicester’ or ‘Dishley sheep’, a new breed created by intensive selection and inbreeding; 1843 when Gregor Mendel was admitted to the monastery of St Thomas in Brno (Brünn) and first came under the influence of Abbot Cyrill Napp (1792-1867). As a prominent member of the Moravian and Silesian Agricultural Society, Napp had made several published pronouncements on heredity, all of which arose out of debates on sheep breeding.

The word heredity (hérédité) was not in vogue before the nineteenth century. It is nevertheless convenient to use it since the ‘conceptual space’ it would later occupy was already recognised. Traits that distinguished one variety from another were referred to as hereditary (Marshall 1790, 1: p. 419). The transfer of such traits between generations was spoken of in terms of leaving or receiving an ‘empression’ (impression) or ‘stamp’ (Marshall 1790, 1: p. 326-7).

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1. THE BACKGROUND ‘Like begets like’ To the eighteenth century farmer heredity meant the property of living organisms by which offspring received the nature of their parents or ancestors. According to received wisdom it was associated in some mysterious way with the animal’s blood and also with its traditional environment. Differences between races (breeds), and also between individuals within a race, were explained ultimately by reference to accidents of development caused by influences over which the farmer had limited or no control. These might be changes of climate, vegetation or mode of life, or even dreams and passions, that deviated the generative process from its established course, that were degenerative.2 The responsibility of breeders lay in regulating the generative process by seeking to preserve or create an ideal type. It was a result to be achieved by a combination of selective breeding, environmental management and other aspects of good husbandry. No good breeder could take the matter of heredity casually.

Blood and seed The hereditary concept of blood, the essence of animal life, rested on the assumption that it must be transformed into semen (seed), the basis of new life.3 As the scripture said, each new being is “fashioned in flesh […], being compacted in blood of the seed”. 4 For many centuries the blood/ seed concept had no serious rival, and it was frequently associated with the idea that “the blood bore in it particles, originating throughout the entire body, that were gathered up by the testicles.”5 But what about the female, did she too produce seminal fluid or substance from her blood? By the eighteenth century, authorities were increasingly ready to postulate that the embryo was formed by the fermentation in the womb of seminal particles, produced by the blood of both sexes.6 The idea of inherited material drawn from all over the bodies of both parents, and united at conception, would be defined later by Darwin (1868) as ‘pangenesis’. Ernst Mayr has explained why this mistaken idea had such a long history: If one believes in the effect of use or disuse or any form of inheritance of acquired characters, as did just about everybody from Hippocrates to the nineteenth century, one is virtually forced to accept such a theory.7

2 3 4 5 6

7

22

For the basis of this idea see Roger (1997), pp. 460 f., quoting the works of Buffon and other eighteenth century authors. Brown (1987), p.184; Wood and Orel (2001), p. 47. Wisdom of Solomon 7: 1-2. Roger (1997), pp. 42-43, quoting Highmore (1651), chap. 5. The concept was presented in the 1730s in a printed booklet for sale in London, The Secrets of Nature revealed, or the Mystery of Procreation and Copulation considered and Explained, by ‘Michael Scott’, supposedly based on a thirteenth century text (Pinto-Correia 1997, p. 86). Mayr (1982), p. 636.

The Sheep Breeders’ View of Heredity (1723-1843)

Blood and locality There was also a strong belief that blood was linked with locality. “Every soil has its own stock” wrote the English estate agent William Pearce in his General Review of the Agriculture of Berkshire.8 The domestic animal could not select its own environment; it inherited it just as truly as it inherited its blood. A local breed’s characteristics depended for their reproduction on the continuity of local conditions.9 Problems with degeneration were expected when a breed was transported into a strange environment. Conventional wisdom insisted that the influence of a new climate and flora could not be escaped despite all efforts by the farmer to mitigate their influence by attempting to isolate his animals from external conditions, and to control mating. It had to be admitted that successful breeds were those in harmony with their environment. We have, at present, through time and the industry of our ancestors, various breeds; some of them adapted, though not perfectly, yet in very considerable degree, to the soil they are upon, and the purpose to which they are wanted […].10

Even so it became the practice to transport the best of sheep breeds more and more widely in the period under review, which allowed the theory to be tested. With experience, the intelligent farmer could establish the limits of natural environmental influence on heredity. Evaluating the connection between blood and locality was to prove a key issue in selective breeding, opening the way, step by step, to a greater understanding both of heredity and of adaptation to the outside world. An explanation was required about how external influences, interacting with the animal’s own ancestral nature, brought about continuity within a race in one circumstance but led to changes in another. The final conclusion drawn from these ‘natural experiments’ would prove to be reassuring, that when mating was controlled, adaptation (degeneration) took place much more slowly than the pessimists predicted.

Maternal impressions and other disputed influences on heredity Some modifying influences on the blood, and thus on heredity, were believed to be at their most effective at the moment of conception. An old and widespread myth, enjoying biblical authority, 11 held that visual impressions received by a mother (‘maternal impressions’) could affect aspects of the outward appearance of her young. Belief in the influence of maternal impressions on patterns and ‘birth marks’ in the new born, led to preventative actions by breeders, well into the nineteenth century.12 In this and other connections, experts in England as influential as W. Youatt were ready to claim wide influences on heredity through the power of the imagination:

8 9 10 11 12

Pearce (1794), p. 46; see also Young (1811), p. 6. The same was believed for varietal differences in plants (Müller-Wille, this volume, quoting Linneaus 1751). Youatt (1837), p. 494. Genesis 30: 39. Wallace (1893), p. 8, referred to by Wilson (1912), pp. 31-32.

23

Roger J. Wood

When in the higher species, the principle [of ‘like-produces-like’] may not at all times seem to hold good, it is because another power, the intellectual- the imaginative- somewhat controls the mere organic one […].13

Another mistaken belief lay in what August Weismann would later call telegony, also referred to as ‘infection of the germ,’ by which was meant the supposed influence of the male upon the female, in such a way as to affect her future offspring by other sires.14 Breeders also felt bound to consider the supposed relationship between healthy conception and the intensity of sexual activity, and whether this related to the degree of heat produced. The ‘hotter’ the male, the more potent he was supposed to be, i.e. more able to transmit his own characteristics. How to recognise ‘hotness’, in this special sense, was a matter of dispute. Concerning the determination of sex, there was a traditional belief in astral or planetary influences.15 Significance could also be attached to the weather, always a favourite topic among farmers. An ancient belief saw great significance in humidity and wind direction. Thus the seventeenth century agricultural pundit William Winstanley advised that “[i]f you would have your Yews [ewes] bring forth Ram-Lambs, then you must put the Ram to the Yew in the dry weather, and observe to drive the Yew towards the blowing of the North Wind.” 16

Nature and husbandry For the conscientious breeder no potential influence on heredity could be ignored except on the basis of proven experience. With so many unanswered questions facing him, he had to be ever watchful to note anything at all that seemed significant. The central issue was the extent to which the blood could be moderated by rearing, i.e. by the farmer’s system of management. As thus defined, rearing could include selective breeding, as well as the control of living conditions and the provision of special food to replace or supplement the natural diet. In this sense, rearing is obviously a more inclusive concept than that which Francis Galton would later define as “nurture”.17 Nature for the farmer was also a different concept from Galton’s, for it included some natural elements added after birth, the most obvious being the native flora (‘natural herbiage’ (sic)) that the animals would be ingesting and assimilating throughout life. Who knew how much 13 14

15 16 17

24

Youatt (1834), p. 522. Charles Darwin (Darwin 1868, pp. 403-4) made the concept scientifically acceptable by giving credit to an example from horses reported by Lord Morton (Morton 1821). The story was that a mare had born offspring by a quagga (a now extinct species related to the zebra) from South Africa and later produced horse colts which showed some striping. The fact that Darwin was prepared to believe the story suggests that the concept was accepted by the practical breeders with whom he was in contact. See also Davenport (1907), pp. 185-189; Burkhardt (1979). Roger (1997), p. 52. Winstanley (1679), p. 118. Galton did not accept that external nature (a change in conditions) had much impact on heredity during an individual’s lifetime. In Galton’s eyes nature referred principally to ‘those inborn or congenital peculiarities that were also congenital in one or more ancestors’ (Galton 1876, p. 329). He later added ‘the unforeseen appearance of ‘sports’ or ‘mutations’ of a kind not hitherto observed, but which, for all of that, may become hereditary’. (Galton 1908, chapter 21). For Galton, selective breeding was simply a manipulation of nature. Nurture referred to non-heritable modification of inherited “germs ” caused directly by the individual’s conditions of life, so that “the law of heredity goes no further than to say that like tends to produce like ” (Galton 1876, p. 338, my emphasis).

The Sheep Breeders’ View of Heredity (1723-1843)

this and other natural influences would affect the nature of future generations? Experience suggested that environmentally induced changes would become increasingly fixed in heredity the longer the animals were bred in that environment.

Blood and grading The most direct way for a farmer to obtain domestic animals of higher quality than those he already possessed, was to breed exclusively from blood stock of both sexes imported from wherever the best could be found. If this course of action had to be rejected, either on grounds of cost or caution (about possible degeneration), the alternative was to introduce superior blood by importing only males, to be crossed to native females and to their progeny for several successive generations. With each generation of such crossing, the proportion of superior (‘noble ’) blood would increase: 1/2 in the first generation, 3/4 in the second, 7/8 in the third, 15/16 (93.75%) in the fourth, rising to more than 99% in the seventh. This technique, known as ‘grading’ or ‘grading up’, based on a proportionate concept of heredity, as a fraction of the blood, probably had a long history, perhaps even back to Roman times.18 It had gained special significance in English racehorse breeding in the seventeenth and early eighteenth centuries, when great sums of money and aristocratic patronage rested on the breeders’ skill with Arab, Turkish and Berber livestock. 19

2. ALSTRÖMER ’S ACHIEVEMENT When Spanish Merino blood-stock finally became available to countries north of the Pyrenees, the grading technique found a new application. This was after Jonas Alströmer, who had risen from humble beginnings in Sweden to become a wealthy businessman in London with British citizenship, decided to import Merino sheep into Sweden directly from Spain. His growing knowledge of the wool trade convinced him that the best way ahead for Sweden lay in producing the finest wool in home territory, from the best foreign sheep. With a flash of genius and a good measure of optimism, he shipped into the port of Göteborg a small consignment of Merino sheep taken from their long established habitat in Spain. Landed in 1723, they were housed, like English racehorses, in clean, dry, well-insulated stables. Only in the warmest weather would they be let out. Despite early losses of stock, Alströmer had the faith to persist and, on the basis of further imports, to build up Swedish fine wool production into a thriving business, under Royal patronage. 20 He and his successors were able to demonstrate that even a radical change in the Merino sheep’s conditions of life was insufficient to cause degeneration of their wool when accompanied by good husbandry practices. Their satisfactory state of health demonstrated that they could attain a degree of harmony with their new environment. Shepherds were trained under Alströmer’s guidance in the best techniques to ensure that numbers of the ‘pure’ Merino type would expand year by year. A ‘mixed breed’ was also created, the product of grading crosses.

18 19 20

Anonymous (1811). Russell (1986), p. 86. Schulzenheim (1797), pp. 314-7; Wood and Orel (2001), pp. 126-7.

25

Roger J. Wood

Alströmer’s enterprise paralleled that of his compatriot and friend Carl von Linné (Linnaeus), striving to establish exotic plants in Sweden. By the time Alströmer was showing off his remarkable achievement to Linnaeus in 1746,21 he had flocks at various stages of improvement. He was hoping that those he had built up on the basis of crosses with Swedish sheep, would be better suited to local conditions and thus less expensive to maintain.22 Linnaeus deplored the xenophobia of Swedish farmers opposed to using the Merino stock for crossing. 23 The Swedish Merino experience inspired other countries to follow Alströmer’s example. For grading local Swedish sheep he recommended that none but ‘foreign’ (i.e. Merino) rams should be used for three generations, advice to be repeated many times by other writers, of various nationalities, in the coming years. Eventually, however, the number of generations the experts considered necessary to make a suitable transformation rose from three to four and then to five. 24 Even after five generations it proved necessary to practise selective breeding if stability was to be achieved. The risk of degeneration was ever present.

Degeneration and selective breeding The constant threat of degeneration presented a challenge to the best skills of breeders, out of whose successes there developed a growing conviction that control of rearing conditions and selective breeding were complementary techniques to oppose it. In harsh Scandinavian conditions, the type of Merino sheep that finally emerged at the end of the century was larger and stronger than the original Spanish imports but still yielded wool of excellent quality, equalling or even excelling that from direct Spanish imports.25 In parallel with the Swedish experience, although starting later, came the development of specialised Merino varieties in Prussia, Saxony, Austria/Hungary and France, each one with its own particular characteristics. How to maintain the stability of selected strains of Merino sheep, to yield high quality wool on a consistent basis, never ceased to be a matter of debate, as information was exchanged. Experience of grading German sheep with Merinos convinced J. H. Fink (1730-1807), the King of Prussia’s head bailiff, that fineness of wool was attributable more to breeding than to environmental influences.26 The point was noted with approval by the English farmer George Culley in the fourth edition of his book Observations on Livestock.27 Earlier published work had been inclined to stress the supremacy of environmental influences on wool, a belief that he and his mentor Robert Bakewell had already learned to question from their own experience.

21 22 23 24 25 26 27

26

Suneson (2001). Lasteyrie (1802); Schulzenheim (1797), pp. 307, 319-321; Kjellberg (1943), pp 304-5; Schulzenheim (1804), pp. 172-3; Culley (1807), p. 238; Rees (1819), section: Sheep; Martin (1849), p.61. In 1746, Suneson (2001), p. 15. Hastfer (1752a), (1752b), (1756); Daubenton (1782); Stumpf (1785); Fink (1799), p. 48; Parry (1806), p. 339; Tessier (1811). Schulzenheim (1797), pp. 315-6; Lasteyrie (1802); Rees (1819), section: “Sheep”. Fink (1797), p. 278, (1799), p. 54. Culley (1807), p. 260.

The Sheep Breeders’ View of Heredity (1723-1843)

3. BAKEWELL’S REVELATION Under the influence of a succession of breeders,28 it came to be recognised that the force of heredity could be strengthened and also directed, trait by trait. Certain traits seemed to be inherited together, others to vary independently. Taking the lead in this activity was Robert Bakewell who farmed at Dishley in Leicestershire. By paying exceptional attention to detail, he was able to regulate the process in a manner that Walton (1983) has described as giving selective breeding “a formal shape”, and to demonstrate publicly how rapidly a transformation could take place. Among his achievements was the creation of a new breed of sheep, known as the ‘Dishley’ or ‘New Leicester’, derived from rare individuals showing unusual or extreme characteristics (‘accidental varieties’).29 As the new strain was being perfected, individuals within it could be compared and evaluated as breeding stock according to the quality of their progeny.

Male and female No successful breeder from Bakewell’s time onwards was in doubt that both sexes contributed to inheritance,30 but less certainty was expressed about whether the two parents contributed equally.31 Of course it was easier to evaluate a good male than a good female because of the much larger number of progeny he could produce. However by the 1830s most British experts were in agreement with Youatt who stipulated, with direct reference to sheep, that “no certain degree of excellence can be attained unless the female possesses an equal degree of blood with the male.” 32 Both sexes contributed to inheritance of the same characters.

Individual trait selection, inbreeding and progeny testing Bakewell had startled the farming world in the 1770s and 80s with the inflated prices he could demand for his breeding stock. What made his highly selected Dishley sheep so specially valued was their rapid growth rates, associated with a unique body shape, designed to minimise unprofitable parts of the carcass and maximise joints of meat that would gain the highest prices. The difference in their appearance from traditional breeds was remarked upon with astonishment. It seemed that he worked like an artist who created an image in his mind and then transformed it into reality.33 His startling achievement depended on more than a proportionate, ‘holistic’ view of heredity such as underpinned the principle of grading. His was a ‘trait-based’ approach, one in which defined characteristics could be followed in families, sometimes independently, sometimes in association with one another.34 By taking practical advantage of such hereditary interactions, 28 29 30 31 32 33

34

Marshall (1790), 1: p. 338; Culley (1786), p.112; Trow-Smith (1959), pp. 45-69. Bakewell’s selective breeding of other domestic species is not being considered. Berry (1826); W. Youatt, writing as Lincolnshire Grazier (1833), p. 171; and Sir John Sinclair, President of the Board of Agriculture in London (Sinclair [1817] 1832, p. 97-8). Boswell (1829); see also Youatt (1834), pp. 523-4; discussion by Sinclair [1817] 1832, p. 97. Lincolnshire Grazier (1833), p. 238. Pitt (1809), p. 249; Somerville (1806); Wood and Orel (2001), p. 69. Despite the power of the artistic image in relation to Bakewell, it was not original. The same idea had inspired the Paris physician C. A. Vandermonde half a century earlier, in respect to the creation of new races of cats, dogs and horses, when he likened the breeder to a sculptor (Vandermonde 1756, p.155, quoted by Terrall 2002). Culley (1786), p. 186; Marshall (1790), 1: p. 298; Wood and Orel (2001), pp. 6-7.

27

Roger J. Wood

e.g. between small leg bones and rapid maturation,35 he could push his selection programmes forward without delay. As a natural accompaniment to selective breeding for growth and body shape, Bakewell carried out feeding experiments, by comparing different breeds side by side, to discover the best regime to maximise productivity at minimum cost. Because he was selecting intensively he was also inbreeding very closely, father to daughter, mother to son and brother to sister. His technique, given the name ‘breeding in-and-in’,36 had to be used with great care, for it ran the risk of concentrating deleterious traits as well as favourable ones. And just as the latter were to be preserved, the former had to be rigorously excluded. Bakewell’s own attitude on the matter was well known. Only animals in robust good health, as well as of the highest quality, judged by touch as well as eye, were considered suitable for breeding at this level. To minimise the potentially injurious effects of breeding in-and-in, Bakewell developed a progeny-testing programme in cooperation with fellow farmers who formed the Dishley Society in 1783. Through the interchange of stock, Bakewell and his friends were able to test their rams against a much wider variety of ewes than they could possibly have kept on their individual farms.37 Through his skill in defining economically significant traits, and employing the techniques of inbreeding and progeny testing, Bakewell became acknowledged as a true ‘Prince of Breeders’. 38 His Dishley sheep stock was used by many other sheep owners for crossing into almost every British breed, and many abroad, to introduce its quick fattening quality. 39

Heredity as a changing concept When fellow farmers were asked about Bakewell’s breeding philosophy, they would frequently stress his great faith in the old adage that ‘like produces like’: The simple observation that domestic animals possess a tendency to produce animals of a quality similar to their own was the groundwork of all Bakewell’s proceedings.40

By observing successive generations of his breeding stock under controlled conditions, he determined which traits were most strongly inherited. The result, as William Marshall reported, was that “a number of traits were found, in some considerable degree at least, to be hereditary.” 41 Bakewell’s confidence in heredity extended to unmeasurable or less readily defined traits, including resistance to bad weather, tolerance of poor food (“hard fare”) and even propensities to certain disorders. Visible traits of no economic value in themselves (‘nicks’ or ‘marks’) could be used as inherited markers of useful properties and propensities. 42 In no case could it be doubted that heredity arose from both sexes.43 35 36 37 38 39 40 41 42

28

Young (1771), p. 111; Wood and Orel (2001), p. 75. Marshall (1790), 1: pp. 300-301; Wood and Orel (2001), pp. 70-74. Pitt (1809), p. 256 ff.; Wood and Orel (2001), pp. 82-85. Young (1791). Young (1791); Trow-Smith (1959), pp. 66-9, 269-74; Walton (1983); Wood and Orel (2001), pp. 85-8, 145-6, 148. Berry (1826), quoted by Youatt (1834), p. 522. Marshall (1790), 1: p. 419. Wood and Orel (2001), p. 78; Culley (1786), p. 186.

The Sheep Breeders’ View of Heredity (1723-1843)

Population thinking All Bakewell’s actions tell us that his aim was to produce a breed in which the same characteristic traits would appear in every individual, despite the uncertainties of heredity. To be ‘well bred’ from good parentage was not enough. Breeding stock had to be selected carefully on the basis of ‘form’, i.e. the expression of desired traits, as well as blood (i.e. ancestry). This had to be done in every generation, supported by wise and consistent husbandry. Only then would it be possible to ‘concentrate’ the desired traits within the breed and avoid detrimental ones, with the necessary degree of efficiency.44 In his appreciation not only of parentage but also of form in a consistent environment, Bakewell avoided falling into the extreme hereditarian position taken by some other breeders of his day. Although the ancestry (blood) of an animal was of serious importance to Bakewell, particularly as his breed became progressively refined, its individual form had also to be considered and, more importantly, the form of its already existing progeny. It was on this broadest possible basis that every individual animal used for breeding, female as well as male, had to be selected most carefully.45 Through this course of action, successful breeders like Bakewell came to appreciate the significance of the whole flock as the ultimate target for improvement. Their attitude to breeding exemplifies what Ernst Mayr has defined as ‘population thinking’. He believes that breeders were the first group to gain an understanding of the concept. 46 Bakewell’s attitude on the matter was revealed in a conversation with Young, who quoted him as saying: The merit of a breed cannot be supposed to depend on a few individuals of singular beauty: it is the larger number that must stamp their character on the whole mass: if the breed, by means of that greater number, is not able to establish itself, most assuredly it cannot be established by a few specimens.47

To “establish itself” fully, in Bakewell ’s terms, meant that the breed had to prove successful commercially in competition with other breeds, particularly the one it replaced. 48 Furthermore, as Marshall pointed out, a breed had to be adapted to “the farmer’s climature, soil and system of management,” otherwise, as he stated, “if we reason from analogy, the improver appears to be setting himself up against nature, a powerful opponent.”49 How was opposition to nature to be avoided? Marshall had no doubt what Bakewell and his friends would answer: by carefully selecting all breeding stock against that particular environmental background. Only then could the highest levels of breeding excellence be achieved, when what we may call ‘internal’ and ‘external’ nature existed in harmony. “By this process […], the term blood became distinctively applied. When reference could be made to a number of ancestors of distinguished excellence the term blood was admitted.”50 43 44 45 46 47 48 49 50

Marshall (1790), 1: p. 481. Ibid., pp. 464-5. Lawrence (1809), p. 25. Mayr (1971), (1972). Young (1791), p. 570. Wood (1973), p. 238. Marshall (1790), 1: p. 464-5. Berry (1826), quoted by Youatt (1834), p. 52.

29

Roger J. Wood

Here the term is used in the sense of ‘full blooded’ or thoroughbred. The concept was becoming obvious to other farm animal breeders but Bakewell’s superior breeding success was reflected in the uniquely high prices paid (or claimed to have been paid) for his breeding stock.

4. AFTER BAKEWELL Races and species When breeders like Bakewell began to compare races native to different environments, side by side at a single location but isolated reproductively, they noted that they remained distinct for as many generations as there was time to observe them. The obvious conclusion was that any environmental effect on heredity must be slow and gradual. How then had the different races 51 come about? The influence of earlier human selection was a clear possibility. Another explanation to be considered was that races/varieties were natural divisions, like species, incapable of merging permanently into a single type.52 William Marshall, the prominent agricultural writer from Yorkshire, felt bound to consider the idea: Whether in the Animal Kingdom varieties are altogether accidental or artificial, or whether they are not, or have been originally natural subdivisions of species, would, with respect to domestic animals be now difficult to determine.53

In his own opinion, which he believed to be “supported by naturalists,” varieties arose “by climature [sic], soil, accident or art, under the guidance of reason or fashion, during a succession of centuries.” Marshall’s conclusion, supported by a mass of breeding evidence, was significant as a reaction against the widely held opinion by naturalists that members of a species differed only in superficial, non-essential characters.54

Inbreeding and prepotency Farmers recalled Bakewell ’s attitude to inbreeding by quoting his supposed dictum that “inbreeding gives prepotency and refinement.”55 “Prepotency” was the ability of certain individuals to pass on their traits with extra certainty. Inbreeding was seen to increase potency and thus accelerate the progress of improvement. It was recognised to be a result of hereditary stability, and thus as a “concentration of the blood.” ‘Concentration of the blood’ became a breeding maxim demanding an explanation from science. In a pamphlet published in 1812 the surgeon John Hunt, a supporter of the ‘Dishley System’, deplored the idea of blood being the actual vehicle 51 52 53 54

55

30

The terms ‘race’, ‘variety’ and ‘breed’ seem to have been used interchangeably by animal breeders at this time. Home (1776), p. 309. Marshall (1790), 1: p. 462, his emphasis. Mayr (1972). Referring to British cattle, Darwin would later write: ‘A large part of the difference [between domestic races], no doubt, may be due to descent from primordially distinct species; but we may feel sure that there has been in addition, a considerable amount of variation’ (Darwin 1868, 1: pp. 86-87). Earlier, in a letter to A.R. Wallace in 1857, he had placed the emphasis in the opposite direction (Darwin 1887, 2: pp. 95-96), as Marshall had done in 1790. Lush (1951), p. 501.

The Sheep Breeders’ View of Heredity (1723-1843)

of heredity as “far exceeding the laws of nature”. He agreed with the Merino breeder, Dr Parry of Bath, that “the word blood is nothing more than an abstract term expressive of certain external and visible forms which from experiment we infer to be separately connected with those excellencies which we most covet.” Similar sentiments were expressed by other breeding experts at the time.56 All that could be said for sure was that blood, in the hereditary sense, was divided between the parents in proportion. Thus a son mated to his mother would cause her to produce lambs with six parts of herself and only two of his father.57 As noted earlier, the same argument was used to calculate the proportion of high (noble) blood in grading crosses. By 1800 two traditions in animal breeding had been established: 1) the grading technique with its proportionate view of heredity; 2) selective breeding from accidental varieties, usually isolated from the progeny of controlled matings, sometimes between members of different geographical races. On the basis of experience, breeders were guided to success by a number of valuable breeding maxims, but even the deepest thinkers among them could suggest no causal explanation for heredity. Dr James Anderson, a Scottish farmer and scholar, who did business with Bakewell, 58 tried to face the issue in some of his writings but had to confess it to be a mystery, both in “origin and perpetuation”.59 For practical purposes, the priority for a breeder in the Bakewell tradition was to possess breeding stock with a recognised capacity to transmit desirable traits as surely and certainly as possible, through either sex. Theoretical explanations could wait till later.

The Bakewell approach to breeding is evaluated A number of breeders in Britain benefited from Bakewell’s example although by no means everyone who wished. For those with only a shadow of his judgement, and without the equivalent of a Dishley Society to evaluate their breeding stock, inbreeding could bring drastic penalties. Potential problems included loss of fertility and various constitutional weaknesses and disorders not easily eliminated. As more and more breeders jumped onto the inbreeding bandwagon, British experts like Sir John Saunders Sebright and Sir John Sinclair recommended caution. 60 Too many breeders were finding that close inbreeding “impairs the constitution and affects the procreative powers.”61 The care Bakewell had taken to avoid these deleterious effects by evaluating rams and exchanging stock with other Dishley breeders was widely recognised. Bakewell’s original and unusually systematic approach to breeding was also being evaluated on the other side of Europe. His combination of techniques was being taken most seriously in relation to fine wool production, even though his own interest did not lie particularly in that direction. By the end of the century the technical term ‘breeding in-and-in’ had entered the French, Austrian and German breeding literature, either in English or in translation. 62 In a German textbook on animal improvement, published in 1785, the author, Christian Baumann 56 57 58 59 60 61 62

Parry (1806); Lawrence (1800), p. 44. Sebright (1809). Pawson (1957), p. 106. Anderson (1799), p. 87; see also Marshall (1818 ), 1: p. 43. Sebright (1809); Sinclair ([1817] 1832), pp. 93-5; Blacklock (1838), pp. 106-7; see also Knight (1799). Berry (1926), quoted in a footnote to Youatt (1834), p. 526; Berry (1829). Fink (1799), p. 73; Bourde (1953), p. 145; Klemm and Meyer (1968), pp. 139, 179; C.C.André (1804); Thaer (1804); Wood and Orel (2001), pp. 140,157,164,167, 212, 215, 231.

31

Roger J. Wood

(1739-1803), a Cistercian monk, who spent most of his life in Würzburg, wrote in favourable terms about a fellow Bavarian who was evaluating his rams in conscious imitation of the famous Englishman. He was referring to the “talented economist” von Bori (Borie) who farmed at Neuhaus, close to Bad Neustadt. “Mr Bori hired his rams to his neighbours to evaluate their offspring, following the example of Bakewell.”63

5. SHEEP BREEDING IN MORAVIA Towards the end of his life, Baumann moved to Moravia where he produced a new textbook on agriculture published in 1803.64 Describing fine wool production in Central Europe, he picked out for special mention the estate of the noble Ferdinand Geisslern at Ho £tice in Moravia.65 Baron Geisslern made a practice of exchanging rams with neighbouring landowners, mating ram with ewe in carefully controlled matings.66 By that time Geisslern had attracted sufficient notoriety to be called the “Moravian Bakewell.”67

The Brno Sheep Breeders’ Association Most of what we know about sheep breeding in Moravia comes from the proceedings of an “association” for sheep breeders, created in Brno, the major Austrian centre for wool cloth production, in 1814, as a section of the Moravian and Silesian Agricultural Society (AS). The Association of Friends, Experts and Supporters of Sheep Breeding (hereafter abbreviated to Sheep Breeders’ Association or SBA) was set up to discover “incontrovertible principles to ensure favourable results in sheep improvement.”68 Meeting annually for nearly thirty years, it stimulated numerous publications, written or edited by its secretary C. C. André. Unique in Europe not only for its length of existence, but for having among its membership representatives of every profession with an interest in the wool business, the SBA drew together progressive individuals from throughout the region, from Silesia, Bohemia, Hungary and Austria, as well as from Moravia itself. Several members had extensive knowledge of foreign breeding literature, including a mass of English books and journals through access to major castle libraries. 69 The 63 64 65 66 67 68 69

32

Baumann (1785), pp. 217, 273. Wood and Orel (2001), pp. 157-8, 211-2. Baumann (1803), p. 706. Köcker (1809); ‘K in Mähren’ (1811); C.C. André (1812); R. André (1816); Wood and Orel (2001), pp. 231-2. Köcker (1809). R. André (1816); Wood and Orel (2001), p. 229f. One of these was Count H. F. Salm-Reifferscheidt (1778-1836), first President of the AS and a leading innovator in industrial textile production. His strong British connections included his marriage to a Scot. At his castle at Rájec (Raitz), a few kilometres north of Brno, the Count had a library of 59,000 volumes, rich in works on natural science and technical subjects, to which C.C.Andre, secretary of the SBA, had full access from 1811-1820, as Salm’s economic advisor (Wirtschaftrath). Another member well placed to communicate foreign ideas was Count Imre (Emmerich) Festetics (1769-1847), from a Hungarian family with an exceptionally rich agricultural library owned by his brother Count György Festetics (1755-1819) at Keszthaly on Lake Balaton (Kurucz 1990). The library contained most of the major English agricultural works, including publications by Young, Culley, Sinclair and Marshall and, most importantly, the County Surveys produced by the Board of Agriculture in London. When dealing with animal breeding, these publications gave a major emphasis to Robert Bakewell.

The Sheep Breeders’ View of Heredity (1723-1843)

secretary’s son, R. André, produced a practical manual, Instructions for the Improvement of Sheep, published in Prague, based directly on Geisslern’s methods.70

The Brno inbreeding debate By 1817 an area of disagreement was developing within the SBA about the value of inbreeding as a means of ‘fixing’ traits, as a route to more constant inheritance. On the assumption that the continuity of heredity was determined partly by an “inborn component (theils angeboren)” and partly “by rearing (durch Erziehung)”71, the question resolved itself into whether, and how, inbreeding affected these two influences. A leading expert from the Vienna region Baron J. M. Ehrenfels (1752-1843) called for caution, claiming that the pairing of nearest blood relatives must lead to “natural climatic degeneration (natürliche klimatische Rückbildung)” by disrupting “the principal plasma of the animal’s organisation (Hauptplasma der thierischen Organisation)”.72 We may note how Ehrenfels linked inconstancy of inheritance with greater sensitivity to climatic influences because of a supposed physiological disturbance. Reporting on the debate, the SBA’s secretary, C.C.André, was sure that deeper study would show consanguinity to have predictable consequences, behaving according to some “physiological, natural law”, knowledge of which could bring benefit: What subtle problems are here to be solved before we can approach nearer to the truth [about inbreeding] with confidence. Here we are penetrating into the innermost secrets of nature.73

His son Rudolf had already suggested, in his Instructions (1816), that the answer lay in regulating the intensity of inbreeding according to the degree of improvement already achieved. He claimed this to reflect Geisslern’s own views on the subject, i.e. that the more “noble” the stock, the more important it becomes to improve such animals “purely and simply through each other.” 74 Bartenstein, the SBA’s president, wrote that he was convinced that inbreeding need not produce the ill effects that Ehrenfels feared, if members would follow the Bakewell principle of assessing the value of an animal not simply for its own qualities but for those of its parents and descendants.75 As Bakewell had shown, an exchange of breeding stock of the same highly selected race, to ‘progeny test’ the males, also helped to prevent unwise inbreeding. A regular exchange of rams between stockowners had been a feature of the SBA from the beginning. Like Bakewell, the best breeders among SBA members were ready to adopt “population thinking” when dealing with selected stock, female as well as male, in order to produce the highly uniform product the cloth manufacturer demanded. As R. André had written in his Instructions: No animal should be perceptively better or worse than the others, particularly in wool. In this way one should work from the beginning of the improvement programme.76

70 71 72 73 74 75

R.André (1816); Wood and Orel (2001), pp. 197-204. Festetics (1819a). Ehrenfels (1817). C.C.André (1818), original in German; see also Wood and Orel (2001), p. 235. R. André (1816), pp. 41-42, 94-96. Bartenstein (1818).

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‘Genetic Laws of Nature’ C.C.André persisted in pressing for a compromise and ultimately a law (or laws) of breeding to guide them. Writing in 1818 on the effectiveness of the Agricultural Society, he identified a number of outstanding questions concerned with the supposed ‘weakening’ effects of inbreeding, in particular on wool fineness and susceptibility to disease. He was concerned about how to retain the characteristics of a pure breed without reversion while still retaining the potential for improvement. Count Festetics, a Hungarian member, stated his opinion that the potential for weakness was present within any non-inbred race before inbreeding took place, from which he concluded that weakness, if it occurred, was created not by the inbreeding process but by incorrect selection.77 Sebright in England had made the same point earlier although he did not believe that breeding-in-and-in could be continued indefinitely without detriment. 78 Festetics summarised his own 15 years of experience of inbreeding under the heading “genetic laws of nature (genetische Gesetze der Natur)” in four parts.79 Here he (1) associated pure “noble” breeding with good health, (2) appreciated that inherited traits could be recessive for one or more generations, (3) recognised the extent of variation, even within seemingly pure breeds, (4) stipulated that the precondition for applying inbreeding safely must be scrupulous selection of stock animals. As an aid to selection, R. André had refined a scheme to evaluate wool into seven grades, using a newly developed micrometer.80 Festetics greeted André’s “mathematical” evaluation of wool quality with enthusiasm: “It will be judged as marking the beginning of an epoch in the science of breeding that in 1819 the grades of wool fineness were established and defined with mathematical precision.” 81 In reflection of the successes achieved by Geisslern and his followers, the SBA continued to recommend close inbreeding for the totality of its existence (1814-1845). A later member J.K. Nestler (1783-1841) defended the practice on the basis of English experience when he wrote: “Without this technique [breeding in-and-in] Bakewell could never have existed nor could one exist in the future.”82

The pure race concept and chance deviations One of the lessons of experience from the SBA was that sheep derived from several generations of grading crosses, what the breeders called ‘noble sheep stock (edles Schafvieh)’, did not breed as true as those descended from uncrossed imported sheep, the ‘pure noble race stock (reines edles Racevieh)’. Achieving the noble state by grading was evidently far from straightforward. Even when the sheep looked externally the same as the pure breed, they proved to be less consistent in breeding. In his Instructions, R. André wrote of how nobility was a relative matter, depending on ancestry. Optimistically he stated that the traits of the original pure breeding stock imported from Spain “remain constant even when external conditions are unfavourable for their preservation.” 83 76 77 78 79 80 81 82

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R. André (1816), p. 37, original in German; N.B. the population concept is here restated; see also Wood and Orel (2001), pp. 201-2. Festetics (1819a). Sebright (1809), p. 11. Festetics (1819b); Orel and Wood (2000); Wood and Orel (2001), pp. 237-8. R. André (1819). Festetics (1820), p. 33, original in German. Nestler (1839), original in German.

The Sheep Breeders’ View of Heredity (1723-1843)

Like other breeders of his time, he pictured heredity in terms of the ‘strength’ of an animal’s internal nature, represented in its blood. If blood was the factor by which to judge a pure noble race, a major question for sheep breeders was whether a graded noble stock could be converted into a ‘pure noble’ one. Rudolf André was sure that Geisslern had proved it possible with fine wooled sheep in Moravia, based on Merino crosses, and he described the process: With care and attention a merely noble flock can be raised to the pure race if one refrains from mixing alien bloods and, through an appropriate control of pairings, brings together specific characteristics of body build and wool, to be transmitted to the progeny, and preserved to the same degree. In this way something constantly unique (constant originelles) arises, something fixed in the organisation of these animals, something derived totally and exclusively from pure blood relatives (aus lauter Blutsverwandten hergeleitete), which is characteristic of the lineage.84

This then was the secret, as it had been before to Bakewell and his followers, to match the parents for their traits, to practise rigorous selection and to fix the type by inbreeding. Individually controlled matings (Sprung aus der Hand) were the answer. Even so, as Bakewell had shown earlier, racial stability could never be absolute. The Moravian experience revealed that selective breeding was required even in the ‘pure noble race’. Constancy of inheritance could only be maintained by matching the best rams to their close female relatives, each ram forming “a sire’s family”.85 Even an apparently “genetically fixed race (genetisch befestigte Rasse)” was expected to degenerate. It could not be doubted that eventually a reversion (Rückschlag), a chance deviation (Natursprung) or freak of nature (Spielart, Naturspiel) would appear and multiply, and would have to be removed to regain the original “principal racial form (Hauptsgeschlechtsform)”.86 A potential existed for genetic changes to occur which in those days were attributed to a direct influence of the new environment in one of its many aspects. Hence the firm rejection by the SBA of the popular idea of inherent racial constancy, proposed by Justinus (1815) with reference to horses, that the noble race stays for ever noble when the purity of the blood is maintained, (i.e. without the need for persistent selective breeding). The concept was simply not justified by the experience of Moravian sheep breeders.

Improvement of the pure race Sometimes a chance deviation could be advantageous, by providing an opportunity to make an improvement to the race, to add to its economic value. Selection would then be directed towards making such favourable accidental varieties endure (zufällige Varietäten bleiben zu machen).87 It was a strategy that had its roots with Bakewell and his followers in the British Isles. Marshall had referred to Bakewell as having used accidental varieties as the basis of his selection of long horned 83 84 85 86 87

R. André (1816), pp. 5-6. Ibid., pp. 6-7, original in German. R. André (1816), pp. 6-7; C.C. André (1804); Petersberg (1815); Nestler (1836); Stieber (1842), pp. 4144; d’Elvert (1870), 2: pp. 48-49; Wood and Orel (2001), pp. 195-6, 244-5, 268. ‘Irtep’ (1812); Wood and Orel (2001), pp. 224-5. ‘Irtep’ (1812); R. Andre (1816), pp. 94-96; Nestler (1829); Wood and Orel (2001), pp. 224, 243.

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cattle: “solicitously seizing the superior accidental varieties produced.” 88 The possibility of exploiting favourable variants implied that there could be no fixed limit to the improvement of a race, taking into account the “predispositions for a higher perfection (die Anlagen für eine höhere Vollkommenheit)”.89 The possibility of such improvement was becoming evident all over Europe where different forms of so-called pure Merino stock were established, varying in size, degree of skin wrinkling and length of staple. To increase the efficiency of selective breeding, the SBA recommended that all animals in a flock should be numbered, with all parents and their progeny recorded, as a standard procedure.90 After 1819 when C.C. André’s position was made difficult because of political opposition, requiring him eventually to move to Stuttgart, the discussion of breeding theory and heredity in the SBA lapsed for a brief time. It was revived by J. K. Nestler who occupied the Chair of Natural History and Agricultural Science at the Moravian University of Olomouc (Olmütz), from 1823 until his death in 1841. Nestler was the first in the Society to write about heredity (Vererbung).91 He noted that resemblance between generations was most certain when parents were of the same sort (Art), and he also made clear that defects and weaknesses could be inherited. 92 Other active new members included the lawyer and estate manager F. J. Teindl and the recently appointed abbot of the Monastery of St Thomas, C. F. Napp. Together with Bartenstein and Ehrenfels, they took the discussion of sheep breeding and heredity into fresh areas.

6. NAPP’S SCIENTIFIC CURIOSITY Abbot Cyrill Franz Napp (1792-1867) was a dignified scholar, expert in Oriental Languages and the Old Testament, but also an energetic administrator with a shrewd sense of business and a strong interest in the useful applications of science. A contemporary wrote that he was “the supporter of every scientific effort.”93 His interest in breeding was long-standing and broadly based,94 for although he took a particular satisfaction in plant breeding, it was sheep that provided the major source of income from the extensive monastic lands. The results of new approaches to breeding fired his intellectual curiosity, and made him determined to make practical use of them. Within a year of his arrival in Brno, he had been elected a member of the Agricultural Society (1825); two years later he was on its organising committee. Later he would become its president. By the time Mendel entered the monastery in 1843, advanced breeding was well established on efficiently managed monastic estates. In the SBA, discussions had recently been held on the nature of heredity, in which Napp had taken a leading part. 88 89 90 91

92 93 94

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Marshall (1790), 1: pp. 383-4; see also Anderson (1799), p. 23, referring to variations ‘accidentally produced’ being inherited, as well as the ‘general characteristics of the parental breed’. R. André (1816), pp. 95; Ehrenfels (1837); Wood and Orel (2001), p. 257. Köcker (1809); R. André (1816), pp. 7, 35; Petersberg (1816), p. 113; Wood and Orel (2001), pp. 194, 2002, 232-3. McLaughlin (this volume) has pointed out that Vererbung in a biological sense was used by Immanuel Kant in 1793, who applied it not just to individual peculiarities but to sub-specific varieties that bred true, as in human races. McLaughlin notes that Vererbung takes the perspective of the donor and might best be translated as ‘bequeathment’. Nestler (1929). Rohrer (1830). Orel (1975b); Orel and Wood (2000b).

The Sheep Breeders’ View of Heredity (1723-1843)

Inner and outer organisation SBA experience had shown beyond doubt that constancy of “type (Art)” in a sheep, which was believed to reside in its “internal organic structure”, was more influenced by selective breeding than by favourable conditions.95 In one of the debates Napp had commented that when pure breeding was the aim, it was important not only that parents should be of the same type, but that they should also correspond in “both inner and outer organisation”: The highly esteemed Abbot and Prelate of St Thomas’s Monastery, Cyrill Napp, asserts that, according to his view, heredity of characteristics from the ‘engenderer’ (Erzeuger) to the ‘engendered’ (Erzeugten) consists above all in the ‘mutual affinity by kinship’ (gegenseitige Wahlverwandschaft) of paired animals. As a result of this, a ram chosen for the ewe should correspond to it in both inner and outer organisation. This process deserves to be the subject of an important physiological study.96

The use of the term “inner and outer organisation” does not appear to be a reference back to traditional ideas about separate influences coming from the two sexes, because the ram and ewe are expected to correspond in both respects. Superficially it might seem to recall a statement made thirty years earlier by the English surgeon Henry Cline, that “the external form [of a sheep] is only an indication of the internal structure.”97 However, Napp’s words are probably more in tune with those of B. Petri expressed in a lecture to the 1837 meeting of German Naturalists and Physicians in Prague. Speaking on the subject of animal breeding, he attached particular significance to selection from crosses, i.e. blending (Vermischung), “where the inner cohesion of the external formation of the individual in its different varieties has to remain hidden from the eye.” 98 Neither Petri nor Napp seem to be assuming that the inner organisation is determinable simply by examining the animal’s outward appearance. Napp sees the relationship between inner and outer organisations as a problem for physiological study.

Inheritance capacity Among the mysteries of heredity to be investigated was the observation of differences in the capacity of individual animals to transmit their traits. On this basis British breeders had coined the term ‘prepotent’ for the most reliable males for breeding. Under the influence of Nestler, members of the SBA became convinced of the importance of determining the ‘inheritance capacity (Vererbungsfähigkeit)’ of each individual used for breeding, measured as the strength of transfer of inherent characteristics from that individual. Professor Nestler believes that the most essential matter of all, as well as the most pressing question at this time, in relation to improved sheep breeding, is the ‘inheritance capacity’ of noble stock animals.99

95 96 97 98

R. André (1816); Ehrenfels (1831); Wood and Orel (2001), pp. 225-6. Teindl et al. (1836), quoting Napp, original in German. Cline (1805); Sinclair ([1817] 1832), p. 87. Petri (1838).

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Could this capacity be enhanced by inbreeding? Nestler (1839) asked. His conviction about the significance of inheritance capacity stimulated Napp to bring discussion back to the subject in relation to what might be its physiological basis: Prelate Napp continued the course of the continuing debate with rigorous brevity…and drew attention to the fact that they had completely deviated from the proper theme of inheritance capacity. ‘What we should have been dealing with’, he pointed out, ‘is not the theory and process of breeding. But the question should be: what is inherited and how?’100

Napp’s words, briefly reported, do not allow us to distinguish between the inherited traits themselves and what lies behind them, in terms of inner organisation. However the context makes clear that he was reacting against an inductive approach to investigating heredity, based on examinations of existing sheep breeding records. Attempts by the breeders to trace patterns of inheritance from their stock registers were proving incapable of solving the problem. It seems clear from Napp’s various reported statements that he is hoping that further understanding of hereditary transmission will result from new developments in physiology.

Crossing/hybridisation At a time in biological history when the boundary between race and species was not always clearly defined, the progeny of crosses between races/varieties could be referred to as hybrids. 101 It had long been recognised that whereas the first generation from an inter-racial cross (hybridisation) could be quite uniform, and often more or less intermediate between the parents, the second generation could be highly variable and often unpredictable: “It is a universal property of hybrids that in their progeny there appear traits reminiscent of the parental forms with great variability.”102 The breaking down of essential characteristics in the progeny of hybrids was a strong argument against hybridisation beyond the first generation, and in favour of grading a ‘common’ flock with a ‘noble’ race to ‘fortify’ or ‘fix (befestigen)’ noble traits in the progeny. The supposed long history of the major races advised a conservative approach to breeding from hybrids between them. The fact remained, however, that many highly bred races could be crossed and it was intriguing to discover whether something economically advantageous might arise from their progeny. Instructive examples of crosses between highly bred races were being reported from the British Isles where farmers competed for access to Dishley stock to cross to their local breeds, their aim being to develop new breeds combining the best characteristics of both parent races. The derivation of one of these, the ‘Improved Cotswold’, has been traced by Walton (1983), whose analysis of breeding stock sales reveals the first example of the new breed in 1811.

99

Teindl et al. (1836), original in German; see also Bartenstein (1837); Bartenstein et al (1837); Wood and Orel (2001), pp. 246-7. 100 Bartenstein et al., original in German. 101 As by Mendel (1866). 102 Elsner (1826), original in German; see also Wood and Orel (2001), p. 251.

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The Sheep Breeders’ View of Heredity (1723-1843)

Thinking along similar lines, SBA members recognised that hybrid progeny provided the basis for selection in new directions, e.g. to improve wool, body form and meat quality together, or to associate superior wool with a high fleece weight.103 The first of these two aims was claimed to have been achieved in Moravia before 1811, by Geisslern.104 Later there were crosses made between the Negretti and Escurial races of the Merino, to create the highly successful EscurialNegretti Vollblut race in Austrian Silesia. Partly based on Geisslern’s stock, it was eagerly purchased by certain Australian colonists and greatly admired by the Prussia-based breeding expert Hermann Settegast.105 Attempts to analyse patterns of heredity in crosses were strongly encouraged by Nestler but were largely unsuccessful.106 All that could be concluded was that any two sets of parental characters, considered as a whole, appeared to “blend (verschmelzen)” in the progeny.107 Nestler led the way in calling for specially designed experimental crosses. 108 He and other enlightened members of the SBA, including Napp in particular, hoped that an understanding would be reached concerning the actual basis of heredity, not only its principles and rules but its physiological basis, and that such knowledge would aid in breed improvement. 109 However, as Nestler admitted, most breeders were still stumbling in the dark.

Co-operation with plant breeders The Sheep Breeders were not the only group to react to a commercial stimulus. A Pomological and Oenological Association (POA) was established in Brno in 1816. Instigated by C. C. André, it drew inspiration from British achievements in producing new varieties of fruit by controlled pollination,110 reported by T. A. Knight (1759-1838), president of the London Horticultural Society, and on the application of hybridisation in creating new varieties of cereals, reported by the secretary of the Pomological Society of Altenberg (near Leipzig), C. C. L. Hempel. 111 An article by Hempel published in 1820 included a call to uncover “the law of hybridisation” 112 Sedlá™ek von Harkenfeld, second president of the POA, published an article on inter-varietal crosses based on his personal experience with vine hybridisation. He expressed his expectation that new varieties derived in this way would be capable of producing better quality wine than that from any known varieties, “even from abroad”.113 The membership of the POA and SBA overlapped, with Napp highly active in both of them from 1825 onwards. Under his influence Franz Diebl was called to the Chair of Agricultural Science at the Brno Philosophical Institute. Nestler, and others proposed that further enlightenment about heredity in sheep would be gained by swapping information with plant 103 104 105 106 107 108 109

Bartenstein, reported by Löwenfeld (1835). ‘K in Mähren’ (1811); R. André (1816), p. 7; Wood and Orel (2001), pp. 185. Settegast (1861); Wood and Orel (2001), pp. 184,185, 188-9. Nestler (1837). Nestler (1829), pt. 1. Nestler (1837); Orel and Wood (2000); Wood and Orel (2001), pp. 252-9. Nestler (1839) and (1841), p.337, quoting Napp; Bartenstein (1837), quoting Napp; Teindl et al. (1836), quoting Napp; Wood and Orel (2001), pp.246-7, 258. 110 See ‘Irtep’ (1812); Wood and Orel (2001), p.24. 111 Hempel (1818); Orel (1978); Mylechreest (1988); Anonymous (1816). 112 Hempel (1820); Orel (1974). 113 Sedlá™ek (1826); Wood and Orel (2001), pp. 239-40.

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breeders, placing faith in “the power of science”. Also recommended were contacts with naturalists, anatomists and physiologists, in order to put sheep breeding on a “rational” basis. 114

The physiological basis of heredity By 1843 when Mendel entered the monastery, it was accepted in Moravia that heredity was brought about by the meeting of two parental ‘germs’, located in the egg of the female and the semen or pollen of the male.115 The subject was still wide open for investigation if a way could be found. It continued to puzzle Napp that “nothing certain can be said in advance as to why production through artificial fertilisation remains a lengthy, troublesome and random affair.” He remarked on a problem that all breeders faced when attempting to produce new varieties by artificial fertilisation, the significance to be attached to mere chance. 116 The chance element demanded to be understood and, if possible controlled. Napp’s statement of 1836 that the process of heredity “deserves to be the subject of an important physiological study,” 117 has obvious significance in the light of further events, instigated by his dependant Mendel.

7. SUMMARY The traditional concept of heredity as functioning through the blood, which was converted into semen (seed) at the start of each new generation, had logical consequences in relation to what was supposed to affect genetic continuity. Any change in an animal’s physiological state, for whatever reason, was perceived to have a good chance of influencing either the blood itself or its seminal derivative. A shift of emphasis, leading to a ‘harder’ view of heredity, came as a result of practical experience when breeds were moved over long distances into new locations, to be maintained under carefully controlled conditions of housing and nutrition. This happened with Merino sheep transported north of the Pyrenees, first to Sweden in 1723 by Jonas Alströmer, later to Saxony, Prussia, France and Austria. Such economic experiments established the extent to which breed characteristics might be maintained in the face of nutritional and climatic pressures. With growing confidence that external pressures could be contained, breeders began a trait-based approach to selective breeding, which further established the limits within which breed characteristics were able to vary. The most adventurous among them found ways to exploit some aspects of variability to create new breeds, even exceeding the best traditional races in economically important characteristics. Even so, their selected breeds had to exist in harmony with (be adapted to) the natural environment. Strictly regulated, trait-based selective breeding at an intensive level introduced the potential for inbreeding depression, which had to be avoided without disrupting progress towards the improvement desired. The answer was found to lie in interchanging stock with other farmers, and selective breeding for fitness traits, as well as those of specific economic interest. Fitness was assessed by progeny-testing which, to be efficient, required the organised co-operation of farmers willing to enlarge their knowledge about heredity. Robert Bakewell and fellow members of the 114 115 116 117

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Nestler (1829); Diebl (1839); Fraas (1852); Orel (1975a) and (1978); Wood and Orel (2001), p. 255. Purkyne (1834). Nestler (1841), p. 337, quoting Napp. Teindl et al. (1836).

The Sheep Breeders’ View of Heredity (1723-1843)

Dishley Society, formed in 1783 in England, were pioneers in this respect. Progeny-testing revealed that certain males, referred to in Britain as ‘prepotent’, were better at transmitting desired traits than others. In Moravia interest developed in the ‘inheritance capacity (Vererbungsfähigkeit)’ of noble stock animals. Nestler speculated on whether this capacity could be enhanced by inbreeding, just as Bakewell claimed with respect to prepotency. Inter-racial crosses took two forms. From the beginning of the eighteenth century the grading technique was much in evidence, being the means by which a local breed was improved by successive generations of crosses to males of a superior one. With growing experience the first attempts were made to combine the qualities of two superior breeds by selecting among the progeny of their hybrids. In practice it proved very difficult to select the right combination of traits among the great array of possible types that segregated in the second and subsequent hybrid generations. However there were some significant successes, including the Escurial-Negretti Vollblut Merino race in Silesia. Unpredictable variability observed in crosses confirmed in the minds of breeders the difficulty of formulating any general law of heredity. Even so the practical advantage of discovering such a law drove them to further effort, and encouraged animal and plant breeders in Moravia to join together to see if they could resolve the problem by co-operation. By 1843 when Mendel arrived on the scene, discussions on this important topic had been intensively pursued in the Brno Sheep Breeders’ Society for many years, with Abbot Napp recently playing a leading part, but had reached no obvious conclusion. It seemed that although heredity could be manipulated by selective breeding, following certain defined procedures, there was no escape from the element of chance invisible in the germ.

Acknowledgements I am greatly indebted to my friend and collaborator Vítezslav Orel for his enthusiasm, encouragement and wise counsel during the past 30 years, and for his generosity in providing me with photocopies of early published material from Moravia, unobtainable in England. The paper has been slightly amended since it was presented at the MPI Workshop.

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Abbreviations Folia Mendeliana. Published yearly by the Moravian Museum in Brno since l966. Issues l-9 appeared as special volumes. Issues 10-20 were published as a supplement to Acta Musei Moraviae, sc. nat. with the pagination of these volumes. The part Folia Mendeliana were also reprinted separately. Folia Mendeliana Volume 21 and further volumes are being published again as special volumes with their own pagination. Hesperus. Belehrung und Unterhaltung für Bewohner des österreichisches States - Hesperus, Prag. Mittheilungen. Mittheilungen der k.k.Mährisch-Schlesischen Gesellschaft zur Beförderung des Ackerbaues, der Natur- und Landeskunde in Brünn. ONV. Oekonomische Neuigkeiten und Verhandlungen, Prag PTB. Patriotisches Tagesblatt oder öffentliches Correnspondenz- und Anzeiger-Blatt für sämtliche Bewohner aller kais. köng. Erbländer über wichtige, interessierende, lehrreiche oder vergnügende Gegenstände zur Beförderung des Patriotismus, Brünn.

References Anderson, J. 1799. “An inquiry into the nature of that department of natural history which is called varieties.” Recreations in Natural History 1: 49-100. André, C.C. 1804. Cited in: K in Mähren [Köller, M.] (1811), and republished by d ’Elvert (1870), Volume 2, 145-152. _____. 1812. “Anerbieten, Gutbesitzern auf dem kürzesten und sichersten Wege zur höchsten Veredlung ihrer Schafherden behülflich zu seyn.” ONV. 181-183. _____. 1818. “Terminologie für Woll-industrie.” ONV. 342-343. André, R. 1816. Anleitung zur Veredelung des Schafviehes. Nach Grundsätzen, die sich auf Natur und Erfahrung stützen. Prague: J.G. Calvé. _____. 1819. “Meine Ansichten und Bemerkungen über organische Schwäche, besonders bei feinwolligen Schafen; veranlasst durch den Aufsatz des Herrn Grafen Emmerich von Festetics im Jännerheft 1819.” ONV. 161-162. Anonymous. 1811. On the Nature and Origin of the Merino Breed of Sheep. London: J. Hardling. (British Library, Banks Library B514 [14]). Anonymous. 1816. “Künstliche Befruchtung des Obstes.” ONV. 447. Bajema, C.J. 1982. Artificial Selection and the Development of Evolutionary Theory. Stroudsburg, PA: Hutchinson Ross Co. Bartenstein, E. 1818. “Bericht des Herrn Präses Baron Bartensteins an die k.k. Ackerbaugesellschaft.” ONV, ausserordentliche Beilage. 81-84. _____. 1837. “Äusserungen von Bartenstein über das von dem Herrn Professor Nestler bei dem SchafZüchter-Verein im Jahre 1836 aufgestellte und debatirte Thema der Vererbungsfähigkeit edler Stammthiere.” Mittheilungen. 9-10. Bartenstein, E., F. Teindl, J. Hirsch, and C. Lauer. 1837. “Protokol über die Verhandlungen bei der Schafzüchter-Versammlung in Brünn in 1837.” Mittheilungen. 201-205, 225-231, 233-238. Baumann, C. 1785. Nothwendige Anstalten zur Vermehrung, Verbesserung und Verschönerung der PferdRindvieh- Schaf- Geiss- und anderer Thierzuchten ohne Ausartung. Frankfurt/Leipzig. _____. 1803. Der Kern und das Wesentliche entdeckter Geheimnisse der Land- und Hauswirtschaft, zur bequemern Uebersicht und zum ausgebreitetern Gebrauch, mit der neunsten bewährten Versuchen und Nahrungsquellen, Liebhabern zum Handbuch gewidmet. Brünn: F.K. Siedler. Berry, H. 1826. “Whether the breed of live stock connected with agriculture be susceptible of the greatest improvement, from the qualities conspicuous in the male or from those conspicuous in the female parent.” British Farmer’s Magazine 1: 28-36. _____. 1829. Prize essay on “Whether the breed of live stock connected with agriculture be susceptible of the greatest improvement, from the qualities conspicuous in the male or from those conspicuous in the female parent.” Transactions of the Highland Society of Scotland 7 (New series 1): 39-42. (shorter, rewritten version of Berry 1826). Blacklock, A. 1838. A Treatise on Sheep. London. Boswell, J. [1825] 1829. Prize essay on “Whether the breed of live stock connected with agriculture be susceptible of the greatest improvement, from the qualities conspicuous in the male or from those conspicuous in the female parent?” Transactions of the Highland Society of Scotland, 7 (new series volume 1): 17-39.

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Bourde, A.J. 1953. The Influence of England on the French Agronomes 1750-1789. London: Cambridge University Press. Brown, R.D. 1987. Lucretius on love and sex. Leiden: E.J. Brill. Burkhardt, R.W. Jr. 1979. “Closing the door on Lord Morton’s mare: the rise and fall of telegony.” Studies in History of Biology 3: 1-21. Cline, H. 1805. “On the form of animals.” Communications to the Board of Agriculture 4: 440-446. Culley, G. [1786] 1794. Observations on Livestock; containing hints for chusing and improving the best breeds of the most useful kinds of domestic animals. London: G.G. & J. Robinson. _____. [1794] 1804. Culley George über die Auswahl und Veredlung der vorzüglichen Hausthiere. German translation of the 2nd edition from 1794. Berlin: F. Maurer. _____. 1807. Observations on Livestock; containing hints for chusing and improving the best breeds of the most useful kinds of domestic animals. 4th edition including appendix. London: G Wilkie, J Robinson, J Walker, G Robinson. Darwin, C.R. 1868. TheVariation of Animals and Plants under Domestication. London: Murray. Translated into German in 1868. Das Variiren der Tiere und Pflanzen im Zustande der Domestication. Stuttgart. Darwin, F., ed. 1887. Life and Letters of Charles Darwin, including an autobiographical chapter. 3 Volumes. London: John Murray. Daubenton, L.J.M. 1782. Instructions pour les bergers et pour les propriétaires des troupeaux. Paris, Dijon. Translated into German by Ch. A. Wichmann with added notes in 1784. Katechismus der Schafzucht zum Unterricht für Schäfer und Schäfereiherrn. Leipzig. Davenport, E. 1907. Principles of Breeding. Boston MA: Ginn & Co. Diebl, F. [1835-1841] 1844. Abhandlungen aus der Landwirtschaftskunde für Landwirthe, besonders aber für diejenigen, welche sich der Erlernung dieser Wissenschaft widmen. 4 Volumes. Brünn. Ehrenfels, J.M. 1817 “Ueber die höhere Schafzucht in Bezug auf die bekannte Ehrenfelsische Race. Belegt mit Wollmustern, welche die dem Herausgeber in Brünn zu sehen sind.” ONV. 81-85, 89-94. Ehrenfels, J.M. 1831. “Fortsetzung der Gedanken des Herrn Moritz Beyer über das Merinoschaf.” Mittheilungen. 137-142. Ehrenfels, J. M. 1837. “Schriftlicher Nachtrag zu den Verhandlungen der Schafzüchter-Versammlunmg in Brünn, am 10. Mai 1836.” Mittheilungen. 2-4. Elsner, J.G. 1826. Beschreibung meiner Wirtschaft. Prague: Calvé. d’Elvert, C. 1870. Geschichte der k.k. mähr. schles. Gesellschaft zur Beförderung des Ackerbaues, der Natur- und Landes-kunde, mit Rücksicht auf die bezüglichen Cultur-Verhältnisse Mährens und Oestrr. Schlessiens. Brünn: M.R. Rohrer. Festetics, E. 1819a. “Erklärung des Herrn Grafen Emmerich von Festetics.” ONV, ausserordentliche Beilage. 9-12, 18-20, 26-27. _____. 1819b. “Weitere Erklärungen des Herrn Grafen Emerich Festetics über Inzucht.” ONV, ausserordentliche Beilage. 169-170. _____. 1820. “Bericht des Herrn Grafen Emerich Festetics als Repräsentaten des Schafzüchter-Vereins in Eisenburger Comitate.” ONV. 25-28. Fink, J.H. 1799. Verschiedene Schriften und Beantwortungen betreffend die Schafzucht in Deutschland und Verbesserungen der groben Wolle, aus eigener Erfahrung und Thathandlung, zusammengetragen in Frühjahr 1799. Halle. _____. [1797] 1804. “Answers to questions posed by Sir John Sinclair, concerning the breeding of sheep, particularly in upper Saxony and neighbouring provinces.” In Communication to the Board of Agriculture. Volume 1, edited by A. Young. London: W. Bulmer a Co. 276-294. Fraas, K.N. 1852. Geschichte der Landwirtschaft, oder: Geschichtliche Übersicht der Fortschritte landwirtschaftlicher Kenntnisse in den letzten 100 Jahren, Volume 1 and 2. 2nd edition. Prague: F. Tempsky. _____. 1865. Geschichte der Landbau- und Forstwirtschaft. Seit dem sechzehnten Jahrhundert bis zur Gegenwart. München: Gottasche Buchhandlung. Galton, F. 1852. Unterricht von der Zucht und Wartung der besten Art von Schäfe. Leipzig. _____. 1856. Instruction sur la manière d’élever et de perfectionner les bêtes a laine. Dijon. _____. 1876. “A theory of heredity.” Journal of the Anthropological Institute of Great Britain and Ireland 5: 329-348. _____. 1908. Memories of my Life. London: Methuen. Hastfer, F.W. 1752. Ütförlig och omständig underrättelse om fullgoda fårs ans och skjötsel, til det all männas tjänst sammanfalttad af Fried. W. Hastfer. Stockholm: J Merckell. Hempel, G.C.L. 1818. “Horticultural Societät in London.” ONV. 408. _____. 1820. “Ueber die Entstehung und Wichtigkeit der verschiedenen Sorten der Getreidearten.” ONV. 161-165.

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Highmore, N. 1651. The History of Generation. London: John Martin. Home, H., Lord Kames. 1776. The gentleman farmer. Edinburgh: W. Creech & T. Cadell. “Irtep” [Petri]. 1812. “Ansichten über die Schafzucht nach Erfahrung und gesunder Theorie.” ONV. 1-5, 916, 21-23, 27-28, 45-48, 60-61, 81-85, 91-92, 106-107. “K in Mähren” [Köller, M.]. 1811. “Ist es nothwendig, zur Erhaltung einer edlen Schafherde stets fremde Original-Widder nachzuschaffen, und artet sie aus, wenn sich das verwandte Blut vermischet?” ONV. 294-298. Kjellberg, S.T. 1943. Ull och Ylle bidrag til den Svenska Yllemannfakturens historia. Lund, Sweden. Klemm, V. and G. Meyer. 1968. Albrecht Daniel Thaer. Halle: UEB Max Niemeyer. Knight, T. A. 1799. “An account of some experiments of the fecundation of vegetables.” Philosophical Transactions of the Royal Society of London 89: 195-204. Translated into German in 1800. “Versuche über die Befruchtung der Pflanzen”. Oekonomische Hefte. Leipzig. 322-340. Köcker, M. 1809. “Auszüge aus Briefen des Herrn Oekonom Köcker, auf den fürst. Salmischen Herrschaft Raiz in Herbst 1808 an den Herausgeber des letzten.” Hesperus 277-303. Kurucz, G. 1990. “The literature of the new agriculture in the Festetics library.” (English translation of Hungarian title). Magyar Knyuszenle, Hungary 106 (1-2): 32-44. Lasteyrie, C.P. 1802. Histoire de l’introduction des moutons à laine fine d’Espagne dans les divers états de l’Europe, et au Cap du Bonne-Espérance. Paris. Translated into German by G. Fleischer in 1804. Geschichte der Einführung der feinwolligen spanischen Schafe in die verschiedenen europäiachen Länder. Leipzig. Translated into English by B. Thompson in 1810. An account of the introduction of Merino sheep into the different states of Europe, and at the Cape of Good Hope. London. Lawrence, J. (A Farmer and Breeder) 1800. The New Farmer ’s Calendar ... comprehending all the material improvements in the new husbandry with the management of livestock. London: C. Whittingham. _____. ([1805] 1809). A General Treatise on Cattle, the Ox, the Sheep and the Swine. Comprehending their breeding, management, improvement and diseases. Dedicated to the Rt. Hon. Lord Somerville. London. Lincolnshire Grazier (William Youatt). 1833. The Complete Grazier. 6th edition. London: Baldwin and Craddock. Löwenfeld, R. 1835. “Andeutungen um die Veredlung der Schafe in allen vorzüglichen Eigenschaften nach geregelten, aus der Natur und Erfahrung abgeleiteten Grundsätzen mit dem besten und sichersten Erfolge vollkommen auszuüben.” Mittheilungen. 1-6, 9-13, 54-56. Lush, J.A. 1951. “Genetics and animal breeding.” In Genetics in the 20th century. Edited by L.C. Dunn. New York: MacMillan Co. 493-525. Marshall, W. 1790. The Rural Eeconomy of the Midland Counties. 2 volumes. London: G. Nicol. _____. [1818] c.1968. The review and abstract of the county reports to the Board of Agriculture. 5 volumes. York: T. Wilson. (Volumes published separately between 1808 and 1817, reissued in 1818). Reprint, Newton Abbot: David and Charles. Martin, W.C.L. Undated, ca.1849. “The Sheep.” In The Farmer’s library, Animal Economy. Volume 2. London: Charles Knight. 1-220. Mayr, E. 1971. “Open problems of Darwin research.” Essay review. Studies in the History and Philosophy of Science 2(3): 273-280. _____. 1972. “The nature of the Darwinian revolution.” Science 176: 981-989. _____. 1982. The Growth of Biological Thought: Diversity, Evolution and Inheritance. Cambridge MA: Harvard University Press. Mendel, G. 1866. “Versuche über Pflanzen-Hybriden.” Verhandlungen des Naturforschenden Vereines, Abhandlungen, Brünn 4: 3-47. (Editions in different languages were reviewed by Matalová (1973). They include the English translation commissioned by W. Bateson, 1959, reproduced in Classic papers in genetics, ed. J.A. Peters. Englewood Cliffs, NJ: Prentice Hall. And a second English translation published by C. Stern and E.R. Sherwood, ed., 1964, The origin of genetics: a Mendel source book. San Francisco: W.H. Freeman.) Morton, Earl of. 1821. “A communication of a singular fact in natural history.” Philosophical Transactions of the Royal Society 111: 20-22. Mylechreest, M. 1988. “Thomas Andrew Knight (1759-1838) and the Altenburg connection in the origin of Mendelism.” Folia Mendeliana 23: 27-32. Nestler, J.K. 1829. “Ueber den Einfluss der Zeugung auf die Eigenschaften der Nachcommen.” Mittheilungen. 369-73, 377-80, 394-8, 401-4. _____. 1836. “Ueber die Andeutung zur Veredelung der Schafe, von Herrn Rudolf v. Löwenfeld.” Mittheilungen. 153-155, 163-164, 173-176, 185-189, 205-208. _____. 1837. “Ueber Vererbung in der Schafzucht.” Mittheilungen. 265-269, 273-279, 281- 286, 289-300, 300-303, 318-320.

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_____. 1839. “Ueber Innzucht.” Mittheilungen. 121-128. _____. 1841. Amts-Bericht des Vorstandes über die vierte, zu Brünn von 20. bis 28. September 1840 abgehaltene Versammlung der deutschen Land-und Forstwirthe. Olmütz: A.Skarnitze. Orel, V. 1974. “The prediction of the laws of hybridization in Brno already in 1820.” Folia Mendeliana 9: 245-254. _____. 1975. “Das Interesse F. C. Napps (1792-1867) für den Unterricht der Landwirtschaftslehre und die Forschung der Hybridisation.” Folia Mendeliana 10: 225-240. _____. 1978. “The influence of T.A. Knight (1759-1838) on early plant breeding in Moravia.” Folia Mendeliana 13: 241-260. Orel, V., and Wood, R.J. 1998. “Empirical genetic laws published in Brno before Mendel was born.” Journal of Heredity 89: 79-82. _____. 2000. “Essence and Origin of Mendel’s Discovery.” C.R. Acad. Sc. Paris/Life Sciences 323: 1037-1041. Parry, C.H. 1800. Facts and Observations tending to show the practicability and advantage to the individual and the nation, of producing in the British Isles clothing wool, equal to that of Spain, together with some hints towards the management of fine-woolled sheep. Bath. (Reviewed in PTB, 1800, 222-223. Reviewed again in PTB 1801, under the title “Ueber die spanische Schafzucht in England ”, 907-909.) _____. 1806. “An essay on the nature, produce, origin and extension of the Merino breed of sheep.” Communications to the Board of Agriculture 5: Part 1, (18): 337-541. Pawson, H.C. 1957. Robert Bakewell, Pioneer Livestock Breeder. London: Crosby Lockwood & Son. Pearce, W. 1794. General view of the agriculture of the country of Berkshire. Drawn up for consideration by the Board of Agriculture and Internal Improvement, London. London: Nicol. Petersburg, J. 1815.”Veredlung des Schafviehs in der Blutverwandschaft betreffend.” ONV. 1-4. _____. 1816. “Äusserung des Repräsentanten für den Olmützer-Kreis Herrn Wirtschaftsraths Petersburg über die acht ausgestelleten Hauptpunkte.” ONV. 113-115. Petri, G. 1838. “Ueber die Inzucht.” Versammlung der Naturforscher und Aerzte zu Prag vom 18. bis 26. September 1837. Sektion für Landwirstchaft, Pomologie, Technologie und Mechanik. Pinto-Correia, C. 1997. The Ovary of Eve. Egg, Sperm and Preformation. Chicago and London: Chicago University Press. Pitt, W. 1809. General View of the Agriculture of the County of Leicester. Drawn up for consideration by the Board of Agriculture and Internal Improvement, London. London: Nicol. Purkinje, J.E. 1834. “Erzeugung (generatio, genesis, procreatio).” Encyclopedisches Wörterbuch der medicinischen Wissenschaften. Berlin. 11: 515-549. Rees, A.D.D. 1819. “Sheep.” In Cyclopaedia. Volume 22. (unpaginated). Roger, J. 1997. The Life Sciences in 18th Century French Thought. Edited by. K. R. Benson. Stanford: Stanford University Press. Röhrer, R. 1830. “Botanische Notitzen.” Mittheilungen. 120. Russell, N. 1986. Like Engend’ring Like. Heredity and animal breeding in early modern Europe. Cambridge: Cambridge University Press. Schulzenheim, Baron D. Schulz. de. [1797] 1804. “Observations on sheep, particularly those of Sweden.” Communications to the Board of Agriculture 1: 306-324. Sebright, J. [1809] 1982. The Art of Improving the Breeds of Domestic Animals. London: John Harding. Reprinted in Bajema (1982), 93-122. Sedlá™ek, H. 1826. “Zustand des mährischen Weinbaues und Vorschläge ihn durch Einführungder Rebenschulen von den edelsten Sorten zur Vervollkommen.” Mittheilungen. 161-7. Settegast, H. 1861. Die Zucht des Negrettischafes und die Schäfereien in Mecklenburgs. Berlin: G. Bosselmann. Sinclair, J. 1832. The Code of Agriculture. 5th edition. Sherwood, Gilbert & Piper, London. (The 1st edition 1819 was reviewed in Möglin Annalen, the review being reproduced in ONV, 1820, 201.) Somerville, J. [1803 and 1806] 1809. Facts and Observations relative to sheep, wools, ploughs and oxen: in which the importance of improving the short wooled breeds by a mixture of the merino blood is deduced from actual practice. Together with some remarks on the advantages, which have been derived from the use of salt. London: W. Miller. Spary, E. 1996. “Political, natural and bodily economies.” In Cultures of Natural History, edited by N. Jardine, J.A. Secord, and E.C. Spray. Cambridge: Cambridge University Press. Stieber, F. 1842. “In welchem Alter und unter welchen Lebensverhältnissen zeigt sich die Vererbung des Schafbockes und der Schafmutter am kräftigsten und sichersten?” Mittheilungen. 41-44. Stumpf, J.G. 1785. Versuch einer pragmatischen Geschichte der Schäfereien in Spanien, und der Spanischen in Sachsen, Anhalt-Dessau etc. Leipzig. Stumpf, J.G. 1800. An essay on the practical history of sheep in Spain and on the Spanish sheep in Saxony, Anhalt-Dessau etc. Translated by Dr. Lanigan. Dublin.

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Suneson, K.-H. 2001. 1700-talets fårimporter. Institutionen för landskapsplanering Ultuna. Agrarhistoria no 3. Uppsala. Teindl, F.J., J. Hirsch and J.G. Lauer. 1836. “Protokol über die Verhandlungen bei der SchafzüchterVersammlung in Brünn am 9. und 10. Mai 1836.” Mittheilungen. 303-309, 311-317. Terrall, M. 2002. “Speculation and Experiment in Enlightenment Life Sciences.” In A Cultural History of Heredity I: Seventeenth and Eigtheenth Centuries. Preprint 222. Berlin: Max Planck Institute for the History of Science Berlin. 27-41. Tessier, H.A. 1811. Instruction sur les bêtes à laine et particulièrement sur la race des mérinos. 2nd augemented edition. Paris. Translated into German by W. Witte in 1811. Über die Schafzucht, insbesondere über die Race der Merinos. Berlin. Thaer, A. 1804. Einleitung zur Kenntnisse der englischen Landwirtschaft. Volume 3. Hanover: Gebrüder Halm. (the final of three volumes on English agriculture, 1798-1804). Trow-Smith, R. 1959. A History of British Livestock Husbandry 1700-1900. London: Routledge & Kegan Paul. Vandemonde, C.-A. 1756. “Essai sur la manière de perfectionner l’èspece humain.” 2 volumes. Paris. Wallace, R. 1893. Farm Live Stock, Edinburgh. Walton, J.R. 1983. “The diffusion of improved sheep breeds in eighteenth and nineteenth century Oxfordshire.” Journal of historical geography 9: 115-155. Weckherlin, A. von. 1846. Die landwirtschaftliche Thierproduktion. Volume 1. Stuttgart and Tübingen: G. Gotta. (Later editions in 1851, 1857, 1865). Wilson, J. 1912. The Principles of Stock Breeding. London: Vinton & Co. Ltd. Winstanley, W. 1679. The Countryman’s Guide. London. Wood, R.J. 1973. “Robert Bakewell (1725-1795) pioneer animal breeder and his influence on Charles Darwin.” Folia Mendeliana 8: 231-243. Wood, R.J. and Orel, V. 2001. Genetic Prehistory in Selective Breeding: a prelude to Mendel. Oxford: Oxford University Press. Youatt, W. 1834. Cattle, their Breeds, Management and Diseases. London: Robert Baldwin. _____. 1837. Sheep: their breeds, management and diseases to which is added the mountain shepherd’s manual. London: Baldwin and Cradock. Young, A. 1771. The Farmer ’s Tour through the East of England. London. _____. 1791. “A month’s tour to Northamptonshire, Leicestershire, etc.” Annals of Agriculture 16: 480-607. _____. 1801. “Report of experience of Mr Crowe in breeding greyhounds and then sheep.” Annals of Agriculture 37: 439. _____. 1811. On the Husbandry of three Celebrated British Farmers, Messrs Bakewell, Arbuthnot and Duckett: being a lecture read to the Board of Agriculture. London.

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Characters written with invisible ink. Elements of Hybridism 1751-1875 Staffan Müller-Wille

In his inaugural lecture for the Collège de France, held December 2, 1970, Michel Foucault characterized Gregor Mendel as “a veritable monster (un monstre vrai)”, who “talked about objects, put methods to work, placed himself on a theoretical level, which all were alien to the biology of his epoch.” Mendel was not “in the truth (dans le vrai)” of his time, said Foucault, borrowing a concept from Canguilhem.1 But why a monster? Foucault brought up Mendel’s “discovery,” as we are used to call it, in a discussion of the notion of “discipline” as the “principle of control in the production of discourse.”2 “Within its limits, each discipline knows of true and false propositions; but it repels, to the other side of its margins, an entire teratology of knowledge.”3 Mendel seems a prominent case in point. Foucault’s remarks about Mendel were directed against those historians “who ask themselves how botanists and biologists of the nineteenth century could have done so much as not to see that what Mendel said was true.”4 However, inadvertently, I guess, Foucault’s answer reproduced a view wide spread among historians of science: the view of Mendel as an isolated figure tragically ahead of his time. Due to the historical work of Viczeslav Orel (1996) we now know that this view is thoroughly flawed: that Mendel’s work was contiguous on the thorough and up-to-date scientific education he had received at the universities of Olomouc and Vienna, that he was in close contact with a local community of naturalists and scientifically educated breeders, and that the questions he was trying to answer had been raised in this community. Other contributions to Mendelian scholarship have discussed late eighteenth and early nineteenth century “hybridism,” the work of Linnaeus, Kölreuter, Gärtner, as a research tradition that framed the questions guiding Mendel’s research.5 If, in view of these findings, we should still want to follow Foucault’s characterisation of Mendel as a “monster,” it would have to be in the sense of Mendel’s own theory: in his work we find an exceptional combination of disciplinary elements – statistics, breeding techniques, cell-theory, to name just the most obvious – none of them, taken in isolation, being abnormal for their time, all of them together, however, bringing about an extraordinary constellation. My aim in this paper is rather modest in trying to sketch out the genesis and historical legacy of one of these elements. It has sometimes been maintained that one of the things that made Mendel exceptional was that he did not – as the hybridists before and around him – care for the distinction between species and varieties in his experiments: He chose to experiment with plants which did not differ in a complex of characters – as “good” species should – , but in single and easily identifiable character pairs, while his predecessors had always been occupied with the

1 2 3 4 5

Foucault (1971), pp. 36-37. Ibid., p. 37. Ibid., p. 35. Ibid., p. 36. Olby (1979); Monaghan and Corcos (1990); cf. Orel and Hartl (1994), pp. 451-457 for a critical review.

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hybridisation of “good” species.6 Mendel himself pointed out explicitly in his 1866 paper that the distinction of species and varieties was irrelevant to his experiments: Uebrigens bleibt die Rangordnung, welche man denselben [i. e., the Pisum-varities used in Mendel’s experiments] im Systeme gibt, für die in Rede stehenden Versuche völlig gleichgiltig. So wenig man eine scharfe Unterscheidungslinie zwischen Species und Varietäten zu ziehen vermag, eben so wenig ist es bis jetzt gelungen, einen gründlichen Unterschied zwischen den Hybriden der Species und Varietäten aufzustellen.7

This statement is troublesome to those interpreters who have maintained that Mendel, as the hybridists before him, was primarily interested in the question if new species could arise from hybridisation.8 For if the role of hybrids in the genesis of new species was indeed at stake, then the taxonomic status of the plants used for and produced in the experiments should matter. Moreover, the resolution of this problem seems pertinent to the related question if Mendel was interested in inheritance as such at all, or only in so far as it “bore on his analysis of the evolutionary role of hybrids.”9 It is a well-known fact that Mendel used the term “inherit (vererben)” only once in his paper.10 If, however, the species-variety distinction did not matter to him, then it seems that he should have been ready to discard the age-old distinction of specific and individual inheritance as well, as he, and others before him, had also done in regard to the distinction of paternal and maternal inheritance. “Pure” inheritance – inheritance of particular elements abstracted from the generic forms they belong to – would then have been the object of his experiments. One possibility to resolve these problems is to doubt the assumption that plant hybridists before and around Mendel were working on the basis of a simple opposition of specific and individual (varietal) difference. And indeed, Mendel did address a current taxonomic determination of his experimental plants that clearly undercut this distinction: Wollte man die schärfste Bestimmung des Artbegriffes in Anwendung bringen, nach welcher zu einer Art nur jene Individuen gehören, die unter völlig gleichen Verhältnissen auch völlig gleiche Merkmale zeigen, so könnten nicht zwei davon [i. e. of the Pisum-varieties used in Mendel’s experiments] zu einer Art gezählt werden.11

In the Linnaean tradition, species and varieties had been distinguished along a nature-nurture divide: species comprise individuals characterised by a form that remains constant, i. e. does not change in the course of generations, no matter what external conditions individuals are subjected 6 7

8 9 10 11

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See, e.g., Stubbe (1963), p. 108; Dunn (1965), p. 18 quoting Correns (1905); Jacob (1970), pp. 220-222; Bowler (1989), p. 99. Mendel (1866), pp. 6-7; translation by Bateson (Mendel [1866] 1902), 44: “The positions, however, which may be assigned to them in a classificatory system are quite immaterial for the purposes of the experiments in question. It has so far been found to be just as impossible to draw a sharp line between the hybrids of species and varieties as between species and varieties themselves.” See, e.g., Olby (1979), p. 67. Olby loc. cit. Mendel (1866), p. 14. Mendel (1866), p. 6; translation by Bateson ([1866] 1902): “If we adopt the strictest definition of a species, according to which only those individuals belong to a species which under precisely the same circumstances display precisely similar characters, no two of these varieties could be referred to one species.”

Characters written with invisible ink. Elements of Hybridism 1751-1875

to from generation to generation and from place to place. These constant forms obey, as Linnaeus put it, “inherent laws of generation (generationis inditas leges).” They are due to internal forces that lie hidden in the organisation of living bodies, ultimately in the interaction of the generative substances produced by the male and female reproductive organs in fertilisation. Varieties, on the other hand, comprise individuals of the same species that differ in form due to the influence of external factors, as climate and soil, varying with time and place. Varieties, that is, are due to the physical forces governing the environment of organisms.12 From this distinction of species and varieties it would follow, that members of a species brought under a regime of perfectly homogenous external conditions should be perfectly identical in all respects – the species criterion Mendel refers to in the above quote. Any individual difference – as the character pairs round-wrinkled, yellow-green etc. used by Mendel in his experiments – separating individuals under such circumstances would, therefore, have to count as a specific difference. The Linnaean species concept thus offers, in principle, the possibility to distinguish species according to differences in individual characters: if plant forms, cultivated and propagated under conditions homogenous at any given point in time, should happen to exhibit an individual difference, in all other respects remaining perfectly identical, they should be counted as belonging to different species. Specific heritable difference, that is, may, under this condition, coincide with individual heritable difference. And this is exactly the condition under which Mendel referred to the plants he experimented with as different ‘species (Arten)’: In the paragraph preceding the one quoted above he made sure to stress, that the 22 pea varieties (Erbsensorten) he selected for his experiments had proven good under two years to constantly produce identical descendants (durchaus gleiche und constante Nachkommen).13 Throughout his paper, Mendel should refer to his experimental plants as different “species (Arten)”14 in this sense, using the taxonomically neutral expression “forms (Formen)” interchangingly with it.15 In the following I want to sketch out, how the tension of individual and specific difference came into play in the “hybridist” tradition before Mendel. My aim with this is to show, that Mendel’s step to deal with varieties different in respect to single character pairs was not as dramatic as it seems in retrospect, but rather contiguous on the hybridist tradition.

1. Constant Varieties The first systematic discussion of plant hybrids was published in 1751 by Linnaeus under the title Plantae hybridae. As it formed the major target of critique in Kölreuter’s experiments from the 1760s, it can be regarded as the founding document of the hybridist tradition. In stark contrast to that later tradition, however, it shows a surprising lack of experimental evidence: The only experiment in plant hybridisation, that Linnaeus should ever perform, dates from a much later time, 1758/59, and was executed in answering a prize question of the St. Petersburg Academy of Sciences asking for proofs of plant sexualtiy.16 The hybrid plants of the 1751 Plantae hybridae were 12 13 14 15 16

Müller-Wille (1998), pp. 351-358. Mendel (1866), p. 6. E.g. ibid., p. 6, 9, 11. E.g. ibid., p. 24; Bateson’s translation is sometimes misleading in this respect, by using “variety” (Mendel 1866 [1902], 47) or “stock” (65), where Mendel has Art. For a discussion of this experiment see Müller-Wille (1998), pp. 370-372.

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surmised on a completely different basis, which, in general terms, was explained by Linnaeus in the following words: Cardo argumenti pro statuendis plantis hybridis, videtur tamen a posteriori corroborari. Constat enim omnibus, Botanicos, utpote Tournefortium, Boerhavium, Mitchelium, Pontederam, Helwingium cum reliquis sub finem prioris seculi, & quidem ad tempora Cl. Praesidis, hujus seculi, innumeras collegisse novas plantas Europaeas, veteribus ignotas; sed quod observandum, qua maximam partem similes veterum, ut vix figuris distingui potuerint. Cum itaque characteres sufficienter defecerint in hisce plantis, non potuit non Cl. D. Praeses has referre ad matres suas, sub nomine VARIETATUM, quae colore, odore, sapore, magnitudine, tempore, pubescentia &c. differebant, quod acutissimi quoque Botanici, ut Jussieus, Royenus, Gronovius, Hallerus, Gmelinus, Guettardus, Dalibardus, Wachendorffius, Gorterus, Bütnerus &c. post illum approbarunt & continuarunt. Dictum quidem est has varietates ex solo, loco, climate ortas esse, ideoque tantummodo accidentales; Interea tamen huic reformationis variae subjectae sunt, quae Botanicos plurimum sollicitabant in determinandis speciebus, ad quam scilicet speciem merito referendae essent, quando duabus similes erant, & quidem ita, ut ne unica nota in planta quaestionis, etiamsi distinctissima, & maxime singularis inveniatur, quae non in alerutra ambarum occurit. Crescuitque haec hybrida in eodem solo cum ambabus & tamen a veteribus non observata. Haec primum nos ad hoc Problema deduxerunt, utrum duae diversae plantae tertiam produxissent, an ut hybridae considerandae essent, cum Botanici his primum temporibus generationem plantarum optime norint, & quales mutationes in plantis hinc deducantur, observarint in Brassica aliisque.17

This passage testifies that the problem of hybrids was not constituted for Linnaeus by a record of more or less authorative tales and reports. It rather resulted from a systematic and continuous taxonomic practice that tried to correlate the reproduction of plants, a vital process, with the structural differences they exhibit: Differences among individuals related by reproduction – individuals referrable “to their mother” – are varietal differences in the sense of depending for their reproduction on the continuity of certain local environments – “soil, locality, climate” – while differences among species depend on reproduction as such, or, as Linnaeus put it in a paragraph preceding the one quoted above, depend “on that natural law”, according to which 17

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Linnaeus [1751] 1764, pp. 32-33: “The main argument for proposing hybrid plants, however, seems to be corroborated a posteriori. It is certain to everyone, that botanists like Tournefort, Boerhaave, Michel, Pontedera, Helwig and others at the end of the last century and certainly in these days have collected innumerable new European plants, unknown to antiquity; but that it was observed, that these were so similar to those of antiquity, that they could hardly be distinguished from them by figures. As therefore sufficient characters were missing in them, [Linnaeus] could only refer them to their mothers under the name of varieties, which differ by colour, odour, taste, magnitude, time, thorns, etc., and this was continued and attested after him by the more acute botanists, like Jussieu, Royen, Gronovius, Haller, Gmelin, Guettard, Dalibard, Wachendorff, Gorter, Büttner. It has been said, that these varieties were brought forth by soil, locality, climate, so that they by all means are accidental. Meanwhile, however, various [plants] have been subjected to this reform, which have troubled botanists in determining species, as to which species they should be rightfully referred, when they were similar to two others, in such a way, that no single character of the plant in question, even if most distinct and singular, could be found, which not also occurred in one of the two others. These hybrids also grow in the same soil as the others, though they were not observed by the old. This has first lead us to the problem, if two different plants have produced a third, or if they can be considered hybrids, as botanists only now are learning about the generation of plants, and deduce such mutations in plants, as had been observed for cabbage and others.”

Characters written with invisible ink. Elements of Hybridism 1751-1875

members of a species “celebrate their marriages and propagate their families, such that they rarely deflect (abeunt) from that law to other species.”18 Taxonomic analysis, carried out along these lines, however, reaches a twofold limit in phenomena whose very possibility is determined by the presupposition that guides it. On the one hand, plant forms may exhibit a constellation of specific differences, which appears as a mere combination of other plant forms, making it impossible to identify them as really distinct forms: each of their individual characters is not only reproduced within their own species, but will also be found to be reproduced in either one of two others. On the other hand, reproductively contiguous plants – “growing in the same soil” – may exhibit differences which do not depend on external factors for their reproduction as they remain different under homogenous conditions and, each respectively, identical under heterogenous conditions. This point is only implicitly hinted at in the passage quoted above, but made perfectly clear in another essay on the “metamorphosis of plants” which Linnaeus published in 1754: Cum botanici vidissent eandem speciem in diverso climate vel solo, variare, primum ex illis novas species constituerunt; quo accidit, ut numerus plantarum nimis augeretur, nulii dum limites essent. Recentiores itaque botanici varietates ad species suas reducere coeperunt, ne entia praeter necessitatem multiplicarentur. Botanici, qui videbant solum & coelum tam multas fecisse varietates, intelligebant quoque, quod solum & coelum eas reducerent, quam ob caussam in hortis botanicis eas seminabant; at, cum viderent nonnullas in uno eodemque solo & climate, aeque constantes esse, contendebant aliqui, eas non pro varietatibus, sed distinctis speciebus habendas esse […].19

As in the Plantae hybridae, this observation is used as a starting point to speculate about the hybrid origin of such “constant varieties”. The two cases of dubious species, “constant varieties” and species of a combinatory appearance, served Linnaeus in the enumeration of 101 hybrid plants contained in the Plantae hybridae as a principle for their classification: The hybrids no 1 to 17, and the hybrids no 18 to 34 were called Bigeneres and Congeneres, expressing their origin from the hybridisation of plants of different genus and species respectively, while the hybrids no 35-40 were classified as “deformed (deformatae)”, in as much as “individuals of the same species had acquired a structure different from their mother or ancestors.”20 The remaining sixty hybrid plant species were classified as “obscure (obscurae)” and “suspect (suspectae),” obviously as a result of the fact that they could not as easily be referred to one of the preceding classes. 21 While Linnaeus’s descriptions of hybrid species were based on a highly unclear and speculative theory of sexual reproduction – “adventurous and against all reason (abentheuerlich und wider alle 18 19

20 21

Ibid., p. 30. Linnaeus, ([1755] 1788), p. 380: “When botanists saw that the same species varied in diverse climate or soil, they first made different species of them; with the consequence that the number of plants was so excessively multiplied that there were no limits. Recent botanists have therefore begun to reduce varieties to their species, as to not multiply beings beyond necessity. Botanists, who saw that soil and sky make so many varieties, understood also, that soil and sky would reduce them, and therefore sowed them out in botanical gardens. However, when they saw that some nevertheless remained constant in one and the same soil and climate, some contended that they should not be held for varieties but for different species.” linnaeus ([1751] 1787), p. 50. Müller-Wille (1998), pp. 366-368.

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Vernunft laufend)” as Kölreuter would call it in 1766,22 – they disclose the highly specific institutional conditions that allowed for their detection. Specific forms of living beings, especially of exotic animals, had been assumed to hybridise and transmute since antiquity, but such events were tied to the exceptional nature of certain environments, as the oasis of the Sahara, or conditions of domestication and cultivation. Even hybrids had their “natural places”, so to speak. In the system of botanical gardens, to which Linnaeus referred in the above quote form the Metamorphoses plantarum, this close dependence of place and the reproduction of forms was loosened in a regulated practice by which different plant forms were actively reproduced under a variety of controlled environments.23 Plant form, under this condition, was broken down to a system of minute structural differences that proved to be distinctively reproducable under all circumstances; and plant reproduction was broken down to individual genealogical lines populating the controlled beds of a variety of garden localities. The dichotomy of specific (essential) and varietal (accidental) form thus opened to include a complex array of phenomena where the reproduction of plants as such, independent of place and time, seemed to engender variation.

2. Hybrid-like Nature Joseph Gottlieb Kölreuter may very well have been inspired to his hybridisation experiments by the prize essay Linnaeus had submitted to the St. Petersburg Academy of Sciences: with this essay, Linnaeus had sent seeds of the Tragopon-hybrid he had produced experimentally, so that the hybrid might be reproduced in the botanical garden of St. Petersburg, which it was. Kölreuter discussed the specimens raised there in his Vorläufige Nachricht von einigen das Geschlecht der Pflanzen betreffenden Versuchen (1761),24 doubting, however, their hybrid nature by arguing that they were “half-hybrids”, that is, that male pollen from the maternal species had inadvertently been involved in their production. Kölreuters publications on hybridisation experiments, which appeared with three further Fortsetzungen can, over all, be read as an attempt of refuting Linnaeus’s claim that new species can arise from hybridisation. Expressing a veritable fear for the “astoninshing confusion (erstaunliche Verwirrung)” and the “monstruous swarm of imperfections (ungeheuren Schwarm von Unvollkommenheiten)” that would be the consequence of plants “unchangingly and constantly conserving their bastard species”25 he basically followed two lines in demonstrating experimentally the impossibility of hybridisation giving rise to new species: Firstly, by demonstrating, that plants raised from artificially hybridised species do not exhibit a combination of parental and maternal traits, as Linnaeus had ventured to say, but that their traits occupied the “middle proportion” between the parental traits. And secondly, by demonstrating, that true species hybrids always are infertile, and thus cannot reproduce, a fact, that Linnaeus sometimes noted, but seems to have been surprisingly indifferent to.26 22 23 24 25 26

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Kölreuter (1766), p. 37. Cf. Müller-Wille (2001). Kölreuter (1761), pp. 41-42. Kölreuter (1763), p. 9. Ibid., p. 37.

Characters written with invisible ink. Elements of Hybridism 1751-1875

Kölreuter summarised the results of his experiments in a classification of hybrid plants which, other than the one proposed by Linnaeus, was not based on the kinds of evidence he had for them – a quite external criterion – but on the dependence of the fertility of hybrid offspring on the similarity of their parental species, which formed the core of Kölreuters theoretical claims: Nachdem nun alle die Bastarte, die ich hervorzubringen und zu erziehen das Glück gehabt habe, angezeigt worden, so will ich sie nach ihrer verschiedenen Natur in folgende Klassen, Ordnungen, Geschlechter und Gattungen abtheilen. Erstlich theile ich sie in drey Klassen: unter die I. Kl. Gehören die vollkommenen Bastarte, die aus zwo oder drey verschiedenen natürlichen Gattungen eines Geschlechts entstanden sind, und bey deren Erzeugung der eigene männliche Saame gänzlich ausgeschlossen worden. Unter der II. Klasse hingegen stehen die unvollkommenen, die zwar aus zwo verschiedenen natürlichen Gattungen eines Geschlechts entstanden sind, bey deren Erzeugung sich aber außer dem fremden auch noch etwas weniges von ihrem eigenen männlichen Saamen zugleich mit eingeschlichen hat. Die III. Kl. begreift die Bastartvarietäten unter sich, die aus zwo Varietäten einer natürlichen Gattung entstanden sind, und bey deren Erzeugung der eigen männliche Saame gänzlich ausgeschlossen worden ist.27

This classification is based on a distinction of “natural species (natürliche Gattung)”28 and “varieties of a natural species (Varietäten einer natürlichen Gattung)” that the ensueing “systematic table of all hybrids hitherto produced by art” discloses: While the perfect and imperfect hybrids, both produced from different “natural species”, are further subdivided according to way in which they exhibit infertility or strongly reduced fertility – from “maternal”, “paternal” or “both sides” – hybrid varieties are characterised as “perfectly fertile”. By adopting an additional criterion for specific contiguity beyond mere genealogical descendence (as in Linnaeus), namely the production of fertile offspring, Kölreuter is able, so to speak, to save nature from the “monstruous swarm of imperfections” that would follow for her if Linnaeus had been right: Hybrid varieties do reproduce successfuly, but as they always occupy the “middle proportion” between their parental traits, their unions remain within its bonds. “Perfect” hybrids, on the other hand, uniting two “natural” species, do not reproduce successfully. There is a natural limit to the reproductive union of plant forms so dissimilar as to constitute different species, a central tenet that Kölreuter expressed in the following words: Bey vielen […] Pflanzen […] habe ich ihrer ziemlich nahen Anverwandtschaft ungeachtet , doch durch dergleichen Versuche nicht das geringste ausgerichtet, und es ist, in Absicht auf den Erfolg, eben so viel gewesen, als wenn ich sie gänzlich verschnitten, oder gar nicht mit Saamenstaube belegt hätte: Woraus ich zur Genüge ersehen, daß sich die Bastartpflanzen 27

28

Ibid., pp. 47-48: “Since all the hybrids, which I happened to produce and educate, have now been indicated, I want to divide them according to their different nature into the following classes, orders, genera, and species. Firstly, I divide them into three classes: to the first class belong the perfect hybrids, which originated from two or three different natural species, and where their own male seed was completely excluded from their production. Under the second class stand the imperfect, where although generated from two different species of a genus something of their own male seed, besides the alien one, crept in durign generation. The third class comprises hybrid-varieties, which were generated from two varieties of a natual species, and where their own male seed was completely excluded during generation” The term Gattung designated species in eighteenth century German, other than in modern German, where it designates genera. For “genus” Geschlecht was used, while Art, corresponding to “species” in modern German, was used to refer to kinds in general.

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nicht so leicht erzeugen lassen, als sich manche einbilden mögen, und daß eine widernatürliche Befruchtung eine weit größere Ähnlichkeit voraussetzt, als sie von einigen, wider alle Wahrscheinlichkeit, als hinreichend angenommen.29

As with Linnaeus distinction of species and varieties, Kölreuters classificatory move produced, however, the very possibility of observing problematic cases. For one thing, he operated in his hybridisation experiments with what Linnaeus had designated as “constant varieties”, i. e. varieties which preserved their distinctive characters in the course of generations. In some cases he expressly addressed them as such – natürliche und beständige Varietät – , and expressed himself to be unable to “state the effective cause (wirkende Ursache) of this minute difference” as they often occured side by side in the field;30 in others he seems to have instituted control experiments to assure himself of the constancy of their characters.31 Kölreuter thus was forced to admit the omnipresence of a “generation among varieties”, with admittedly unforeseeable consequences. 32 And in one of the hybridization experiments he instituted with colour varities of Dianthus he did encounter unexpected results: So sicher sich sonst bey denjenigen Bastarten, deren natürliche Mutter und Vaterpflanzen, sie seyn nun verschiedene Gattungen, oder nur bloße Varietäten, noch auf keinerlei Weise aus der Art geschlagen sind, die mittlere Farbe einzufinden pflegt: so unregelmäßig scheint es in diesem Stücke bey solchen herzugehen, die, wie z. E. die Gartennelken und mehrere andere Gattungen aus diesem Geschlechte, durch die Cultur auf mannigfaltige Art verändert werden. Es erhellet solches nicht nur aus den gegenwärtigen Beyspielen, sondern auch vornehmlich daraus offenbar, daß von einer aufs sorgfältigste mit ihrem eigenen Saamenstaube belegten Blume dieser Art öfters nicht eine geringe Anzahl ganz verschiedener Sorten entspringen, wie ich aus einer zuverlässigen Erfahrung versichern kann. Vielleicht giebt die mannigfaltige Veränderung, die in der Natur fast aller, seit einer langen Reihe von Jahren her einer widernatürlichen Behandlung und Lebensart unterworfener Pflanzen und Thiere vorgeht, zu Aufhebung des Gleichgewichts bey der ordnungsgemäßen Erzeugung nicht nur in Absicht auf die Farbe allein, sondern auch so gar in Ansehung der Gestalt, Lage, Zahl und Proportion aller Theile untereinander selbst, eben so viel Anlaß, als der erste ab- oder aufsteigende Grad bey der Bastardzucht. Wenigstens lassen sich viele dergleichen Varietäten und Mißgeburthen so wohl im Thier- als Gewächsreiche aus der ungleichen Mischung einer Saamenfeuchtigkeit mit der anderen, und aus ihrer wechselweisen ungleichen Wirkung und Einfluße auf einander, auf eine ganz ungezwungene Weise herleiten. Sollte wohl z. E. die größere oder geringere Ähnlichkeit der Kinder bald mit dem Vater, bald mit ihrer Mutter, und die denselben zu Theil gewordene größere oder geringere Fruchtbarkeit, nebst verschiedenen anderen Eigenschaften mehr, einen anderen Grund haben? Die Natur der Thiere und Pflanzen wird gewissermaßen bastartartig, so bald sie sich auf irgend eine Weise von derjenigen Bestimmung entfernen, zu der sie eigentlich erschaffen worden. Und wer weiß, ob unter den Menschen selbst eben so gar viele vorkommen, die in diesem Verstande nicht halbe Bastarte sind? 33 29

30 31 32

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Kölreuter (1761), p. 44: “In many plants, despite their rather close affinity, I have not been able to effect anything by such experiments, and it is, in regard to their success, as if I had castrated them completely or had not pollinated them at all. From which it follows with sufficient reason, that hybrid plants are not produced as easily, as some have imagined, and that an unnatural conception presupposes a much closer resemblance as it it is supposed to be sufficient by some, against all propbability.” Kölreuter (1766), p. 35. Kölreuter (1763), p. 46; Idem. (1766), p. 35. Kölreuter (1763), p. 44.

Characters written with invisible ink. Elements of Hybridism 1751-1875

The source for Kölreuter’s dismay with the result of his Dianthus-cross – which found its expression in an eloquent and speculative digression unusual for the otherwise very sober descriptions of experimental results – is stated in the first sentence: Counter to Kölreuters theoretical expectation,34 the offspring from this cross proved to display an array of varieties, combining traits of their parental species in various ways, instead of occupying the “middle proportion”. Kölreuter was used to this phenomenon from the “first descending or ascending grade in hybrid breeds” or “half-hybrids” – i. e., what we nowadays would call a first back-cross – but he accounted for these cases by the “unequal mixture” and disruption of the “balance” of the male and female “seed fluids (Samenflüssigkeiten)”, which, in “perfect hybrids”, resulted from the union of different “natural species” and was also responsible for their infertility. 35 In terms of fertility, the Dianthus-cross behaved like a “hybrid variety”, in terms of the distribution of character traits like a hybrid of two “natural” species. The only resort from this consequence is the relegation of such cases to conditions of domestication and cultivation. The same is the case with an experimental result, that formed the inverse of the one just discussed: In crossing two species of Cucurbita, which, in regard to their dissimilarity, Kölreuter had obviously expected to behave as belonging to two different “natural” species, he acquired perfectly fertile offspring, accounting for this by designating the parental forms as “essentially as little different, as a lap-dog and an English mastiff”.36 As in the case of Linnaeus, the opposition of species and varieties is opened in a regulated experimental practice – a “workshop of varieties (Varietätenwerkstätte)”, as Kölreuter once designated his experiments in producing “half-hybrids” by back-crossing – this time characterised by the attempt to correlate the physiological function of fertility with taxonomic affinities in hybridizatrion experiments. And as with Linnaeus’s “constant varieties”, it is opened to include an apparent oxymoron, the bastardartig nature of plants, animals, and humans.

33

34 35 36

Kölreuter (1766), p. 85 (transl. partly based on Mayr 1986, 170): “As surely as otherwise the middle colour occurs in those hybrids, whose natural mother- and fatherplants, be they different species, be they different varieties, have degenerated in no way; as irregular does the play go on in those hybrids, which, as in the case of garden carnations and other species of this genus, which have been changed by cultivation in manifold ways. This is not only illuminated from the present examples, but also clear from the fact, that a flower of this kind, most carefully impregnated with its own pollen, often produces a considerable amount of very different sorts. Perhaps the manifold changes which take place in the nature of nearly all plants and animals subjected for a long series of years to an unnatural treatment and mode of living give cause to an unsettling of the balance in regular generation not only in respect to colour, but also with respect to form, position, number, and proportion of all the parts to each other, as much as in the first de- or ascending degree in hybrid breeding. At least it seems possible to explain quite naturally many such varieties and monstrosities among animals as well as plants as caused by an unequal mixture of their seed fluids and by the reciprocally unequal effect and influence on each other. Why should the greater or lesser similarity of the children with either father or mother, and the greater or lesser fertility that fell onto them, as well as other attributes, have any other cause? The nature of animals and plants becomes, so to speak, hybrid-like, as soon as they have departed in some manner from that destination, for which they had been created. And who knows whether even among humans themselves many occur which, in this sense, are half-hybrids?” Mayr calls the last statement “rather prophetic […] when translated into Mendelian language.” Cf. Kölreuter (1764), p. 11. Cf. Mayr (1986), pp. 168-170. Kölreuter (1766), p. 119; cf. p. 93.

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3. Types of Hybrids When Carl Friedrich Gärtner published his Versuche und Beobachtungen über die Bastarderzeugung im Pflanzenreich in 1849, he could, other than Kölreuter, already look back on a rich and varied literature on plant hybridisation, both produced experimentally and inferred from taxonomic analysis. Sageret had presented his results on the hybridization of melons and pumpkins (Cucurbitaceae), rejecting the view that the “similarity of hybrids to their two ascendants [consists] in the intimate fusion of diverse characters proper to each of them in particular, but in a distribution, equally or unequally, of the same characters.” 37 And William Herbert had proposed to view genera as the stable units of living nature “disport[ing] themselves in numerous forms of species and local or accidental varieties, which are more or less capable of intermixture according to their constitution and diversity, with various degrees of fecundity and sterility in the united produce.”38 Kölreuter’s simple dichotomy of “perfect” hybrids and “hybrid varieties” had given way to a complex picture of various degrees in the combination, segregation and atavistic reappearance of varietal traits on the one hand, and various degrees in the physiological capacity of hybrids to produce fertile offspring. In attempting to clarify this picture, Gärtner made sure to choose a different perspective than Kölreuter had done, though he acknowledged his authority. Instead of viewing hybridisation as a basically chemical process, as Kölreuter had done, Gärtner chose a “biological” point of view). 39 Most explicetly, this point of view came to the fore, when Gärtner discussed the distinction of species and variety: Die Frage, worin sich die Art von der Varietät unterscheide, ist daher, wie E[lias] Fries bemerkt, eine rein biologische: indem ein sicherer Grund der Artbestimmung nicht bloß in der Abstraktion gefunden werden kann, weder in den Merkmalen noch in den Uebergangsformen, sondern man muss ihn in der Reflexion suchen, d. h. in der individuellen Geschichte einer jeden Art, deren ganzen Entwikkelung, und nicht in einem gewissen Moment.40

In other words: there is no external criterion by which species are distinguished from varieties, but it is rather the life of the species itself, its “individual history” and “whole development” that determines it as a particular species. This is to be sought “in reflection” as its history and development has to be seen in relation to other species, as the definition of species, which Gärtner provided after all, testifies: Das Wesen der Art besteht daher in dem bestimmten Verhältniss ihrer sexuellen Kräfte zu anderen Arten, welches Verhältniss neben der specifischen Form bei jeder Art ein eigenthümliches, besonderes und constantes ist; Form und Wesen sind in dieser Beziehung Eins.41

The consequence of this species definition is a distinction of two kinds of “affinities (Verwandtschaften)” among plants: an “external (äußere)” one, consisting in the “conformity of 37 38 39 40 41

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Sageret (1826), pp. 300-302. Herbert (1837), p. 17. Gärtner (1849), p. x. Ibid., p. 151. Ibid.

Characters written with invisible ink. Elements of Hybridism 1751-1875

habitus, i. e. in its growth form, in the figure and form of leaves and in the harmony of flowers and reproductive organs;” and an “internal (innere)” one, consisting “in the lesser or greater tendency (Geneigtheit) for sexual union among species in hybrid fertilization.”42 The “chemical” view, “in analogy to dead nature,” would suggest “that similar substances produce similar forms and forces.”43 Yet the fact that both varieties and species always produce the same “types of bastards (Bastardtypen)” speaks against this analogy.44 One has to suggest, therefore, as Gaertner put it […] dass in den beiden Substraten der Geschlechter der Pflanzen und in ihrer gegenseitzigen Anziehung der Grund der Fähigkeit zur Bastardzeugung liegt: worin aber die specielle Beschaffenheit des einen wie des anderen Faktors besteht, wird weder durch mikroskopische, noch durch chemische Untersuchungen zu beantworten sein; indem es sich hierbei um eine rein vitale Thätigkeit handelt, welche wir mit keinem passenderen Wort, als mit dem der Wahlverwandtschaft zu bezeichnen wissen.45

Gärtner therefore suggested a new classification of hybrids in place of the one proposed by Kölreuter, which had been based on the distinction of (fertile) variety bastards and (infertile) “perfect hybrids” between species and which Gaertner considered as “unnatural (nicht naturgemäß)”. Instead, the “natural classification (naturgemäße Eintheilung)” should be established according to the “composition and descendence (Zusammensetzung und Abstammung)” of hybrids solely. The taxonomic status of the parental forms does not play a role anymore in this classification, and Gärtner made sure, in a separate chapter on “varieties and variety bastards”, to point out that hybrids of varieties, as long as the latter prove to be “stable” and “constant” behave in many respects exactly in similar ways as species bastards. 46 And it is in this respect, that “species constancy” counted for Gaertner, namely in as much as the “types of hybrids are not vague and variable, but constant, obeying certain invariable laws of formation (Bildungsgesetzen),” and in as much as they are “constantly and regularily (constant und gesetzmäßig) produced from the same factors again and again.”47 Species appear as a reservoir of “factors”, which combine, reappear, and distribute in a regular fashion in hybridisations. As a consequence, Gaertner accepted all but one – which had proved to be variable under cultivation – of Kölreuter’s varietal hybrids as species hybrids.48 What distinguishes species, is their “inner nature” by which each of them produce “different types of hybrids”.

4. Concluding Remarks In this paper, my main aim was to draw a more complex picture of the work of the hybridist tradition of the late eighteenth and early nineteenth century, especially in regard to the status of the taxonomic categories employed in this tradition. In each case, the apparent oxymorons of “constant varieties”, “hybrid-like natures”, and “types of hybrids” resulted from a use of these 42 43 44 45 46 47 48

Ibid., p. 166. Ibid. Ibid., p. 168. Ibid., p. 186. Ibid., pp. 576-577. Ibid., pp. 243-235. Ibid., p. 581.

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categories in attempts to structure nature not by identifying the “natural places” of things, but by identifying regularities in their motion. The apparent stability of a hierarchy of living forms broke down under this condition, laying bare heredity as one of the central forces in the development of life, both as a source of its diversity and its regularity. Mendel’s contribution was, in this specific respect, perfectly “in the truth” of this tradition, as shown by his own conviction to have contributed to the Entwicklungsgeschichte der organischen Formen. The decision to deal – in a Detailversuch (even that a Gärtnerian expression),49 as he made sure to specify – with “constant varieties” was not as dramatic a deflection from the tradition culminating in Gärtners voluminous book. In Gärtners understanding, the pea varieties Mendel experimented with, though only differing in respect to single character pairs, were different species, and the conclusions Mendel drew from them in regard to his predecessor’s work in the “concluding remarks” of his paper were therefore quite naturally phrased with respect to “species”. That Mendel did take a decisive step towards twentieth century Mendelian genetics is therefore not simply a myth, but based on a historical coincidence, that did, in fact, occur. For the theme of this conference, the “cultural” history of heredity, there is one important consequence that I would like to draw from the “internal” history I have tried to sketch out: Heredity becomes central in the early nineteenth century not as a result of social processes in which dichotomic exclusions become cemented, but rather as a result of processes in which such structures become dissolved. Science played an important role in this, though obviously not science alone. In no other statements from the hybridist tradition, I believe, does this come to the fore more clearly than in the sentences with which Charles Darwin summarised the results of the chapter on reversion or atavism in his Variation of animals and plants under domestication: The fertilized germ of one of the higher animals, subjected as it is to so vast a series of changes from the germinal cell to old age – incessantly agitated by what Quatrefages well calls tourbillon vital [the maelstrom of life] – is perhaps the most wonderful object in nature. It is probable that hardly a change of any kind affects either parent, without some mark being left on the germ. But on the doctrine of reversion, as given in this chapter the germ becomes a far more marvellous object, for, besides the visible changes which it undergoes, we must believe that it is crowded with invisible characters, proper to both sexes, to both the right and left side of the body, and to a long line of male and female ancestors separated by hundreds or even thousands of generations from the present time: and these characters, like those written on paper with invisible ink, lie ready to be evolved whenever the organization is disturbed by certain known or unknown conditions.50

49 50

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Ibid., pp. 291, 580. Darwin (1875), 2: pp. 35-36.

Characters written with invisible ink. Elements of Hybridism 1751-1875

References Bowler, Peter J. 1989. The Mendelian Revolution: The Emergence of Hereditarian Concepts in Modern Science and Society. Baltimore: John Hopkins University Press. Correns, Carl, and Gregor Mendel. 1905. “Gregor Mendels Briefe an Carl Naegeli 1866-1873. Ein Nachtrag zu den veröffentlichten Bastardisierungsversuchen Mendels.” In Gesammelte Abhandlungen Zur Vererbungswissenschaft. Aus periodischen Schriften 1899-1924. Berlin: Julius Springer, 1233-97. Darwin, Charles. 1893. The Variation of Plants and Animals under Domestication. 2nd edition. 2 volumes. London: John Murray. Dunn, Leslie Clarence. 1965. A Short History of Genetics. The Development of Some of the Main Lines of Thought: 1864-1939. New York: McGraw-Hill. Foucault, Michel. 1971. L’ordre de discours. Leçon inaugurale au Collège de France prononcée le 2 décembre 1970. Paris: Gallimard. Gärtner, Carl Friedrich. 1848. Versuche und Beobachtungen über die Bastarderzeugung im Pflanzenreich. Stuttgart: Auf Kosten des Verfassers. Herbert, William. 1837. Amaryllidaceae: Preceded by an Attempt to Arrange the Monocotyledonous Orders, and Followed by a Treatise on Cross Bred Vegetables and Supplement. London: Ridgway. Jacob, François. 1970. La logique du vivant. Paris: Gallimard. Kölreuter, Joseph Gottlieb. 1761. Vorläufige Nachricht von einigen das Geschlecht der Pflanzen betreffenden Versuchen und Beobachtungen. Leipzig: Gleditschischen Handlung. _____. 1763. Fortsetzung Der vorläufigen Nachricht von einigen das Geschlecht der Pflanzen betreffenden Versuchen und Beobachtungen. Leipzig: Gleditschischen Handlung. _____. 1764. Zweyte Fortsetzung der vorläufigen Nachricht von einigen das Geschlecht der Pflanzen Betreffenden Versuchen und Beobachtungen. Leipzig: Gleditschischen Handlung. _____. 1766. Dritte Fortsetzung der vorläufigen Nachricht von Einigen das Geschlecht der Pflanzen Betreffenden Versuchen und Beobachtungen. Leipzig: Gleditschischen Handlung. Linnaeus, Carl. [1751] 1764. “Plantae Hybridae [Diss., Resp. J. J. Haartmann].” In Caroli Linnaei Ammoenitates Academicae, seu dissertationes variae physicae, medicae, botanicae antehac seorsim editae. Holmiae: Salvius. 28-62. _____. [1755] 1788. “Metamorphoses Plantarum [Diss., Resp. N. E. Dahlberg].” In Caroli Linnaei Amoenitates Academicae, seu dissertationes variae physicae, medicae, botanicae antehac seorsim editae. Erlangae: Jo. Jacobus Palm. 367-86. Mendel, Gregor. 1866. “Versuche über Pflanzen-Hybriden.” Verhandlungen des Naturforschenden Vereins zu Brünn 4 (1865): 3-47. Monaghan, Floyd V., and Alain F. Corcos. 1990. “The Real Objective of Mendel’s Paper.” Biology and Philosophy 5: 267-92. Müller-Wille, Staffan. 1998. “‘Varietäten auf ihre Arten zurückführen’ - Zu Carl von Linnés Stellung in der Vorgeschichte der Genetik.” Theory in Biosciences 117: 346-76. _____. 2001. “Gardens of Paradise.” Endeavour 25 (2) : 49-54. Olby, Robert C. 1979. “Mendel No Mendelian?” History of Science 17: 53-72. Orel, Vítezslav, and Daniel L. Hartl. 1994. “Controversies in the Interpretation of Mendels Discovery.” History and Philosophy of Life Sciences 16: 436-55. Orel, Vítezslaw. 1996. Gregor Mendel: The First Geneticist. Oxford: Oxford University Press. Sageret, Augustin. 1826. “Considérations sur la production des hybrides, des variantes et des variétés en général, et sur celles des cucurbitacées en particulier.” Annales des Sciences Naturelles, 1st ser. 8: 294-314.

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Comments on the papers given by Roger Wood and Staffan Müller-Wille Raphael Falk

I wish to start by noting the major contribution of Víczeslav Orel to our understanding of the history of heredity at and prior to Mendel’s time, and especially of heredity and of breeding, dealt with in this session. It is with much frustration that I deplore his inability to attend this meeting and I wish to extend to him my – and I am sure all of your – wishes for better health and many more contribution to our understandings of the cultural history of heredity. In a way, this session, and perhaps the whole workshop may appear to be disappointing to a Mendel scholar: It was nice to stick to the legend that Mendel’s work sprung out of nowhere, like deus ex machina. But, of course, every serious person knows that this was not the case. Foucault’s comment on Mendel as un monstre vrai notwithstanding, outstanding an individual as Mendel was, his contributions were not “alien to the biology of the epoch,” but rather deeply rooted in his intellectual and social surroundings – as Orel has been emphasizing for many years. The topic of this session, “Heredity and breeding,” on which I wish to comment, was appropriately represented by a two prong assault: Roger Wood’s, from the perspective of the breeder, and Staffan Müller-Wille’s from that of the academics. Not less significant, both approaches utilized hybridization as the major tool of their assault.

* Carlos López-Beltrán, in his Introductory Remarks, pointed out that interest of scientists in heredity happened only towards the beginning of the nineteenth century, when, from a secondary, disposing component it became a main, if not the main, cause of human “nature”. Obviously, no single event in the eighteenth century could bring about the shift that precipitated such a centrality of the problem of hereditary. Developments like Altströmer’s 1723 initiative to breed high quality Spanish sheep in Sweden in spite of this country’s inhospitable environment, the publication of Linnaeus’s 1735 Systema naturae on the classification of animals and plants according to their immanent characteristics, as well as Kant’s 1775 Races of Men notion of the organism as an autonomous entity, were undoubtedly major consequences of the causes that changed the attitude to the notion of intrinsic, essential characters of living beings. Heredity was always a notion of a duality of both continuity and change; and I would like to suggest that it was an increasing consciousness of the inherent variability of human nature during the eighteenth century that was the process, which eventually propelled heredity to the center of intellectual discourse towards the end of the century. Stephen Toulmin epitomized this process in his thought-provoking book, Human Understanding, by an event that took place on April 13 1769. On that day Captain James Cook arrived in Tahiti on H.M.S. Discovery. This event acquired “a retrospective significance that no one could have recognized at the time.” This was not so much because of the party of scientists assembled under the auspices of the Royal Society for astronomical observations “needed to make good one last key detail hitherto missing in the orthodox Newtonian picture of God’s astronomical Creation,” as to the discovery that

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Life on Tahiti, though tranquil enough, was surprisingly free and easy: the inhabitants got along happily, disregarding many of the taboos and commandments regarded in Europe as essential elements of god’s Order […] From this point on, it became progressively more difficult to square men’s factual knowledge about theories and practices of their own fellow humans with accepted views about the impartial standpoint from which rational judgement and appraisal were to be conducted.1

This increasing awareness of natural variation between human societies, and particulate uniqueness of the individual, at odds with the prevailing social and scientific law-given Godly classifications culminated a century later, on November 24, 1859 in the publication of Charles Darwin’s The Origin of Species by Means of Natural Selection, and on February 8 and March 8, 1865 in Mendel’s presentation of his Versuche über Pflanzen-Hybriden. Darwin as well as Mendel coped with the mechanisms of inherent change and continuity on a new ideological and theoretical background than their predecessors a century earlier. Of course: The duality of continuity and change of living beings struck humans as far back in history as we can follow. In biblical times, however, these were conceived as patterns of reproduction, denoting repetition of conserved qualities, and generation, denoting creation of new qualities: “And Adam lived thirty and a hundred years, and begot a son in his own likeness, after his image; and he called him Seth.”2 Inheritance was a concept of material and social transmission of commodities, land, and eventually titles, from person to others: And the Lord gave unto Israel all the land which he sware to give unto their fathers.3 But Zelophehad […] had no sons, but daughters; […] And they came before […] Joshua the son of Nun, […] saying, “The Lord commanded Moses to give us an inheritance among our brethren.” Therefore, according to the commandment of the Lord he gave them an inheritance among the brethren of their father.4 Rachel and Leah answered [Jacob] and said to him, “Is there yet any portion or inheritance for us in our father’s house? Are we not counted of him strangers? […] For all the riches which God hath taken from [Laban] our father, that is ours, and our children’s”.5

Only rarely did inheritance refer to the transmission of traits of creatures, and then as a rule, metaphorically. Until the eighteenth century passive conservation as opposed to active creation had been the leitmotif in the notion that is subsumed toady under “biological heredity”. As Wolfgang Lefèvre has already indicated in the “Abstracts” of the Workshop, the scientific interest in heredity has always been shaped by the social and political theoretical context in which it acted. It appears that the “biologization” of heredity reflected the traditional society’s powercenters embracing themselves, through a kind of God-given typology, against the imminent rocking of the social and political boat; claiming for the inborn nature of their privileged status. I 1 2 3 4 5

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Toulmin (1972), p. 42. Genesis 5:3. Joshua 21:43. Joshua 17: 3-4. Genesis 31: 14-16.

Comments on the papers given by Roger Wood and Staffan Müller-Wille

think that this is also what Staffan Müller-Wille meant in his conclusion that “Heredity becomes central in the early nineteenth century not as a result of social processes in which dichotomic exclusions become cemented, but rather as a result of processes in which such structures become dissolved”.

** Hybridization, the time-honored tool of animal and plant breeders, was considered an act conflicting with the God-given order of natural reproduction. However, just because hybridization violated a basic notion of life that like engenders like, it also high-lightened the important property of living matter that whereas progeny resemble their progenitors in some properties, they may vary from them in others. It was this realization that eventually turned hybridization into a tool for defining discrete characteristics and provided a heuristics for investigating them. Müller-Wille traced the academic prong of the employment of hybridization. The problem of the fixity of the “species” concept in spite of the observed variation became a central issue of academic research in the century following Linnaeus’s Plantae hybridae of 1751. To the extent that it was considered at the level of experimental manipulation, the main tool available was the breeders’ hybridization: Linnaeus reasoned that the only way to change God’s species was by such non-godly acts as hybridization. Indeed, by the very nature of the problem, when dealing with the transformation of species, most of the crosses were made between species. And although the definitions of species and varieties were based on other criteria than the current ones, the dominating experience was that the great majority of the crosses was sterile or produced sterile offspring. Of the 62,688 flower hybridizations carried out by C. F. Gärtner in the years 1823-1848 only 23,335 or 37.22% gave fruits, and in only 257 of these, or 0.41%, the fruits were normal in form and seed number.6 This kind of academic research could hardly resolve the problems of the hereditary identity or dissimilarity of “element characters.” Although Kölreuter tried to discern between “natural species” and “varieties of natural species,” the breakthrough came only in 1859 with Darwin, who, not surprisingly, took his cues from “variation under domestication,” i.e., from the breeders’ rationality. Darwin abandoned the Linnaean distinction between species constant characters and nonspecies’ variable characters, which led him to doubt the demarcation of species and subspecies. Yet, it must be recalled that Darwin never challenged the conception of natural types. His notion was essentially neo-Aristotelian: The properties that maintained species as distinct typological entities came from within, and were inherent in the hereditary-sociological relationship of the members of the species in their environment. His attack on the problem of heredity, like that of Kölreuter or Gärtner, was an academic one, and not a too successful one at that. Now, breeders too encountered from the beginning of time the paradoxical reality of constancy and change of animals and plants. If there was no constancy, breeding must be started anew every generation. Indeed, they believed that unless you provide the bred animals and plants with the proper conditions all the time, they would not yield the expected products. Yet breeders 6

Falk, (1991), p. 460.

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were able to demonstrate some hereditary stability. Whereas the scientists got stranded with problems of fertility, like Kölreuter, with his Dianthus color variations, breeders were practical persons emphasizing the constancy of their products over generations, i.e., biological inheritance sensu stricto. Breeding and controlling the environments were going hand in hand. Their practices, although affected by scientific ideas, were mainly based on traditions and common beliefs and personal experiences and constrained by economic demands. As has been pointed out by Roger Wood the tradition of transmission of traits through the blood and then through the semen was more or less the unchallenged breeders’ concept of inheritance until the eighteenth century. It must also be kept in mind that although many of the more successful breeders, like Bakwell, were versed in the developments in the sciences especially of physiology and embryology, and in the methodology of controlled experiments, their work as a rule did not allow them to apply many of these: Certainly they could not afford the luxury of running “controls”, as every biologist trained in the reductionist tradition of the physical sciences would have been expected to do. No wonder that Wood repeatedly referred to Bakwell as working “like an artist”, rather than as a scientist. In line with this, Bakwell’s approach to problems of heredity was more ‘holistic’ than that of the academics who strove to reduce problems to their elements, or as Müller-Wille called it, to the “atomization” of species, shifting “the level of analysis from specimens to differential characters as the ‘elements’ of the species”.

*** Thus, the changes in eighteenth century European breeders’ concepts were largely technologydriven and toward improved rearing conditions rather than theory-driven. By and large, they could not be expected to keep pace with discoveries in physiology. On this background the one exception in Roger Wood’s description is most remarkable. As he notes, the Moravian “Association of Sheep Breeders” established in Brno in 1814 was interested in genetische Gesetze der Natur. Thanks to some outstanding persons, Brno became the hotbed for a new science of heredity, combining the traditions of the art of animal and plant breeding and the newly achieved developments in scientific reasoning and experimentation. The exertion for the conceptual bridging of the dichotomy may best be summarized in the quotation that Wood brings from Napp in 1837: “What we should have been dealing with is not the theory and process of breeding. But the question should be: What is inherited and how?” Being a member of the Brno Society, Mendel the scientist had access to the breeders’ mentality and, contrary to the academic hybridists before him, he was not handicapped by the taxonomic status of individual variation and the question of whether new species could arise from hybridization. On the other hand, being trained in the physical sciences, Mendel was able to sublime the breeders’ holistic conceptual framework and to adopt a strict reductionist analytic attitude, following the inheritance of discrete characteristics per se. Thus, coming back to Foucault, even on this background, Mendel’s work, exceptionally combining diverse disciplinary elements, such as statistics, cell-theory with advanced breeding techniques, tags him not as a “monster” but as a real giant, in his attempt to discover God’s laws “written in the language of mathematics.”7

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References Falk, Raphael. 1991. “The dominance of traits in genetic analysis.” Journal of the History of Biology 24 (3): 457-484. _____. 2001. “Mendel’s hypothesis.” In G. E. Allen and R. M. MacLeod, eds. Science, History and Social Activism: A Tribute to Everett Mendelsohn. Dordrecht: Kluwer Academic Publishers. 77-86. Toulmin, S. 1972. Human Understanding. The Collective Use and Evolution of Concepts. Princeton NJ: Princeton University Press.

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Falk (2001).

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Acquired character: the (pre-genetic) material of the ‘self-made man’ Paul White

It is Character which builds an existence out of circumstance. Our strength is measured by our plastic power […] thus it is that in the same family, in the same circumstances, one man rears a stately edifice, while his brother […] lives for ever amid ruins. (G. H. Lewes, Life of Goethe)

Historians have located a widespread shift in Victorian Britain from models of hierarchical order based on innate capacity – the blue blood of nobility – to one in which character of the highest sort could be acquired. Debates of this sort were much older, but the story is often told as part of the rise of the middle-classes and the professions and the consequent decline of the hereditary aristocracy.1 This social transformation was far from straightforward, and models of inborn and acquired character often overlapped or were employed in different ways according to setting. Ongoing debates about the role of ancestry, of landed property, of money, and of moral conduct in the constitution of social status often focussed on the figure of the gentleman. The inheritance of acquired characteristics, prior to the isolation of chromosomes as the material of hereditary transmission, has often been treated virtually as a timeless theory with an inherent plausibility, and therefore requiring no historical explanation.2 And yet the natural historical concept of character as developed in studies of breeding and hybridization was closely linked to prevalent concerns among the middle classes of the Victorian period about inborn or acquired character in the wider, moral sense. Research on heredity in Victorian Britain may thus be situated within a wide-ranging set of debates about self-formation, individual improvement, and the formation of gentlemanly character. The concept of the hereditary transmission of acquired characters, so crucial to progressive theories of evolution and to theories of degeneration in the period, must itself be understood in relation to the broader controversy over the acquisition of (moral) character.

Breeding character The term “character”, together with “characteristic”, has been used in natural historical contexts to signify a distinctive feature of a species or variety since the mid 18th century, although its wider meaning as an “essential quality” and its use with reference to moral and mental attributes begins much earlier. In Victorian discourses of breeding, these two senses, the natural historical and the moral, are considerably intertwined. What was “character” for the breeders? At one level, it was simply a physical or behavioural trait, taken as distinctive of a given variety: a particular shape of beak or horn, the sheer size and mass of an animal’s body, a manner of masculine display, such as the inflating of the chest. Such traits were not only variable, but mutable, and breeders often 1 2

For example, Perkin (1989); and Reader (1966). On the inheritance of acquired characteristics, see Bowler (1989); Churchill (1976) and (1987); Lomax (1979); Robinson (1979); Windholz (1991); and Zirkle (1946).

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presented themselves as able to shape a plant or animal’s character, almost at will. Thus the veterinary surgeon, William Yuoatt, spoke of selection as “that which enables the agriculturalist, not only to modify the character of his flock, but to change it altogether. It is the magician’s wand, by means of which he may summon into life whatever form and mould he pleases.” 3 This power to mould character through the manipulation of conditions and the exercise of will, was in fact the operative principle of another Victorian creature: the self-made man. Elaborated in numerous didactic works and novels of the early Victorian period, the culture of self-help and self-fashioning was epitomised in the writings of Samuel Smiles, first in his biography of the manufacturer George Stephenson, in many subsequent lives of working men-made-good, and a series of companion volumes on particular virtues such as thrift, duty and, in 1871, on Character itself.4 According to Smiles, character was “formed by a variety of minute circumstances, more or less under the regulation and control of the individual.” Every action, thought, and feeling contributed to its formation: “man is not the creature, so much as he is the creator, of circumstances […] energy of will – self-originating force – is the soul of every great character.” By locating the virtues of moral character in individual deeds and habits, rather than in ancestry, Smiles legitimized, indeed lionized, the rapid social ascent of Britain’s industrial and commercial leaders, while holding out the promise of improvement to many. Revealed in the “transactions and commonplaces of daily duties”, a respectable character could be acquired by anyone.5 How then was this culture of individual improvement and character formation operative in the discourse and practice of breeders? One way of getting at this connection is to look at the various forms by which character, both in its natural historical and moral senses, was displayed.

Fig. 1: Short-horn bull from Bates, The history of improved short-horn or Durham cattle, and of the Kirklevington herd, (1871), p. 354 facing.

3 4 5

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Youatt (1837), p. 60. For a general discussion of Smiles’s work, see Jarvis (1997). On the literary construction of the self-made man by Smiles and others, see Meckier (2001); Pettitt (1999); Rodrick (2001); and Wyke (1999). Smiles (1871), pp. 1-6.

Acquired character: the (pre-genetic) material of the ‘self-made man’

Fig. 2: Pedigree from Biddell, The Suffolk stud-book: a history and register of the county breed of cart horses, (1880), p. 654 facing.

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This prize short-horn bull (fig. 1), the VIIIth Duke of York, is exhibited here in Thomas Bates’s 1871 history of the improved Durham cattle. Harriet Ritvo’s work on animal husbandry in Britain has shown that breeders often affixed their own social status to that of their animals; in breed characteristics, no less than prize competitions, the criteria of social hierarchy were defined, and the boundaries established between a person (or animal) of noble pedigree and a parvenu. 6 The nobility of a creature was always demonstrated by its ancestry and progeny, as in this pedigree (fig. 2) of Canterbury Pilgrim from the Suffolk Stud Book. Such registers typically added a further layer of representation, in the form of biographical entries, noting the prizes that the animal had won, the prices it had fetched, and also its leading characteristics. Thus Cup-Bearer (fig. 3) was depicted as “a large horse with a grand fore end, great depth of girth, and splendid muscular shoulders. […] although not an elegant walker,” its sons were legion, numbering many prize winners. The animal is shown here with its owner, Mr. Crisp.

Fig. 3: “Cup-Bearer” from Biddell, The Suffolk stud-book: a history and register of the county breed of cart horses, (1880), p. 166 facing.

6

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Ritvo (1987), pp. 45-63. On general breeding practices in Britain, see also Russell (1986).

Acquired character: the (pre-genetic) material of the ‘self-made man’

Fig. 4: Prize bull of Sir Charles Morgan from Ritvo, The animal estate: the English and other creatures in the Victorian age, (1987), p. 49.

Fig. 5: Charles and Robert Colling from Shorthorn transactions, (1868-9), frontispiece.

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This sort of iconography could assume a more elaborate and costly form, as in this oil on canvas (fig. 4), drawing on the tradition of gentry portraiture, in which lords of the manor were foregrounded against their estates.7 But this animal, the prize bull of Sir Charles Morgan, baronet, is not just part of the gentleman’s large holdings; its distinguishing properties – its formidable size, beauty, and elegance – were those of nobility, and were also distinctive of its owner. Corpulence, in conjunction with elegance, was a virtue among the English gentry, signifying power, wealth, and generosity.8 Typically displayed by a rounded, decorous body on slender legs, as in this portrait of Charles and Robert Colling (fig. 5). These “fathers” of the short-horn cattle manifest the same prodigious girth and delicacy of limb (or calf) as their noble breed. To complete this system of representation, on the page opposite in this volume of Shorthorn Transactions, their own moral pedigree is given. Thus Robert, “from his enlarged views, generous disposition, love of order, nobility of bearing, and extensive means […] was called the‘Prince of Skerne’”. 9 The role of breeders in modifying structure through habit and conditions thus bore equally on their own status as gentlemen and conservative improvers. These forms of representation, the family tree, portraiture, and biography, together enforced the view that gentlemanly character was natural historical – composed of heritable traits, rooted in good breeding – and yet highly malleiable in the breeders’ art. The great estate owners who commanded this culture of improvement, some of whom were new members of England’s ruling classes, were, as much as their prize animals, living examples of the power of selective breeding to modify character, and thereby enrich the stock of the nation.

Powers of discrimination In response to criticism that in Origin of Species he had described natural selection as an agent, productive of species, Darwin replied that he had merely adopted the language of breeders. 10 James Secord has shown the attention to the work of animal and plant hybridists in the framing of Darwin’s theory; and indeed, the skill and art of breeders in fashioning their stock was repeatedly emphasized in Darwin’s acknowledgements of individual fanciers and horticulturalists in the Variation of Plants and Animals Under Domestication.11 As Darwin wrote in Origin, they “habitually speak of an animal’s organisation as something quite plastic, which they can model almost as they please.”12 In drawing the analogy between natural and artificial selection, Darwin conceded a power to breeders that was a source of consternation among some of his gentlemanly correspondants, such as Charles Lyell. On reading the first chapter of Origin, Lyell remarked: Your comparison of Selection to the Architect […] I […] cannot accept. The architect who plans beforehand & executes his thoughts & invents […] must not be confounded in his functions with the humble office of the most sagacious of breeders […] it is the deification of Natural Selection. 7 8 9 10 11 12

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Barrell (1986). On the virtue of corpulence, see Porter and Rousseau (1998). Shorthorn transactions, (1868-9), frontispiece. Darwin (1859), p. 30; Darwin to Charles Lyell, 22 January 1865, in Burkhardt et al. (1985-2002), vol. 13. Secord (1981), and (1985). Darwin (1859), p. 31.

Acquired character: the (pre-genetic) material of the ‘self-made man’

The idea of [selection] being ever allowed to play such pranks as the breeder has played & to sport with God’s creatures & the laws of reproduction so as to perpetuate pouter pigeons, & other monstrosities, would have been scouted by a philosopher of the Wealden Period.13

The deificiation of natural selection, and by implication, the elevation of breeders to creative status, remained an ongoing dispute. Darwin actively defended his analogy as appropriate. Yet Darwin’s views on the relative power of external agency to shape the character of species and varieties differed substantially from those of the leading breeders whom he consulted. While agreeing that the conditions of life, especially those of domestication, had profound effects on the variability of character, Darwin urged “we must never forget that the nature of the organisation which is acted on, is by far the most important factor in the result.” 14 Unlike the horse and cattle breeders, the majority of the poultry men with whom Darwin associated were or had been artisans, a point of considerable importance to a community which prided itself on gentlemanly character. Though a tailor and thus an artisan by trade, John Matthews Eaton, one of Darwin’s leading authorities in Variation, clearly regarded himself and his fellow fanciers as gentlemen – not by birth, or wealth, but by virtue of their pasttime. Some gentlemen, even nobleman, were indeed breeders – a fact reiterated in countless treatises on domestic animals; and breeding pigeons did allow tradesmen and others of more humble status to imitate the genteel and noble. Yet the crucial point of Eaton’s claim, however, was that the practice of breeding was itself gentlemanly in character. It was a “study and science” especially “adapted to the professional gentlemen of law, physic, and divinity, or any other person engaged in long continued and excessive exertion of the intellectual faculties.” 15 Breeders were renowned for their unsurpassed powers of observation and discrimination, able to detect differences of character between individuals that were invisible to anyone not a master of the art. Their way of seeing, their watchfulness, vigilance, and discrimination were as important to Darwin as their knowledge of variation and inheritance; for they embodied nature’s selective power. Darwin’s own account of the changes wrought by breeding practice in Origin of Species culminated in a character-sketch of the breeders as self-made men, succeeding according to the Smilesian virtues of perseverance, hard work, and the resolute pursuit of a single goal: Not one man in a thousand has accuracy of eye and judgment sufficient to become an eminent breeder. If gifted with these qualities, and he studies his subject for years, and devotes his lifetime to it with indominatable perseverance, he will succeed, and may make great improvements; if he wants any of these qualities, he will surely fail. Few would readily believe in the natural capacity and years of practice requisite to become even a skilful pigeon-fancier.16

And yet Darwin’s own relations with breeders such as Eaton clearly indicates that he regarded them as social inferiors, joking privately about their diminutive stature and their bad grammar. To his eldest son, he wrote: “NB all Pigeon Fanciers are little men.” 17 When the zoologist Thomas 13 14 15 16 17

Charles Lyell to Charles Darwin, 15 June 1860 and 19 June 1860, in Burkhardt et al. (1985-2002), vol. 8. Darwin (1868), 2: p. 502. See also Winther (2000). Eaton (1851), pp. iii-vi. Darwin (1859), p. 32. Charles Darwin to William Erasmus Darwin, 29 November 1855, in Burkhardt et al. (1985-2002), vol. 5.

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Huxley asked Darwin for information on breeding for a talk at the genteel Royal Institution, Darwin included some extracts from Eaton’s treatise on the Almond Tumbler that he thought “would make the audience laugh.” After citing passages that displayed the gentlemanly pretenses of the fancy, together with the “little tailor’s grammar” to comic effect, Darwin cautioned Huxley not to mention Eaton’s name, “as he is my friend.”18 As a man of wealth and property, and the recipient of a traditional classical education, Darwin could have rested his superiority to the breeders on these grounds, and yet he made common cause with Huxley, of relatively humble birth, poor, and largely self-educated. That Darwin and Huxley could laugh at the breeder’s expense, and yet regard them as friends, and look to them as authorities on questions of science, raises questions about the social status of their own experimental practice, and the grounds on which they distinguished themselves from such associates. The particular gifts that Darwin praised in breeders, their accuracy of eye, and powers of concentration and discrimination, were also widely regarded as characteristic of the Victorian ‘man of science’.

Hereditary genius How then did the ‘man of science’ stand in relation to models of innate and acquired character? In fact, just like the breeders, the status of men of science as gentlemanly, and as self-made, was also uncertain. A romantic tradition was still operative in Victorian scientific biography and selfpresentation, in which the practitioner was distinguished by specially endowed powers of mind. In his essay Characteristics, Thomas Carlyle described genius as a heroic intellectual force: innate, like nobility, but often residing in those of humble birth. Its characteristics – intuition, mental suppleness, refined discrimination – marked out a few individuals as destined to be leaders of men and spirits of the age.19 Such romantic models were appropriated by practitioners such as Huxley, and John Tyndall, whose backgrounds did not confer gentlemanly status. And yet in the Victorian period, genius was transformed by the culture of industry and work.20 What had been conceived as a largely effortless and innate capacity by 18th century writers became an endowment of the self-made man, who had to labor and struggle after truth. Even Darwin, a gentleman of secluded leisure, continually characterized his scientific activity as “hard work” and “hard labour”, tallying up the months and years, right up to the day, at which he spent in the production of each of his books. 21 Francis Galton’s 1874 survey of men of science compiled over a hundred comparable accounts: men who walked 50 miles a day without fatigue in search of specimens, men who worked habitually until two or three in the morning. Using autobiographical testimony, Galton documented their leading characteristics as perseverance, steadiness, determination, and “the secretion of nervous force” – that is, energy: “many have laboured as earnest amateurs in extra professional hours long into the night […] they have climbed the long and steep ascent from the lower to the upper ranks of life.” 22 18 19 20 21

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Darwin to Thomas Huxley, 27 November 1859 and 13 December 1859, in Burkhardt et al. (1985-2002), vol. 7. Carlyle (1831). On the cult of genius and work amongst Victorian men of science, see White (2002), ch. 1. See also Tyndall (1868). Darwin (1958).

Acquired character: the (pre-genetic) material of the ‘self-made man’

Yet, despite this emphasis on self-fashioning through hard work, certain characteristics of men of science remained inborn. In his 1869 work, Hereditary Genius, Galton used his own family history, and the pedigrees of others, to argue that intellectual aptitude was hereditary. Galton drew heavily on the tradition of family trees to map out the inheritance of illustriousness over generations, as in this pedigree of the natural philosopher, Robert Boyle (fig. 6). Like Carlyle, Galton viewed men of genius as great forces of the age, but recast their influence in terms of hereditary transmission, as the sports of nature by which the new higher types of mankind were established. In his theory of sports, a new character suddenly made an appearance in one individual and was transmitted, causing a change in the typical centre of the race and a step forward in the course of evolution.23

Fig. 6: Pedigree of Robert Boyle from Galton Hereditary genius, an enquiry into its laws and consequences, (1869), p. 197.

Diseases of character In the Preface to the 1892 edition of Hereditary genius, Galton noted recent work by Lombroso and others that traced the proximity of genius to insanity, commenting that he did see some evidence in families for a “painfully close relation between the two” conditions. 24 The very characteristics of mind possessed by leaders of humanity, the geniuses and self-made men, brought them close to less favourable qualities, commonly identified with degeneration. I want to look at one form, or stage, of degeneration – alcoholism – for it became in the Victorian period a particular malady of the self-made man. This appears expecially prominent in the medical literature and novels of the mid-Victorian period, decades of great prosperity for the middle classes. 22 23 24

Galton (1874), p. 75. Galton (1892), pp. xvii-xix. See also Cowan (1985). Galton (1892), p. ix.

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Anthony Trollope’s Doctor Thorne problematizes the simple Smilesian version of character as a product of self-determination, as well as the traditional equation of gentlemanly character with noble descent.25 The novel contains two self-made men: one a paragon of Smilesian heroism, a lowly stonemason who rises to be a captain of industry, building bridges, canals, and railroads across the British Empire, and is made a baronet – a hereditary title, though significantly not an aristocratic one. Though conquering the world, Roger Scatcherd succumbs to the habit of drink and dies, leaving a sickly son, who inherits the craving for drink, suffers delirium tremens, and dies at the age of twenty, extinguishing the family line. The other, of ancient family, who prides himself on being a poor man of high birth, and yet who rises to eminence as a physician through displays of great generosity, honesty, and duty. It is Thorne’s belief in his superior birth, rather than any necessary inherited tendency, that motivates his gentlemanly conduct. Similarly, Scatcherd’s hereditary malady is closely linked to his own entrepeneurial manner of self-fashioning. Thus Trollope implies that inheritance can operate as a cushion against degeneration in a period of rapid social change; but it does so as a form of culture – as a structure of beliefs, values, and behaviours. The problem of alcoholism is medicalized to some degree in the novel. Dr. Thorne is Scatcherd’s physician. The physical effects of the toxin are delineated and inescapable. And yet the doctor is ultimately powerless to effect a cure without the patient’s cooperation. His methods, consisting largely in severity and admonishment, are ultimately ineffectual either on the father or the son. Nor is the condition, following the pattern of a congenital disease, clearly separated from the social world it inhabits. As Trollope remarks, had Scatcherd or his son been born a duke or an earl, provision would have been made for such behaviour – it would have been redefined and accomodated within the culture of aristocracy: great bouts of drinking were appropriate at the parties and banquets of great houses, and within gentlemen’s clubs. Within the highest circles, occassions for such social drinking could be multiplied almost indefinitely. But Scatcherd has no place in this level of society. He is an alcoholic, at least in part, because he is not, despite his social climb, a gentleman. Alcoholism is in fact a disease of the self-made man. Though a propensity in Scatcherd when still an apprentice mason, the condition is worsened during the course of his rise to eminence, with each bout of heavy drinking coinciding with the winning of a new government contract. Trollope’s account is borne out by medical literature of the period, which linked the onset of the condition to the pressure of work, noting its high concentration among the merchant, manufacturing, and professional classes. Once widely regarded as a problem of the lower orders, of the poor and destitute, medical theory now suggested that poverty was a result of habitual drinking, not a cause. Widely treated as a hereditary or congenital condition from the early Victorian period onwards, alcoholism remained the most controversial of ‘diseases’ because of its relationship to the will. Medical treatments and physiological research that identified alcohol with a material toxin nevertheless continued to regard alcoholic subjects as morally responsible for their illness or cure.26 In an 1850 essay, the physiologist and medical author William Carpenter, characterized the disease as a premature exhaustion of nervous power, a weakening of the 25 26

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Trollope (1858). Bynum (1984); Huertas (1993); Levine (1978); Valverde (1997); and Warner (1994).

Acquired character: the (pre-genetic) material of the ‘self-made man’

controlling power of the will, and an augmenting of the automatic or impulsive part of nature, giving a constitutional predominance to the lower feelings and passions. Habitual intemperance thus produced a progressive degradation of moral character by repeated excitement of the lower propensities and diminution of the will, leaving the person in “a state of complete [self-induced] slavery”.27 Alcoholism was a preeminent case illustrating how habits acquired by individuals, and transmitted to their offspring could progress over generations into more degenerate conditions, such as criminality and madness.28 In the second half of the century, a medical speciality grew up around the condition, closely allied with alienists and asylum culture, and more elaborate medical treatises were produced over the course of the second half-century. Perhaps the most exhaustive study, by the physician Norman Kerr, pronounced that no “natural law was more patent than [that] of alcoholic heredity”. Men and women of the highest culture, the most irreproachable morals, and the strongest will, had come to know that they could “never dally with strong drink”, and that doing so would in fact alter their character, and turn “perfect gentlemen” into criminals or lunatics. And yet, despite the inexorable material and hereditary basis of the disease, Kerr cast the alcoholic within the terms of the Smilesian self-made man, his cure an epic struggle of will against an oppressive disease: “The continuous and victorious struggle of such heroic souls with their hereditary enemy – an enemy the more powerful because ever leading its treacherous life within their breasts, presents to my mind such a glorious conflict, such an august spectacle, as should evoke the highest efforts of the painter and the sculptor.” 29 Other treatises on alcoholism typically included chapters on the treatment of the disease through a strict moral regime, involving the modification of habits and conditions of life in order to reinvigorate the will. 30 Such medical accounts of alcoholism bring to the surface the fundamental circularity of the Smilesian account of character: it was both a property (a “noble estate”) and a product, a cause and an effect. The self-made man was formed through a process of struggle, and the overcoming of hardships; and yet one could only succeed amidst life’s difficulties if one’s heart was strong and upright, and one’s spirit lively – in other words, unless one was already endowed with a noble character.

Conclusion: a new biometrics Tying together theories of heredity, acquired character, and degeneration, the medical literature on alcoholism would figure prominently in early eugenic writings. Like the pedigrees of prize animals, noble families, and hereditary genius, eugenics-inspired studies of the hereditary effects of alcoholism were displayed as family trees (fig. 7). This chart of alcoholic degeneration was drawn at the beginning of the 20th century by the research committee of the Eugenics society. Significantly, the other modes of representation through which the characters of the great and good had been displayed – biography and portraiture – were now absent.

27 28 29 30

Carpenter (1850), p. 47. Huertas (1993); Lubinsky (1993). Kerr (1894), pp. 17, 55. Carpenter (1853); Clouston (1884); Maudsley (1874); Sankey (1884); Talbot (1898).

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The families themselves were not sired by self-made men, merchants, or professionals, but by paupers. Like the social surveys and industrial novels of the 1840s, the problem of alcoholism at the end of the century, a period which saw the rise of socialism, of the labour movement, and the formation of a Labour Party, was again located in the lowest ranks of society. 31 And the pedigrees themselves, which had served partly as forms of self-fashioning for members of the gentry, professionals, and some artisans, were now generated in biometrics laboratories, asylums and work houses, with the aim of enlisting the State to direct the improvement of the race or nation. In order to determinde the laws of heredity, Galton urged, it was essential both to deal statistically with populations, and to regard human characters like the traits of a pea (figs. 8 and 9). These tables from Galton’s Natural Inheritance show the distribution of eye colour across three generations, and the diameters of parent and filial seeds of the sweet pea. This paring down of ‘data’, in favour of symbols denoting the presence or absence of a single characteristic, is of course a style of representation that would become fundamental in modern genetics and population biology. The hyperbolic claims of Victorian breeders to shape an animal’s nature would make few genetic engineers of today blush, while physicians, alienists, and eugenicists like Galton, the new custodians of the poor, seemed to appreciate that good breeding depends very much on who descriminates between desirable and undesirable characters, and on what grounds.

Fig. 7: Pedigree from Mazumar, Eugenics, Human genetics and human failings: the Eugenics Society, its sources and its critics in Britain, (1992), p. 81. The original source is A. F. Tredgold, Mental Deficiency, (London, 1908).

31

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Hawley (1989); Mazumar (1992).

Acquired character: the (pre-genetic) material of the ‘self-made man’

Fig. 8: Table over eye-colour from Galton, Hereditary genius, an enquiry into its laws and consequences, (1892), p. 206.

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Fig. 9: Table over seed-size from Galton, Hereditary genius, an enquiry into its laws and consequences, (1892), p. 226.

References Barrell, John. 1986. The political theory of painting from Reynolds to Hazlitt: ‘the body of the public’. New Haven: Yale University Press. Bates, Thomas. 1871. The history of improved short-horn or Durham cattle, and of the Kirklevington herd. Newcastle: Robert Redpath. Biddell, Herman. 1880. The Suffolk stud-book: a history and register of the county breed of cart horses. Diss: Francis Cupiss. Bowler, Peter J. 1989. The Mendelian revolution: the emergence of hereditarian concepts in modern science and society. Baltimore: Johns Hopkins University Press. Burkhardt, Frederick, et al. 1985-2002. The correspondence of Charles Darwin. 13 volumes. Cambridge: Cambridge University Press. Bynum, W. F. 1984. “Alcoholism and degeneration in nineteenth-century European medicine and psychiatry.” British Journal of Addiction 79: 59-70. Campbell, Margaret. 1983. “The concepts of dormancy, latency, and dominance in 19th-century biology.” Journal of the History of Biology 16: 409-431. Carlyle, Thomas. 1831. “Characteristics.” Edinburgh Review 51: 351-383. Carpenter, William. 1850. On the use and abuse of alcoholic liquors, in health and disease. London: Charles Gilpin. _____. 1853. The physiology of temperance & total abstinence. London: Henry Bohn.

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Churchill, Frederick B. 1976. “Rudolf Virchow and the pathologist’s criteria for the inheritance of acquired characteristics.” Journal of the History of Medicine and Allied Sciences 31: 117-148. _____. 1987. “From heredity theory to‘Vererbung’: the transmission problem, 1850-1915.” Isis 78: 337-364. Clouston, T. S. 1884. The effects of the excessive use of alcohol on the mental functions of the brain. Edinburgh: Andrew Elliot. Cowan, Ruth Schwartz. 1985. Sir Francis Galton and the study of heredity in the 19th century. New York: Garland. Darwin, Charles. 1859. On the origin of species by mean of natural selection. London: John Murray. _____. 1868. The variation of animals and plants under domestication. 2 vols. London: John Murray. _____. 1958. The autobiography of Charles Darwin. Edited by Nora Barlow. London: Collins. Eaton, John Matthews. 1851. A treatise on the art of breeding and managing the almond tumbler. London: J. M. Eaton. Galton, Francis. 1869. Hereditary genius, an enquiry into its laws and consequences. London: Macmillan. _____. 1874. English men of science: their nature and nurture. London: Macmillan. _____. 1889. Natural inheritance. London: Macmillan. _____. 1892. Hereditary genius, an enquiry into its laws and consequences. 2nd edition. London: Macmillan. Hawley, John C. 1989. “Mary Barton: the inside view from without.” Nineteenth Century Studies 3: 23-30. Huertas, Rafael. 1993. “Madness and degeneration, part II: alcoholism and degeneration.” History of Psychiatry 4: 1-21. Jarvis, Adrian. 1997. Samuel Smiles and the construction of Victorian values. Stroud: Sutton. Kerr, Norman. 1894. Inebriety or narcomania, its etiology, pathology, treatment and jurisprudence. 3rd edition. London: H. K. Lewis. Levine, Harry. 1978. “The discovery of addiction.” Journal of Studies of Alcohol 39: 143-74. Lewes, George. 1875. The life of Goethe. 3rd edition. London: Smith, Elder, & Co. Lomax, Elizabeth. 1979. “Infantile syphilis as an example of 19th-century belief in the inhertiance of acquired characteristics.” Journal of the History of Medicine and Allied Sciences 34: 23-39. López-Beltrán, C. 1994. “Forging heredity: from metaphor to cause, a reification story.” Studies in History and Philosophy of Science 25: 211-235. Lubinsky, Mark S. 1993. “Degenerate heredity: the history of a doctrine in medicine and biology.” Perspectives in biology and medicine 37: 74-90. Mazumar, Pauline. 1992. Eugenics, Human genetics and human failings: the Eugenics Society, its sources and its critics in Britain. London: Routledge. Maudsley, Henry. 1874. Responsibility in mental disease. London: Henry King & Co. Meckier, Jerome. 2001. “Great Expectations and self-help: Dickens frowns on Smiles.” Journal of English and Germanic Philology 100: 537-54. Pettitt, Clare. 1999. “‘Every man for himself, and God for us all!’ Mrs Oliphant, self-help, and industrial success literature in John Drayton and The Melvilles.” Women’s Writing 6: 163-79. Porter, Roy, and Rousseau, G. S. 1998. Gout: the patrician malady. New Haven: Yale University Press. Ritvo, Harriet. 1987. The animal estate: the English and other creatures in the Victorian age. Cambridge Mass.: Harvard University Press. Robinson, Gloria. 1979. A prelude to genetics: theories of a material substance of heredity, Darwin to Weismann. Lawrence, Kansas: Coronado Press. Rodrick, Anne Baltz. 2001. “The importance of being an earnest improver: class, caste, and self-help in midVictorian England.” Victorian Literature and Culture 29: 39-50. Russell, Nicholas. 1986. Like engend’ring like: heredity and animal breeding in early modern England. Cambridge: Cambridge University Press. Sankey, W. H. O. 1884. Lectures on mental disease. London: H. K. Lewis. Secord, James. 1981. “Nature’s fancy: Charles Darwin and the breeding of pigeons.” Isis 72: 163-86. _____. 1985. “Darwin and the breeders: a social history.” In The Darwinian heritage, edited by D. Kohn. Princeton: Princeton University Press, pp. 519-42. Shorthorn transactions. 1868-9. London: John Thornton. Smiles, Samuel. 1871. Character. London: Murray. Talbot, Eugene. 1898. Degeneracy, its causes, signs, and results. London: Walter Scott Ltd. Trollope, Anthony. 1858. Doctor Thorne. London. Tyndall, John. 1868. Faraday as a Discoverer. London: Longmans. Valverde, Mariana. 1997. “’Slavery from within’: the invention of alcoholism and the question of free will.” Social History 22: 251-268.

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Waller, John C. 2001. “Ideas of heredity, reproduction and eugenics in Britain, 1800-1875.” Studies in history and philosophy of biological and biomedical sciences 32: 457-489. Warner, Jessica. 1994. “’Resolv’d to drink no more’: addiction as a pre-industrial construct.” Journal of Studies on Alcohol: 685-91. White, Paul. 2002. Thomas Huxley: making the‘man of science’. Cambridge: Cambridge University Press. Windholz, George. 1991. “Pavlov’s view of the inheritance of acquired characteristics as it relates to theses concerning scientific change.” Synthese 88: 97-111. Winther, Rasmus G. 2000. “Darwin on variation and heredity.” Journal of the history of biology 33: 425-455. Woiak, Joanne. 1994. “‘A medical Cromwell to depose king alcohol’: medical scientists, temperance reformers, and the alcohol problem in Britain.” Social History 27: 337-365. Wyke, Terry. 1999. “The culture of self-improvement: real people in Mary Barton.” Gaskell Society Journal 13: 85-103. Youatt, William. 1837. Sheep. London: Baldwin & Cradock. Zirkle, Conway. 1946. “The early history of the idea of the inheritance of acquired characters and of pangenesis.” Transactions of the American Philosophical Society N. S. 35: 91-151.

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Kant on Heredity and Adaptation Peter McLaughlin

The consideration of heredity in the latter eighteenth century seems to have focussed primarily on the transmission of defects, especially hereditary disease. Immanuel Kant, too, uses the concept “Vererbung” in this context: he analyses the discussion of the transmission of defects as carried on in the three “higher” university faculties (law, medicine, and theology) with their notions of Erbschuld, Erbkrankheit and Erbsunde (hereditary debt, hereditary disease, and original sin). But he also deals with the concept of “Vererbung” in biology, where it was used in an area with decidedly positive connotations: adaptation. My presentation will deal with the relation of heredity to adaptation in the biological writings of Kant. Individual adaptations – as opposed to the appropriateness of the species form for its place in nature – did not constitute a central theme in eighteenth century biological thought. And even for Lamarck they are of only secondary interest. Kant’s position may be an important point of departure for the study of adaptive heredity and for the quite different relation of nineteenth century thought to the problem of adaptation.

Heredity and Adaptation Neither the heredity nor the adaptation of individual traits was a primary concern of eighteenth century biology. The fit between organism and environment was most often conceptualized as an intrinsic aptitude (of the species form) for life accompanied by a contingent and superficial adaptation of individual characters to aspects of the environment. The realm of the regular, the lawlike, the vitally significant was the basic species form – and this at the time had little to do with heredity. It was unusual to speak of individual traits as adaptations – in fact the first use of the word “adaptation” in this sense mentioned by the Oxford English Dictionary is Darwin’s Origin of Species (1859). Seventeenth and eighteenth century science inherited from classical antiquity a threefold classification of kinds of inter-generational similarity. Individual peculiarities, species form, and sex mark off different ways in which progeny can be similar to parents. The distinction between the first two forms – the lawlike transmission of species form and the contingent disturbance or supplementation of the results of this transmission through the transmission of individual traits – structured early modern theorizing. For instance, the spermatic parts could form an outline that is then fleshed out by the sanguinous parts (Everard), or the inherited species soul could direct the pangenesis of the individual body (Gassendi), or the pangensis of the somewhat etherial species form by “spiritual” or “formal” atoms might govern the pangenesis of the body by the “material atoms” (Highmore), or the species form is encased in the germ while individual peculiarites are transmitted pangenetically with the first nourishment (Bourguet). 1 As a rule the two levels were kept distinct and the species form was what was scientifically interesting. Buffon’s moule interieur, for instance, guaranteed the unity of the species and the boundary lines between different species 1

Gassendi (1658); Highmore (1651); Bourguet (1729), pp. 154-156.

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in spite of all the individual peculiarities that can arise and can be accumulated. Even Lamarck’s later transformation theory still displays this distinction in the two basic processes of organic change: the progressive tendency to increased complexity of organic structure – so to speak a great escalator of being – and the collective adaptation of the individuals to changing environmental conditions that distorts the otherwise lawlike linear progression of forms. In all these theories the species form is fixed either by divine preformation or by natural law; and pangenesis provides a simple mechanism for the transmission of acquired characters: either directly to the newly formed germ or indirectly to a preformed germ’s first nourishment. But the individual peculiarities that may genuinely be said to be inherited are just that – individual and peculiar; they are important not for theoretical science but for the application to individual cases in medicine and agriculture. The species characters, on the other hand, were not in any sense products of a contingent process of heredity. The individual hereditary traits were also normally not viewed as being particularly beneficial to their bearers. The best studied cases in the species that interests us most were the transmission of sixth digits and of hereditary diseases. The fitness of an organism for its environment, adaptation, purposiveness, etc. were properties of the species form. And here too there was a certain tension – at least in the materialistic theories that dominated the second half of the eighteenth century. Whereas deistic preformation theories could imagine that there were innumerable possible organic forms that might have been actualized and that God elected only some of these (presumably the better ones), the materialistic theories of Buffon, Blumenbach and others were committed to the natural necessity of all actually existing species. Their emphasis was on viability not adaptedness, on living, not on living well. According to Buffon, every kind of organism that can exist does exist. The sloth, for instance, is a border-line case; it has just barely everything that it needs for life, but if it had one more defect, it would become extinct. 2 In such a scheme the nature of matter determines species form and its continuity over time: if a catastrophe were to occur and wipe out some or all species, they would return again by spontaneous generation as soon as the environment returned to normal. There is no serious role for heredity or adaptation in such a scheme. We might want to look for the beginnings of adaptationism in physico-theology; the purported influence of Paley’s “argument from design” on Darwin’s theorizing is widely cited. And it is true that this argument did increasingly put single adaptive traits at the focus of attention – though there were limits. While the detailed fit of particular traits to aspects of the environment might evoke amazement at the skill of a technician, it was the greater systems that inspired awe at the grandeur of the Creator. Although there were originally two basically different versions of the design argument – one involving the “design” or plan of the world system and the other involving the “designs” or intentions of the Creator for particular organs –, it was the final cause of particular traits not the formal cause of an orderly world system that to some extent survived Hume’s critique of the design argument.3 But here, too, it is species characters that give occasion to speculation 2

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“We formerly remarked that every thing that possibly could be, really did exist; of which the sloths are a striking example. They constitute the last term of existence in the order of animals endowed with flesh and blood. One other defect added to the number would have totally prevented their existence” (Buffon (1765), p. 40. [“The Sloth,” Smellie translation, 9: pp. 7-8]). See Hume (1779), pt. 2 and 3.

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about the intentions of the Creator. Even physico-theologians had difficulty with individual peculiarities. Albrecht von Haller at one point in his preface to the German translation of Buffon’s Histoire naturelle has God’s general plan produce the species, while his special providence produces the individual details and differences. At other times (in the same text), however, he leaves the details to chance and only the species form to God. What remains constant in Haller’s exposition is the distinction between the two levels of transmission. 4 There wasn’t much theological milage to be gotten out of the individual peculiarities that could be inherited. Since the physico-theologically-minded tended to be preformationists even well into the nineteenth century, and since preformation excludes the species form from the domain of the hereditary, we should not expect much in the way of inheritance of adaptations from this field.

“Vererbung” The concept of heredity seems to have entered German science in the same breath as the concept of race – in the writings of Immanuel Kant. Kant lectured on physical geography once every year for half his life, and in these lectures the concept of race played an important role. In 1775 – announcing what seems to have been his twentieth run through the material – he published a short broschure on human races and their origin: On the different human races.5 Ten years later he published a paper in a Berliner journal trying to defend and clarify his concept of a human race “Determination of the concept of a race of humans”.6 When this paper occasioned severe criticisms by Georg Forster, Kant returned again to the subject in a third essay entitled “On the Use of Teleological Principles in Philosophy” published in early 1788 in the journal Teutscher Merkur. In this paper Kant uses the German term vererben for the first time (in the verb form).7 There is some reason to believe that his use of the term is prototypical – at least I know of no previous biological uses outside of medecine. In the Critique of Judgment of 1790, which contains his philosophy of biology, Kant says little or nothing on the subject of either race or heredity. Finally in a slightly later work entitled Religion within the Bounds of Mere Reason (1793) Kant also uses the concept Vererbung (in the noun form) in a context in which it was already somewhat familiar. Much of the discourse on heredity in latter eighteenth century France was focussed primarily on the transmission of defects, especially hereditary disease. In this work on religion, in the course of an analysis of the origin of moral evil and its transmission, Kant discusses the explanations of this phenomenon offered by the three so-called “higher” faculties of the university (law, medicine, and theology). Each of these faculties, he tells us, has developed a notion of the inheritance of defects: the concepts of Erbschuld, Erbkrankheit, and Erbsunde (hereditary debt, hereditary disease, and original sin).8 But the question of biological mechanisms of heredity is 4

5

6 7 8

Haller (1752), vol. 2: “If nature were not the hand of creative wisdom, then there would be differences just as much in the basic constitution as in the small and numerous parts of the structure, but nonetheless the latter occurs constantly and the former never at all.” Kant ([1775] 1964), 6: pp. 9-30. This was Kant’s only publication during his so-called “silent decade” preceding the Critique of Pure Reason. It was revised and republished in Der Philosoph für die Welt (ed. by J.J. Engel) Leipzig 1777. Kant ([1785] 1964), 6: pp. 63-82. Originally published as “Bestimmung des Begriffs einer Menschenrasse.” (Berliner Monatsschrift, Nov. 1785). The term Vererbung also occurs in the Physical Geography published in 1801 based on Kant’s lectures. Kant ([1793] 1964), 4: p. 689.

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only obliquely raised in long and speculative footnotes on how various theories of generation might handle the inheritance of original sin and how they might explain the lack of transmission of this defect to Jesus. There also exist some elaborate manuscript drafts of these passages where Kant weighs the merits of epigenesis and the ovist and spermist forms of preexistence on this question; but I am not sure how seriously all this should be taken – it all seems to me to be somewhat facetious.9 What is important is that with Kant the notion of the hereditary moves from the higher or applied faculties to the lower, philosophical or theoretical faculty and that it is not confined to or even oriented towards defects. The German term for heredity (or inheritance), as I have mentioned, is Vererbung. Vererbung is slightly different in meaning than the English “inheritance” inasmuch as it takes the perspective of the donor not the recipient: it should actually be rendered as “bequeathment” – but this is seldom done. In the mid-eighteenth century the concept Vererbung was a somewhat complicated notion in German property law. An heir to property had a calendar year in which to accept or to reject the inheritance definitively. Problems could arise if the prospective heir were to die during this grace period. In such cases the tentatively inherited property was passed on to his or her heirs. It was specifically this procedure that was called Vererbung. Thus, use of the legal term Vererbung was restricted to situations in which one person received from another something that person had received by inheritance.10 There can, for instance, be no Vererbung erworbener Eigentümer, no inheritance of acquired properties because only property acquired by birth falls under the concept as defined. Now this is precisely the meaning that Kant wanted for the biological concept he was groping for in the race papers.11 Far more interesting than the religious speculations using the term is Kants use of the concept of heredity within the philosophical faculty – in biology –, where he uses it in an area with decidedly positive connotations, in connection with adaptation, and applies it to traits that are neither species typical nor merely individual peculiarities. Kant’s biological writings bring together heredity and adaptation. He develops his ideas on heredity and adaptation on the example of human races and all the context of anthropological discourse is relevant to an historical 9

10

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Kant (1964), 4: p. 736: “For, according to the hypothesis of epigenesis, the mother, who is descended from her parents through natural generation, would still be tainted with this moral blemish, and would bequeath [vererben] it to her child, at least half of it, even in case of a supernatural generation. Consequently, in order for this not to be the case, the system of preexistence of the germs in the parents must be assumed, but not the [system] of development on the female side as well (because through this the above consequence is not avoided) but only on the male side (not the [system] of ova but of spermatic animalcules); which side is omitted in case of a supernatural pregnancy, and thus this way of representing [generation] can be defended theoretically appropriate to that idea [i.e., virgin birth].” See also the drafts in AA 23, pp. 105-108. See Zedler’s Lexikon (1746): “Vererbung […] is when an heir, before he accepts or rejects the inheritance accrued to him, dies and that inheritance which he neither really accepted nor rejected is transmitted and further bequeathed […] to his heirs. […] And this occurs first of all when someone receives an inheritance from his parents or from elsewhere […] and dies within the year’s period after acquiring notice of the inheritance […] in such a case this kind of tranmission is called in Latin transmissio ex jure deliberandi.“ As we know, the ploy didn’t quite work. It may be a contradiction in terms to speak of the inheritance of acquired properties in an eighteenth century German courtroom, but in the outside world it is hard to impose the meaning constraints of technical terms out of their technical context. Even Kant himself relented: in his own last will and testament he speaks of “bequeathing his household goods” (von der Vererbung meines übrigen Hausgerätes) few of which, I presume, were acquired by inheritance.

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explanation of Kant’s theorizing here. But I shall be dealing only with one small aspect of his racial thought, an aspect that applies just as well to dogs and to pigs. Races of dogs for Buffon, races of pigs for Blumenbach and races of men for Kant were also occasions to discuss the question of subspecific variations that breed true. I shall stick to the more narrow biological context.

Hereditary Adaptations In the three papers on race Kant is primarily concerned with introducing and justifying a distinction between various kinds of subspecific classes: in particular between races and varieties. The theoretical background of the distinction is “Buffon’s rule” that animals that can produce fertile progeny together belong to the same species. The ostensible empirical phenomenon that Kant takes as the point of departure for his reflections (besides massive amounts af anecdotal travel literature) is the experience of the Portuguese colonists in Africa and of black Africans transported to Europe. Conclusions are drawn about other races as well, but the central question is about blacks and whites. Theoretical background – From Buffon’s breeding criterion and the ability of all humans to interbreed Kant draws the conclusion that all humans whatever their morphological differences should be viewed as descendants of one common stock and thus not only to belong to the same species but to constitute one family. Kant wants to distinguish between two kinds of disciplines: a purely descriptive and classificatory discipline – which he calls Naturbeschreibung – and a causal explanatory discipline – which he calls Naturgeschichte. (Linnaeus and Buffon provide the prototypes of these two disciplines.) The first gives us a school system for our memory, the second a physical system for our understanding. But Kant’s causal, physical natural history also has something contingently historical about it. The classificatory traits he is interested in are not explained only by natural law but also by the contingent family history of man. Empirical Phenomenon: skin color – The direct effect of the climate in Africa according to Kant is to darken the skin, but the Portuguese who had been living in tropical Africa for many generations were just as white at birth as their countrymen in Lisbon. And the blacks transported to Europe did not seem to be turning white – there was much literature from Paris about blacks and one of the Hessian Dukes had his own little imported colony of blacks established near Kassel. Furthermore, it is assumed that dark skin is beneficial in the African climate and white skin in the European, that is, that skin pigmentation is adaptive. The basic questions that arose were: Why don’t the Europeans in Africa turn black? Why don’t the Africans in Europe turn white (again)? And how did the blacks get black in the first place? The easy way out of course would be just to deny the phenomenon and assume that the timespan needed to adapt skin color is very very great. But Kant takes the occasion to ask a principled question: How can something purposive like a new adaptive trait arise, become heritable, and become heritable in such a way that it cannot be removed? Kants first step is to distinguish at least four kinds of subspecific taxa, which can be arranged on a sort of scale according to how invariably (unausbleiblich is Kant’s term) the identifying characters are transmitted:

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Strain (Schlag)

Variety (Varietät)

Sport (Spielart)

Race (Rasse)

Low

High Variable, dependent on Environment

Generally transmitted

Invariably transmitted in in-crosses

Invariably transmitted even in out-crosses

The most important difference is between races and the other categories: varieties, sports, and strains, in as much as this difference also correlates more or less with that between natural history and natural description. The possession of a character defines a race only if it is invariably bequeathed to the next generation even when the organisms are transplanted to a new environment and even when they are crossed with individuals from another race. In the latter case all progeny are halfbreed (Kant’s term is halbschlächtig); that is they, express a middle position between the two racial characters. In sports some traits, like blond hair and blue eyes, are passed on invariably and independent of change in environment, but they don’t produce halfbreed forms when crossed with other (say brown eyed) types. And in varieties the characters are generally but not always passed on in in-crosses; not all children of brown eyed pairs have brown eyes. In what Kant calls a strain some traits may be dependably passed on and even invariably appear when their possessors are crossed with similar individuals and always produce halfbreed traits when the bearers are crossed with other forms, but they may just disappear if the organisms are transferred to a different environment. Kant then tells a speculative story about how the original (and presumably white) humans spread out from their original habitat to all parts of the globe. On the way they adapted to the local climate. The problem is how to explain this adaptation in such a way that it is compatible with his other ideas about the explanation of organisms. Kant doubts that the laws of mechanics can explain the first origin of a purposive organic form – though they might be able to explain its replication – and thus he is committed to some kind of original organization in animals. If the environment could directly change organisms to their advantage in such a way that the changes could be passed on, preserving or even improving the purposivensss of their structure, there is no reason why it should not in time also be able to change the organism to any arbitrary extent (even beyond the species boundary) or, for that matter, why it could not have produced them in the first place. There would be no limit to the bizarre forms that creatures might take and it might become impossible to reconstruct the original species form. But Kant claims that it is impossible to imagine how environmental conditions could be able to cause beneficial changes that can be passed on. Thus, he insists that all beneficial and hereditary traits must have been included in the original purposive organization. All adaptations to any particular environment must have been part of the original species equipment; they were just not expressed – or, in Kant’s terminology, they had not yet been “unfolded”. Kant asserts that the lineage must have contained “germs” or “natural dispositions” for any adaptive trait that arises and then breeds true. The germs unfold into particular organs or traits; the natural dispositions are responsible for new relations among given parts. Once brought to expression they are permanent and thus other alternative possibilities are

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excluded for the future: the unfolding germs for black skin, for instance, close off the developmental possibilities for other colors. But only beneficial traits already contained potentially in the species form will breed true. The environmental, climatic circumstances cannot produce the new traits they can only trigger their release. Everything passed on invariably in generation is itself inherited. In somewhat anachronistic terms, every hereditary adaptation is a preadaptation. As Kant puts it in the first race paper: Accident or universal mechanical laws cannot produce such fitnesses [Zusammenpassungen]. Therefore we must consider such occasional unfoldings [Auswickelungen] as preformed. However even where nothing purposive is displayed, the mere capacity to propagate its particular assumed character is itself proof enough that a special germ or natural disposition was to be found in the organic creature. For external things can be occasional causes but not productive causes of what necessarily is propagated and breeds true [anerbet und nachartet] Just as little as accident or physical-mechanical causes can produce an organic body can they add something to its generative force, that is, effect something which propagates itself, if it is a particular shape or relation of the parts.12

The adaptive traits that Kant is interested in here do not occur in every current individual of the species nor did they occur at all in the original ancestral form: they are historically new, but now permanent. But they are also not mere individual peculiarities. These are not the adaptations of one Immanuel Kant to the cold winter in Königberg of 1774. It would have been fairly implausible to assume millions upon millions of little germs for this or that possibly useful adaptation. The skin is seen to be the major adaptive interface between organism and environment and it is skin color and texture that make up the primary racial character. As it turns out Kant assumes only four racially relevant sets of germs in humans. He maintains that the human species possessed germs and dispositions for traits preadapted to four basic climates. Where Linnaeus explains his four races by means of the distibution of four galenic humors, so that the dominance of sanguinous or phlegmatic aspects might even be taken causally to explain the racial differences between whites and blacks, Kant on the other hand goes back even farther to the presocratic doctrine of elements and the qualities derived from their combinations – cold and wet, hot and wet, hot and dry, cold and dry – and uses them as part of a teleological explanation of races. Europe is of course cold and wet; Africa hot and wet; and the other races are somehow assigned to the other two climates. (And just exactly what the two non-white, non-black races are changes over time). As humans radiated out of the Garden of Eden – or whatever it was that Kant thought was located between the 31st the 32nd parallels in the Old World – the germs and dispositions appropriate to the new climate they encountered were unfolded and their skins assume the appropriate color and structure. Kant also stipulates that this is an historically irreversible choice. The chosen germs shut off the others – that is why the blacks cannot become white again. The process of adaptation is irreversible even if the end product, due for instance to later transplantation, should turn out to be detrimental.

12

Kant ([1775] 1964), 6: p. 18.

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To recapitulate briefly: the relevant assertions are: 1. 2. 3. 4.

that all humans belong to one species, that racial differences are primarily due to adaptation to different climates, that any trait that is dependably passed on in this way is presumed to be adaptive, and that any trait that arises at some point in time and afterwards breeds true must have been contained as a potency in the original equipment of the species. In trying to deal with these questions Kant introduces a complicated and basically untranslatable jungle of special terms to distinguish various aspects, permutations and combinations of hereditary phenomena. (And thus it is not surprising that the term that later came to be used for heredity is among the crowd.) Here is a list of the major entries: erblich – hereditary (traits are hereditary) erben – to inherit (an organism inherits a trait) ererben – to inherit (an organism inherits a trait) anerben (intrans) – to be propagated (a trait is propagated) sich vererben – to pass on (a trait passes itself on) vererben – to pass on (an organism passes on a trait) forterben – to continue (a trait is continued) Abartung / abarten– to degenerate (deviate from the original form) Ausartung / ausarten – to deviate beyond the possibility of reversal (literally to speciate, but used here for change within the species establishing a race)13 Nachartung / nacharten (trans.)– to take after (F2 trait takes after an F1 trait) Nachartung / nacharten (intrans.)– to continue Anartung / anarten (intrans) – to be propagated (said of a trait) Anartung / anarten (with dat. object or with an + dat./acc) – to adapt to (said of an orgnism) eingeartet – acclimatized (adapted to an environment) What is important in this terminological morass is that Kant is systematically groping for conceptual tools to deal with phenomena that we conceptualize using the notions of heredity and adaptation. For our purposes the following results of Kant’s speculations seem to be historically relevant: 1. The characters focussed on are not species characters determined by divine preformation or physical natural law. The germs and dispositions still fit the deterministic eighteenth century model of pre-given potentialities, but at least the fact of their expression as traits is a product of contingent history. 2. Some of the contingent traits that an organism possesses solely because its parents possessed them are not just accidental modifications but rather choices among species-given possibilities and thus are perhaps susceptible to rules and may be explainable by the same kinds of mechanisms as species-wide traits. 3. The realm of the hereditary is not confined to defects or mere peculiarities but also includes beneficial traits – and, as pursued in the “philosophical” faculty, it includes only beneficial traits.

13

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Note that this does not refer to the sterility of hybrids but only to the fixation of the racial character. Kant at one point calls real speciation “wahre Ausartung”; see Kant, ([1775] 1964), 6: p. 19.

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4. Hereditary characters of the kind discussed must themselves be inherited. Genuinely acquired characters are not hereditary. It is true that for Kant historical contingency remains within the species lineage, but Kantian races have all the characteristics of traditional species except the sterility of hybrids. They provide examples of lawlike purposiveness conceptualized as improvements on a given species form relative to contingent actualizations of basic climatic possibilities. The racial characters are neither completely determined by species membership nor are they merely arbitrary or accidental. They present an opportunity or occasion for the application of available concepts to situations for which they were not originally devised or intended and thus may force some change or development in these concepts.

References Gassendi, Pierre. [1658] 1966. Syntagma, Opera. In Marcello Malpighi and the Evolution of Embryology. Excerpted and translated by Howard B. Adelmann. Ithaca, NY: Cornell University Press. Highmore, Nathaniel. 1651. The History of Generation. London. Bourguet, Louis. 1729. Lettres philosophiques sur la formation des sels et des crystaux et sur la génération & le mechanisme organique des plantes et des animaux. Amsterdam: L’Honoré. Buffon, Georges Luis Leclerc de. 1765. Histoire naturelle. Volume 13. Paris. _____. 1790-1799. Natural History, General and Particular. Translated by William Smellie. ) vols. London David Hume. 1779. Dialogues concerning Natural Religion . Edinburgh. Haller, Albrecht von. 1752. Preface to Allgemeine Historie der Natur, by G.L.L.Buffon. Volume 2. Hamburg. Kant, Immanuel. [1775 and 1777] 1964. “Von den verschiedenen Rassen der Menschen.” Revised and republished by J.J. Engel. In Der Philosoph für die Welt. Leipzig. In Werke in sechs Bänden. Volume 6. Edited by. W. Weischedel. Darmstadt: Wissenschaftliche Buchgesellschaft. 9-30. _____. [1785] 1964. “Bestimmung des Begriffs einer Menschenrasse.” Berlinische Monatsschrift (Nov. 1785). In Werke in sechs Bänden. Volume 6. Edited by W. Weischedel. Darmstadt: Wissenschaftliche Buchgesellschaft. 63-82. _____. [1793] 1964. “Die Religion innerhalb der Grenzen der bloßen Vernunft.” In Werke in sechs Bänden. Volume 4. Edited by W. Weischedel. Darmstadt: Wissenschaftliche Buchgesellschaft. _____. [1798] 1964. "Über den Gebrauch teleologischer Prinzipien in der Philosophie." In Werke in sechs Bänden. Volume 4. Edited by W. Weischedel. Darmstadt: Wissenschaftliche Buchgesellschaft. 139-170.

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Inheritance of Acquired Characters in Lamarck’s and Geoffroy Saint-Hilaire’s Zoology Wolfgang Lefèvre

INTRODUCTION The question of heredity became only late a scientific topic that was systematically pursued by sustained research – probably not before the turn of the nineteenth century, that is, not before the discovery of Gregor Mendel’s (1822-84) discoveries. Up to this point of time, heredity was only occasionally a subject matter of scientific considerations. Moreover, it was almost never such a subject in its own right but only as an aspect of other issues of interest such as breeding, race, actually or allegedly hereditary diseases, investigations into hybridisation or generation and development.1 Thus, the issue of heredity often was not more than a facet of conceptions about races, diseases or generation, and, accordingly, inseparable from those concepts. Our recognition of concepts of heredity in eighteenth and nineteenth century theories, thus, may be slightly anachronistic since marked-out concepts of heredity came into being only later. Because of this danger of an anachronistic approach, it seems well-advised to state first what is understood by heredity in this paper. In the following, I take such concepts to be concepts of heredity that assume that traits of living beings are dependent on traits of their parents or even farther ancestors. Such conceptions need not comprise explanations of these dependencies. On the contrary, I just would like to distinguish heredity proper from all past and present physiological theories of generation, genetics, and development that provide an account of the mechanism by which the transmission of traits is brought about. True, this separation of heredity proper from explanatory theories and, consequently, its confinement to patterns of trait occurrences among living beings of subsequent generations is conceivable only after Mendel. It is no actors’ category with respect to life scientists of the eighteenth and nineteenth century. Rather, it is an analytical category of the historian that allows both, the comparison of otherwise incomparable theories, which imply assumptions about heredity, and their distinction from ideas that only seemingly do so. The topic of this paper is heredity in the zoological theories of Jean Baptiste Lamarck (17441829) and Etienne Geoffroy Saint-Hilaire (1772-1844). These theories, remarkable in so many respects, deserve attention not at least as the first serious attempts to explicate and explain adaptation to changing environmental conditions. Today, no such theory of adaptation is conceivable without heredity concepts in the tradition of population genetics. The more remarkable is the casual manner in which Lamarck and Geoffroy touched the question of heredity. It is for this casualness, that I regard their theories a noteworthy example of the manner in which heredity was treated in scientific discourses before the end of the nineteenth century. With respect to a history of heredity, these theories deserve attention not so much for what they state about the

1

Bowler (1989), p. 23.

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inheritance of acquired characters than as symptoms of the inferior significance of heredity concepts in biological theories of this age. In my view, this inferior, casual, and secondary status in and for biological theories constitutes a characteristic feature of heredity concepts before the discovery of Mendel that deserves to be explicitly addressed by a workshop dedicated to this period. For this reason, I will not immediately start with Lamarck and Geoffroy. Rather, I will first touch briefly upon some of the other contexts mentioned above – breeding, the problem of race, theories of generation and development – in which the question of heredity became a topic of scientific inquiry although only as a secondary aspect of the main conceptions in question. The purpose of these preliminary excurses is the indication of a broader frame for my argument regarding Lamarck and Geoffroy.

1. OCCASIONS FOR REFLECTIONS ON HEREDITY Since time immemorial, man, himself an animal who propagates his genus through sexual reproduction, experienced phenomena of heredity. However, the experiences made with the own biological procreation did not yield a clear pattern. This is not at all surprising in view of the complexity of the generation process. True, the offspring very often resembles its parents, but equally often it does not. There is something like family resemblance and even tribe resemblance, but there are also a lot of individuals who do not fit this pattern. Moreover, regarding the plants and animals men are familiar with, it seems safe to claim as a rule that the progeny will always belong to the species of the parents. But what about malformations, and what about hybrids? Do they not violate this rule? Obviously, every-day experiences are not sufficient for singling out stable patterns of heredity, and, one should add, every-day practice is probably not in need of a more distinct picture of this realm of phenomena.

1.1 Breeding One would expect that the professional practice of breeders2 must have brought about a base of experiences broad and stable enough for correcting wrong assumptions of the laymen as well as for establishing certain rules of heredity – and perhaps even a stock of experiences on which scientific investigations into heredity could start and rest. In a way, this was indeed the case. Mendel’s research as well as that of his re-discoverers around 1900, that is, the very beginnings of the modern biological science of heredity, was imbedded in the context of the breeding practice of this age. Although this relation between professional breeding and the beginnings of modern genetics seems quite natural, it deserves doubtless attention for many reasons and provokes many questions.3 With respect to the topic of my talk, it is of special interest to ask whether heredity did become a scientific topic in its own right just because scientists became curious of heredity patterns when the practice of professional breeders provided essential preconditions for a promising research. Was this scientific interest not also, and perhaps even primarily, structured and driven by open questions in other fields of biological research – for instance, by the question whether or not Charles Darwin’s (1809-82) assumption of minimal variations is viable as part of his account of 2

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In this paper, the term breeding is used for both, breeding of animal stocks and growing of plants.

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the process of speciation?4 To put it more general: were certain theoretical constellations in late nineteenth century biology responsible for the commencement of not only periodical but continuous and sustained research activities on heredity? Or, to put the question more speculative: Would the practice of nineteenth century breeders have caused scientists to study heredity as a subject matter in its own right if these scientists had not had their own theoretical reasons for an exploration of this question?

1.2 Human Races Whatever the answer to these questions may be, the professional practice of breeding was in any case more than just an occasion for addressing scientifically some related issues of heredity. Heredity constituted the very centre of this practice, which provided additional means and ways for its investigation. This does not hold for the debates on races in the context of anthropology. These debates were certainly only an occasion for contemplations on patterns of heredity; and perhaps, one can add, an unfit one. Modern anthropology originated in the colonial conquest and domination of large parts of Africa, America, and Asia by European nations and was therefore coloured by ideological, political, economic, and moral discourses from its beginnings. Even the question whether and in which manner issues of heredity became prominent in this context 3

Why did the experiences of professional breeders become subject of scientific consideration only that late? Where those experiences not ripe for scientific scrutiny before the second half of the nineteenth century? If so, what is so special with the breeding practice of the nineteenth century in contrast with the more than ten thousand years of breeding that preceded it? Or, to face an alternative possibility, were perhaps the life sciences not ripe for, or not interested in, the heredity topic before the second half of the nineteenth century? If so, why exactly did breeding attract the attention of scientists in the end, and, more important, what did enable them to tackle the subject successfully? Recently, in a “heredity reading group” here at the Max Planck Institute for the History of Science, we studied a series of articles about sheep breeding written, in the 1830s, by a certain J.K. Nestler, an Austrian professor for agriculture and farming, and published in the journal of the Moravian-Silesian society for the advancement of agriculture. (Mitteilungen der k. k. Mährisch-Schlesischen Gesellschaft zur Beförderung des Ackerbaues, der Natur- und Landeskunde in Brno. Issues 34ff.) Apart from all their details and right or doubtful observations, what these articles conveyed impressively was a picture of the high complexity of the breeding business. Obviously, breeding is more than just organizing the mating of individuals with desired properties. Thousands of other things were regarded of equal significance by the professionals – the right season for mating, the age of the two mates, whether the male should be younger, older or of the same age as the female, the question of the best food, influence of the environment when a stock was moved into a new locality, the problems of selection in view of the principal fact that the breeding goal almost never consists in the preservation, transmission and enhancement of just one single trait but in that of a complex of traits which could only be approached by way of compromises, etc., etc. Leafing through these articles, it became immediately clear that stable patterns of heredity could not be established without a radical reduction of this complexity, and that detachment from the immediate practical goals of the breeding business was an essential precondition of such a reduction. This leads to further questions. How exactly did breeding practice and science interact and contribute to the emergence of modern genetics? Certainly, what regards the distance to immediate practical goals needed for the reduction of complexity, breeding experiments like those of Mendel show detachment from practical goals of growers. The colours of pea blossoms were without any practical value. But does this mean that we must credit the scientific side alone with the reduction of complexity necessary for the establishment of heredity patterns? Did the breeding practice contribute nothing to it? What about the keeping of pedigrees by the breeders? Is a scientific examination and contemplation of breeders’ experiences conceivable without such pedigrees? And what about the scientific breeding experiments? Were they not based on techniques of professional breeders? For an expert’s view of these issues, see the paper of Roger J. Wood in this volume.

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depended to a large extent on ideological orientations. For those who regarded the various aboriginals of the colonised countries different species rather than varieties of the human species, heredity questions came up only because of the so-called half-bloods which raised the special question of hybridisation if the contended species difference was taken consequently. For those, on the other hand, which believed in the one human species and, additionally, that the variety of human races was brought about by a historical process of divergent developments of the descendents of a single primal tribe, this variety could, indeed, provoke speculations on issues of heredity. The anthropological context appears thus as an ambiguous cause and rather sterile soil for raising and pursuing scientifically questions of heredity. It is probably not by chance that no continuous exploration of these questions flourished on this background, and also not that the trend of biological anthropology drifted in the direction of a science of measurement that engendered eventually the “mismeasured man” – to allude to a famous book of the late Stephen J. Gould. I could, therefore, leave anthropology as an occasion for studies on heredity at this point. But I want to mention additionally at least one example of an anthropologically informed speculation on heredity. My example is the speculations on heredity comprised in Immanuel Kant’s (1724-1804) anthropology. Since Peter McLaughlin’s paper explores these speculations, I can confine myself to one single aspect, which is of interest for my story on heredity in Lamarck’s and Geoffroy’s zoology. Kant considered the human races as varieties of a single human species. 5 For him, these races descended from one single primal tribe and developed their distinctive features under the impact of the different climates of the continents in which they finally, at the end of a long period of migration, dwelled. But – and this is the interesting point – Kant did not assume that men acquired the genetic dispositions – natürliche Anlagen (natural dispositions) in his words6 – for these distinctive features under the influence of those climates. Rather, he supposed that all men possessed originally the dispositions for all of these different features. The role of the climate consisted in stirring the actual development of the best fitting features. With other words, Kant put forward a speculation on adaptation to changing environmental conditions that did not include a change in the stock of hereditary dispositions. He assumed, however, that the actual development of features under given climatic conditions leads to an irreversible fixation of this developmental choice, which is passed on to the progeny. – I will briefly come back to this sensible speculation when discussing certain assertions of Geoffroy as well as Lamarck.

1.3 Generation and Development The breeding practice and the debates on human races stimulated from outside the life sciences to reflections on questions of heredity. However, in eighteenth and nineteenth century life sciences, issues of heredity were also raised from within, namely in different, although closely connected, fields of theoretical and experimental investigations. There was a broad spectrum of theoretical 4 5 6

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For the spectrum of interests supposed by historians as having driven Mendel, see Sapp (1990). For Kant’s notion of race and its distinction from that of strain (Schlag), variety (Varietät), and sport (Spielart), see the paper of Peter McLaughlin. Kant ([1775] 1912), p. 434.

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issues, which touched more or less explicitly questions of heredity. Just to remind of three famous debates in the seventeenth and eighteenth century life sciences: • The debate about symmetric or asymmetric roles of the two sexes in generation, that is, the question of whether the germ is provided by both sexes or only by one of the two – either by the male whereas the egg serves only as a soil for its development or, alternatively, by the female whereas the semen plays only the role of a stirring principle; • The debate whether or not plants engender their progeny sexually; • The debate on development, that is, whether it is a process of growing, in the sense of blowing-up, of an already finally structured miniature organism (doctrine of preformation) or rather a modelling process by which an initially almost unstructured germ develops step by step the complex structure of the organism (doctrine of epigenesis). It goes without saying, that each of these debated views had crucial consequences for the understanding of heredity regardless of whether or not their champions dealt explicitly with heredity issues. But it is also clear that basic preconditions of the very possibility of heredity were at stake in these debates, with the consequence that details like differential reproduction were almost eclipsed. It comes therefore without surprise that explicit discussions on heredity came only rarely down to details and did almost never go beyond the limits of ad-hoc arguments. The debate between preformationists and epigenesists deserves particular attention in the context of this talk because both, the preformist and the epigenetic view, 7 when pushed to their extremes, could lead to the denial of the inheritability of individual traits or even of the very possibility of inheritance as such. According to the ontogenesis conception of the epigenesist Caspar Friedrich Wolff (1738-94), for instance, almost unstructured organic matter provided by the parental organism develops step by step into the organism’s final structure by virtue of an innate vis essentialis (essential force). What the parents bequeath to their progeny is the initial organic matter with its essential force, which brings about the species-specific organism under suitable environmental conditions. The likeness between parents and offspring rests on the uniformity of this initial matter’s development under comparable environmental conditions. Being equally the effect of equal causes rather than cause and effect of each other, the traits of individuals of subsequent generations are not linked by a relation of dependency. 8 On the side of the preformationists, the conception of hereditary traits was entirely erased if the preformist view of the germ was combined with the assumption of its pre-existence, that is, with the assumption that the germs of the entire progeny of a species were created in the beginning along with its primal couple, encased in each other like Russian dolls. In the framework of this understanding of generation and development, no place was reserved for the issue of heredity, not even for the inheritance of the species-specific characters. This issue simply did not exist. Apart from being designed by the same divine creator, there were no relations between the characters of subsequent generations. With other words, we come here across a biological debate that implied the possibility to question heredity as a meaningful concept and, thus, investigations of patterns of heredity as a reasonable enterprise.9 Ironically, the preformist view of generation and development stimulated interest in heredity 7 8 9

Peter McLaughlin’s paper for the 1st workshop on A Cultural History of Heredity discussed the bearing of the two views for the understanding of heredity. Bowler (1989), p. 40f.; Stubbe (1965), pp. 73ff. This seems noteworthy because, in my view, theories that negate explicitly or implicitly heredity deserve no less attention in a history of heredity concepts than theories that shape the meaning of such concepts.

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issues just because of its negation of heredity – namely on the side of epigenesists who combined their view of the developmental process with pangenetic accounts of the formation of the germ. Actually or allegedly obvious facts of heredity were picked up as arguments against the extreme preformist view.10 In particular, hybrids and descendents of parents of different races could be adduced as striking examples of mixed traits that seemed incompatible with the assumption of pre-existent preformed germs.11 However, this critique resulted only in ad-hoc theories on heredity and not in detailed investigations. It is a telling fact in this respect that, in the course of the argument between adherents and opponents of the preformist doctrine, experiments for confirming or refuting certain claims about heredity patterns were proposed but obviously never carried out.12 I will come back to these doctrines in connection with Lamarck.

2. HEREDITY IN LAMARCK’S AND GEOFFROY SAINT-HILAIRE’S ZOOLOGY The arguments put forward so far pursued the purpose to mark a distinctive feature of scientific occupations with heredity issues before the end of the nineteenth century. This feature could be described negatively by stressing the lack of continuity of these studies, their ad-hoc character, or by pointing to the fact that heredity became no scientific subject in its own right. When heredity attracted the attention of life scientists in this age, it did so always as a concomitant of inquiries on issues of practical (e.g., breeding and allegedly hereditary diseases) or social-political concern (e.g., the question of human races) or of theoretical problems with implications for heredity (e.g., generation and development). These inquiries and their different backgrounds shaped of course the understanding of heredity as well as the selection of the aspects that were subjected to scrutiny. Being not a subject of scientific investigations in its own right, it depended entirely on certain constellations of practical, ideological, and theoretical views and interests whether or not heredity was dealt with at all, and, if dealt with, which of its aspects were investigated and by which means. This feature is essential for my story. Lamarck’s and Geoffroy’s theories of adaptation to changing environmental conditions formed such a constellation that opened a window for the heredity question. These theories integrated results from different fields of research – biogeography, geology, palaeontology, teratology, and physiology – and constituted thus themselves constellations of theories. However, this case confirms not only once more that the understanding and treatment of heredity depended on such a constellation. It proves furthermore that the question of heredity could even be part and parcel of fiercely debated theoretical positions without getting investigated in more detail by any of the opposed parties.

10

11 12

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On their part, preformationists tried to avoid the extreme consequences of their view and to allow for the inheritance of individual traits by assuming a transmission of characters through the nutrition of the germ – by the seminal fluid of the father as well as the mother’s womb. Charles Bonnet (1720-93), for instance, developed ideas that resulted even in a combination of preformism and pangenesis. (Bowler, p. 30f.; Zirkle, p. 143) See also Peter McLaughlin’s paper for the 1st workshop on A Cultural History of Heredity. Jacob (1971), p. 80f. Ibid., p. 91f.

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2.1 Adaptedness and Adaptation The adaptation of animals and plants to their habitats was of course no new discovery of Lamarck’s and Geoffroy’s days, although the awareness of such adaptations had certainly been sharpened by the increase of knowledge about “exotic” faunae and florae through European colonialism. The eighteenth century was almost obsessed with wondering about these indeed striking phenomena. The reasoning of this century’s physico-theology dwelled on them excessively as an inexhaustible source of proofs of the divine creator’s wisdom and providence. With hindsight, it is obvious that these admirers of the creation dealt only with adaptedness and not with adaptation. They supposed that animals and plants fit so admirably well their environments because the divine creator designed them suitably. But if plants and animals were originally created as well adapted to their habitats, no process of adaptation needed to be assumed to account for their adaptedness. Living beings appear, thus, as being adapted to their habitats but not as adapting themselves to them. Adaptation as a natural process was not yet a needed concept. It became such a concept when naturalists were confronted with facts that indicated strongly to the possibility and even high probability that some, many or perhaps even all species of plants and animals had not always lived in their present habitats. With other words, with the emergence of doubts in stable relations between organic forms and habitats, the concept of adaptation to a new environment advanced as a possible natural account of adaptedness under changing environmental conditions. The facts that induced doubts in unchanged environmental conditions were put forward firstly by the results of biogeographical investigations. The findings in this field of naturalistic explorations, which experienced a first flourish just in the second half of the eighteenth century, indicated strongly to the probability that closely related animals and plants from different regions of the world might be derivatives of one and same parental form which adapted themselves to different climates when spreading by migration. In Kant’s anthropology, we encountered already such considerations about migration and adaptation, which also played a prominent role in the theories of the two most eminent figures of eighteenth century biology, namely Carl Linnaeus (1707-78) and George-Louis Leclerc, Comte de Buffon (1707-88). Buffon, who has been celebrated as the founding father of modern biogeography, combined additionally the biogeographical view with that of a historic geology. In the middle of the eighteenth century, the question whether or not the Earth’s surface underwent historical change was definitively given a positive answer by expert geologists, how much ever they dissented with respect to the extent and to the supposed causes of this secular change. Buffon went even so far as to put forward the bold particular hypothesis that Africa and South America formed formerly a united continent. Interestingly, this speculation drew less from geological evidences than from zoological ones, which indicated that different adaptations to unlike environmental conditions might account best for the differences between closely related species or genera of the two continents.13 The dramatic consequences of a historical view of the Earth for the biological understanding of adaptation unfolded finally just in the days of Lamarck and Geoffroy when modern palaeontology celebrated its first triumphs. The findings of George Cuvier (1769-1832) and William Smith (1769-1839) showed unmistakably that the different strata of the Earth contained

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entirely different faunae and florae. This was really revolutionary and bewildering news. All naturalists, regardless to which extent they accepted these findings, agreed immediately upon one point, namely that these surprising facts must be subsumed under the adaptation concept: the strata contained different faunae and florae because they represented different life conditions in the Earth’s past. The dissent came with the next question: must these different worlds of living beings be considered different creations which were completely wiped out by sudden changes of the Earth’s surface or can they be taken to be different developmental stages of the same creation which, notwithstanding the extinction of a large amount of species, continued to exist although radically altered by its adaptations to drastic changes? It goes without saying, with this alternative, really big questions were at stake and not just a detail of a theory – questions with far reaching consequences for religious and philosophical convictions and orientations which touched social and political interests. And for the life sciences, a cornerstone of their theoretical edifice was touched and endangered by this alternative, namely the concept of the constancy of species. It is therefore not surprising that the majority of naturalists preferred the first view and regarded the different faunae and florae as adapted to the conditions of their respective stratum but not as a result of a natural process of adaptation. The awkward consequence of a series of life erasing catastrophes and independent new creations seemed to them obviously less horrifying than the assumption that the species of living beings can transform themselves like a Proteus.

2.2 Lamarck and Geoffroy on Adaptation However, as almost always in real life, tertium datur, that is, a middle position could be taken in this situation. Naturalists could defer judgment on the big questions raised with the palaeontologic discoveries – all the more as almost every conclusion from these discoveries could reasonably be questioned at that time – and confine themselves to the assumption that adaptation to changing environmental conditions engenders alterations of varieties, species, or even genera within certain limits. Even the radical Lamarck followed in a way this line. His theory of species transformation was not an attempt at keeping up with the history of life on this planet as suggested by palaeontology. He elaborated it independently of the palaeontologic discoveries of his days. These discoveries constituted a disturbance rather than a support for his theory of the historical development of organic forms, and he tended, therefore, to play down their significance and implications, and denied, for instance, the extinction of species apart from a few single cases. Lamarck conceived of the transformation of species as a process that is primarily determined by inner-organismic laws and only secondarily and additionally by the environment. According to his theory, it is the interaction between the fluid and the solid parts of an organism what effects not only the ontogenetic development but also the gradual advancement of the adult structure of a species. His assumption that the species of the animal Kingdom, and the same holds for the 13

See Roger (1997), pp. 331ff. – Speculations about adaptations of animals and plants to supposed changes of the Earth’s surface had become wide-spread since the appearance of Benoit de Maillet’s (1656-1738) Telliamed in 1748. In these speculations, adaptation was treated as a matter of fact that needs no explanation. An explication of the notion of adaptation to changing environmental conditions was seriously attempted first by Lamrack.

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vegetal Kingdom, constitute different successive developmental stages of one and the same primal species14 did not draw on any external factors. It was derived from supposed inner-organismic processes, and only this internal determination warranted the regularity and lawfulness of the species transformation. In the framework of this theory, external factors played the role of interferences from outside that disturb the natural way of development and alter its results. The unavoidability of such interferences rested, for Lamarck, on the spreading of the species throughout the different climates of the world rather than on historic alterations of the surface of the Earth.15 And it is exactly at this point where the concept of adaptation surfaces in this picture. Adaptation to changing external conditions has the function to account for the deviations from the natural line of species transformation. As is well known, Lamarck’s and Geoffroy’s concepts of adaptation differed considerably, but they did so on the basis of a shared presupposition that sounds strange today. Both men assumed that the alterations of an organism that effect its adaptation to new environmental conditions are induced, either directly or mediated, just by these new conditions. In order to mark the peculiarity of this presupposition, it may be convenient to remind briefly of the understanding of adaptation after Darwin’s theory of evolution. According to this theory, a species’ adaptation to changing conditions is effected by the natural selection of its individual variants that fit best the new conditions. However, this theory does in no way presuppose that fit individual variants are brought about by the changed environment, all the more not since the environment is viewed as being made up in the first place by other species and not by inorganic factors. Whether or not suitable and thus selectable variants come into being is an entirely open question in the framework of the theory of evolution, and thus the survival of the species under the new circumstances. In contrast to this, the alterations with an adaptive value are induced just by the new environmental conditions in Lamarck’s and Geoffroy’s conceptions 16 – although in different ways. Geoffroy supposed a direct impact of changed external conditions upon the organism that causes its modification.17 A background of this assumption was Geoffroy’s studies on “monsters,” as malformations were called then.18 He pursued these teratologic studies not only as a comparative anatomist but combined morphologic investigations with embryologic experiments on how physical parameters like chemical composition of the air, temperature, etc. alter the development of a germ.19 The results led him to the view that organisms are surprisingly flexible beings that are able to react, within certain limits, to changed physical conditions by developing modified organic structures in ontogenesis. And he considered this flexibility the basis of adaptations to changed environmental conditions.20

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15 16 17 18 19 20

More precisely, Lamarck assumed that all higher classes of the animal Kingdom arose from one of the four classes of the most primitive animal organisms, namely from the class of worms, whereas the three other primitive classes (infusorians, polyps, and radiarians) seem unrelated to the line of development that connects the worms with the mammals, and, thus, with man. See Lamarck (1809), 2: p. 457. Lamarck (1809), 1: pp. 266f. And just because of this, Lamarck needed not to be worried about the erasing effects of interbreeding – see Burckhard (1995), p. 181. Russell (1916), p. 68; Appel (1987), pp. 132ff. Ibid., pp. 75ff. Ibid., pp. 128ff. Cahn (1962), pp. 237ff.

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Lamarck, on the other hand, denied such an immediate effect of changed conditions on the structure of organisms. He elaborated instead his famous theory according to which such conditions induce living beings to change their behaviour. Change of behaviour entails a different use of organs. New employments or refrain from former employments of organs lead to physical modifications of both the organs that are suitable for fulfilling the new tasks and the organs that come gradually out of use. Thus, the structure of an organism is modified by its active response to a new environment.21 One has to admit that this concept of adaptation has the advantage to explain why just those alterations of the organism are induced by a new environment that are needed for the organism’s coping with it.

2.3 Adaptation and Heredity It goes without saying that Lamarck’s theory of adaptation presupposed the inheritance of the modifications with an adaptive value obtained by one generation. The finally adapted shape of the organism’s structure could only be achieved gradually through the accumulation of adaptive improvements of many generations. The famous or notorious inheritance of acquired characters was thus a necessity in Lamarck’s theoretical framework. It is not equally clear why Geoffroy followed the example of Lamarck what regards heredity. His theory does not necessarily imply an accumulation of alterations. Its distinctive feature consisted just in a supposed flexibility of organisms that enabled them to develop different structures under different conditions. This theory seems, thus, not in need of the assumption that advantageous characters acquired by one generation must be passed on to the progeny. Each generation seems able to develop itself the most advantageous structure. Being myself no expert what regards Geoffroy’s zoology and having not been able to find information about this point in the secondary literature, I have to leave open this question.22 Anyway, Geoffroy followed Lamarck what regards the inheritance of acquired characters and referred explicitly to the two laws that the latter put forward in his Philosophie zoologique.23 These laws explicate with desirable clarity how Lamarck viewed the connection between adaptation to changed environmental conditions and heredity and are, thus, worth to be quoted: First Law: In every animal which has not passed the limit of its development, a more frequent and continuous use of any organ gradually strengthens, develops and enlarges that organ, and gives it a power proportional to the length of time it has been so used; while the permanent disuse of any organ imperceptibly weakens and deteriorates it, and progressively diminishes its functional capacity, until it finally disappears. 21 22

23

Lamarck (1809), vol. 1, chap. 7. However, I would like to indicate the conjecture that Geoffroy assumed the inheritance of altered structures among other things because of experiences gained by his embryologic experiments, namely that the flexible responses of organisms to altered conditions lack reversibility. This conjecture is suggested by an assumption of Kant mentioned in the first part of this paper, namely that man was initially endowed with the genetic dispositions for the features of all human races but could not go back to this initial universal state once specific features had become actualised under a certain climate. In both cases, an initially universal genetic disposition, which is open to different developments under different physical conditions, would loose its full plasticity after the realisation of one of the developmental possibilities. Subsequently, a limited plasticity is passed on to the progeny. Geoffroy (1826).

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Second Law: All the acquisitions or losses wrought by nature on individuals, through the influence of the environment in which their race has long been placed, and hence through the influence of the predominant use or permanent disuse of any organ; all these are preserved by reproduction to the new individuals which arise, provided that the acquired modifications are common to both sexes, or at least to the individuals which produce the young. 24

The qualifications at the end of the second law are explicated in a short passage that reads as follows: […] in reproductive unions, the crossing of individuals who have different qualities or structures is necessarily opposed to the permanent propagation of these qualities and structures. Hence it is that in man, who is exposed to so great a diversity of environment, the accidental qualities or defects which he acquires are not preserved and propagated by reproduction. If, when certain peculiarities of shape or certain defects have been acquired, two individuals who are both affected were always to unite together, they would hand on the same peculiarities; and if successive generations were limited to such unions, a special and distinct race would then be formed. But perpetual crossings between individuals, who have not the same peculiarities of shape, cause the disappearance of all peculiarities acquired by special action of the environment. Hence, we may be sure that if men were not kept apart by the distances of their habitations, the crossing in reproduction would soon bring about the disappearance of the general characteristics distinguishing different nations.25

To my knowledge, this short passage is the most extended and elaborated one on heredity one can find in the Philosophie zoologique.26 Out of the fifteen pages in which Lamarck expounds the two laws, this quarter page contains all he has to say about the inheritance of acquired characters. Almost all of his explanatory efforts are dedicated to the elucidation of his conception of the acquirement of new characters in response to new challenges of the environment. He sensed

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Lamarck (1963), p. 113; Lamarck (1809), 1: p. 235: “Première Loi. Dans tout animal qui n’a point dépassé le terme de ses développemens, l’emploi plus fréquent et soutenu d’un organe quelconque, fortifie peu à peu cet organe, le développe, l’agrandit, et lui donne une puissance proportionnée à la durée de cet emploi ; tandis que le défaut constant d’usage de tel organe, l’affoiblit insensiblement, le détériore, diminue progressivement ses facultés, et finit par le faire disparoître. Deuxième Loi. Tout ce que la nature a fait acquérir ou perdre aux individus par l’influence des circonstances où leur race se trouve depuis long-temps exposée, et, par conséquent, par l’influence de l’emploi prédominant de tel organe, ou par celle d’un défaut constant d’usage de telle partie ; elle le conserve par la génération aux nouveaux individus qui en proviennent, pourvu que les changemens acquis soient communs aux deux sexes, ou à ceux qui ont produit ces nouveaux individus.” Lamarck (1963), p. 124; Lamarck (1809), 1: pp. 261f.: “Au reste, dans les réunions reproductives, les mélanges entre des individus qui ont des qualités ou des formes différentes, s’opposent nécessairement à la propagation constante de ces qualités et de ces formes. Voilà ce qui empêche que dans l’ homme, qui est soumis à tant de circonstances diverses qui influent sur lui, les qualités ou les défectuosités accidentelles qu’il a été dans le cas d’acquérir se conservent et se propagent par la génération. Si, lorsque des particularités de forme ou des défectuosités quelconques se trouvent acquises, deux individus, dans ce cas, s’unissoient toujours ensemble, ils reproduiroient les mêmes particularités, et des générations successives se bornant dans de pareilles unions, une race particulière et distincte en seroit alors formée. Mais des mélanges perpétuels entre des individus qui n’ont pas les mêmes particularités de forme, font disparoître toutes les particularités acquises par des circonstances particulières. De là on peut assurer que si des distances d’habitation ne séparoient pas les hommes, les mélanges pour la génération feroient disparoître les caractères généraux qui distinguent les différentes nations.” See also the short passage in Lamarck (1802), p. 61 and the thought experiment ibid., pp. 53f.

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obviously no urgency to explicate equally elaborately the question of heredity. 27 The fact that he confined himself to the adduction of some speculations about human races is really a telling one. Thus, one gets unavoidably the impression that Lamarck regarded the assumed inheritance of acquired characters the least risky part of his transformation theory. And this impression is strengthened by the literature on Lamarck, a large part of which takes this assumption as a commonly shared one at that time,28 invoking, as a rule, the example of Erasmus Darwin (17311802).29 I do not know whether contemporary laymen believed generally in such an inheritance, but would not be surprised if they did so notwithstanding the innumerable counter-instances, which could not hinder this belief since times immemorial. There are, however, good reasons not to presuppose a consensus on such questions in the community of life scientists. 30 Already the quoted passage on interbreeding of human races should caution us against rash judgements. Its assertions are put forward as if everybody was familiar and agreeing with them. But we noticed already that a well-informed writer like Kant adduced opposite assumptions what regards the possibility to wipe out certain distinct racial features by crossing. Obviously, questions like this were far from being answered definitively. And the fact that such contrary assumptions could be put forward like evident facts seems only to indicate that such assumptions were still not in the focus of biological conceptualisations. What regards the assumption of the inheritance of acquired characters, however, there was an immediate connection to heatedly debated theories that occupied the attention of life scientists since almost half a century. For this assumption was only theoretically intelligible in the framework of an epigenetic conception of development that included a pangenetic understanding of the germ’s generation – a conceptual framework that was not at all generally accepted among life scientists around 1800. Both, Geoffroy and Lamarck, were adherents of such an epigenetic understanding of generation and development. This conception formed the theoretical base for the teratological studies of the former31 and constituted the very backbone of the transformation theory of the latter. Both men were familiar with the French tradition of epigenesis theories, which goes back to Maupertuis and Buffon.32 It is Maupertuis who deserves particular attention in our context. For, in the frame of his theory of generation, the inheritance of acquired characters appeared not only natural but even inevitable. In contrast with the preformist view, the doctrine of epigenesis had the advantage to be in good conformity with embryological results as well as with then known phenomena of regeneration. However, its disadvantage became obvious when it was about an explanation of how the particles of a developing organism manage to arrange themselves in such a way that they form exactly its complex organic structure. How can one account for this miracle if only the then known mechanical and chemical laws are admissible? As a way out, Buffon proposed moules intérieures in his Histoire des Animaux from 1749, that is, interior molds by means of which the particles are cast mechanically into the structure of the germ.33 Four years earlier, Maupertuis had devised a 27 28 29 30 31 32

Burckhard (1995), pp. 179f. Zirkle (1946), p. 91. Ibid, p. 115. Ibid., p. 117. Appel (1987), pp. 127f. For this tradition, see Roger (1998) part 3.

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different explanation, namely a pangenesis hypothesis that reminds one very much of that of Charles Darwin.34 He supposed that all organs of the parent organisms contribute to the formation of the germ through the production of specific genetic particles. Such specific genetic particles from a certain parental organ were thought to be responsible for the formation of just the same organ of the young. He even ascribed to the genetic particles a kind of memory. Anyway, the assumed transmission of properties of a parental organ to the corresponding one of the offspring via specific genetic particles is mechanically enough conceptualised as to make sure that individual and not generic properties are transmitted.35 It is, furthermore, a transmission of individual traits of the parents that does not discriminate between innate and acquired ones. Inheritance of acquired characters, thus, is a natural consequence of this theory. There were no much improvements of the epigenetic interpretation of generation and development after Maupertuis, and it is clear with hindsight that the followers of the doctrine of epigenesis had no chance to put forward better explanations on the base of the chemistry and physiology of this age, without our distinction between phenotype and genotype, and so on. It was, therefore, certainly not in the first place because of a strong belief in the preformationists’ view when Cuvier, probably the most influential naturalist in the first two decades of the nineteenth century, rejected the doctrine of epigenesis.36 In his view, this doctrine was a deterring instance of the wild speculations characteristic of the eighteenth century that science ought to overcome for the sake of its advancement. He sensed rightly that the epigenesists’ view was part and parcel of broader speculations on how to bridge the gap between the inorganic and the organic world. And Lamarck’s Philosophie zoologique could have served him as an example for the substantiation of this assessment. It contains, apart from other traits of an spirit à la Buffon or à la Maupertuis, a chapter on spontaneous generation,37 in which Lamarck tries to demonstrate that spontaneous generation and generation by fertilisation are essentially processes of one and the same kind that differ only with respect to the stirring principles. As is well known, Lamarck’s theory of species transformation did not meet with much approval among the contemporary professional naturalists. It may be less known, that these naturalists correctly perceived it as part of his Physique terrestre, that is, a planned but only partly completed work, in which Lamarck intended to give a comprehensive theoretical account of the interrelated processes in the atmosphere, the lithosphere, and the biosphere. 38 The knowledge of this embedment of his zoological theory may lead to a better understanding of the contemporary 33

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Buffon (1749), chap. 2. Though throughout mechanical, the idea of such a interior mold was much more intricate than the metaphor of a mold might suggest – see the paper of Mary Terral for the 1st workshop on A Cultural History of Heredity. Maupertuis (1768), Stubbe (1965), pp. 65ff. The idea of a pangenetic generation of the germ is as old as that of the inheritance of acquired characters, and variants of it can be traced back to classic antiquity and even farther backwards – see Zirkle (1946). For Darwin’s pangenesis hypothesis, see Darwin (1894), vol. 2, chap. 17. As Peter McLaughlin’s has stressed in his paper for the 1st workshop on A Cultural History of Heredity, the distinction between the transmission of species-specific and individual traits is of great significance for an adequate historical understanding of eighteenth century heredity conceptions. These conceptions focus almost exclusively on the transmission of the former and neglect the latter nearly completely. However, in mechanical pangenesis hypothesis like that of Maupertuis, it is hard to discern this difference. Species-specificity is only present as a limit of individual deviations. Appel (1987), pp. 49f. Lamarck (1809), vol. 2, chap. 6.

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naturalists’ sceptic attitude towards Lamarck – not towards the expert for zoological classification, but towards the philosophe. Rejecting the entire enterprise of the Physique terrestre, they did not bother themselves with criticising its details, and at least such a comparative trifle like the assumption of the inheritance of acquired characters. The silence about this assumption must, therefore, not be mistaken as approval. Heredity in Lamarck’s and Geoffroy Saint-Hilaire’s zoology, thus, proves to be only a further instance of the peripheral existence of heredity as a subject matter of scientific research before Mendel and the discovery of his discoveries. Also in this case, it surfaces only as a concomitant of other subjects of interest, this time as a complementary aspect of the question of adaptation to changing environmental conditions. It comes therefore without surprise that the question of heredity is treated casually in Lamarck’s and Geoffroy’s zoology and did not lead to investigations into its details, neither by these two scientists nor by their critics. My story about heredity in Lamarck’s and Geoffroy Saint-Hilaire’s zoology turns out to be the story of a missed opportunity.

References Appel, T. A. 1987. The Cuvier-Geoffroy Debate. French Biology in the Decades before Darwin. Oxford: Oxford University Press. Bowler, P. J. 1989. The Mendelian Revolution. The Emergence of Hereditarian Concepts in Modern Science and Society. Baltimore: The Johns Hopkins University Press. Buffon, Georges Louis Leclerc de. 1749. “Histoire des Animaux.” In Histoire naturelle, générale et particulière. Volume 2. Paris: Imprimerie Royale. Burkhardt, R. W., Jr. 1995. The Spirit of System. Lamarck and Evolutionary Biology. Cambridge MA: Harvard University Press. Cahn, T. 1962. La vie et l’oeuvre d’Etienne Geoffroy Saint-Hilaire. Paris: Presses Universitaires de France. Darwin, C. 1894. The variation of animals and plants under domestication. 2nd edition. New York: Appleton. Geoffroy Saint-Hilaire, E. 1826. “Mémoire sur l’anatomie du Crocodile.” Séance de l’Academie des Sciences du 11 avr. 1825. Mémoires de Académie des Sciences 1826. Jacob, F. 1971. La logique du vivant: une histoire de l’ hérédité. Paris: Gallimard. Kant, I. [1775] 1912. “Von den verschiedenen Racen der Menschen.” In Kant’s gesammelte Schriften. (Acc. Edition). Volume 2. Berlin: Reimer. Lamarck, J. B. 1802. Recherches sur l’organisation des corps vivants: précédé du discours d’ouverture du cours de zoologie donné dans le Museum d’Histoire Naturelle. Paris: Maillard. _____. 1809. Philosophie zoologique, ou, Exposition des considérations relative à l’histoire naturelle des animaux. Paris: Chez Dentu [et] L’Auteur. _____. 1963. Zoological Philosophy. New York and London: Hafner Publishing Company. Lefèvre, W. 1984. Die Entstehung der biologischen Evolutionstheorie. Frankfurt am Main: Ullstein. Maupertuis, P. 1768. “Vénus physique (1745).” In Oeuvres. Volume 2. Lyon: J.-M. Bruyset. Roger, J. 1997. Buffon. A Life in Natural History. Ithaca and London: Cornell University Press. _____. 1998. The life sciences in eighteenth-century French thought. Stanford: Stanford University Press. Russell, E. S. 1916. Form and Function. A Contribution to the History of Animal Morphology. London: John Murray. Sapp, J. 1990. “The nine lives of Gregor Mendel.” In Experimental Inquiries, edited by H. Le Grand. Amsterdam: Kluwer. 137-166.

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To my knowledge, Sinai Tschulok was the first who reconstructed the fact and realised the significance of this embedment of Lamarck’s zoological theory – see Tschulok (1937).

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Stubbe, H. 1965. Kurze Geschichte der Genetik bis zur Wiederentdeckung der Vererbungsregeln Gregor Mendels. Jena: Gustav Fischer. Tschulok, S. 1937. Lamarck. Eine kritisch-historische Studie. Zürich: Niehans. Zirkle, C. 1946. “The Early History of the Idea of the Inheritance of Acquired Characters and of Pangenesis.” Transactions of the American Philosophical Society 35(2): 91-151.

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Erasmus Darwin on Hereditary Disease: Conceptualizing Heredity in Enlightenment English Medical Writings Philip K. Wilson

Introduction From ancient times, natural philosophers and physicians have questioned and offered explanations of the passage of hereditary traits including disease. The Hippocratic writers, for instance, attributed a hereditary character to epilepsy, claiming that it arose during the production of semen (a general reproductive fluid that the ancients claimed resided in both males and females) when “diseased parts of the parental body send off diseased seed.” 1 This essay focuses on the period roughly a century before the rediscovery of Mendel – a time of considerable turbulence and discord in both societal and intellectual thought – the shifting era between the Enlightenment and the Romantic Period. My geographical focus – the seedbed of the European Industrial Revolution – lies in England’s West Midland county of Staffordshire. There, I turn to an individual who offered considerable written insight into contemporary thoughts on heredity and disease. This individual, the physician, industrialist, educator of females – all around polymath – Erasmus Darwin. As our subject was better known to his contemporaries as “Dr. Darwin,” I, too, will continue to use the sobriquet “Dr. Darwin,” especially when distinguishing him from other family members. Despite his reputation – at least in some circles – during his own lifetime, he remains to us, at best, the ‘other’ Darwin. Erasmus Darwin has long been a favored study in pedigree analysis, particularly in relation to the intellectual predisposition of his grandson, Charles Darwin. Historians typically cite Dr. Darwin’s section, “On Generation”, from his Zoonomia, or, The Laws of Organic Life (1794-96) to establish an intellectual background for his grandson’s thoughts on adaptation, sexual selection, and evolution.2 Such foreshadowing misses the opportunity to analyze Erasmus Darwin’s thoughts on heredity within their own Enlightenment and early Romantic era contexts. 3 To begin filling this lacuna, this essay will examine several key influences upon Erasmus Darwin that are evident from his own writings on reproductive generation. In particular, I will focus upon Dr. Darwin’s writings on human heredity, an area that even Erasmus Darwin enthusiasts have neglected in favor of his more voluminous and lusty botanical writings. 4 First, I’ll concentrate upon two of his writings that look particularly at human heredity and disease. Then, I’ll examine

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Temkin (1971), pp. 4, 31, 52. Many authors have forged connections on evolution between the two Darwins including Glass, Temkin, and Straus, Jr. (1959); Darlington (1961), esp. pp. 7-13; King-Hele (1963), esp. pp. 63-96; and Harrison (1971). In one early notable exception, Bowler (1974) carefully contextualized eighteenth century views of evolutionary thought. Porter (1989) also addressed this point properly. Sheffield (2002) incorporated Dr. Darwin’s evolutionary views into his historical science fiction. For an overview of Erasmus Darwin and human reproductive generation, see Wilson (Forthcoming). Hassler (1973) convincingly portrays Dr. Darwin as a bridge between Enlightenment and Romantic literati. Schiebinger (1993) has surveyed Erasmus Darwin’s botanical writings in reference to gender.

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briefly the views that several of his contemporaries held regarding hereditary disease in order to better contextualize Dr. Darwin’s own expressed beliefs. Disease was not, to the industrial-minded Dr. Darwin, an entity that invaded the body. Rather, sharing the convictions of his Edinburgh medical school professors, he viewed disease more in constitutional terms as the result of a “malfunction” of the healthy motions within the body. 5 In particular, disease expressed its mechanical defect as a “disturbance in one or more of the classes of fibrous activity.” The physician’s role, therefore, was “to apply remedies which would restore [... the body’s] normal functioning,” or even better, to prevent any malfunction in the first place. 6 Building upon the Enlightenment projects of ordering and classifying information into new knowledge, Dr. Darwin envisioned his Zoonomia as a Linnaean classification for disease.7 “There is need of a theory in the medical profession,” Darwin argued, “a theory founded upon nature, that should bind together the scattered facts of medical knowledge and converge into one point of view the laws of organic life.”8 The passion to reclassify medicine had also been the long-term goal of Edinburgh Professor of Chemistry and Medicine, and later the Institutes (i.e., theory) of Medicine, William Cullen. Like many during this reign of neurophysiological nosologists, Cullen deemed “almost the whole of diseases of the human body might be called Nervous.”9 However, he distinguished his 1769 Synopsis Nosologiae Methodicae from the taxonomies proposed by François Boissier de Sauvages (1763), Carl von Linné (1763), and Rudolph August Vogel (1764) by labeling the “Neuroses” or Nervous Diseases a class of diseases by itself. Dr. Darwin’s nosology, published a quarter of a century later, differed even further. For consistent with his view of society in general, he viewed diseases more in terms of their dynamic, evolving qualities rather than as the static entity more typical of Enlightenment taxonomies.10

Dr. Darwin, Heredity & Disease Moreso than many of his contemporaries, Dr. Darwin readily acknowledged a hereditary predisposition to disease. In Zoonomia and in the Temple of Nature: or, The Origin of Society (1803), he argued that consumption (i.e., tuberculosis), scrofula (the “King’s Evil”), gout, epilepsy, and insanity were hereditary. Ever prone to versify, Darwin stated his argument thusly: E’en where unmix’d the breed, in sexual tribes Parental taints the nascent babe imbibes; 5

6 7

8 9 10

Motions within the fibrous part of the constitution were the key to Dr. Darwin’s understanding of physiology and pathology. Many scholars have pigeonholed Erasmus Darwin as a mechanist without equivocation. His mechanistic inclinations should not surprise us given all of the machines he contrived for his fellow Lunar Society members. However, from a close scrutiny of Zoonomia, I discern his support of a deistic vitalism (or a vitalistic deism). In particular, I find that he proposed many vitalistic notions, albeit generally construed within a linguistic framework of mechanism. McNeil (1987) and King-Hele (1999) provide well contextualized overviews of Dr. Darwin’s industrial-mindedness. Crum (1931), p. 124. Uglow (2002) depicted ordering and classifying as among the key projects of the Enlightenment, p. 266. For a highly readable overview of eighteenth century nosologies, including Dr. Darwin’s, see King (1958), pp. 193-226. E. Darwin (1809), 1: p. viii. Bynum (1993), p. 152. See, for example, Bowler (1974), pp. 166-179; and McNeil (1987), pp. 86-124.

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Eternal war the Gout and Mania wage With fierce unchecked hereditary rage; Sad Beauty’s form foul Scrofula surrounds With bones distorted, and putrescent wounds; And, fell Consumption! Thy unerring dart Wets its broad wing in Youth’s reluctant heart.11

Why did Darwin pay special attention to hereditary disease? In part, he had considerable experience with diseases that appeared in the same family – his own family. Throughout his life, Dr. Darwin suffered from stammering (i.e., stuttering). This “defect” was so pronounced that some said it to be “painful” to hear. Others, however, claimed that Darwin “repaid his auditors [i.e., attentive listeners] so well for making them wait for his wit or knowledge, that he seldom found them impatient.”12 When one young man questioned him as to whether he found this “deficit” to be inconvenient, Dr. Darwin replied, with typical wit, “No, sir, it gives me time for reflection, and saves me from asking impertinent questions.”13 Darwin’s eldest son, Charles, was also afflicted with stammering. Darwin sent this son, when 18, to France, in hopes that “if he was not allowed to speak English for a time, he would be cured of [… stammering]. Charles returned a year later, spoke French fluently for the rest of his life, but continued to stammer in English.” 14 For Dr. Darwin, this served as evidence that at least some hereditary diseases could be modified in their expression. His further investigation into this familial linkage was stopped when Charles, as a prize-winning medical student at Edinburgh, died from blood poisoning following a finger cut at the dissecting table. Dr. Darwin also suffered from gout, a disease thought by many to be as inherited as titles among the upper class. William Darwin, Erasmus’s great-great-grandfather, had died in 1644, it was claimed, “from gout.” It was “therefore probable,” claimed Erasmus’s grandson, the gouty naturalist Charles Darwin, that Dr. Darwin, as well as many other members of the family, “inherited from this William, or some of his predecessors, their strong tendency to gout.” 15 Dr. Darwin also gathered considerable case study evidence from his wide-ranging practice, predominantly though not exclusively among the middle and upper ranks of society, that he interpreted as further proof of a hereditary transmission of gout.16 It was also an “early attack of gout” that turned Dr. Darwin into a “vehement advocate of temperance” for the rest of his life.17 Darwin viewed intemperance (i.e., alcoholism) as the foundational hereditary disease. In the words of his grandson, “No man ever inculcated more persistently and strongly the evil effects of intemperance than did Dr. Darwin […].” 18 Dr. Darwin 11 12 13 14 15 16

17 18

E. Darwin (1804), Reproduction of Life, Canto II. Pearson (1930), p. 42. Krause (1879), p. 40. For an historical overview of stammering, see Compton (1993). Pearson (1930), pp. 9-10. C.Darwin (1879), p. 1. Although based in Lichfield and later Derby, Dr. Darwin’s practice was based over a large geographical area. He traveled extensively, either in his carriage or on horseback, to see patients. In 1766 alone, he calculated having traveled over ten thousand miles. C. Darwin (1879), p. 1. C. Darwin (1879), p. 56. As Pearson (1930) noted (p. 27), the novelist Maria Edgeworth, daughter of Lunar Man Richard Edgeworth, also testified that Dr. Darwin “believed that almost all the distempers of the higher classes of people arise from drinking […] too much vinous spirit.”

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himself argued that all the “hereditary diseases of [Britain]” originated as “the consequence of drinking much fermented or spiritous liquor.”19 Consequently, he argued that overcoming the hereditary tendency to intemperance was “perhaps […] the most practical line of attack” against all “ill-health.”20 Turning again to his own family lineage, Darwin shared thoughts about intemperance as a hereditary disease. In a letter to his son Robert Waring Darwin, Erasmus discussed his own fatherin-law, Charles Howard. Howard, he claimed, was “a drunkard both in public and private – and when he went to London he became connected with a woman and lived a deba[u]ched life in respect to drink, hence he always had the Gout of which he died.” 21 These effects carried forth into the life of Mary Howard, Dr. Darwin’s first wife. Mary Darwin began to drink excessively early in their married life – though attempting to conceal this from her husband – in order to overcome episodes of “temporary delirium, or […] insanity.”22 Enlightenment medical authorities including William Heberden, William Hunter, and William Cullen – all with whom Dr. Darwin had studied – as well as the physician Thomas Beddoes, founder of the Pneumatic Institute in Bristol, and the Birmingham physician and fellow Lunar Society man, William Withering, also viewed what were called “the drunken diseases” [including gout, epilepsy, and insanity] as “hereditary in some degree.” 23 The Scot-born gouty physician and literati, Tobias Smollett, also featured the hereditary passage of disease in his popular late eighteenth century novels, The Adventures of Peregrine Pickle (1751) and The Adventures of Humphrey Clinker (1771). Consistent with these authorities, Dr. Darwin claimed that it was the tendency, the diathesis, the predisposition to disease rather than the particular disease itself that was hereditary.24 Contemporaries described “hereditary predisposition” as an “original conformation of the body, transmitted from the parent to the offspring” that, “when particular exciting causes are applied, a similar train of morbid phenomena takes place” in the child as was experienced by the parent.25 Predisposition, the precise factor that was inherited, was deemed as “the medium between health and disease.”26 Thus the susceptibility to be afflicted with a hereditary disorder like gout, epilepsy, insanity, or consumption could be enhanced when an individual was exposed to certain triggering environments. Or, using the terminology of Dr. Darwin’s day, the remote cause of disease (its inheritance) could be triggered by the proximate or “exciting” cause (cold, heat, some spasm or other debilitating disturbance in one’s environment).27 The clime unkind, or noxious food instills To embryon nerves hereditary ills; 19 20 21 22 23 24 25 26 27

E. Darwin (1804), p. 178. C. Darwin (1879), p. 56. E. Darwin ([1792] 1958), p. 224. Colp (1977), p. 119. E. Darwin ([1792] 1958), p. 224. For complementary accounts of the Lunar society, see Schofield (1963) and Uglow (2002). For an overview on diathesis, see Ackerknecht (1982); and for an elaboration on the importance of “constitution” in regard to hereditary disease, see Olby (1993). Trotter (1808), p. 169. Ibid., p. 204. Bynum (1994), p. 19.

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The feeble births acquired diseases chase, Till Death extinguish the degenerate race.28

Like the claims of many natural philosophers both before and after this time, Dr. Darwin viewed disease itself, even when labeled as a hereditary disease, as the result of both nature and nurture. 29 Specifying the particular cause of disease was, to a nosologist like Darwin, of utmost importance for the precision of his diagnoses. Identifying the causal factors, however, seemed less significant in determining the actual treatments he proffered. Rather, by implicating the degree to which nature –versus– nurture was viewed as the chief “cause” of a particular disorder helped Dr. Darwin target how to best prevent the further occurrence of this disease within a particular family. If a patient’s past pedigree suggested that he was the likely carrier of a hereditary propensity to disease, then it was the physician’s role to prevent the patient from being exposed to the precipitating factors that would most likely induce the onset of disease. Neglecting to treat the symptoms of a reputed hereditary disease could also present a problem. For if “improperly treated,” a hereditary disorder like the gout may be “diverted from its proper course” such that “the miserable patient has a chance to be ever after tormented with head-achs, coughs, pains of the stomach and intestines.”30

Dr. Darwin on Consumption and Gout Darwin’s thoughts on hereditary disease, especially consumption and gout, were influential to several contemporaries including the physician, Thomas Trotter, of Newcastle on Tyne. Trotter, whose mind was fixed on the nervous temperament as the exciting, proximate cause to all disease, also incorporated the predisposition to hereditary disease in his diagnoses. Indeed, Trotter argued that the inherited predisposition itself may become visibly apparent long before the patient complained of specific symptoms. Such was the case, he argued, of those “phthisically disposed,” i.e., those marked with the predisposition to phthisis (i.e., consumption or tuberculosis). Markings of phthisis – the pallid, gaunt face, and the sunken chest – may frequently appear in the first stages of infancy, late in infancy, or even later in development in patients afflicted with a latent form of disease. The onset of puberty, for example, was a particular time in later life where “changes […] in the constitution, conjoined to the quick growth of the body” proved to be “a most critical period” for the onset of symptoms of phthisis as well as other hereditary diseases. 31 According to Dr. Darwin, hereditary pulmonary consumption was known to “attack” patients “so infallibly a few years after puberty, that it does not appear to depend much on external circumstances.”32 Thus, at times, nature appeared to have heavily outweighed nurture in precipitating the onset of hereditary disease.

28 29

30 31 32

E. Darwin (1804), Reproduction of Life, Canto II. López-Beltrán (1994) has described that a malleable admixture of nature and nurture existed in the “soft hereditarianism” beliefs of the early nineteenth century in contrast to the more objective qualifications of a nature-based, “hard hereditarianism” later in the century. Buchan (1828), p. 277. T. Trotter (1808), p. 172. For descriptions of later views of hereditary tuberculosis, see Wilson (Forthcoming). E. Darwin (1809), 2: p. 255.

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Coupled with the evidence for hereditary disease garnered from his own family lineage, Dr. Darwin gathered additional evidence from many of the families for whom he offered medical care. “I have seen epilepsy,” one of the “drunken diseases,” “produced […] very often” in the same family. As a treatment, he noted from his own success that “one sober generation” can cure epilepsy. Insanity, another ”drunken disease,” could be stopped in a similar manner. 33 Indeed, Dr. Darwin claimed to have treated many people ”who had insanity [… on] one side” of their family, yet whose “children now old people have no sign of it.” His practice demonstrated that diseases, like gout and intemperance, though inheritable, were not inevitable. Consistent with the views of later historians of the Enlightenment, hereditary diseases could, at times, be disinherited. 34 In particular, hereditary predispositions “cease,” Darwin claims to have observed, “if one or two sober generations succeed; otherwise the family becomes extinct.” 35 If the “drunken diseases” were not curable by imposing one sober generation upon the family, then, Darwin exclaimed, “there would not be a family in the kingdom without epileptic, gouty, or insane people in it.”36 Not everyone shared Dr. Darwin’s belief in the transmission of hereditary predispositions, or even in hereditary diseases themselves. The irascible Edinburgh physician, John Brown, whom Darwin admired for quantifying “excitation” and “stimulation” in his explanation of the actions of human physiology, disease, and medications, disapproved of applying the concept of “hereditary” to any disease. Brown’s theory – known widely, especially throughout continental Europe as Brunonian medicine – was based upon the belief that every individual’s temperament or constitution was the same. Drawing upon Isaac Newton’s revolutionary claim that “one single principle” governed all motion in “the whole planetary systems of the universe,” Brown conjectured that the human form was similarly organized around one single human constitution.37 Disease, according to Brown, an ex-pupil and extramural rival of William Cullen, resulted from either an excess or deficiency of “excitability” (i.e., the capacity to react to external stimuli).38 To claim that “a taint, transmitted from parents to their offspring” should be “celebrated under the appellation of hereditary” was, Brown argued, “a mere tale.” For true Brunonians, “there is nothing to the fundamental part of [… the] doctrine of hereditary disease.”39 As evidence, he speculated that the “sons of the rich,” who “succeed to their father’s estate, succeed also to [… their] gout. Those [sons] who are excluded from the estate, escape the disease also, unless they bring it on by their own conduct.” It is interesting that despite his adamant disbelief in the concept of hereditary disease, his recommendations for preventing such disorders were not that dissimilar from those of Dr. Darwin. In one of his rationalized case examples, Brown argued that although “Peter’s father may have been affected with the gout, it does not follow that Peter must be affected; because, by a proper way of life, that is by adapting his excitement to his stamina, he may have learned to evade his father’s disease.” “If the same person,” he continued, “who from his own fault and improper management, has fallen into the disease; 33 34 35 36 37 38 39

E. Darwin ([1792] 1958), p. 224. Porter and Rousseau (1998), p. 117. E. Darwin (1809), 1: p. 414. E. Darwin ([1792] 1958), p. 225. Brown (1803), pp. 242-243. Bynum (1994), p. 17. Yet, as Lawrence (1988) has insightfully pointed out, identifying the essential characteristic(s) of Brunonian medicine has been a difficult task for over two hundred years. Brown (1803), p. 396.

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afterwards, by a contrary management, and by taking good care of himself, prevents and removes the disease, […]. What then [… has] become of [the] hereditary taint?” Alas, there is nothing whatsoever hereditary about this or any other disease. “Whatever produces gout in one [individual], will produce it in another. And whatever cures it in any one [individual], cures it also in every other.”40 Another contemporary, William Cadogan, physician to the Foundling Hospital in London, an institution formed upon Enlightenment ideals, also questioned the claim that diseases were hereditary. Like Brown, he argued that if a disease was truly hereditary, it would “necessarily be transmitted from father to son, and no man whose father had it could possibly be free from it.” His own extensive practice and readings had provided him “many instances to the contrary.” Cadogen explained further, “Our parents undoubtedly give us constitutions similar to their own, and, if we live in the same manner they did, we shall very probably be troubled with the same diseases; but this by no means proved them to be hereditary: it is what we do to ourselves that will either bring [… the diseases] on, or keep us free.”41 Cadogan argued his case specifically in reference to gout.42 This select paragraph exemplifies his numerical-based (i.e., statistical) critique. Those, who insist that the gout is hereditary, because they think they see it so sometimes, must argue very inconclusively; for if we compute the number of children who have it not, and women who have it not, together with all those active and temperate men who are free from it, though born of gouty parents; the proportion will be found at least a hundred to one against that opinion. And surely I have a greater right from all these instances to say that it is not hereditary, than they have from a few to contend that it is. What is all this, but to pronounce a disease hereditary, and prove it by saying that it is sometimes so, but oftenernot so? Can there be a greater absurdity.”43

Although many, including Darwin, concurred with Cadogan on temperance, they opposed his view regarding gout and heredity. William Falconer of Bristol rebuked Cadogan in a 1772 treatise as did Aberdeen practitioner William Grant in an essay seven years later. Grant firmed up a general view between constitution and hereditary disease by arguing that just as “constitutional diseases are often hereditary, […] the hereditary diseases are always constitutional.” 44 The Lichfield luminary, Samuel Johnson, praised Cadogan’s work as “a good book in general” but criticized it as a “foolish one in particulars. […] ‘Tis foolish, as it says, the gout is not hereditary.” 45 Consistent with his deistic beliefs that development on the earth followed no fixed plan, Dr. Darwin argued that disease, too, was not predestined. Following the dissenter’s practice of working to improve one’s lot during his earthly existence, one could also work to overcome or prevent disease, including diseases to which one might be hereditarily predisposed. To achieve these aims, Darwin argued, one must learn how best to exert power over nature. Such objectives 40 41 42 43 44 45

Ibid., p. 243. Cadogan (1771), p. 70. Porter and Rousseau (1998), p. 104, call Cadogan’s view on gout and heredity as the “paradigm-shift framing his entire theory.” Cadogan (1771), p. 70. Porter and Rousseau (1998), p. 115. Ibid., p. 81.

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also represented Dr. Darwin’s critique on Britain’s traditional social fabric that enforced aristocratic, class-based structures of inheritance.

Darwin & Buchan: Progressive-minded Reforms of Inheritance Darwin’s views were most consistent with the progressive-mindedness of his fellow Edinburgheducated physician, William Buchan (1729-1805). Indeed, Buchan and Darwin were likeminded in many ways. They both supported the revolutionary action in the American colonies and in France; they both promoted the diffusion of useful knowledge; they were both avidly protemperance decades before any formal organization was to be found in Britain. In short, they were both liberal, progressive-minded reformers who exemplified that doctors, like politicians, could also be crusaders for social improvement. Buchan, however, distinguished himself from Darwin through the aim of his medical writings. Unlike Darwin, whose chief medical writing, Zoonomia, was addressed to his fellow practitioners, Buchan, in the true spirit of Enlightenment’s view of the democratic power of nature, sought to put medical wisdom in the hands of the public. His principle work, Domestic Medicine, first appeared in 1769 and was reprinted throughout the nineteenth century. Like John Wesley’s Primitive Physic (1747), and the anonymously written midwifery manual, Aristotle’s Masterpiece, Domestic Medicine served many needs of the lay public. Some claim that Buchan’s book partnered with the Bible in its popularity.46 As a means of appreciating what much of the public in Darwin’s Britain understood regarding hereditary disease, Buchan’s Domestic Medicine continues its usefulness in serving us as a valuable guide as well. Based upon his own extensive practice and reading, Buchan claimed to have uncovered “manifold and decisive proofs” of the heritable transmission of disease. In particular, he noted that during “certain periods of life,” children “become liable to the diseases of their parents, and consumption, gout, or dropsy makes its appearance, the germs of which must have lain in the system from the earliest periods of existence, although they did not disclose themselves till their due season.”47 But once a disease is “contracted and riveted in the habit,” it became “entailed on posterity.”48 Expanding beyond Darwin, Buchan suggested how to anticipate the future occurrence of hereditary disease. He informed readers that children were “particularly prone” to the diseases of the parent “to whom they bear the greatest personal similarity.” Such direct linkage was not the sole key, however, for he warned that just as “we occasionally perceive the resemblance of some more remote ancestor break forth” in the personality and characteristics of a child, “so we shall [also] find the constitution and diseases of that child […] in the nature of the progenitor whom it most resembles.”49 Further noting that this “point of similarity between parents and children” regarding the hereditary disease had “not hitherto been sufficiently attended to,” Buchan focussed considerable attention to this familial connection in discussing diseases including consumption, gout, scrofula, 46 47 48 49

Porter (1992), p. 217. Buchan (1828), p. 162. Ibid., pp. 441-442. Ibid., p. 162.

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scurvy, and dropsy – all disorders that he identified as having a hereditary predisposition. 50 These disorders must, he continued, “be the result of a certain combination of habits, continued, perhaps, from one generation to another, combined with the peculiar circumstances in which the individual is placed. It is reasonable to suppose that, by altering the former, and counteracting the latter, the general constitution might be changed”51 But just what did Buchan mean by “altering the former” – that is nature’s contribution – regarding the cure of hereditary disease? Recalling Jean-Jacques Rousseau’s analogies that were popular in England at this time, Buchan reminded readers of the farmers’ inability to reap “a rich crop from a barren soil.” Delicate women and men who pursue “irregular” habits may, he acknowledged, be able to bring forth children, but such children are “hardly […] fit to live.” The “first blast of disease” tends to “nip the tender [child] in the bud.” Although some of these children may be able to “struggle through a few years’ existence,” their “feeble frame[s], shaken with convulsions from every trivial cause,” leave them “unable to perform the common functions of life.” In short, they “prove a burden to society.” 52 This burden, Buchan was quick to note, was not perpetuated only amongst the lower classes. “How happy had it been,” he charged, “for the heir of many a great estate had he been born a beggar, rather than to inherit his father’s fortunes at the expense of inheriting disease.” 53 Indeed, the lower classes might actually hold some predilection against the onset of the hereditary diseases that were precipitated by the Epicurean living of England’s upper class. Beyond the diagnostic significance of identifying hereditary predisposition, Buchan, perhaps more than any other contemporary, challenged readers to accept the responsibility they held, individually and collectively, in improving future generations. Most pointedly, Buchan proclaimed, it was in the name of progress that any “person laboring under any incurable [hereditary] malady ought not to marry.” Doing so, he continued, not only “shortens his own life, but [also] transfers its misery to others.” When “both parties are deeply tainted” with hereditary diseases, their offspring, he prognosticated, “must be very miserable indeed.” 54 Working to improve one’s circumstances beyond what nature provided was consistent with industrial, progressive-minded, Enlightenment ideals. And by placing further responsibility for the future in the hands of the public, Buchan believed himself to be enhancing the progressive cause. “Want of attention to these things” in the past, he proclaimed, had “rooted out more families than plague, famine, or the sword.” And unless individuals became more informed and active in suppressing the propagation of the diseased, the “evil” associated with such disorders was bound to continue.55 Thus for Buchan, as for Dr. Darwin, the social reality of hereditary disease had significantly prohibited human progress. To enhance a more positive evolution of humanity, he argued that people must assist and encourage nature to work in a progressive manner. Believing that particular diseases were introduced via marriage lines, Darwin, too, suggested that more attention should be directed to one’s choice of marriage partner. The “art to improve the sexual progeny” in humans would, he argued, follow the choice of “the most perfect of both 50 51 52 53 54 55

Ibid., p. 162. Ibid., p. 163. Ibid., p. 441. Ibid. Ibid., p. 442. Ibid.

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sexes.” Fertility itself, Dr. Darwin claimed, was hereditary and subject to selection. 56 Noting that some families had “become gradually extinct by hereditary diseases,” he offered the cautionary note that “it is often hazardous to marry an heiress, as she is not infrequently the last of a diseased family.”57 Offspring would also be less liable to hereditary disease, Dr. Darwin argued, it they married into different families. Such views contrasted traditional aristocratic family structure in which “like” tended to marry “like.” They convey Dr. Darwin’s social progressive-mindedness, and they were consistent with what Lawrence Stone (1977) and others have characterized as contemporary attempts to break from the traditional notions of fixed patterns of inheritance. These views, proposed in Dr. Darwin’s last work, The Temple of Nature, were also foundational to what, a century later, became incorporated into programs of both positive eugenics (i.e., promoting the marriage and proliferation of “good stock”) and negative eugenics (i.e., the prohibition of marriage and breeding between “defective stock”). Few physicians in Britain expanded more upon the particulars of what might be termed the “eugenics” of this era than did William Buchan. Buchan noted that responsible actions were required of both women and men towards shaping the future in a progressive-minded manner. If women would “reflect on their own importance and lay it to heart, they would embrace every opportunity of informing themselves of the duties which they owe to their infant offspring.” It was “their province,” Buchan argued, “not only to form the body, but to give the mind its most early bias.” Women “have it very much in their power” to make their children “healthy or valetudinary, useful in life or the pests of society.”58 “Were the time that is generally spent by females in the acquisition of trifling accomplishments employed in learning how to bring up their children […] so as best to promote their growth and strength […], mankind would derive the greatest advantage from it.” But until the education of females expands beyond “what relates to dress and public show, we have nothing to expect from them but ignorance” in important concerns such as their role in preventing the propagation of hereditary disease.59 It should be noted that Darwin, too, devoted considerable attention to improving the education of women. His chief work, A Plan for the Conduct of Female Education in Boarding Schools, written, ironically, for his two illegitimate daughters, the “Misses Parkers,” was aimed at improving the minds and bodies of girls attending the boarding school that he had established at Ashbourne in Derbyshire. Progress in the body, as in society, was, in part, achievable through improving nurture.60 Buchan argued that men, too, must become more attentive to their role in improving their offspring. Why is it that men are “not ashamed to give directions concerning the [… breeding of] dogs and horses, yet would blush were he [… to perform] the same office for that being who derived its existence from himself, who is the heir of his fortunes, and the future hope of his country!” 61 Darwin similarly noted that “those who breed animals for sale” had long known that “the sexual progenies of animals” were “less liable to hereditary diseases, if the breeding took place between families rather than within the same family.”62 For when both parents suffer from the 56 57 58 59 60 61

Darlington (1961), p. 11. E. Darwin (1804), p. 179. Buchan (1828), p. 440. Ibid. For further discussion of Erasmus Darwin’s model of female education, see Schiebinger (1993). Buchan (1828), p. 440.

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same hereditary disease, he argued that the disease was more likely to descend to their posterity. Thus, another way to achieve progress among humanity resulted from improvements upon nature. Both Darwin and Buchan popularized the proscription of marriage when hereditary disease was a family concern. Buchan also charged physicians as being negligent in showing attention to hereditary disease. This inattentiveness, he argued, was due largely to their unwillingness to attend to the sickness, management, and “preservation” of children. How striking the “labour and expense [that] are daily bestowed to prop up an old tottering carcase for a few years, while thousands of those who might be useful in life perish without being regarded! Mankind [is] too apt to value things according to their present, not their future usefulness.”63 Such characterization symbolizes a difficulty that had long plagued preventative thinking in medicine. Focussing on the here and now in medical practice had been (and continues to be) the norm, thereby leaving little energy (and funding) to expend on improving the lot of humanity for the future. 64 Darwin, Buchan and many of their contemporaries based their medical practices, in part, on altering one’s physical constitution in order to enhance resistance to disease. Habits, characteristics – indeed – physical constitutions were thought to be subject to alteration. Subsequently, the altered or acquired forms of a constitution were heritable. Such a “transformation of descent”65 exemplified what has been labeled “trans-generational progress.” 66 In the minds of some, the altered constitutional makeup was thought to be passed along intact to the next generation. Thus Darwin anticipated what Gottfried Reinhold Treviranus and, more notably, Jean-Baptiste Lamarck would later argue about the passage of acquired characteristics. In his Philosophie Zoologique (1806), Lamarck provided considerably more evidence drawn from nature than did Dr. Darwin. Moreover, he argued for a directional development that Darwin had not done. Still, in the context of industrial revolutionaries, Darwin’s arguments set forth in Zoonomia “biologised the concept of progress.”67 The progressive developments in organic life were carried forth into the embryonic development of the next generation, or in other words, inherited. It might be said, contrary to the usual phrasing, that Lamarck was actually Darwinian in his thinking.68

Nineteenth century Heredity & Disease after Dr. Darwin Darwin routinely had the ear of fellow Lunar Society men like Josiah Wedgewood, Matthew Boulton, and James Watt, industrialists whose success depended upon unprecedented demands upon the work force. More than some, Wedgewood appreciated that his maximal financial gain depended in large part upon maximizing the health of his workers. In part, through Dr. Darwin’s 62 63 64 65 66 67 68

E. Darwin (1804), p. 178. Buchan (1828), p. 441. Rosen (1977), pp. 69-77. Darlington (1961), p. 10. McNeil (1987), p. 112. Ibid., p. 123. Krause (1879), p. 133, made a similar comparison, rightly noting that it is “more proper” to view Jean Lamarck as a Darwinian of the “older school” than to characterize Dr. Darwin as a Lamarckian. Drachman (1930), p. 88, also addressed this point.

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efforts, public health and hygiene became critical concerns within the industrial Midlands. In the generation immediately following these Lunar luminaries, public health concerns became increasingly addressed in these heavily populated towns-turned-cities. The divide between the “haves” and the “have nots” intensified. From an environmental perspective, the financially dependent were crowded into unsanitary areas that soon became known for their high morbidity and mortality. From a hereditary perspective, these rates were thought to increase even further for these environments could easily excite those with predispositions to particular diseases into full blown manifestations of their disorders. It could be argued – though with sweeping generalizations – that concern about hereditary disease became relatively inconsequential in England during the decades immediately following Dr. Darwin. With the rise of industry, inheritance was no longer the only way to achieve substantial financial gain. The growth of industry also diminished the significance of inheritance within the minds of many medical reformers. Individuals including William Wilberforce, Lord Shaftsbury, Jeremy Bentham, and Edwin Chadwick focused much of their reform efforts on improving the environmental factors related to disease and public health. Although some claimed Dr. Darwin to be a pioneer of temperance reform, later temperance campaigns operated from the premise that drinking was more of a societal ill than an inherited one. 69 The rise of bacteriology later in the nineteenth century further diverted the medical gaze, focusing more upon the germ than the germ cell. With efforts aimed at cleansing society from germs, the concept of hereditary disease might appear to have lain dormant within British medical and popular writings throughout much of the later half of the nineteenth century. According to standard historical accounts, heredity was only excited into action in after another British polymath, Francis Galton, a grandson of Erasmus Darwin, drew attention to what he called the “study of agencies, under social control, that may improve or impair the social qualities of future generations either mentally or physically.”70 Encouraging a shift away from the primary focus on bacteriology, Galton proposed that massive efforts should be undertaken to improve the human reproductive stock of society by giving “the more suitable races or strains of blood a better chance of prevailing speedily over the less suitable.”71 Although some truth lies within such explanations, we are only beginning to see more clearly through works of Carlos López-Beltrán (1994), John C. Waller (2002) and others that support for hereditary explanations of disease remained strong throughout nineteenth century Britain. Charles Darwin’s son, Leonard Darwin – a key spokesperson for British eugenics in the early twentieth century – claimed that this interest in heredity, both as related to disease and to societal well-being, was itself a commonly noted trait passed along through family pedigrees. For his family, this may well have been true. However, our continually refined focus upon the passage of ideas, or influence, of one generation of medical writers upon succeeding generations will further illuminate areas of continuity and areas of change regarding more general thoughts about the possible connections between heredity, environment, and disease in the nineteenth century.

69 70 71

Pearson (1930), p. 27. Galton ([1909] 1984), p. 81. Galton (1883), p. 24.

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References Ackerknecht, E.H. 1982. “Diathesis: The Word and the Concept.” Bulletin of the History of Medicine 56: 317325. Bowler, Peter J. 1974. “Evolutionism and the Enlightenment,” History of Science 12: 159-183. Brown, John. 1803. The Elements of Medicine. Portsmouth, N.H.: William & Daniel Treadwell. Buchan, William. 1828. Domestic Medicine; or, A Treatise on the Prevention and Cure of Disease. Exeter: J. & B. Williams. Bynum, W.F. 1993. “Cullen and the Nervous System.” In William Cullen and the Eighteenth Century Medical World, edited by A. Doig et al. Edinburgh: Edinburgh University Press. _____. 1994. Science and the Practice of Medicine in the Nineteenth Century. Cambridge: Cambridge University Press. Cadogan, William. [1771] 1925. A Dissertation on the Gout and All Chronical Diseases Jointly Considered. Boston: Henry Knox. Reprinted in John Ruhräh. “William Cadogan and His Essay on Gout.” Annals of Medicinal History 7: 64-92. Compton, David. 1993. Stammering: Its Nature, History, Causes, and Cures. London: Hodder and Stoughton. Colp, Ralph, Jr. 1977. To Be An Invalid: The Illness of Charles Darwin. Chicago: University of Chicago Press. Crum, Ralph B. 1931. Scientific Thought in Poetry. New York: Columbia University Press. Darlington, C.D. 1961. Darwin’s Place in History. New York: MacMillan. Darwin, Charles. 1879. “Preliminary Notice.” In Erasmus Darwin, by Ernst Krause.Translated by W.S. Dallas. London: John Murray. Darwin, Erasmus. 1804. The Temple of Nature; or, The Origin of Society. New York: T. and J. Sword. _____. 1809. Zoonomia: or, The Laws of Organic Life. In Three Books. Boston: Thomas and Andrews. _____. [1792] 1958. “Letter to Robert Waring Darwin, 5 January.” In The Autobiography of Charles Darwin, 1809-1882. Edited by Nora Barlow. New York: W.W. Norton. Drachman, Julian M. 1930. Studies in the Literature of Natural Science. New York: MacMillan. Galton, Francis. 1883. Inquiries into Human Faculty and Its Development. London: J.M. Dent. _____. [1909] 1984. “Probability the Foundation of Eugenics.” In Essays on Eugenics. New York: Garland. Glass, Bentley, Owsei Temkin, and William L. Straus, Jr., eds. 1959. Forerunners of Darwin, 1745-1859. Baltimore: Johns Hopkins University Press. Harrison, James. 1971. “Erasmus Darwin’s View of Evolution.” Journal of the History of Ideas 32: 247-261. Hassler, Donald M. 1973. Erasmus Darwin. New York: Twayne Publishers. King, Lester S. 1958. The Medical World of the 18th Century. Chicago: University of Chicago Press. King-Hele, Desmond. 1963. Erasmus Darwin. New York: Charles Scribner’s Sons. _____. 1999.Erasmus Darwin: A Life of Unequalled Achievement. London: Giles de la Mare Kraus, Ernst. 1879.Erasmus Darwin. Translated by W.S. Dallas. London: John Murray. Lawrence, Christopher. 1988. “Cullen, Brown, and the Poverty of Essentialism.” Medical History Supplement 8: 1-21. López-Beltrán, Carlos. 1994. “Forging Heredity: From Metaphor to Cause, a Reification Story.” Studies in the History and Philosophy of Science 25: 211-235. McNeil, Maureen. 1987. Under the Banner of Science: Erasmus Darwin and His Age. Manchester: Manchester University Press. Olby, Robert C. 1993. “Constitutional and Hereditary Disorders.” In Companion Encyclopedia to the History of Medicine, edited by W. F. Bynum and R. Porter. New York: Routledge. 412-437. Pearson, Hesketh. 1930. Doctor Darwin. London: J.M. Dent. Porter, Roy. 1989. “Erasmus Darwin: Doctor of Evolution?” In History, Humanity and Evolution: Essays for John C. Greene, edited by James R. Moore. Cambridge: Cambridge University Press. _____. 1992. “Medicine in Georgiand England.” In The Popularization of Medicine 1650-1850, edited by Roy Porter. London: Routledge. Porter, Roy and G.S. Rousseau. 1998. Gout: The Patrician Malady. New Haven, CT: Yale University Press. Rosen, George. 1977. Preventive Medicine in the United States 1900-1975: Trends and Interpretations. New York: Prodist. Schiebinger, Londa. 1993. Nature’s Body: Gender in the Making of Modern Science. Boston: Beacon Press. Schofield, Robert E. 1963. The Lunar Society of Birmingham. Oxford: Oxford University Press. Sheffield, Charles. 2002. The Amazing Dr. Darwin. The Adventures of Charles Darwin’s Grandfather: Its All in the Family. Riverdale, NY: Baen Books.

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Stone, Lawrence. 1977. The Family, Sex and Marriage in England, 1500-1800. New York: Harper Colophon. Temkin, Owsei. 1971. The Falling Sickness: A History of Epilepsy from the Greeks to the Beginnings of Modern Neurology. Baltimore: Johns Hopkins University Press. Trotter, Thomas. 1808. A View of the Nervous Temperament; Being a Practical Inquiry into the Increasing Prevalence, Prevention, and Treatment of those Diseases commonly called Nervous, Bilious, Stomach & Liver Complaints; Indigestion; Low Spirits, Gout, &c. Troy, NY: Wright, Goodenow, & Stockwell. Uglow, Jenny. 2002. The Lunar Men: The Fiends Who Made the Future. London: Faber and Faber. Waller, John C. 2002. “’The Illusion of an Explanation’: The Concept of Hereditary Disease, 1770-1870.” Journal of the History of Medicine and Allied Sciences 57: 410-448. Wilson, Philip K. Forthcoming. “Erasmus Darwin on Human Generation: Placing Reproductive Generation within Historical and Zoonomian Contexts.” In The Genius of Erasmus Darwin, edited by C.U.M. Smith and R. Arnott. Aldershot, Hampshire: Ashgate. _____. Forthcoming. “Confronting ‘Hereditary’ Disease: Eugenic Attempts to Eliminate Tuberculosis in Progressive Era America.” Journal of Medical Humanities.

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Introduction To understand the widespread popularity of the notion ‘pathological heredity’, we must first identify the ways in which physicians acted in specific conditions to pursue their interests and ideals and to spread their convictions. By the end of the 18th century, heredity was a topic of debate not just among philosophers but among plant and animal hybridists as well. Philosophers speculated about the continuity and progress of mankind. Most of their theories were dominated by fantastical representations of generation. Historically, breeders had fundamental concerns focused on the generation and transmission of peculiarities. Using empirical methods, they sought to isolate and preserve acquired features among animals and plants. Meanwhile, influential physicians saw the question of heredity as a physiological problem rather than a pathological one. They considered the conundrum of heredity as being “beyond [their] immediate grasp.” In 1815, when young physicians were discharged from Napoleon’s Armies, they faced a problem of overcrowding in the medical profession. Like other middle-class professions in Western Europe between 1815 and 1848, medicine was suffering from the discrepancy between demand for services and supply (i.e. medical school graduates). Paris certainly was saturated, with only 3 percent of the national population but 13 percent of all doctors. The result was bitter competition for patients and for salaried hospital positions. In 1819, the Dictionnaire des Sciences Médicales had an entry for “Médecine politique”, described in normative terms as “the series of relations that doctors ought to have with governments in the interest of the governed.” It was noted that most of these ‘relations’ were not highly developed – a situation that was soon to change. After Fodéré had published his Traité de Médecine légale et d’Hygiène publique, in 1813, no well-informed physician spoke about hereditary diseases in the same way as his predecessosrs had done some 20 or 30 years before. Then the same disease with the same symptoms had been thought to appear in children at the same age and in the same circumstances as it had appeared in their affected parent. Now, they would use the key terms, “hereditary predisposition”, “hereditary taint” and “predisposing factors” to talk about hereditary disease. Heredity transmitted an organic disposition not only to one particular disease but also to a special condition that could produce a variety of illnesses or disabilities. As this new model of thinking developed and spread through the medical corps, an unprecedented notion took root: chronic diseases would appear in those with an inherited constitutional weakness. This constitutional weakness was inherited at the time of the conception, birth or during weaning. At this stage, hypotheses diverged but the physicians’ basic argument was simple: Given that the transmission of external physical characteristics seemed to be constant, there could be no doubt that the same transmission process was at work in determining man’s internal constitution. So, when the same chronic disease affected several

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members of the same family, transmission by heredity was involved. It was convenient in medical language to mix up the use of the terms “inherited diseases” and “family diseases”. When referring to contagious illnesses, observers, not being able to explain why contagion did not affect every individual in a group to the same extend, began to write passionately on this question of transmission by heredity. The question I want to address is the following: at a time when progress was the driving force of history and science, was there a group of influential physicians who were willing to adopt a new attitude on such essential issues as the status of medical theories and the role of the medical corps in the new political context, the insuperable obstacles to real knowledge of the causes of illnesses – a new attitude, then, which replaced the vagueness of theory with the positive and sure light provided by statistical data, held as a fundamental value?

1. Medicine is a science Now, only science could guarantee progress in knowledge. Physicians had been able to observe that great improvements were being made in sciences like chemistry, physics and astronomy but progress in medical science was almost nil. So, anxious to ensure the scientific stature of medicine, they expected that medicine could be modeled on the physical sciences and could achieve the same kind of certainty. The new medicine would indeed then be a science, rather than a conjectural art as it had been considered in the past. Physicians, like many other scientists and publicists, were intellectually influenced by Pierre-Simon Laplace, then Europe’s most famous figure in mathematical probability theory.1 After having claimed that the regularity of phenomena, established with rigor, permitted investigation of causes, Laplace also asserted that his theory could be extended to the human and moral sciences. From 1800 on, numbers had begun to play a prominent role in all social activities where control and prediction were sought. The range of phenomena and activities under methodical observation was widened to encompass anything and everything, and particularly, aspects of society such as conscription, demography, suicide and criminality. Overall, the two main features in this endeavor were the great importance given to studying the poor, and the focus on seeking causes (in general terms). In public opinion, physicians were having a lot of trouble solving medical problems. They needed a strategy which • would create a stir in the medical field, • boost their claim to specialized knowledge, • and, in this way, enhance the social authority of their profession. Physicians began to place an emphasis on statistics.2 They thought that this approach to seeking constant causes could eliminate uncertainty in sicknesses. They also felt that a tabular form of representation lent them an appearance of scientific objectivity. Hence, their use served their pretensions to scientificity well. Physicians had to find a cause. Calling on a material and physical explanation of illnesses like phthisis, madness or syphilis which afflicted many and had 1 2

Laplace’s Essai philosophique sur les probabilities was published in 1814 but this work was well-known long before. The work had gone through five editions before the author’s death in 1827. In the 18th century, German intellectuals coined the noun Statistik but the thing had existed under the name of “political arithmetic” in England in the 17th century. By the 19th century, the label from the Germans was adopted.

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no cure, could enable them to convince the French government and public opinion that physicians alone could diagnose and cure diseases. The refusal to consider contagion as a cause of illness led to the search for etiological explanations: heredity was the prime candidate. But it remained to be proven. The development of medical statistics could not have occurred without hospital medicine. 3 Physicians were to take the hospital as their experimental site par excellence. The notion of pathological hereditary predisposition emerges at the point where clinical teaching converged with the use of medical statistics. Clinical observation of numerous cases – the basis of scientific medical knowledge – could lead to certainty only if it was completed by statistics from which doctors could extract a sufficient mass of probabilities. Administered by the State since the Revolution, the hospitals kept careful registers of patient admission and check out. Some practitioners, who had begun to specialize in treating madness, decided to make more use of the registers than just for calculating the cost of inmates. As Baillarger would say: “Asylums have hundreds and hundreds of the insane all in one place. Thus in La Salpêtrière and Bicêtre, there are more than two thousand of the mentally ill. Nothing is easier than to collect numerous observations in a few years.” The moral physicians, as they called themselves, were well placed to collect rich data. And Baillarger added: “Indeed, consumptive, scrofulous and gout-afflicted persons are scattered here and there and a single observer would have much difficulty and spend much time collecting enough data.”4 For the French State, mental illness was the disease of the 19th century. Of the many medical topographies, newspapers and chronicles, the majority of texts that evoke physiological heredity were by physicians who had responsibilities in hospital asylums. These physicians were in the front line in opposing the religious orders who had been in charge of the hospitals and hence the insane before the Revolution. The religious orders were now recovering their former prerogatives, thanks to the monarchy’s return. Here, we will examine only works written by the most influential doctors of the time. Physicians knew little about what caused madness, what its ultimate nature was, or how to cure it, but they still insisted that diagnosis and cure had to be made by members of organized medicine.

2. Hereditarianism and professional legitimacy Within the framework of his project of “scientising” his moral treatment of the insane, Philippe Pinel (1745-1836) set out to use quantitative thinking. On the basis of his research, he concluded in the first edition of his Traité medico-philosophique de l’aliénation mentale (1801), that the “calculus of probabilities” could be used to determine the effectiveness of various therapies by counting the number of times a treatment had a positive effect. Although Pinel alluded to the mid18th century work of the mathematician Daniel Bernoulli, there is little evidence that he meant anything more by the term “calculus of probabilities” than accurate record keeping. It is from

3 4

“La clinique” in the M. Foucault’s words. (En effet tandis que) “les phtisiques, les scrofuleux, les goutteux sont disséminés çà et là (et) un seul observateur ne pourrait qu’avec beaucoup de peine et de temps arriver à rassembler un nombre suffisant d’observations. Les aliénés au contraire sont réunis par centaines dans les hospices, ainsi à la Salpêtrière et à Bicêtre il y a plus de 2000 aliénés et rien de plus facile que de recueillir en quelques années des observations très nombreuses” (Baillarger 1844).

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1807 onward that we can consider Pinel a real user of statistics. Speaking at the Institut de France, he advocated that the use of the “calculus of probabilities will always be an easy and simple application when a distinct knowledge of the number of events, favorable and contrary, has been acquired”.5 In the second edition of this Traité in 1809, he included a section entitled “Résultats d’observations et construction de tables pour servir à déterminer le degré de probabilité de la guérison des aliénés.”6 This section, more than two hundred pages in the new edition, was devoted to commenting on figures characterizing his patients in Bicêtre and La Salpêtrière and the circumstances of their illness. Pinel showed the importance of the patient’s family background and his relationships to other patients. In the study of causes of madness, he defined two kinds: predispositional (or latent) and chance. By starting his discussion of causes with heredity, he drew attention to this as a major predispositional cause, although his figures did not explicitly show it: “It would be difficult not to admit the hereditary transmission of mania when we see everywhere that in certain families, some members are affected by the disease, over several successive generations.” It is interesting to note that, at the same time as Pinel was asserting that medical therapy could attain the character of a true science only “by the application of the calculus of probability”, he abandoned all reference to the “concierges” method of curing.7 In the field of pathological heredity, the most influential voice was Esquirol (1772-1840). He was the first practitioner to specialize in the cure of madness only (Pinel was also a teacher at the École de Médecine). We know that his calculus had served as the premises for the 1838 law on the insane. His role as a statistician was not just an administrative one. Between 1810 and 1820 he made three research trips at his own expense to collect data. Esquirol argued that observation in medicine could be improved only by statistics. And when the Académie de Médicine was created anew in 1820, Esquirol was named a member of its Statistics Commission. It is in the framework of this institution that Esquirol played a part in the debate on numerical method. He spoke up on behalf of his colleague, Pierre Louis,8 saying: What is experience if not observation of facts, often repeated and stored in memory? But sometimes, memory is unreliable. Statistics record and don’t forget. Before a physician puts forward a prognosis, he has done a mental calculus of probability and solved a statistical problem. Notice that he has observed the same symptoms, ten, thirty, a hundred times in the same circumstances and from this, he draws his conclusion. If medicine had paid attention to this tool of progress, it would have acquired a greater number of positive truths. It would not be taxed with being a vague and conjectural science lacking strong principles.9

5 6 7

8 9

Pinel (1807), p. 199. Pinel (1809), pp. 402-403. In the introduction to the first edition, Pinel had explained that his “moral treatment” was drawn from healers’ practice. “Men who are strangers to medical principles and are guided solely by sound judgment or humble folk tradition have dedicated themselves to curing the insane and have brought about the recovery of a great many.” Pierre Louis had found that in a sample of 28 phtisics, 18 had at least one parent who was or had been ill with the same disease. Esquirol (1835), pp. 5.

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For Esquirol, this was why statistics had to lead to an evaluation of the relative importance of the various causes of madness. And the statistical procedures he used would be a demonstration of the “investigation of causes”. An essential premise of Esquirol’s reasoning was the validity of comparative studies of the insane made according to rigorous observations over time, and in which the variables of social context, climate, and same conditions of observation were similar or identical. From such studies, he suggested: “[…] what invaluable results could be had for understanding madness and its causes!” As well, Esquirol warned against using statistics to justify a priori knowledge but he was himself a prisoner of his own convictions. On the basis of a 3-year survey (1811 to 1814) in La Salpêtrière, Esquirol found heredity to be the cause of mental illness in 105 out of 466 cases. This impressive proportion, almost 23 %, did not satisfy him and he made assertions that his figures did not, in fact, support: “I am sure that heredity predisposes to mental illnesses more frequently than my figures show here.” Comparing the figures from La Salpêtrière to those from his private clinic, he concluded that hereditary mental disease was more frequent in rich people than in poor people. His ratios were 1 to 2 for the rich and 1 to 4 for the poor. Esquirol did allow that the proportion for the poor would probably be greater. At La Salpêtrière, poor female patients “often did not even know their parents’ name,” but in the private clinic where the rich were situated, doctors had more background information on the patients’ families. From 1820 on, Esquirol’s figures were repeated in many works. Etienne Georget, used them, for example, in his famous treatise De la folie. Moreover, on several occasions, Esquirol had laid great stress on the large proportion of the “idle rich” (3 out of 4), as he had concluded that mental illness was also caused by ennui. The fact that only rich people were in a position to pay for medical treatment explains why there was such a focus on the number of rich who were mentally ill. It is also why physicians found it important to impress upon them the major risk of hereditary predisposition. The medical profession in Paris, towards the end of the Restoration (1815-1830) had a pyramid shaped organization. At the very top, we find a notorious and inescapable personality, Antoine Portal (1742-1832): •

•

Before the Revolution, he was Professor of Natural History to the Dauphin. Afterwards, he was to hold eminent positions under all the different political systems that France would devise. In 1818, he was First Physician to the King, Louis XVIII, and in 1824, he held the same position under Charles X.

•

And in the 1820’s, he was also the President of Académie de Médecine, a member of the Institut de France, and Professor of Medicine at the Collège de France. This incredible career was due to his work as an anatomo-pathologist and his publications. He wrote more than 40 volumes on a wide variety of medical subjects, including surgery. As he had performed many autopsies without catching any chronic disease, he concluded that such diseases were a matter of heredity rather than of contagion. He thus advocated the heredity of diseases in most of his works, especially in Considérations sur la nature et le traitement des maladies héréditaires ou maladies de familles (1808). In less powerful intellectual spheres, there were other circles where social connections counted heavily in selecting candidates. Esquirol was a strong patron. His circle was very close to and

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became solidly anchored in the state system. On his visits to provincial asylums, he was able to strike up relations with the physicians in charge. They were to reserve positions for Esquirol’s disciples from the 1830s on. Jan Goldstein has compiled a list of nineteen doctors who were members of Esquirol’s circle. In time, writing in the Annales de Médecine Legale et d’Hygiene Publique, founded by Esquirol in 1829, they would become advocates of Parisian moral medicine and work to spread the statistical methodology and conclusion on pathological heredity. As Félix Voisin (1794-1872) was to say: “From every disease afflicting mankind, there is none which is transmitted more easily by heredity than madness. So, starting from this mathematical truth […].”10

Conclusion When Portal died in 1832, a group of physicians, indebted to their patron for their hospital position and convinced that the inheritance of mental illness had been proved by statistics, could take over in defending the theory of inheritance of diseases. The notion of latent hereditary predisposition allowed physicians to judge someone as potentially unwell even if he appeared completely healthy. This induced three significant developments: 1. with their claim to understand the causes of illness, physicians showed the French State that they had an important part to play in setting up the public health administration. 2. In the judicial system, alienists used the notion of pathological heredity to compete with judges in the courtroom, providing another understanding of criminal and destructive behaviors. Their expert knowledge could lead to the exculpation of criminal behavior or more humane methods of handling deviants who were deemed a danger to society. 3. The conviction that illness and vice could “run through generations of marriage” obviously had had implications for ideas about the role of physicians in the management of populations. Thus, doctors stressed their role as counselors to the upper classes on the advisability of particular marriages. They extended their role to the keeping of marriage registers, in the same way as lawyers kept records of property transactions. The growth in heredity studies corresponded to a time of increasing fragility in the relationship between social controls and individual rights. The delicate balance was to be severely affected. The medical profession succeeded in slanting power in favor of social identity over that of the individual. It is thus no surprise that during the July Monarchy (1830-1848) we see physicians entering government “en masse”.11

10 11

Voisin (1826), p. 287. Bajard, Dupuytren, Hahneman, P. Ricord, G.L Bayle, Beclard, Gall, Raspail, Villermé, Trousseau. In contrast, during the Restoration, the elected Chambre des Pairs had only 3 physicians who moreover did not practice medicine; Berthollet and Chaptal were chemists and Porcher-Dupleix, ennobled by Napoleon was practically the ‘lord and master’ in La Chatre (Indre).

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References Aubanel, H. and Thore.(1841). Recherches statistiques sur l’aliénation mentale faites à l’hospice de Bicêtre, Paris: J. Rouvier. Baillarger. 1844. “Recherches statistiques sur l’hérédité de la folie.” Annales médico-psychologiques (mai 1844). Cabanis, Georges. 1956. “Rapports du physique et du moral de l’homme.” In Œuvres philosophiques de Cabanis. 2 volumes. Paris: Presses Universitaires de France, Corpus des Philosophes français. Cadoret, Michèle. 1969. Esquirol et la statistique médicale. Thèse de médecine. Paris. Caffé, P. 1832. Considérations sur l’histoire médicale et statistique du cholera-morbus de Paris. Paris: Tilliard. Charuty, Giordana. 1985. Le Couvent des Fous. Paris: Flammarion. Céard, Jean, ed. 1985. La Folie et le corps. Paris: Presses de l’ENS. Desrozieres, Alain. 1991. “Masses, Individus, Moyennes: la statistique sociale au XIXe siècle.” In Moyenne, milieu, centre: histoire et usages, edited by J. Feldman, G. Lagneau and B. Matalon. Paris: EHESS. 249273. Esquirol, Jean. 1835. “Mémoire historique et statistique sur la Maison Royale de Charenton.” Annales d’Hygiène Publique et de Médecine Légale.Volume 13. Paris: J. B. Bailliere, Esquirol, Jean and Dominique Etienne. 1819. Des Etablissements des Aliénés en France et des moyens d’améliorer le sort de ces infortunés. Mémoire présenté à son Excellence le Ministre de l’Intérieur en septembre 1818. Paris: Imprimerie de Madame Huzard. _____.[1824] 1832. “Mémoire sur cette question: existe-t-il de nos jours un plus grand nombre de fous qu’il n’en existait il y a 40 ans?” Lu dans la séance publique de l’Académie Royale de Médecine, le 23 juillet 1824. Reproduit dans Des maladies mentales…. Volume 2. 723-742. _____. 1832. Des maladies mentales considérées sous les rapports médical, hygiénique et médico-légal. 2 volumes. Paris: Baillière. Falret, Jean-Pierre. 1988. “Observations sur le projet de loi relatif aux aliénés, 1837.” In C. Quétel. La loi de 1838 sur les aliénés. Volume 1, L’élaboration. 93-125. Ferrus, Marie-André. 1834. Des Aliénés. Paris: Imprimerie de Madame Huzard. Fischer, Jean-Louis. 1991. Monstres, Histoire du corps et de ses défauts. Paris: Syros-Alternatives. Fodéré, François-Emmanuel. 1813. Traité de médecine légale et d’hygiène publique. Volume 5. Paris: Mame. _____. 1832. Essai médico-légal. Strasbourg: LF. Le Roux. Foucault, Michel. 1972. Histoire de la Folie à l’âge classique. Paris: Gallimard, Collection Tel. _____. [1963] 1997. Naissance de la clinique. Paris: Presses Universitaire de France, Réédition Quadrige. _____. 1999. Les anormaux. Paris: Gallimard, Le Seuil. Gall, François-Joseph. 1825. Sur l’origine des qualités morales et des facultés intellectuelles de l’homme, et sur les conditions de leur manifestation. Volume 1. Paris: J. B. Baillière. Gauchet, Marcel. 1985. Le désenchantement du monde: une histoire politique de la religion. Paris: Gallimard. _____. 1992. L’Inconscient Cérébral. Paris: Le Seuil. Gauchet, Marcel and Gladys Swain. La Pratique de l’esprit humain, l’Institution asilaire et la Révolution démocratique. Paris: Gallimard, Bibliothèque des sciences humaines. Georget, Etienne-Jean. 1820. De la folie: considérations sur cette maladie. Paris: Crevot. Goldstein, Jan Ellen. [1987] 1997. Consoler et classifier, l’essor de la psychiatrie française. Paris: Institut de Synthélabo, Les Empêcheurs de Penser en rond. Gouveritch, M. 1964. “Esquisse d’une mythologie de la santé et de la maladie.” Encéphale 2: 437-477. Haustgen, Thierry. 1983. Observations et certificats psychiatriques au XIXe siècle. Thèse de médecine. Paris. Havelange, Carl. 1990. Les figures de la guérison (XVIIIe – XIXe siècles) : une histoire sociale et culturelle des professions médicales au pays de Liège. Liège: Bibliothèque de la Faculté de Philosophie et Lettres de l’Université de Liège. Herzlich, Claudine and Janine Pierret. [1984] 1991. Malades d’hier, malades d’aujourd’hui. Paris: Payot. Hildesheimer, Françoise. 1993. Fléaux et société: de la Grande Peste au choléra, XIVe-XIXe siècle. Paris: Hachette. Jacob, François. 1970. La Logique du Vivant. Paris: Gallimard. Louis, Pierre-Charles-Alexandre. [1825] 1843. Recherches sur la phtisie. Paris. Murat, Laure. 2001. La Maison du docteur Blanche. Paris: JC. Lattès. Petit, Antoine. 1817. Essai sur les maladies héréditaires. Paris: Gabon.

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Pinel, Philippe. 1807. “Resultats d’observations et construction des tables pour servir à déterminer le degré de probabilité de fla guérison des aliénés.” Mémoire classe des sciences mathématiques et physique de l’Institut de France 8: 199. _____. 1809.Traité médico-philosophique sur l’aliénation mentale. Paris. _____. 1815. Médecine clinique. 3rd edition. Paris. Piquemal Jacques. 1993. Essais et leçons d’histoire de la médecine et de la biologie. Paris: Presses Universitaires de France. Portal, Antoine. 1809. Observations sur la nature et le traitement de la phtisie pulmonaire. Paris. _____. 1827. Observations sur la nature et le traitement de l’épilepsie. Paris: JB Baillière. Postel, Jacques and Claude Quetel, eds. 1994. Nouvelle Histoire de la Psychiatrie. Dunod. Quétel, Claude. 1988. La loi de 1838 sur les aliénés. Volume 1, L’élaboration. Paris: Frénésie Editions. Régnault, Elias. 1828. Du degré de compétence des médecins dans les questions judiciaires. Paris. Rey, Roselyne. 1985. “L’approche de la folie chez quelques médecins vitalistes du XVIIIe siècle.” In La Folie et le corps. Paris: Presses de l’ENS. 111-140. Semeleigne, René. 1888. Ph. Pinel et son œuvre. Paris: Imprimeries réunies. _____. 1894. Les Grands aliénistes français: Ph. Pinel, Esquirol, Ferrus, Jean-Pierre Falret, F. Voisin, Georget. Volume 1. Paris. Simonnot, Anne-Laure. 1999. Hygiènisme et eugénisme au XXe siècle à travers la psychiatrie française. Paris: Solilan. Swain, Gladys. 1994. Dialogue avec l’Insensé. Paris: Gallimard. _____. [1977] 1997. Le sujet de la folie: naissance de la psychiatrie. Paris: Calmann-Lévy. Voisin, Félix. 1826. Des causes morales et physiques des maladies mentales. Paris: Baillière.

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Poor Old Ancestors: The Popularity of Medical Hereditarianism, 1770-1870 John C. Waller

In 1769 the popular British medical writer William Buchan issued a stern warning to his many thousands of lay readers. “When a disease is contracted and riveted in the habit it is entailed on posterity. What a dreadful inheritance is the gout, scurvy, or the king’s evil, to transmit to our offspring.”1 A few years later he continued on the same theme, “How happy had it been for the heir of many a great estate had he been born a beggar, rather than to inherit his father’s fortunes at the expense of inheriting his diseases!”2 From these short excerpts we can distil three of the primary characteristics of medical hereditarianism in the period stretching, very roughly, from 1770 to 1870. First, hereditary maladies were typically assumed to have their origins in acquired conditions. Second, once a predisposition to a disease became ‘rooted’ in the constitution it was considered to be difficult to remove. Third, so resistant to therapeutics were heritable diseases deemed to be that sexual prophylaxis was regarded as the only reliable means of obviating their spread. Accordingly, those who had children despite haling from scrofulous families were routinely accused of having a callous indifference to the laws of life. When it is remembered that the conditions eighteenth and nineteenth century physicians labelled ‘hereditary’ (in particular, gout, scrofula, consumption and insanity) were prevalent and devastating, one can begin to understand why Buchan and hundreds of others recommended sexual forbearance with such pious conviction. But there is something curious about the conceptualisation of hereditary malady I have just outlined: if new hereditary characteristics could be acquired, as most doctors agreed they could, then why were inherited maladies associated with such dismal prognoses? If the constitution was labile and susceptible to alteration, why would remedies be of no avail? Or, to put it another way, why was the origin of inherited disease judged to be compatible with soft hereditarianism, whilst hereditary maladies, once contracted, were typically described in all-but hard hereditarian terms. In trying to resolve this tension, one thing is immediately apparent. Ideas of hereditary disease drew upon but were seldom derived from empirical data. The pedigree evidence was slight and, as several doctors noted, highly ambiguous.3 Nor can we attribute the genesis of the concept of hereditary disease to the weight of medical tradition; for, as has been stressed on several occasions, the idea of the specific disease diathesis was in part a creation of eighteenth century medical discourse.4 In fact, as I aim to show, medical hereditarianism comprised an assemblage of largely unconscious and unexamined assumptions, sutured together by loose associations and certainly not by a process of logical deduction. Exploring these associations will, I hope, provide the key to understanding the origins of the concept of hereditary disease and explaining why medical hereditarianism achieved such 1 2 3 4

Buchan (1791), p. 9. William Buchan (1809), p. 15. See, for example, White (1784), p.10; and Henning (1815), p. 19. See Carlos López-Beltrán (1992) and López-Beltrán’s contribution to this volume.

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paradigmatic importance in eighteenth and nineteenth century medical discourse. I will begin this analysis by substantiating the claim that heredity and relative fixity were virtual synonyms in medical parlance. Then I will go on to show how this non-obvious association arose, and why it proved so congenial to the medical profession.

To Palliate or to Cure? The association between heredity and incurability was a touchstone for both sides of the eighteenth century debate as to the proper treatment of gout. In 1772 the Bristol-born physician William Cadogan irritated many of his peers by stating that given simple changes in lifestyle noone need suffer the agonising paroxysms of gout. Yet in pressing home this view, Cadogan felt it imperative to refute the idea that gout is heritable; for, he explained, heritability automatically implied that any remedy would be to no avail. Cadogan argued with Puritanical zeal that the condition is caused by improper living, or “the mistaken habits of life.” Talk of diatheses was dangerous, believed Cadogan, because having received a diagnosis of inherited gout the afflicted promptly abandoned any hope of cure and continued to live dangerously indulgent lives. The possibility that gout was both hereditary and curable apparently never occurred to him. “Diseases really hereditary,” he wrote, “I fear are never cured by any art or method whatsoever.” 5 In accusing medical hereditarians of fatalism, Cadogan hardly exaggerated. As early as 1734 the English physician Thomas Bennet was explicitly linking heritability with the inefficacy of medicine. “The Acquir’d Gout may be cur’d,” Bennet wrote, “the Hereditary reliev’d.” Or, as he went on to say, “Hereditary Gout […] may be at least palliated, the Intervals lengthened out, or the Paroxysm render’d less VIOLENT; and the Acquir’d, if taken in Time may be always with Safety, and generally with Certainty, cur’d.”6 This binary opposition between heredity and curability persisted throughout the eighteenth century, and beyond. Sir James Jay, for example, did not deny that “palliation” was possible, but he was adamant that “gout must be looked upon as a hereditary disease,” in consequence of which, “a radical cure is not to be expected.” Few medical hereditarians had much more to recommend to their seriously gouty readers than the virtues of a stoical frame of mind. Only charlatans and quacks, they agreed, would claim that gout can be any more than palliated by the available materia medica.7 To an equal if not greater level, fatalism permeated discussions of the scrofulous maladies. The scrofulous diathesis was considered by most doctors to be the underlying cause of not only scrofula, but also consumption and, in some cases, secondary and syphilis too. Accordingly, all called forth gloomy prognoses. London’s George Henning made the cognitive ties between inheritance and incurability most explicit. “I am now to advert,” Henning wrote in a treatise of 1815, “to an opinion concerning the nature of scrofula, which has long been entertained, and which, if it was well founded, would supersede all reasoning, and render the practice of medicine in it of no avail; namely, that it originates, and wholly depends on a something, which is transmitted through parents to their progeny.”8 The London surgeon Thomas White reckoned 5 6 7

Cadogan (1772), p. 18-19. Having cited evidence for the curability of gout, Cadogan also noted, “this is another strong argument that proves it not hereditary” (p. 7). See also Stevenson (1779), pp. 6-7. Bennet (1734), p. 30. Jay (1772), pp. 43-4; Berdoe (1772), p. 9.

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that medical hereditarians had cost “many thousands of lives” because their pessimism had forced desperate patients into the hands of charlatans dispensing heroic remedies. 9 White demurred from conventional wisdom believing that scrofulous diseases could be controlled through changes in lifestyle and the taking of specifics. As with Cadogan, his therapeutic optimism impelled him to inveigh against the doctrine of hereditary disease.10 Crucially, neither he nor Cadogan ever doubted that heritability implies difficulty of cure. The same assumptions pervade Sir James Clark’s 1835 Treatise on Consumption. Clark explained that standard therapies usually fail to arrest this terrible malady because they are directed at superficial symptoms rather than at the diathetical cause of all scrofulous maladies. For Clark, as for the majority of his fellow physicians, the alleged fact that consumption was “a hereditary disease” readily explained the difficulty they had in treating it. Condemning the panaceas of quack medicine, Clark asserted that the hereditary taint is so difficult to eradicate that, even in ideal conditions and with the most appropriate regimen, it still takes at least a “few generations” to effect a full cure.11 But there were many different forms and degrees of scrofulous disease, some of which carried rosier prognoses than others. It is therefore even more significant that it was only the incurable forms of the malady that tended to be placed within the hereditary category. William Stokes, another influential British writer on consumption, instructed doctors to attempt a cure only in “the absence of the strumous diathesis, or hereditary predisposition.” Where there is evidence of a diathesis, Stokes continued, stick to “palliatives.”12 Richard Payne Cotton, physician to Brompton’s Hospital for Consumption, also noted that “cases arising from hereditary taint [are] more intractable, of shorter duration, and less amenable to remedial agents, than those in which the tuberculous diathesis has, from any cause, been acquired.” 13 James Whitehead, author of a treatise on hereditary diseases, emphatically agreed. “When a disease,” Whitehead wrote, “the elements of which were inherited, has once been developed into palpable form, all efforts at what is termed a radical cure, will probably be unsuccessful.”14 The terms of this debate were recapitulated among early nineteenth century alienists. Despite his firm anti-hereditarian stance on gout, William Cadogan had described “mania” as unquestionably heritable because of its apparent incurability.15 Similarly, those psychiatrists who made a distinction between the heritable and environmental causes of madness were mostly convinced that the more intractable forms of madness involved hereditary taint. Briton’s George Man Burrows stated baldly: “Hereditary insanity protracts.” And, although alleviation is sometimes possible, Burrows added that “relapses and recurrences are more often to be expected.”16 Not all alienists, however, agreed. James Cowles Prichard, for instance, deemed 8 9 10 11 12 13 14 15 16

Henning (1815), p. 49. White (1784), p. 12. Cadogan (1772), p. 14. Cadogan also considered the scrofulous maladies so hard to treat that they had to be heritable (Ibid., p. 17). Clark (1834), p. 63. Stokes (1844), pp. 417-9. Cotton (1858), p. 86. Whitehead (1851), p. 66. Cadogan (1772), p. 19. Burrows (1828), p. 568.

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hereditary madness to be “not less curable than a disease having symptoms of the same description.” But Prichard recognised that he was at fault with general psychiatric opinion. 17 In America, most early Victorian doctors shared Cadogan’s and Burrows’ attitude of qualified pessimism towards cases of allegedly heritable insanity. Presenting an overview of psychiatric thought in 1847, George B. Wood noted, “Inherited insanity is less easily and permanently cured than that arising without such predisposition.”18 William H. Stokes, a Maryland psychiatrist, predicted that “ninety-nine in a hundred can be radically restored” from madness, but he lodged an important caveat: such improvements could not be expected in individuals with “a strong constitutional [i.e. hereditary] tendency to mental disease.”19 These confessions of impotence in the face of hereditary insanity are all the more significant because they coincided with a period of intense optimism in psychiatry. But the conceptual link between heredity and incurability was so deeply entrenched in the logic of medical practice that in the case of inherited madness optimism was often conspicuous by its absence. During the second half of the nineteenth century, there are indications that psychiatric theory became generally more fatalistic. As asylums became overcrowded and filled up with the incurably insane and large numbers of socially marginalized immigrants, it is said that psychiatrists were no longer able to sustain the soaring optimism of the days of Phillipe Pinel and Daniel Tuke. An indicator of this, several historians have argued, was an increase in the invocation of heredity as a causal factor in mental illness.20 Of course, heritability had been identified as a chief predisposing factor in madness since the late eighteenth century and had always been associated with ineradicability. But it does indeed seem that the 1860s witnessed a growing fascination with essentialistic psychology. Dozens of examples, from Britain and America, indicate that Henry Maudsley’s extreme mental hereditarianism of the 1860s and 1870s was far from exceptional. Even Maudsley’s well known eugenic proposals were adumbrated by several prominent medical and psychiatric writers during and after the late 1700s.21 This fatalism is no less perceptible in non-professional discourses. The heritability and persistence of gout was so firmly established that by the late 1700s the gouty diathesis had become a primary cultural motif for notions of tradition and pedigree.22 And the fear of marrying into hereditarily tainted families was well entrenched by the late eighteenth century. 23 There is much evidence to suggest that parents often sedulously examined the relatives of their children’s suitors for evidence of chronic disease; and after titles, cash and connections, the family’s record of health was often the next most serious criterion used when deciding whom their progeny should marry. It was rumoured, for instance, that the poet William Cowper’s hopes of marrying his cousin were vetoed on the grounds that hereditary insanity existed in both lineages. 24 And the theme of ‘prudent’ reproduction is explored in several eighteenth and nineteenth century novels.

17 18 19 20 21 22 23 24

Prichard (1837), p. 122. Wood (1847), p. 759. Cited in Earle (1887), p. 40. See, for example, Dain (1964), especially chapter 4 and Jacyna (1982). See Waller (2001). See Porter and Rousseau (1998) and Macaulay (1834), p. 285. See Percy ([1609] 1930), p. 12 and Gregory (1774), p. 9. King (1986), p. 24.

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In his last work, Sir Charles Grandison (1759), Samuel Richardson explicitly linked heredity and untreatable insanity. Referring to the marital aspirations of two of the principal characters, he narrated: “The Count found an opportunity to let me know his unabated passion for Clementia and that he had lately made overtures to marry her, notwithstanding her malady, having been advised, he said, by proper persons, that as it was not an hereditary, but an accidental disorder, it might be, in time, curable.”25 Unsurprisingly given medical orthodoxy of the time, many children born to insane parents lived in considerable fear of what one novelist described as “the evil destiny” of hereditary madness. So much so, in fact, that Charles Dickens classed inherited insanity as a taboo subject unsuitable for mention in the journal he edited, Household Words. Thus when in 1847 his friend, the crime novelist Wilkie Collins, asked him to publish a short story on the subject of hereditary insanity, he was peremptorily turned away. 26 Claims that insanity is heritable clearly troubled Charles Dickens. A decade before fobbing off Collins, he had written The Pickwick Papers in which he referred to the “well-known medical theory” that “hereditary madness” exists in certain families. Dickens lodged his disapproval of the idea by explaining how one particular madman fell into “morbid insanity” and eventually “raving madness” because he became obsessed with the “strange delusion” that he had inherited the seeds of mania.27 As Dickens saw, the adjective ‘hereditary’ struck terror into those whose madness was so labelled because it suggested that all hope of cure was vain. Dickens, however, was probably being over-protective of his readers, as during the following decades a spate of novels were published, all presenting the same gloomy conception of hereditary insanity. Geraldine Endsor Jewsbury’s 1855 Constance Herbert was based entirely around the theme of hereditary insanity and the moral duties of the afflicted. In this novel, the feckless and irresponsible father of Constance, Charles Herbert, is redeemed only by recognising the folly that led him to reproduce. Bemoaning the fate of his daughter, he sobs: “To know that she is doomed to live under the shadow of madness, and that it is I, her father, who have entailed it upon her; - it is this, this that is the bitter sting in my grief.”28 Most famous of all nineteenth century treatments of hereditary insanity was Mary Elizabeth Braddon’s 1862 crime thriller Lady Audley’s Secret. Lady Audley had many secrets, not least her bigamy, her abandoned child, and the attempted murder of her first husband and his best friend. But there was one secret that first propelled her into the career of lying and deception that resulted in her eventual incarceration in a Belgian asylum: the fact that her mother had “died mad” in a “mad-house” and that, therefore, she too probably carried what the novel’s alienist referred to as the “taint of hereditary insanity.”29 Lady Audley’s Secret, one of the most popular books of the Victorian age, turned on the assumption that heredity implies incurability. To its readers, the description of Lady Audley’s pedigree at the denouement of the book gave them the final piece of information they required in order to understand the reasons for Lady Audley’s career of reckless and murderous deceit.

25 26 27 28 29

Richardson (1753-4), vol. 4, letter 40. See House and Storey (1965), p. 45. Dickens (1839), p.146. Jewsbury (1855), 2: p. 23. Braddon (1862), 3: p. 231.

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Nor were these the only treatments of hereditary insanity in Victorian fiction. Jane Margaret Winnard’s 1854 Lady of Darkness, Holme Lee’s 1862 Gilbert Massenger and Wilkie Collins’ 1859 Mad Monckton all focused on the theme of inherited mania and all characterised it as an irrepressible and uncontrollable morbid taint.30 These authors were equally undivided in declaring the marriage of the hereditarily mad to be morally repugnant. Clearly, to the popular novelist and to the professional psychiatrist, sexual prophylaxis was one of the few guaranteed means of obviating the risk of future lineages being condemned to madness. And this observation brings us back to the seeming paradox highlighted at the beginning of this paper. If it was generally assumed that the heritable material was susceptible to alteration, why did the term ‘hereditary’ so frequently carry connotations of destiny and incurability? In short, how and why did this cluster of associations arise?

Of essence and experience The biological concept of heredity originated from the desire to label, if not explain, the phenomenon of offspring more closely resembling their parents than more distant relations or, for that matter, different species. It was an essentialistic concept invoked to describe the stability and/ or fixity over the course of generations of certain species, racial or familial attributes. Yet a recognition that the extraordinary variety of heritable traits could not have been present in the progenitors of each species pushed natural philosophers into accepting various forms of soft hereditarianism. The hereditary material became plastic, and acquired features and traits heritable. This, however, had significant repercussions for the concept of heredity: its meaning was detached from its anchorage among the Platonic essences and allowed to drift towards the opposite pole. In other words, the belief in the inheritance of acquired characteristics blurred the distinction between what one is born with and what one acquires during a lifetime of experiences. It followed logically from this shift that nothing inscribed in the hereditary fabric had to be considered indelible. Nevertheless, the association between heredity and relative fixity persisted. Indeed, so strong did it become that for centuries, possibly for millennia, the direction of inference ran in both directions: fixity began to imply heredity almost as much as vice versa. The very fact that an attribute seemed resistant to change over an individual’s lifetime time indicated that it was also very likely to be heritable. This confusion, or conflation, of meanings underpins much of the tradition of hereditarian thought and, I believe, helps explain why heredity was so routinely applied to mental characteristics and disease states even when there was no evidence that an individual’s parents were similarly endowed. A representative example is provided in Alexander Walker’s book Intermarriage. This London doctor wrote in 1839 of an anonymous Frenchman who learned to speak English with breathtaking fluency in less than two years. Such a pronounced aptitude seemed inexplicable in terms of learning and environment: something deep-rooted and (therefore) hereditary had to be at work. Sure enough, upon investigation it transpired that Walker’s linguist had an English-speaking grandmother. The Frenchman had never met his

30

Winnard (1855); Collins (1859).

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grandmother but he had inherited a voice box adapted to speaking her native tongue. How else, Walker noted, would he have been able to pronounce ‘Thistlethwaite’? 31 In the same way, when centuries of writers spoke of the “nature” or “disposition” of their characters, referring to the stable elements of personalities that might be refined or modified but not erased, it required only a small cognitive step to make them hereditary as well. One did not need direct evidence of hereditary transmission in such cases, the intractability of the quality in question was usually evidence enough. Conversely, heredity has often been invoked by writers to imply a lack of personal control over deep-seated thoughts, feelings and actions, and especially the inability to suppress sexual desire. The contradiction arising between the Christian morality of self-denial and the more primitive, animalistic instincts that people observed in themselves convinced many that mankind was beyond redemption, and theories of hereditary taint rationalised people’s sense of their desires and drives being beyond their conscious governance. Similarly, poets and playwrights implied that original sin is somehow part of man’s physical fabric, a hereditary entailment against which he is utterly defenceless. It was as a means of emphasising the fixity of traits that Victorian writers so frequently introduced the idea of heritability. George Eliot used the motif of heredity in her 1868 The Spanish Gypsy to signify the irresistibility of her heroine’s tragic fate. Fedalma had been “chosen”, she wrote, “not by any momentary arbitrariness, but as a result of foregoing hereditary conditions.” 32 Novelists, including George Eliot, found the idea of heredity just as useful; for it instantly symbolised to the reader that the traits in question defined the individual and set the boundaries of their potential thoughts and actions. Indeed, it was because of her fascination with the insufficiency of the individual will that Eliot made such extensive use of essentialistic concepts. In Middlemarch, for example, Dorothea’s irrepressible humanitarian zeal was attributed to a “strain” of Puritanical energy that ran through her family and “glowed alike through [all her] faults and virtues.”33 So deeply inlaid in her nature was this zeal that she could in no way resist her urge to improve the conditions of her fellow creatures. Such a basal quality, it seemed to Eliot, had to be traceable to Dorothea’s ancestry. Likewise, when Dickens wished to emphasise Martin Chuzzlewit’s propensity for getting into violent scrapes, he imputed a long tradition of Chuzzlewit pugnacity extending back to “their Great Ancestor beneath the vaults of the Parliament House at Westminster.” In this novel Dickens came close to identifying the underlying rationale of imputing heredity. Seeking to account for the murderous tendencies of Jonas Chuzzlewit, he remarked, “the more extended the ancestry, the greater the amount of violence.”34 True bellicosity, it seemed, had to have a pedigree. Likewise, it was in order to explain the intractably immoral character of Becky Sharp that William Thackeray raised the possibility of hereditary taint in Vanity Fair. Miss Pinkerton, her school mistress, comments on her ward’s pedigree: “My dread is, lest the principles of the mother – who was represented to me as a French Countess, forced to emigrate in the late revolutionary horrors; but who, as I have since found, was a person of the very lowest order and morals – should at any time prove to be hereditary in the unhappy young woman whom I took as an outcast.” 35 The 31 32 33 34

Walker (1839), p.108. Eliot (1868), 3: p. 22. Eliot, (1872), p. 13. Dickens (1844), p. 12.

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misdemeanours that Becky Sharp would go on to commit were so wicked that they required a hereditarian rationale. Of course, heredity has never been applied solely to malign qualities: any marked characteristic has tended to earn the adjective ‘hereditary’. The parents of geniuses, for instance, have been expected to be prodigiously able themselves. And Francis Bacon’s remark “genius continueth not” arose from his surprise that so many sons were actually disappointments to their fathers (though Bacon could hardly have felt let down by Nicholas). Indeed, the failure of some of the greatest minds, from Shakespeare to Cromwell and Pope, to produce talented offspring caused considerable puzzlement. In writing his biography of the famously low-born Michael Faraday, John Tyndall struggled to explain his lack of eminent forbears. How could so pronounced a quality as Faraday’s intellect, he asked, have arisen de novo? “I once used the privilege of my intimacy with Mr. Faraday,” Tyndall wrote, “to ask him whether his parents showed any signs of unusual ability. […] He could remember none. His father, I believe, was a great sufferer during the latter years of his life, and this might have masked whatever intellectual power he possessed.” 36 Because it was hard to imagine anything as distinctive as Faraday’s intellect being other than heritable, nature was emphasised over nurture despite all the evidence pointing in the opposite direction. In 1857 G. H. Lewes chided those who, like Tyndall, ignored counter-examples. 37 But only three years later even Lewes affirmed the heritability of high intellect by listing about a dozen notable cases, such as the Bachs and Tassos, in which talented fathers sired famous sons. 38 Those historians who claim that concepts of heredity pre-Weismann were non-deterministic typically refer to the hereditarianism of Victorian optimists such as Spencer, Robert Chambers and an array of professional and amateur phrenologists. And, to be sure, these figures did stress the capacity for the individual’s heredity to be improved from generation to generation. Nevertheless, one must ask why these prophets of progress seized upon hereditarian concepts with such alacrity. The answer is I think straightforward: If intellectual and moral improvements could be rooted in the hereditary makeup this implied that any gains would be much more permanent than if they rested on emulation and education alone. Hereditarianism appealed to the social reformers precisely because it suggested that progress was an additive process and that achievements were hard to reverse. This is also precisely what the large readership of phrenological works wanted to be told. Their own attainments, they were delighted to find out, would lead to the birth of more intelligent and moral children whose capacity for edification would exceed their own. There would be no risk of affirming the old maxim ‘rags to rags in three generations’, for backsliding was barely even a possibility.

Explaining failure The conceptual suture we have been exploring between fixed traits and heritability provides the key to understanding why certain conditions were labelled heritable. History’s doctors were confronted with a wide range of persistent maladies, like gout, scrofula and consumption that 35 36 37 38

Thackeray (1847-8), p. 90. Tyndall (1868), p. 15. Lewes (1857), p. 384. Lewes (1859).

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failed to respond to anything within the available pharmacopoeia. In attempting to explain the tenacity of chronic diseases, the concept of heredity was an obvious recourse. The Greeks might have said little about the inheritance of specific maladies, but the conceptual framework in which chronic ailments came to be associated with heritability was of extremely ancient provenance. So, just as with Alexander Walker’s anglophone Frenchman, pedigree itself was not in the first instance important. The mere fact that chronic maladies refused to respond to medical care was enough to suggest a corrupted inheritance. But once invoked it was easy to make the label ‘hereditary’ stick. For gout, scrofula and consumption were so common that they could be expected to appear every two or three generations in most families. Even where the pedigree evidence was very weak indeed, the doctor could declare that the family carried a latent taint, or predisposition, that had remained dormant for many generations. In fact, of course, the pedigree evidence was open to multiple interpretations. But the strong tendency to conflate heritability with destiny became a critical factor in sustaining the concept of hereditary disease. There was also another way in which the conflation of heredity with fixity helped forge the concept of hereditary disease. It is vital to realise that physicians almost universally agreed that the locus for chronic diseases like scrofula, consumption, gout and, in some cases, madness, was the inborn constitution. The idea of the individual constitution, or temperament, derived from a Hippocratic-Galenic tradition that remained vibrant during the 1700s and 1800s despite the rise of localism. The concept itself was predicated on the idea that some individual aspects of character, health and appearance are highly resistant to change over time. A person, it was believed, was born and died with the same temperament, whether lymphatic, bilious, sanguineous, phlegmatic or, in some typologies, nervous and athletic. This constitutional arrangement linked together general bodily condition, upon which the individual’s health depended, with such apparently fixed properties as physiognomy, hair and skin colour, the general shape of the body, mental acuity, and the individual’s type and degrees of intellect and emotionality. Seen as core elements of individuality, these traits lent coherence and continuity to an individual’s existence in the midst of a daily life of seemingly bewildering flux and unpredictability. Clearly, that the constitution itself could be significantly altered, for good or for ill, was in large part antithetical to the very idea of a physiological constant. As such, those born with constitutional imbalances, doctors claimed, could only remain healthy through the constant efforts of their doctors to maintain equilibrium. Enduring changes would take many years to accomplish. Considering that the constitution was deemed this hard to alter, it should come as no surprise to learn that it was also generally assumed to be heritable. “That a peculiar temperament distinguishes a nation, no one who will consult history, or look through the world, at the Turks, the Dutch, the Spaniards, can deny,” explained George Bancroft in a lengthy 1829 essay entitled the Doctrine of Temperaments.39 Likewise, the American doctor John B. Lewis noted that the “peculiar condition of organism we denominate temperament, is remarkably subject to continuation in the offspring.”40 Adhering to Ancient medical tradition, most doctors accepted that the constitution was no less heritable than hair colour, eye colour and physical form. Indeed,

39 40

Bancroft (1829), p. 142. Lewis (1866), pp. 87-105, p. 96. See also Anonymous (1833), p. 122.

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those inheriting “fair hair, light blue or light grey eyes, and a fair, soft skin” were for long believed especially likely to be carriers of the scrofulous taint.41 This provides the final key to explaining how the concept of hereditary malady thrived despite the paucity and ambiguities of the available pedigree data. As we have seen, in the shape of gout, consumption, scrofula and insanity doctors confronted on a regular basis a range of serious and apparently incurable conditions. In explaining their resistance to medical therapeutics, one option was to invoke environmental causes alone, such as unhealthy lifestyles capable of producing stubborn build-ups of morbid poisons. But this explanation must have seemed implausible to many doctors because modulating the sufferer’s habits and environment, or subjecting them to courses of bleeding and purging, rarely seemed to have a beneficial effect. This being the case, a rather more persuasive rationalisation was that chronic diseases were caused by forms of constitutional disturbance. Since the constitution was deemed to be largely unchanging, it shared with chronic, hard-to-treat maladies a fundamental property: altering environmental conditions seldom produced lasting change. Diseases like insanity, scrofula and gout, on the one hand, and the individual constitution, on the other, were both characterised by relative permanency. Grafting irremediable illnesses onto the pre-existing notion of the inborn constitution was a means of explaining the hitherto inexplicable. This, however, reinforced the association between chronic illness and inheritance. For making incurable maladies constitutional implied that they were also, at least some of the time, hereditary. It was a view neatly expressed by the New York doctor Daniel Haynes in 1838. Having argued, that “hereditary diseases are never cured,” Haynes justified his belief on the grounds that some illnesses are so firmly fixed within the body’s organisation that they are bequeathed alongside its essential corporeal structures. “I believe it will be generally admitted,” he wrote, “that a disease contracted, not inherited, by what we call a healthy man, may become so confirmed and fixed in his system, as to be by him transmitted to his offspring, and become a hereditary disease.” 42 The primary suture here was between the ideas of incurability and the individual constitution. The connection between heredity and incurable disease was to some extent a consequence of this linkage. Once again, of course, this was an association of ideas underpinned by the conflation of fixity with heritability in Western biological thought. But the concept of the disease diathesis not only had the power to rationalise incurability. Perhaps just as importantly it was also capable of legitimating it. Encountering, over a lifetime’s practice, hundreds of desperate patients with persistent, chronic maladies, few doctors could have relished explaining to them that for reasons they could not fathom, medical science could treat some cases of chronic malady but not others. One means of upholding the dignity of the profession was to insist that diseases like gout, scrofula and phthisis were the result of faulty constitutions over which doctors could hardly be expected to exercise any curative authority. The relative fixity of the individual constitution was an accepted principle of both professional and popular medical thought, so locating chronic disease within the constitution promised to assuage any doubts the public might have had in the competence of the medical profession. Once more, a

41 42

See Phillips (1846), p. 10. Haynes (1838), p. 198.

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bi-product of forming this conceptual linkage was that gout, scrofula, phthisis and insanity inadvertently became hereditary maladies. This explanation was also suggested by a handful of eighteenth and nineteenth century medical writers. Thomas White rhetorically asked why doctors assumed that scrofula was hereditary. “To deem it hereditary,” he explained, “was the best apology that ingenuity could devise” for the “great difficulty of curing those afflicted with it.”43 The poet John Byrom versified the same suspicion: When our distempers did their names receive, (One instance more, good doctors, by your leave), Some chronic matters, such as gout and stone, That would the fare of no arcana own, To save their Credit, these, the learned dons, Cried out, were fix’d hereditary ones: If a man’s father, grand- or great-grand sire; Had the same, ‘twas needless to enquire; Plain was the case, and safe the doctor’s fame; The poor old ancestors bore all the blame.44

George Henning put the case with the greatest tenacity. “It is a fact, on every account worthy of observation” he wrote in 1815: […] that gout and mania, scrofula and phthisis, together with epilepsy are the only diseases […] which are acknowledged to be incurable by the means of medicine, and are the only ones that have acquired the character of being inheritable. A fact that begets some suspicion that the medical world has taken sanctuary under this term hereditary, to shelter themselves from the opprobrium of not having devised remedies for these obstinate maladies. For surely, if it can be rendered plausible, that these infirmities are so intimately blended, by nature, so interwoven, as it were, with our fabric, as to be inextricable form it by any art, we vindicate our profession from censure, although we add nothing to the reputation of it.45

Revealingly, in the literature of the 1700s and 1800s there is barely a single case of an author claiming a chronic malady to be non-hereditary as well as incurable. This is not surprising. For, to have held this unhappy position, would have been to acknowledge the limitations of ones professional competence whilst depriving oneself of any face-saving rationalisation for failure. Cadogan, White and Henning were happy to repudiate the concept of hereditary disease, but only because they shared the conviction that the profession was selling itself short and that chronic diseases frequently “give way to medicament and skilful regimen.” 46

43 44 45 46

White (1784), pp. 15-16. Byrom (1899), p. 70. Henning (1815), p. 58. Cadogan (1772), p. 34.

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“Fix’d hereditary ones” “Fix’d hereditary ones” – this cluster of words from Byrom’s poem crisply conveys the associative relationship that underpinned the idea of hereditary disease. By the late 1700s, heredity had come to be invoked, as little short of a cognitive reflex action, whenever a characteristic seemed unusually stable over long periods. In the case of inherited malady, this false syllogism had a double bind. Within this theoretical framework, because certain maladies were resistant to medical intervention, they could automatically be considered heritable. At the same time, the failure of therapeutics to cure many chronic diseases called forth another essentialistic category, constitutional disease, which because of its association with fixity, in turn also came to imply heritability. Small wonder that it proved extremely hard to dislodge the notion that conditions such as consumption were heritable, even after Robert Koch’s famous bacteriological investigations. Likewise, we can now see why, when psychiatrists assembled clear evidence to show that insane individuals with insane relations were no less treatable than those without, 47 the association between heredity and incurability remained unbroken. Broader inferences may also be drawn from this analysis. An over-emphasis on how doctors, naturalists, physicians and breeders imagined new hereditary variants to arise has, in my opinion, obscured the implicit fatalism of much hereditarian thought prior to the 1880s. Soft hereditarian assumptions may have been routinely invoked to explain the differences among species, races, tribes and families, but inherited mental traits and disease conditions were assumed to be extremely stubborn once engendered. Indeed, according to many physicians, only a distinct physical principle, usually dubbed the ‘law of atavism’ or ‘variation’, could explain why hereditary diseases often seemed to disappear and why mankind was not freighted with a far heavier load of inherited flaws. Finally, once we appreciate this powerful current of fatalism I think it becomes easier to account for both the hard hereditarianism of Charles Darwin and the recurrence of eugenic ideologies at irregular intervals throughout modern European history.

References Anonymous. 1833. Review of Henry Belinaye’s “The Sources of Health and Disease”. American Quarterly Review 14: 120-123. Bancroft, George. 1829. “Doctrine of Temperaments.” American Quarterly Review 5: 118-143. Bennet, Thomas. 1734. An essay on the gout; in which a method is propos’d to relieve the hereditary, and to cure the acquir’d. Particularly, the method of diet, exercise, &c. to be observ’d in this, and most other chronical disorders, is laid down…. London: Richard Ford. Berdoe, Marmaduke. 1772. An Essay on the Nature and Causes of the Gout. Bath: S. Hazard. Buchan, William. 1791. Domestic Medicine: or, a treatise on the prevention and cure of diseases by regimen and simple medicines. London: A. Strahan, T. Cadell and J. Balfour. _____. 1809. Advice to Mothers, on the subject of their own health; and of the means of promoting the health, strength, and beauty of their offspring. Boston: Joseph Bumstead. Braddon, Elizabeth. 1862. Lady Audley’s Secret. 3 volumes. London: Tinsley Bros.

47

See, in particular, Stewart (1864).

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Burrows, George Man. 1828. Commentaries on the Causes, Forms, Symptoms and Treatments, Moral and Medical, of Insanity. London: Thomas and George Underwood. Byrom, John. 1899. “Drink.” In The Poems of John Byrom. Edited by A. W. Ward. Manchester: The Chetham Society. Cadogan, William. 1772. A Dissertation on the Gout, and all chronic diseases, jointly considered, as proceeding from the same causes; what those causes are; and a rational and natural method of cure proposed. London: J. Boyles. Clark, Sir James. 1834. A Treatise on Tubercular Phthisis, or Pulmonary Consumption. Reprinted from the Cyclopaedia of Practical Medicine. London: Marchant. Collins, Wilkie. 1859. “Mad Monkton.” In The Queen of Hearts. London: Hurst and Blackett. Cotton, Richard Payne. 1858. On Consumption: its Nature, Symptoms and Treatment. London: John Churchill. Dain, Norman. 1964. Concepts of Insanity in the United States, 1789-1865. Rutgers University Press. Dickens, Charles. 1839. The Pickwick Papers. London: E. Grattan. _____. 1844. The Life and Adventures of Martin Chuzzlewit. London. Earle, Pliny. 1887. The Curability of Insanity: A Series of Studies. Philadelphia: J. B. Lippincott Co. Eliot, George. 1868. “The Spanish Gypsy.” The Atlantic Monthly 22 (September 1868): 380-384. _____. 1872. Middlemarch. Edinburgh: Blackwood. Gregory, John. 1774. A Father’s Legacy to His Daughters. Dublin: Thomas Ewing and Caleb Jenkin. Haynes, Daniel. 1838. “Address delivered before the Rensselaer county Medical Society … June 3, 1838.” New York State Medical Society 4: 196-204. Henning, George. 1815. A Critical Inquiry into the Pathology of Scrofula, in which the origin of that disease is accounted for on new principles; and a new and much improved method is recommended and explained for the treatment. London: Callow. House, Madeline, and Graham Storey, eds. 1965. The Letters of Charles Dickens. Oxford: Clarendon Press. Jay, Sir James. 1772. Reflections and Observations on the Gout. London: Kearsly. Jacyna, L. S. 1982. “Somatic Theories of Mind and the Interests of Medicine in Britain, 1850-1879.” Medical History 26: 233-258. Jewsbury, Geraldine Endsor. 1855. Constance Herbert. 3 volumes. London. Macaulay, Alexander. 1834. “Gout.” In A Dictionary of Medicine, Designed for Popular Use. Edinburgh: Adam and Charles Black. King, James. 1986. William Cowper: A Biography. Durham: Duke University Press. Lee, Holme. 1855. Gilbert Massenger. London. Lewes, G. H. 1857. “Hereditary Influence.” Westminster Review 16 april 1857. 384. Lewis, John B. 1861. “Hereditary Predisposition.” Annual Dissertation read before the Convention May 22nd 1861. _____. 1859. Physiology of the Common Life. Edinburgh and London. López-Beltrán, Carlos. 1992. Human Heredity 1750-1870: The Construction of a Domain. Unpublished Ph.D. diss., London: King’s College. Percy, Henry. [1609] 1930. Advice to his son, by Henry Percy, ninth earl of Northumberland. Edited with a biographical introduction by G. B. Harrison. London: E. Benn. Phillips, Benjamin. 1846. Scrofula; its Nature, its Causes, its Prevalence, and the Principles of Treatment. London: Bailliere. Porter, Roy, and G. S. Rousseau. 1998. Gout: The Patrician Malady. New Haven: Yale University Press. Prichard, James Cowles. 1837. A Treatise on Insanity and Other Disorders Affecting the Mind. Philadelphia: E. L. Carey and A. Hart. Richardson, Samuel. 1753-4. The History of Sir Charles Grandison: in a series of letters published from the originals. Volume 4. London: Printed for S. Richardson. Letter 40. Stevenson, William. 1779. A Successful Method of Treating the Gout by Blistering: with an introduction, consisting of miscellaneous matter. Bath: R. Cruttwell. Stewart, Hugh Grainger. 1864. “On Hereditary Insanity.” Journal of Mental Science 10: 50-66. Stokes, William. 1844. A Treatise on the Diagnosis and Treatment of Diseases of the Chest. Diseases of the lung and windpipe. Philadelphia: E. Barrington & G. D. Haswell. Thackeray, Arnold. 1847-8. Vanity Fair. London. Tyndall, John. 1868. Faraday as a Discoverer. London: Longmans, Green and Co. Walker, Alexander. 1839. Intermarriage; or the mode in which, and the causes why, beauty, health and intellect result from certain unions, and deformity, disease and insanity from others. London: J. Churchill.

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Waller, John C. 2001. “Heredity, Reproduction and Eugenics: from the early nineteenth century to Francis Galton.” Studies in History and Philosophy of Biological and Biomedical Sciences 32: 20-52. White, Thomas. 1784. A Treatise on Struma or Scrofula, commonly called the king’s evil; in which the impropriety of considering it as an hereditary disease is pointed out; more rational causes are assigned; and a successful method of treatment is recommended. London: J. Murray and R. Turner. Whitehead, James. 1851. On the Transmission, from Parent to Offspring, of some Forms of Disease, and of Morbid Taints and Tendencies. London: J. Churchill. Winnard, J. M. 1854. The House of Raby; or, our Lady of Darkness. 3 volumes. London. Wood, George B. 1847. A treatise on the Practice of Medicine. Philadelphia: Grigg and Elliot.

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Since this workshop is part of an ongoing series of conferences, I will include in my comments the essay by Carlos López-Beltrán “Natural Things and Non-natural things. The Boundaries of the Hereditary in the 18th Century”, that was presented at the preceding conference,1 and that in my view gives important background information for a better understanding and discussion of the essays by Wilson, Waller, and Cartron. 1. López-Beltrán has pointed out a decisive shift in the conceptualisation of heredity in the early 19th century. He argues that after the 1830s, first in French and then in other European languages, one can notice a transition from a typically adjectival use of the term (the hereditary) to a nominal use (hérédité, heredity). According to López-Beltrán, this reification, or “hardening”, of a concept that had been formerly used in a loose metaphorical sense, implied the creation of consensus over the “facts of heredity” as a commonly recognised set of observed regularities, a growing attribution of causal value to genealogical links and the progressive transformation of heredity into a central aspect of physiology and pathology.2 López-Beltrán contrasts this new strength and salience of the concept of “heredity” with the marginality of the “hereditary” in early modern medicine.3 Before the early 19th century, he argues, the hereditary was not a stable domain, an object of debate rather than general consensus, nor was it a key-element of physiological and pathological theories. A set of phenomena called hereditary (resemblance to parents, origin and transmission of monstrosity, etc.) was discussed in the 17th and 18th centuries in relation to the theory of generation. But the transmission of form from generation to generation was seen as due to a constant invariable cause that gave the species its fixity and stability, and had nothing to do with the contingencies of genealogy. For either preformationists or epigenetists, there was a basic common structure for each species (due to a pre-existent germ or to some epigenetic principle) over which the singular, accidental characters of the ancestors had no permanent influence. It was considered impossible that variations due to external influences (such as climate, nutrition, etc.) would be integrated into the lineage and eventually adopted in non-accidental manner. In consequence, López-Beltrán argues, the hereditary was seen as belonging to the domain of the accidental. To use his own words, “there was a deep conceptual hiatus between the explanation of organization (taxonomic similarities) and of accidental individual peculiarities (variations).” According to López-Beltrán, this conceptualisation of the hereditary as accidental explains why from Antiquity to the Renaissance and well into the early modern age, Western medicine paid little attention to the heritability of disease. In the 18th century, innovators such as the “solidists” (who believed that disease was caused by physical or mechanical lesions of the solid body parts – fibers, tissues, organs) did not believe in hereditary diseases, since they did not believe that 1 2 3

López-Beltrán (2002). López-Beltrán (1994). But see the entry “morbi hereditarii” in Lipen (1679), s.v.

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external factors could affect the body irreversibly. In contrast the humoralists, building on the ancient Galenic notion of individual temperament, or constitution, thought it possible that the embryo could be altered by peculiarities of the parental seed (paternal or maternal) due to external influences, and that such alterations could be transmitted through the lineage. How were these alterations produced? López-Beltrán explains at length how the notion of the six “non-naturals”, deeply rooted in humoralism, was used to explain the link between body and environment. The Galenic distinction between the seven “natural things” (elements, temperaments, parts, humours, spirits, faculties, actions) and the six “non-naturals” (air, food and drink, work and rest, sleep and vigil, retentions and excretions, the passions of the soul) allowed the conceptualisation of the boundary and interplay between internal and external determination of bodily features. “Nonnatural” factors (such as diet, exercise, mental and emotional states, etc.) could affect the internal balance of the humours that was the main determinant of individual temperament. The disease (conceptualised in Galenic terms as “praeternatural”, that is a deviation from nature’s customary ways) appeared when something went wrong in the interaction between naturals and nonnaturals. According to López-Beltrán, the concept of heritability of disease was developed in connection with a stronger emphasis on the heritability of temperament. If temperament could be modified by the “non-naturals”, that is by behavioural and environmental factors, this modification could be transmitted across generations as a particular susceptibility to disease. López-Beltrán stresses, however, that this was at best a very soft form of hereditarianism. Within humoralism, what was inherited from the parents was only a small portion of the complex web of influences that acted over each body, and it could in any case be further modified by non-natural factors. The hereditary was the consequence of non-natural factors, and it was not irreversible because they could again affect it. In this framework, hereditarianism was strongly subordinated to environmentalism. More importantly, the hereditary was conceptualised as unstable: there was always the possibility that hereditary traits could be discontinued or even reversed by behavioural and environmental changes in the non-naturals. At the end of the 18 th century there was considerable variety of opinions on this issue. Kant, for instance, believed that variations due to inheritance of traits derived from external influences could disappear in changed circumstances. For Blumenbach instead hereditary variation, including hereditary disposition to disease could become a stable part of temperament, an irreversible “second nature”. So López-Beltrán stresses that within a humoralist framework, the notion of hereditary transmission of disease did not go beyond a “soft hereditarianism”, which went hand in hand with a strong emphasis on behavioural and environmental factors as the crucial determinants of temperament. The notion of hereditary transmission of physical and moral characters did not preclude the possibility of improvement through hygienic measures – a view that French vitalists preserved throughout the 19th century. According to López-Beltrán, a breakthrough occurred only in the first half of the 19th century, when solidists accepted the evidence of hereditary transmission of diseases. At this point a distinction was developed between properly hereditary transmission and congenital influences derived from external factors. Only at this point could “hard hereditarianism” develop. 2. The papers by Wilson, Waller and Cartron provide further evidence of the transition from “soft” to “hard” hereditarianism in English and French medical thought over the 18 th and 19th

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centuries. Wilson describes in detail Erasmus Darwin’s notion of hereditary diseases, which seems to have been typical of the “soft hereditarianism” of 18th century doctors. Like many of his contemporaries, Dr Darwin adopted the view that diseases such as gout, consumption, scrofula, epilepsy, insanity were hereditary. What was inherited however was a “tendency” – the diathesis, a predisposition to disease – that could be effectively stopped by behavioural changes (the nonnaturals). Not only disease in general but even hereditary disease was amenable to treatment through changes in the “non-naturals”. That behaviour was considered crucial in explaining and treating these “hereditary diseases” is shown by the fact that they were also called the “drunken diseases”. Gout, in particular, was explicitly seen as the consequence of “intemperance in drinking much spirituous liquor,” to use Dr Darwin’s own words. In fact Dr Darwin’s views on hereditary diseases provide a very clear example of the primacy of environmental factors over strictly hereditary ones that is typical of 18th century “soft hereditarianism”. The chief, original cause of disease was environmental, as stated in the couplet from Dr Darwin’s Temple of Nature: “The clime unkind, or noxious food instills / To embryon nerves hereditary ills.” This soft, environmental hereditarianism was inherently optimistic. Precisely because the original cause of hereditary disease was environmental, its transmission could be stopped by behavioural and environmental changes brought about by preventive medical measures. Hereditary diseases could be “disinherited” thanks to behaviour properly regulated by medical advice. Waller’s paper describes the development, over the period 1770-1870, of a harder, more pessimistic form of medical hereditarianism. In contrast with the 18 th century optimistic view of hereditary diseases as amenable to cure or prevention, he stresses how 19 th century English and American doctors established a self-evident link between the heritability and incurability of the very same diseases (gout, epilepsy, scrofula, consumption, insanity) that their 18 th century predecessors had considered hereditary and curable. For instance, in marked contrast with Dr Darwin, William Cadogan, already in the 1770s, stressed that the gout should not be considered hereditary, because it was curable by change in behaviour. This implied that, by definition, hereditary diseases could not be cured. Waller provides various examples of this link between heredity and incurability of disease, especially from early and mid-19 th century sources. It seems to me, however, that he tends to over-stress this link, which often seems less strong, on the evidence of his own sources, than he takes it to be. He quotes for instance, as evidence of the presumption of incurability for hereditary diseases, Sir James Clark’s argument, in his Treatise on Consumption (1835), that a full cure of hereditary consumption can be effected over “a few generations”. This seems more an instance of soft hereditarianism: the idea seems to be that the transmission of disease can be interrupted thanks to medical intervention and changes of behaviour. Going over the medical texts quoted by Waller, what strikes me is that the sources that stress more clearly the link of heredity and incurability date typically from the mid 19 th century. This seems to confirm López-Beltrán’s thesis that a “hardening” of the notion of heredity occurred after the 1830s. Waller himself, however, seems to have a different view. He argues that the link of heredity and incurability was firmly in place already by the mid 18 th century, and that “implicit fatalism” characterized European hereditarian thought well before the 19 th century. He argues that “hard hereditarianism” and eugenic ideologies as they developed at the end of the 19 th

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century were no novelties, but represented in fact the resurfacing of a trend that recurred “at irregular intervals throughout modern European history.” Indeed, he argues that “the conceptual framework in which chronic ailments came to be associated with heritability was extremely ancient.” This conceptual framework, he says, was provided by the Hippocratic-Galenic idea of temperament: since temperament was thought to be inheritable, incurable diseases, which were thought to be constitutional, were also seen as hereditary. In other words, Waller seems to think that “hard”, pessimistic hereditarianism is not a 19th century development but a deeply seated feature of Western medical tradition. I find this thesis unconvincing, especially in view of the arguments and evidence presented by López-Beltrán and Wilson. It seems to me that the humoralist notion of temperament provided at most the mental outillage for a soft kind of hereditarianism. It also seems to me that Waller at times misinterprets his sources by reading more hereditarianism into them than they actually contain. For instance, he quotes George Bancroft’s belief that different people are distinguished by a peculiar national temperament. This seems to him clear evidence that temperament was thought to be inheritable. But did this view necessarily imply strong hereditarianism? Commonality of national temperament was just as easily attributable to shared environmental conditions, especially within a humoralist framework. The notion of national temperament and national diseases is not a 19th century development: we find it already in early modern medicine, the most obvious example being the 17th, 18th century medical literature on morbi iudeaorum, the diseases of the Jews. What is stressed in this literature, however, is that such diseases are rooted not so much in hereditary transmission as in the “non-natural” factors of temperament shared by a population, namely work and living conditions, diet etc.4 It seems to me that Waller underestimates the extent to which the humoralist notion of temperament was environmental rather than hereditary, or rather was based on a very blurred distinction between environmental and hereditary factors. Though Waller, in my view, fails to recognize some important differences between 18 th century and 19th century versions of medical hereditarianism, his paper provides much persuasive evidence of the popularity of medical hereditarianism in the 19th century. What explains the popularity of medical hereditarianism in this period? Waller suggests that this popularity had to do with the medical practitioners’ anxiety about their performance. By arguing that diseases like gout, scrofula and consumption were hereditary, doctors could justify their failure at curing them. As the poet John Byrom shrewdly pointed out, by blaming the “ancestors” doctors managed “to save their credit.” This argument is developed in Cartron’s paper. Like López-Beltrán and Wilson, and differently from Waller, Cartron sees medical hereditarianism as a new 19 th century development. In 18th century France heredity had been a physiological rather than a pathological problem, an object of interest to philosophers and naturalists rather than to physicians. But in the first decades 4

See for instance the chapter on the “diseases of the Jews” in Ramazzini (1700), chap. 32. It was argued for instance that the Jews were particularly subject to haemorrhoids as a hereditary condition, but several early modern medical authors questioned this view, arguing instead that the disease derived from external factors such as diet (see for instance Frommann (1677), pp. 147-49, who argues that not the Jews but the English are disproportionately afflicted by this disorder because of the excessive amount of sugar in their diet).

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of the 19th century there is a new emphasis on “hereditary taint, or predisposition to disease”, exemplified by Fodéré’s Traité de Médecine légale (1813). What explains the rise of this new medical hereditarianism? Cartron suggests that the emphasis on heredity supplied a theoretical alternative to the notion of contagion that was considered outdated (and hard to reconcile, as Edwin Ackerknecht has shown, with the policies of economic liberalism). Another element, according to Cartron, was the development of medical statistics that seemed to provide hard evidence of the hereditary nature of some diseases, in particular insanity. It must be noted, however, that the kind of statistical argument used by Esquirol – as quoted by Cartron – to prove that insanity was hereditary, seems hardly an instance of the primacy of numbers. Finding that 105 out of 466 cases of insanity at La Salpêtrière were hereditary, Esquirol proceeded to argue that in fact “heredity predisposes to mental illness much more frequently than my figures show here.” For Esquirol, a theoretical presumption that insanity was hereditary clearly overrode the force of statistical evidence. Cartron argues that another factor of the diffusion of medical hereditarianism in early 19 th century France was the corporate spirit of a medical profession that had emerged newly organized from the crisis of the old regime. The emphasis on heredity helped to legitimize a new and more influential social and political role for the medical men. Since physicians could not cure some diseases, they could justify their failure by arguing that such diseases were hereditary. On the other hand, thanks to their soft hereditanianism, they could claim that the hereditary transmission of disease was preventable thanks to a newly-medicalized social hygiene, including for instance, as Fodéré insisted, pre-nuptial medical advice. Since what was inherited was a predisposition to disease rather than the disease itself, medical preventive treatment could interrupt the hereditary chain. By diagnosing a latent hereditary disposition to disease in some families or sectors of the population, physicians could thus contribute decisively to public health. The medical men’s claim to this new public role was furthered by their intensive participation in liberal politics: Cartron points out that physicians entered the government en masse during the years of the July Monarchy (1830-48). 3. All these papers (with the exception of Waller) present further evidence and arguments supporting the chronology suggested by López-Beltrán. Roughly, the chronology is the following. From Antiquity to the early modern period hereditarianism had a very limited role in European medicine. In the 18th century, in contrast, there are very clear signs that a “soft” but widespread hereditarianism developed in medical circles – a version of hereditarianism based on the humoralist notion of temperament and characterised by a very blurred distinction between environmental, behavioural and hereditary factors. In the mid 19 th century there is another fundamental shift, with the development of a “hard”, or at least harder form of hereditarianism. It would be interesting to compare in detail the soft, 18th century version of hereditarianism with the hard 19th century version. This could be done, for instance, by comparing Erasmus Darwin’s medical hereditarianism with Francis Galton’s eugenics. My impression is that much changed between the two, and that such a comparison would be highly instructive. If this chronology is correct (and it is of course only a rough sketch) some interesting questions emerge. First of all: why was hereditarianism so unimportant in European medical tradition before the 18th century? And why did it become significant in the 18th century? I think that, in

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looking for an answer to these questions, we need to go beyond the intellectual reasons explored in these papers. There is a social element that is very obvious in the sources, though neither Wilson, nor Waller nor Cartron make much of it. A recurrent feature of the medical discourse on heredity in the 18th and early 19th century is the critique of the aristocratic family. Some very significant examples of the link between the emphasis on the heritability of disease and the rejection of aristocratic culture are given by Wilson. Not by chance they concern especially gout, the “patrician malady” par excellence. Wilson quotes William Buchan’s view that it would be better “for the heir of many a great estate” to be “born a beggar, rather than to inherit his father’s fortunes at the expense of inheriting disease.” Erasmus Darwin stressed that “it is often hazardous to marry an heiress, as she is not infrequently the last of a diseased family.” Buchan noted that “once a disease is contracted and riveted into habit,” it became “entailed on posterity.” Entail of course is a word coming directly from the language of legal inheritance, and by the late 18 th century it had acquired a strongly negative connotation as a symbol of the inequities of property devolution within the aristocratic family (such negative connotations are obvious for example in John Galt’s novel, The Entail, 1823). “Entailing” disease on posterity was presented by 18th century and early 19th century doctors as a reprehensible aspect of aristocratic behaviour. The association between the aristocratic family model and hereditary disease was so strong that even John Brown used it to prove that gout was not hereditary, by pointing out that eldest sons succeeded not only to their fathers’ estates and titles but also to their gout, while cadets escaped the disease (which went to show, of course, that gout was caused by behavioural and not by hereditary factors.) This suggests that when we try to understand the origins of medical hereditarianism in the 18 th century we should probably connect it with the crisis of the aristocratic family model – a crisis that was a prominent aspect of European culture in the Enlightenment and Romantic period. In this perspective, we can also understand why medical hereditarianism developed only at this point, though the intellectual framework on which it was based (the Galenic notion of temperament) was extremely ancient. In the late Renaissance and in the 17th century the aristocratic “culture of lineage” was at the peak of its influence over the family structures and strategies of the European elites, as shown by the diffusion of primogeniture. Within the “culture of lineage”, marital choices were firmly dictated by a rigid policy of class endogamy and wealth acquisition, that gave strategic prominence to the marriage of eldest sons and often required the enforced celibacy of cadets and daughters. There was not much space, in this perspective, for health or even fertility considerations in the choice of a marriage partner. Also, what travelled along the generational chain, in this view of the family, was exclusively charged with positive value: ancestors were the fountainhead of family honour, patrimony, glory – it was hardly conceivable to think of them as the source of disease. Even admitting this possibility probably required the crisis and even the breakdown of the aristocratic family model, and more in general the collapse of the cultural hegemony of the aristocracy. By asserting the heritability of disease, physicians expressed a new vision of the family based on bourgeois rather than aristocratic strategies and values. But what about the transition from soft to hard hereditarianism? Cartron’s paper suggests that a class element was involved also in this shift. It is significant that most of the 19 th century doctors’ interest in hereditary diseases shifted from gout, the patrician malady, to insanity – a disease considered to be endemic at the other end of the social ladder, among the “dangerous” classes, the

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urban poor. These classes, rather than the aristocracy, were perceived as a threat to the social order at this point. Furthermore, this new threat was perceived as a much more serious menace to the social fabric. When dealing with aristocratic diseases, physicians had taken an optimistic view of hereditary stock as improvable over the generations, but in dealing with the disease of the urban poor, in contrast, they adopted a much more negative attitude, one that emphasized the social need to prevent the reproduction of people from tainted stock. Erasmus Darwin’s insistence on the need to include health considerations in the choice of marriage partner is a far cry from 19 th century medicalized social hygiene and especially 19th century eugenics. This does not mean that the transition from soft to hard hereditarianism can be wholly explained by social factors. There is one intellectual development that seems to have been obviously related to it. The “hardening” of medical hereditarianism should be understood in connection with the changing notions of nature in this period. Medical thought in the 18 th century was still deeply inspired by a benevolent view of nature and a collaborative model between nature and medical art. Within this view, there was as yet no clear-cut distinction of the natural, on one side, from the conventional, the artificial, the social on the other. As rightly stressed by LópezBeltrán, the natural/non-natural distinction in humoralist medicine was very different from our nature/nurture distinction. In late 18th century medicine, nature and nurture, though increasingly distinct, were not yet seen as mutually exclusive. The boundary between them remained elusive because nature was still understood first of all as a prescriptive, not a descriptive concept. Nature still meant a telos, an intention never fully realized in actuality. Such teleological view of nature was at the core of the co-operative model of nature and art (including medical art): art could improve on nature by seconding and furthering nature’s own goal. Nature therefore did not imply destiny: it was understood as a pliable set of potentialities (like the humoralist notion of temperament), rather than a reality inexorably, unalterably fixed. Like nature, so heredity: both were conceived as malleable, susceptible to improvement or deterioration brought about by behaviour. Medical hereditarianism assumed much stronger form in the 19 th century, when nature turned from a benevolent, purposeful entity to assume the harsh face of physical necessity. The “hardening” of heredity seems to have been part and parcel of the process that led to the hardening of the face of nature.

References Frommann, J.C. 1677. Tractatus singularis de haemorrhoidibus. Norimbergae. Lipen, Martin. 1679. Bibliotheca Realis Medica. Frankfurt. López-Beltrán, Carlos. 1994. “Forging Heredity, from metaphor to cause: a reification story.” Studies in the History and Philosophy of Science 25. _____. 2002. “Natural Things and Non-natural things. The Boundaries of the Hereditary in the 18th Century.” In A Cultural History of Heredity I: 17th and 18th Centuries. Berlin: Max-Planck-Institute for the History of Science Preprint 222. Ramazzini, Bernardino. 1700. De morbis artificum diatriba. Modena.

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Morel’s works can be viewed as an attempt to unify, within the emerging mental medicine, notions and issues belonging to natural sciences, medicine and philosophy. Two fundamental questions are at the center of his work. His first concern is to explore the reasons why an increasing number of individuals suffer from mental disorders and why these disorders remain permanently with these individuals.1 The second concern deals with the impact of mental diseases on social life and its future. These questions are directly linked to his philosophical belief that we live in a world threatened by a series of morbid forces.2 This paper seeks to present Morel’s answers to these questions compiled in two books, which have been among the most influential works of 19 th century French psychiatry. They were written at the time of the French Second Empire, which spans from 1852 to 1870. Born in Vienna in 1809, Morel is placed in a seminary located in the west of France where he spends some ten years.3 Influenced by Félicité de Lammenais’s (1782-1854)4 reformist catholic views, he is destined to become a priest. Instead, he chooses to study medicine and settles in Paris. He arrives there just after the Revolution of 1830, which paved the way to a new constitutional monarchy. These liberal trends spread among university students from all political convictions who supported the new king, Louis-Philippe.5 Morel meets several doctors and intellectuals close to the Catholic reformist movement promoted by Philippe Buchez (1796-1865) who is to become Morel’s great friend. Beside his medical practice, Buchez is considerably active as a journalist and as a free writer. Hostile to the former royal power, he then becomes more and more critical of the new regime, accusing it of promoting exclusively material instincts and the supremacy of money. Morel never got as much involved into philosophical reflections as his friend Buchez did during this period of the July Monarchy but Morel was very much under his influence. 6 Morel gets his medical degree in 1839 at a time when the battle between the physiologists and the psychologists, 1 2 3 4

5 6

“A mesure que j’avançai dans la carrière […] je ne tardai pas à m’apercevoir que la curabilité des affections mentales était un problème de plus en plus difficile à résoudre” (Morel 1857, p. 341). Morel (1857), p. 8. For a more complete biographical information, see Lasègue (1873); Motet ([1874] 1894); Constant (1970). As a priest Lammenais promoted religious freedom, Christianity involved in social matters but he was denied by the pope Grégoire XVI in 1832. He was deputy in 1848 and 1849 after the birth of the Second Republic. Let me precise that there is a ‘conservative’ Lamennais mostly before the 30s – very much for instance in favor of an increasing power for the Pope and in favor of a kind of theocracy and then a liberal after he became very deluded with the Restauration regime in France. The historian Jean-Claude Caron considered that the University took a large part in the Revolution of 1830 (Caron 1991, pp. 295-317). First of all a carbonaro, then close to the Saint-Simonian movement, Buchez attempted to unify the message of the Gospels and the ideas of the French Revolution. After the 1830s, he returned to a more classical catholic faith. Buchez has developed throughout his books a true phisophy of history of great influence on Morel. Buchez’s books used for instance by Morel in his own works later on are: Introduction à la science de l’histoire, ou science du développement de l’humanité (Paris, 1833); Essai d’un traité complet de philosophie au point de vue du catholicisme et du progrés, 3 vol. (Paris, 1840). There is some literature on Buchez; on his specific ties with medical thinking see Isambert (1967).

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which started in the 1820s, was coming to a close. Morel clearly supports the psychologists at the time.7 Morel sets up a practice but fails to attract patients. In these difficult times he has the opportunity to meet the psychiatrist8 Jean-Pierre Falret (1794-1870) who is introduced to him by his friend and former Claude Bernard. Falret is the leading alienist of the 40s in France after the death of Esquirol, the spiritual son of Philippe Pinel’s himself the emblematic figure of French psychiatry, who died in 1840. Morel becomes Falret’s secretary and translator of German texts. Upon Falret’s suggestion, Morel travels throughout Europe visiting asylums. In the first years of the new decade, he visits Italy, the Netherlands, Germany, and Switzerland. From theses places he sends articles and detailed accounts of his tour which are published in the newly founded Annales médico-psychologiques.9 In these writings, he reports upon the state of psychiatry in these countries and offers a thorough knowledge about theorists and practitioners there. He returns to France still without a permanent job, before the Revolution starts in February 1848. Although he does not get involved into politics, he takes advantage of the revolutionary climate to call for the establishment of a social medicine truly orientated to the poors in several articles. 10 In May 1848 his friend Buchez becomes elected president of the National Assembly. Thanks to Buchez’ patronage Morel is granted a job as chief doctor at Mareville asylum (close to Nancy) in 1848 and thus integrates the public national asylum system. He then moves to Rouen (Normandy) in 1856 where he dies in 1873. In Nancy he becomes a local public figure but suffers from jealousy and criticism from the asylum director, who thinks that he is too iconoclast and lacks the constitutive characteristics of a good civil servant.11 Apparently, Rouen asylum atmosphere is more in line with his personal frame of mind. He rapidly becomes a member of the selective local Academy of Sciences and Humanities/Letters and integrates the local bourgeoisie circles. At this point we may note that his rebellious attitude and ‘revolutionary’ calls for reform are somewhat attenuated, all the more so, that the Imperial Regime has honored him with the Legion d’honneur in 1864.

The Treatise on Degeneracy If Rouen is a better place for him, others do not share this bliss. Fast-growing industrialization, terrible working conditions in the predominant garment industry, in that part of Normandy, extended poverty of the lower classes is what he finds in the Rouen vicinity. Like Alexandre ParentDuchâtelet (1790-1836)12, Louis-René Villermé (1782-1863)13 and others from the movement for social observation,14 Morel is concerned with the effects of social pathologies on workers’ physical 7

8 9 10 11 12

Morel (1842). For an exponent of the psychologist trend see Jouffroy (1839), vol. 2. For a good analysis of the clear-cut intellectual atmosphere of the time see Braunstein (1986). For this study, I use the words “materialism”, “spiritualism”, “dualism” with the meanings given by the actors of that time. Though, we have to keep in mind that these words are also categories which need historical and critical investigations and as such have not necessarily to be taken for granted. For my part, I consider Morel as a spiritualist doctor which means for me his refusal to reduce the activity of the mind to mere physiological processes. The expression mainly used at the time was ‘alienist’ rather than ‘psychiatrist’. Annales médico-psychologiques, (1844), vol. 3 and (1846), vol. 6. For instance: Revue nationale (1848), 2 (2). According to Motet (1894), p. 17. Parent-Duchâtelet wrote on public hygiene and on prostitution. See Parent-Duchâtelet ([1831] 1981).

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and moral health. At the beginning of his book entitled Treatise on the Physical, Intellectual, and Moral Degeneracy of the Human Race, Morel writes: Le nombre toujours croissant des suicides, des délits, des crimes contre les propriétés, sinon contre les personnes, la précocité monstrueuse des jeunes criminels, l’abâtardissement de la race, qui dans beaucoup de localités, ne peut plus remplir les anciennes conditions exigées pour le service militaire, sont des faits irréfragables.15

Further down in the introduction Morel reminds that poverty is always linked to moral flaws. Disciplines such as natural sciences, ethnography, geography, anatomy, physiology and history are present in this book. So are the leading scientists from the 18th and the first quarter of the 19th that he quotes at length to the point that the book looks like a catalogue of portraits. Most data contained in the book go back a while and are secondary sources; for instance, Buffon holds a predominant place but he is mentioned through Flourens’ (1794-1867) comments who is an academic, member of the Collège de France, and professor of physiology. 16 But Morel’s book also contains more immediate knowledge, for instance through the influence of Henri Ducrotay de Blainville (1777-1850) whose courses he attended at the Museum of Natural History. 17 If Morel plans to raise the issue on the relations between the physical and the moral aspects of man, his understanding of Cabanis – author of a book precisely on that question 18 – is through the Italian doctor Cerise (1807-1869), another friend of Buchez who considers Cabanis’s treatment of the question much too materialistic.19 Morel, like his friends from the spiritualist movement, is a dualist. A state that is illustrated by his point that “the brain is the organ of the soul”.20 Morel shows some interest in the Austrian Franz Joseph Gall (1758-1828), but not for his invention: phrenology, but for his book on the relationships between harmony and disease that Gall wrote at the end of the 18th century.21 Last but not least, Morel refers several times to Claude Bernard’s experimental physiology. More particularly, Morel uses Bernard’s work to analyze the influence of toxic substance on the degeneration process. Nonetheless, Morel’s text is by no means 13 14 15 16

17

18

19

20

Villermé is the author of Tableau de l’état physique et moral des ouvriers dans les fabriques de coton, de laine et de soie, (1840). He is also known for his books on prisons and on the physical constitution of workers. Coleman (1982). Morel (1857), p. ix. Morel uses Flourens’s book Histoire des travaux et des idées de Buffon. He mentions 1850 as the date of publication. This is actually the book’s second edition. (Paris: Hachette). The first edition was printed in 1844. Morel himself acknowledges this influence in his introduction, but he never mentions any particular de Blainville’s publication in the rest of his treaty. It is thus not easy to evaluate this influence. De Blainville’s interest for the milieu might have had some ‘resonance’ on Morel’s own interpretation on that topic. The first edition was published in 1802. Cabanis (1757-1808) has been a professor at the Paris medical school and has been deputy during the revolutionary period and a senator of the Council of the Five Hundreds after the coup d’état of the 18th Brumaire (1799). His book was written in a materialist tone and Cabanis was considered then as a fervent materialist. Nonetheless, he moderated his own views and he became a member of the Ideologue group. During the 1820s and on, his book was still regarded as one of the terrible example of materialist ideas. He had a large influence on the ‘physiological medicine’ throughout the 19th century. See Génil-Perrin (1910). See also the classical work by Staum (1980). L. Cerise (also written Cerisi) wrote a preface to the re-edition of Cabanis famous book newly published in 1843. Moreover he wrote: Essai sur les principes et les limites de la science des rapports du physique et du moral, (Paris, 1855). See Bourdin (1872). Morel (1857), p. 56.

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a “Bernardian” manifesto. We can note that the majority of scientists present in Morel’s book constitute a homogeneous group representing a science, which is sharply put into question in the mid 19th century.

Teratology, Pathology and Milieu The purpose of the book is to present a theory entitled the theory of degeneracy. 22 The length of the volume is not due to the complexity of the theory; it is the result of the many examples of degeneration chosen by Morel from numerous locations in the world and from every kind of population. Degeneration is, according to Morel, a disease. Thus, the book is build up around the study of the causes of the disease, the description of its symptoms, its evolution and finally the therapy/the cure. His definition of degeneracy is briefly explained in the first pages of the book: degeneracy is a “morbid deviation from the primitive human type”. 23 These deviations are produced by a range of causes and Morel is particularly eclectic since he writes: “[…] the contribution of external circumstances, social institutions, and any other occasional influence […].”24 The deviation process bears some characteristics. It is hereditary and it contributes to the gradual weakening of the reproductive functions of the individual every time it is passed on. These deviations result in the physical and moral modifications on the individuals. The modification becomes apparent through physical stigmata on the body and curious abnormal behaviors. For instance, he writes: […] la petitesse ou la mauvaise conformation de la tête, la prédominance d’un tempérament maladif, des difformités spéciales, des anomalies dans la structure des organes, l’impossibilité de se reproduire; mais aussi à des aberrations les plus étranges dans l’exercice des facultés intellectuelles et des sentiments moraux. 25

The signs of degeneration are mainly physical but not always external. Cerebral lesions could also be interpreted as signs of the degeneration process at work. But Morel seeks to get away from the sole anatomical-pathological model to explain madness. He applies a new definition to the word 21

22

23 24 25

F. J. Gall, Recherches médico-philosophiques sur la nature et sur l’art dans l’état de santé et de maladie chez l’homme, a book also mentioned by Claude Bernard in his Introduction to the study of experimental medicine. The original title is: Philosophisch-medicinische Untersuchungen über Natur und Kunst im kranken und gesunden Zustande des Menschen. Gall came to Paris in 1807 and acquired the French citizenship in 1819. He gave lectures on phrenology and he became quickly successful. But his reputation vanished rapidly; nonetheless phrenology was not completely forgotten after his death. To know more on the French episode of this European movement see Renneville (2000). And on Gall: Lantéri-Laura (1965). There is a growing literature on the subject, not always from an historical approach, though. The oldest analysis – to my knowledge – in French medical literature is Georges Génil-Perrin’s clever and welldocumented thesis, Histoire des origines et de l’évolution de l’idée de dégénérescence en médecine mentale, (Génil-Perrin 1912-13); He also wrote a more specific study entitled “L’idée de dégénérescence dans l’œuvre de Morel”, (Génil-Perrin 1911). On secondary sources, see Friedlander (1973); Dupeu (1976), a text which still deserves to be cautiously read. Martin (1983); Peset (1986); Carlson (1985); Nicasi (1986); Hoochman (1992). And for the cultural dimension of degeneration theory, Daniel Pick’s book remains an excellent and brilliant synthesis (Pick 1989). Morel (1857), p. 5. Ibid. Morel (1857), p. 62.

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‘lesion’: degeneration causes functional anomalies within the body. Thus it is not surprising if a doctor doesn’t find cerebral lesions (in the anatomical sense of the word) among hereditary madmen. Lesions are functional and result from the slow process of the toxic influence of the natural environment and/or individual living conditions. These deviations gradually produce morbid varieties within the species. Pathology has a proximity with teratology in Morel’s treatise. Morel acknowledges the view of biologists that monstrous characteristics occur through the stoppage of physical and mental growth in the individual. But I think that Morel did not completely give up the former conception of teratology. In a certain extent, Morel remains ‘fascinated’ by the monstrous. In that sense it could be argued that he perpetuates the ancient teratology influenced by the demonology.26 Morel’s theory is made up of creationist philosophy and the monogenist conception of the human race. In Morel’s beliefs, the origin of man is God, hence the question why does the human race degenerate? He rejects Rousseau and Condillac’s views and in particular their critics concerning the role of the institutions in the decadence and prefers pointing out “the original degradation of the human nature”.27 But this process is rarely possible without any other cause. And if, for instance, the role of the original sin is fundamental, it is not considered by Morel as unique.28 It needs to be associated with other causes coming from the pathogenic natural and social environment to carry on influence. As I wrote previously, the notion of milieu is important within his treatise, which can be regarded as representative of the environmental medicine trend.29 Neo-hippocratism is largely explicit in Morel’s works.30 As a matter of fact, his books rely on medico-topographical investigations. And one of the origins of his theory comes from the personal investigations he conducted on cretinism some years earlier.31 On the basis of this work, he then moves on to demonstrate that cretinism as a degeneration is caused by the natural environment, more particularly by the mineralogical constitution of the soil. Following the same line of thinking, he attributes a great role to climate32 and in a lesser extent to the quality of air. All these factors together cause what he labels “toxic degeneration”. I would argue that he represents milieu as a fluid. This shows how much he is influenced by the mechanical conception stemming from Newtonian physics. 33 On the other hand, Morel never makes clear if the living has any impact on the milieu. Therefore it is difficult to measure what influence dynamic conception may have had on his interpretation. But the influence of Buffon on his work is clear as Buffon’s interpretation relied likewise on the mechanistic and geographical 26 27 28

29 30

31

Canguilhem (1975), especially pp. 176-80, has written interesting observations on that point. Morel (1857), p. 3. This is of course Morel who gives that interpretation of Rousseau and Condillac’s thinking. A link between original sin and French legitimists of the Restauration government is suggested by Locke (1974), p. 141. Here, there is no evidence of a link between the cultural atmosphere of the Restauration and Morel. He remains closer to the social catholicism of his youth and gives apparently any approval to the catholic traditionalism, predominant at that time. In the tradition initiated by the Société royale de médecine at the end of the 18th century. See for instance Peter (1967). For a general analysis see Jordanova and Porter (1979), especially pp. 119-46. Many medical texts were influenced by former hippocratism. Emile Littré (1801-1881), doctor, former activist of the Revolution of 1830 before becoming a positivist, had previously published Hippocrat’s works. The first volume was issue in 1839. See also Jouanna (1992), especially, p. 512. Morel wrote: “Le crétinisme avait été la première variété dégénérée sur laquelle s’étaient fixées mes recherches”(Morel 1857, p. 357).

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dimension. Former and classical interpretations of the milieu are predominant in Morel’s treaty. He never mentions Auguste Comte’s conception and there is absolutely no evidence that he may rely on ideas developed in the Cours de philosophie positive, for instance.34 Although the milieu is very important, heredity is the focal point of Morel’s interpretation of degeneracy.

Morel and Heredity Morel’s work on the topic of heredity is witness to an increasing interest in the issue of the role of heredity in the development of madness. While alienists are arguing on the question whether the nature of insanity is either a disease of the soul or a cerebral disease, Morel prefers to work on the causes of insanity rather than on its precise nature. Until the 1840s, alienists are convinced that hereditary pathological predispositions are frequent. The psychiatrist Jules Baillarger, one of the directors of the Annales médicopsychologiques declares at the Academy of Medicine in 1844: “[…] c’est d’ailleurs une opinion populaire et très ancienne que celle de l’hérédité de l’aliénation mentale et les relevés statistiques publiés depuis vingt ans ne font que le confirmer.”35 The tone of certainty that he adopts here should not mislead us. Heredity was and remains still very mysterious. As one of his colleagues, the catholic alienist Alexandre Brierre de Boismont, asserted that heredity raises difficulties and problems, rarely solutions. 36 Thus, the 1850s witnessed the emergence of heredity as a focal point of interestand psychiatrists put hereditary mechanisms on their agenda.37 A growing number of psychiatrists were ready to consider heredity as the cause of the cause of insanity. The psychiatrist and republican Prosper Lucas (1808-1885) published a book in two volumes between 1847 and 1850 entitled Philosophical and Physiological Treatise on Natural Heredity. More philosophical than physiological, his book is praised by the Academy of Sciences38 and becomes rapidly a reference among psychiatrists.39 Morel’s Treatise of degeneracy raises directly the question of heredity in the second part of his book although he only dedicates a few pages to the subject.40 Nonetheless, he introduces a new 32

33 34

35 36 37

38 39

Morel (1857), p. 31; his sources come mainly from the English doctor James C. Prichard (1786-1848) who studied at length human races. Morel mentions two of his books in his own treaty: his Natural history of man and his Researches as to the Physical History of Man. Prichard is also know to have coined the expression ‘moral insanity’ which Morel as well as Prichard will then use and consider as a distinct disease. In Morel’s view, people suffering from that type of insanity belong to the large class of degenerates. In his own treaty, Morel also did a review of Prichard’s On the different forms of insanity, in relation to jurisprudence, designed for the use of persons concerned in legal questions regarding unsoundness of mind. See Annales médico-psychologiques (1843), 1: pp. 329-37. On this influence on the notion of milieu, see the masterpiece written by Canguilhem (1952). Auguste Comte discusses about the notion of milieu in the 41st, 42nd and especially 43rd lesson entitled “Considérations philosophiques sur l’étude générale de la vie végétative ou organique” and written according to Comte in 1837. On some influence of Comte’s notion of milieu, see Braustein (1997), pp. 557-71. Baillarger (1844), p. 158. “L’étude de cet important sujet soulève une foule de problèmes dont la solution a présenté jusqu’alors des difficultés insurmontables.” (Boismont 1849, p. 221). See Laure Cartron’s paper and Carlos López-Beltrán’s contributions to the first and to the second workshop, and especially López-Beltrán (1994) and Bénichou (1989). On the specific context of psychiatry Dowbiggin (1991). Comptes-rendus hebdomadaires de l’Académie des sciences, 35, 1853. Bernard Balan (1989), pp. 49-71. His judgment on Lucas is nonetheless too severe according to me.

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interpretation. It is time, he declares, “to give the word heredity a greater meaning than assigned to it ordinarily.”41 Psychiatrists usually consider heredity as a mechanism of transmission of similar characteristics. Morel on the contrary, conceives morbid heredity as the transmission of a predisposition to disease, rather than the transmission of a specific disease entity. 42 Consequently, this conception leads to the creation of a morbid variety which encapsulates very different individuals, though all of them are linked by the hereditary predisposition and /or diseases. The second main characteristic of the hereditary mechanism is its progressiveness. Each new generation received a heavier and more destructive dose of morbidity. This process is nonetheless not infinite throughout history. Sterility indeed intervenes at the fourth generation, subsequently stopping the process of degeneracy. Thus, Nature returns back to its natural order. Harmony is, at the end, stronger than inevitability and I think that this notion of harmony is predominant within the entire works of Morel.43 The hereditary predisposition needs to be connected to a vicious or a pathogenic milieu. This is what he labels the “law of the double fertilization”.44 The best illustration is alcoholism. Individuals drink because of their moral misery and drinking has negative physiological impacts on the body.45 Then, children from this heritage are particularly exposed to degeneration. Organic predisposition and heritage of a vicious milieu will indeed favor the disease. What can be done against this hereditary process? In 1857, Morel declares himself very concerned with the question of the therapeutic program and announces a treaty of regeneration on the subject. But there is no trace of the project. Nonetheless, he dedicates some pages on the issue in his Treatise of degeneracy. The therapeutics he elaborates are made of three major measures: crossbreeding, moral treatment and prophylaxis. He is particularly concerned with crossbreeding and as such fights racial determinism. 46 His belief stems from his monogenist conception.47 Then, he asserts that “the most active means of the regeneration of the species is crossbreeding.”48 But crossbreeding will have a positive impact only if other recommendations are followed.49 For instance, after crossbreeding, moral treatment should be particularly developed to reach a perfect regeneration of the former insane individual. For the rest, he remains fairly vague. In 1860 he declares that hereditary madness is curable 50 upon

40 41 42

43 44 45

46

While he has put a lot of footnotes throughout his book, he mentions very few writers on the topic of heredity. For instance there is no reference to medical texts except Lucas’s treaty. Morel (1857), p. 565. “Nous n’entendons pas exclusivement par hérédité la maladie même des parents transmise à l’enfant. […] Nous comprenons sous le mot hérédité la transmission des dispositions organiques des parents aux enfants” (Morel 1857, p. 565). We know that the idea of ‘une marche de la nature’ is spread over the life sciences throughout the 19th century. See for instance Conry (1980). Morel (1857), p. 567. The link between alcoholism and degeneration is promised to a long life throughout centuries. See Jacques Borel, Du concept de dégénérescence à la notion d’alcoolisme dans la médecine contemporaine, (Montpellier, 1968). Also: W.F. Bynum, “Alcoholism and Degeneration in 19th Century European Medicine and Psychiatry”, British Journal of Addiction, 79, 1984. For instance, he rejects Gobineau’s ideas on race. (Précis analytique des travaux de l’Académie des sciences, belles-lettres et arts de Rouen, 60, 1857-58, p. 152). Man of letters and diplomat, Arthur de Gobineau (1816-1882) became famous after his essay entitled Essai sur l’inégalité des races, published between 1853 and 1855. He asserted that the decay of the western civilization was caused by the lack of purity of the white race. Crossbreeding was then regarded as the most irrelevant thing to do.

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the expressed conditions that prophylaxis and good hygiene are respected. What seems clear in his mind is not expressed from a practical point of view. Moral life may be what is lying under this rhetoric.

The Treatise on Mental Disorders After the publication of the treatise on degeneracy Morel carries on explaining why degeneration should be important to psychiatrists.51 In the treatise he writes: “insanity is predominantly a degeneration”52, and this is what he will seek to demonstrate in a new book entitled Treatise on Mental Disorders in 1860. The book is a manual for psychiatrists but also for general practitioners. He introduces among other novelties a new nosology. Within his classification based on the aetiological dimension of the disease, he labels a new medical entity: folie héréditaire, hereditary madness. Individuals which belong to this entity are divided in four groups. This division is based on the graveness (seriousness) of the disease, that is to say, in Morel’s view, on the degree of hereditary intensity. Indeed, theses groups are organized along the same structure previously used by Morel. Consequently, the first group gathers individuals with nervous temperament. In the second one, we find alcoholics, individuals suffering from aberration of the moral sense, and moral insanity. In the third group, we find the same individuals but with a more pronounced taint; as well as eccentric, disorderly and dangerous individuals because of their total lack of sense of Good and Bad. In the last group are gathered cretins and idiots. We can note that, in a way, hereditary madness is close to degeneration. There seems to be the idea of a vital force in this morbid heredity. The level of intensity of heredity determines the intensity of mental disorders. How are these ideas received? Following the publication of Morel’s last treatise, the Société médico-psychologique53 (SMP) opens the debate on the classification of mental diseases and discusses the treatise. Buchez has already introduced Morel’s books to their colleagues of the SMP when the debate on classification begins. In his introduction, Buchez is as enthusiastic as he had been when he explained the Treatise of degeneracy to his fellows some three years ago, (always in the SMP). Nevertheless, Buchez’s colleagues are not entirely convinced by Morel’s book and Buchez’s argumentation.54 A majority of the members praises his ambition to suggest another 47

48 49

50 51 52 53 54

Morel (1857), p. 349: “L’homme est un, l’espèce est une. Il ne peut y avoir, pas plus entre les races humaines qu’entre les variétés maladives de ces races, de distances infranchissables telles qu’il en existe entre les espèces et les règnes que renferme la nature.” At that time, Armand de Quatrefages, professor of anthropology at the Museum of Natural History and member of the Academy of Sciences strongly supported the unity of the human species. (“Du croisement des races humaines”, Revue des DeuxMondes, 1-3-1857.) For more information on the scientific and intellectual debate on monogenism and polygenism in France, see Blanckaert (1996), 2: pp. 3021-37. Morel (1857), p. 524. He previously reported negative examples of mixed races in others pages of his book. See Morel (1857), p. 413. Thus crossbreeding could provide opposite results: breeding in (what he labels “remonter vers le type supérieur” – Morel 1857, p. 517 ) or breeding out. Morel (1860), p. 646. Morel (1860), p. iii. Morel (1857), p. 345. This association has been founded in 1852 and was opened to non-practitioners. It was at the time the only structure which gathered the profession of médecins-aliénistes. The first paper read during the days dedicated to the debate was by Louis Delasiauve – a colleague appreciated by Morel – who summarized most of the critics pronounced later on.

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methodology for classification. Still, they think that an aetiological classification is unrealistic given the state of their knowledge. The sharpest criticism concerns the entity of the hereditary madness. Psychiatrists do not understand how Morel can assert an entity strictly based on heredity since heredity concerns different types of mental disorders. What could be the criteria to define such a category? Morel argues that this new category gathers specific individuals who suffer special forms of delirium, a mix of calm and of furor etc.55 Since the nature of the pathological process is specific, madness of that origin is specific as well. Morel is unsuccessful in convincing the majority of his colleagues. The difficulty to trace a borderline between hereditary madness and other insanities where heredity plays a role is seen as too subtle. Besides, some members of the SMP have the feeling that this interpretation may lead to the assimilation of heredity and pathology. As a conclusion, we can say that the members of the SMP in 1860 do not endorse this new medical entity.56 In the years following the debate, Morel keeps on publishing on the subject of heredity. As a tribute to Flourens, he publishes a book on different case studies on toxic degeneration. 57 He also publishes a series of new articles on hereditary mechanisms and the process of degeneration. In these, he does not change his views but tries to be more specific and to attenuate some of their stronger aspects. For instance he points out again that the transmission of dissimilar characters is the rule in the mechanisms of the heredity of mental pathologies, but that the rule is not exclusive.58 Nevertheless, he remains adamant on the specificity of heredity of mental disorders and never gives up the notion of hereditary madness. He just tends to moderate the fatalistic trend that some of his colleagues see in his general framework.59

Conclusion Milieu and the original sin cause abnormality; heredity maintains and accelerates the abnormality and there is no cure because this is the tragedy of the human race. That is what Morel tells us. This is not to be discarded as merely simplistic. Morel is indeed of great importance in the development of mental medicine within the more general domain of medical and biological sciences. To a certain extent, his own intellectual evolution from spiritualist psychology to physiological pathology could be seen as an illustration of the evolution of psychiatry itself. 60 It may be said that he provides a wonderful model to explain a bizarre process. In a way, he gives a solution to the question of insanity and therefore opens up a new path for the development of mental medicine. His Treatise of degeneracy reveals a large influence from vitalism and from teleology. But in his attempt to put forward a solution, he does not restrain himself to these philosophies and mixes diverse intellectual influences on the nature of insanity. In doing so, he 55 56 57 58

59 60

Morel (1860), p. 252. When he wrote his review of Morel’s book, Buchez himself hesitated between his friendship and his doubt about Morel’s attempts. Buchez (1860). Morel (1864a). Morel (1867). This aspect remains very successful among psychiatrists after Morel; it is of great importance, according to me, to understand the intellectual orientations of French psychiatry. See Dupeu (1976). And also Gilman (1985). Morel (1864a), p. 17. As one of his colleagues declared: “[…] quoique spiritualiste, je n’ai pas peur de la matière;” (Société médico-psychologique 1860, p. 325).

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puts forward an interpretation, which can satisfy those who defend the model of brain lesions and those belonging to the psychologist trend (which was kept alive mainly because of the weakness of the opposite interpretation). But his role has not only been to unify the different interpretations of insanity of his time. Also, he has suggested that mental pathology is governed by its own specific rules. And the hereditary mechanism as viewed by Morel contributed to a new interpretation of the pathological. Far from being determined by a link of degree with the normal, the pathological becomes determined by a difference of nature. Therefore there is no possibility to go back to normality and this is why hereditary madness cannot be curable. Although hereditary madness is central to Morel’s view, it cannot be argued that Morel is a mere exponent of biological psychiatry. Heredity and milieu are not opposed in Morel’s framework. On the contrary they can be considered as very much intertwined, linked into solidarity. In a way, Morel prolongs and extends the role of Milieu as a cause menacing the harmony of the body. Because for the Fin de siècle psychiatrists, hereditary madness means organic causality, Morel’s interpretation of heredity has been traditionally included in the positive age of psychiatry61 and praised by a new generation of psychiatrists convinced that insanity is a pure organic disease. Also, they have seen in Morel a precursor of organic psychiatry and evolutionism. But the interpretation of hereditary mechanisms as shaped by Morel and praised by French psychiatrists after his death, may indeed have not favored the coming together of psychiatry and positive biology. It can be argued on the contrary that it has reinforced the specificity of psychiatry, or at least the particularity of the exponents of the notion of hereditary madness. They primarily defined a pathology of heredity rather than explaining in what sense insanity could be considered a hereditary disease. The exponents of the physiological orientation of mental medicine support an interpretation of hereditary mechanisms, which is not particularly influenced by Claude Bernard’s experimental medicine principles. This is a kind of paradox. But after all, degeneration has been largely praised during the 19th century, which is also the century of evolutionary ideas and progress.

References Baillarger Jules. 1844. “Recherches statistiques sur l’hérédité.” Annales médico-psychologiques 3: 328-59. Bénichou, Claude, ed. 1989. L’ordre des caractères: aspects de l’hérédité dans l’histoire des sciences de l’homme. Paris: Vrin. Blanckaert, Claude. 1996. “Monogénisme et polygénisme.” In Dictionnaire du darwinisme et de l’évolution, edited by P. Tort. Volume 2. Paris: Presses Universitaires de France. 3021-37. Boismont, Alexandre Brierre de. 1849. “Analyse de l’ouvrage de Prosper Lucas, Traité philosophique et physiologique de l’hérédité naturelle dans les états de santé et de maladie du système nerveux.” Annales d’hygiène publique et de médecine légale 42: 221-39. _____. 1874. “Morel. Fragments de son œuvre en aliénation mentale, l’hérédité morbide, les dégénérescences.” L’Union médicale 81 (july): 25-35. 61

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Bourdin, Claude-Etienne. 1872. Etudes médico-psychologiques: Cerise, sa vie, ses œuvres. Paris. Braunstein Jean-François. 1986. Broussais et le matérialisme. Médecine et philosophie au XIXe siècle. Paris: Méridiens-Klincksieck. _____. 1997. “Le concept de milieu, de Lamarck à Comte et aux positivismes.” In Jean-Baptiste Lamarck, edited by G. Laurent. Paris: CHTS. Buchez, Philippe. 1838. Introduction à l’étude des sciences médicales. Paris: Eveillard. _____. 1838. Introduction à la science de l’histoire, ou science du développement de l’humanité. Paris: Paulin. _____. 1838-40. Essai d’un traité complet de philosophie au point de vue du catholicisme et du progrès. 3 volumes. Paris: Eveillard. _____. 1857. “Rapport sur le Traité des dégénérescences.” Annales médico-psychologiques, 3rd ser. 3: 456-67. _____. 1860. “Rapport sur le Traité des maladies mentales de Morel.” Annales médico-psychologiques, 3rd ser. 6: 613-30. Canguilhem, Georges. 1975. La connaissance de la vie. 2nd edition. Paris: Vrin. Carlson, Eric T. 1985. “Medicine and Degeneration: Theory and Praxis.” In Degeneration, the Dark Side of Progress, edited by J. Chamberlin and S.L. Gilman. New York: Columbia University Press.120-140. Caron, Jean-Claude. 1991. Générations romantiques. 295-317. Paris: A. Colin Cerisé, Laurent. 1872. Oeuvres (publiés par la famille et les amis). 2 volumes. Paris: Masson. Coleman, W. 1982. Death is a social disease: public health and political economy in early industrial France. Madison: University of Wisconsin Press. Comte, Auguste. 1975. Cours de philosophie positive. Philosophie première: Leçons de 1 à 45. Introduced and annotated by M. Serres, F. Dagognet, and A. Sinaceur. Paris: Hermann. Conry, Yvette. 1980. “L’idée d’une ‘marche de la nature’ dans la biologie pré-darwinienne.” Revue d’histoire des sciences 33 (2): 97-149. Constant, Françoise. 1970. Introduction à la vie et à l’œuvre de Bénédict-Augustin Morel. Ph.D. diss., ParisCochin: Faculty of medicine. Dowbiggin, Ian. 1991. Inheriting Madness. Professionalization and Psychiatric Knowledge in Nineteeth Century France. Berkeley: California University Press. Dupeu, Jean-Marc. 1976. La dégénérescence. Figure et doctrine de l’aliénation. 2 volumes. CES Psychiatrie. Paris: Faculty of medicine. _____. 1976. La question de l’hérédité dissimilaire dans la pathologie mentale, Ph.D. diss., Montpellier: Faculty of medicine. Flourens, Pierre. 1844. Buffon, histoire de ses travaux et de ses idées. Paris: Paulin. _____. 1864. Examen d’un livre de M. Darwin sur l’origine des espèces. Paris: Garnier. Friedlander, Ruth. 1973. B.A. Morel and the Theory of Degenerescence: the Introduction of Anthropology into Psychiatry. Ph.D. diss., San Francisco: University of California. Génil-Perrin, Georges. 1910. “L’œuvre de Cabanis dans la psychiatrie.” Revue de psychiatrie (oct. 1910). _____. 1911. “L’idée de dégénérescence dans l’œuvre de Morel.” Revue de psychiatrie 15: 134-56. _____. 1912. Histoire des origines et de l’évolution de l’idée de dégénérescence en médecine mentale. Ph.D. diss., Paris: Faculty of Medicine. Gilman, Sandor L. 1985. Difference and Pathology: Stereotypes of Sexuality, Race and Madness. Ithaca, NY: Cornell University Press.Isambert, François-André. 1967. Politique, religion et science de l’homme chez Buchez (1796-1869). Ph.D. diss., Paris: University of Sorbonne. Hoochman, Jacques. 1992. “La théorie de la dégénérescence de B.A. Morel, ses origines et son évolution.” In Darwinisme et société, edited by P. Tort. Paris: Presses Universitaires de France. 402-412. Jordanova, Ludmila and Roy Porter, eds. 1979. Images of the earth. Essays in the History of the Environmental Sciences. London: The British Society for the History of Science. Jouanna, Jacques. 1992. Hippocrate. Paris: Fayard. Jouffroy, Theordore. 1839. “De la légitimité de la distinction de la psychologie et de la physiologie.” Mémoires de l’Académie des sciences morales et politiques. Section de philosophie. Volume 2. 1-42. Lantéri-Laura, Georges. 1965. Histoire de la phrénologie. Paris: Presses Universitaires de France. Lasègue, Charles. 1873. “ Morel, sa vie, et ses œuvres.” Archives générales de médecine. Volume 1. 588-600. Locke, Robert. 1974. French Legitimists and the Politics of Moral Order in the early Third Republic. Princeton: Princeton University Press. López-Beltrán, Carlos. 1994. “Forging heredity: From Metaphor to Cause: a Reification Story.” Studies in History and Philosophy of Sciences 25 (2): 211-35. Lucas, Prosper. 1847-1850. Traité physiologique et pratique de l’hérédité naturelle dans les états de santé et de maladie du système nerveux, avec l’application méthodique des lois de la procréation au traitement général des affections dont elle est le principe. 2 volumes. Paris: Baillière.

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Martin, Claude. 1983. La dégénérescence dans l’œuvre de B.A. Morel et dans sa postérité. 2 volumes. Thèse de 3e cycle. Paris: EHESS. Moreau de Tours, Joseph. 1852. “De la prédisposition héréditaire aux affections cérébrales.” L’Union médicale 6 (8): 195-6. _____. 1859. La psychologie morbide dans ses rapports avec la philosophie de l’histoire ou de l’influence des névropathies sur le dynamisme intellectuel. Paris: Masson. Morel, Bénédict-Augustin. 1842. “Mémoire sur la manie des femmes en couche.” Bulletin de la Société médico-pratique de Paris January: 222-235, April: 550-73. _____. 1848. “La médecine sociale dans ses rapports avec l’avenir de l’amélioration sociale.” Revue nationale 2 (2). _____. 1855. Influence de la constitution géologique du sol sur la production, lettres de Mgr Billiet,... réponses de M. le Dr Morel. Paris: Masson. _____. 1857. Traité des dégénérescences physiques, intellectuelles et morales de l’espèce humain et des causes qui produisent ces variétés maladives. Paris: Baillière. _____. 1859. “Des caractères de l’hérédité dans les maladies nerveuses.” Archives générales de médecine 14: 257-81. _____. 1860. Traité des maladies mentales. Paris: Masson. _____. 1864a. De la formation du type dans les variétés dégénérées ou Nouveaux éléments d’anthropologie morbide pour faire suite à la théorie des dégénérescences dans l’espèce humaine. Paris: Baillière. _____. 1864b. De la folie héréditaire. Paris: Masson. _____. 1864c. Du goitre et du crétinisme. Etiologie, prophylaxie, traitement, programme médico-administratif. Paris: Asselin. _____. 1866. Traité de médecine légale. Paris: Masson. _____. 1867. “De l’hérédité morbide progressive ou des types dissemblables et disparates dans la famille.” Archives générales de médecine, 6th ser. 9: 385-401 and 564-96. _____. 1868. Analogies entre les dégénérescences intellectuelles, physiques et morales des habitants des contrées paludéennes et des habitants des contrées goitrigènes. Paris: Asselin. Motet, Auguste. 1894. Notices biographiques. Paris: Baillière. _____. [1874] 1894. “Eloge de Morel.” Annales médico-psychologiques, 5th ser. 12. Reprinted in Notices biographiques. Paris: Baillière. Nicasi, Stefania. 1986. “Uomini e animali nella teoria della degenerazione.” Intersezioni 6(3): 557-75. Parent-Duchâtelet, Alexandre-Jean-Baptiste. [1790-1836] 1981. La prostitution à Paris au XIXe siècle. Introduced and annotated by Alain Corbin. Paris: Le Seuil, coll. L’Univers historique. Peset, José-Luis. 1986. “Del ‘angel caìdo’ al enfermo mental: sobre el concepto de degeneracion en las obras de Morel y Magnan.” Asclepio 38: 215-240. Peter, Jean-Pierre. 1967. “Une enquête de la Société Royale de Médecine (1774-1794).” Annales ESC XXII. Pick, Daniel. 1989. Faces of Degeneration: an European Disorder 1848-1918. Cambridge: Cambridge University Press. Piorry, Pierre-Adolphe. 1840. De l’hérédité dans les maladies. Paris: Bury & Baillière. Quatrefages, Armand de. 1857. “Du croisement des races humaines.” Revue des Deux-Mondes 1 march 1857, 159-88. Renneville, Marc. 2000. Le langage des crânes: une histoire de la phrénologie. Paris: Les Empêcheurs de pesner en rond. Ribot, Théodule. 1873. L’hérédité. Etude psychologique sur ses phénomènes, ses lois, ses causes, ses conséquences. Paris: Librairie philosophique de Ladrange. Société médico-psychologique. 1861. “Classification des maladies mentales.” Annales médicopsychologiques, 3rd ser. 7: 128-43, 145-77, 316-32, 456-71, 642-57. Staum, Martin S. 1980. Cabanis: Enlightenment and Medical Philosophy in the French Revolution. Princeton: Princeton University Press.

164

George Combe’s law of hereditary descent John van Wyhe

In this paper I would like to make two main points. The first is historical and the second theoretical or methodological. The historical point is that the phrenologist and philosopher George Combe played a disproportionately major role in popularizing some forms of heredity and proto-eugenic concerns in the mid-nineteenth century. The second or methodological point I would like to make is that the heredity of the ideas of inheritance themselves is as important as their context in determining their specific characteristics. George Combe was a young Edinburgh lawyer when the German phrenologist Dr Johann Gaspar Spurzheim came to Edinburgh to face down an acrimonious critic in 1816. Through skilful dissections and naturalistic rhetoric Spurzheim was widely seen to have defeated his opponent and Spurzheim’s reputation, for some, quickly swung from that of an outrageous foreign quack to a serious man of science to be respected. Combe was among those Spurzheim converted to the new science of phrenology during a seven month stay in Edinburgh. Spurzheim was at that time the only advocate in Britain of the system founded by Dr Franz Joseph Gall formerly of Vienna and later residing in Paris. Gall called his system Die Schädellehre and later in Britain it was named phrenology. Gall recognized that the brain was the organ of mind and furthermore he argued that the mind must be composed of multiple distinct innate faculties. Therefore, each faculty must have a distinct seat or ‘organ’ in the brain. The size of an organ, other things being equal, must be a measure of its power. Hence, as the skull follows the shape of the underlying brain, the exterior of the head serves as an indicator of the shape of the brain underneath and reveals an accurate index of psychological aptitudes and tendencies. 1 George Combe began by purchasing some plaster busts to study the science and these soon attracted friends and visitors who looked to Combe to explain the details. This was Combe’s first real taste of speaking authoritatively on a scientific subject and he was soon addicted. Over the next few years Combe and his fellow Spurzheim acolytes created the British hybrid doctrine we now recognize as phrenology with its white plaster busts with black markings and the lists of 35 or so renamed cerebral organs with their corresponding mental faculties arranged into orders and genera. Combined with these was a version of the traditional doctrine of the four humours or temperaments – as the phrenologists termed them the lymphatic, bilious, sanguine and the nervous temperament. Combing estimations of temperament and the shape of the skull, phrenologists diagnosed character, aptitudes and mental powers and inclinations. The authority to classify and define the powers and abilities of all humans was a uniquely authoritative role for a phrenologist and it does not seem coincidental that all of the major phrenologists were ambitious, egotistical and often quite arrogant men. George Combe was perhaps the most ambitious of them all. He first set his sights on becoming a man of science. This he to some measure achieved by co-founding the Phrenological Society in 1820, by writing scientific publications on the subject – his first publications – and by giving public 1

van Wyhe (2002).

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lectures. For Combe the direction in which phrenology was to be taken next was to continue to emphasise and develop the already marked naturalistic themes, which had become part of phrenology. For example, phrenologists used an appeal to Nature (usually with a capital N) to defend themselves from all matter of criticism of their science. Phrenology must be true, phrenologists argued, because Nature showed it to be true. This sort of tactic, reiterated and relied on throughout the 1820s as it was, became a constant accompaniment and eventually a typical aspect of phrenology. In the face of critics who quipped: “Fool and Phrenologist are terms nearly synonymous,” phrenologists replied that: “phrenological […] is another word for natural,” or “whatever is natural is just to the same extent and in the same degree phrenological.” 2 Combe was deeply impressed by the natural philosophical and phrenological works of his mentor, including Spurzheim’s A view of the philosophical principles of phrenology (1825) and his Philosophical catechism of the natural laws of man (c1824). Combe wrote to Spurzheim: “[Your book the Philosophical principles of phrenology] has afforded me more delight than I ever received from any book on any subject whatever.”3 Spurzheim, then living in Paris, was deeply influenced by Enlightenment writers like Baron d’Holbach, Constantin François de Volney and Pierre-JeanGeorges Cabanis. Spurzheim reiterated many of their Enlightenment themes such as the progress of knowledge through naturalism, the supreme reign of natural laws, anti-clericalism and also a better morality – which for Spurzheim was not incompatible with Christianity. Spurzheim also wrote about the inheritance of healthy or sickly constitutions and mental characteristics. Perhaps the details of Spurzheim’s hereditarian comments owe as much to earlier writers as do his phrenological and philosophical ideas. It is difficult to trace Spurzheim’s sources with no papers and few letters known. Reading Spurzheim along with Combe, however, there can be no doubt about the formers’ influence on the latter. Combe borrowed a radically naturalistic, progressive and normative philosophy full of scientistic elements such as hierarchical classifications, nested realms of natural laws and the unanswerable force of causal necessity. From these materials Combe eventually formulated what he called, after Spurzheim, the ‘doctrine of the natural laws’. This doctrine was expressed in many of Combe’s works but its primary showplace was Combe’s masterpiece The constitution of man (1828). This book went on to become one of the most widely read books of the nineteenth century. It sold more than 350,000 copies by the end of the century and was continuously in print from 1828 until 1899 with more than one hundred publishers in half a dozen languages.

2 3

See “Anti-Phrenologia” in Blackwoods Edinburgh Magazine 13 (1823), pp. 100-8, 199-206, p. 100; The Phrenological Journal, 1, 1823/4, pp. xxi, 94. Combe to Spurzheim, 18 February 1826.

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Fig. 1: Sales of George Combe, The Constitution of Man, Robert Chalmers, Vestiges of the natural history of creation, and Charles Darwin, The origin of species.

If we compare the English-language sales of Constitution with those of the better-remembered Vestiges of the natural history of creation (1844) or The origin of species (1859), we get a sense of how much more common The constitution of man was in the nineteenth century than its reputation today would lead us to believe (see fig. 1). As I have argued elsewhere Constitution did not just sell more copies but probably created more controversy and inspired more subsequent writers to imitate it than Vestiges and the Origin combined during the nineteenth century.4

The doctrine of the natural laws So what was Combe’s doctrine of the natural laws? The doctrine was a systematic, though somewhat vague and amateurish, bid to provide an alternative for traditional Christian systems as guides to conduct and especially as an alternative to beliefs of the fallen state of Nature and Man, the sufficiency and necessity of the Bible as a guide to daily living and as a moral, philosophical, and epistemological authority. Briefly stated, the doctrine went thus: if Man were to devote himself to understanding and following the natural laws, all would live in a happier, healthier world and experience the greatest possible joys and satisfactions as civilization, and individuals, progressed ever farther towards perfection. All the evils in the world follow from disobedience to the natural laws and all pleasure and progress follows from knowledge of and obedience to them. As ‘the true science of mind’, phrenology could be the key to unlocking this doctrine, but Combe was explicit in referring to phrenology and the doctrine of natural laws as distinct enterprises. Little in Combe’s account was very new but he did arrange his pieces into a convincing and provocative order which countless thousands of nineteenth century readers found profoundly moving. According to Combe, Nature was designed benevolently by a deistic creator according to a progressionist principle. It was not ruled by divine intervention but by a complex set of natural laws. The main categories of natural laws were physical, organic and intellectual or moral (the 4

van Wyhe (forthcoming 2003).

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wording varied). The three realms were reflected in Man’s constitution and corresponded to the “three classes of [phrenological] organs, the animal, moral, and intellectual.” 5 The laws were the regularities of matter and mind which the creator had willed at the beginning. Combe never tired of reiterating that the realms of natural laws which he preached acted independently of each other; although this too he borrowed from Spurzheim. 6 The physical laws included traditional notions of natural laws such as chemical properties, gravity, and other consistencies of Newtonian physics. The organic laws were a realm devoted to the unique properties of living organisms and Combe used them to espouse the importance of cleanliness, hygiene, adequate ventilation, guarding against rapid bodily temperature changes, bodily exercise, moderation in exertion, and sufficient rest. Obeying the moral and intellectual laws would lead to the enjoyment of “a fountain of moral and intellectual happiness”.7 New and higher pleasures awaited those who could bring their faculties into harmony with Nature. Therefore, apart from promoting good health, Combe valued intellectual more than physical pursuits. Modes of behaviour proscribed by the moral and intellectual laws included greed and corruption, employing people to do things for which they were not naturally suited, and capital punishment (then a common sight in Edinburgh). 8 Most of all respect for Combe’s style of secular natural philosophy was enjoined as progressive and just.

The law of hereditary descent One law within the realm of the Organic laws was what Combe called the ‘law of hereditary descent’, referring to the fact that offspring acquire characteristics from their parents – both of what would today be called heritable traits and Lamarckian inheritance. Combe’s law of hereditary descent was essentially his description of the fact that heredity occurred and that a good or bad constitution was inherited by offspring. According to Combe, the qualities of children were determined jointly by the constitution of the parents (though often to a varying degree) and by the faculties, which predominated in power and activity in the parents at the time of conception. Combe divided heredity into two basic kinds: the inheritance of inborn mental and physical characteristics 2. the transmission of acquired and even momentary mental and bodily qualities and conditions. Like Darwin’s Variation of Animals and Plants under Domestication of 1868, Combe provided many cases of heredity of various kinds. For example, Combe wrote that mothers have a strong hereditary impact on offspring, particularly if the mother is marked by strong mental or physical qualities. A mother’s state of mind, especially any strong impressions like fear or horror at the sight of a cripple could specifically imprint themselves on the offspring. (Incidentally, for this fact Combe cited the authority of Dr Darwin.) In general both parents contributed characteristics to offspring and it seemed likely to Combe that fathers contributed more to sons and mothers more to daughters. If a clever man married a dull woman then their children, because mixed, would be 1.

5 6 7 8

Combe (1828), p. 181. Spurzheim (1825). Combe (1828), pp. 19, 37. Gatrell (1996).

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less clever than the father. Close relations should not marry. In addition the too young and too old should not reproduce because their imperfect condition would be passed on to their children. The prime of health and vigour was heritable for Combe. The same inheritance of mental qualities was attributed to different human races – hence “each Hindoo, Esquimaux, Peruvian, and Carib, obviously inherits from his parents a certain general type of head; and so does each European.” In intermarriage between Europeans and Hindoos or native Americans similar effects would occur as when a clever man and dull woman reproduce – pureness of virtue would be diluted and lost – but the offspring would still be superior to those of pure native ancestry. According to Combe, mankind was arranged in a hierarchical scale of superiority and inferiority. The scale began with non-European races, especially those with dark skins “whose brains are inferior” at the bottom, and western Europeans, like Combe himself, at the top. 9 European interbreeding in India, for example, would lead to a mixed-race that would eventually rule the native inhabitants. Combe’s belief in distinct human ‘races’ and in a scale of their superiority were wholly unoriginal points. Despite the low value attributed to other ‘races’, Combe was vehemently opposed to chattel slavery and was an early critic of colonialism. For Combe non-European races were emphatically human, as phrenology proved, by possessing the same cerebral organs. Nevertheless in all things there were degrees of power or virtue. A phrenological brain organ could be well or poorly developed – but it was still the same organ. In the same sense all humans were ranked according to their natural gifts – some were intelligent, others stupid, some healthy, others sickly. Human races were essentially the same for Combe, though some were better than others. Combe asserted that vital and long-term conclusions were to be drawn from the law of hereditary descent. He thought that heredity alone enabled the progress of Man to occur in the long run, as each individual could increase his or her physical and mental powers through proper use and exercise and as the actual heightened state of physical and mental virtue could be passed on to offspring. Combe theorized that each generation could be given a head start by beginning at the heightened state of perfection reached by its parents. European society could gradually increase its concentration of intelligent and moral beings and lower races could gradually improve their stock and thus climb the scale of civilization. In order to best progress societies must practice improvement breeding almost like that done for domesticated animals. Only the fittest people in the prime of their lives, who were in perfect health and could afford to support a family should be allowed to breed. In support of the view that careful improvement breeding should be applied to humans Combe cited, among others, Horace, John Gregory10, Voltaire11, Dr James Gregory,12 John Mason Good13, Albrecht von Haller, and an unnamed medical friend. Interestingly in the marginalia to the 6th edition of Constitution held at the Whipple library in Cambridge an unnamed evangelical reader left traces of his reactions to Combe’s work. Among 9 10 11 12 13

Ibid., p. 194. John Gregory (1766). Voltaire (1766), s.v. “Cato.” James Gregory [c. 1780]. Good (1825), vol. 5.

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these are some scribbled responses to Combe’s hereditarian pronouncements. The evangelical reader objected to Combe’s proposal to improve mankind because of “[the] original depravity Adam fell!” The human species could not be improved because “the germ of all is bad.[…] hence Sin has descended to us all!”14 Combe’s lengthy focus on heredity, which totalled about 10,000 words in The Constitution of Man, more than was devoted to phrenology, brought to a vast reading audience many of the themes for which Darwin is now better known. Constitution brought these subjects from specialist contexts into the home as Jim Secord has shown that Vestiges brought evolution into the home.15 Darwin’s comment that Vestiges helped prepare readers for his Origin of species could have included Constitution which for thirty years had taught countless thousands of readers to think in terms of selective breeding and the cumulative effects of the early death of the sick or infirm: When we reflect on the transmission of hereditary qualities to children, we perceive benevolence to the race, in the institution which cuts short the life of an individual in whose person disease of essential organs has exceeded the limits of the remedial process: it prevents the extension of the injurious consequences of his errors over an innumerable posterity […] the race is guaranteed against the future transmission of his disease by hereditary descent.16

However a focus on Combe’s great influence should not lead us to think of him as the source of these ideas. Very similar passages can be found in Spurzheim’s earlier and less widely-read books. For example, Spurzheim wrote: “Since beggars, and those with hereditary dispositions to diseases, only propagate to the detriment of society and to entail misery on their progeny, were it not better to prevent them from marriage altogether?” The future of society is dependent on only “the stoutest and best made men” propagating and not those with “bodily weakness and disease”. 17 In his turn Spurzheim had been influenced to the same extent by Gall and the physician Pierre-JeanGeorges Cabanis as Victor Hilts has shown.18 These remarks by Spurzheim and Combe sound remarkably like Darwin’s view in The descent of man (1871): Yet [one] might by selection do something not only for the bodily constitution and frame of his offspring, but for their intellectual and moral qualities. Both sexes ought to refrain from marriage if they are in any marked degree inferior in body or mind; but such hopes are Utopian and will never be even partially realised until the laws of inheritance are thoroughly known. […] all ought to refrain from marriage who cannot avoid abject poverty for their children; for poverty is not only a great evil, but tends to its own increase by leading to recklessness in marriage.19

All three of these passages, from Spurzheim, Combe and Darwin are what Francis Galton would later call “eugenics” in his Inquiries into Human Faculty and Its Development (1883) as “the study 14 15 16 17 18 19

G. Combe (1836), 6th ed., pp. 109, 123, WSM Store PH-52. Secord (1989). This point was made earlier by Chadwick ([1975] 1995). p. 165. G. Combe (1828), pp. 247-8. Spurzheim (1825), pp. 178, 179. Hilts (1982), pp. 62-77. Darwin (1882), pp. 618-9.

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of agencies under social control which may improve or impair the racial qualities of future generations either physically or mentally.” Victor Hilts observed that, “Spurzheim placed most of his faith upon the regulation of marriage, whereas Combe resurrected Lamarck by teaching that parents could transmit good qualities to their offspring by perfecting those same qualities in their own persons.” To my knowledge there is no evidence that Combe borrowed this from Lamarck’s writings. Instead Combe seems to have expressed a common-sense impression of heredity.

Descendants of Combe’s hereditarianism I have argued elsewhere20 for the extraordinary influence of Combe’s writings, especially the Constitution of man. Combe also promulgated his laws of inheritance in his Moral Philosophy: or the Duties of Man Considered in His Individual, Social, and Domestic Capacities (1840). His brother, Andrew Combe, a well-known physiologist and phrenologist also stressed the importance of the transmission of characteristics via heredity.21 But Combe’s hereditarianism was spread much more widely as it was picked up by many other writers, especially in America by popular authors such as the phrenological Fowlers whose works were often little more than paraphrases of Combe’s writings. A number of their works dwelt particularly on the subjects of marriage and heredity including Orson Squire Fowler’s The practical phrenologist (1869), his Matrimony: or Phrenology and Physiology Applied to the Selection of Companions for Life (1842[?]) and especially his Hereditary Descent: Its Laws and Facts Applied to Human Improvement (1852). There was also Lorenzo Fowler’s popular Marriage: its history and ceremonies: with a phrenological and physiological exposition of the functions and qualifications for happy marriages (1847). The Fowler publishing industry also distributed similarly Combean works by the Rev. George S. Weaver such as Hopes and helps for the young of both sexes. Relating to the formation of character, … and marriage (1854). Furthermore, as Victor Hilts observed, two English writers most associated with hereditarian ideas in the latter nineteenth century, Herbert Spencer and Francis Galton were both influenced by Combean phrenology. The Combean flavour of Spencer’s hereditarianism is unmistakable. Galton was less influenced by Combean hereditarianism, but was nevertheless influenced by it. Therefore from Volney, Gall, Cabanis and others we can trace a direct line of descent for these hereditarian concerns through Spurzheim to Combe and from Combe to the Fowlers, Spencer and Galton and from them to a larger audience than ever before.

Methodology and conclusion John Waller remarks in his important recent article22 that in the past we tended to conceive of eugenics popping into existence in late Victorian Britain from a context of “rival economic superpowers and an increasingly volatile metropolitan underclass.” Waller is quite right to conclude that this was not the case and that Galton was not a founding father as he has been portrayed. Waller goes on to declare: 20 21 22

van Wyhe (forthcoming 2003). A. Combe (1834). Waller (2001), pp. 289-457.

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To identify Galton as a primary causal agent in the early history of eugenics is to overpersonalise an episode and period in which, as I have sought to show, individuality is insignificant in comparison with context. […] By attaching too much importance to individuals we lose sight of the fact that¯¯in terms of causal agency¯¯the idea of eugenics arose from a general fascination for […] and a particular set of social, institutional and political circumstances of the mid-Victorian period.23

I completely agree that Galton should not be identified as an original source, but I think it is unjustified to conclude from this fact that “the idea of eugenics arose from a general fascination.”24 Ideas cannot arise in social contexts themselves – literally speaking ideas arise only in individuals’ heads. Of course all individuals are within a social context and their ideas reflect their context – but to say that the ideas arose from the context is to be ahistorical. It is ahistorical because if ideas are spontaneously generated by contexts then they would have no history, that is they would not contain elements or characteristics of past ideas. We would not see the gradual cultural change, which we observe and the affinities between ideas in different periods. My discussion of the history of Combe’s hereditarian ideas and more so Hilts’ and Waller’s work show that ideas of heredity developed gradually over time from person to person and from decade to decade. I have suggested that Combe played a disproportionately major role in the propagation of hereditarian ideas in the nineteenth century due to the propitious success of his Constitution of Man. More people encountered and were influenced by his versions of heredity than other versions. No one proposes returning to writing only histories of Great Men but we cannot overlook that humans are the agents in history. The hereditarian ideas that were available in the late nineteenth century had the characteristics that they did, including notions of acquired characteristics, purity of blood, responsibility to breed only with healthy mates etc. not just because of the context of the moment, but also because of their history, that is, where they came from, or what they were before, because of the people who had possessed and promulgated them at an earlier time in different contexts. I think Waller is correct that the different context of the latter decades of the century explains why eugenics became the widespread phenomenon familiar to us but I think we must always consider context as well as the ancestry or earlier sources or inspirations of the ideas in a context.

23 24

Ibid. Ibid. Italics mine.

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References Chadwick, Owen. [1975] 1995. Secularization of the European mind in the nineteenth century. Cambridge: Cambridge University Press. Combe, Andrew. 1834. Principles of Physiology Applied to Health. Edinburgh. Combe, George. 1828. The Constitution of man. Edinburgh/London. _____. 1836. The Constitution of man. 6th edition. WSM Store PH-52. Darwin, Charles. 1882. The descent of man. 2nd edition. London: John Murray. Gatrell, V.A.C. 1996. The hanging tree: execution and the English people 1770-1868. Oxford: Oxford University Press. Good, John Mason. 1825. Study of Medicine. 5 volumes. London. Gregory, James. c.1780. Conspectus Medicinæ Theoreticæ. Edinburgh Gregory, John. 1766. Comparative View of the State and Faculties of man with those of the Animal World. 3rd edition. London. Hilts. 1982. “Obeying the Laws of Hereditary Descent: Phrenological Views on Inheritance and Eugenics.” Journal of the History of the Behavioral Sciences (january ): 62-77. Secord, James A. 1989. “Behind the veil: Robert Chambers and Vestiges.” In History, humanity, and evolution: essays for John C. Greene. Cambridge: Cambridge University Press. Spurzheim, Johann Gaspar. c.1824. Philosophical catechism of the natural laws of man. London. _____. 1825. A view of the philosophical principles of phrenology. 3rd edition. London. van Wyhe, John. 2002. “The authority of human nature: the Schädellehre of Franz Joseph Gall.” British Journal for the History of Science (march): 17-42. _____. Forthcoming 2003. Phrenology and the origins of naturalism in Victorian Britain. Voltaire. 1766. Philosophical Dictionary. London. Waller, John. 2001. “Ideas of heredity, reproduction and eugenics in Britain, 1800-1875.” Studies in History and Philosophy of Science Part C: Studies in History and Philosophy of Biological and Biomedical Sciences 32(3): 457-489.

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Majorat: Literature and the Law of Succession in the 19th Century Ulrike Vedder

This paper explores the lines of transmission between generations. It deals with what is passed on from one generation to another, in which ways, and who the agents mediating between the generations are. In dealing with these questions, I shall be looking more closely at the legal system, respectively the law of succession, as the principal cultural system of regulation that seeks to determine the transfer of material possessions and, as such, the relationships underlying such material transfers. Since we are dealing with definitions of the family and of kinship order, and how these are reproduced, respectively cast into the future, the discourses of nature and law clash time and again. The figure of the bastard, which I shall be considering briefly in the narratives of inheritance which this paper discusses, is a valid case in point, since the bastard recurs as a figure of conflict along the lines of inclusion and exclusion; of what is proprietary and native that is, and what is not and is therefore alien. As I intend to approach the discourse of inheritance and succession as it stands in the law by considering how it is ‘breached’ in literature, I shall focus on literary texts rather than legal ones. We shall see that the authority of the law, that is its power to determine and sanction, is challenged in literature, just as the boundaries that are drawn by those laws of nature considered valid at any point in time are unsettled when those forces are brought into play that cannot be separated from the processes of transmission between generations: passion, guilt and power.

1. Law, Nature, Literature The law of succession governs the relationship between genealogy and legitimacy, kinship and property, and affiliation and transfer between generations. Genealogical and legal relationships are structured by semiotic systems, however, that need to be interpreted so that they become valid. This is one reason why literature is a preferred arena to debate issues concerning genealogy and the making over into law of property and kinship relationships. On the other hand, it is worth noting that a fictionalizing impetus takes effect time and again in the law and its dispensation. Specific ways of dealing with authority and authorship, with rhetoric and fiction, with interpretation and the production of meaning, to mention just some issues, play their part in the complex relationship of law and literature. Both – law and literature – develop strategies to posit and repeal what has been statuted. Both apply different strategies to make things of, and exist in, time. Both adopt a range of procedures to typify and represent events as what comes to be known as a case, a case history and a narrative; to make reality unequivocal or ambiguous; to bring into relation the normal case and the exception; to enforce, differentiate and abolish legal and aesthetic norms. These various aspects of law and literature form a rich nexus of issues concerning the law of succession, where the problems of defining basic terms such as kinship and property become particularly apparent; likewise, so do the intricate entanglements of property and passions that are involved in the transmission between generations, or the difficulties of determining points of

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rupture and devolution, of negotiating inclusion and exclusion, affiliation and alienness. These difficulties also include the fundamental doubt that turns every indication of paternity into a fiction or, to put it differently, that amounts to the potential conflict between maternal knowledge about the events of family reproduction, on the one hand, and the bequeathing of name and property along patrilinear lines of succession, on the other hand (whose principal interest it is to pass on the father’s estate and affiliation to social rank to his own heirs on a continual basis). As we shall see in one of Balzac’s novels, the gender war about fiction and reproduction, which no legal settlement can resolve, is waged in literature. The law of succession expands through time to take up the past, build on tradition and determine directives for the future of a family order. Another element of its impetus to fictionalize lies in the fact that it casts existing law into the future, since each last will reckon with the case that has not (yet) occurred: death. This problem is aggravated by a specific settlement of inheritance that is projected onto the open, unfinished future and remains nonetheless tied to a lineage endowed with a past that is still effective, and that is considered great and worth preserving: the majorat (in Germany and France, that is the entailed estate). The majorat is intended to transmit and make immortal the dignity, virtue and glory of a family by subjecting all future generations to a form of succession that privileges the “first born of the paternal line” (by way of the authority of the estate’s founder) and preserves the indivisibility of the estate. This is meant to safeguard a family estate – real estate, in most cases – against fragmentation and loss, as much as against financial speculation and indebtedness. Thus, according to whichever economic point of view is taken, family property is transformed either into a security with maintained value or into dead capital. Instituting a majorat (which became more and more common in Germany from the seventeenth century onwards, for example) thus determines that commodities and assets (special funds) are to be indivisible and unalienable – through a private declaration of volition, that is through a legal transaction.1 According to its reformulation in the General Common Law of the Prussian States (of 1 June 1794), assets would go to “the next in line in the family, according to grade, […] among several close enough, however, to the elder”. A later commentary states: “Individual-Succession innerhalb des Agnaten-Stammes; denn nur jene erhält den Reichtum, nur dieser den Namen.”2 What we recognise here is that the law produces a ‘rhetorical’ community – the family that is to be perpetuated – and points to the rhetorical essence of supposedly natural communities, while the law of succession and the majorat in particular are supposed to naturalise legal communities. The legal institution of the majorat indicates the status of ‘Nature’ in two directions: first, in that recourse to nature is taken in determining succession in terms of blood relationship, if succession is only able to occur within the male line of relatives, that is within the patrilinear filiation of the same blood; secondly, by taking recourse to naturalisation, which lies in the majorat being perpetuated forever, ‘for all eternity’, in that property is seen to be virtually sanctioned by the law of nature. The majorat constitutes a regulated means of consolidating a family estate, and of bringing material goods into time. In turn, such an entailed estate materializes time. As a “materialized 1 2

Cf. Eckert (1992), p. 24. Gierke (1909), p. 104.

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medium”3 for the continuity of a family and its assets, the majorat establishes a particular form of genealogy, which is charged with a high conflict potential, that makes it of interest for literature. This potential results first from the injustice that only the first-born inherits, secondly from the founder’s claim to authority by committing succeeding generations to a specific form of tradition for all eternity through an immutable majorat, and thirdly potential conflict results from the problems of defining terms such as family, paternity, parental love and the concomitant difficulties of producing signs and their interpretation. Literature makes use of these uncertainties and complications. This can go as far as questioning not only the legitimacy of progenies, which means calling into question family affiliation, but, as we shall see in one of Achim von Arnim’s stories, it can go as far as challenging the legitimacy of the laws of nature, which amounts to calling into question affiliation to the human species. During the French Revolution, legislation refers to the new general principles of freedom and equality, thus aspiring to a “morcellement”, a dividing up of the large estate holdings in the hands of the landed aristocracy. Accordingly, the so-called “substitutions” are abolished. There is a conspicuous rhetoric of excess and unnaturalness in the ensuing debates: “monstre des substitutions”, “monstrueuse inégalité”, “monstruosité politique”. 4 In these debates, furthermore, the present is set against the adherence to the past as well as against the majorat being cast into the future, to which it keeps being sacrificed: Thus, there is mention of the Ancien Régime’s regulations of the law of succession “qui subordonnent les intérêts du peuple vivant aux caprices du peuple mort” and where “la génération qui est se trouve constamment sacrifiée à celle qui n’est point encore”.5 The present’s ‘gap in time’ becomes an insoluble problem in the literary texts embracing the majorat. The revolutionary legal texts, by contrast, look for a solution that is both unequivocal and final. Thus the Convention states in Article One of the Décret sur les substitutions of 14 November 1792: “Toutes substitutions sont interdites et prohibées à l’avenir.” Now this did not mean that majorats had been done away with for good, however. There are numerous clashes in the first half of the nineteenth century, both in France and in the German states, such as Prussia, which lead to the alternating abolition and the (more or less clandestine) reconstitution of majorats.6 Arguments in favour of and against the majorat are also debated in the literary texts which I wish to look at. The rhetoric of the law and of jurisdiction come into specific operation in these texts, be it in trial scenes or in drawing up settlements. Beyond such instances of legal discourse, the narratives of inheritance and succession of the first half of the nineteenth century deal with the law’s power to define and interpret what constitutes family and the linking together of generations; vice-versa, these narratives are also about how legal norms are made legitimate in the face of the threatening disruptions of property situations and the succession of generations, arising part and parcel from the transmission and recurrence of the guilt, deaths and passions inscribed in the majorat’s very conception of succession. In the following, I shall discuss the legal-literary order and how genealogies run their course 3 4 5 6

“[…] das Majorat in seiner Funktion als materiales Medium der Kontinuität der Familie […]” (Mangold 1989, p. 228). Cf. Eckert (1992), pp. 180-185. Ibid., p. 202. Cf. Dietze (1926); Sybel (1870); Eckert (1992).

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in three texts that deal with the antiquated legal institution of the majorat. Around 1800, legally sanctioned paternal authority enters not only a state of crisis, but leads straight to the downfall of families, as we shall see in E.T.A. Hoffmann’s Das Majorat (1817). Thus, an entirely different social principle takes the place of the law: the power of money, respectively capital. This process is demonstrated in a polarised manner in Achim von Arnim’s Die Majoratsherren (1819). The power of money, however, makes the exclusiveness of the ‘old family’ in some way contingent, as in Achim von Arnim’s story and then particularly in Honoré de Balzac’s novel Le Contrat de mariage (1835). Both narratives feature main characters that are hybridized, respectively creolized, that are contaminations of the legal discourse and of the discourse of race. This will amount to an account of the end of genealogies, that is the triple demise of old lineages.

2. The Genealogy of Death: Law, Writing, Fiction (E.T.A. Hoffmann) In E.T.A. Hoffmann’s Das Majorat (1817), the lineage of “Freiherr Roderich” comes to an end without any descendants; all members of the family have either been murdered treacherously, have fallen in battle, or have died in accidents or from grief. The ancestral castle lies in ruins and the rich landholding has fallen into the hands of the state. The splendor familiae et nominis, which is mentioned repeatedly in all arguments in favour of the majorat, and which features as a dazzling objective over this story, too, has become more and more tainted precisely by the majorat and has ultimately evaporated. “Eine verfehlte Operation”, a failed operation which is not explained in any detail, but has obviously occurred in the field of “geheimer Wissenschaft”, 7 leads Freiherr Roderich to seek the cause of his misfortune in his “ancestors’ guilt”, who had left the ancestral castle. Furthermore, he determines it to be a majorat, “um […] wenigstens das Haupt der Familie an das Stammhaus zu fesseln.” Determining the future through the law of succession is thus supposed to bind together house and lineage, even more so since “deren Zweige schon in das Ausland hinüberranken.”8 Besides the injustice of the majorat and its consequences, though, shackling all these creeping branches to one place and its inherent laws, fatally causes them and hence the entire family pedigree to break off. In Hoffmann’s story, the majorat is marked by its almost imminently lethal effect, taking hold of all the founder’s descendants. Both sons and their four descendants, as well as two wives and the manor’s steward die due to the majorat, whereas not one single birth or fathering is recorded. The majorat thus consists in the recurrence of death: The sons die their fathers’ deaths, and the first son’s murderer keeps returning to the room of death – first as a sleepwalker, then as a dying man, and finally as a ghost. What this amounts to is in fact a genealogy of death and of the undead. In other words: Just as the naturalisation of the legal institution of the majorat turns into a rhetoric of unnaturalness and the monstrous, so does what has been regulated in supposedly ‘rational’ legal terms in E.T.A. Hoffmann, and later in Achim von Arnim, go over increasingly to the ‘irrational’, that is specifically to the genre of ghost and visionary stories. Accordingly, the new beginning that Wolfgang, the founder’s son, hopes for by taking on the name “Born” – he had married abroad against his father’s will and under a false common law name – is impossible, since he has named his own son Roderich again – after his father. Besides 7 8

Hoffmann (1985), p. 200. Ibid.

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names and properties, there are various other agents that mediate between the generations, such as letters – letters announcing death, letters written from the deathbed – and written documents negotiating the past and future: certificates of baptism, certified excerpts from church registers, the sovereign’s confirmation of the endowment, tally books and documents left behind, such as wills, collections of correspondence, or confessions. These documents compete for interpretation, only from which the factualness of the law begins to transpire. Thus, litigation proceedings are required to establish how the endowment of the entailed estate and the will are connected, as much as to ascertain the conclusiveness of the various proofs of identity, when the legitimacy of Wolfgang’s son, who appears by surprise as an issue from the secret marriage and who is initially suspected of being a bastard not entitled to inheritance, is interpreted in court. When the first-person narrator, a young “Justizmann”, visits the ruins of the castle at the end of the text as the sole survivor of the story, it is not as if he has claimed victory in a historical process, as one might be tempted to gather from the analogy of the collapsed ancestral castle with the demise of the aristocracy. Much rather, he is caught up in the narration of a genealogy of death, a fact that is made concrete so to speak by Roderich inviting the narrator to be buried alongside the noble family in the same tomb.

3. Mixture and Exclusion (von Arnim) In Achim von Arnim’s story Die Majoratsherren (1819), which was almost written at the same time as E.T.A. Hoffmann’s Das Majorat, the narrator seeks to elude such entanglements. Despite the many blendings and meddlings that traverse the text, Arnim’s narrator is separated by a manoeuvre that cuts him off sharply from the age of the majorat and the lineage that collapses with it. He situates himself after the French Revolution, when, in the frame of the story, he discusses the changing of times that the French Revolution entailed and that severed the colourful and fulfilled pre-revolutionary past – the age of the majorat – from the post-revolutionary present – a monotonous and poor age. This shift is symbolized in the majorat house, which, by the end of the story and of the French Revolution, will have fallen into the hands of Vasthi, an old unscrupulous Jewess. In dealing with Achim von Arnim’s story, I shall focus on the key role of time, respectively temporality, on the one hand, and the mixing and separation of what is native and proprietary from what is not, on the other. Both aspects are crucial to considering the issues of inheritance and transmission by succession, since both are to be made governable by the law of succession. The organising principle of the majorat fails, however; for one, within the family, as the majorat order of succession is ignored in treacherous ways; for the other, failure occurs in historical-systematic terms, since the economy displaces the law – in its function as a dominant principle which rules the organisation of society. The story of the young, indecisive majorat lord, who comes to his substantially older cousin – a grotesque, decrepit lieutenant, who spends his life waiting for a spouse and an inheritance – , only lasts for four days, during which the past history, that has occurred thirty years before, is revealed. At that time, the old majorat lord, who was without a male heir, had given away his only child, a new-born daughter, who was not entitled to inherit, to a Jewish “Roßtäuscher”, 9 a horsetrader, who called the child Esther and took her in as his foster daughter. Concurrently, the old

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lord secretly adopted the illegitimate son of a lady-in-waiting as his own to cheat the rightfully entitled cousin out of the majorat. This cousin is portrayed as being as immutable as the majorat house that has been unoccupied for thirty years, but whose clocks are still wound up, and as lifeless, too, as he goes out on the same walks in his worn out uniform coat and clings to unchanging snuff and visiting habits. Neither his preoccupation with heraldry nor his collection of coats of arms are able to enfuse his life with any of the old glory or hope for the future, as these interests are contaminated by economic necessity, which requires him to copy the collection, stick the coats of arms on with care and sell them to earn his crust. The protection that the majorat sets against any form of change thus also amounts to a constraint of not being able to change anything. Both elder protagonists are subjected to this constraint – the cousin is stuck with the old lady-in-waiting (i.e. the illegitimate son’s mother), for whose love he has been waiting in vain for thirty years, just as he has for the entailed estate – as well as the two younger protagonists, the young majorat lord and the horse-trader’s foster daughter, Esther, who are both permeated by the ghosts of the past. They both fail to live in the present, which is not theirs, and which will ultimately lead to their deaths. The old cousin does indeed go on to take possession of the entailed estate and marry the lady-in-waiting, moving into the majorat house with her and a horde of animals – instead of a swarm of children, notably. However, life there, which he has desired so much for thirty years, is hell; the majorat lord is humiliated by his spouse and her horde of animals until one day he passes away unnoticed and ingloriously. This draws attention to the issue of the present in genealogical and law-of-succession narratives. The generations that the majorat chains together are conceived as a continuum of the past and the future – without assigning more to the present than maintaining the majorat in due form. In Hoffmann, the present consists of administering the majorat and its documents and tally books. In Arnim, the loss of the present that is so fixed on the past brings forth a “Hohlraum in der Zeit”,10 a hollow space which can be identified in topographic terms as the unoccupied majorat house with its clocks ticking away emptily. Vasthi, the old Jewess, Esther’s step-foster mother, makes use of the hollow space of this empty present. In the wake of the Revolution, after the abolishment of the majorat, the release of the Jews and, as we are told explicitly, “unter der Herrschaft der Fremden”, 11 Vasthi emerges as a warprofiteer to take over “das ausgestorbene Majoratshaus durch Gunst der neuen Regierung zur Anlegung einer Salmiakfabrik.” And the last sentence of the story reads: “[…] es trat der Kredit an die Stelle des Lehnrechts.”12 In Achim von Arnim’s story, the economy comes to occupy the place of the law, monetary transaction (with its Jewish connotation) takes the place of (feudally marked) real estate holding, and unalienable real estate – charged with family history – is displaced by its speculative, abstracting monetary value. This makes contingent affiliation with the family/the lineage, a contingency which the text produces as a blurred distinction between what is proprietary and what is alien. Thus, the young majorat lord, who as a visionary is hardly capable of distinguishing between 9 10 11 12

Arnim (1991), p. 217. Oesterle (1988), p. 29. Arnim (1991), p. 250. Ibid., p. 251.

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reality and vision, is a foisted bastard in terms of matrimonial law and the law of succession. Esther is not only the foster daughter of a Jewish “Roßtäuscher” – who earns a living not necessarily from ‘täuschen’ (deceit) but from ‘tauschen’ (exchange) – but she is also an image of her Christian mother before she is cast – as she is dying – as a mythical, hybrid creature consisting of a Jewish angel of death and a Christian shining light. The two illegitimate children, the young majorat lord and Esther, reveal their exchanged and deceptive origin in a vision: “Ich bin Sie und Sie sind ich.”13 So what is proprietary can no longer be identified clearly, neither within family boundaries – given the multiplication of father- and motherhoods as natural, adoptive, step and foster parents – nor within the human species: In the end, the cousin, caricatured as a turkey, is obliged to wait upon the lady-in-waiting’s cats and dogs at the majorat banquet table. Given such narrative blendings of boundaries, one particular boundary that is brought into focus time and again in the text gains significance and explosive force: that between the town and the Jewish ghetto, which can only be crossed at the expense of death. The boundary between Christians and Jews, which first is introduced in topographic terms, is then sharpened and naturalised in a racist manner as that between the human and the non-human, f.e. when the Jewess Vasthi is strangling Esther, her Christian foster daughter, and when she appears as a monstrous sharpened silhouette: […] wie die ausgeschnittenen Kartengesichter, welche einem Lichte entgegengestellt […]: sie erschien nicht wie ein menschliches Wesen, sondern wie ein Geier, der lange von Gottes Sonne gnädig beschienen, mit der gesammelten Glut auf eine Taube niederstößt.14

The crisis of family authority and its inheritance is transferred into anti-Jewish denunciation to shift the decrepitness of feudal-patrilinear transmission, caricatured in the grotesque drawings of the elderly cousin and his lady-in-waiting, onto the takeover by the “foreigners” and their financial resources. As a universal means of exchange, money abstracts any tangible value (such as, for example, that of entailed real estate, with its family-historical significance). That it is exchangeable in general terms makes money function as that which basically devalues the specifically individual (and proprietary); in Lacan’s terms, it becomes “the most annihilating signifier”. This process becomes even more ‘unnatural’ in Achim von Arnim’s story as family disorder, destruction and annihilation radiates from the mother’s position in the system (Esther’s mother, the lady-inwaiting, Vasthi). Honoré de Balzac’s novel Le Contrat de mariage, which I would like to look at next, offers another account of the way in which the discourse of law (and lawfulness) and the discourse of nature (and race) compete with each other through money, which is in turn bound up with gender.

4. Classification and Gender War (Balzac) In Balzac’s novel (first published in 1835, then republished in 1842 in the Comédie humaine, in the Scènes de la vie privée), a contract between spouses is concluded in which the establishment of a majorat features as a key clause that will ultimately ruin the marriage. What is highly significant 13 14

Ibid., p. 232. Ibid., p. 245.

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about this contract is that it is concluded between two ‘hybrids’, of which one, as we are told, is assigned to the social order – “un métis social” – and the other to the natural order – “la créole […] une nature à part”. One of the parties to the contract is Paul de Manerville, a rich heir from landed gentry stock, gentle and naïve, whom the text names as “un métis social”, 15 a social mestizo, a bastard in other words. Paul owes this designation to his uncapability of exercising any male-aristocratic power. Though he is forced to follow the customs of the aristocratic habitus, he pursues the entirely unaristocratic and quite bourgeois-sentimental ideal of love, desiring wedlock with a woman who stands by him, shares his thoughts and secrets, forms a unity with him in order to lead a married life shaped by intimacy, love and naturalness: “Un cœur à qui confier mes affaires et dire mes secrets.”16 Provincial society nicknames Paul “Fleur des pois”, “pea-flower” (which was what the novel was first called), which is an early eighteenth century designation for an “homme à la mode”. At the beginning of the nineteenth century, when the novel is set, however, this name is no longer “à la mode”, but an anachronism that the novel uses as a marker of a political-social conflict: to be named a “fleur des pois” is considered an honour by the “royalistes”, whereas it evokes irony among the “société liberale”17 – not least, perhaps, because the pea is a hermaphrodite. The peaflower is known to have acquired a key systemic position at the end of the nineteenth century, among others in Mendel’s experiments and creation of “plant-hybrids” (recorded in his Versuche über Pflanzenhybride, first published in 1866). Mendel chose the fleur des pois, because it allows one to reconstruct and calculate – and thus produce by experiment – the purity and hence proportions of a mixture in an optimal manner. This admittedly anachronistic association draws attention to Balzac’s arrangement of the novel as an experiment, specifically as an experiment in the domain of nature, and in the social and legal spheres – as an experiment that concerns the unity (of the species) and the difference (between the genders), where two hybrids are supposed to unite. As it happens, the “métis social” happens to choose a stunningly beautiful Creole, whose inheritance from her deceased father, a businessman, has long been squandered and who is thus in search of a marriage that will bring her profit in economic and social terms. She and her mother are not only Creoles, but female, and these two features of ‘nature’ make up her dangerous character: “La créole est une nature à part […]; nature gracieuse d’ailleurs, mais dangereuse.” 18 These dangerous hybrid-formations transcend the modes of the natural and the social, since their boundaries and transformations are constantly subject to discursive negotiations that Balzac explores in various different ways: as those occurring in the area of the law (within the novel) and as those in the sciences (in the programmatic preface to the entire Comédie humaine). In the “Avant-propos” (written 1842), he ponders his dream – a “chimère” 19, as he calls it: the comparability of “l’Animalité”, and the human, “l’Humanité”. As is well-known, Balzac also refers to Cuvier’s dispute in the Académie with Geoffroy Saint-Hilaire (1830) about the relationship between unity and variety; he follows Geoffroy Saint-Hilaire’s position of a unité de composition 15 16 17 18 19

Balzac (1976b), p. 530. Ibid., p. 534. Ibid., p. 537. Ibid., p. 605. Ibid., Balzac (1976a), p. 7.

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when he states: “Il n’y a qu’un animal. […] un seul et même patron pour tous les êtres organisés.”20 Tracing back all milieu-determined species to a fundamental, unified model encourages Balzac to undertake his programme of a comprehensively catalogued society, of classifying the social, in the Comédie humaine, on the analogy of the organising principles of Buffon’s Zoologie. He then goes on to cite the “drames”, “hasards” and “confusion” that stand in the way of such organising principles. Contrary to the writing of literature, which he at first conceives as the application of a theory, Balzac’s novels in the Comédie humaine are precisely about the inscrutabilities of classification, about how coincidences and passions intervene in the drawing up of boundaries and distinctions between the various genera, and about the inconsistency and lack of uniformity of the composition. This becomes apparent in Le Contrat de mariage, not only by featuring hybrid characters, but by negotiating a contract between the two family lawyers that reaches more and more into the arena of warfare. And this is where we find the calculating – and in this way ‘unnatural’ – Creole, who will emerge victorious from the battle waged over the contract between spouses, in which the newly inserted majorat is supposed to act as a barrier preventing the flow of money from the male to the female side. The Manerville’s longstanding family lawyer had after all managed to bring the heir’s estate through the French Revolution and the wars unscathed. But neither the lawyer nor the amorous husband are up to the gender war that the young wife and her mother wage, which entails the husband’s complete and utter financial ruin, and ends in his departure for Calcutta, while his mother-in-law takes possession of his rich commodities and assets. 21 Classifying human beings into social species, on the analogy of classifying animals into zoological species, which Balzac plans in his preface to his cycle of novels, comes up immediately against the difficulty of gender difference, which compounds his classification: “La description des Espèces Sociales était donc au moins double de celle des Espèces Animales, à ne considérer que les deux sexes.”22 Whereas Buffon had managed to describe the females of a particular species in a few lines in his “Zoologie”, Balzac makes it quite clear that one annotation on the female ‘variant’ will not do in cataloguing the social species; moreover, Balzac’s basic hypothesis, which follows Geoffroy Saint-Hilaire’s, that there is a unity of the genus, whose wealth of variations is developed by each particular milieu and which can, given the analogies between these variations, ultimately be put down to the notion of unity again – this hypothesis can no longer be sustained once the gender issue is taken into account: “[…] dans la Société la femme ne se trouve pas toujours être la femelle du mâle. Il peut y avoir deux êtres parfaitement dissemblables dans un ménage.” 23 The programmatic text of the “Avant-Propos” does not go into more detail about this discrepancy, which explodes the analogy of humanité and animalité and thus, too, the notion of the unity of the human being as a basis of a social world that can be catalogued. By contrast, the novel acts out this area of conflict, by inserting a legal construct – in the shape of a contract between spouses – in the place of a mediating natural entity. Within this contract, the majorat appears as the unifying central character, with whose assistance all differences between the sexes 20 21 22 23

Ibid., p. 8. In other words, and to quote Balzac’s: “J’ai peint toutes les infortunes des femmes: il est temps de montrer aussi la douleur des maris.” (Balzac: “Lettres à l’Etrangère”, 1: p. 275; cf. Perrod 1968, p. 221.) Balzac (1976a), p. 9. Ibid., pp. 8 f.

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and families are supposed to be resolved by way of the law of succession, but which actually causes the conflict, respectively the war to erupt with all possible passion and to ultimately put it beyond reconciliation. It is the Creole’s young lawyer – a modern-type profiteer of speculative transactions, who later goes on to marry “une mûlatresse riche” 24 – who manages to pull off a surprise coup by inserting a clause quite by the bye that the old lawyer fails to take seriously because of its entirely fictitious character. This clause sketches the case of an absence of descendants, in which case the majorat is to be transferred into the spouses’ community of property; as it happens, the young wife brings about this very case and makes this fiction a reality by evading descendancy. The descendants no longer die of the majorat, but they are even not brought into life. And just as it is merely a clause that is slipped in ex post facto, inserted into a comprehensive majorat contract like a foreign body, and becomes the decisive moment in the spouses’ sexual and genealogical life, thus it is but one single word on the morning of the wedding with which the young bride tells her mother “que si Paul avait gagné la partie au jeu du contrat, sa revanche à elle commençait.” For she had already gained her husband’s complete obedience – “la plus parfaite obéissance”25 – in the wedding night. We are not told which word the bride tells her mother, but the fact that her husband’s ruin goes ahead according to plan is clear enough evidence that it was one of negation, such as ‘no’, perhaps even ‘never’, that expresses her refusal of ‘natural reproduction’ and denial of conjugal rights. To conclude: Balzac scholarship has shown that the old lawyer would have been able to prevent the husband’s downfall with legal means, since the subsequently inserted contractual clause is in fact ineffective.26 Balzac, however, demonstrates that it is the institution of the majorat itself that is ineffective and that it is the fact that it is drawn up at all that leads to destruction. But this means, for Balzac, Hoffmann and Arnim, giving up the conception of the direct and linear transmission, first of property and title, that is position; secondly, of name and affiliation to what is proprietary, that is identity; thirdly, of resemblance and essence, that is nature; and finally, of time and future, that is perpetual continuity. Troubling, even thwarting the genealogical thinking is highlighted in the literary texts with their specific means in three ways: first, by the unpredictable passions and entanglements that are always also transmitted and that the three texts that I have considered set in motion; secondly, by the increasing abstraction of the property that it is to be entailed, which has a tangible shape as real estate and gold nuggets in Hoffmann, features as the transition from a feudal system to credit in Arnim, and appears as a speculative capital investment in the shape of bonds and shares in Balzac; and thirdly, by the fact that all three texts present figures of the hybrid, the bastard, the undead, since these are all characters that thwart the majorat and its underlying principles, namely its determination of affiliation, its determination of the perpetuation of the proprietary, and its inherent logical consistency in one generation succeeding another.

24 25 26

Ibid., 3: p. 623. Ibid., p. 618. Cf. Perrod (1968).

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References Arnim, Achim von. 1991. “Die Majoratsherren.” In Erzählungen. Edited by Gisela Henckmann. Stuttgart. 211-251. Balzac, Honoré de. 1976a. “Avant-Propos.” In La Comédie humaine. Volume 1. Edited by Pierre-Georges Castex. Paris. 7-20. _____. 1976b. “Le Contrat de mariage.” In La Comédie humaine. Volume 3. Edited by Pierre-Georges Castex. 527-653. Dietze, C. von. 1926. “Fideikommisse.” In Handwörterbuch der Staatswissenschaften. 4th edition. Volume 3. 993-1006. Eckert, Jörn. Der Kampf um die Familienfideikommisse in Deutschland: Studien zum Absterben eines Rechtsinstituts. Frankfurt/M. u.a. Gierke, Otto. 1909. “Fideikommisse (Geschichte und Recht der Fideikommisse).” In Handwörterbuch der Staatswissenschaften. 3rd edition. Volume 4. 104 ff. Hoffmann, E.T.A. 1985. “Das Majorat.” In Sämtliche Werke. Volume 3, Nachtstücke und andere Werke 18161820. Edited by Wulf Segebrecht and Hartmut Steinecke. Frankfurt/M. 199-284. Mangold, Hartmut. 1989. Gerechtigkeit durch Poesie. Rechtliche Konfliktsituationen und ihre literarische Gestaltung bei E.T.A. Hoffmann. Wiesbaden. Oesterle, Günter. 1988. “Illegitime Kreuzungen : Zur Ikonität und Temporalität des Grotesken in Achim von Arnims Die Majoratsherren.” Etudes Germaniques 169: 25-51. Perrod, Pierre-Antoine. 1968. “Balzac et les ’Majorats’.” In L’Année balzacienne. Paris. 211-240. Sybel, Heinrich von. 1870. Geschichte der Revolutionszeit von 1795-1800. Volume 4. Düsseldorf.

Translation: Mark Kyburz

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“Victor, l’enfant de la forêt” – Experiments on the Heredity of Human Nature in Savage Children Nicolas Pethes

1. Observing the innate: cases of wolf children At the end of the 18th century heredity was not discussed within the empirical sciences. Although natural historians such as Linnaeus, Buffon, or Maupertuis already suggested theories on the generation of human beings and the inheritance of their traits, it was a philosophical debate that dominated the field: the discussion surrounding the innateness of human faculties. Before the rise of the life sciences, evolution theory, and genetics, ‘heredity’ is a problematic concept not only within the realm of experimental science, but also within the realm of ideas. The philosophical concept of innateness addresses two questions: Is there a ‘human nature’ common to all of humanity? And are the individual faculties of a single human acquired or inherited? The latter distinction hints at the discursive context in which the question of inheritance is addressed. This context is the distinction between ‘nature’ and ‘culture’ and the various evaluations both concepts experience in the course of the 17 th and 18th centuries. Schematically speaking, the notion that culture provides the best possible state for human beings, as represented by Samuel Pufendorf, is opposed to Jean-Jacques Rousseau’s pessimistic account of the depravation of man’s nature within society.1 Transferred to the philosophical issue of heredity, this opposition reads as follows: Is the education within society responsible for every positive trait within otherwise ‘savage’ humans? Or do we have to strip away everything society has taught and trained us in order to find our ‘essential nature’, the state in which none of the current political depravations have spoiled our innate knowledge of a good life? Rousseau, of course, was well aware of the methodological problems associated with the latter choice, given the fact that all of humanity seems to live within larger or smaller social units. Consequently, he asks at the beginning of his Discours sur l’origine et les fondements de l’inégalité parmi les hommes from 1755: “What experiments would be necessary to achieve knowledge of natural man? And what are the means for making these experiments in the midst of society?” 2 How can we validate the idealistic claim of the ‘naturalness’ of human nature? Can we think of any empirical data, let alone experimental research, able to support the theory of innate human qualities? Obviously, human nature itself provides the answer: No! Due to the lengthy time spans between two generations, the transmission of characteristic traits between two generations and the comparison of their development within humanity as a whole cannot be analyzed under laboratory conditions.3 Inheritance between humans seems to be a field of speculation or, at best, analogous reasoning. Twin research, the main field for studying inheritance in humans today, was introduced only in 1893 by Francis Galton. Before him, one of the central areas of research in the 1 2

Cf. Baecker (2000), p. 44. Rousseau ([1755] 1992), p. 13.

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19th century, heredity in human individuals or races, was not able to meet the standards of the main epistemological paradigm of the time: the experiment. There seems to be one exception, and Rousseau – after admitting that not even natives from the Carribean islands qualify as “noble savages” because they too tend to live in communities – mentions it in a footnote of his Discours.4 The only possible way to address the problem of human heredity by using – however extremely restricted – empirical data is research on the cases of so called ‘wolf children’ or ‘savage children’ that were documented numerous times in the 18 th and 19th centuries. Referring to rumors about the deliberate isolation of children by pharaoh Psammetich, the Mongolian king Akbar, or emperor Friedrich II, as well as to the early example of the wolf boy of Hessia (1344), authors report on Wild Peter of Hanover (1724), MarieAngélique de la Champagne (1731), Victor de l’Aveyron (1800), Kaspar Hauser (1828), and the wolf children of Sultanpur (1843, 1848), to name only the most prominent cases before Mendel, Darwin and Galton.5 As the natural historian Johann Friedrich Blumenbach wrote in his chapter on Wild Peter’s growing “Celebrität” in the middle of the 18th century: “Sie traf in die Zeit, wo gerade der Streit über die Frage: ob es angeborene Begriffe gebe, mit voller Lebendigkeit und respective Hitze geführt ward. Und da schien Peter ein erwünschtes Subject zur Entscheidung derselben.”6 In this paper, I will analyze one of these case studies in order to demonstrate how the philosophical discourse on heredity at the end of the 18th century implemented, for the first time, the method of the new experimental sciences of the same period. Earlier reports about wolf children had not yet addressed the problem of heredity. Instead, they were interested in the kinship and differences between humans and animals, and savage children seemed to be a kind of hybrid ‘in-between-state’, or ‘monster’, in Foucault’s sense of the word. 7 Nevertheless, Linnaeus subsumed the homo ferus as a subspecies in his classification and mentions nine examples of such homines feri in the twelfth edition of his Systemae Naturae from 1766. He attributes Latin classification names to each one of them (Juvenis lupinus hessensis, Puella campanica, etc.), and points out their major traits: they are unable to speak, covered with fur, and walk on all fours.8 Obviously, from this description, wolf children seem to be more closely related to the animals, who had been their companions, than to the human beings, who were their parents. Transferred to a model of inheritance, this implies that it was their way of life rather than their genealogical descent that influenced the faculties as well as the place within nature’s order of these children. It 3

4 5

6 7

Müller-Wille (2002), p. 14, shows in a similar sense that the synthesis of empirical data from single case studies into a theory of heredity posed a problem “that could not be solved in anthropology”. The field for experimental studies in heredity was restricted to plants and minor animals. For the relation between speculation and experiment in 18th century theory of human heredity, see Terral (2002). Maupertuis’ studies on inheritance in families that Terral mentions (pp. 36f.) are not experimental insofar as they do not enable Maupertuis to manipulate the objects of observation. Maupertuis’ suggestions for actual experiments in human-animal-hybridization were not carried out. Rousseau ([1755] 1992), pp. 68-70. For a survey on all reported cases cf. Singh/Zingg (1942), and Malson ([1964] 1972), pp. 62-82. Seitter (1986), pp. 45ff., points out the intrinsic power-relation within this attempt to create knowledge about humans. Blumenbach (1811), p. 18. Cf. Douthwaite (1997). Clark (1948), p. 42ff., reckons the case studies on wolf children in the 18th century, focusing mainly on the connection between men and apes, among the “precursors of Darwin.”

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was only at the turn of the 19th century that this focus on the natural history of man was replaced by the focus on human beings as objects of experimental science. This shift also redefined the image of wolf children as well as the significance that was attributed to them with regard to the question of human heredity. In the chapter of their Practical Education from 1798 that deals with language teaching, Maria and Richard Edgeworth report about an experiment they conducted with Wild Peter: In 1779 we visited him, and tried the following experiment. He was attended to the river by a person who emptied his buckets repeatedly after Peter had repeatedly filled them. A shilling was put before his face into one of the buckets when it was empty; he took no notice of it, but filled it with water and carried it homeward: his buckets were taken from him before he reached the house and emptied on the ground; the shilling, which had fallen out, was again shown to him, and put into the bucket. Peter returned to the river again, filled his bucket and went home; and when the bucket was emptied by the maid at the house where he lived, he took the shilling and laid it in a place where he was accustomed to deposit the presents that were made to him by curious strangers, and whence the farmer’s wife collected the price of his daily exhibition. It appeared that this savage could not be taught to reason for want of language.9

In using Peter’s lack of abstract reasoning as proof for Condillac’s anthropological claim that man can have no ideas without words, the Edgeworths consider the wild boy – however hopelessly impaired – a representative of humanity. Ever since, savage children who grew up isolated from any social influence were regarded as “‘experiments with nature’” 10 – experiments that were supposed to answer the question as to which of our faculties were ‘human’ from the core: reason, language, memory, consciousness of the self, morality, social skills? 11 If any of these features were found in children that had never experienced any kind of educational ‘input’, they could rightfully be called part of the basic and general heredity of humanity. The traits that the children lacked, however, had to be attributed to society, culture, and education alone. Instead of stating human nature as Rousseau had done, experimental anthropologists now began to question and test its genealogy. And instead of trusting in the god-given innateness of human faculties, the possible influence of education becomes an important element within the theories of the progressive sciences.12 Of course, the proclamation one chose to follow – Rousseau’s ‘Back to Nature!’ or Helvetius’ ‘Ahead towards Education!’ – depended on specific 8

9 10 11

It is precisely the last aspect Rousseau ([1755] 1992), pp. 69f., refers to in his footnote mentioned above, claiming however that the evidence of a few human beings using their hands for the purpose of walking because of a different upbringing does in no way contradict the natural law according to which humans are meant to walk upright. Jean-Pierre Bonnaterre, to whom I will return later, uses the example of Victor de l’Aveyron to prove Linnaeus wrong as far as the traits of furriness and animal-like walking are concerned. However, it is still within this context that the third feature, the language acquisition, is going to be the experimentum crucis of Victor’s treatment. Edgeworth and Edgeworth ([1798] 1815), pp. 61f. Malson ([1964] 1972), p. 49. It is the interest of this examination itself that makes it a “forbidden experiment” for Shattuck ([1980] 1994), p. 41. Shattuck hints at the tradition of deliberately isolated children as well as at the modern sciences that emerged from discourses like the ones circulating around wild children. “What I call the forbidden experiment is one that would reveal to us what ‘human nature’ really is beneath the overlays of society and culture. Or at least an experiment that could tell us if there is any such thing as human nature apart from culture and individual heredity.”

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interests. In the same way, interpretations of case studies on wolf children are – up to today – structured according to current scientific ideologies. Just as in the late 18 th century wolf children were used as proof for the opposing claims that there is both an essence and an absence of human nature, 20th century research has been split on this point according to the ruling paradigm: The rise of molecular biology in the middle of the last century (as well as its recent success) established the belief that there are genetic patterns that make us human. The foreword of the report on the wolf children of Midnapore reads: It must also be pointed out that although extreme distortions can be produced, both mentally and physically by unusual elements in the environment, it by no means follows that heredity is eliminated either under the normal or the abnormal condition. On the contrary, genetic experiment makes it clear, that heredity is at work in both cases, and that the fundamental bases even for quite small physical and mental differences are determined by heredity, however they may happen to be distorted or suppressed in an unusual environment.13

In the 1970s, however, when behaviorist approaches and social theories dominated, scholars dealing with wolf children rejected any notion of human faculties developing without environmental influence and adaptation pressure: “If any further proof is needed that the term ‘human nature’ is completely devoid of sense, the following study of wolf children will provide it.”14 In any case, these examples show that wolf children are subjected not only to the scientists experimenting on them, but also to the various interpretations of these experiments, and I will compare some of these experiments and interpretations in the following pages. But in doing so, I will refer not merely to philosophical or scientific sources. The appearance of a savage child is not only a spectacular event for scientists, it is also ‘spectacular’ in the core meaning of the word: it is a sensation, an attraction, public entertainment. In most reports on wolf children – be it Peter at King George’s court, Victor in Rhodez, or Kaspar Hauser in Nürnberg– it is pointed out that they spend the first days in society like animals in a zoo or, for that matter, like the native inhabitants of European colonies, who were put on display at fairs, in vaudeville acts, and at world exhibitions later in the 19th century. But it is not just the wild child who is a spectacular phenomenon. The new methods of his examination are, too. As Barbara Maria Stafford has shown, the early 19 th century demonstrations

12

13

14

It is important to note that both attitudes are part of enlightenment thought: Rousseau’s claim of the innate nobility of every human (”nature is everything”) constitutes the idea of equality, whereas Helvetius’ belief in the significance of education follows from the principle of progress and perfectibility (”education is everything”) – a belief that was not at all alien to Rousseau himself, as I shall point out below. Singh/Zingg (1942), p. xvi. Cf. from today’s point of view Pinker (1994), p. 282: “The muteness of wild children in one sense emphasizes the role of nurture over nature in language development, but I think we gain more insight by thinking around the tired dichotomy. If Victor or Kamala [the girl from Midnapore] had run out of the woods speaking fluent Phrygian or [the assumed first language] ProtoWorld, who would they have talked to?” Malson ([1964] 1972), p. 35. Malson claims that there is no human life outside of society and therefore no such thing as a “pre-cultural state”, either. Wolf children, who missed the time-window for language acquisition, demonstrate neither typical species-behavior nor are they in any way similar to the myth of the ‘noble savage’.

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of experimental science were received like magic shows in front of stunned audiences. 15 Moreover, the exotic and spectacular dimension of experiments on wolf children is met by a response to these discoveries that can be found within popular culture: Theatre-presentations, cabaret-shows, and novels regularly took up on and popularized the observations made in anthropological case studies.16 The scenarios of these popular performances did not necessarily respond to actual discoveries, but sometimes even preceded them. In his analysis of the various documents, texts, and discourses that paved the way and accompanied the discovery of Kaspar Hauser, Jochen Hörisch states: “Tatsächlich entspricht sein Nürnberger Erscheinen nur den ihm vorausliegenden diskursiven Ereignissen, die es verdoppelt.”17 Wild children, who are not able to speak, enter a universe of discourse which, in this sense, always precedes them. The constellation of scientific experiments on heredity and the popular discourse on human nature is precisely what interests me here: A cultural history of heredity must take into account the discourses that accompany and structure a certain scientific practice. Such discourses, as is the case with both empirical and fictive commentaries on wolf children, present conceptions and reconceptualizations of what is considered to be ‘human’ at a certain time. 18 Around 1800, the question of heredity is an anthropological, physiological, linguistic, psychiatric, and pedagogical one and therefore cannot be answered by any of these disciplines alone. What I offer in the following is therefore an analysis of three different discourses on the isolation of humans and experiments that attempt to undo the effects of this isolation. These three discourses present three possible solutions to the problem of whether human faculties develop by hereditary or educational means – or both. My analyses of a novel, a case study, and a scientific treatise will also show, however, that the notion of ‘heredity’ that circulates within a number of discourses at the turn of the 19th century is very different from the current scientific notion. While striving to be experimental, it nonetheless remains a ‘macro-discourse’ on ‘human nature’ and ‘humanity’ as a whole, instead of being an analysis of individual traits, their genealogy and inheritance. Only with the historicization that Foucault pointed out for the knowledge systems of the 19 th century was the synchronic concept of innateness replaced by the modern concept of heredity.

2. Isolation and education: the experimental paradigm of anthropology around 1800 Before I turn to the analysis of the three examples for a discourse on heredity concerning wolf children, I shall elaborate briefly the notion of experimentation within these studies. One of the main features that actually connects the experimental approach and the discovery of savage 15 16

17

18

Cf. Stafford (1994). Daston (1998) argues that the description of ‘strange facts’ – among others monstrous appearances – is the take-off of modern natural philosophy in the 17th century. Cf. Gineste (1993), pp. 29-32. Malson ([1964] 1972), pp. 37f., emphasizes the mingling of fact and fiction in any of the reports on wolf children insofar as they “involve not only metamorphoses of nature but human dramas as well. One should hesitate, therefore, before rejecting a story just because part of it is invented. Hoaxes here are as common as they are anywhere else: every so often the press reports the discovery of yet another Mowgli waiting for his Kipling”. In the same way, Lane ([1976] 1977), pp. 185ff., stresses that a case such as the Wild Boy of Aveyron’s produces “a legend, a tale of epic proportions”. Hörisch (1979), pp. 270f. Since discursive signifiers precede Kaspar’s actual appearance, to Hörisch Kaspar Hauser marks the switch from the subject-centered paradigm of enlightenment to the modern semiotic paradigm. On the popular “re-writing” of the savage to this end, cf. Douthwaite (1994-95).

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children is the element of ‘isolation’ that, when taken as a discursive concept, seems able to combine various approaches at the end of the 18th century. On the one hand, both the rise of the experimental sciences and the emphasis on controlled observation require the elimination of unwanted influence and the reduction of any distracting context – in a word, they require the isolation of the subject of observation; we have learned to call the realm of this experimental isolation the ‘laboratory’.19 On the other hand, but in a structurally very similar manner, Rousseau’s ‘natural state’ of humans is emphatically situated opposite any kind of ‘social state’. Primarily, this opposition merely serves the function of hinting at the ‘original’ qualities of human beings before their cultural and societal transformation. But on a different level, the opposite of ‘being social’ is, obviously, ‘being alone’, and there are several passages within Rousseau’s Discours that struggle with the logical consequences of his argument: We have to picture the ‘natural man’ per definitionem as an isolated man, because any contact with another human being establishes the basic structures of a community and all the mischief it causes, such as possession, inequality, and effemination. According to Rousseau, as long as man is alone, he lives without knowledge of good and bad and the feeling of want. There are only two elements that distinguish him from being a mere animal: the instinct of compassion (responsible for his pre-moral ‘nobility’) and the perfectibility of his reason (responsible for his eventual abandonment of the state of nature and communitybuilding – the first step of the depravation of mankind). These two faculties – for Rousseau the only innate ones – contribute to a twofold characterization of human nature: On the one hand, Rousseau attributes a basic dignity to every human being – especially the wild, natural, and uneducated. On the other hand, he admits the possible education of human beings. Though he paints a distinctly pessimistic picture on the outcome of education in his Discourse, at the beginning of the fictive pedagogical experiment on Émile he admits: “Under existing conditions a man left to himself from birth would be more of a monster than the rest.” 20 Thus, the principle of isolation and the principle of education do not exclude each other. Rather, at the end of the 18th century, they both contribute to a common discursive matrix that is especially successful within revolutionary France. The revolution was based on Rousseau’s idea of the original equality of man. The privileges that used to come with a noble descent were replaced by the notion that anybody had a natural right and chance to lead a good life, regardless of his birth. Conversely, there were various proposals to re-educate the dethroned nobles and turn them into ‘natural’ humans. The attempt to give the dauphin a ‘civil’ education after the execution of his mother, Marie Antoinette, or the scenario of Sylvain Maréchal’s Le jugement dernier des rois (1793), in which a group of royals is stranded on a lonely island and forced to cope with the law of nature, equally express the pedagogical optimism that accompanies the belief in the natural state of mankind.21 The constellation of isolation and education within an experimental paradigm that aims to produce insights into human nature can thus be considered a basic idea within late 18 th century 19 20 21

For the emergence of the experimental paradigm and its socio-historical contexts – especially the modern discourse of political control – see Shapin and Shaffer (1985). Rousseau ([1762] 1911), p. 5. Cf. Gumbrecht ([1981] 1992), pp. 206-211. Sanja Perovic is currently preparing a dissertation on the trial of Marie Antoinette and Maréchal’s plays at Stanford University.

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anthropology. The latter term seems to indicate the new type of science that comes along with this aim, but it is interesting to see how differently it is used: Whereas in the German tradition ‘anthropology’ means to reason on human nature on a conceptual level, the French (as well as the English) notion of ‘anthropology’ is a much more empirical one, often referring to ethnological and experimental research.22 In 1799, the Société des Observateurs de l’homme is founded, declaring human experiments as the main method of the science of man. In his opening address, one of the founding members, Louis-François Jauffret, names the aims of the society as follows: “Tatsachen zu sammeln, die Beobachtungen zu erweitern und zu vermehren und dabei alle leeren Theorien, alle verwegenen Spekulationen beiseite zu lassen.”23 The society’s interests concerned the physiological, intellectual, and moral nature of humans, and the two main research projects Jauffret proposed are firmly grounded within the discursive scenario unfolded so far: One is called Considérations sur les diverses méthodes à suivre dans l’observation des peuples sauvages, but it is the other in which Jauffret really invests his hopes, and it is worth quoting in its entirety: Eines Tages muß die Gesellschaft wohl prüfen, ob es, um die fortschreitende Entwicklung der physischen, intellektuellen und moralischen Fähigkeiten des Menschen auf eine so neue wie umfassende Weise zu verfolgen, nicht günstig wäre, mit Genehmigung der Regierung ein Experiment über den Naturmenschen zu unternehmen, das darin bestünde, während zwölf oder fünfzehn Jahren vier oder sechs Kinder, zur Hälfte männlichen, zur Hälfte weiblichen Geschlechts, sorgfältig zu beobachten, nachdem man sie von Geburt an am selben umfriedeten Platz, fern jeder gesellschaftlichen Einrichtung ausgesetzt und die Entwicklung ihrer Ideen und ihrer Sprache dem natürlichen Instinkt überlassen hätte. Zweifellos erhielte man zahlreiche außerordentlich nützliche Beobachtungen, um uns mit Sicherheit über die Entwicklung unserer Fähigkeiten aufzuklären, indem die Philosophie einige von Geburt an von unseren Sitten, unseren Einrichtungen, unseren Vorurteilen und sogar von unserer Sprache getrennte Kinder beobachtete, die nicht anders als auf Grund des allen Menschen gemeinsamen natürlichen Instinkts und Naturzustandes handeln und sich ausdrücken würden.24

Although such an experiment would demand the “sacrifice of an entire life” – the life of the researcher, not of the research subject, nota bene – it would enable science to solve “die schwierigen Probleme der Entstehung der Sprachen und der Ideen überhaupt sowie der Grundbegriffe des menschlichen Geistes” for the first time.25 Jauffret refers to the historical precedent of such an experiment mentioned above – what he really seems to refer to, however, is the discursive matrix that connects insights into the human’s true nature with the examination of his isolated state. In his analysis of the emblematic architectural design of the new scientific approach to humans, Jeremy Bentham’s prison that allowed permanent surveillance of the inmates, Michel Foucault suggests the same connection:

22

23 24 25

There is a short period of an empirical approach to anthropology in Germany, too, connected with the names of Platner, Wezel, and Moritz who develop a systematic notion of ‘observation’ within the sciences of man. I tackle this approach in my current book project, Fallgeschichten. Die literarische Anthropologie des Menschenversuchs. Cf. also Eckardt, e.a. (2001). Quoted from the German translation of Jauffret’s address in Moravia ([1970] 1973), p. 209. Ibid., p. 215. Ibid., p. 216.

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But the Panopticon was also a laboratory; it could be used as a machine to carry out experiments, to alter behavior, to train or correct individuals. […] To try out pedagogical experiments – and in particular to take up once again the well-debated problem of secluded education, by using orphans. One would see what would happen when, in their sixteenth or eighteenth year, they were presented with other boys or girls; one could verify whether, as Helvetius thought, anyone could learn anything; one would follow ‘the genealogy of every observable idea’; one could bring up different children according to different systems of thought, making certain children believe that two and two do not make four or that the moon is a cheese, then put them together when they are twenty or twenty-five years old; one would then have discussions that would be worth a great deal more than the sermons or lectures on which so much money is spent; one would have at least an opportunity of making discoveries in the domain of metaphysics.26

As long as the Société lacked both a government grant and suitable experimental subjects, the only way to realize Jauffret’s proposal, however, was fiction. As the example of Maréchal’s play shows, literature at the end of the 18th century reinvestigated the topic of isolation on lonely islands that Defoe’s Robinson Crusoe had introduced.27 And it is not by chance that novels were the primary formal vehicle: As an art form that is produced and received in the lonely states of reading and writing, it seems to present the best possible solution for dealing with the question of human behavior under perfectly isolated circumstances. The foundling topic in the first half of the 19 th century – e.g. in Heinrich von Kleist’s Der Findling, Walter Scott’s Legend of Montrose or George Sand’s François le champi – is one example of this. It is important to note, though, that the renaissance of the genre takes place earlier, contributing to the discursive matrix of isolation and education in post-revolutionary France as well as allowing the staging of experiments hardly possible in reality.

3. Fanfan et Lolotte: case study on a lonely island In 1797, François-Guillaume Ducray-Duminil published a popular novel that will serve as my first example for the common discourse on isolation and education within the emerging practice of experiments on human heredity though it was published before any such experiment was actually conducted. Fanfan et Lolotte is the story of two English children who were washed up on a desert island in the Carribean at the age of three and manage to survive the fierce conditions and the proximity of native inhabitants until they are discovered four years later by the person who was to become their enlightened teacher of civilization. The Carribean setting shows how close at hand Rousseau’s vision of the “noble savage” was to any scenario set to examine the ‘natural’ faculties of humanity as opposed to the ones provided by culture. The two children from England are experimental subjects who are isolated but still dispose of a background of Western culture. At the same time, lacking any instructions from Western culture, the natives on the surrounding islands are forming communities, while living much closer to a natural state. The ‘scientific’ question addressed by Ducray-Duminil’s fictive case study is obvious: Will the children turn out to be savages, reduced to their natural faculties? Do 26 27

Foucault ([1976] 1977), p. 204. For the connection of Robinson Crusoe as well as the social experiment reported in Schnabel’s Insel Felsenburg with the scientific debate of the time, cf. Campe (2002), pp. 188-276.

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they possess such faculties at all or will they be dependent on the native’s help? Or will they instead be able to develop some of the basic principles of the culture from which they stem? Although Lucy Peacock, who translated Fanfan et Lolotte into English, omitted from the original many passages supposedly unsuitable for younger readers, she fully subscribes to DucrayDuminil’s experimental interest in the relation between human nature and the influence of education that can be analyzed in isolation. In her preface she describes the topic of the novel as “the native feelings of the heart unadulterated by vice, docility and industry of two children abandoned to themselves at an early age, the lessons of a watchful and enlightened preceptor.” 28 This “preceptor” is the shipwrecked Colonel Carlton who discovers the children. At first, they take him to be their long lost father and express hope that now finally they will be “loved” – a first hint that there are indeed basic emotions that can be developed without instruction. But apart from that, the children speak a “jargon he could just distinguish to be English” 29 and know nothing about their family and home country. The Colonel was astonished to conceive it possible that two children, so young and so delicate, should provide for all their wants at an age when others scarcely know how to walk or think: he wished to discover by what means this miracle had been effected, but they expressed their ideas very imperfectly.30

Here, the novel presents an issue that we will encounter time and again in the debate on human heredity in savage children: Do isolated children possess natural faculties, or is education necessary for their development? Peacock’s translation of Ducray-Duminil’s novel demonstrates that both alternatives are able to coexist. Carlton finds that his “pupils of Nature” 31 are normally developed as far as their “heart” and their “intelligence” are concerned. 32 They established a Robinson-like calendar and expressed basic religious feelings towards the sun – two cultural phenomena developed on their own. Also, they greedily eat the meat the Colonel offers them, although they never had it before and – as evidenced by Victor de l’Aveyron and Kaspar Hauser – should detest it. At the same time, the Colonel decides that the children nevertheless need further instruction: “It is time, my children, that I think of the great work to which God has appointed me, that of your education.”33 He teaches them to bury the dead (they had kept the corpse of their sole travel companion in their hut for over four years), to build a house (in which the bedrooms are separated by sex), and to tell their story (which he takes as evidence for the providence of God, who provided the children with the natural instincts that helped them survive). After a few days, the acculturation of the savage children is perfect: The Colonel takes the boy hunting, while the girl is “employed in baking the bread or other little offices of domestic oeconomy.”34 The goal of Western education seems to be the separation of the sexes. The report 28 29 30 31 32 33 34

[Ducray-Duminil (1797)]/Peacock (1807), p. iii. Ibid., p. 3. Ibid., p. 6. Ibid., p. 10. Ibid., p. 31. Ibid. Ibid., p. 51.

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about the progress also makes a clear statement on the relationship between heredity and culture in humans: human beings are born with innate faculties sufficient to keep them alive and to develop the basic moral and religious ideas that distinguish them from animals. But they need further education in order to develop higher cultural qualities that distinguish them from the savages who visit the island every now and then. Carlton’s education serves the cultural development of humanity whereas his observations of the children prove that the biological development is provided by nature. The Colonel teaches his charges farming, gardening, reading, writing, and religion. Thus, the novel suggests that the evolutionary elements of humanity can be repeated and taught to single individuals in the very same order they developed on a macroscopic level. Consequently, the last lesson the children learn is politics: “Men living in society, my son, said the Colonel, in which there are good and bad, it is necessary to establish rulers, that order and peace may be maintained.” 35 Witnessing the children’s natural detest of his mentioning the bad, the Colonel observes how much better these “rational and attentive” children of nature are compared to the “depravity of human nature” in society.36 The fictive experiment on isolation in Fanfan et Lolotte reinforces Rousseau’s conviction of the good nature of humanity that can also be detected in the natives of the neighboring islands: They do not like to be called savages; this name, they say, belongs to beasts of prey; they equally detest that of cannibal, which is with them equivalent to man eater; but they are very fond of the title of Carib, because in their primitive language this word signifies a good warrior, a courageous man.37

At the same time, the novel emphasizes the importance and necessity of an education that develops the higher faculties within humans. In a way, Fanfan and Lolotte, who are confronted with the civilized world on a journey at the end of the novel, combine the better elements of both human heredity and education: They lead a natural as well as a cultural life – both without the depravations of modern society.

4. Victor de l’Aveyron: idiot or pupil? Ducray-Duminil’s novel is not merely a romantic escapist fairy tale; it contributes to the debate on human heredity and its possible observation in isolation at the time. Another of his novels is also worth mentioning here en passant. It tells the story of a boy who grew up among a band of robbers in the woods where he is found by a nobleman, who grants him higher education without depriving him of the natural instincts he acquired in his earlier life. This plot does not seem to address exactly the issue we are interested in here. However, the title of the novel that came out in 1796 – four years before Jauffret proposed his experiment to the Société – is the precise prophecy of how Jauffret’s wish would be fulfilled just a few months after his opening address. DucrayDuminil’s earlier novel was called Victor, ou l’enfant de la forêt – and after January 8th, 1800, almost everybody in Paris knew what it really was about. 35 36 37

Ibid., p. 54. Ibid., pp. 59 and 56. Ibid., p. 96.

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On January 8th, 1800, a boy of about 11 years of age, who had been seen running about naked in the woods like a wild animal on several prior occasions, entered a farmer’s workshop and started his career as the only actual experimental subject of the Société des Observateurs de l’homme. From all evidence, it seemed that the boy – who could not speak, wore no clothes, preferred smelling to his other senses, and appeared to be unable and unwilling to engage in any kind of contact with other human beings – must have spent most of his life alone in the forest, living on roots and nuts. He was the perfect enfant de la forêt,38 and although his teacher Jean Itard would later report different reasons for the name he gave his pupil, it seems only too obvious why the name he did chose had to be ‘Victor’.39 The various reports of the initial examination of the savage child from Aveyron indicate the shift of interest in human nature at the turn of the 18th century. The Abbé Pierre-Joseph Bonnaterre who is taking care of the child in Rodez reports to the Société that Victor shows “purely animal functions”: He has no knowledge, no wants, no memory, no imagination, and no moral sentiment whatsoever. “You might say his mind is in his stomach.” 40 Bonnaterre’s interest is mainly in natural history, and he uses the common comparison to animals in order to classify the savage child. The two experiments he conducts on the boy emphasize this interest: Bonnaterre strips his charge naked, exposes him to temperatures below zero Celsius and notes with amazement that the boy “appeared glad indeed to be rid of these garments.” 41 On another occasion, he tests the ability of the “child of nature” to share food with others and observes: “[H]e has no idea of property, wants everything for himself, because he thinks only of himself.” 42 The Abbé concludes his report with a far-seeing remark: Such an astonishing phenomenon will furnish philosophy and natural history with important ideas about the essential nature of man and the development of his intellectual faculties, provided that the state of imbecility we have noticed in this child places no obstacle in the way of his instruction.43

The two options presented here – a possible education or the diagnosis of idiocy – are precisely the ones the wild boy from Aveyron faces when he is brought to Paris on request of the Société. Two 38

39

40

41 42 43

There was, in fact, a novel that dealt with the case of Victor de l’Aveyron itself: Lane ([1976] 1977), p. 352, refers to a book by J.A. Neyer: Rodolph ou le sauvage de l’Aveyron. Paris: Jouanaux 1800, that I could not track down in any library. Cf. Gineste (1993), p. 482: “Ce roman n’a pas été retrouvé. L’annonce de son publication figure pourtant dans les Journal général de la littérature française du 2 Octobre 1800 (10 vendémiaire an VII): ‘Rodolphe ou le sauvage de l’Aveyron’ par J.A. Neyer, auteur des Extraits d’Hervey etc... vol in 18, fig. Prix 60c et 75c. franc de port. Paris, Garnier impremeur, rue du Gd. Hureleur no 5; et Sombert lib., boulevard Saint-Martin’.” Itard ([1801 and 1964] 1972), p. 119, does not refer to Ducray-Duminil but writes instead he chose the name because the vowel ‘o’ was the only one the boy reacted to – a hardly convincing explanation, considering the very different vowel the name Victor begins with. One should think that names such as Jean or Georges should have been more appropriate. The assumption that Itard’s choice was influenced by Ducray-Duminil’s novel and its adoption for theatre is also expressed in Shattuck ([1980] 1994), p. 92. Quoted from the first English translation of Bonnattere’s Notice historique sur le sauvage de L’Aveyron et sur quelques autres indivus qu’on a trouvés dans les forêts à différentes époques (Paris: Pancoucke 1800) in Lane ([1976] 1977), pp. 35-54, here pp. 41 and 42. Ibid., p. 49. Ibd., p. 47. Ibd., p. 53.

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of the leading Observateurs represented the two different approaches the savage child was confronted with: On the one hand, the Abbé Roche-Ambroise Sicard, principal of the first school for deaf and dumb children in Paris, one of the most progressive teachers of his time. On the other hand, Philippe Pinel, the renowned reformer of mental hospitals, who had been systematizing diagnostic methods for the retarded. The wild boy’s trip from Bonnatere’s Rodez to Pinel’s Paris is also the journey from natural history to natural science. Whereas Bonnaterre had likened the boy to animals, Pinel, in his report to the Société from 1800, added a different comparison: “[H]e is much inferior to many individuals who are locked away in our hospitals. I should not be afraid to say that in this respect even elephants have a marked advantage over him.” 44 The imbeciles Pinel knows so well show “clear parallels with that of the child of Aveyron”. Like him, they are mute, emotionally unstable, without memory, etc.45 But more important than the parallels is the verdict that follows immediately from the chosen comparison: Since there is no hope of ever educating or integrating the wild boy into society, he ought to be locked away with his fellow sufferers at Bicêtre. Obviously, both Bonnaterre’s and Pinel’s diagnoses make an implicit statement on the hereditary outfit of the boy from the woods. Bonnaterre’s approach is still part of the 18 th century’s tradition of dealing with homines feri: his experiments attempt to discern the relation between the boy’s behavior and animal behavior. Pinel, on the contrary, replaces this approach with clinical diagnosis46 and an interest in the boy’s relation to society rather than nature. The question he raises with regard to heredity now reads: “finding out the nexus of ideas and moral sentiments which are independent of socialization.”47 Since he is unable to find this nexus, Pinel concludes that he is not a suitable subject for further experimentation. 48 At this point Jean Itard came on the scene, protesting Pinel’s view vehemently without siding with Bonnaterre. Itard, who shortly after Pinel’s report took the boy into his household and kept him there for six years, conducted one of the first and longest systematic educational experiments and simultaneously introduced a new approach to human nature: He agreed with Pinel that Rousseau’s belief in the ‘nobility’ of savage humans had to be rejected. Obviously, the intimidated, 44

45 46

47

As in the case of Bonnaterre, I quote Pinel’s Rapport fait al la Société ddes Observateurs de l’homme su l’enfant connu sous le nom de Sauvage de L’Aveyron (1800, deuxieme partie 1801) from Lane ([1976] 1977), pp. 64-79, here p. 66. Lane discovered the report which, until than, had been thought missing in a Journal of Anthropology from 1911. The original text (with a different date) is now available in Gineste (1993), pp. 249-260 and 271-278. Quoted from Lane ([1976] 1977), p. 69. Cf. the title of Gineste (1993), Dernier enfant sauvage, premier enfant fou, as well as Douthwaite (1997), p. 191: “Earlier, the wild child had been embraced as an example of nature, but by the 19th century his antisocial habits and insensitivity gave rise to claims of mental pathology.” Quoted from Lane ([1976] 1977), p. 64. Authors at the turn of the 19th century even extended this diagnostic approach to earlier examples. Cf. Edgeworth and Edgeworth ([1798] 1815), p. 61: “Peter, the wild boy […] had all his senses in remarkable perfection. He lived at a farm house within half a mile of us in Hertfordshire for some years, and we had frequent opportunities of trying experiments on him. He could articulate imperfectly a few words […]; he could in a rude manner imitate two or three common tunes, but without words. Though his head […] resembled that of Socrates, he was an idiot: he had acquired a few automatic habits of rationality and industry, but he could never be made to work at any continued occupation.” Blumenbach (1811), pp. 26f., comments on the same case: “Kurz als Ende vom Lied, das vermeinte Ideal des reinen Naturmenschen, wozu spätere Sophisten den wilden Peter erhoben hatten, war durchaus nichts weiter, als ein stummer, blödsinniger Tropf.” That is the reason why “von dem blödsinnigen Buben für Psychologie oder Anthropologie eben keine bereichernde Ausbeute zu erwarten sei” (ibid., p. 22).

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spastic, and autistic child did not provide much evidence that human beings develop best in a natural state. What he refused, moreover, this time contrary to Pinel, was the belief in the innateness of the boy’s defects readily observable in the wild boy, to whom he will, as a first step toward humanization, give a name. To Itard, the fact that Victor was a child of nature proved all the more greatly how much the development of our faculties depends on training. Drawing both from Condillac’s theory of sensory education and from Sicard’s success with the deaf and dumb, Itard planned to prove this theory by turning Victor into a well-educated, conversational, social being. Should he succeed, he would have evidence for his basic hypothesis on human heredity – or non-heredity, for that matter: Cast on this globe, without physical powers, and without innate ideas; unable to obey the constitutional laws of his organization, which call him to the first rank in the system of being; man can find only in the bosom of society the eminent station that was destined for him in nature, and would be, without the aid of civilization, one of the most feeble and least intelligent of animals – a truth which, although it has often been insisted upon, has not as yet been rigorously demonstrated.49

5. Itard’s sincere observations: laboratory nature Itard’s two case studies on Victor from 1801 and 1806 will serve as my second example for the discourse on isolation, education, and experiment surrounding the concept of heredity at the turn of the 19th century. It is important to note that Itard, despite his refusal of the Société’s recommendation to leave Victor in the asylum, still shared the basic methodological approach sketched out by Jauffret. After the news about the wild boy had spread from Aveyron to Paris, Jauffret wrote a letter to the orphanage of Saint-Afrique where the spectacular discovery was initially kept: If it is true that you have currently in your orphanage a young wild boy, twelve years old, who was found in the woods, it would indeed be important for the progress of human knowledge that a zealous and sincere observer take him in charge and, postponing his socialization for a little while, examine the totality of his acquired ideas, study his manner of expressing them, and determine if the state of man in isolation is incompatible with the development of intelligence.50

This “zealous and sincere observer” turned out to be Itard. Whether he was in fact “postponing” Victor’s progress cannot be decided retrospectively. In any case, he shared the experimental attitude implied in his proposal: to design a realm in which a savage child could be fruitfully 48

49

50

Cf. Moravia ([1970] 1973), p. 116: “Nachdem man den Knaben, der einer der sensationellsten wissenschaftlichen und kulturellen Fälle zu Beginn des neuen Jahrhunderts gewesen war, vom ‘Wilden’ zum ‘Idioten’ deklassiert hatte, verlor er für die spezifischen Interessen der ‘Société des Observateurs de l’homme’ an Bedeutung.” Itard ([1801 and 1964] 1972), p. 91. Itard’s first report was published as De l’éducation d’un homme sauvage ou des premiers développements physiques et moreaux du jeune suivage de l’Aveyron with Goujon in Paris. Quoted from Lane ([1976] 1977), p. 15. Lane also quotes various contemporary newspaper articles that express hopes for ‘pure’ laboratory conditions for the study of human nature in Victor (ibd., p. 20).

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observed. As Itard notes at the beginning of his first report, this was a rare opportunity that had remained unused on all earlier occasions. People were interested only in ‘seeing’ the spectacular discoveries, whereas “actual observation was reckoned of no value; and these interesting facts tended little towards improving the natural history of man.”51 Remarks like these reveal that Itard’s incentive for taking care of the child was motivated not only by his philanthropy. Or, to put it differently: it was definitely motivated by the love for humans, but this love was much more directed toward the laws of human nature and less toward individual representatives of the species. Whereas Pinel’s devastating diagnosis had at least been directed on the boy, for Itard he is but a means to gain general knowledge. 52 In order to contradict Pinel’s verdict effectively, Itard had to revise the diagnosis of the wild boy’s idiocy and trace instead back Victor’s developmental backwardness to his isolation: “[T]he Savage of Aveyron is much less a simple youth, than an infant of ten or twelve months old.” 53 His attempt to avoid Pinel’s medical rhetoric, however, drives him back into the realm of colonial projections similar to those already encountered in Ducray-Duminil: “[L]ike some savages in the warmer climates, he was acquainted with four circumstances only; to sleep, to eat, to do nothing, and to run about in the fields.”54 Itard’s principle of education entailed first tending to these basic – obviously innate – needs and thus mitigating the hostility the boy felt toward the society that ended the life he was used to in such an aggressive and irretrievable way. A further method of education Itard applied to his increasingly accepting charge, though, took no regard of any kind of need the boy might feel. Rather, Itard designed a series of experiments to verify his basic theoretical conviction, according to which human senses and understanding are not innate but – as Locke and Condillac had claimed – subject to experience and training. When he first met the boy, he had no sense for heat and cold, no fear of anything, and no sense of justice. Itard, consequently, exposes him to warm baths, extreme height, calculatedly unfair treatment and many more stimuli intended to awaken his dormant senses. If we summarize, rather than enumerate these various experiments, there are two general conclusions that can be drawn from Itard’s approach: The five steps of his educational program – socialization, sensualization, development of ideas and concepts, language acquisition, general education and instruction – mirror the order that Condillac had introduced in his theory of human development: first the establishing of sensation and perception, then consciousness and attention, and finally imagination and memory – the latter requiring the ability to use signs. 55 Viewed from this model, Victor’s progress is a complete repetition of the process of becoming human. Just as in Fanfan et Lolotte the process of education followed the steps of humanity’s cultural evolution, Victor’s education mirrors the order of each individual human being’s intellectual evolution as stated by Condillac. The second anthropological implication of Itard’s experimental set is closely related to the first: The experiments resulted in Victor’s growing ability to distinguish his impressions and 51 52 53 54 55

Ibid., p. 92. For a critique of Itard’s methods cf. Mannoni ([1965] 1972); Moravia ([1970] 1973), p. 102f.; and Lane ([1976] 1977), p. 104. Itard ([1801 and 1964] 1972), p. 101. Ibid., p. 103. Condillac ([1746] 2001), pp. 15-40.

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reactions: hot and cold, pleasant and unpleasant, comfort and fear, love and anger. These distinctions are the ones that ‘civilized’ and ‘socialized’ the wild boy in the first place. Itard’s experiments were a means not only of observing Victor, but also of training and shaping him. Scholarly criticism of Itard’s work so far has focused on the therapeutic nihilism expressed in Victor’s treatment that had rather produced an “experimental neurosis” 56 within the boy instead of helping him to fight his previous ones. But from a theoretical perspective what is even more interesting is that Itard’s experiments in fact produced what they were supposed to analyze: ‘human nature’. Itard’s own account of his first results does not go that far: After one year of treatment, he notes the “mental equality between the boy and the brute”57 – to which Victor had been inferior upon his arrival in Paris. The crucial distinction applied to Victor in order to measure his humanness is the distinction between ‘speaking’ and ‘non-speaking’. The question “‘Does the savage speak’?”58 is at the core of Itard’s experiments – as well as of Condillac’s and Rousseau’s theoretical treatises on the development of language in the ‘natural state’, well known to Itard. Victor could hear and was able to distinguish sounds, but he did not utter them. Still, Itard rejected the assumption of hereditary defects and attempted to prove that Victor did have the language competence necessary to express his needs and wishes: I satisfied myself of this one day by an experiment of the most conclusive nature; I chose, from amongst a multitude of others, a thing for which I was previously assured that there did not exist between him and his gouvernante any indicating sign; such was, for example, the comb, which was kept for his purpose, and which I wished him to bring to me. I should have been much mistaken, if, by disordering my hair with my hand, and showing him my hand in this state, I had not been understood. Many persons see, in all these proceedings, only the common instinctive actions of an animal; as for myself, I confess, that I recognize in them the language of action, in all its simplicity; that primitive language of the human species, originally employed in the infancy of society, before the labour of many ages had arranged and established the system of speech, and furnished to civilized man a fertile and sublime means of indefinite improvement, which calls forth his understanding even in his cradle, and of which he makes use all his life without appreciating what he is by means of it, and what he would be without its assistance if he were accidentally deprived of it, as in the case which at present occupies our attention.59

Again, it is Condillac’s description of a “language of action” that accounts for Itard’s judgment, and – just as in Condillac’s example for the development of human ideas from the acquisition first of sign and then of spoken langue – it is a judgment derived from experimental observation. Condillac, in his Essai sur l’origine des conaissances humaines, proposed a thought-experiment: “But I am assuming that two children, one of either sex, sometime after their deluge, had gotten lost in the desert before they would have known the use of any sign.” 60 He concluded that the two 56 57 58 59 60

Lane ([1976] 1977), p. 136; cf. Mannoni ([1965] 1972). Itard ([1801 and 1964] 1972), p. 104. Ibid., p. 106. Ibid., p. 126. Condillac ([1746] 2001), p. 113. Condillac also mentions a Lithuanian wolf child in his Essai. For the relation of fictive thought-experiments to scientific research, cf. Moser (1989).

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children would habitually develop signs for situations that repeated themselves, such as the search for food: These details show how the cries for passion contributed to the development of the operations of the mind by naturally originating the language of action, a language which in its early stages, conforming to the level of this couple’s limited intelligence, consisted of mere contortions and agitated bodily movements.61

In Itard’s closely related observation of an actual experiment, it is again the experiment itself that produces the anthropological feature: Humans are humans as long as they engage in language, and if Victor demonstrates sign usage, he has entered the realm of human nature. For the time being, he is at the stage of primitive man – a stage identical to the stage of the isolated children in Condillac –, but Itard planned to demonstrate by further educational experiments that the inherited faculties of the species can be acquired by the single specimen under proper circumstances.62 That Itard himself called his ensuing attempts to teach Victor how to talk a failure is mainly due to the very stable theoretical concepts that guided his observations. Victor did indeed learn to use written letters in order to denote certain objects, but Itard did not acknowledge this achievement because his concept of language requires spontaneous voiced articulations. 63 While conceding his failure, he concludes his report with the remark that there are “most important inferences relative to the philosophical and natural history of man, that may be already deduced from this first series of observations!” Itard claims that Victor is already “endowed with the free exercise of all his senses; that he gives continual proofs of attention reflection, and memory; that he is able to compare, discern and judge, and apply in short all the faculties of his understanding to the objects which are connected with his instruction.” 64 Thus, both Pinel’s rejection of Victor’s possible education and Ducray-Duminil’s Rousseauian belief in innate human qualities are repudiated. “The moral superiority which has been said to be natural to man, is merely the result of civilization.”65 Itard calls this statement on human heredity the destruction of “prejudices […] which […] constitute the most amiable, as well as the most consoling illusions of social life.”66 But it is only through his own prejudices that the experimental production of human qualities within Victor succeeded to a certain degree. The second report Itard submitted five years later at the behest of the French Secretary of State (the Société had ceased to exist two years earlier) provides even more examples for this productive 61 62 63

64 65 66

Ibid., p. 115; cf. Lane ([1976] 1977), pp. 92f. This is, if you will, the complementary view to Lamarck’s theory of heredity, according to which acquired faculties can be handed on to descendants. The “language of action”, on the contrary, is supposed to be but a deficient supplement for true human language that not only serves as communication, but also enables the development of ideas. This is why Edgeworth and Edgeworth ([1798] 1815), p. 60, call Wild Peter an idiot – precisely the diagnosis Itard wanted to avoid, though referring to the same theoretical background: Condillac. Mannoni ([1965] 1972) criticizes the way Itard subsumes all of his observations under Condillac’s principles and, by doing so, completely ignores the suffering he caused Victor – a suffering that indeed can only be read between the lines of Itard’s report. Itard ([1801 and 1964]1972), p. 137. Ibid., p. 138. Ibid., p. 140.

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use of experiments. This time, Itard immediately begins with the acknowledgement that his report is “less a story of the pupil’s progress than an account of the teacher’s failure.” 67 Victor’s language training failed whenever he was supposed to perform something beyond mere imitation. Itard, consequently, closed this case and turned to another essential realm of human nature: You have seen, my lord, how civilization awoke the intellectual faculties of our savage from their lethargy first by applying them to the satisfaction of his needs, then by extending the scope of his ideas beyond his animal existence. Your Excellency will now see the same order of development in his emotional faculties, first aroused by the feeling of need inspired by the instinct of self-preservation, then stirred by less selfish feelings, by more generous impulses and finally by some of those noble feelings which are the happiness and glory of the human heart.68

Again, in order to be successful, these “emotional faculties” must be reconsidered as the result of education rather than heredity, since Victor was emotionally completely indifferent when first found. Therefore, Itard engaged Victor in a training of his emotions, e.g. the feeling of remorse after his final attempt to escape. He is brought back and encounters his benefactor: “But when he saw that instead of going to him I stood where I was with a cold demeanour and an angry face, he […] began to cry.”69 Itard deliberately provoked Victor’s emotional reaction by staging an appropriate stimulus. The same holds true for the second example, this time aimed at Victor’s sense of justice: After Victor successfully performed an achievement, Itard refused to congratulate him but instead criticized him. As a result of this “test”,70 Victor bit Itard: I could only delight me, for the bite was a legitimate act of a vengeance; it was an incontestable proof that the idea of justice and injustice, the permanent basis of the social order, was no longer foreign to my pupil’s mind. By giving this feeling to him, or rather by stimulating its development, I had raised savage man to the full stature of moral man through the most striking of his characteristics and the most noble of his powers.71

As in the case of his sensual and intellectual training, Itard’s experiments provoked in Victor those emotional reactions claimed to be absent in the first place. This entanglement of observation mode and observation object, however, makes it impossible to distinguish between what was actually observed and what was generated by the observation. There is a general conclusion that can be drawn from this analysis of my second example: the example of a plea for education alone. The omnipresence of the experimental gaze in Itard’s pedagogical laboratory prevented the visibility of any kind of ‘natural’ processes whatsoever. Victor, the savage child, can only be analyzed as such if he is, at the same time, deprived of this genuine savagery – or, as Sergio Moravia has phrased it: “Jetzt, da der immer fluchtbereite, immer gewaltsam zurückgehaltene und gezwungenermaßen als ‘Gefangener’ lebende Victor ein Leben führte, dem das typischste Merkmal des Naturzustandes fehlte, nämlich die Freiheit?” 72 This 67 68 69 70 71

Itard ([1806 and 1964] 1972), p. 141. Ibid., p. 168. Ibid., p 170. Ibid., p. 173. Ibid., p 174.

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paradox is not an avoidable misconception, but the result of the experimental approach itself: the need to replace the wild boy’s isolation from society with Victor’s isolation from nature. Itard had to draw “Victor back into a cocoon in which life alternated between the household and the classroom, nearly abolishing all Victor’s opportunities for unstructured contact with the natural and social environment; instruction became the predominant means of learning.” 73 Thus, the question as to whether innate human faculties can be experimentally refuted cannot be answered precisely because laboratory observation fundamentally changes the situation to be observed. Contrary to Ducray-Duminil’s example, the experiment takes place in ‘culture’, not in ‘nature’ – and consequently results in the assumption that education is more significant than heredity. Indeed, the only possible experiment that could reveal Victor’s nature and hereditary outfit would bring about the deconstruction of its intrinsic observational hierarchy: Had Itard followed Victor into the woods in order to see him in his ‘natural state’, it would soon have become clear that the deficiency in coping with the environment changed sides. In the woods of Aveyron, Itard would have learned from Victor that it is less important whether one’s faculties are inherited or acquired, but imperative that they adjust to the requirements of the immediate environment, be it cultural or natural.74

6. The first principles of anthropology: Gall and Spurzheim visit the ‘pretended savage’ I will now turn to my final example for the various discourses on experiments in heredity with savage children, and this time it will be neither a fictive nor an actual case study, but a systematic scientific treatise. From the perspective of contemporary science, there was another problem with Itard’s hypotheses, aside from the methodological concerns just mentioned: they were all wrong. Parallel to Itard’s efforts, the early 19th century witnessed amazing progress in physiological and neurological research that arose from revolutionary models of physical and cognitive functions of humans. The result of these models was a renewed optimism in the possibility of explaining human nature according to its own principles and without the need to refer to external factors such as society and education. It was the beginning of the positivistic age that would in fact develop theories that were able, for the first time, to suggest a scientific model of heredity. Returning to the concept of innate human faculties, but altogether abandoning the philosophical realm in which authors such as Rousseau had been claiming this concept, observations of wolf children headed in a new direction. In 1809, Johann Christoph Spurzheim publishes a volume that summarizes his and Franz Joseph Gall’s physiological theory of the nervous system. There is no doubt that this field is connected directly to the topic we have dealt with thus far: “The first question perhaps in Anthropology is, Whence has man his faculties? Is man born indifferent; or does he come into the world endowed with determinate faculties?”75 There are two possible answers. One is the position as represented by Itard:

72 73 74 75

Moravia ([1970] 1973), pp. 103f. Lane ([1976] 1977), pp. 191f. Cf. ibid., p. 198; as well as Mannoni ([1965] 1972), p. 229. Spurzheim ([1809] 1815), p. 53.

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According to this opinion, not only man but also animals are born without determinate faculties – indifferent – as tabulae rasae or blank paper. All the instincts and aptitudes of animals, from the insect to the dog and elephant, are the effects of instruction. […] It must be answered that neither in animals nor in man does education produce any faculty whatever.76

The other one is represented by Gall’s and Spurzheim’s attempt to “demonstrate that external influences are not the cause of the internal faculties of the mind.” 77 Spurzheim mentions the physical condition of humans and their unconscious “automatic life” as an example for innate qualities. With regard to the aspects of conscious “animal life”, this evidence also seems to apply to faculties such as motion and the senses. They are obviously “determined by creation”, insofar they also can be found in animals. 78 But what about qualities specific to human beings, such as affects and understanding? They can be explained “either by external impressions, or by internal causes”. Spurzheim argues that the former, as random events, cannot account for the emergence of structures: It is certain that external circumstances must be presented, otherwise internal faculties cannot act; but opportunities do not produce faculties. Without food I cannot eat; but I am not hungry because there is food. […] How many children are exposed to the same influences without manifesting the same energy of faculties?79

But at the same time, Gall and Spurzheim are strongly opposed to the assumption of a ‘natural state’ drawing from this rejection of ‘cultural’ influences. The claim of a natural state lacks empirical evidence and is therefore absurd: “According to this hypothesis, man is made for solitude.”80 That man is never – or, for that matter, only very seldom – found in solitude is not proof for the depravation of his natural instincts, but the very result of these instincts. Spurzheim claims: “society itself is a natural institution,” following the “social instinct” of human nature. 81 This is, of course, nothing less but a complete reversal of Rousseau: the natural heredity of human beings is his striving to surpass the merely natural state. Society is part of human nature and the faculties we need to be able to live in it are not acquired but inherited. Spurzheim presents two empirical proofs for this claim. One is the example of geniuses: “Children sometimes show particular disposition and faculties before they have received any kind of instruction. Almost every great man shows in his infancy the character of future greatness. […] Nero was cruel from the cradle.”82 We encountered the other proof earlier, albeit in a different sense: 76 77 78

79 80 81

82

Ibid., p. 68. Ibid., p. 60. Ibid., p. 59. Spurzheim emphasizes: “It results from these considerations, that the comparison of man with other beings (not only with animals, but also with plants and minerals) must be admitted, and cannot be repugnant to our feelings.” Ibid., p. 62. Ibid., p. 63. Ibid., p. 64. In the same way, Blumenbach (1811), p. 44, in his reconstruction of the story of Wild Peter of Hanover, rejects any notion that children who were exposed at an early age would develop back towards an “ursprünglich wilde Stammrasse” of humans: “Überhaupt läßt sich für den zum Hausthier geborenen Menschen gar kein ursprünglich wilder Naturzustand gedenken.” Spurzheim ([1809] 1815), p. 70. Moreover: “Why are we not all men of genius” and perfectly peaceful and social behavior (p. 70) if these faculties could be acquired and taught?

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In order to prove that man acquires all his moral and intellectual faculties by education, it is asserted that savages who are found in woods, destitute of all human faculties, are like beasts, only because they have not received any education. This objection is refuted, as soon as the condition and state of these pretended savages are known. These unfortunate creatures may be referred to in two classes. Ordinarily they are wretched persons of a defective organization, with heads too large, being increased in size by dropsy of the brain; or they have heads too small and deformed. These individuals have almost always scrofula, hanging lips, a thick tongue, a swollen neck, a bad constitution in general, a wavering and unsteady gait; they are more or less completely idiots. They generally consist of persons who have been exposed and given up to care of Providence, because they were burdensome to their parents. The pretended savage of Aveyron, who is kept in the Institution of Deaf and Dumb at Paris, is an idiot in a high degree. His forehead is very small and much compressed in the superior region: the eyes are little, and lay in the orbits. We could not convince ourselves that he hears; for it was impossible to make him attentive to our calling him, or to the sound of a glass struck behind him! His attitude and manner of sitting are decent, but his head and body are incessently in motion from side to side. He knows several written signs and words, and points out the objects noted by them. The most remarkable instinct in him is the love of order. As soon as any object is displaced, he puts it in order. Such unfortunate creatures therefore are idiots, not because they have not received any education, but because they cannot be educated on account of their imbecility. It is difficult to conceive that in our populous countries, a well organised person should long wander about like a savage, without being discovered. However, if a well organised individual, who has escaped in his infancy, be discovered in a forest, though he cannot be acquainted with our manners and determinate education, yet he will manifest the essential and characteristic faculties of mankind; and such an individual, living in society, will soon imitate the manners and receive the instruction of others. The girl of Champaigne proves this assertion.83

It is interesting to see how Spurzheim, in the last sentence, replaced an inadequate experimental subject – the “pretended savage” Victor – with a more suitable one, Marie-Angélique de la Champagne84, without changing the basic principle, previously employed by Pinel and Itard: to test and observe wild children in order to draw conclusions about the natural outfit of human beings. It is stunning to see to what extent the interpretation of the observations of the same experimental subject differ from one another: What for Itard proves the influence of education on human behavior is for Gall and Spurzheim an example of the determination of the same behavior by innate faculties. Education is replaced by physiology, and physiognomic details illustrate the description of Victor, the idiot.85 For Spurzheim, there is no doubt that the “first principle of anthropology”, according to which “every special faculty is innate,”86 holds true even when confronted with the case of wild children. Individual deviations such as Victor’s prove the influence of education no more than 83 84

85

Ibid., pp. 71f. For the various discourses on Marie-Angélique within the paradigm of natural history, cf. Douthwaite (1994-95). The fact that the wild girl of Champaigne had a companion – which could well account for her ability to learn how to speak – is not mentioned by Spurzheim. Blumenbach (1811), pp. 19ff., also collects the physiognomic evidence from various reports on savage children and concludes, obviously influenced by the teaching of Gall and Spurzheim, “daß das samt und sonders naturwidrige Mißgeschöpfe waren” (p. 41). It is only at the end of the 19th century that Rauner (1885) suggests abandoning the deterministic discourse of biology on wolf children and reflecting, instead, on cultural influence.

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cultural differences. Only the use of faculties may be dependent on external circumstances and the existence of these faculties is granted by heredity. “In one word, all that man does he did at first through nature alone.”87 Both the strict opposition between Itard’s and Spurzheim’s interpretations of Victor’s behavior and the fictive combination of both view-points in Lolotte et Fanfan hint at one important observation: As soon as scientific observation takes the stage, there is nothing left in isolated children that can be called their ‘natural inheritance’. All that remains are theories, and Victor is judged far more according to the principles of these theories than according to the principles guiding his behavior. Although wolf children seem, for the first time in the history of studies on human heredity, to provide empirical material for experimental evidence, at the turn of the 19th century the relation between innate and acquired faculties result in various interpretations of the experiments on human nature. Therefore, instead of trying to add yet another interpretation to the endless and irresolvable debate on whether Victor was genetically or socially deprived, it is – as I have done here – more rewarding to take a step back and examine the circumstances under which all these interpretations arose: It is the coincidence of experimental observations and scientific ideologies that contribute to the concept of human nature at a given time. The result of this coincidence either combines heredity and education (Rousseau/DucrayDuminil), claims the sole influence of education (Condillac/Itard), or emphasizes the exclusive significance of heredity (Gall/Spurzheim). The three discourses on human heredity lead to the development of three models for a philosophy of innateness. Human faculties are: 1) the result of noble nature plus education; 2) the result of savage nature and education only; or 3) the result of only nature without education. The fact that the last model is most recent and established scientific model does not imply that it is necessarily the ‘master discourse’ on heredity in the first half of the 19th century. Despite the rise of the age of biology, all three discourses are perpetuated on the level of popular communication. George Sand’s foundling-novel François le champi begins with the author’s statement: “I have educated several foundlings of both sexes, who have turned out well physically and morally. It is no less certain, however, that these forlorn children are apt, in rural districts, to become bandits, owing to their utter lack of education.” 88 The book continues with the introductory confession of the narrator: I should like to be what the existing state of society allows a great number of men to be from the cradle to the grave – I should like to be a peasant; a peasant who does not know how to read, whom God has endowed with good instincts, a serene organization, and an upright conscience; and I fancy that in the sluggishness of my own useless faculties, and in the ignorance of depraved tastes, I should be as happy as the primitive man of Jean-Jacques’s dreams.89

86 87 88 89

Spurzheim ([1809] 1815), pp. 94 and 74. Cf. ibid., p. 83: “How indeed could the Creator abandon man in the greatest and most important occupations, and give him up to chance?” Ibid., p. 92. Sand ([1852] 1894), p. 5 Ibid., p. 18.

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And at the beginning of the main plot the miller’s wife remarks, after finding the boy who is not able to tell her about his whereabouts: “It is a pity […] that he seems to be so idiotic.” 90 As for the debate on further discoveries of savage children, the contribution of such cases to a theory of heredity is limited. When Kaspar Hauser appeared in 1828, he too became subject to experiments by his foster father Georg Friedrich Daumer. The goal of these experiments, however, was no longer the identification of the innate faculties of human nature. Instead, Daumer tested Kaspar’s magnetic abilities to detect metal and to react to gestures that had not actually touched him.91 But Daumer did not speculate as to whether these abilities are innate or result from the circumstances of Kaspar’s imprisonment, during which he had no contact with humans for over ten years. Rather, the evidence he reports was designed to reject assumptions that Hauser’s story may have been a hoax.92 Thus, a new discourse arises that addresses the issue of savage children: the legal discourse. Kaspar’s exposure is considered not an experiment in human nature, but a Verbrechen am Seelenleben des Menschen, as Anselm Feuerbach called his case study from 1832. Instead of questioning Kaspar’s natural faculties, this discourse focused on the biographical childhood that was stolen away from him. The question of heredity enters a different realm here: The main issue surrounding the discovery of Kaspar Hauser was his possible descent from the Duke of Baden. Where experiments fail to prove that savages are noble, they might at least demonstrate that nobles can be savages.

References Baecker, Dirk. 2000. Wozu Kultur? Berlin: Kadmos. Blumenbach, Johann Friedrich. 1811. Beyträge zur Naturgeschichte. Zweyter Theil. Göttingen: Heinrich Dieterich. Campe, Rüdiger. 2002. Spiele der Wahrscheinlichkeit. Literatur und Berechnung zwischen Pascal und Kleist. Göttingen: Wallstein. Clark, Robert E.D. 1948. Darwin: Before and After. The Story of Evolution. London: Paternoster Press. Condillac, Etienne Bonnot. [1746] 2001. An Essay on the Origin of Human Knowledge. Translated from the French and edited by Hans Aarslef. Cambridge/New York: Cambridge University Press. Daston, Lorraine. 1998. “The Language of Strange Facts in Early Modern Science.” In Inscribing Science. Scientific Texts and the Materiality of Communication, edited by Timothy Lenoir. Stanford: Stanford University Press. 20-38. Daumer, Georg Friedrich. [1832] 1983. Mitteilungen über Kaspar Hauser. Edited by Peter Tarkowsky. Dornach: Rudolf Geering.

90 91

92

Ibid., p. 31. Cf. Daumer ([1832] 1983), p. 28: “Auf einem Spaziergange machte ich einst im Beisein Herrn Prof. Wurms zu Nürnberg folgenden Versuch. Ich ließ ihn in ziemlicher Entfernung vor mir hergehen und sagte ihm, ich wolle gegen ihn mit der Hand herabfahren und er solle sagen, wann er etwas empfinde. Ich fragte ihn zweimal, ob er nichts spüre, so daß es schien, als mache ich hinter ihm die Bewegung, die ich unterließ, worauf er verneinend antwortete. Als ich aber wirklich, und zwar sehr schnell, mit der Hand herabfuhr, sah man in diesem Augenblick die Äußerung des Frostschauders an ihm, worauf er sich umdrehte und sagte, nun sei ich mit der Hand herabgefahren.” Cf. the documentation in Hörisch (1979), pp. 214-222.

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Douthwaite, Julia. 1994-95. “Rewriting the Savage: The Extraordinary Fictions of the ‘Wild Girl of Champagne’.” Eighteenth Century Studies 28 (2): 163-192. _____. 1997. “Homo ferus: Between Monster and Model.” Eighteenth Century Life 20: 203-221. [Ducray-Duminil, François-Guillaume. 1797. Fanfan et Lolotte.] Translated from the French with alterations, adapting it to the perusal of youth by Lucy Peacock. 1807. Ambrose and Eleanor. 3rd edition. London: Johnson & Harris e.a. Eckardt, Georg, John Matthias, Temilo van Zantwijk, and Paul Ziche. 2001. Anthropologie und empirische Psychologie um 1800. Köln/Weimar/Wien: Böhlau. Edgeworth, Maria and Richard Lovell Edgeworth. [1798] 1815. Practical Education. Volume 1. 2nd American edition. Providence: J. Francis Lipitt / Boston: T.B. Wiat & Sons. Foucault, Michel. [1976] 1977. Discipline and Punish. The Birth of the Prison. New York: Vintage. Gineste, Thierry. 1993. Victor de l’Aveyron. Dernier enfant sauvage, premier enfant fou. Édition revue et augmentée. Paris: Hachette. Gumbrecht, Hans Ulrich. [1981] 1992. “Outline of a literary history of the French Revolution.” In Making Sense in Life and Literature. Minneapolis: Minnesota University Press. 178-225. Hörisch, Jochen. 1979. Ich möchte ein solcher werden wie ... Materialien zur Sprachlosigkeit des Kaspar Hauser. Frankfurt/M.: Suhrkamp. Itard, Jean. 1801. “On the First Developments of the Young Savage of Aveyron.” Translated from the French by Joan White. In Malson ([1964] 1972). 95-140. _____. 1806. “Report on the Progress of Victor of Aveyron.” Translated from the French by Joan White. In Malson ([1964] 1972). 141-179. Lane, Harlan. [1976] 1977. The Wild Boy of Aveyron. New York: Bantam. Malson, Lucien. [1964] 1972. Wolf Children and the Problem of Human Nature. Translated from the French by Edmund Fawcett and Peter Ayrton. New York/London: The Monthly Review Press. Mannoni, Octave. [1965] 1972. “Itard und sein ‘ Wilder’.” Translated from the French by Eva Moldenhauer. In Die Wilden Kinder. Lucien Malson, et al. Frankfurt/M.: Suhrkamp. 223-245. Moravia, Sergio. [1970] 1973. Beobachtende Vernunft. Philosophie und Anthropologie in der Aufklärung. Translated form the Italian by Elsiabeth Piras. München: Hanser. Moser, Walter. 1989. “Experiment and Fiction.” In Literature and Science as Modes of Expression. Edited by Frederick Amrine. Dordrecht/Boston/London: Kluwer. 61-80. Müller-Wille, Staffan. 2002. “Cabbages, Tulips, Ethiopians – ‘Experiments’ in Early Modern Heredity.” In A Cultural History of Heredity I: 17th and 18th Centuries. Preprint 222. Berlin: Max-Planck-Institute for the History of Science. 7-25. Pinker, Steven. 1994. The Language Instinct. How the Mind Creates Language. New York: Harper-Collins. Rauber, August. 1885. Homo sapiens ferus oder die Zustände der Verwilderung und ihre Bedeutung für Wissenschaft, Politik und Schule, Biologische Untersuchungen. Leipzig: Denicke. Rousseau, Jean-Jacques. [1762] 1911. Émile. Translated from the French by Barbara Foxley. London: Dent/ New York Duton. _____. [1755] 1992. “Discourse on the Origins of Inequality. (Second Discourse).” Translated from the French by Judith R. Bush et al. In The Collected Writings of Rousseau. Edited by Roger D. Masters and Christopher Kelly. Hanover/London: University Press of New England. Sand, Georges. [1852] 1894. François the Waif. Translated from the French by Jane Minot Sedgwick. Boston: Little Brown and Company. Seitter, Walter. 1986. Menschenfassungen. Studien zur Erkenntnispolitikwissenschaft. München: Boer. Shapin, Steven and Simon Shaffer. 1985. Leviathan and the Air-Pump: Hobbes, Boyle and the Experimental Life. Princeton: Princeton University Press. Shattuck, Roger. [1980] 1994. The Forbidden Experiment: the Story of the Wild Boy of Aveyron. New York/ Tokyo/London: Kodansha International. Singh, J.A.L. and Robert M. Zingg. 1942. Wolf-Children and Feral Man. New York: Harper Row. Spurzheim, Johann Christoph. [1809] 1815. The Physiognomical System of Drs. Gall and Spurzheim; founded on anatomical and physiological examination of the nervous system in general, and of the brain in particular; and indicating the dispositions and manifestations of the mind. London: Baldwin, Cradock, and Joy. Stafford, Barbara Maria. 1994. Artful Science. Enlightenment, Entertainment and the Eclipse of Visual Education. Cambridge, MA: MIT Press. Terral, Mary. 2002. “Speculation and Experiment in Enlightenment Life Sciences.” In A Cultural History of Heredity I: 17th and 18th Centuries. Preprint 222. Berlin: Max-Planck-Institute for the History of Science. 27-41.

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Sunday, January 12

Session IV: Heredity, Man & Society

9:30

Constitution, Diathesis, and Genetic Susceptibility: An Aspect of the Cultural History of Medical Genetics Robert Olby (University of Pittsburgh)

Max-Planck-Institut für Wissenschaftsgeschichte Max Planck Institute for the History of Science

14:30 George Combe’s Law of Hereditary Descent or Hints on the Heredity of Ideas of Heredity John van Wyhe (University of Cambridge)

15:15 Acquired Character: The (Pre-Genetic) Material of the 'Self-Made Man' Paul White (University of Cambridge)

A Cultural History of Heredity II 18 th and 19 th Centuries

10:15 Coffee Break

10:45 Heredity, Milieu and Sin: The Work of B.A. Morel (1809-1873) Jean-Christophe Coffin (Centre Alexandre Koyré Paris)

16:00 Comments Sigrid Weigel (Zentrum für Literaturforschung Berlin)

16:30 Coffee Break 11:30 Majorat: The Law of Succession and Literature in the 19 th Century Ulrike Vedder (Zentrum für Literaturforschung Berlin)

17:00 General Discussion

12:15 Comments David Sabean (University of California)

13:00 Lunch Break

A

MAX-PLANCK-INSTITUT FÜR WISSENSCHAFTSGESCHICHTE

Wilhelmstraße 44 D– 10117 Berlin Telefon (+ 4930 ) 22667 – 0 Telefax (+ 4930 ) 22667 – 299 www.mpiwg-berlin.mpg .de

January 10 – 12, 2003

Friday, January 10

Saturday, January 11

14:30 Welcome and Introduction

fterno Session II: Heredity & Biology

15:00 Hérédité, Old and New Carlos López Beltrán (Universidad Nacional Autónoma de Mexico)

9:00

Adaption and Heredity in Lamarck’s and Geoffroy Saint-Hillaire’s Biological Theories Wolfgang Lefèvre (MPIWG Berlin)

9:45

In Search of Lost Generations. Life-Cycles and Organisms between 1800- 1860 Ohad Parnes (University of Bern)

Session III: Heredity & Medicine

14:00 Erasmus Darwin on Hereditary Disease: Conceptualising Heredity in Enlightenment English Medical Writings Philip Wilson (Penn State’s College of Medicine)

15:45 Coffee Break

Session I: Heredity & Breeding

16:15 The Sheep Breeder’s View of Heredity (1723-1843) Roger Wood (University of Manchester)

17:00 Characters Written with Invisible Ink. Elements of Hybridism 1751-1875 Staffan Müller -Wille (MPIWG Berlin)

17:45 Comments Raphael Falk (Hebrew University Jerusalem)

14:45 “Poor Old Ancestors”: The Popularity of Medical Hereditarianism 1770-1870 John C. Waller (Wellcome Trust Centre London)

10:30 Coffee Break 15:30 Coffee Break 11:00 Heredity and Adaptation in Kant Peter McLaughlin (MPIWG Berlin)

11:45 Comments Pietro Corsi (Centre Alexandre Koyré Paris)

12:30 Lunch Break

16:00 Pathological Heredity as a Bid for a Greater Recognition of Medical Authority in France, 1800-1830 Laure Cartron (University of Paris I – Sorbonne)

11:45 Comments Gianna Pomata (University of Bologna)

19:30 Dinner

Fridday March 2nd, 2001

MAX-PLANCK-INSTITUT FÜR WISSENSCHAFTSGESCHICHTE

M a x P la n c k I n s titu te fo r th e His to r y o f Sc ie n c e

2005

PREPRINT 294

Co nf e re nc e

A Cultural History of Heredity III: 19th and Early 20th Centuries

Table of Contents

Introduction Staffan Müller-Wille and Hans-Jörg Rheinberger

3

Mendel’s impact Raphael Falk

9

The Biometric Sense of Heredity: Statistics, Pangenesis and Positivism Theodore M. Porter

31

Sources of Johannsen´s Genotype Theory Nils Roll–Hansen

43

Inheritance of Acquired Characters: Heredity and Evolution in Late Nineteenth-Century Germany Wolfgang Lefèvre

53

Darwinism versus Evo–Devo: a late–nineteenth century debate Jeffrey H. Schwartz

67

Message in a Bottle: The Business of Vaccines and the Nature of Heredity after 1880 Andrew Mendelsohn

85

The Chromosomal Theory of Heredity and the Problem of Gender Equality in the Work of Theodor and Marcella Boveri Helga Satzinger

101

Hugo de Vries's transitions in research interest and method Ida H. Stamhuis

115

Herbert Spencer’s two editions of the Principles of Psychology: 1855 and 1870/72. Biological heredity and cultural inheritance Snait B. Gissis

137

Writing Heredity: Emile Zola’s Rougon-Macquart and Thomas Mann’s Buddenbrooks Ulrike Vedder

153

Heritage – Appropriation – Interpretation: The Debate on the Schiller Legacy in 1905 Stefan Willer

167

From pedigree to database. Genealogy and Human Heredity in Germany, 1890–1914 Bernd Gausemeier

179

Bismarck the Tomcat and Other Tales: Heredity and Alcoholism in the Medical Sphere, The Netherlands 1850–1900 Stephen Snelders, Frans J. Meijman and Toine Pieters

193

A Changing Landscape in the Medical Geography of ‘Hereditary’ Disease: Syphilis, Leprosy, and Tuberculosis in Hawai‘i (1863-1903) Philip K. Wilson

213

How Cultural Is Heritage? Humanity’s Black Sheep from Charles Darwin to Jack London Marianne Sommer

233

Race and Kinship in Anthropology: Morgan and Boas Staffan Müller–Wille

255

Introduction Staffan Müller-Wille and Hans-Jörg Rheinberger

The contributions to this volume were prepared for the workshop “A Cultural History of heredity III: Nineteenth and Early Twentieth Centuries” that took place at the Max-Planck-Institute for the History of Science January 13-16, 2005. The workshop was part of a long-term interdisciplinary project dedicated to the cultural history of heredity. Concentrating in turn on a succession of time periods in chronological order, this series attempts to uncover and relate to each other the agricultural, technical, juridical, medical, and scientific practices in which knowledge of inheritance was materially anchored and in which it gradually revealed its effects. 1 The two previous workshops took place in May 2001 and January 2003, each devoted to what we identified as ’historical epochs’ in the history of hereditary thought.2 What follows is a summary of the results reached so far in our project, of the questions we wanted to address with the third workshop, and the answers suggested to these questions by the contributions collected in this preprint.3 In his book “The Logic of Life”, François Jacob already pointed out that the concept of reproduction, and by extension, that of heredity were virtually absent from speculations about generation until the eighteenth century. This we found largely corroborated. It may be worthwhile to stress that heredity as a biological concept originated as a metaphor: prior to the nineteenth century it was used as a synonym of “inheritance” in legal contexts only, a sense which it since then has lost. It was only in the early nineteenth century, in medical contexts, that metaphors of heredity began to gain currency. To be sure: Phenomena that nowadays would count as hereditary had by no means gone unnoticed before. It seems, however, to be a simple matter of historical fact that these phenomena were not addressed in terms of inheritance. The conceptual reason for this was the lack of some fundamental distinctions in pre-modern theories of generation. Before the end of the eighteenth century hereditary transmission was not a domain regarded as separate from the contingencies of conception, pregnancy, embryonic development, parturition, and even lactation. Similarity between progenitors and their descendants was thought to come about as a result of the similarity in the constellation of causal factors involved in each act of generation. Parental organisms were thought to actually make their offspring, without the intervention of a specific hereditary substance transmitted from generation to generation. So if speculations into generation did not provide the context in which biological heredity originated, what did? When Buffon, Maupertuis, and Kant were addressing heredity in the second half of the eighteenth century, they were referring, alongside the discourses of natural history and breeding, to a discourse of highly idiosyncratic origin, namely the Latin-American system of racial 1 2 3

See http://www.mpiwg-berlin.mpg.de/HEREDITY/ for further information on the project. Contributions to the first two workshops have been documented in the Max Planck Institute’s preprint series (no. 222 and 247). A more extensive summary of the resultsfrom the project so far, including a bibliography, has been published as in the Max Planck Institute’s preprint series no. 276.

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Staffan Müller-Wille and Hans-Jörg Rheinberger

classification, known as las castas, and usually presented in paintings arranged in a tabular form. This scheme was primarily based on a classification according to skin color, to a lesser degree also on hair form and eye color. Children resulting from mixed marriages were positioned in this scheme in analogy to the simple mechanism of color mixing, implying “blending” as the causal relation connecting traits of parents with traits of their offspring. The castas classification originated from attempts in early colonial Latin America to find a measure by which legal and social status could be allocated, while colonial society experienced constant flux due to intermarriages and migration. This indicates an important general result of our project: the problem that biological heredity came to address was not the constancy of species; it were the patterns and processes that structure communal life at a sub-specific level. This shift of attention was associated with a mobilization of early modern life within various largely independent cultural domains, all of which also saw a growing concern with hereditary phenomena towards the end of the eighteenth century: Breeding new varieties for specific marketable characteristics, the exchange of specimens among botanical and zoological gardens, experiments in fertilization and hybridization of plants and animals, the dislocation of Europeans and Africans that accompanied colonialism, and the appearance of new social strata with their particular pathologies in the context of industrialization and urbanization, all of these processes interlocked in relaxing and severing cultural and natural ties to provide the material substrate for the emerging concept of heredity. The various domains in which the knowledge regime of heredity took shape during the eighteenth century did not cohere, however, after the model of an overall “influence” that a fundamental idea of heredity gained over them. Rather, conjunctions between them came about by a kind of domino effect that mobilization in one field had on another. The growth of a class that depended on mobile property evoked a culture of leisure collecting and breeding. The import of plants for collection purposes of natural history in turn inspired attempts at their acclimatization for economic purposes. Several, highly specific, and largely independent cultural sub-fields were thus subsequently conjoined to form a field of phenomena which only eventually, in the mid-nineteenth century, came to be addressed by the concept of biological heredity. Although this makes it difficult, even impossible, to draw a general picture of the historical development that led to the formulation of full-fledged theories of heredity in the mid-nineteenth century, it is possible, in hindsight, to characterize the result of that development. As a point of departure, let us take Darwin’s theory of pangenesis. Two aspects of that theory, if compared with premodern theories of generation, are remarkable. First of all, Darwin endorsed a view of heredity that abstracted, to some extent at least, from the personal relation between parents and their offspring. While conceding an inheritance of acquired properties, Darwin believed that the true carriers of the properties to be inherited are not the parents themselves, but submicroscopic entities — “invisible characters” as he called them — which circulate, from generation to generation, among individuals within one and the same species. Secondly, Darwin’s theory of heredity shows a peculiar inversion in comparison with early modern theories of generation: while the latter emphasize the vertical dimension of lineal descent — where ancestral organisms bring forth their offspring — Darwin invoked an image where the horizontal dimension dominates, the dimension of a common reservoir of dispositions, passed down from the sum total of ancestors,

4

Introduction

redistributed in each generation among individuals, and competing now, in the present, for their realization. We take these two aspects, which accord well with the long term development of Western European kinship systems as social anthropologists have reconstructed them, to be the fundamental hallmarks of modern hereditary thought. The complex constellation that Darwin evisaged — thousands of generations represented virtually in the microscopic space of each fertilized egg — points to the complexity of the biological concept of heredity. In this respect, one can speak of an “epistemic space” of heredity that came into being in mid-nineteenth century. In contrast to other subjects of biological research, which are determined within individual experimental settings, heredity depended on a vast, spatial configuration of distributed technologies and institutions connected by a system of exchange: botanical gardens, hospitals, chemical and physiological laboratories, genealogical and statistical archives. It is this “space” and its development during the late nineteenth century that we wanted to explore in the third workshop in our project. It was planned to occupy itself — chronologically speaking — with the latter part of the nineteenth century and the first years of the twentieth. Roughly, this period can be regarded as one in which various attempts were made to thoroughly theorize heredity on the basis of observation, experiment, and statistical analysis, but in different directions. For orientation, we demarcated the period by two publications: Francis Galton’s Hereditary Talent and Character of 1865 and Wilhelm Johannsen’s Über Erblichkeit in Populationen und in reinen Linien of 1903. Obviously, it could be asked why we did not take Mendel’s 1864 paper and the year 1900, the annus mirabilis of the birth — or re-birth — of genetics as our points of orientation. We did not do that because we did not intend to focus on the triumphal advent of classical genetics, breeding practice included, in this workshop. Rather, we wanted to locate this event, this advent — the “(re-)discovery” of Mendel — in the much broader spectrum of theories and practices of inheritance that had already been consolidated in such widely different contexts as evolutionary and developmental biology, cytology, bacteriology and epidemiology, eugenics and anthropology by the end of the nineteenth century. What we did do, however, is to ask Rafael Falk to talk to us on Mendel’s impact as a sort of introductory lecture, an exergon so to speak for — and before — this conference, and then turn to the other topics. We plan to come back, as already said, to Mendel, breeding, and classical genetics in a future workshop. Following an earlier suggestion from Jean Gayon, we can draw the following scheme for the development of hereditary knowledge in the second half of the nineteenth century: 4 With the rise of heredity as a central biological problem by the middle of the nineteenth century, the question of its material basis and of its mechanism had taken shape. In the second half of the nineteenth century, two major frames were proposed to deal with this question. The first one saw heredity as a force, whose strength could be accumulated over the generations, and which, as a measurable magnitude, could be subjected to statistical analysis. This concept was particularly widespread among nineteenth-century breeders, and it influenced Francis Galton and the so-called “biometrical school.” The second saw heredity as residing in matter that was transmitted over the 4

For references see our entry on “Gene” in the Stanford Encyclopaedia of Philosophy at http:// plato.stanford.edu/entries/gene/.

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Staffan Müller-Wille and Hans-Jörg Rheinberger

generations. Two major trends can be differentiated in this latter case. One of them conceived of hereditary matter as particulate and amenable to breeding analysis. Charles Darwin called the presumed hereditary particles “gemmules”; Hugo de Vries, “pangenes”; Gregor Mendel, “elements”. None of these authors, however, thought of associating these particles with a particular hereditary substance. They were seen as microscopic entities, which, if accumulated en masse, would make the particular traits visible for which they stood. A second category of biologists in the second half of the nineteenth century, to whom Carl Naegeli and August Weismann belonged, distinguished a specific hereditary substance, the “idioplasm”, or “germ plasm”, which they assumed to be responsible for generational hereditary continuity, on the one hand, and the body substance, the “trophoplasm” or “soma”, on the other. While considerable work has already been done in terms of a history of these ideas, comparatively little effort has been made to explore the links of the above mentioned late nineteenth century developments with the notion of heredity itself. Most studies, which paid attention to the contexts of, e.g., anthropological or medical work have tended to take inheritance as the necessary, yet in itself unquestioned prerequisite, upon which their actors relied to promote their hierarchies of human inequality. With the variety of hereditary theories we encounter around 1900 it seems, however, that the very notion of inheritance was rather deeply troubled at this time. The workshop was planned to explore the sources for these troubles in a (certainly incomplete) number of fields in which hereditary considerations played a role: anthropology, evolutionary and developmental biology, cytology including microbiology, plant and animal breeding, epidemiology including bacteriology, eugenics, and juridical and sociological concepts of hereditary transmission. Not all of these fields could be covered with equal density, others came into sharper focus during the preparation of the conference, and in putting together the proposals, we had to make our choices. So we ended up with six sessions. The first set the stage of theorizing heredity in the later nineteenth century; the second looked at heredity in the context of evolution and development; the third occupied itself with the physiology of heredity in a broad sense; the fourth brought together aspects of social, psychological, and cultural heritage; the fifth was devoted to heredity and genealogy; and the sixth, finally, thematized heredity and anthropology. One of the main outcomes of the workshop was that the two dichotomies developed by historians so far with respect to late nineteenth-century theories of heredity, namely soft vs. hard (Ernst Mayr) and blending vs. non-blending inheritance (Robert C. Olby) do not seem to work well to fully capture the variety of theoretical approaches in that time period. There are two reasons for this: (1) It became apparent that up until the very end of the nineteenth century speculations into heredity either conflated these oppositions, or remained rather indifferent with respect to them. Within the medical community, for example, the belief was widespread that diseases leave their “stamp” upon offspring or that substances like alcohol “poison” the germinal substances. This position reflects a conception of heredity both “soft” and “hard” at the same time — soft, in as much as environmental factors induce a change in the hereditary substance; hard, in as much as this change, once it has occurred, is irrevocably passed on in the germinal line (in obvious analogy to original sin). Likewise, the “blending” of parental characters in the offspring was often seen as

6

Introduction

compatible with a view of the hereditary substance as being composed of “non-blending”, particulate elements, as notably in Darwin’s theory of pangenesis and Galton’s theory of the stirp. Both oppositions, “soft” vs. “hard” and “blending” vs. “non-blending” seem to have become serious issues of dispute and dissent only after the onset of Mendelism in 1900. (2) The oppositions of “soft” vs “hard” and “blending” vs “non-blending” heredity cover up or cut across more fundamental, and more hotly debated, oppositions with regard to the material make-up and causal agency of hereditary material. Was heredity to be conceived as a natural force or as an organic structure? What was the source of the hereditary material, the ancestral organism itself or something passed on independently? Was there a particular substance that formed the germ plasm, and where was it possibly located? And could it be related to particular cellular structures? What, if any, were the elements of the hereditary material, and how did these elements relate to each other? Did they fuse, or just mix? And in what manner did they determine the future organism, directly throughout the individual life-span, or only by determining the first steps of development? Finally, what, if any, where the distinct roles the two sexes played in inheritance? The nineteenth century did not come up with concluding answers to these questions, as is well known, and the contributions to the workshop showed the considerable breadth of positions with respect to inheritance that persisted well into the first decade of the twentieth century. Yet the very nature of these questions points to a decisive trend in the period studied by the workshop. Inheritance was increasingly seen not as a relation between individual organisms – ancestors and descendents – but as a relation of populations to a shared, germinal substrate. Various contributions pointed to two important, but until now under-researched, sources of models that shaped representations of this substrate: genealogy and epidemiology. Genealogy developed in the late nineteenth century into a tool for analyzing populations rather than individual ancestry, and brought to the fore both the openness of family relations and the quasi-mathematical closure of genetic relationships. In epidemiology, heredity, infection, and vaccination intersected to produce what Jean-Paul Gaudiellière and Ilana Löwy have called the “impossible separation” of horizontal and vertical dimensions in the transmission of diseases. The reason for the “devaluation of ancestry” in favor of a view that sees (cultural as well as biological) inheritance as a common stock of dispositions seems to lie in the association of heredity with the future rather than the past, with projection rather than with legitimization, that occurred in the context of the all-pervading late-nineteenth century theme of progress. Even where the past entered hereditary discourse, it did so either as a threat to the present, in form of “atavisms”, “throw-backs” and “degeneration”, or as a “heritage”, “stock”, or “capital” to be appropriated anew by each generation. Breeding plants and animals provides the obvious model here, and Mendelism with its close connection to the breeding industry took the decisive step to attack hereditary phenomena by “deducing forward”, as Raphael Falk put it during the workshop. Studying the development of Mendelism against the background explored by this workshop will be the agenda for the next workshop, which is planned to take place at the University of Exeter in autumn 2006.

7

Mendel’s impact Raphael Falk

Abstract Mendel introduced a reductionist methodology to the study of inheritance. By 1900, reduction of biology to the laws of chemistry and physics was not self evident: Although developmental bottom-up preformation was largely inadequate, organismic epigenetic hypotheses often fall prey to metaphysical assumptions. The achievements of Mendel’s reductive research methodology in genetics were increasingly extended to reduction at the conceptual level, not only with respect to transmission genetics but similarly in dealing with problems of development and evolution. Genetics became extremely genocentric in its explanatory arsenal, and accordingly it was divorced from the top-down, life-as-organized-systems’ notion of embryology. Similarly, evolution was conceptually reduced to population genetics in terms of gene-frequencies. It was, however, the increasing attempts to ground the bottom-up approach at the molecular level that eventually pushed conceptual reductionism to its crisis. Modern developments of molecular and computational methods finally forced, or allowed, genetics and development to apply reductionist methods to top-down systems’ analysis, thus to close the cycle of the adoption of reductionist methodologies to top-down conceptions. Gregor Johann Mendel’s personality unfolded and matured in a community of scientifically minded breeders. We are naturally most interested in the scientific impact of Mendel. However, Mendel apparently had also an impact on breeders, as was unexpectedly brought home to me through a story that a bio-ethicist colleague, Dan Wikler, told me a couple of years ago, about his Himalayan holiday in 1993. On his tour of northern India Dan met a local scholar, Tshering Dorje, a specialist in pre-Buddhist religions of Tibet, who functioned also as his group’s guide. The man came from a valley, named Lahoul, located in the Western Himalaya, north of a pass that leads to Kulu, Manali, and further down to Punjab (fig. 1). The inhabitants of that valley were quite well off ever since a road that connected them with the rest of the world had been built. This allowed them to export the crop of peas that they had specialized in growing and that has been known for its quality. According to the guide, a Moravian missionary, called Francke, introduced pea growing into the area at the end of the nineteenth century. Francke related that he learned the secret of growing peas from his cell mate in the monastery, whose name was Mendel, “who was quite good at growing peas.” Dan suspected that the guide did not know much of Mendel and his impact on science.

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Raphael Falk

Fig. 1.: The Lahoul valley in the Himalaya. Now, this story is too good not to be true. I tried to follow the story somewhat further: A Moravian missionary by the name August Herman Francke wrote a history of Ladakh (Francke, 1977). He was however too young to be Mendel’s cell mate: He was twelve when Mendel died. In his book, A History of Ladakh there is no hint to that story, but Moravian missionaries had been active in the area already before Francke: An earlier history of the country was written by another Moravian missionary by the name Karl Marx (of all possible names!). I finally got hold of an e-mail address of the Moravian mission in Ladakh, and got a prompt response from the reverend Elijah Gregan, a pastor of the Moravian church, who turned out to be a cytogeneticist by training. He wrote: “I have never heard of the story that you wrote about Mendel … However, Moravians are well known in the area for having introduced vegetables in the Lahoul and Ladakh valleys.” I hope you will agree with me that this adds a piquant and unexpected flavor to the story of the origins of genetics, although unfortunately at the moment it seems not to be true.

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Mendel’s impact

Genetics: Coming of age of heredity Mendel’s impact on science, I wish to propose, was by introducing a reductionist agenda to the research of heredity. This notion of Mendel I believe was based primarily on a deep religious conception of the world as governed by laws of nature formulated in numerical principles (Falk, 2001). Mendel, however, did not introduce a particulate material notion into the study of heredity – he talked of Faktoren. As pointed out by Gayon (2000, 71-73), it was Darwin who rejected the prevalent notion of hereditary force and promoted a notion of material particles, gemmules, and de Vries who turned these particles into pangenes, which in 1900 he associated with Mendel’s factors. De Vries (1848-1935) was primarily interested in backing up his own hypothesis of intracellular development and evolution, rather than in establishing pangenes as foundations of a theory of particulate heredity along Mendel’s insight. Notwithstanding, his role in promoting Mendel’s insight was crucial, though indirect. As for the other two traditional “rediscoverers” of Mendel’s work: The younger Carl Correns (1864-1933), who was less ambitious in formulating universal laws of nature, it must be admitted, did not grasp in due time the potential of Mendel’s work as the dawn of a new science of heredity. After reading de Vries’s paper, he related that he had obtained similar results, and indeed, was aware that he “had found something new,” but added “that Mendel’s Law of segregation cannot be applied universally,” and accordingly, he “did not consider it necessary to establish” his priority (Correns 1900, in Stern and Sherwood, 1966, 120). With regard to the third “rediscoverer” of Mendelian segregation of characters, Erich von Tschermak-Seysenegg (1836-1927), it was noted by Curt Stern that “his publications in 1900 show him not to have been a discoverer of Mendelism but only an experimenter whose understanding … had ‘fallen short of the essential discovery’” (Stern and Sherwood, 1966, xi-xii) . Or, as pointed out by Bob Olby, his paper “reveals very clearly how Mendelian results could be treated in terms of the orthodox conceptions of heredity in the nineteenth century, even after reading Mendel’s paper” (Olby, 1985). Although he never claimed to have had discovered either the Mendelian ratios or their explanation, it was William Bateson (1861-1926, fig. 2) who conceived of Mendel’s experimental work as a founding contribution to the new research discipline of heredity and variation, which in 1906 he called genetics. Contrary to de Vries, who superimposed his notions of pangenes’ function on the Mendelian factors in the service of his theory on the origin of species by mutations ( Die Entstehung der Arten durch Mutation, de Vries, 1902-03, II, xii), Bateson, like Correns, was interested in the experimental discovery of the nature of these factors of inheritance. Notably, both adhered to material preformationist notions that did not discriminate conceptually between the factors for traits and the traits proper: This was most obvious from their respective notions of dominance-recessivity relationships: Correns interpreted dominance in terms of the physiological action of the factors, whereas Bateson attributed it to the material presence of the factors of inheritance, as formulated in his Presence-Absence Hypothesis (see Falk, 2001; Rheinberger and Müller-Wille, 2004). When Wilhelm Johannsen (1857-1927) formally introduced in 1909 the distinction between the “unit character” and the gene he was most careful not to attribute any explicit material meaning to his gene-concept (Johannsen, 1909). Johannsen’s gene was merely an intervening variable. But with the establishment of the Chromosomal Theory of Inheritance by Morgan

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Raphael Falk

(1866-1945) and his students, the Mendelian units of inheritance became hypothetical constructs, and for at least one of his students, Hermann J. Muller (1890-1967), the genes were discrete material entities, the atoms of inheritance, the physical-chemical properties of which he set out to study (Falk, 1986).

Fig. 2.: William Bateson, the founder of the science of Genetics (on the left) and Wilhelm Johannsen, who segregated the phenotypic traits from the genotypic potential (to Bateson’s right). Thus, I repeat, Mendel’s scientific impact was by introducing a reductionist research agenda, and genetics became the prima facie reductionist life-science. Now, from the privileged perspective of

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Mendel’s impact

somewhat more than one century later, I wish to examine the achievements and the shortcomings of the Mendelian reductionism and its conceptual extensions on the science of inheritance. This, however, must be done in the context of the aggressive developments in the life sciences at the turn of the twentieth century, especially in notions of development and evolution. Although by1900 reduction of biology to the laws of physics and chemistry had already been accepted among physiologists (even though the breakthrough of biochemistry was still decades ahead), this notion still struggled for its application in studies of development and evolution. Mendel’s reductionist research methodology thus was to confer on the study of inheritance a special status among the life-sciences, namely that of providing for the extension of reductionism to embryological and evolutionary studies. I wish to claim that genetics as the science of heredity was never divorced from development and evolution. Rather, we have been witnessing in the nineteenth and the twentieth century a full cycle, from a holist view of heredity as an aspect of development and evolution that necessarily fell back on irrational assumptions, to the higher level of a rational though reductionist view of heredity, back to a holist perspective of heredity at the level of systems’ analysis of development and evolution. In the footsteps of Mendel, who heuristically sublimed the details of the characters that he judiciously selected for his experiments on transmission between generations, geneticists viewed the specific cases of development and evolution which they happened to study as samples of generalized – methodologically and eventually conceptually – reducible units. Starting with Bateson’s presence- absence hypothesis of dominance (Bateson, 1905 (1928)), through Muller’s interpretation of dominance as precision of genetic functional adaptation (Muller, 1950), to Beadle and Ephrussi’s study of specific discrete gene-effects that culminated in the one-gene – one-enzyme thesis (Beadle & Ephrussi, 1936; Beadle & Tatum, 1941), and to the formulation of the New-Synthesis of evolution in term of populations’ gene frequencies (see Provine, 1971), geneticists inferred rules of development and evolution in which the gene provided “a kind of inertia principle” against which effects could be measured (Gayon quoted in Rheinberger & Müller-Wille, 2004).

Genetics: Reducing inheritance with development Reduction involves the explanation of laws or phenomena in one realm by those of another. Reductionism is the thesis that reductions between two realms are always (or, at least, are always likely to be) successful as explanations. I wish to differentiate between methodological reductionism and conceptual reductionism. Methodological reductionism is the belief that empirically following single (experimental or observational) variables (other variables being kept constant or randomized) is the effective design to bridge realms. Conceptual reductionism assumes that phenomena may be reduced to a component or components at a more basic realm, that individually or interactively bridge the phenomenon at the higher realm.

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Raphael Falk

The distinction is one between explanation and resolution. Methodological reductionism may be identified as an epistemological statement, whereas conceptual reductionism is essentially an ontological statement (Sarkar, 1998, 17ff). Methodological reduction would best correspond to Sarkar’s (1998, 43-45) abstract hierarchical reduction, which assumes that the rules of one realm are more fundamental than those governing another realm. Conceptual reduction would correspond to his strong or approximate strong reduction, in which the hierarchy referred to is also a hierarchy in physical space. Modern science has been methodologically reductionist, ever since Galileo introduced the study of one variable at a time, by keeping all other variables as much as possible constant (or, at least, let the other variables vary randomly with respect to the variable studied). It became also increasingly Cartesian, or conceptually reductionist. The achievements of reductionist methods and conceptions in physics, and eventually also in chemistry encouraged researchers to extend reductionism to the life-sciences. But the life sciences of the eighteenth century were demarcated at the outset from physics and chemistry as immanently non-reductionist. In spite of nineteenth century biologists’ inroads in extending reductionism to the life sciences (Lenoir, 1982), life remained immune to complete conceptual reduction. Something else was needed, whether it was Caspar Friedrich Wolff’s vis essentialis, Johann Fiedrich Blumenbach’s Bildungstrieb, Carl Nägeli’s Vervollkommnungskraft, Theodor Eimer’s Orthogenesis, Hans Driesch’s Entelechy, or Niels Bohr’s search for special laws of physics expressed only in living systems, or the modern theories of complexity. The wish to reduce living being to systems that may be completely explained bottom-up, countered the recognition that living being are the fundamental entities that should be explained top-down in order to unravel the properties of their components. The three major conceptual revolutions in the life sciences of the nineteenth century, the cellular theory (1838-1839), the theory of evolution by natural selection (1859), and the theory of particulate inheritance (1865), all reflect the craving for reductionist explanations for living systems. Of the three, Darwin’s hypothesis of The Origin of Species by Means of Natural Selection was actually a top-down theory. But Darwin’s juxtaposition of the organism versus its environment, independently of what the organism “wants” or needs – contrary to conceptions up to and including the Lamarckian hypothesis of the organism in its environment – inserted a conspicuous reductionist element into his hypothesis. Although Darwin repeatedly tried to back out of this notion of the organism-environment segregation, the reductionist aspect of Darwinism, of ‘nature vs. nurture,’ as phrased by Francis Galton in 1865, was firmly established once formulated by August Weismann (1893) as the germ-plasm theory of heredity. Of the other two conceptions, Schleiden´s and Schwann’s theories of the cell as a universal reductionist idea gained its success with Virchow’s 1855 doctrine of omnis cellula e cellula, whereas Mendel’s reductionist theory gained recognition at the turn of the twentieth century. But it was in the area of development and inheritance in which the reductionist and holist conceptions confronted each other most directly. Wilhelm His and Wilhelm Roux introduced the mechanics of development (Entwicklungsmechanik) largely in the hope to repeat the achievements of the physiologists’ reductionist methods:

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Mendel’s impact

Developmental mechanics is the doctrine of the causes of organic forms. Form represents the most essential attribute of the organism. … In accordance with Spinoza’s and Kant’s definition of mechanism, every phenomenon underlying causality is designated as a mechanical phenomenon; hence the science of the same may be called mechanics. Since … physics and chemistry reduce all phenomena, … to movement of parts, … the words ‘developmental mechanics’ agree with the more recent concepts of physics and chemistry, and may be taken to designate the doctrine of all formative phenomena. … In so far, however, as forces or energies are only known by their effects, … the problem may be defined as the ascertainment of the formative modi operandi. (Roux, 1894, 107-108)

Yet, as Roux himself was conscious of, even if a bottom-up mechanistic methodology was imperative, the conceptual reduction of development was more problematic and controversial: … developmental mechanics must from the start be guided by the conviction that organic structure is mainly due to the operation of components which at present are so complicated as to exceed the limits of our observation. … Although according to our immediate conception of the matter, even these components depend in the last instance on inorganic modi operandi, nevertheless the complexity of the composition lends them attributes which often differ so widely from those of inorganic modi operandi that they are not only very dissimilar but even appear to contradict in part functions of these same inorganic modi operandi. (Roux, 1894, 111)

For embryologists the issue of the mechanism of development became part of the discussion of preformationism versus epigenesis, i.e., of the extent of the immanent inheritance of developmental processes. Although a long way from the old notion of Hartsoeker’s (1694) homunculus, preformation, or the bottom-up view of development as the sum total of the physical and chemical properties of its building blocks was the reductionist notion that prevailed. Those who adhered to the opposite conception of epigenesis, which conceived of the organism as the frame of reference that assimilated the environmental, including the inherited cues in development, could not construe it in terms of additively reducible physico-chemical terms (as preformationist hoped was feasible). Thus, epigenesis repeatedly implicated, sometimes inadvertently, scientists with sensu stricto meta-physical notions of development. The predicament of the analytical biologist, confronted with his experimental results obtained in the best tradition of the reductionist methodology, was presented most eloquently by Edmund B. Wilson in his 1893 lecture at the Marine Biological Laboratory in Woods Hall: Many leading biological thinkers now find themselves compelled to accept a view that has somewhat in common with the theory of prae-formation … Every one of [the] hereditary characters is … represented by definite structural units in the ideoplasm of the germ cell, which is therefore conceived as a kind of microcosm, … [T]he so-called mosaic theory of Roux and Weismann … is essentially a whole arising from a number of independent self-determining parts, ... . [Yet, a] more decisive result was reached in 1891 by Driesch, … [which] the writer repeated … and found that … [t]he isolated blastomere behaves from the beginning, like an entire ovum of one-half or one-fourth the normal size… .

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These facts are obviously a serious blow to the mosaic theory, and the efforts of Roux and Weismann to sustain their hypothesis in the face of such evidence only serve to emphasize the weakness of their case. (Wilson, 1893, 69-72)

The problem was a conceptual one, rather than one of methodological particularization. It is here that Mendel’s accomplishment, of judiciously selecting appropriate markers for his experiments in the 1860s, applying the reductionist methodology, without making an explicit statement on the conceptual problem, is fully paying off. This was noted already in 1902 by Udny Yule: The experimental plants must, Mendel states … “possess constant differentiating characters.” … The races for crossing were thus chosen with the greatest care and patience so as to be absolutely distinct; an A individual mated with an A never producing a’s, nor vice-versa, for, as I understand, the whole period of ten years (two years of preliminary trials and eight years of experiment). (Yule, 1902, 222-223)

De Vries, however, from his preformationist perspective, claimed that determinants were universals, for all characteristics, though unlike the Roux-Weismann hypothesis, he did not believe that these determinants were unequally distributed to the different blastomers: According to him differentiation was achieved by activation-inactivation of latent particles, pangenes. This, by the way, was why de Vries needed the notion of dominance as if it were one of Mendel’s laws, to replace his activation-latency terminology (see Stamhuis, 2003; Stamhuis, Meijer, & Zevenhuizen, 1999). Once Bateson adopted the Mendelian notion of units of inheritance, a preformationist notion of units of development was for him the consequence, and the Mendelian factors were unitcharacters in potency. To the extent that the morphologist’s or physiologist’s unit character did not agree with those of Mendelian segregation units, the former were not really “unit characters,” but rather complex characters. Mendelian segregation was for Bateson a device to assess spurious morphological unit characteristics, such as in the case of chickens’ comb morphology: The “walnut” comb was a compound of two unit-characters, the “pea” comb and the “rose” comb (Schwartz, 1998, 2002).1 Indeed, to a large extent, the dispute between the Mendelians led by Bateson and the Biometricians led by Pearson may be traced back to the Mendelian factors being conceived as preformed unit-characters. Thus, Darbishire’s hybridization experiments with mice, 1

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Bateson’s ignoring any chromosomal association with Mendelian factors has recently been explained by Patrick Bateson as being due to Bateson’s irritation about “the glibness of the explanation offered for the role of chromosomes. The chromosome was portrayed not simply as a structural site of the gene but almost as the gene itself. Worse, the chromosome was treated as the pre-formed version of the character influenced by the gene on that chromosome” (P. Bateson, 2002, 54). Indeed, reviewing Morgan’s Mechanism William Bateson wrote that “it is inconceivable that particles of chromatin or any other substance, however complex, can possess those powers which must be assigned to our factors. … The supposition that particles of chromatin, indistinguishable from each other and indeed almost homogenous under any known test, can by their material nature confer all the properties of life surpasses the range of even the most convinced materialism” (Bateson, 1916). Bateson may have shared Samuel Butler’s rage against machine-aged materialism, crass professionalism, and unabashed utilitarianism (Harman, 2004, 29), but he certainly was looking for a more sophisticated material preformationism. I tend to accept Punnett’s response to the question of how he and Bateson missed the tie-up of linkage phenomena with the chromosomes: “The answer is Boveri. We were deeply impressed by his paper ‘On the individuality of the chromosomes’ and felt that any tampering with them by way of breakage and recombination was forbidden” (Punnett, 1950, 10).

Mendel’s impact

in which there were unexpected variations in fur color, whereas a preformationist principle of dominance predicted uniformity of color for unit characters in the F 1, were taken by Biometricians as evidence against Mendelian determinism, and Bateson had to come forward with various excuses for the variability of fur color of the alleged dominants (Ankeny, 2000, 328). Similarly, William Castle’s (1867-1962) explanation of the variation in color-patterns of hybrid rats by a model of allele-contamination in heterozygotes (Castle, 1906) was based on a preformationist notion that did not distinguish between traits and the factors for traits. My own experimental studies of heredity, begun in 1902, early led me to observe characters which were unmistakably changed by crosses and so I have for many years advocated the view that the gametes are not pure in the sense expressed by Bateson. (Castle, 1919, 126)

Not so for Johannsen, whose notion was one of an essentialist typologist (Roll-Hansen, 1978). His was a top-down view, the essence of which was an Aristotelian genotype, the earthly appearance of which is the phenotype. Johannsen emphasized the status of the genotype as the essential entity, independent of the environment and the circumstances in which it developed. Sodann erkennen wir, daß der “Typus” im Quetelet’schen Sinne nur eine Erscheinung oberflächlicher Natur ist, welche täuschen kann; erst durch weitere Untersuchungen wird entschieden, ob ein einziger oder mehrere biologisch verschiedene Typen vorhanden sind. Darum könnte man den statistisch hervortretenden Typus passend als Erscheinungstypus bezeichnen oder, kurz und klar, als “Phaenotypus.” (Johannsen, 1909, 123)

Johannsen’s essentialist notion of the genotype is made most explicit by confronting it with that of Woltereck (1877-1944), who conceived of the organism in its environment to be the top-down essence (Woltereck, 1909). Woltereck offered a dynamic, non-typical norm of reaction of the hereditary element of the organism instead of the constant essential genotype. Woltereck’s “phenotype curves” are graphical schemes of the degree of a “particular character as it manifests itself under different conditions.” Johannsen the essentialist protested: Of course the phenotypes of the special characters, i.e., the reactions of the genotypical constituents, may under different conditions exhibit all possible forms of transition or transgression – this has nothing at all to do with constancy or inconstancy of genotypical differences. … Nobody will assume that there should be genotypical transitions here! Pure lines of beans may in one year be different in size … Differences of soil may produce something similar, and it is well known to breeders that some strains of wheat yield relatively much better than others on rich soil, while the reverse is realized on poorer soils. … The genotype-differences are nevertheless constant; the “Reaktionsnorms” of the organisms in Woltereck’s cases, … are of course eo ipso “constantly different” just as well as the “Reaktionsnorms” of different chemical compounds” (Johannsen, 1911, 145-146).

The genotypes are as constant as are chemical elements, whatever the properties of the compounds these elements are involved in might be. It is important to note, however, that for Johannsen the major distinction was that between the phenotype and the genotype; his introduction of the “gene” was a bow to Mendel and Weismann, in spite of his holist conception. From the perspective of one decade later he asserted:

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My term “gene” was introduced and generally accepted as a short and unprejudiced word for unit-factors … but originally I was somewhat possessed with the antiquated morphological spirit in Galton’s, Weismann’s and Mendel’s viewpoints. From a physiological or chemicobiological standpoint … there are no unit characters at all! … We may in some way “dissect” the organism descriptively, using all the tricks of terminology as we please. But that is not allowed in genetical explanation. Here, in the present state of research, we have especially to do with such genotypical units as are separable, be it independently or in a more or less mutual linkage. (Johannsen, 1923, 136-137)

Hence, by his insight Johanssen provided the framework that allowed to severe the Gordian knot of the unit character and the Mendelian factor of Bateson and his school, paving the way for the new science of genetics to become phenomenologically reductionist without being preformationist, or adopting an “instrumental reductionism” without adopting a “material reductionism.” This was taken up by another ardent opponent of Mendelian preformationism, namely Thomas Hunt Morgan. Morgan repeatedly tried to maintain the distinction between methodological reductionism – which he accepted – and conceptual reductionism – which he tried to postpone, if not reject (see, e.g., Falk & Schwartz, 1993). Although Morgan eventually had to give way to the technical hurdles that this position put in his thinking, he maintained the “instrumental reductionist” viewpoint throughout his career (Allen, 1978; Falk, 1986). However, among his students and colleagues, the distinction between the reductionist conception and the reductionist methodology often became rather equivocal. Thus, by the beginning of the second decade of the twentieth century, when the focus of genetic research moved from Europe across the Atlantic to the United States, and especially to Morgan’s “Fly-Room,” the lines began to be drawn: Contrary to the Central European tradition of a strong philosophically, worldview-driven research, whether holistic or reductionist (for an extensive discussion, see Harwood, 1987), the American tradition was more a pragmatic, methodological reductionism, which, however, degenerated into conceptual reductionism. Confronted with the achievements of reductionist genetics of the Morgan school, Johannsen’s appeal for a genotypic – we would call it nowadays, genomic – perspective, was to no avail: We are very far from the ideal of enthusiastic Mendelians, viz. the possibility of dissolving genotypes into relatively small units, … Personally I believe in a great central “something” as yet not divisible into separate factors. The pomace-flies in Morgan’s splendid experiments continue to be pomace-flies, even if they loose all “good” genes necessary for a normal fly-life. … But however far we may proceed in analyzing the genotypes into separable genes or factors, it must always be born in mind, that the characters of the organisms – their phenotypical features – are the reaction of the genotype in toto. The Mendelian units as such, taken per se are powerless.(Johannsen, 1923, 139)

Obviously, Johannsen was increasingly alarmed by genetics becoming more and more conceptually reductionist or genocentric. Instead of characters being “markers” for genes, genes became entities “for” characters. Any attempts to maintain a top-down conception were marginalized.

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The genocentric notion probably was at its peak when Oskar Vogt attempted in 1926 to reintroduce essentialist notions of discontinuity of taxonomic entities. He and TimoféeffRessovsky (1900-1981) in his footsteps, dismissed phenotypic deviations of the pre-determined genetic essences by trivializing the variations as due to phenotypic “penetrance” or “expression” of the mutants (Vogt, 1926). In the same vein, geneticists were deliberating on whether pleiotropy – the production by one particular mutation of apparently unrelated multiple effects – was genuinely “primary,” one gene producing fundamentally many products, or “secondary,” according to which all the multiple effects of a gene should be reduced to a single effect (see Falk, 2000). As Laubichler and Sarkar (2002) note, these concepts “are invoked when any simple relation between a specific genotype and phenotype break down.” They argue, furthermore, that even today “the basic role that ‘expressivity’ and ‘penetrance’ play in contemporary human behavioral genetics is ideological: they help maintain a genetic etiology for traits in the face of recalcitrant detail.” It is instructive to recognize even in our present conceptions the remnants of these genocentric considerations of pleiotropy, though in the reverse direction, namely in the efforts to identify the DNA-sequence which may be authoritatively defined as a gene. Should the multiple alternative messenger-RNAs extracted from a given DNA-sequence be all considered the products of one gene, or may a DNA-sequence embrace multiple, partly overlapping genes? In 1910 Morgan asserted that “When we speak of the transmission of characters from parent to offspring, we are speaking metaphorically” (Morgan, 1910), and in 1934, in his book Embryology and Genetics, he still continued to claim that “The story of genetics has become so interwoven with that of experimental embryology that the two can now to some extent be told as a single story.” He even came up with an important model – not testable at the time – of development through differential activation of the constant hereditary entities, universally present in all somatic cells (Morgan, 1934, 9-10). But by that time Morgan apparently found out that he could not reconcile the old couple of embryology and genetics. As Jane Maienschein noted: “In the end, with his book Embryology and Genetics of 1934, he did not even find it profitable to try very hard” (Maienschein, 1987, 92): The “and” in the book’s title turned out to be more a dissociate- than an associate-conjunction. Herman J. Muller’s identification of genes as the atoms of inheritance and variation, the physico-chemical properties of which should be elucidated, gained increasing acceptance. As stressed by Allen (2002, 27) “the strong emphasis on discrete and separable units interacting additively, as opposed to synergistically, posed serious problems for understanding developmental processes.” Non-genocentirc notions, like Richard Goldschmidt’s (1878-1958) top-down philosophy, just as Barbara McClintock’s (1902-1992) genomic – sometimes mystic – conception, were politely but emphatically rejected. On the other hand, Lewis Stadler’s (1896-1954) conceptual holism yet methodological operationalism (Stadler, 1954) was gracefully acknowledged. Yet, the involvement of geneticists of the twentieth century in issues of development was not less than that of the biologists in the late nineteenth century who insisted that heredity and development were inseparable (Maienschein, 1987; Sandler & Sandler, 1985). In the footsteps of Mendel, problems of development were skillfully converted to problems of individual gene functions. Haldane (1892-1964) described the function of specific genes in the synthesis of

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antocyanines that determine the flower color in Pelargonium (Haldane, 1954, 52-58) and Sturtevant (1891-1970) described the function of specific genes in the development of coiling in the snail Limnea (Sturtevant, 1923) or of organ differentiation in Drosophila (Sturtevant, 1932). Beadle (1903-1989) and Tatum (1909-1975) rightfully saw the direct line from Garrod’s (18571936) “inborn errors of metabolism,” which assigned specific malfunctions in metabolic pathways to specific human diseases, to their experimental evidence that reduced the metabolism of Neurospora crassa to that of a “one-gene – one-enzyme” relationships (Beadle & Tatum, 1941). Significantly, however, Alfred Kühn (1885-1968) expressed a wider awareness of the need to refer to the organism when discussing genes in development. Based on his work on the pigmentation of the moth Ephestia küniella, he pointed out that our methodological experimental reductionism should not distract us from a conceptual holistic notion of the organism: [Our] apprehension of the expression of hereditary traits is changing from a more or less static and preformistic conception to a dynamic and epigenetic one. The formal correlation of individual genes mapped to specific loci on the chromosomes with certain characters has only a limited meaning. … One trait appears to have a simple correlation to one gene only as long as the other genes of the same action chain and of other action chains which are part of the same node, remain the same. (Rheinberger and Müller-Wille, 2004, quoting Kühn, 1941, 258)

Genetics: The taming of Darwinism Whereas heredity and development could arguably be viewed as separate or identical disciplines, depending on whether the emphasis was on the physiological aspects of developmental changes or on the generation-stability of development, the relation of heredity and evolution was unequivocal as has been laid down by Darwin’s “provisional hypothesis of pangenesis” in The Variation of Plants and Animals Under Domestication (1868). Bateson adopted Mendel’s hypothesis immediately upon reading de Vries’s 1900 paper because it was consistent with his notions of evolution by discontinuous steps. Contrary to Bateson’s conclusions that relied on the a priori theory of “Mendel’s Law of Hybridization,” concerning the organization of biological material, Karl Pearson (1857-1936), the student of positivism, argued for a position that was based on statistical observations, ostensibly without prior theoretical assumptions. This, he claimed, was represented in the “Law of Ancestral Heredity” (at least, once the parameters introduced by Galton had been eliminated from it). However, already in 1902 it was shown by Udny Yule (1871-1951) that mathematically one notion could be reduced to the other. Sixteen years later R. A. Fisher (1890-1962) too showed that mathematically the Law of Ancestral Inheritance can be reduced to that of Mendelian Inheritance (Fisher, 1918). Yule and Fisher, however, had different agendas. A careful analysis of Yule’s comments (Tabery, 2004) shows that the dispute between Pearson’s biometricians and Bateson’s Mendelians was one between an organismic, top-down phenomenological descriptive analysis of the variability of a race (= breeding population) and a bottom-up particulate reductive analytical interpretation of individual (or pure line) character differences in hybridization experiments.

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There has always been a good deal of misunderstanding between biologists … due in great part, I believe, to the fact that [they] do not use such terms as heredity, variation, variable, variability, in precisely the same signification. … The employment of quantitative methods necessarily leads to the use of such expression in a more precise signification. Quite generally, the statistician speaks of a character as inherent whenever the number or “constant” B [in the equation Y=A+B.X] is greater than zero. … The distinction between continuity and discontinuity of variation, between inheritance of attributes and of variables do not seem to me to be of necessary importance for the theory of heredity; … The real and important distinction seems to lie between the phenomena of heredity within the race, and the phenomena of hybridization that occurs on crossing two races admittedly distinct. (Yule, 1902, 195-199)

Mendelism is concerned with hybridizations. Heredity represents the population-aspect of inheritance, whereas hybridization is the method to study the inheritance of specific difference characteristics between individuals. The Law of Ancestral Heredity is concerned with heredity, it is a law that regards the correlation of variance in one generation of the population with that of another; we would say that it is a law in population genetics. Yule made this most explicit a year later, opening his paper: “The statistical theory of heredity, as developed in the work of Galton and Pearson, concerns itself with aggregates or groups of the population and not with single individuals” (Yule, 1903). The statistician’s notion of “ancestral heredity” is: “will a knowledge of the grandparent’s character enable one to increase the accuracy of the estimate” of the character of the grandchildren above that obtained from the knowledge of the character of the parent? “If the answer to the question be in the affirmative, … then there is what may be termed a partial heredity from grandparent as well as from parent” (Yule, 1902, 201). The law that may be deduced is “that the mean character of the offspring can be calculated with the more exactness, the more extensive our knowledge of the corresponding characters of the ancestry, may be termed the Law of Ancestral Heredity” (Yule, 1902, 202). This is an early version of the breeders’ concept of later years of heritability, which was formulated in terms of Mendelian genetics as the ratio of genetically caused (additive) variability to total variability of a character in a population (Rieger, Michaelis, & Green, 1991). Noticeably, Yule pointed out that “it is difficult to suppose that the weight attached to pedigree is based on nothing but illusion” (Yule, 1902, 202) – it begs for a hypothesis, and this was provided by Mendel’s theory of hybridizations.. Fisher is generally regarded as “the first to successfully put forth a theory of the relationship between biometry, Darwinian evolution, and Mendelian inheritance,” and his paper “is considered to be a direct descendent of Yule’s earlier suggestion” (Tabery, 2004, 82). However, contrary to Yule’s perspective of the demographic fact of the inherited variation in the population for which Mendelism may or may not provide a good theoretical explanation, Fisher was an ardent Mendelian reductionist who endeavored to explain inherited variance of populations from his genocentric perspective. As put by Tabery, “For Fisher the ancestral law was a special case of the Mendelian principle. For Yule, the Mendelian principles were a special case of the ancestral law” (Tabery, 2004, 90-91). Fisher claimed that “if one first supposes Mendelian inheritance, then one can derive the correlation between relatives, resulting in the ancestral law of heredity”

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(Tabery, 2004, 83). In other words, Fisher provided a conceptual reduction of biometry’s law of ancestral heredity: He took the Mendelian principles of inheritance as the explanatory base and then derived the biometric law of ancestral heredity to show that the statistical law was just a special case of the physiological law (Sarkar, 1998, 106). Fisher was aware that he came upon “the Law of Ancestral Heredity as a necessary consequence of the factorial mode of inheritance” (Tabery, 2004, 83, quoting Fisher, 1918, 421). In the era when genocentric reductionist conceptions gained primacy, Mendelian explanation made the Law of Ancestral Heredity redundant. Thus Fisher laid the foundation for the Modern Synthesis of Huxley and colleagues two decades later, which turned the theory of evolution into a theory of gene frequencies or population genetics. But Yule, by juxtaposing Pearson’s support for the Ancestral Heredity Theory and Bateson’s claim for the Mendelian theory, also laid bare important aspects of the way scientific deductions are carried out. The fact that although the theory of ancestral contribution to heritage implies the law of ancestral heredity, the converse is not true: the law of ancestral heredity need not in any way imply actual physical contributions of the ancestry to the offspring. The ancestry of an individual may serve as guides to the most probable character of his offspring simply because they serve as indices to the character of the germplasm as distinct from his somatic characters. (Yule, 1902, 206)

As for Yule, Mendel’s Laws and their relation to the Law of Ancestral Heredity, cannot at least be “absolutely inconsistent” with each other, as Mr. Bateson contends. The Law of Ancestral Heredity is certainly a law of nature of wide generality which cannot be dismissed in such a fashion. Mendel’s Laws I assume to be true also. The problem is to delimit their respective spheres, and shew in what way the one type of law may pass into the other, or the two even coexist. (Yule, 1902, 207)

Whereas Pearson and Galton deduce (describe) backwards, to ancestral generations, presumably with no hypothesis implied, Mendel’s laws, ex hypothesis deduce forward, to future generations (Gayon, 2000). Starting with Galton-Pearson’s descriptive law: The value of the work of Mendel and his successors lies not in discovering a phenomenon inconsistent with that law [of Ancestral Heredity], but in shewing that a process, consistent with it, though neither suggested nor postulated by it, might actually occur. (Yule, 1902, 227)

No doubt, conceptual reductionism or genocentricity that became the dominant driving force in genetic research was a hypothesis-driven project at the price of excluding other alternative hypotheses. However, the positivists’ claim of merely describing the phenomena with no hypothesis involved is misleading and similarly a dangerous illusion. The evolution of science along eras of paradigmatic science, punctuated by periods of scientific revolutions, reflects the struggle inherent in these perceptions. It may, however, be claimed that these alternations of paradigmatic and revolutionary science are only the phenotypes of a more constant scientific-type

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that obtains a wide range of norms of reaction of conceptual struggles, which, even if explicitly denied, are constantly there.

Genocentricity and its discontents I cannot elaborate on the development of the conception of genetic reductionist determinism over the later decades of the twentieth century, and it is out of the scope of this conference. It must, however, be kept in mind that the origins of genocentricity or more accurately hereditarydeterminism, so dominant a factor in biological thought and practice throughout most of the twentieth century, may be traced to the late nineteenth century. With the intensification of the interrelationship of science and society, and the appearance of the scientists as professionals in their own right, biological determinism became a weapon, as well as a shield, of the growing community of scientists that considered it their duty to emphasize the social relevance of their science, yet had to face their inability to provide unequivocal answers. Eugenics is, of course, the first that comes to mind. Galton’s Law of Ancestral Heredity, much like Darwin’s pangenesis, was an ad hoc hypothesis, in which environmental effects had to be explained away. Ruth SchwartzCowan noted in 1977 that of all things that Galton wrote in Heredity Talent and Character in 1865, “he ended his argument against the inheritance of acquired characteristics with a rough statement of the continuity of germ plasm, the principle that we usually associate with the name of August Weismann and with the year 1883, …[as] the result of sociopolitical rather than biological imperatives” (Schwartz Cowan, 1977, 142, my emphasis). A few years later Galton created a physiological theory of heredity, the stirp theory that embodied the principle of continuity of germ plasm. “The idea of continuity of germ plasm was absolutely essential for Galton’s scheme” of human breeding if it had to compete with that of indoctrination or education in averting the social problems of his time (Schwartz Cowan, 1977, 143). More recently, John Waller pointed out that “heredity came to imply relative fixity and resistance to therapeutic intervention. Moreover, the concept often provided physicians and alienists with a useful rationalization when therapeutics proved impotent” (Waller, 2001, 461). He quoted a poem of John Byrom, probably written in the 1730s, but published in 1894, that beautifully exposed the inclination of scientists to explain away phenomena by hereditary determinism, which was formulated by providing names: When our distempers did their name receive, (One instance more, good doctors, by your leave), Some chronic matters, such as gout and stone, That would the fare of no arcana own, To save their Credit these, the learned dons, Cried out, were fix’d hereditary ones: If a man’s father, grand- or great-grand sire; Had the same, ‘twas needles to enquire; Plain was the case, and safe the doctor’s fame; The poor old ancestors bore all the blame.

Whichever way we take this, the differentiation of the phenotype and the genotype was superposed on the Weismannian segregation of the germ-line from the soma, and it made sense of “transmission genetics” – a study of heredity independently of development. But not less

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significant a consequence of this reductionist, bottom-up disjunction that Entwickelungsmechanik forced on embryology was the “remarkable awakening of interest and change of opinion” among working embryologists with respect to evolution. Leo Buss in 1987 in his The Evolution of Individuality noted that an “ugly fact” which remained from this bottom-up approach to biology has been that “evolutionary biologists have only rarely been able to make specific predictions regarding the patterns studied by the reductionist research tradition.” This situation changed only when “molecular biology has suddenly become a comparative, and inevitably evolutionary discipline. A new ‘fossil record,’ writ in the genome, is now accessible and being read in a necessary piecemeal fashion.” Noticeably, the breakthrough has been technological before it became conceptual (Buss, 1987, vii). Let me, then, round up my discussion by pointing out that with the presentation of the structural organization of DNA in 1953, the notion of genetic reductionism, and inadvertently also that of genetic determinism, got a strong boost. As Watson and Crick (1953) pointed out at the conclusion of their first paper, “It has not escaped our notice that the specific pairing we have postulated immediately suggests a possible copying mechanism for the genetic material,” both for replication and for catalyzing specific gene products. Notwithstanding, molecular biology, the triumph of genetic reductionist determinism, already carried the seeds of discontent. The detailed power of the molecular analysis challenged the old concepts by confronting them to the utmost. Many human and other organisms’ traits which follow Mendelian patterns of inheritance have been labeled “genetic,” and many of the genes involved have been mapped, their mutations identified, and screening programs for them instituted. Such a “geneticization,” or the tendency to reduce trait differences to genetic ones is expressed in the increasingly popular notion of “the gene for” (Gannett, 1999; see also Nelkin and Lindee 1995). It reflects the extreme deterministic image that genetics acquired. However, an increasing awareness of the more complex traits, ones for which environmental contributions are known to be significant, and whose genetic component could not be attributed to a single gene, emphasized the urgent need for theoretical and empirical methods that would allow a system approach. As it turned out, toward the end of the century, the ambitious technical project of mapping the complete human genome, that had raised many eye-brows among theoretical scientists and philosophers, generated – out of necessity – new tools, empirical as well as theoretical, to meet the challenge. Consequently, we enter the twenty-first century to a large extent with a challenge similar to that of the beginning of the previous century, but this time we are much better equipped to face it both empirically and theoretically. Comparing the wording of a recent statement by Francis S. Collins, head of the National Human Genome Research Institute at a workshop on Human Genome Variation and ‘Race’ at Howard University, on May 15, 2003, with one of ninety years earlier, which expresses exactly the same notion, by the eugenics-determinist Charles B. Davenport, makes this amply clear: In many instances, the causes of health disparities will have little to do with genetics, but rather derive from differences in culture, diet, socioeconomic status, access to health care, education, environmental exposure, social marginalization, discrimination, stress and other factors. Yet it would be incorrect to say that genetics never has a role in health disparities. …

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The question of whether genetics will explain a substantial proportion of health disparities … is largely unanswered … (Collins, 2004)

And: In the study of human heredity it has first of all to be recognized that progress will be made only as traits are studied one at a time. The modern science of heredity indeed seeks as the element of study the “unit character.” What are unit characters can, however, be told only by breeding experiments in which the true units reveal themselves as relatively, if not absolutely, constant, unalterable, indivisible things. … The first step in the resolution of human traits is, then, a primary rough analysis into fairly simple traits and, second, the study of the behavior of these traits in heredity. … …The criticism may be made that not all the traits, especially the diseases, given here have any hereditary basis. It is not affirmed that this is the case and yet it cannot be denied that all have an hereditary basis. Even tuberculosis, syphilis, and the plague are the product of a specific germ acting on a susceptible protoplasm and it is this susceptibility that is the inheritable factor. (Davenport, 1912)

To close the circle, we have to return to Roux’s emphasize of the need for simple methodologies, and of the caution that must be exerted in interpreting the data as simple concepts: Among biologists there is a tendency derived from inorganic sciences, to regard the hypothetical deductions which appear to us to be the “simplest” as having the greatest probability for the very reason that they seem so simple. [But a]lthough much has been done on this assumption, and unfortunately must be done, … nevertheless this method must always be applied with great reserve to normal biological phenomena, … Thus we suppose that we are really simplifying matters when, e.g., we attribute in consequence of functional adaptation many typical and purposive forms to the self-constructive effects of use. The correctness of this principle and its application in many cases has long been capable of direct proof. Nevertheless, we observe that many structures which might be the result of this principle, … are already established before there is an opportunity for them to exercise their definite functions. (Roux, 1894, 119)

Only “forward deductions” from experiments that control the causal force, can be “certain.” There is no sense (at least in living systems) in isolating one causal force, independently of other causes, but this was the only way that a cause could be studied directly. Roux would have been happy if it were possible “to reproduce synthetically in an inorganic way structures, forms, and processes which resemble as closely as possible, those of the organic world.” But as he noted, “even in cases where it is claimed that such a reduction has been brought about, it appears that the part which the simple components contribute to the formation in question, as compared with that of the coöperant complex components, has been considerably overestimated” (Roux, 1894, 128129). Embryologists, in view of such a program as Roux’s, were caught between holistic concepts of organisms as complex systems that cannot be simplified and studied effectively by the methods of physicists and chemists, and the reductionist need to apply experimental mechanics in order to study “real” causes. Geneticists by having been offered distinct and discrete entities, the genes,

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believed that they could successfully apply not only methodological, but also conceptual reductionism. Since then we have advanced in our molecular and computational technologies, and the old dream may again appear luring to the modern genetic engineer. Yet, the same problem may exist in today’s studies in genomics which try to follow simultaneously many individual causes. Unless a conceptual change takes place, the risk remains that the methodologically isolated forces would be inadvertently identified with the conceptually assumed forces. The challenge, as I see it, is to sort out the reductionist methods of empirical research – which have been and will be irreplaceable in research in the life-sciences in general, and in genetic research in particular – from the notions of conceptual reductionism that may have had tactical, heuristic and instrumental value, but must be replaced by systems’ conception, which are, after all, that unique specificity of life on earth. Griesemer’s (2000) suggestion that we should think of both development and inheritance as processes reducible to reproduction may be a step in the direction of maintaining methodological reduction as an efficient research instrument within the framework of organisms as integrated systems. If we return, for a moment to the 1920s and 1930s, when Niels Bohr and Erwin Schrödinger wondered “What is Life,” and physicists, in the footsteps of Max Delbrück flocked into biological research, to discover the specific Laws of Nature hidden in living creatures, we may say now, with some confidence: There are no special laws of the living world, neither Lebenskraft, nor Entelechy, nor any need for an Anthropomorphic Principle or divine power. It is just the evolution of the material world, as put forward by Darwin, and contrary to that believed by Mendel. Raphael Falk, Hebrew University, Jerusalem [email protected]

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Mendel’s impact

References Allen, G. E. 1978. Thomas Hunt Morgan: The Man and His Science. Princeton, NJ: Princeton University Press. –––. 2002. The classical gene: Its nature and its legacy. In L. S. Parker & R. A. Ankeny (Eds.), Mutating Concepts, Evolving Disciplines: Genetics, Medicine and Society. Pp. 11-41. Dordrecht: Kluwer. Ankeny, R. A. 2000. Marvelling at the marvel: The supposed conversion of A. D. Darbishire to Mendelism. Journal of the History of Biology, 33, 315-347. Bateson, P. 2002. William Bateson: a biologist ahead of his time. Journal of Genetics, 81(2), 49-58. Bateson, W. [1905] 1928. A suggestion as to the nature of the “walnut” comb in fowls. In R. C. Punnett (Ed.), Scientific Papers of William Bateson. Vol. II, pp. 135-138. Cambridge: The University Press. –––. 1916. The Mechanism of Mendelian Heredity: A Review. Science, 44, 536-543. Beadle, G. W., & Ephrussi, B. 1936. The differentiation of eye pigments in Drosophila as studied by transplantation. Genetics, 21(3), 225-247. Beadle, G. W., & Tatum, E. L. 1941. Experimental control of development and differentiation: Genetic control of developmental reactions. The American Naturalist, 75, 107-116. Buss, L. W. 1987. The Evolution of Individuality. Princeton: Princeton University Press. Castle, W. E. 1906. Yellow mice and gametic purity. Science n.s., 24, 275-281. –––. 1919. Piebald rats and the theory of genes. Proceedings of the National Academy of Science, Washington, 5, 126-130. Collins, F. S. November 2004. What we do and don’t know about ‘race’, ‘ethnicity’, genetics and health at the dawn of the genome era. Nature Genetics, 36(11), s13-s15. Correns, C. 1900. G. Mendel’s Regel über das Verhalten der Nachkommenschaft der Rassenbastarde. Berichte der deutschen botanischen Gesellschaft, 18, 158-168. Darwin, C. 1868. The Variation of Animals and Plants under Domestication (Vol. 1 & 2). London: Murray. Davenport, C. B. 1912. The Trait Book. Cold Spring Harbor: Eugenics Record Office. de Vries, H. 1902-3. Die Mutationstheorie: Versuche und Beobachtungen Über die Entstehung von Arten im Pflanzenreich. Leipzig: Von Veit. Falk, R. 1986. What is a gene? Studies in the History and Philosophy of Science, 17(2), 133-173. –––. 2000. Can the norm of reaction save the gene concept? In R. S. Singh & C. B. Krimbas & D. B. Paul & J. Beatty (Eds.), Thinking About Evolution: Historical, Philosophical and Political Perspectives. Vol. 2, pp. 119-140. Cambridge: Cambridge University Press. –––. 2001. Mendel’s hypothesis. In G. E. Allen & R. M. MacLeod (Eds.), Science, History and Social Activism: A Tribute to Everett Mendelsohn. Pp. 77-86. Dordrecht: Kluwer Academic Publishers. Falk, R., & Schwartz, S. 1993. Morgan’s hypothesis of the genetic control of development. Genetics, 134, 671674. Fisher, R. A. 1918. The correlation between relatives on the supposition of Mendelian inheritance. Transactions of the Royal Society, Edinburgh, 52, 399-433. Francke, A. H. [1907] 1977. A History of Ladakh. New Delhi: Sterling. Gannett, L. 1999. What’s the cause? The pragmatic dimension of genetic explanation. Biology and Philosophy, 14(3), 349-374. Gayon, J. 2000. From measurement to organization: A philosophical scheme for the history of the concept of heredity. In P. J. Beurton & R. Falk & H.-J. Rheinberger (Eds.), The Concept of the Gene in Development and Evolution: Historical and Epistemological Perspectives. Pp. 69-90. Cambridge and New York: Cambridge University Press. Griesemer, J. R. 2000. Reproduction and reduction of genetics. In P. J. Beurton & R. Falk & H.-J. Rheinberger (Eds.), The Concept of the Gene in Development and Evolution: Historical and Epistemological Perspectives. Pp. 240-285. Cambridge and New York: Cambridge University Press. Haldane, J. B. S. 1954. The Biochemistry of Genetics. London: George Allen & Unwin. Harman, O. S. 2004. The Man Who Invented the Chromosome: A Life of Cyril Darlington. Cambridge, MA: Harvard University Press. Hartsoeker, N. 1694. Essay de Dioptrique. Paris: Jean Anisson. Harwood, J. 1987. National style in science: Genetics in Germany and the United Stated between the World wars. Isis, 78, 390-414. Johannsen, W. 1909. Elemente der exakten Erblichkeitslehre. Jena: Gustav Fischer. –––. 1911. The genotype conception of heredity. The American Naturalist, 45(531), 129-159. –––. 1923. Some remarks about units in heredity. Hereditas, 4, 133-141.

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Kühn, A. 1941. Über eine Gen-Wirkkette der Pigmentbildung bei Insekten. Nachrichten der Akademie der Wissenschaften in Göttingen, Mathematisch-Physkalische Klasse, 231-261. Laubichler, M. D., & Sarkar, S. 2002. Flies, genes, and brains: Oskar Vogt, Nikolai Timoféeff-Ressovsky, and the origin of the concepts of penetrance and expressivity. In L. S. Parker & R. A. Ankeny (Eds.), Mutating Concepts, Evolving Disciplines: Genetics, Medicine and Society. Vol. 75, pp. 63-85. Dordrecht: Kluwer. Lenoir, T. 1982. The Strategy of Life. Chicago: University of Chicago Press. Maienschein, J. 1987. Heredity/development in the United States, circa 1900. History and Philosophy of the Life Sciences, 9, 79-93. Morgan, T. H. 1910. Chromosomes and Heredity. The American Naturalist, 44, 449-498. –––. 1934. Embryology and Genetics. New York: Columbia University Press. Muller, H. J. 1950. Evidence of the precision of genetic adaptation. The Harvey Lectures, 43, 165-229. Nelkin, D., & Lindee, M. S. 1995. The DNA Mystique. The Gene as a Cultural Icon. New York: W. H. Freeman and comp. Olby, R. 1985. Origins of Mendelism (2nd ed.). Chicago: University of Chicago Press. Provine, W. B. 1971. The Origins of Theoretical Population Genetics. Chicago and London: The University of Chicago Press. Punnett, R. C. 1950. Early days of genetics. Heredity, 4(1), 1-10. Rheinberger, H.-J., & Müller-Wille, S. 2004. Gene, Stanford Encyclopediaof Philosophy. (in press) Rieger, R., Michaelis, A., & Green, M. M. 1991. Glossary of Genetics and Cytogenetics (5th ed.). Berlin: Springer-Verlag. Roll-Hansen, N. 1978. The genotype theory of Wilhelm Johannsen and its relation to plant breeding and the study of evolution. Centaurus, 22, 201-235. Roux, W. 1894. The problems, methods, and scope of developmental mechanics. In J. Maienschein (Ed.), Defining Biology: Lectures from the 1890s. Pp. 104-148. Cambridge, MA: Harvard University Press. Sandler, I., & Sandler, L. 1985. A conceptual ambiguity that contributed to the neglect of Mendel’s paper. History and Philosophy of the Life Sciences, 7, 3-70. Sarkar, S. 1998. Genetics and Reductionism. Cambridge: Cambridge University Press. Schwartz Cowan, R. 1977. Nature and nurture: The interplay of biology and politics in the work of Francis Galton. In W. Coleman & C. Limoges (Eds.), Studies in the History of Biology. Pp. 133-208. Baltimore and London: The Johns Hopkins University Press. Schwartz, S. 1998. The significance of the trait in genetics, 1900-1945. Unpublished Ph.D. Dissertation, The Hebrew University of Jerusalem, Jerusalem. –––. 2002. Characters as units and the case of the presence and absence hypothesis. Biology and Philosophy, 17(3), 369-388. Stadler, L. J. 1954. The gene. Science, 120, 811-819. Stamhuis, I. H. 2003. The reactions on Hugo de Vries’ Intracellular Pangenesis; the discussion with August Weismann. Journal of the History of Biology, 36(1), 119-152. Stamhuis, I. H., Meijer, O. G., & Zevenhuizen, E. J. A. 1999. Hugo de Vries on heredity, 1889-1903: Statistics, Mendelian laws, pangenes, mutations. Isis, 90(2), 238-267. Stern, C., & Sherwood, E. 1966. The Origin of Genetics: A Mendel Sourcebook. San Francisco: W. H. Freeman. Sturtevant, A. H. 1923. Inheritance of the direction of coiling in Limnaea. Science, 58, 269-270. –––. 1932. The use of mosaics in the study of the developmental effects of genes, Proceedings of the 6th International congress of Genetics, Ithaca, 304-307. Tabery, J. G. 2004. The “evolutionary synthesis” of George Udny Yule. Journal of the History of Biology, 37(1), 73-101. Vogt, O. 1926. Psychiatrisch wichtige Tatsachen der zoologisch-botanischen Systematik. Zeitschrift für die gesamte Neurologie und Psychiatrie, 101, 805-832. Waller, J. C. 2001. Ideas of heredity, reproduction and eugenics in Britain, 1800-1875. Studies in History and Philosophy of Biology and the Biomedical Sciences, 32(3), 457-489. Watson, J. D., & Crick, F. H. C. 1953. Molecular structure of nucleic acids. Nature, 171, 737-738. Weismann, A. 1893. The Germ-Plasm. A Theory of Heredity. New York: Charles Scribner’s Sons. Wilson, E. B. 1893. The mosaic theory of development. In J. Maienschein (Ed.), Defining Biology: Lectures from the 1890s. Pp. 67-80. Cambridge, MA: Harvard University Press. Woltereck, R. 1909. Weitere experimentelle Untersuchungen über Artveränderung, speziell über des Wesen quantitativer Artunterschiede bei Daphnien. Verhandlungen der Deutschen Zoologischen Gesellschaft, 19, 110-173.

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Yule, G. U. 1902. Mendel’s laws and their probable relations to intra-racial heredity. The New Phytologist, 1(9 & 10), 193-207 & 222-238. –––. 1903. Professor Johannsen’s experiments in heredity: A review. The New Phytologist, 2, 235-242.

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The Biometric Sense of Heredity: Statistics, Pangenesis and Positivism Theodore M. Porter

The first commentators on almost every scientific episode are the participants themselves, whose categories of analysis provide a starting point for proper historical studies. But one task of the history of science is to gain some distance from scientific debates and to see them against a wider background in ways the protagonists could not. This generally means, at a minimum, refusing to pass along as historical truth the judgment of the winners. In fact there was no clear winner, or at least no loser, in the controversy between Biometricians against Mendelians. But the statistical program of population genetics has remained mostly distinct from laboratory studies at the level of individual genes, which now usually means molecular genetics, and accordingly, somewhat discrepant histories have also been perpetuated. Add to this the intensity of the controversy at the beginning of the twentieth century and some bitter rivalries within the statistical program involving such leading figures as Karl Pearson, Udny Yule, R. A. Fisher, and Sewall Wright, and we have a landscape littered with polemical arguments to seduce and sometimes mislead the historian.

Blending inheritance or blended characters? The great evolutionist Theodosius Dobzhansky identified the deficiencies of the theory of blending inheritance in his magnum opus of 1937, Genetics and the Origin of Species. This concept, he explained, “was based on the assumption that the germ plasms of the parents undergo a sort of amalgamation in the hybrid.” They dissolve as dye mixes into water, uniformly. “The amount of variation present in a sexually reproducing random breeding population must be halved in every generation. Given a population which exhibits a large variability at the start, we are bound to observe a progressive, rapid, and irretrievable decay of the variability, until a complete homogeneity is reached.” Darwin and his follower were “inexorably driven to this conclusion,” Dobzhansky continued, and this is why the new Mendelian approaches after 1900 were so urgently needed to complete the theory of evolution. The “particulate” theory of the gene allowed evolution to be understood in a new way. “Even where the hybrid or heterozygote is intermediate between the parents, no contamination of the genes takes place, and the homozygotes recovered from the hybrids are like the parental races.” A particulate germ plasm automatically maintains variability “on an approximately constant level despite the interbreeding.” 1 Historians of science have come to recognize blending inheritance as one of the principal obstacles to an adequate theory of evolution. Robert Olby, for example, explains: “It is now well known that the most unfortunate of the assumptions underlying Darwin’s mechanism of evolution was that of blending heredity; i.e. that parental differences are merged in the offspring of bisexual reproduction so that variation is constantly being diminished.” 2 Peter Bowler seems to 1 2

Dobzhansky (1937), pp. 121– 122. Olby (1966), p. 55.

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adopt a completely different view: “We shall see that Darwinism would function quite effectively with a model of blending inheritance.” Yet in his interpretation of the disagreement between biometricians and Mendelians, he sounds very much like Dobzhansky: “In effect, Mendelism undermined Fleeming Jenkin’s claim that natural selection was ineffective because new characters will be swamped by interbreeding with unchanged individuals. If heredity is particulate rather than blending a favourable new character will be preserved intact so that its frequency can be gradually increased within the population.”3 And Alfred Sturtevant, the eminent geneticist turned historian of genetics, explained how Darwin’s reliance on blending implied that favorable variations would be swamped, retarding or even blocking evolutionary change. 4 Some other biologist–historians have been more perceptive about blending. Ernst Mayr, in a review, called Olby’s treatment of blending inheritance “thoroughly confused,” and referred to the “myth,” left intact by Olby, that Mendel’s greatest contribution was his “concept of particulate inheritance, in opposition to the ‘prevailing’ idea of blending inheritance.” There is, he continued, abundant evidence that Darwin did not support blending inheritance. 5 Michael Bulmer, in a recent biographical study of Francis Galton, comments that this historical interpretation of blending inheritance originated with R. A. Fisher, whose 1930 book The Genetical Theory of Natural Selection is the most complete expression of his successful effort to integrate Mendelian genetics into a mathematical version of Darwinian evolution. Bulmer calls the claim misleading and complains that it has “been uncritically accepted by many evolutionary biologists.” Dobzhansky, as we can see, was among them, for he too cited Fisher’s book as well as an earlier paper in Russian by Sergei Chetverikov on the baleful effects of the blending theory on early Darwinian theories of evolution. Bulmer offers an important distinction between “physical blending of the hereditary particles during fertilization, which is what Fisher meant, and the blending of phenotypes in the sense that offspring are often intermediate between their parents.” 6 Fisher’s analysis of Darwin’s assumptions about inheritance was based on a reading of manuscripts from 1842 and 1844 and letters to Huxley as well as Darwin’s books, and is not without merit as historical interpretation. The conclusions he drew, however, went well beyond Darwin. Since half of all variance will disappear in each generation in a sexually–reproducing population that mates randomly, a huge mutation rate is required to maintain variability. This, he suggested, was why Darwin put so much emphasis on the causes of variation. The concepts Fisher invoked, obviously, are anachronistic. He was more profoundly misleading, however, in leaping directly from Darwin’s era to his own, as if nothing but Mendelian genetics had happened since 1859 (or at least since Darwin’s Variation of Plants and Animals under Domestication in 1868), until Fisher himself stepped heroically forward to demonstrate the need for particulate rather than blending inheritance and to demonstrate the mathematical advantages of the substitution. 7 The concept of “blending inheritance,” unknown to Darwin, was not an actor’s category even in the era of Bateson and Pearson. Indeed, Fisher may have coined the phrase. “Blended inheritance,” which sounds at first like the same thing, was a well–known biological concept 3 4 5 6 7

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Bowler (1989), pp. 48, 63, 139. Sturtevant (1965), pp. 20– 21. Mayr (1973), p. 141 Bulmer (2003), p. 139. Fisher (1930), chapter I.

The Biometric Sense of Heredity

before 1900. Galton deployed the term in 1889 in his Natural Inheritance in a way that suggests something new. The children of a white and a negro, he wrote, “are neither wholly white nor wholly black, neither are they piebald, but of fairly uniform mulatto brown. The quadroon child of the mulatto and the white has a quarter tint; some of the children may be altogether darker or lighter than the rest, but they are not piebald. Skin colour is therefore a good example of what I call blended inheritance.” He immediately made it clear that blending implied no theory of heredity, but only described the results of certain crosses. “It need be none the less ‘particulate’ in its origin, but the result may be regarded as a fine mosaic, too minute for its elements to be distinguished in a general view.”8 A few pages back, Galton had explicitly endorsed a theory of “particulate” inheritance—the word, he said, is “good English” though the quotation marks embracing it implied unfamiliarity. There were no grounds for supposing unfamiliarity when Fisher called for a particulate theory of inheritance in 1930. Galton illustrated particulate inheritance by comparing the formation of an organism to one of those Italian buildings whose elements, stones and columns and pediments, had been pillaged from previous structures, its progenitors.9 There was no theory of blending inheritance in the early twentieth century, but only a distinction between those characters that blend in the offspring of hybrids and those that “alternate,” which meant to assume one or another discrete form. The entry on heredity in the eleventh edition of the Encyclopaedia Britannica, published in 1910, explains that the discrete forms studied by Mendel as well as the saltatory “mutations” made famous by Hugo de Vries pertain “almost exclusively to the crossing of artificial varieties of animals and plants” and “point strongly to the occurrence of alternate inheritance instead of blended inheritance for artificial varieties. On the other hand, in the case of natural varieties it appears that blended inheritance predominates.”10 This identification of mutations and Mendelian factors with hybridity was widely shared in the first decade of the twentieth century, before T. H. Morgan’s fly lab redefined mutation as a change at the level of a single gene. Karl Pearson’s student Udny Yule, in his effort to reconcile Mendelian with biometric studies of heredity, argued that Mendelians and statisticians used the same words in different ways. Statisticians, he explained, understand heredity as the study of how the divergence of an individual from a racial mean relates to the divergence of its offspring. This was not the object of investigation for Mendelians. “On passing from the Law of Ancestral Heredity to Mendel’s Laws, we are passing from a law of intra–racial individual heredity to a series of laws based solely on hybridisation–experiments, and clearly stated by their discoverer as laws of hybridisation only.”11 These remarks suggest that, to allude to a classic paper by Olby, Mendel may have been a Mendelian after all, since the first generation of Mendelians was scarcely less preoccupied with hybrids than he was.12 In a 1905 symposium in Philadelphia on the mutation theory, William Castle followed Darwin and de Vries in their use of artificial selection as a source of insight into biological evolution. Nature and breeders alike rely overwhelmingly on mutations. 8 9 10 11 12

Galton (1889), p. 12. Ibid., pp. 7– 8. Mitchell (1910), pp. 353–354. Yule (1902), pp. 196, 222. See Tabery (2004). Olby (1979).

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The latter do not simply wait for these to happen, but work actively to create them through cross– breeding.13

Blending and swamping As Dobzhansky’s argument exemplifies, the significance of blending inheritance has from the beginning been associated with an argument that, absent some mechanism of preserving variants, they would disappear due to “swamping” as exceptional individuals bred back into the parent population. In 1867, Fleeming Jenkin offered what is still often regarded as a damaging critique of Darwin’s theory of evolution by natural selection. He argued that indiscriminate mating would swamp the effects of any new variant, which would ineluctably be averaged away in subsequent generations unless the exceptional individual could find other similarly–endowed specimens with which to reproduce.14 In the era of Fisher and Dobzhansky, Mendelian genetics was celebrated as the first adequate response to this conundrum. It is, in its modern form, a curious argument, since it seems to suppose that variants leading to new species must be discrete and exceptional, yet subject to attenuation through continuous processes of mixing or blending. Jenkin, who understood the argument better than most of his interpreters, did not apply it to continuous variation of the sort that would be found in every generation. Although he doubted the capacity of new species to arise through selection on continuous variation, this was for other reasons. He argued that such variation would run up against limits, and could not support the substantial changes of form required for a new species. This doctrine of limits to variation was a common and enduring reaction to Darwin. Skeptics of Darwinian evolution in the early twentieth century, when the mutation theory of de Vries was all the rage, distinguished between continuous or “fluctuating variations,” which go nowhere, and mutations, which can provide a new stable center for a species. Some, including Castle, made the distinction virtually a matter of definition: “mutations are permanent, variations transitory;” and some naturalists sympathetic to Darwin such as Asa Gray as well as opponents like Louis Agassiz believed that small variations were mere oscillations around a normal state which could not be inherited.15 De Vries and his contemporaries did not rely on the argument of swamping to discredit evolution by selection on small variations. Neither did Jenkin, who instead used his swamping argument to deny the possibility that new species could arise through discontinuous variation or “sports.” More than forty years ago, Peter Vorzimmer showed that Darwin perfectly understood this aspect of Jenkin’s argument, and that if confirmed him in his gradual shift away from any reliance on sports and toward a belief in nature without leaps: Natura non facit saltum. Saltations, which he already regarded as too rare and too often infertile to be the main engine of species change, now appeared unlikely to perpetuate their own kind even when they did survive and reproduce. The objection applied most forcefully to sports that blended, and Huxley held out hope that new species might form as a result of nonblending ones. Darwin, however, emphasized more than ever the gradual change that could

13 14 15

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Castle (1905). Jenkin (1867). Castle (1905), p. 524; Vorzimmer (1963), p. 380.

The Biometric Sense of Heredity

occur as selection acted on those small favorable variations that should appear relatively abundantly in every generation.16 Galton, progenitor of biometry, wrote of blending and discrete inheritance as the radical alternatives and argued that few if any traits conform entirely to either extreme. He minimized the force of Jenkin’s critique, even for the case of hybridization as in the imagined presence of a lone white individual within a wholly black population, concluding “that the establishment of a somewhat rare variety as that of white men naturally suited to thrive and multiply in tropical climates” was not such an improbability as some supposed. While he recognized that “mutually– exclusive heritages” such as eye color will be distributed differently from blended traits, his example was about hybridization (at least in a loose sense). Nowhere did he consider that blending must lead to collapse of variation within the race or species, as argued by Fisher and Dobzhansky, and there is no hint of a suspicion that he thought micro–particulate inheritance was needed to preserve variability.17 Karl Pearson, similarly, worked out a descriptive account of gradual biological change, a detailed quantitative elaboration of evolutionary mechanisms that could work on continuous variation even under conditions of indiscriminate mating. He argued that it did not depend on any particular mechanism of heredity. Like Darwin, he supposed that biological variation was natural and fundamental, and not at all exceptional in the way that Fisher understood mutations to be. Pearson’s solution was in some ways a graphical version of Darwin’s, translated however into a statistical idiom of frequency distributions that was mostly alien to Darwin.18 Pearson’s version was enhanced to include the quantitative effects of heritability and differential fertility as well as survival. Not the appearance of mutant individuals but a slowly shifting frequency distribution was the mark of evolutionary change for Pearson. The raw material of evolution appeared like statistical clockwork in each generation. 19

Pangenesis and statistics Although Darwin placed increasing emphasis in the 1850s and 1860s on continuous variation, he was not in any straightforward way an advocate of blending inheritance. Bulmer shows how he restated Jenkin’s (flawed) quantitative demonstration that a singular variation, even if advantageous, must disappear over time, so that half the offspring would possess the variation and half not rather than, as in Jenkin’s version, having the variation attenuate by half in each successive generation.20 The “Provisional Hypothesis of Pangenesis” that he published in his big book on variation in 1868 was itself, after all, a particulate theory. His gemmules, to be sure, were very numerous, perhaps one for every cell in the body, and examples like the blending of skin color in mulattoes convinced him that these gemmules must merge as they form such tissues, so that in the next cycle of reproduction they would be intermediate in color.21 Others, most notably his cousin Galton, rejected all blending in favor of a mosaic theory of the relations of gemmules. Galton’s model of hereditary transmission was thus inspired by Darwin’s and used the same terms, but his 16 17 18 19 20 21

Vorzimmer (1963). See Bulmer (2004) for some clarifications and corrections. Galton (1887), pp. 400– 402. Gayon (1992), pp. 95– 112. Pearson (1900), chapters X–XI. Bulmer (2004), p. 290. Olby (1966), pp. 70– 71.

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gemmules did not blend. On the contrary, Galton conceived mechanisms by which gemmules could join forces to produce visible features recognizably derived from one ancestor or the other. In terms of his architectural analogy, this involved the columns or porticos that were passed along intact to a new structure. In statistical terms, this insight provided the basis of his biological theory of correlation, which was based on gemmules that tended to stay together. 22 More fundamentally, the statistical study of heredity was anchored from the beginning in the theory of Pangenesis. In 1870, after all, statistics was the quantitative science of human populations, and Galton depended on analogies between gemmules and individuals in society in order to imagine a statistics of human heredity. Much of the historiography of heredity links the quantitative biometric approach to a strong descriptive positivism which dismissed as meaningless the idea of genetic elements. Pearson’s Grammar of Science is Exhibit One for the prosecution,23 but Pearson never rejected genetic units, and sometimes worked at mathematical methods to try to count them. As Ida Stamhuis has shown, the association of Pangenesis with statistics was not limited to Britain, but was also central to the work of de Vries in Amsterdam, both before and after he became familiar with Mendel’s work.24 Was Mendelian genetics required for an adequate theory of evolution? Bowler, as we have seen, suggests that it was, that Mendelism at last enabled biologists to escape Jenkin’s critique. But in the same breath he explains how T. H. Morgan and coworkers came to the crucial insight that evolution might occur through the accumulation of small mutations so as to appear continuous. 25 This last formulation, absent the language of mutations, is indistinguishable from the biometric version of evolution. In 1909 (repeated as late as 1925), Wilhelm Johannsen identified Galton’s pathbreaking statistical methods, more than Mendel’s discoveries, as the basis for a hereditary mechanism that could make sense of Darwinian evolution.26 And in fact the two were in many respects compatible, as the leading biometricians generally recognized. The Darwinian gemmules on which rested Galton’s statistics of heredity could readily be replaced by Mendelian genetic elements without fundamentally revising the statistics. In principle, statistical approaches favored a quantitative positivism, in which phenomena were counted and analyzed, and the underlying mechanisms put aside. As we will see, Pearson often proceeded this way, sometimes to the chagrin of his allies as well as the bewilderment of his opponents. The statistical study of heredity, however, was initially not at all positivistic, and an ontology of heredity particles was long connected to it. In our terms, the hypothesis of Pangenesis was perhaps more like a model than a theory, particularly as wielded by Galton. Nobody had ever detected a Galtonian gemmule or a pangene of de Vries, and nothing was known about the physiological processes that might produce them. Galton undertook, unsuccessfully, to demonstrate their presence in the bloodstream through transfusions among rabbits. Otherwise they were mostly theoretical entities, whose presumed properties were inferred in a somewhat circular fashion from the phenomena they were designed to explain, rather than tangible objects susceptible to laboratory manipulation. 22 23 24 25 26

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Porter (1986). The classic statement is by Norton, e.g. (1975a) and (1975b). For example, Stamhuis (2003). Bowler (1989), p. 138– 139. Johannsen (1926), p. 5. On Johannsen see work by Nils Roll–Hansen.

The Biometric Sense of Heredity

Like Darwin, Galton supposed there must be a very large number of gemmules, enough to permit reliable statistical conclusions to be drawn. He used an extended analogy with processes of political representation to develop the statistical aspects of heredity. The gemmules in a newly– fertilized egg would undergo first a segregation or “election” to determine which would form themselves into the embryo, and, later, a second segregation to sort out the ones that would be transmitted by this individual to the next generation. There was, he explained, no need for a special mechanism to assure that every part of the body was represented; the familiar regularities of statistics would take care of this. When he first outlined this model, Galton regarded the transmission of this genetic inheritance from generation to generation as a random process, akin to fair games of chance, like drawing balls from an urn. The sorting that determined which of these elements should “represent” the whole population of gemmules by determining the bodily characteristics of the offspring, on the other hand, was subject to the elective affinities of the gemmules. Such affinities were required to assure that a tall individual would tend to be more or less uniformly so rather than having very long legs but very short arms. 27 As Galton’s program of experimental breeding got underway, and again when he applied his analysis to a burgeoning collection of records of family traits, he revised his account of these biological processes and of the statistical methods appropriate for dealing with them. His first important mathematical result on heredity, based on pea–breeding experiments, was a law of “reversion,” a quantitative relationship between parent pea seeds and their descendants, which he announced in 1877. The plant grown from a large pea seed tended itself to produce large peas, though these were on average less exceptional than their parent. Using deviations from the mean of the whole population as his measures, he found that the average deviation of the offspring from a given size class of parent was a constant fraction of the deviation of the parent. Given that there was also considerable variation among offspring of parent peas from every size class, there were mathematical reasons that the fraction must be less than one if the overall variability is stable from generation to generation. But there remained, he thought, a biological aspect to the question: why were the offspring seeds on average less exceptional than their parents? His answer followed from his hereditary model. According to his version of Pangenesis, the gemmules that formed a given individual derived not only from their parents, but also from grandparents and great grandparents. The more remote these ancestors, the more numerous they must be, and hence the more similar to the general population. This shift toward the mean of the “race” was literally a reversion in the biological sense, a partial return to the traits of these more remote ancestors. 28 A decade later, when he revisited these questions of inheritance armed with a mass of human records, he was able to apply the same mathematical formulation. He changed the word, however, to reflect a new understanding of the biology of heredity. “Regression,” his new term, might involve the influence of ancestral forms, but designated now an innate tendency to return to type, which would correspond with the mean for the population. He explained to the Anthropological Section of the British Association in 1885: “The type is an ideal form towards which the children of those who deviate from it tend to regress…. The stability of a type would, I presume, be measured by the strength of its tendency to regress.”29 Or, in terms of Pangenesis as political 27 28

Galton (1873). Galton (1877); Porter (1986), p. 270– 296.

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Theodore M. Porter

analogy, the affinities that formed an individual out of genetic materials meant that the selection of gemmules to be expressed in the offspring was not random, but favored the prevailing type, the dominant party, or the race, from which most gemmules would have derived. This made Darwin’s scheme of evolutionary change, without leaps, problematical, because small variations would tend to return to the racial mean unless displaced by a sport or pushed away to a new point of stability by strong and insistent selective pressures. Regression, in this version, was closely allied to the biological correlation of parts, which was about harmonies or similarities among different organs or parts of a single organism. In 1888, just as he was finishing his book Natural Inheritance, Galton saw in a flash that the correlation of measurable traits could be understood with the same mathematics as he had developed for regression. As so often in his mathematical work, he came to this realization in the context of a specific quantitative problem, but it also made biological sense to him, for correlation and regression both followed from the same structure of elective affinities among gemmules.30 The strange positivism of Karl Pearson allowed, and was perhaps even favorable to a hypothesis of Pangenesis, on condition that the gemmules be regarded only as conceptual, and not projected into the world of sensations. Although Pearson’s working methods were often positivistic in the sense of not presuming any particular theory of heredity, he did not reject genes on epistemological or metaphysical grounds. He criticized Mendelism rather for its failure, as he saw it, to meet the standard of adequate agreement with the results of measurement and calculation. Genes and Mendelian factors, too, were scientifically legitimate, but only as handy formulas that are valuable to the extent they lead to correct conclusions. Already in 1900, before Mendel meant anything to him, he complained of August Weismann’s projection of Darwin’s gemmules “into the phenomenal world,” as if we can see or touch them. We should not succumb to mechanistic delusions that would transmute these inventions of mind into material explanations or causes of life.31 But Pangenesis was Pearson’s preferred model of hereditary processes, as it was Galton’s. He never acquired the same enthusiasm for Mendelian units, but there is simply no basis for the supposition that he rejected them on grounds of positivist philosophy.32 Although his quantitative research program was indeed positivistic in important respects, this did not exclude physiological mechanisms of heredity. The statistical “law of ancestral heredity” was potentially consistent with a plurality of models of heredity. 33 By 1900, when Pearson issued this complaint, he and his biological ally W. F. R. Weldon were already somewhat embattled over the proper uses of statistics in biology. The rapid rise of Mendelism, beginning that same year, sharpened the antagonisms. Pearson repeatedly emphasized that Mendelism was, in principle, scientifically legitimate, and should be judged according to its fruits. The gene, in this respect was no different from the pangene or the gemmule, except that some bold Mendelians were soon announcing a host of one–gene traits. For Pearson, as for many of his antagonists, eugenics gave particular urgency to the new science of heredity, and 29 30 31 32 33

38

Galton, (1885). Galton (1886). See also Galton (1889); Galton (1888); Galton (1890). Pearson (1900), pp. 335–337, from chapters that appeared only in the second edition of this work. I thus take issue with Norton (1975). MacKenzie offers the beginning of a critique in (1981), pp. 140– 141; Porter (2004), p. 269. Provine (1971), p. 25.

The Biometric Sense of Heredity

Pearson thought that irresponsible Mendelian claims “in the name of eugenics” were casting discredit on it. He argued that one–gene traits were very rare, and he devoted almost all of his own efforts to statistical investigations of complex quantifiable traits. The logic of his program reduced Mendelian efforts to a very minor role in evolutionary or eugenic studies, which mostly would be subsumed into the statistics. In the 1890s he worked out methods for ascertaining the number of gemmules that determined a limb or organ from the specific form of the asymmetrical frequency curve, and in the early twentieth century he tried to determine the number of genes required for such traits, as evidence that biometry was a necessary tool even for Mendelians engaged in studies of evolution and eugenics.

The graphical basis of the science of heredity The gap between biometry and Mendelism was not mainly one of philosophy, but of skills and practices. The statistical study of heredity was graphical and sometimes highly mathematical, and did not involve tight experimental control. It was characteristic of Pearson’s statistics, and a point on which he differed from his successor and bitter antagonist R. A. Fisher, that not too much was made of the difference between observation and experiment. Fisher also took a more sympathetic interest in Mendelism, as indeed did Pearson’s student, with whom however he was for some years at odds, Udny Yule. Although Pearson did not reject Mendelism, it was for him a narrow and rather uninteresting special case of a program that proceeded with other tools using other sorts of data. Like Galton, and for similar, mostly eugenic reasons, Pearson was very much concerned to distinguish environmentally–conditioned traits from those inherited biologically. But his preferred methods for this work involved instruments of measurement and quantitative analysis, not laboratory interventions. The distinctive strengths of zoology and botany, displayed in efforts to connect mechanisms of heredity with careful microscopy and investigation of cellular physiology, meant little to the biometricians. Pearson’s biometric style owed much to Galton, whom he learned to appreciate as a result of collaborations with his University College colleague Weldon. This much is well known, and should not be minimized. In my recent book on Pearson I try to show that his quantitative program derived also, and no less fundamentally, from his role as a professor of applied mathematics, which in practice meant engineering mathematics, and that his Biometric Laboratory drew inspiration from an Engineering Laboratory established by another University College colleague, Alexander Kennedy. There students were trained in the use of instruments to trace and measure curves, and in graphical methods to solve mathematical problems. Pearson originally took up statistics in the form of economic and social statistics as an important case of subjects that could be handled graphically. When, thanks to Weldon, he came to see evolution as one such subject, his excitement about this project grew uncontrollably, yet for three more years he still thought of statistical biology as a chapter in graphical statics. Some of the most fundamental concepts of Pearson’s new statistics derived from engineering graphics. Standard deviation, for example, was mathematically equivalent to swing radius in mechanics, and could be calculated on paper using the very same graphical methods.34 34

Porter (2004).

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Theodore M. Porter

Pearson outlined his graphical vision for the study of heredity and evolution most systematically in his final set of Gresham lectures in 1893 and 1894, and then again in the chapters on evolution he included in the second edition of his Grammar of Science, published in 1900. Galton, too, had relied heavily on graphical presentation, and Weldon used graphs not only to summarize data but also to solve problems. A double–peaked graph was probably responsible for Weldon’s original intuition that the Naples crabs he had been studying were undergoing incipient speciation, and he used graphical techniques to separate the compound curve, as he thought, into two simple ones. This was the problem that fired Pearson’s enthusiasm for the statistical study of evolution and the subject of his first “Contribution to the Mathematical Theory of Evolution” in 1894. There was a sense in which Pearson’s statistical practice was always informed by a geometrical or graphical sensibility, even in writing that took the form of vast algebraic tangles. The graphical structure was most emphasized in relatively popular presentations, as in The Grammar of Science, where he constructed a hypothetical frequency distribution of an unspecified trait and showed how it might change due to differential rates of mortality and then reproduction as functions of this trait. The last stage of the analysis involved heritability, the percentage of individual exceptionality that was passed along to the next generation, and the graphical exercise concluded with a new frequency curve representing the offspring generation. The difference between parental and offspring curves was a direct indication of evolutionary change, in a form that should not be compromised by interbreeding and did not depend on any particular mechanism of hereditary transmission. Pearson published this work in 1900, just before Mendelism became an issue for him.35 Pearson’s research papers for the Philosophical Transactions of the Royal Society, and later for Biometrika, were rarely graphical to the same extent. He came to see that his comparative advantage lay in explicit, algebraic solutions, which few if any biologists could duplicate or even follow. But the relation to mechanisms of inheritance was the same. Pearson’s assumptions about heritability were of a quantitative kind, and did not involve any clear assumptions about how traits were passed from parents to offspring. He could not accept, and indeed actively worked to undermine, Galton’s ideas about stability of type. Mendelian genes presented no particular problem, provided that he was allowed to work on them collectively, not individually. Reconciling Mendelism or Pangenesis with biometry was of moderate interest to him, but was far from central to his research agenda. He needed only to assume that most measurable traits—most traits that mattered for survival and reproduction—were composites of many elements and so could be handled statistically. It was not that genes were inherently unknowable or illegitimate, but that he expected more mileage out of a statistical research program, one that depended on counting and measuring. His studies could be called positivistic in a different, though related, sense, because Pearson was mostly uninterested in the biological details. Pangenesis, too, did not remain very important for Pearson’s biometry, but it had played a considerable role in the creation of the statistical study of heredity, and the rather indirect significance that it retained was supportive of

35

40

Pearson (1900).

The Biometric Sense of Heredity

biometry rather than antagonistic to it. When he discussed physical models of heredity they were always particulate, and theories of blending inheritance, if indeed there were any such in biological work on heredity in the early twentieth century, had no role whatsoever in Pearson’s biometry.

Theodore M. Porter Department of History, UCLA [email protected]

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Theodore M. Porter

References Bowler, Peter. 1989. The Mendelian Revolution. Baltimore: Johns Hopkins University Press. Bulmer, Michael. 2003. Francis Galton: Pioneer of Heredity and Biometry. Baltimore: Johns Hopkins University Press. ———. 2004. “Did Jenkin’s Swamping Argument Invalidate Darwin’s Theory of Natural Selection?” British Journal for the History of Science 37: 281– 297. Castle, W. E. 1905. “The Mutation Theory of Organic Evolution from the Standpoint of Animal Breeding.” Science, n.s. 21: 521– 525. Dobzhansky, Theodosius. 1937. Genetics and the Origin of Species. New York: Columbia Press. Fisher, Ronald Aylmer. 1930. The Genetical Theory of Natural Selection. Cambridge: Cambridge University Press. Galton, Francis. 1873. “On Blood–Relationship.” Proceedings of the Royal Society of London 20: 394– 402. ———. 1877. “Typical Laws of Heredity” Nature 15: 492–495, 512–514, 532–533. ———. 1885. “Address to the Section of Anthropology of the British Association.” Nature 32: 507– 510. ———. 1886. “Regression towards Mediocrity in Hereditary Stature.” Journal of the Anthropological Institute of Great Britain and Ireland 13: 246– 263. ———. 1887. “Address Delivered at the Anniversary Meeting of the Anthropological Institute of Great Britain and Ireland, January 25th, 1887” Journal of the Anthropological Institute of Great Britain and Ireland 16: 387– 402. ———. 1888. “Co–relations and their Measurement, chiefly from Anthropometric Data.” Proceedings of the Royal Society of London 45: 135–145. ———. 1889. Natural Inheritance. London: Macmillan and Co. ———. 1890. “Kinship and Correlation.” North American Review 150: 419–431. Gayon, Jean. 1992. Darwin et l’après Darwin: Une histoire de l’hypothèse de selection naturelle. Paris: Editions Kimé. Jenkin, Fleeming. 1867. “The Origin of Species.” North British Review 46: 277– 318. Johannsen, Wilhelm. 1926. Elemente der exakten Erblichkeitslehre mit Grundzügen der biologischen Variationsstatistik. 3rd ed. Jena: Verlag von Gustav Fischer. MacKenzie, Donald. 1981. Statistics in Britain, 1865–1930: The Social Construction of Scientific Knowledge. Edinburgh: Edinburgh University Press. Mayr, Ernst. 1973. “Essay Review: The Recent Historiography of Genetics.” Journal of the History of Biology 6: 125– 154 Mitchell, Peter Chalmers. 1910. “Heredity.” Encyclopaedia Britannica. 11th ed. Vol. XII: 350– 354. Norton, Bernard. 1975a. “Biology and Philosophy: the Methodological Foundations of Biometry.” Journal of the History of Biology 8: 85– 93. ———. 1975b. “Metaphysics and Population Genetics: Karl Pearson and the Background to Fisher’s Multi– factorial Theory of Inheritance.” Annals of Science 32: 537– 553. Olby, Robert. 1966. Origins of Mendelism. New York: Schocken Books. ———. 1979. “Mendel no Mendelian?” History of Science 17: 53– 72. Pearson, Karl. 1900. The Grammar of Science. 2nd ed. London: Adam and Charles Black. Porter, Theodore M. 1986 The Rise of Statistical Thinking, 1820–1900. Princeton: Princeton University Press. ———. 2004. Karl Pearson: The Scientific Life in a Statistical Age. Princeton: Princeton University Press. Provine, William. 1971. Origins of Theoretical Population Genetics. Chicago: University of Chicago Press. Stamhuis, Ida. 2003. “The Reaction to Hugo de Vries’s Intracellular Pangenesis: the Discussion with August Weismann.” Journal of the History of Biology 36: 119– 152. Sturtevant, A. H. 1965. A History of Genetics. New York: Harper and Row. Tabery, James G. 2004. “The ‘Evolutionary Synthesis’ of George Udny Yule” Journal of the History of Biology 37: 73– 101. Vorzimmer, Peter. 1963. “Charles Darwin and Blending Inheritance.” Isis 54: 371– 390. Yule, G. Udny. 1902. “Mendel’s Laws and Their Probable Relations to Intra–Racial Heredity.” New Phytologist 1: 193– 207, 222– 238.

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Sources of Johannsen´s Genotype Theory Nils Roll–Hansen

The rejection of “soft inheritance” was a crucial step in the founding of classical genetics, according to Ernst Mayr. He acknowledges Wilhelm Johannsen’s central role in clarifying the distinction between “genotype” and “phenotype”, in coining the term “gene”, and in setting up his classical selection experiment on beans demonstrating a striking hardness of the genotype for quantitative characters. But Mayr nevertheless claims that a clear grasp of the distinction was not achieved till the middle of the 20th century, with the discovery that “the genotype consists of DNA. […] In the early years of genetics considerable confusion continued, from which even Johannsen was not exempt” (Mayr 1973, 1982: 782–783). This view is also reflected in the analyses of William Provine (1971), Frederick Churchill (1974) and Garland Allen (1979). However none of these authors have discussed Johannsen’s pre–1903 work, a main reason being that most of it was published in Danish. Wilhelm Johannsen started his education in the best primary and secondary schools of Copenhagen. But his father could only afford one son at the university and Wilhelm took vocational training as a pharmacist. The final part of this training was a one year study at the University of Copenhagen, where he became a favourite student of Eugenius Warming, professor of botany. 1880 Wilhelm passed his final exams as pharmacist, and in 1881, 24 years old, he was appointed assistant at the Carlsberg Laboratory. His job was to make investigations of barley in the chemistry section headed by the analytical chemist Johan Kjeldahl, famous for his method of analyzing organic nitrogen. Under this “unusually liberal boss” Johannsen had ample opportunity to pursue advanced biological studies including periods in Germany and Paris (Johannsen 1910). His other senior colleague was the botanist and yeast specialist Emil Chr. Hansen, known for his “pure lines” of yeast cultures. In 1887 Johannsen left Carlsberg in prospect of a lectureship in plant physiology at the Royal Veterinary and Agricultural College. In 1905 he was called as professor ordinarius in plant physiology to the University of Copenhagen where he spent the rest of his career.

Aristotle, Bacon and Claude Bernard Some characteristic traits of the young plant physiologist appear in an early survey article printed in a popular agricultural magazine (Johannsen 1883). With inspiration from Claude Bernard he praised the experimental method and aimed for the basic physiological principles of life common to plants and animals. Johannsen saw experimental natural science and natural history as united in a comprehensive theory of nature (“naturlære”) (p. 333). He demanded stringent method both in experiment and logic, and pointed to the perennial threat of Bacon’s idols to all good science (p. 360). Already, Wilhelm Johannsen was an independent thinker. An editorial preface (p. 331) warned against some of his claims: That Liebig’s law of minimum factors is useful in practical plant cultivation (p. 356), that heat is the movement of tiny particles (p. 347), and that the idea of

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a “vital force” should probably be banned from science (p. 353). Throughout his career Johannsen was highly active as a popular science writer with clear opinions on controversial topics. The Aristotelian bent of Johannsen is evident in his popular article on “the soul of plants” (“Om Planternes Sjæl”) of 1889. He claims that plants do have a sense of touch (“Berøringsfølsomhed”), exemplified by the mimosa, and thus a soul on the level of the senses (Johannsen 1889: 32). Rephrasing Charles Bonnet he said: “I do not claim to have proved that plants really have consciousness (“Bevidsthed”); but it has certainly not been proven that they do not.” (p. 181). He rebuked Aristotle mildly for his sharp division between the plant and animal kingdoms (p. 163), and quoted Claude Bernard in conclusion: “There is only one kind of life, only one physiology for all living beings” (p. 182).1 A study of the development of the endosperm in barley was Johannsen’s first substantial scientific publication (Johannsen 1884). The search for causes of the difference between “mealy” (“melet”) and “glassy” (“glasset”) barley grains – the first considered to be of superior quality, was one path leading him toward the general problems of variation and heredity. By 1895 Johannsen had developed a well rounded general approach to the study of heredity, with Hugo de Vries and Francis Galton as main sources of inspiration. His views were published in a small popular book, Heredity (“Arvelighet”), and in the third edition of Eugenius Warming’s textbook General Botany (“Den almindelige Botanik”) where Johannsen wrote the chapters on cytology and physiology.

Heredity in evolution and breeding It is Darwin’s eternal claim to fame that he led the idea of organic evolution to victorious breakthrough, wrote Johannsen, but variation and heredity should now be studied in their own right and not merely as subordinate topics to the evolution of species. “Evolution needs theory of heredity; but not vice versa”, he claimed (Johannsen 1896: 12). He recommended the study of pre–Darwinian authors, for instance, the successful practical breeder Louis Vilmorin (p.13). Johannsen presented the normal variation curve of Quetelet and Galton, which described how “the various properties of individuals belonging to a species or race vary around an average” expressing a “type”, and went on to the diverging variation curves of William Bateson and Hugo de Vries. A curve with two peaks is with great probability (“stor Sikkerhed”) due to a mixture of two types (p. 19). Johannsen stressed the fundamental role of causal processes at the individual level in his discussion of statistical methods in breeding. He warned against “German dogmas” like the law of correlation claiming that certain traits were linked and could not be separated. Statistics present average numbers but “do not necessarily say anything decisive about the individuals” (p. 48–50). Recent developments in cytology a main input to Johannsen’s early thinking on heredity. He referred to the work of August Weismann, de Vries and Oscar Hertwig. Weismann’s speculations on the “continuity of the germ–plasm” was essentially a development of Galton’s stirp theory. This idea expressed a perspective on heredity and evolution that was basic to Johannsens thinking from the 1890s on. With two simple drawings he illustrated the difference between the Galton– 1

44

I am translating from Johannsen’s Danish. He gives no reference.

Sources of Johannsen´s Genotype Theory

Weismann view on the one hand and the orthodox Darwinian and Lamarckian ideas of pangenesis and inheritance of acquired characters on the other (Johannsen 1896: 77, figure 4). For Johannsen the Galton–Weismann idea represented what Imre Lakatos would have called the hard metaphysical core of a research program that eventually produced an empirically well–founded theory of the genotype, or of “hard heredity” in Mayr’s terminology. Weismann’s detailed speculations on determinants and the role of reduction division were firmly rejected by Johannsen as excessively preformationist. Weismann’s misleading idea was to “have all possibilities and causes of the later development of the individual compressed into the egg, which accordingly would be at least as complicated in its composition as the fully grown individual” (Johannsen 1896: 73).2 Like Oscar Hertwig (1894) Johannsen held an epigenetic view. He held that the egg did not contain “germs of all possible later properties of the grown individual, but at most certain decisive dispositions (‘Anlæg’) with respect to the earliest steps of development”. The implication was that “differences in the sex cells could … be expressed through diverging reactions … to conditions of development introduced at an earlier or later stage.”3 The sound core of Weismannism was an old idea, “first formulated relatively clearly by Galton more than 20 years ago,” according to Johannsen (1896: 75).

Inheritance of acquired characters Johannsen nevertheless emphasized Weismann’s importance in stimulating the science of heredity. His criticism of claims about inheritance of acquired characters had generally been very skilful and appropriate, but had its limitations. For instance Weismann’s criticism of Brown– Sequard’s experiments of producing hereditary epilepsy in guinea pigs was not convincing (Johannsen 1896: 83). In Johannsen’s view these experiments of Brown–Sequard, Pasteur’s development of an anthrax vaccine, and Schübeler’s experiments on reversible hereditary change of barley subjected to arctic and temperate climates gave strong evidence that individually acquired characters could influence heredity directly (p. 91). For Johannsen a moral implication of such inheritance of acquired characters was that each individual human being had responsibility to future generations for its conduct of life, “a responsibility that would be obliterated by Weismann’s teaching!” 4 In particular, Johannsen was thinking of a damaging effect of alcohol on heredity (Johannsen 1896: 93–94). In General Botany (Den Almindelige Botanik), the text–book co–authored with Warming, Johannsen carefully summarized the most recent discoveries about the behaviour of the nucleus and the chromosomes in reduction division and fertilization. Strasburger, Weismann, and de Vries take these results to show that “the nucleus alone is the carrier of hereditary properties.” 2

3

The whole sentence in Danish: “Disse oppfattelsers fælles ledende, urigtige Grundtanke er den, at Weismann vil have alle Muligheder for og Aarsager til Individets senere Udvikling pressede sammen i Ægget, som saaledes maatte blive mindst lige saa indviklet sammensat somselve det ferdige Individ.” The whole sentence in Danish: “Antage vi derfor ikke i Ægget særligt existerende Smaakim til alle mulige senere Egenskaber hos det voxne Individ, men i det højeste visse afgjørende Anlæg med hensyn til de allerførste Trin af Udviklingen, saa vil ikke desto mindre Forskjelligheder mellem Kønscellerne dog kunne yttre sig ved forskjelligartede reaktioner – I dette Ords videste forstand – overfor Udviklingsfaktorene, være sig paa et tidligere eller senere Trin I Udviklingen.”

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Nils Roll-Hansen

“The idea is seductive and not improbable,” commented Johannsen, but it has not been proved (Warming and Johannsen, 1895: 153–154). There is a lively current debate about the importance of inheritance of acquired characters in the transformation of species, wrote Johannsen. “Probably we come closest to the truth by assuming that both natural selection and direct adaptation is causing transformation” (p. 507). Fifteen years later Johannsen and Warming had split radically on this question. Warming had become a convinced neo–Lamarckian and Johannsen a Mendelian selectionist.

Pedigree Breeding In the mid–1890s Johannsen engaged in breeding high quality barley for brewing. The results confirmed his suspicion that certain Darwinian theories of evolution were an obstacle to plant breeding. For brewing large grain size was generally a positive quality. High nitrogen content was advantageous for some purposes and not for others. The popular idea of correlation, of a tight link between properties, was mistaken in this case, argued Johannsen. Such claims were often supported by population averages, “average numbers of material arbitrarily collected from all over the world,” as he wrote. If individual selection was combined with hybridization the possibilities of combining wanted properties in many different ways were much larger than indicated by the “law of correlation”. “So let us face our tasks without hesitation and beware of German dogmas!” (Johannsen 1898a: 68–78).5 Johannsen pointed to the “theory of heredity and plant breeding before Darwin” (Johannsen 1898b). In particular Louis Vilmorin was a good guide to methods of pedigree breeding and effective use of hybridization as means to increase hereditary variation (Johannsen 1898b: pp. 345–346, 456; 1899: 556). Johannsen cited Hertwig (1898) on the lack of experiments that unequivocally demonstrated the inheritance of acquired characters. However, he also agreed with Hertwig that “a certain degree of heredity for characters developed by external factors” had to be assumed. This phenomenon might be important in the natural evolution of species, but for breeding it was irrelevant: “Nowhere is the difference between theory of evolution and breeding work clearer than precisely in this respect”(Johannsen 1898b: 321). 6 In 1899 Johannsen presented reduction division and fertilization as the basis for understanding heredity. For instance, in distinguishing variation due to internal and external causes he saw “internal” variation as proceeding “quite independently of external conditions” and due to “the ‘imperfect accuracy’ of cell division” (Johannsen 1899a: 452–455). 7 Vilmorin’s old 4

5 6 7

46

The whole paragraph in Danish: “Vi komme altsaa til den Opfattelse, at Individets Livsvilkaar og, hva der naturligvis altid vil staa i Sammenhæng dermed, Individets hele Livsførelse maa kunne faae Indflydelse – om end aldri saa lidt – paa afkommets Beskaffenhed, ogsaa naar vi se helt bort fra Smitteoverførelse. Dette er paa ingen Maade saaledes at forstaa, at en eller anden, være sig god eller slet Egenskab, som man har “erhvervet” sig, just behøver at vise sig hos Afkommet i samme Skikkelse eller i samme Retning. Men paa forskjellige direkte og indirekte Maader kan en Indflydelse, som den nævnte, tænkes virksom. Individet – det enkelte Menneske holde vi os til i dette Øjeblik – faa da i alt Fald et vist Ansvar overfor den kommende Slægt med hensyn til sin Livsførelse, et Ansvar, som efter Weismanns Lære udviskes!” “ – lad os gaa lige løs paa vore Opgaver og vogte os for tyske Dogmer!” (Johannsen 1898a: 78). “Men ingensteds træder Forskjellen mellem Udviklings–Læren og Forædlings–Arbeidet klarer frem, end netop paa det her berørte Omraade.” “Vi antager altsaa, at der kan skje Variation ganske uafhængig af ydre Kaar og nærmest betinget af Celle– Delingens ’ufuldkomne Nøyaktighet’ – om man vil tillade mig Brugen af dette Udtryk.”

Sources of Johannsen´s Genotype Theory

principle that the “hereditary power” of seemingly similar individuals or races could be very different, and that therefore the breeding value of an individual should judged by the qualities of its offspring rather than by the quality of the individual itself, thus had a cytological basis. The importance of this principle was now being rediscovered (Johannsen 1899a: 542), in Johannsen’s own experiments, in the breeding successes of Hjalmar Nilsson at Svalöf, as well as in Hugo de Vries’ experiments with hereditary malformations in plants (p. 547). Johannsen suggested that one reason why Vilmorin’s correct idea had been neglected was his mistaken ideas about fertilization. Like his contemporaries Vilmorin thought that a flower was usually fertilized by its own pollen. But we now know that fertilization by foreign pollen is the rule. Because of this a reliable pedigree is often impossible to establish. Special methods like artificial pollination would have to be used for Vilmorin’s principle to be generally effective. This was the case for important cross–fertilizing plants like rye and sugar beet. However, for self–fertilizers like peas or barley there was no doubt about the “father” and the principle was directly applicable (Johannsen 1899a: 543; Johannsen 1899c: 170–171). Johannsen had worked mostly with barley, and for his classical selection experiment he picked beans, another self–fertilizer.

The analysis of “variability” In the fourth edition of General Botany the treatment of evolution and heredity was reorganized and substantially expanded. The heading of the twelfth and last section of the book was “Theory of Descent” and its one and only chapter, No. 58, was called “Natural Kinship; Variability, Heredity and Phylogenesis”.8 Starting with a discussion of biological systematics Johannsen noted that traditional Linnean species had turned out to contain numerous varieties or subspecies. Some of these were sufficiently stable to be considered independent systematic “types”. In Johannsen’s view these “small” species deserved, more than the Linnean “large” species, to be considered the basic systematic units (Warming and Johannsen 1900: 666). Five kinds of variability (“Variabilitet”) were listed (p. 667): 1) The polymorphism of traditional Linnean species. 2) Differentiation of forms characteristic of the progeny of hybrids (this was further discussed in a subchapter on laws of Mendelian segregation, pp. 679–683). 3) “(I)ndividual or fluctuating variability” in the strict sense. 9 4) Differences developed under “divergent external conditions”.10 (In contrast to de Vries Johannsen wanted to keep a distinction between individual variation and modifications due to the growth environment (“Voksestedsmodifikation”), Johannsen 1902: 558.) 5) So–called mutations by which individuals representing new stable forms, species or subspecies, suddenly appeared.

8 9 10

“Naturlig Slægtskab; Variabilitet, Arveligehd og Fylogenese”. “Den saakaldte individuelle eller fluktuerende Variabilitet, Variabiliteten i snævrere Forstand.” “De ofte iøjenfallende Forskelligheder, som, endog efter Utsæd af samme Frøprøve, vise sig mellom Individer, der udvikle sig under stærkere afvigende ydre Kaar.”

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The first and fifth kinds, species polymorphism and mutation, were directly related to basic systematic groups, and thus by definition hereditary. The second kind, variation following upon hybridisation, was recognized as capable of producing new stable forms by recombining properties of the parent forms. The fourth kind was not hereditary, or at least only partly. But the third kind, “individual variability”, was contentious with respect to its contribution to the formation of new species. Johannsen gave special attention to this third kind, individual variation, defining it as the regularly occurring variation with respect to quantitative measurable properties among closely related individuals belonging to a “subspecies, variety, or cultural form” (Warming and Johannsen 1900: 668).11

The stability of biological types In the mutation theory of Hugo de Vries “small” species did not originate by gradual continuous transitions but through sudden larger or smaller changes, “mutations”. On this theory individual variations “had absolutely no significance for the development of new forms”, according to Johannsen. And he thought it likely that in the Darwinian tradition individual variation had been overestimated (“overvurdert”) with respect to the formation of new types (p. 686–687). By simple investigation of an individual different from the average of the population it was not possible to decide if it represented a different biological type. In particular this was so for quantitative properties. The progeny of the individual had to be studied, wrote Johannsen. If the progeny, on average, has the same character as the parent it represents a mutation, i.e., a different basic systematic class. But if the progeny regresses toward the mean of the population we have to do with an “individual variation” (Warming and Johannsen 1900).12 Thus the purpose of Johannsen’s classical selection experiment on beans 13 was to test the law of regression as formulated by Galton and his followers Karl Pearson and Frank Raphael Weldon, using Vilmorin’s method of analysing the offspring. Johannsen bought sixteen pounds of Princess beans, “well developed” and of “uniform” quality, grown in Fyn in 1900 (Johannsen 1903: 15). In General Botany we find a table showing correlation between width and length of 12000 beans (Warming and Johannsen 1900: 669). This analysis was presumably part of preparations for the first planting made in the spring of 1901.14

11

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13 14

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“Herved forstaas det Forhold at Individerne af en given, snævrere Slægtskabskreds (Underart, Afart, Kultrurform), ’variere’ regelmæssigt med hnesyn til de Egenskaber, som overhovedet kunne udtrykkes i Tal, d.v.s. kunne grupperes omkring typiske Værdier paa lovbestemt maade.” “Først Afkommeets Forhold kan afgøre Sagen: faar Afkommet, gjennomsnitlig set, samme Præg som de paagæedende Individ, da var det Mutation: men slaar Afkommet, gjennomsnitligset, tilbage mot den oprindelige Type, da var det det individuell Variation hos det nævnte Individ. Man forstaar heraf den store principielle Btydning, som en gennemført særskilt Undersøgelse af det enkelte Individs Afkom har for Arvelighedslæren og for Planteforædlingen.” Published in 1903 as “On heredity in Populations and Pure Lines” (“Ueber Erblichkeit in Populationen und in reine Linien”). The preface of Warming and Johannsen 1900 is dated June 1901.

Sources of Johannsen´s Genotype Theory

The bean selection experiment and the Biometricians’ reaction The sharp distinction that de Vries had made between the concepts of “statistical variation” and “mutation” was undoubtedly a substantial step forward, wrote Johannsen in a short “preliminary review” of The Mutation Theory (Johannsen 1901: 8).15 The central question of the bean experiment was apparently sharpened also by Johannsen’s reading of Weldon’s review in Biometrika. Till now, wrote de Vries in the preface to The Mutation Theory, the origin of species has been studied by comparative methods. It has been the general opinion that this phenomenon is inaccessible to direct observation and at least to experiment (de Vries 1901: p. III). 16 It was the ambition of de Vries to change this. But Weldon’s review was very critical: Firstly de Vries had misunderstood Galton’s law of regression, and secondly his results fitted perfectly with Pearson’s interpretation of the law. According to Weldon the successive increase and decrease of average number of seed rows on corn heads through selection formed “a fairly conclusive proof” that the theory of mutations was mistaken (Weldon 1902: 369). Weldon put the burden of proof squarely on his opponents: “A clear proof that Professor Pearson’s view of the facts of regression is wrong … is absolutely essential …” 17 With a high degree of self confidence Weldon dismissed the attempts of de Vries and Bateson “to distinguish between ‘variations’ and ‘mutations’, or between ‘normal’ and ‘differential’ variations”. Their attempts rested on a view “which a little knowledge of the statistical theory of regression will show to be wholly imaginary” (Weldon 1902: 374). This was the challenge that Johannsen picked up in his bean experiment. In a 1902 paper on “The plurality of forms of organisms” (“Organismernes Formrigdom”) Johannsen further developed his views on the nature of “variability” with special focus on “individual variability”. The general question was whether hereditary change in the evolution of species is caused by the direct influence of living conditions, independently of selection, or through mutations (Johannsen 1902: 548). In a note at the end Johannsen remarked that recently the British “biometric” school headed by Pearson had mercilessly revealed the deficiencies in the arguments of Bateson and de Vries. Johannsen nevertheless had no doubt that mutations do occur, though the border between mutations and individual variation might be less sharp than they thought. “Presumably research of the near future will bring some clarification”, he added optimistically (p. 565).18 15

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18

“Denne skarpe Adskillelse mellem Begreberne statistisk Variation og Mutation er utvivlsomt en stor Vinding, og med Mutationslærens Sprog kan man godt som de Vries, sige om en given Underart, at den er ‘völlig konstant und höchst variabel’: Typen er fast, men Svingningen om Typen ere sterke.” The first paragraph runs as follows: “Die Lehre von der Erstehung der Arten ist bis jetzt eine vergleichende Wissenschaft gewesen. Man glaubt allgemein, dass dieser wichtige Vorgang sich der Beobachtung und mindestens der experimentellen Behandlung entziehe.” The whole sentence: “A clear proof that professor Pearson’s view of the facts of regression is wrong although it is in accord both with the theory of chance and with the results of the numerous statistical studies of inheritance which he and his pupils have made during the past seven years, is absolutely essential, if the view held by Professor de Vries is to be maintained.” “Jeg nærer dog aldeles ingen Tvivl om, at Mutationer forekommer, særlig har de Vries Kulturer overbevist mig. Men muligens er Grænsen mellom utvivlsom Mutation og blot og bar individuell Variation næppe altd saa skarp som de Vries og Bateson er tilbøjelige til at antage. Ventelig vil den nærmeste fremtids Forskning her bringe mere Klarhed tilveje.”

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Results of the bean selection experiment Analysis of the harvest of the second planting, in 1902, brought much clearer results than Johannsen had expected. It turned out that within each of the types that he had isolated as pure lines19 there was no hereditary effect of selection. In other words, there appeared to be complete regression to the type of the line. If Johannsen’s interpretation was tenable this was just the kind of disproof of the biometric law of regression that Weldon had asked for. The results appeared so clear cut and convincing, as well as important for the study of heredity, that Johannsen found it right to publish after only two growing seasons. The limited task of this publication, wrote Johannsen in the concluding discussion, is only “to throw light on the Galtonian regression between parents and children” (Johannsen 1903: 64). He had used a material of similar nature to that of Galton, namely beans instead of sweet peas. Like Galton he measured seed in successive generations. But in contrast to Galton he also introduced the pedigree principle. He traced the offspring of each individual first generation parent bean and thus made it possible to test for the presence of distinct types within the population. Galton had taken the whole population to represent a biological unity while Johannsen found that it consisted of many distinct and stable biological types. For the population as a whole Galton’s law of regression was confirmed, but within each pure line it did not hold. The hereditary type of each line was stable, claimed Johannsen, completely unaffected by selection (Johannsen 1903: 57). 20 Thus he had shown that de Vries was right in claiming constancy for the elementary biological types. The Biometricians’ law of regression had been refuted in so far as it was taken to imply a continuous change of heredity. Within pure lines of self–fertilizing plants like peas or beans “individual variations” were not hereditary. Not surprisingly this conclusion did not please Pearson and Weldon. Their reviews in Nature and Biometrika brushed off Johannsen’s results in much the same way as Weldon had rejected de Vries experiments the year before: Johannsen had misunderstood the law of regression and in fact his results were a confirmation of it (Pearson 1903; Weldon and Pearson 1903). Johannsen’s controversy with the Biometricians stimulated his development of the concepts of “genotype”, “phenotype” and “gene” which were first introduced as the basis of a comprehensive theory in his 1909 German textbook (Johannsen 1909: 130). But this is another story (Roll–Hansen 1989). The purpose of this paper has only been to trace the history and background of Johannsen’s concept of stable elementary biological types, leading up to his classical selection experiment of 1901–1902.

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A “pure line” as defined by Johannsen is simply the offspring in successive generations from one single parent. With a regularly self–pollinating organism like beans this was easy to achieve and effects of hybridisation in earlier generations would also be minimal. “Indem ich aber nicht dabei stehen blieb, die Populationen als Einheiten zu betrachten, sondern mein Material in seinen ’reinen Linien’ auflösen konnte, hat es sich in allen Fällen gezeigt, dass innerhalb der reinen Linien der Rückschlag sozusagen vollkommen gewesen ist: die Selektion innehalb der reinen Linien hat keine Typenverschiebung hervorgerufen”.

Sources of Johannsen´s Genotype Theory

Cloncluding remarks The role of pedigree selection in Johannsen’s early work on heredity is striking. Extensive discussions of the implications of recent cytological discoveries for theories of heredity are also clear evidence of his interest in causal factors and processes at the level of the individual organism. His recurring illustration of the Galton–Weismann stirp versus the Darwin–Lamarck pangenesis model for transmission of heredity fits this interpretation of his theory of heredity. Thus it appears a mistake when Frederick Churchill interprets Johannsen’s concepts of phenotype and genotype as primarily statistical when introduced in 1909. According to Churchill “Johannsen was a chemist and statistician first”, and his thinking was basically statistical and “vertical”. It started from the description of population averages, “phenotype”, and positing “genotype” as a theoretical entity with no commitment in cytology. Only later did Johannsen, according to Churchill (1974: 28–30), come around to the cytologically based “horizontal analysis” that was characteristic of mature classical genetics, and as it was formulated for instance by the Drosophila school of T.H. Morgan around 1915. I believe the preceding account shows that “horizontal analysis” examining “directly the genetic composition sundered either conceptually or physically from the Erscheinungsphenomena” (Churchill 1974: 30) was an essential part of Johannsen’s research project on biological heredity from the beginning. This approach was well founded by 1895 and lead up to his 1903 paper and the genotype–phenotype distinction of 1909. Garland Allen, referring to Churchill, also claims that “Johannsen’s eye was focused on the entirety of a population and its range of variation. He was not concerned about distinguishing genotypic from phenotypic variation within the individual organisms, but only within large populations” (Allen 1979: 198). This is a description that fits the biometric view well but seems quite opposed to that of Johannsen. Ernst Mayr in his 1982 book, The Growth of Biological Thought, still takes Churchill’s paper to be authoritative on Johannsen’s view of the genotype. When William Provine (1971: 96–97) found the Pearson–Weldon criticism of Johannsen’s selection experiment to be “powerful”, and held that its neglect by the biological community in general was not justified, the reason may be a similar misinterpretation of Johannsen. To sum up: It seems that the standard interpretation of Johannsen, as promoted for instance by Mayr, Churchill, Allen and Provine, fails to grasp the difference between him and the biometricians.

Nils Roll-Hansen, Institutt for filosofi, ide- og kunsthistorie og klassiske språk, Universitetet i Oslo, [email protected]

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References: Allen, G. 1978. “Naturalists and Experimentalists: The Genotype and the Phenotype”, Studies in History of Biology, 3: 179–210. Bernard, C. 1885. Lecons sur les phénomenes de la vie communs aux animaux et vegetaux. Paris. Churchill, F.B. 1974. “William Johannsen and the genotype concept”, Journal of the History of Biology 7: 5– 30. Dunn, L.C. 1973. “Johannsen, Wilhelm Ludvig”, Dictionary of Scientific Biography 7: 113–115 (New York: Charles Scribner’s Sons). Hertwig, O. 1892 . Zeit– und Streitfragen der Biologie. I. Praeformation oder Epigenese? ———. 1898. Die Zelle und die Gewebe II, Jena. Johannsen, W. 1883. “Plantefysiologiske Meddelelser”, Tidsskrift for Landøkonomi, 2 (1883): 332–365, 792– 795. ———. 1884. “Om Frøhviden og dens Udvikling hos Byg” (About the endosperm and its development in barley), Meddelelser fra Carlsberglaboratoriet (Reports from the Carlsberg Laboratory) 2 (3): 103–133. The results were also published in German: “Einleitende anatomische Studien über die Frage eines milden (mehligen) Gersten–Kornes”, Allg. Zeitschr. f. Bierbr. U. Malzfabr. 12 (1884): 625–750. ———. 1898a. “Nogle Studier over Variation og Forædling med særlig hensyn på Goldthorpe–Byg”, Tidsskrift for Landbrugets Planteavl (senere: Tidsskrift for Planteavl) 5 (1899): 63–90. ———. 1898b. “Om Arvelighedslære og Planteforædling før Darwin”, Ugeskrift for Landmænd 1898: 289– 290, 318–322, 343–346, 428–430, 455–456. ———. 1899a. “Nogle Arvelighedsforhold og deres Betydning for Plantedyrkningen”, Dansk Tidsskrift (Gads Magasin), 1899: 449–464, 542–557. ———. 1899b. “Fortsatte Studier over Kornsortene. I Om Variabiliteten med Hensyn til Forholdet mellem Kornvægt og Kvelstof–Procent hos Byg”, Meddeleser fra Carlsberglaboratoriet 4 (4): 228–313. Also published in German: “Über die Abänderungen der Gerste mit besonderer Rücksicht auf das Verhältnis des Gewichtes der Körner zu ihremGehalt an Stickstoffhaltigen Substanzen”, Zeischrift f. d. ges. Brauw. 23: 487 … 622. ———. 1899c. “Nogle Bemerkninger om Arvelighed og Variabilitet”, Gartner.–Tid. 15: 166–172, 180182. ———. 1902. “Om Organismernes Formrigdom”, Nordisk Tidskrift för Vetenskap (Letterstedske Tidskrift), 1902, pp. 545–565. ———. 1903. Erblichkeit in Populationen und in reinen Linien, Jena: Gustav Fischer. ———. 1909. Elemente der Exakten Erblichkeitslehre. Jena: Gustav Fischer. ———. 1910. Self–biographical note in Festskrift udgivet af Kjøbehavns Universitet i Anledning af Universitetets Aarsfest November 1910, pp. 70–73. Mayr, E. 1973. “The Recent Historiography of Genetics”, Journal of the History of Biology 6: 125–154. ———. 1982. The Growth of Biological Thought, Harvard University Press. Pearson, K. (Anonymously) 1903. “Prof. Johannsen on heredity”, Nature 69 (17 December 1903), 149–150. Provine, W. 1971. The Origins of Theoretical Population Genetics, Chicago: The University of Chicago Press. Roll–Hansen, N. 1989. “The Crucial Experiment of Wilhelm Johannsen”, Biology and Philosophy 3: 303– 329. Vries, H. de. 1889. Intracellulare Pangenesis, Jena: Gustav Fischer. ———. 1901. Die Mutationstheorie. Versuche und Beobachtungen über die Entstehung von Arten im Pflanzenreich, Leipzig: von Veit & Comp. Warming, E. and W. Johannsen. 1895. Den Almindelige Botanikk, 2nd edition, Copenhagen. ———. 1900. Den Almindelige Botanikk, 3rd edition, Copenhagen. Weldon, W.F.R. 1902. “Professor de Vries on the Origin of Species”, Biometrika 1, 365–374. Weldon, W.F.R. and K. Pearson. 1903. “Inheritance in Phaseolus Vulgaris”, Biometrika 2, 499–503.

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Inheritance of Acquired Characters: Heredity and Evolution in Late Nineteenth-Century Germany Wolfgang Lefèvre

INTRODUCTION The assumption of inheritance of acquired characters was a widely shared assumption up to the first half of the twentieth century. Today, inheritance of acquired characters is generally dismissed by biologists. More precisely, it is regarded as being definitely ruled out by the central dogma of molecular genetics as established by Francis Crick in 1958 (Crick 1958): DNA makes RNA makes protein, and never the other way around. Although, probably, little risk would be taken by predicting further revivals of inheritance of acquired characters, in this paper, I will regard the history of inheritance of acquired characters as a closed one. Conway Zirkle traced back this history to classical antiquity, and one can be sure that the assumption of inheritance of acquired characters is far elder than any written record. However, Zirkle’s study documents not only how broadly this assumption was held in all historical periods since antiquity. It also testifies that none of these periods lacked critical voices that doubted or even plainly dismissed inheritance of acquired characters. By this, Zirkle’s study shows thirdly, and probably involuntarily, that inheritance of acquired characters was an indifferent assumption, that is, an assumption that could be held and challenged without raising the need, neither on part of its adherents nor of its critics, to subject it to serious tests. It was no simple task to decide definitely whether or not acquired characters can be transmitted. This became clear when the indifferent status of inheritance of acquired characters ended in the last decades of the nineteenth century and serious efforts were made to prove its groundlessness. “There are three ways,” wrote Ernst Mayr, “to refute an inheritance of acquired characters. The first is to show that the mechanisms by which it is supposed to operate are impossible. This was primarily Weismann’s approach. There is nothing in the structure and division of cells that would make an inheritance of acquired characters possible. […] A second way to refute an inheritance of acquired characters is by experiment. […] Beginning with Hoffmann and Weismann, such experiments were conducted up to the 1930s and 40s and the results were uniformly negative. The third way of refuting the theory of the inheritance of acquired characters is to show that the phenomena that are claimed to require the postulate of an inheritance of acquired characters can be explained equally well or better on the basis of the Darwinian theory. Much of the evolutionary literature of the 1920s, 30s, and 40s was devoted to this third approach” (Mayr 1982, 699ff.). With this summary, Mayr conjures up the theoretical developments before the age of molecular genetics that undermined inheritance of acquired characters step by step. The probably most important, if not even revolutionary, of these developments consisted in the devaluation of the role of the parental organism in the biological transmission process. This role was reduced to that of a mere conveyer and re-combiner of an inheritance that predates it. From different quarters, Francis Galton’s “ancestral line” and August Weismann’s “continuity of the germ

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plasm” (Keimplasma) laid the ground for an understanding of the biological transmission process that left little space, if any at all, for inheritance of acquired characters. That this would be the final outcome, was of course in no way clear in the 1880s when these developments started. At this point of time, it was as open a question as in the past whether or not inheritance of acquired characters can be asserted or must be denied. Yet, what had apparently changed then was the indifferent state of this issue. Now there were biologists like Weismann who did no longer regard inheritance of acquired characters a matter of individual convictions but wanted the question to be answered definitely. One may ask therefore what did transform the assumption of inheritance of acquired characters which had be held so long despite doubts and even complete rejections, into one that must be proved or dismissed? What did lead to an end of the indifferent state of this issue? Or, to put it differently, why did the assumption of inheritance of acquired characters become an urgent point on the agenda of biology in the 1880s? Ernst Mayr, pondering Weismann’s motives for his sudden turn against inheritance of acquired characters, indicated an answer that seems worth to be elaborated. He wrote that it is not clear “whether Weismann had first become convinced of the invalidity of the theory of an inheritance of acquired characters and then adopted the germ-track theory or vice versa. The fact is that he already cites in his 1883 paper [Weismann 1883] so many lines of argument against soft inheritance that one can well imagine that this general conviction preceded the proposal of a specific mechanism. This interpretation is strengthened by the fact that Weismann was a strict selectionist already in the 1870s and presumably had simply no need for an additional mechanism” (Mayr 1982, 701). With the last sentence, Mayr draws our attention to the fact that, since the 1870s, the issue of heredity played a significant role in the debate about evolution. It has, indeed, the appearance that inheritance of acquired characters got then a new state because of its bearing for the understanding of Darwin’s theory of evolution. The goal of my paper is to shed some light on exactly the connection between inheritance of acquired characters and evolution as established by the historical actors in the last quarter of the nineteenth century. However, rather than offering a narrative of the debates by which this connection was brought about by the historical actors, the paper’s focus will be on an analysis of the strategic significance inheritance of acquired characters had within the contemporary framework of competing theories of evolution. In a way, what I present could be regarded a kind of a “rational reconstruction” of this significance, a reconstruction, however, that does not rest on an universal logic of scientific reasoning but on the internal “logic” of the historical theories involved. My paper is ordered as follows: In a very short first part, Darwin’s own relation to inheritance of acquired characters is addressed. In part II, the bearing is discussed inheritance of acquired characters had for the understanding of evolution. In part III, this discussion is exemplified through a sketch of Ernst Haeckel’s Neo-Lamarckian theory of evolution. In the fourth part, finally, the significance of inheritance of acquired characters for conceptions of evolution is shown from a different point of view, namely by citing two contemporary alternatives to Haeckel’s theory of evolution that dismissed inheritance of acquired characters.

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Inheritance of Acquired Characters

I. DARWIN AND INHERITANCE OF ACQUIRED CHARACTERS Today, the assumption of inheritance of acquired characters is equated with a Neo-Lamarckian approach to evolution. However, this assumption was neither first conceived nor given a specific meaning by Jean Baptist Lamarck or one of the known Lamarckians. Rather, as already said, it was a widely shared, though also periodically challenged, assumption from times immemorial up to the end of the nineteenth century and beyond. Against this background, it is in no way remarkable that Charles Darwin himself assumed an inheritance of acquired characters. Darwin devised an elaborate theory of heredity, his renowned or notorious “Pangenesis” theory, and published it in the second volume of The Variation of Animals and Plants under Domestication in 1868. Although all of his assumptions on heredity were dismissed by the subsequent developments of biologists’ understanding of the transmission process, and although not at all original as regards its central physiological hypothesis, Darwin’s theory must be regarded a landmark in the history of heredity theories. For it constituted the very first attempt at a really comprehensive elucidation of the transmission process, based, as is typical of Darwin, on an abundance of empirical material. It is still a first source if one wants to know what nineteenthcentury biologists, breeders, physicians etc. knew or assumed about heredity. At the centre of his theory is the supposition and explanation that and how every “unit” of parental organisms contributes to the formation of the sexual products of the two sexes that constitute the germ when united and contain all that is bequeathed to the offspring. Assuming in this way a direct dependency of the hereditary material on all parts of the parental organisms, this theory entails inheritance of acquired characters quite naturally. If Darwin, on the base of this theory, had wanted to exclude inheritance of acquired characters, he would have been forced to invent a suitable complementary hypothesis. But with inheritance of acquired characters in its indifferent state characterized above, Darwin felt no need to dismiss this assumption. On the contrary, his theory of evolution could even take advantage of inheritance of acquired characters and did so in fact occasionally. There is a long tradition of criticizing Darwin for being not enough a Darwinian but too much a Lamarckian evolutionist. I will not go into this judgment that obviously lacks a historical spirit. 1 Instead, I want to state the remarkable fact that Darwin, despite his assumption of inheritance of acquired characters, based his theory of evolution not on Lamarckian adaptations but on random variations. It was probably his familiarity with the experiences of professional breeders what prevented him from a Lamarckian approach. The fact that Darwin didn’t built upon Lamarckian adaptations needs clarification, and the following section will try to provide it. II. INHERITANCE OF ACQUIRED CHARACTERS AND EVOLUTION Looking back to the first decades after the appearance of Darwin’s On the Origins of Species in 1859, one gets the impression that it took a considerably long span of time before biologists came to terms about what constitutes the very core of Darwin’s theory of evolution. This core became conceivable, or one may even say: was shaped, in a clarification process in which alternative understandings of evolution emerged. Lamarckism or Neo-Lamarckism 2 was one of those 1

For a brief discussion of Darwin’s theory that places it historically, see (Bowler 1989) 54ff.

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alternatives, which contributed a lot to a more adequate understanding of what distinguishes Darwin’s theory from other conceptions of evolution. Ironically, Neo-Lamarckism became first such a clarifying alternative when Weismann attacked inheritance of acquired characters, one of its central elements. It was a shared presupposition of both, Darwinism3 and Neo-Lamarckism, that the historical evolution of plant and animal forms was essentially a result of adaptations to changes in the conditions of life in the course of the historical development of the Earth’s surface. That is to say, both, Darwinism and Neo-Lamarckism, regarded evolution as a process of such adaptations. And it was this shared presupposition by which Darwinism and Neo-Lamarckism distinguished themselves from conceptions of evolution that assumed an inner-organismic formative drive as motor of evolution. Darwinism and Neo-Lamarckism also shared the presupposition that changed conditions of life do not cause directly adaptive modifications of organisms. However, as is well known, Neo-Lamarckians conceived of changed conditions of life as an indirect cause of adaptive modifications. They assumed that organisms react on such changes by changes of their behaviour; that such behavioural changes entail new uses of their organs; that new use, or for that matter disuse, of organs leads to physical modifications of organs and structure; and, finally, that such modifications are transmitted to the offspring and engender, in the course of the succession of generations, modified organisms that are better adapted to the new conditions of life. In our collective memory, the elongated neck of giraffes stands for this Lamarckian explanation, which I will call adaptation by use and disuse hereafter. As is obvious, this explanation of adaptation by use and disuse stands and falls with the assumption of inheritance of acquired characters. This assumption, thus, was rightly equated with a Neo-Lamarckian conception of evolution because the latter cannot be thought without the former. For Darwinians, too, conditions of life have a share in the development of adapted organisms. It is exactly these conditions that assign adaptive value to organismic modifications and, thus, prove to be ultimately responsible for the selection of variations in the “struggle for existence.” However, Darwinism does not presuppose that variations of an adaptive value are directly or indirectly induced by new conditions of life. Rather this conception of evolution assumes that “natural selection” acts upon variations that are essentially random. It presupposes no other causes for variations of an adaptive value than for variations of no such value. In other words, Darwinism does not presuppose any correlation between the (then) unknown causes of variation and the demand for adaptation to changed conditions of life. Variations of an adaptive value are random in that their coming into being has nothing to do with this value. 4

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The label “Neo-Lamarckism” was coined by contemporaries — see, for instance, (Wagner 1908) 121ff. For the problems that made Lamarckism periodically, and not only before 1900, attractive, see (Beurton 2001). Already his contemporaries labelled Charles Darwin’s theory of evolution “Darwinism.” — Notwithstanding the uncountable modifications induced in this theory by biologists during the past 145 years, its core – speciation on the base of minimal random variations and natural selection — also constitutes the core of modern Darwinism. Modern Darwinism is to Darwin’s theory as classical mechanics is to Newton’s theory: Both cannot be found in the writings of their respective patron saint and owe to him nevertheless their decisive conceptual features. For the development of Darwin’s understanding of variation, see chapters 3 and 4 of (Vorzimmer 1972). For ambiguities of Darwin’s understanding of variation, see chapter 3 of (Gayon 1998).

Inheritance of Acquired Characters

Against this background, it becomes understandable why radical challenges of inheritance of acquired characters like Weismann’s didn’t strike Darwinism although Darwin, like most of the contemporary biologists, assumed this inheritance. As became clear in the course of the debate, Darwinism did not stand and fall with inheritance of acquired characters because its explanation of evolution built upon random variations and not on adaptations by use and disuse. In other words, it was Weismann’s attempt at a definite dismissal of inheritance of acquired characters that clarified that Darwin’s Pangenesis theory was not an integral part of his theory of evolution. As regards the issue of heredity, this theory presupposed only something well-confirmed by all breeding experiences, namely that, as a matter of fact, hereditable variations exist. This simple fact did neither imply nor suggest nor exclude any specific theory of heredity. It had been compatible with the assumption of inheritance of acquired characters, as Darwin’s Pangenesis shows, and proved to be in no way at odds with the new understanding of heredity that began to surface with Galton’s statistical theory of heredity and Weismann’s (and others’) cytological distinction between germ and soma plasm. No additional efforts were needed to reconcile it with the understanding of heredity as a process in which a hereditary substance is transmitted and recombined by the parental organisms that remains unchanged by modifications these organisms may undergo. While the development and elaboration of this new understanding of heredity was to undermine inheritance of acquired characters and, thus, the raison d’etre of Neo-Lamarckism, it constituted no challenge to a Darwinian theory of evolution that presupposed random variations. The differences between Darwinism and Neo-Lamarckism with respect to the evolutionary significance of inheritance of acquired characters discussed here are clear for us in hindsight, that is, after nearly 150 years of debates about evolution. But they were far from being clear for biologists in the last decades of the nineteenth century. As already said, the clarification of what is essential for Darwinism and what not took shape only in fierce arguments about evolution which were characteristic of the first 60 years after the appearance of The Origin of Species. Particularly in Germany in the 1870s, when evolution was already accepted by a lot of biologists, and probably by the majority of the younger ones, the discussed differences between a Darwinian and a NeoLamarckian understanding of evolution were almost not recognizable. For, here, this understanding was dominated by a special theory, that of Ernst Haeckel. III. INHERITANCE OF ACQUIRED CHARACTERS IN HAECKEL’S THEORY OF EVOLUTION Haeckel presented the understanding of evolution he had arrived at under the impression of Darwin’s Origins already in 1866, namely in his Generelle Morphologie. This was a book for professional biologists, particularly for comparative anatomists. Two years later, under the title Natürliche Schöpfungsgeschichte, he published essentially the same theory of evolution in a fashion apt for addressing a broader audience. This book shaped the initial view of Darwinism in Germany, not only that of interested non-experts but that of experts as well. Haeckel’s understanding of evolution was embedded in a fundamental theory of the formation of organismic structure. According to this theory, organismic structure results from and is determined by the mutual action of two general “physiological functions,” that is, the mutual action of inheritance (Vererbung) und adaptation (Anpassung) (Haeckel 1866, II 167f.).

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On this base, Haeckel distinguished two kinds of a formative drive (Bildungstrieb) that bring about organismic structure: an inner formative drive (innerer Bildungstrieb) that determines an organism’s development from within according to inherited dispositions; and an exterior formative drive (äußerer Bildungstrieb) that modifies its development in reaction to the outer conditions of life (Ibid. 168). The mutual action of “Vererbung” and “Anpassung” also constitutes the physiological base of evolution: “Die ganze unendliche Mannichfaltigkeit der organischen Formen wird also in letzter Instanz lediglich durch die Wechselwirkung dieser beiden physiologischen Functionen, der Anpassung und der Vererbung hervorgebracht“ (Haeckel 1866, II 169). Natural selection, “der Kampf ums Dasein,” on the other hand, is understood by Haeckel as “Summe der besonderen Verhältnisse, unter denen diese Wechselwirkung überall stattfindet, und von denen sie in hohem Maasse begünstigt wird“ (Ibid.). In Haeckel’s view, this conception rendered adequately the basic conception of Darwin’s theory of evolution (“Grundgedanke(n) von Darwin’s Selections-Theorie.“ — ibid. 167). It may be obvious, though, that the problems Haeckel’s conception tried to solve were not those of Darwin. With respect to what was discussed so far, one particularly striking feature of Haeckel’s theory of evolution is the inferior role individual variations play in its frame, that is, the inferior role of exactly that kind of variation on which Darwin had based his theory. For Haeckel, individual variation (individuelle Abänderung – ibid. 202) is just one out of three instances of what he called “indirecte oder potentielle Anpassung” (ibid.), an adaptation that was taken to be of less significance for the formation of structure than the “directe oder actuelle Anpassung” (Ibid 207ff.). The latter is nothing else than adaptation by use and disuse. Adaptations by use and disuse, not individual random variations, form the centre of Haeckel’s theory of evolution. Having above all the animal kingdom in mind as may seem natural for a comparative zoological anatomist, Haeckel assigned particular importance to the active behaviour with which living beings respond to changing conditions of life: Indem sich der thierische Wille den veränderten Existenzbedingungen durch andauernde Gewöhnung, Uebung u. s. w. anpaßt, vermag er die bedeutendsten Umbildungen der organischen Formen zu bewirken. (Haeckel 1868, 190)5

It is, thus, hardly possible to miss the central role that adaptation by use and disuse played in Haeckel’s theoretical edifice. However, it could play this role only when inheritance of acquired characters was taken for granted. And indeed, Haeckel did take inheritance of acquired characters as an established fact. In his Generelle Morphologie, Haeckel dedicated a sizable chapter of nearly 140 pages to the issue of evolution.6 About twenty pages of this chapter deal with the issue of heredity. 7 In the 5

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The stress Haeckel laid on volition has an almost personal tinge as may be taken from the following passage: “Der Umfang meiner ganz ungeübten Oberarme hatte sich innerhalb eines Zeitraumes von anderthalb Jahren durch fortgesetzte energische Turn-Uebungen fast genau verdoppelt. Dieses enorme Muskelwachsthum und die damit verbundene Uebung der Willens-Vorstellungen wirkte nun mächtig zurück auf die übrigen Vorstellungen meines Gehirns und insbesondere auf diejenigen des Denkens. Ihnen verdanke ich zum grossen Theile (zum großen Theile allerdings auch anderen cumulativ einwirkenden Ursachen), daß die in meinem Gehirne vorherrschenden dualistischen und teleologischen Irrthümer immer mehr den monistischen und causalen Vorstellungen wichen und ihnen zuletzt vollständig das Feld liessen“ (Haeckel 1866, II 213).

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centre of his argument, one finds the distinction between “conservative” and “progressive” inheritance, and both kinds of inheritance are given the form of a law. The law of conservative inheritance reads: Jeder Organismus vererbt dieselben morphologischen und physiologischen Eigenschaften auf seine Nachkommen, welche er selbst von seinen Eltern und Vorfahren ererbt hat. (Haeckel 1866, II 178)

And the law of progressive inheritance reads: Jeder Organismus vererbt auf seine Nachkommen nicht bloss die morphologischen und physiologischen Eigenschaften, welche er selbst von seinen Eltern ererbt, sondern auch einen Theil derjenigen, welche er selbst während seiner individuellen Existenz durch Anpassung erworben hat. (Ibid. 178f.)

Subsequently, a couple of more specific laws follow for each of the two kinds of inheritance. In case of conservative inheritance, these specific laws concern 1.) continual inheritance, 2.) interrupted inheritance (traits are not expressed in each generation), 3.) sexual inheritance (the expression of so-called secondary sexual characters), 4.) mixed inheritance (blending of nonsexual characters), and 5.) abbreviated inheritance (succession in which inherited characters show up in ontogenesis). How interesting ever these laws of conservative inheritance may be, in our context, the specific laws of progressive inheritance are naturally of particular concern. These laws are the following: 6. Gesetz der angepassten oder erworbenen Vererbung: Alle Charactere, welche der Organismus während seiner individuellen Existenz durch Anpassung erwirbt, und welche seine Vorfahren nicht besassen, kann derselbe unter günstigen Umständen auf seine Nachkommen vererben. (Ibid. 186) 7. Gesetz der befestigten Vererbung: Alle Charaktere, welche der Organismus während seiner individuellen Existenz durch Anpassung erwirbt, und welche seine Vorfahren nicht besassen, werden um so sicherer und vollständiger auf alle folgenden Generationen vererbt, je anhaltender die causalen Anpassungs-Bedingungen einwirkten, und je länger sie noch auf die nächstfolgenden Generationen einwirken. (Ibid. 187) 8. Gesetz der gleichörtlichen Vererbung: Alle Organismen können die bestimmten Veränderungen irgend eines Körpertheils, welche sie während ihrer individuellen Existenz durch Anpassung erworben haben, und welche ihre Vorfahren nicht besassen, genau in derselben Form auf denselben Körpertheil ihrer Nachkommen vererben. (Ibid. 188) 9. Gesetz der gleichzeitlichen Vererbung: Alle Organismen können die bestimmten Veränderungen, welche sie zu irgend einer Zeit ihrer individuellen Existenz durch Anpassung erworben haben, und welche ihre Vorfahren nicht besassen, genau in derselben Lebenszeit auf ihre Nachkommen vererben. (Ibid. 190) 6 7

Chapter 19 of the second volume with the title “Die Descendenz-Theorie und die Selections-Theorie” — (Haeckel 1866) II 148-286. (Haeckel 1866) II 171-190; in (Haeckel 1868), see the lectures 8 and 9.

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If one asks on which empirical material or what kind of investigations Haeckel rested these laws, one may be surprised to find that he thought it sufficient to add to each of them merely some remarks. To the first and most fundamental law of progressive inheritance, for example, he added the following assertions and remarks: Gleichwie alle von den Voreltern ererbten, so können auch alle neu erworbenen Eigenschaften der Materie durch die Vererbung fortgepflanzt werden. Es giebt keine morphologischen und physiologischen Eigenthümlichkeiten, welche das organische lndividuum durch die Wechselwirkung mit der umgebenden Aussenwelt erwirbt, mit einem Worte keine "Anpassungen", welche nicht durch Vererbung auf die Nachkommenschaft übertragen werden könnten. Dieses grosse Gesetz ist von der höchsten Wichtigkeit, weil darauf unmittelbar die Veränderlichkeit der Arten, die Möglichkeit, dass verschiedene neue Species aus einer vorhandenen hervorgehen, beruht. Wir kennen in der That keine einzige, in die Mischung, Form oder Function des Organismus eingreifende Veränderung, welche nicht unter bestimmten (uns gewöhnlich ganz unbekannten) Verhältnissen auf wenige, oder auf viele Generationen hinaus vererbt werden könnte. Am leichtesten geschieht dies, wenn die Veränderung sehr langsam und allmählich erfolgt (wie z. B. bei Erwerbung chronischer Krankheiten, die viel leichter als acute vererbt werden). Am schwersten dagegen tritt die Vererbung der Veränderung ein, wenn die letztere ganz plötzlich (z. B. traumatisch) erfolgte.8 Gewöhnlich springen die Fälle, wo eine plötzlich aufgetretene Veränderung auf eine oder mehrere Generationen vererbt wird, sehr deutlich dann in die Augen, wenn die betreffende Veränderung eine "monströse" ist, d. h. einzelne Theile des Organismus in ungewöhnlicher Zahl, Grösse, Form oder Farbe entwickelt zeigt, so z. B. die Fälle, in denen sechs Finger an jeder Hand mehrere Generationen hindurch beim Menschen vererblich blieben, ferner die berühmten StachelschweinMenschen aus der Familie Lambert, wo eine eigenthümliche schuppenähnliche monströse Hautbildung von Edward Lambert an (1735) sich durch mehrere Generationen auf die Nachkommen vererbte, und zwar bloss auf und durch die männlichen Nachkommen. Auch die häufigen Fälle von erblichem Albinismus gehören hierher, ferner die Fälle, wo ein einzelner Schafbock oder Ziegenbock mit keinem oder mit 4 - 8 Hörnern geboren wurde, und nun diesen individuellen Charakter auf seine Nachkommen übertrug. Viel wichtiger, als diese monströsen, auffallend vortretenden Abänderungen, welche durch die angepasste Vererbung übertragen werden, sind die unscheinbaren und geringfügigen Abänderungen, welche erst im Laufe von Generationen durch Häufung und Befestigung ihre hohe Bedeutung für die Umbildung der organischen Form erhalten. Die gesammten Vorgänge der künstlichen Züchtung liefern in dieser Beziehung für das Gesetz der angepassten Vererbung eine lange Beweiskette. (Ibid. 186f.)

That’s all he regarded necessary for making his case. When assessing these remarks, one has certainly to allow for the fact that Haeckel was first of all a comparative anatomist and not a physiologist or cytologist.9 One has, furthermore, to take into account that he could obviously expect his — professional! — audience to accept such 8

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“Gewöhnlich werden bekanntlich traumatische oder durch Verwundung entstandene Veränderungen nicht vererbt. Um so wichtiger ist es, die Fälle aufzubewahren, in denen dies doch bisweilen geschieht. So wurden kürzlich, wie mir Herr Hofrath Stöckhardt als sicherer Gewährsmann mittheilte, auf einem Gute in der Nähe von Jena mehrere schwanzlose Kälber geboren, deren Vater der Schwanz beim unvorsichtigen Zuschlagen eines Thores eingeklemmt und abgequetscht worden war.” (Original note by Haeckel.)

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cursory expositions. This proves again that the issue of inheritance of acquired characters was still in its indifferent stage described earlier. However, as a consequence of this, Haeckel unknowingly ran a high risk in treating the assumption of inheritance of acquired characters that cursory, although this assumption had strategic significance for his theory of evolution. Involuntarily, he contributed by this assumption’s promotion to an urgent topic of biological thought in a twofold way: as a strategic element of a prominent theory that stands and falls with it; and, at the same time, as a weakly defended element that suggested itself for being challenged by critics of this theory. In any case, in Germany, the sustained challenge of the old assumption of inheritance of acquired characters that led eventually to its dismissal began in the 1880s with challenges of Haeckel’s theory of evolution. Two extremely different such challenges may briefly be addressed in the final part of this paper. IV. DISMISSALS OF AN INHERITANCE OF ACQUIRED CHARACTERS

a. August Weismann It is probably not possible to miss the advantages a Neo-Lamarckian theory of evolution would have if it only were compatible with the body of approved biological knowledge, and particularly with genetics. If one could, like Haeckel, presuppose inheritance of acquired characters and, based on it, adaptation by use and disuse as the centre of evolution, it would no longer be a matter of chance whether modifications of structure develop that are adapted to changed conditions of life. Rather, such modifications would be induced just by those changed conditions through the organism’s reaction they provoke. In contrast to this, a Darwinian theory of evolution may appear rather implausible and unintuitive. Regarding the historical development of organismic forms as result of natural selection of random variations, this theory demands to embark in an evolution without guarantees and safety net. It comes therefore without surprise that the idea of a historical evolution of species had first success, not in a Darwinian, but in a Neo-Lamarckian fashion. 10 The rapid victory of the idea of evolution among German biologists since the late 1860s was due exactly to the fact that not a Darwinian theory had been presented to them. What Haeckel successfully offered as Darwinism was, as is clear in hindsight, a Neo-Lamarckian theory of evolution. What is surprising against this background, is the fact that there were at all biologists who read Darwin’s theory differently and attacked exactly the Neo-Lamarckian elements of Haeckel’s theory. The best known of these biologists is certainly August Weismann. It is, however, not clear to me what made him start his critique of Haeckel and try to restore a “pure” Darwinism that seems to be purer than that of Darwin himself. At least his contemporaries conceived of it as a radically new theory of evolution and called it Neo-Darwinism. The centrepiece of Weismann’s critique was his categorical dismissal of inheritance of acquired characters. This rejection and its cytological underpinning count rightly as a decisive step towards modern genetics. 11 Instead of elaborating this well-known story, I want to point to a less known contemporary challenge to 9

10

The physiological theory of heredity he designed nevertheless in later years, his “Perigenesis” hypothesis (Haeckel 1876), reminds, all differences notwithstanding, of Darwin’s Pangensis in that the Perigenesis, too, entails inheritance of acquired characters by implication. See (Stubbe 1965) 156ff. For the success of non-Darwinian theories of evolution in the last decades of the nineteenth century, see (Bowler 1983) and (Bowler 1988).

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Haeckel’s theory, namely two orthogenetic theories of evolution 12 which also dismissed inheritance of acquired characters.

b. Orthogenesis A key to an adequate understanding of Haeckel’s theory of evolution can be seen in the fact that he developed it in the frame of his Generelle Morphologie. From its very beginnings, his conception of evolution was part and parcel of an “Organische Formen-Wissenschaft” as the subtitle of the work spells out the meaning of “morphology.” This science claimed to establish general laws and comprehensive and consistent explanations of the formative processes of organismic structure. The enterprise focussed particularly on actual or seeming regularities of such formative processes as observed by embryologists and anatomists in comparative studies. By such regularities, above all one law of development seemed to be suggested, namely that of a development from simple to complex, from homogeneous to articulated organismic structure with a differentiation of function among its parts and organs. It was not this view of order in the variety of organismic forms that distinguished Haeckel from his colleagues. Embryologists like Karl Ernst von Baer and comparative anatomists like Richard Owen had tried to establish laws of formative processes. However, the regularities, or “logic,” of organismic forms these biologists investigated was understood by them in a way that is usually called that of an idealist or transcendental morphology.13 What distinguished Haeckel’s attempt from these conceptions was his materialist approach. He presented a morphology that claimed to provide a “mechanical”14 explanation of the laws that rule the formative processes. And in this morphology, Darwin’s theory as understood by Haeckel turned out to be the key to such a mechanical explanation. In the late 1860s, under the impact of Darwin’s theory, many biologists were prepared to accept a real, historical evolution as a possible or even likely rationale of the regularities and laws that seemed to connect organismic forms. Yet, what they did reject was Haeckel’s idea that such a lawful development of organismic forms could be explained through a process of adaptations to outer conditions of life, which apparently changed without perceivable regularities. This rejection seems well grounded. For these biologists, Haeckel was linking two issues that couldn’t be linked in this way: namely, on the one hand, adaptation to external conditions of life and, on the other, laws of development of structure that, in the view of these biologists, require inner-organismic causes for their explanation. In what Haeckel had tried to combine, namely laws of formation of structure and adaptation, an increasing number of contemporary biologists saw a clear alternative: If, as for Darwinists, all formation of structure was ultimately brought about by adaptations to external conditions of life, 11 12

13 14

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For the question of whether Weismann’s later theory of “germ selection” contradicts these celebrated achievements, see, for instance, (Ridley 1982) 63ff. The term “orthogenesis” (Orthogenese) was coined by the zoologist Wilhelm Haacke. — For characteristic features of these theories of evolution, see (Mayr 1982) 528ff. And, for a contemporary view, (Wagner 1908) 223ff. For Owen, see for instance (Rupke 2001) 252; for von Baer, see for instance (Muzrukova 2001) 307. See the subtitle of the second volume of (Haeckel 1866): “Kritische Grundzüge der mechanischen Wissenschaft von den entstehenden Formen der Organismen.”

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no laws that rule those formation processes could be assumed. 15 And, vice versa, if these processes are ruled by such laws, they cannot be explained by adaptations to external conditions. Rather, increase of complexity of organismic structure, ever more specialised differentiation of function, in short, the tendency of a progressive development in the realm of organismic forms, seemed to suggest an inner-organismic formative drive. The assumption of such an inner force of development which, in this situation, became attractive to many biologists constituted the core of orthogenetic theories of evolution, which began to challenge Haeckel’s theory from another quarter since the 1870s. I cannot go into the different orthogenetic theories of evolution proposed in the last decades of the nineteenth century. I have to confine myself to naming just two well-known biologists as proponents of this variety of evolutionary thinking, namely Carl Wilhelm von Naegeli und Albert von Koelliker. The two men worked on quite different fields — Naegeli mainly on physiology and anatomy of plants, Koelliker mainly on zoological morphology and embryology — and can in no way be regarded adherents of a certain school. Rather, their research agenda as well as general orientation was quite different. And so are the orthogenetic theories of evolution each of them proposed. 16 There are nevertheless remarkable congruities between their theories besides the shared core assumption of all orthogenetic theories, that is, the assumption of an inner-organismic cause that rules the formative processes of organismic structure. Each of the two men recommended his theory as a truly mechanical-physiological, that is, as a non-teleological one. Both men shared furthermore the conviction that evolution does not necessarily entail a common origin of species and held that each species developed in an independent process of evolution. Finally, though probably not very surprisingly, both men dismissed inheritance of acquired characters. Dismissal, or at least marginalization, of inheritance of acquired characters was characteristic of almost all orthogenetic theories of evolution,17 as was dismissal or marginalization of adaptation by use and disuse, of natural selection, and also of Haeckel’s biogenetic law. All of these items belonged to theories of evolution that conceive of it as essentially a process of adaptation to changing conditions of life. They were hall-marks of theories that regarded organismic structure as evolving in reaction to external conditions, that is, hall-marks of exactly those theories that 15

16

17

In this paper, I cannot go into the issue of undeniable optimisations of “natural technology” (Marx) in the course of evolution. Such optimisations cannot be explained by adaptation processes that take place alone or foremost between living beings and inorganic nature. Rather, first adaptations between living beings themselves assign to them a “teleonomic” character (Mayr) on which such optimisations rest — see for instance (Lefèvre 1984) 252ff. and 260ff. Naegeli published an elaborate version of his orthogenetic theory first in 1884 (Naegeli 1884); yet he presented outlines of it in several talks and articles since 1865 (Naegeli 1865). Koelliker’s orthogenetic theory can be found in part A of (Koelliker 1872). — It is, by the way, remarkable how many botanical physiologists and anatomists endorsed any form of ortogenetic theories. It may suffice to name Eugen Askenasy (Askenasy 1872) and above all Julius Sachs (Sachs 1894) who had long been the most devoted Darwinist among German botanists — see (Höxtermann 2001) and (Junker 1989) 233ff. The latter book is of general interest as regards the reception of the theory of evolution among German botanists. Apart from Naegeli’s and Koelliker’s theories, one should name those of Wilhelm His and Alexander Wilhelm Goette — see (Montgomery 1972) 102f. A noteworthy exception is Theodor Eimer’s orthogenetic theory, which assumed a “stammesgeschichtliches Wachsen” that develops organismic structure in reaction to environmental factors but is restricted by inner forces and tendencies — (Eimer 1897).

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adherents of orthogenetic theories strived to overcome. As regards inheritance of acquired characters, proponents of orthogenesis did not doubt, deny, and dismiss this assumption for special genetical arguments. Rather inheritance of acquired characters was dismissed because it had become part and parcel of rejected theories of evolution. V. CONCLUSION In closing, I would like to sum up my paper by four statements. First, the question of inheritance of acquired characters was put on biologists’ agenda of urgent issues immediately when heredity itself became a major biological topic. (For this, I am assuming that heredity was not such a topic before the last third of the nineteenth century.) Second, biologists tried to decide definitely whether or not inheritance of acquired characters is possible just when first foundations for an answer were laid in physiology, particularly in cytology. The case, thus, seems to confirm Marx’s dictum that mankind never poses a problem before the means for its solution are at hand. However, thirdly, it has the appearance that, in the beginning, inheritance of acquired characters became a fiercely debated issue not so much in the framework of competing theories of heredity but, so to speak, as a collateral damage of a war between competing theories of evolution. Inheritance of acquired characters was (and is) an element of strategic importance for NeoLamarckian approaches to evolution. Fourth and last, against this background, it seems not that surprising that the periodical revivals of inheritance of acquired characters in the twentieth century were often indicative not of new insights in heredity but of lasting difficulties and debates in the framework of evolution.

Wolfgang Lefèvre, MPI für Wissenschaftsgeschichte, Berlin [email protected]

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References Askenasy, Eugen. 1872. Beiträge zur Kritik der Darwinschen Lehre. Heidelberg: Engelmann. Beurton, Peter J. 2001. Hintergründe des modernen Lamarckismus. Deutsche Zeitschrift für Philosophie IL (4): 537-548. Bowler, Peter J. 1983. The Eclipse of Darwinism. Anti-Darwinian Evolution Theories in the Decades around 1900. Baltimore & London: The Johns Hopkins University Press. ———. 1988. The Non-Darwinian Revolution. Reinterpreting a Historical Myth. Baltimore & London: The Johns Hopkins University Press. ———. 1989. The Mendelian Revolution. Baltimore: The Johns Hopkins University Press. Crick, Francis H.C. 1958. On Protein Synthesis. Symposia of the Society for Experimental Biology XII: 138163. Eimer, Theodor. 1897. Die Entstehung der Arten auf Grund von Vererben erworbener Eigenschaften nach den Gesetzen organischen Wachsens. Teil 2: Orthogenesis der Schmetterlinge. Ein Beweis bestimmt gerichteter Entwickelung und Ohnmacht der natürlichen Zuchtwahl bei der Artbildung; zugleich eine Erwiderung an August Weismann. Jena: Fischer. Gayon, Jean. 1998. Darwinism's Struggle for Survival. Heredity and the Hypothesis of Natural Selection. Cambridge: Cambridge University Press. Haeckel, Ernst. 1866. Generelle Morphologie der Organismen. Allgemeine Grundzüge der organischen FormenWissenschaft, Mechanisch begründet durch die von Charles Darwin reformirte Descendenz-Theorie. 2 vols. Berlin: Reimer. ———. 1868. Natürliche Schöpfungsgeschichte. Gemeinverständliche wissenschaftliche Vorträge über die Entwickelungslehre im Allgemeinen und diejenige von Darwin, Goethe und Lamarck, im Besonderen über die Anwendung derselben auf den Ursprung des Menschen und andere damit zusammenhängende Grundfragen der Naturwissenschaft. Berlin: Reimer. ———. 1872. Die Kalkschwämme. 2 vols. Berlin: Reimer. ———. 1876. Die Peregenis der Plastidüle oder die Wellenerzeugung der Lebenstheilchen. Berlin: Reimer. Höxtermann, Ekkehard. 2001. Julius Sachs. In Darwin & Co. Eine Geschichte der Biologie in Portraits, edited by I. Jahn and M. Schmitt. München: C. H. Beck. Junker, Thomas. 1989. Darwinismus und Botanik. Rezeption, Kritik und theoretische Alternativen im Deutschland des 19. Jahrhunderts. Stuttgart: Deutscher Apotheker Verlag. Koelliker, Albert. 1872. Morphologie und Entwicklungsgeschichte des Pennatulidenstammes nebst allgemeinen Betrachtungen zur Descendenzlehre. Frankfurt: Winter. Lefèvre, Wolfgang. 1984. Die Entstehung der biologischen Evolutionstheorie. Frankfurt, Berlin, Wien: Ullstein. Mayr, Ernst. 1982. The Growth of Biological Thought: Diversity, Evolution, and Inheritance. Cambridge, Mass., and London: The Belknap Press of Harvard University Press. Montgomery, William M. 1972. Germany. In The Comparative Reception of Darwinism, edited by T. F. Glick. Austin und London: University of Texas Press. Muzrukova, Elena. 2001. Karl Ernst von Baer. In Darwin & Co. Eine Geschichte der Biologie in Portraits, edited by I. Jahn and M. Schmitt. München: C. H. Beck. Naegeli, Carl von. 1865. Entstehung und Begriff der Naturhistorischen Art. München: Vlg. der Königl. Akademie. ———. 1884. Mechanisch-physiologische Theorie der Abstammungslehre. München und Leipzig: Oldenbourg. Ridley, Mark. 1982. Coadaptation and the Inadequacy of Natural Selection. British Journal for the History of Science XV: 45-68. Rupke, Nicolaas. 2001. Richard Owen. In Darwin & Co. Eine Geschichte der Biologie in Portraits, edited by I. Jahn and M. Schmitt. München: C. H. Beck. Sachs, Julius. 1894. Mechanomorphosen und Phylogenie: Physiologische Notizen VIII. Flora LXXVIII: 21543. Stubbe, Hans. 1965. Kurze Geschichte der Genetik bis zur Wiederentdeckung der Verebungsregeln Gregor Mendels. Jena: VEB Gustav Fischer Verlag. Vorzimmer, Peter J. 1972. Charles Darwin: The Years of Controversy. The "Origin of Species" and its Critics, 1859 - 82. London: University of London Press. Wagner, Adolf. 1908. Geschichte des Lamarckismus. Als Einführung in die psycho-biologische Bewegung der Gegenwart. Stuttgart: Franck'sche Verlagshandlung.

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Weismann, August. 1883. Über die Vererbung. Jena: Gustav Fischer. Zirkle, Conway. 1946. The Early History of the Idea of the Inheritance of Acquired Characters and of Pangenesis. Transactions of the American Philosophical Society New Series XXXV (2): 91-151.

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Darwinism versus Evo–Devo: a late–nineteenth century debate Jeffrey H. Schwartz

Abstract In his notebooks and culminating in the two volume publication on domestication of plants and animals (Darwin 1868), Charles Darwin developed a theory of inheritance, pangenesis, that fit his worldview: through a continuum of variants and intermediate forms, individuals and species graded one into another both syn– and diachronically. Pangenesis accommodated Darwin’s fundamental assumptions: use–disuse, natural selection (as a factor in producing variation and then choosing from the resultant variants), and blending inheritance. He conceived of “gemmules,” which were constantly thrown off from every part of an organism’s body throughout its entire lifetime, as the recorders of everything that impacted an individual during its lifetime. By way of bodily “fluids,” gemmules were conveyed to an individual’s sex organs, to be passed on en masse to offspring, combining upon fertilization with those contributed by the other parent. It was an ingenious idea, especially in its attempt to explain all known phenomena, including the appearance of new or different features (such as sexually dimorphic ones) later in life. Blending inheritance expanded the realm of possible variation in the next generation. The thrust of Darwin’s theory of pangenesis was a justification of his view that the gradual nature of evolutionary change and continuous variation tied all life together in a seamless web. Ironically, the very examples Darwin enumerated for the origin of new breeds or varieties of domesticated plants and animals, and their stabilization through inbreeding and breeding with parental strains, lend themselves not to a model of gradual change and continuous variation, but to a saltationist one, in which morphological novelty emerges abruptly and yet its bearers remain capable of reproducing with the original stock. Victorian saltationism, as articulated by St. George Mivart in (Mivart1871) in On the Genesis of Species, envisioned alterations in development as the basis of major and instantaneous change, but it did not deal with heredity. Mivart pointed out that a major problem with Darwin’s gradualism was that features critical to an organism’s survival and reproductive competence would be useless unless they were functional from the beginning. In addition, as Huxley (Huxley 1860) had argued well before Mivart, natural selection played no role whatsoever in producing change. Since it was obvious that some hereditary process connected successive generations, perhaps saltationists thought it unnecessary to speculate on a mechanism of inheritance because the idea of novelty arising through alterations of developmental processes was not explained by gemmule, germ–plasm, or any other available model of inheritance. Interestingly, the contrasts between Darwin’s theory of heredity and Mivart’s emphasis on organismal change being due to altering developmental processes, is being replayed in the opposing views of present–day Darwinism and evo–devo.

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Introduction A consequence of delving into the history of evolutionary thought is often seeing how a scholar, when confronted with examples of biological phenomena that could lead to diametrically opposed and contradictory models or theories, chooses one set as representing the reality of nature and regards the other as irrelevant. In such instances, the obvious question is: Why? What were the reasons behind the decision? In this regard, an interesting case is the contrast between Charles Darwin (especially his “evidence” for natural selection and gradual evolutionary change as being necessary for the origin of species and the assumptions underlying his theory of inheritance, “pangenesis”) and the saltationists (especially as represented by St. George Mivart).

Darwin’s bias Although one cannot ignore On the Origin of Species (Darwin 1859 et seq.; herewith referred to as the Origin), for this discussion I shall focus on The Variation of Animals and Plants under Domestication (Darwin 1868; herewith referred to as Variation). The rationale for this emphasis is that, in Variation, Darwin not only argues that the character of domesticated plants and animals and the process of domestication via artificial selection make perfect analogues for the character of natural species and the process of evolution via natural selection. In this work Darwin also attempts to articulate a theory of inheritance that embraced his bias toward gradualism. Since the basics of Darwin’s assumptions as presented first in the Origin are well known, I shall summarize them only briefly. 1) Variation is essentially infinite and, in a perfectly preserved synchronous and diachronous world, one would be able to observe continuous variation and insensible and infinitesimal gradation between individuals, sexes, and species. 2) At any point in time, natural selection both produces and then chooses from the resultant more fit or better adapted variations. 3) The consequences of use and disuse contribute significantly to the emergence of variability. 4) Evolutionary change proceeds gradually through the accumulation of continually produced, infinitesimally small variations. In arguing the first assumption, Darwin had to confront the discontinuities between extant species and the “gaps” in the paleontological record between extinct species as well as between extinct and extant species. This absence of evidence was explained neontologically and paleontologically by invoking the extinction or elimination of individuals that would have formed a graded series of intermediates between one species and another. The paleontological conundrum was also addressed from a taphonomic perspective: There had been intermediates, but the deposits containing them no longer existed. With the problems of discontinuity seemingly dealt with, Darwin was then free to attend to other matters, such as the origin of species: The differences between natural varieties are slight; whereas the differences are considerable between the species of the same genus, and great between the species of distinct genera. How do these lesser differences become augmented into the greater differences? How do varieties, or as I have called them incipient species, become converted into true and well–defined species? (Variation, vol. 1, p. 5)

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Although he had already committed himself in the Origin to answering this question by invoking a gradualistic model of change, Darwin was clearly aware of examples, not only from nature, but also from plant and animal domestication, that could have led him to formulate a different model of evolutionary tempo. For example, in volume 1 of Variation (pp. 92–94) he commented on how the niata breed of cattle had appeared suddenly, in the course of one generation, and then described in detail how they differed from common cattle in numerous aspects of their anatomy. For example: In fact, on comparison with the skull of a common ox, scarcely a single bone presents the same exact shape, and the whole skull has a wonderfully different appearance. (p. 94)

Although they appeared suddenly, and their features were so different from those of common cattle, niata cattle could breed successfully with one another, as well as with common cattle. In the latter case, Darwin even discussed the specific characters that were often dominant in offspring when a niata cow was mated with a common bull, and vice versa. Nevertheless, as profound as the differences between niata and common cattle were, and even with the former being reproductively unimpaired, Darwin rejected the case of niata cattle as being representative of how novelty might arise in the wild. His argument was that these cattle were not as adaptable as common cattle when environmental conditions occasionally and drastically changed for the worse, as in periods of drought. As such, Darwin (Variation, vol. 1, p. 94) concluded, “[this] shows us…how natural selection would have determined the rejection of the niata modification had it arisen in a state of nature.” But niata cattle were not the only example of the sudden appearance of marked novelty in animals or plants without loss of reproductive viability of which Darwin was aware. Indeed, it was precisely because these “monstrosities” could mate successfully with one another (as well as with “normal” individuals) that breeders could perpetuate the novelty. For instance: [The sprouting–broccoli] variety is a new one, and bears the same relation to common broccoli, as Brussel–sprouts do to common cabbages; it suddenly appeared in a bed of common broccoli, and was found faithfully to transmit its newly–acquired and remarkable characters (vol. 1, p. 342). Domestic breeds often have an abnormal or semi–monstrous character, as amongst dogs…some breeds of cattle and pigs,–several breeds of fowl,–and the chief breeds of pigeon. In such abnormal breeds, parts which differ but slightly or not at all in the allied natural species, have been greatly modified. This may be accounted for by man’s often selecting, especially at first, conspicuous and semi–monstrous deviations of structure (volume 2, p. 408).

Even in terms of so–called atavistic structures, Darwin knew not only that they re–appeared suddenly, but also that they were morphologically recognizable structures. They were not mere hints of features that had been present in their bearers’ ancestors. As he noted in volume 1 of Variation: Horses have often been observed, according to [the French paleontologist] M Gaudry, to possess a trapezium and a rudiment of a fifth metacarpal bone, so that “one sees appearing by

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monstrosity, in the foot of the horse, structures which normally exist in the foot of the Hipparion [an allied and extinct animal]” (p. 52; comments added).

Yet, in the face of numerous examples of the sudden appearance of novelty in domesticated plants and animals, Darwin argued that they were not reflective of what really occurs in nature: There is a much more important distinction between our several breeds, namely, in some having originated from a strongly–marked or semi–monstrous deviation of structure, which, however, may subsequently have been augmented by selection; whilst others have been formed in so slow and insensible a manner, that if we could see their early progenitors we should hardly be able to say when or how the breed first arose…[But] it is certain that the ancon and mauchamp breeds of sheep, and almost certain that the niata cattle, turnspit, and pug–dogs, jumper and frizzled fowls, short–faced tumbler pigeons, hook–billed ducks, &c., suddenly appeared in nearly the same state as we now see them. So it has been with many cultivated plants. The frequency of these cases is likely to lead to the false belief that natural species have often originated in the same abrupt manner. But we have no evidence of the appearance, or at least of the continued procreation, under nature, of abrupt modifications of structure; and various general reasons could be assigned against such a belief. On the other hand, we have abundant evidence of the constant occurrence under nature of slight individual differences of the most diversified kinds; and we are thus led to conclude that species have generally originated by the natural selection of extremely slight differences. (vol. 2, pp. 409–410) Some naturalists boldly insist that species are absolutely distinct productions, never passing by intermediate links into one another; whilst they maintain that domestic varieties can always be connected either with one another or with their parent–forms. (vol. 2, p. 409)

But does demonstrating “abundant evidence of the constant occurrence under nature of slight individual differences of the most diversified kinds” necessarily contradict inferring from observing the sudden appearance of novel features in domesticated organisms that this process also occurs in nature? Does demonstration of “abundant evidence of the constant occurrence under nature of slight individual differences of the most diversified kinds” necessarily lead only to the conclusion that “species have generally originated by the natural selection of extremely slight differences”? The answer to both of these questions is, I believe, no. But by taking the position he did, Darwin conflated the existence of individual variation, which reflects slight degrees of difference in the expression of a particular feature or array of features, with the advent of the feature itself. And it was this unfounded conflation of two entirely different biological phenomena that informed his belief in continuous variation and continuity via an insensible gradation between species through time and at any point in time. Accordingly, and without any justification or demonstration, Darwin could then claim that “[v]ariations often pass into, and cannot be distinguished from, monstrosities; and monstrosities are of little significance for our purpose” (vol. 1, p. 322). Indeed, for the myriad examples Darwin gives in the Origin and in Variation of “monstrosities” and of slight variations of a feature or features, none supports an assumption of

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gradation. Darwin merely asserts that this is the case, just as, in the same quote, he can so easily declare, “monstrosities are of little significance for our purpose.” But Darwin persists in volume 2 of Variation in asserting the reality of insensible gradation [e.g. “…we so incessantly see in species of the same group the finest gradations between an organ in a rudimentary and perfect state, that we are led to believe that the passage must have been extremely gradual” (p. 308)]. He also attempts to explain why sudden change, at least in wild species, could not occur: “It may be doubted whether a change of structure so abrupt as the sudden loss of an organ would ever be of service to a species in a state of nature; for the conditions to which all organisms are closely adapted usually change very slowly” (p. 308). This is an interesting approach to denying the possibility of sudden change because, here, Darwin focuses on the abrupt loss of structure, not, as with virtually every other example he musters in the Origin and Variation in support of gradual change, the emergence of structure. He then states with unfounded assurance that abrupt change not only would not benefit wild species, but also that it could not occur because organisms adapt gradually to their slowly changing surroundings. Yet Darwin knows that sudden, non–reproductively disruptive change occurs. He also knows, at least from the example of niata versus common cattle responding to drought, that the environment can change abruptly, without provoking visible organismal change at the same time. But, here and elsewhere (see quote further above) he draws a line of distinction between domesticated and wild species, and seeks to justify this distinction by asserting that no one has observed abrupt change in nature. This, of course, is an assertion without basis, because, by definition (and observation in domesticates) there is nothing to observe if change occurs suddenly. There is only the presence of something previously unknown to you whose origin would be a mystery. Thus, while on the one hand, Darwin would like his audience to believe that, as artificial selection can gradually change the character of a domesticated species or breed, so, too, can natural selection act on wild species. On the other hand, Darwin asserts without justification that the sudden appearance (not loss) of novelty in domesticated species – that is, the emergence of “monstrosities” – has no bearing on insights into the workings of the organisms in the “wild.” Still, by asking us to accept the analogy between artificial and natural selection as agents that gradually alter the character of species, and to deny any biological significance to the sudden appearance of novelty in domesticates (and thus natural species), Darwin perpetuates the misconception that individual variation in the expression of a feature is somehow relevant to the origin of the feature itself. That, by artificially shifting the bell curve of expressed variation of a feature, this serves as evidence that, under natural conditions and with enough time, a feature can be transformed into something entirely different.

Darwin’s theory of pangenesis Without doubt, the climax of Darwin’s formulations in Variation was the model of inheritance that he called “Pangenesis.” In many ways, it was quite elegant (see review in Schwartz, 1999). Through pangenesis Darwin could explain, for instance, features that appear early in development as well as those that emerge later in life (e.g. differences in secondary sexual characters). The idea was simple. All parts of an organism issue small particles, which Darwin identified as gemmules,

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throughout the individual’s life. Consequently, the entire life history of an individual, from conception until death, is recorded in a trail of gemmules. Individually unique events, such as those that result from use or disuse, would also be recorded in the gemmules of the affected part or parts and thus affect the pool of potential variation. Traveling by way of some unspecified bodily fluid, gemmules accumulate in an individual’s sex organ. Upon mating, parental gemmules blend, thereby producing additional sources of variation. Gemmules could also become latent and not expressed over a series of generations. But at some later time, they could become active again, which would account for atavisms. Pangenesis solidified Darwin’s ideas on blending inheritance (hinted at in the Origin, although clearly expressed in his notebooks; see Schwartz 1999) and also provided a previously unspecified mechanism for the transmission of acquired characteristics resulting from use or disuse. The theory also increased the number of ways in which individual variation could be produced. Indeed, pangenesis seemed to be able not only to account for the entirety of an organism’s being, but also to provide the fodder necessary for natural selection to slowly transform one species into another.

The opposing saltationist view Three years after the publication of Variation, St. George Mivart, one of England’s leading comparative morphologists, published On the Genesis of Species (Mivart 1871, herewith referred to as Genesis). This was the saltationist’s response to Darwin’s notions of gradual change, the role of natural selection, the essence of variation, and the viability of pangenesis. Although in his review of the Origin, Huxley (Huxley 1860) was clearly strongly opposed to gradualism and a role of primacy of natural selection in producing change, Mivart makes it appear as if his fellow saltationist was not fully committed to this position: Professor Huxley seems now disposed to accept the, at least occasional, intervention of sudden and considerable variations. In his review of Professor Kölliker’s criticisms, he himself, says, “We greatly suspect that she” (i.e. Nature) “does make considerable jumps in the way of variation now and then, and that these saltations give rise to some of the gaps which appear to exist in the series of known forms.” (Genesis, pp. 103–4)

In mounting his case for saltationism, Mivart paralleled Darwin in compiling a massive array of examples from the plant and animal worlds. But instead of focusing on the minutiae of individual differences, Mivart called attention to the major ways in which species differ from one another, whether it be in the stamens and anthers of flowers, the pincers or antennae of beetles, the configuration of the vertebrate versus invertebrate eye, the presence of mammary glands in mammals, feathers in birds, and baleen in whales, or differences between organisms in their reproductive anatomies. In contrast to Darwin’s rejection of sudden change solely on the basis of the argument in Variation on the lack of benefit of losing of an organ, Mivart emphasized the emergence of novel structure. Not, of course, that the loss of structure could not be novel – consider the reduced numbers and generations of teeth in mammals compared to reptiles, of toes in modern horses, and of limbs in snakes. But, inasmuch as Darwin himself presented examples of “gain” rather than “loss” (e.g. the vertebrate eye) in both the Origin and Variation, it was

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appropriate that Mivart do the same, although his interpretation of the requisites for the appearance of novelty was vastly different. Time and time again Mivart discussed a remarkable trait and then raised the question: How could such a functionally important feature have evolved gradually, through an insensible gradation of intermediates, to its present state? How, for instance, could mammals not only have survived, but multiplied in number and become diverse, if the first mammal had merely possessed a vestige of a mammary gland, which, in turn, produced only a drop or two of milk? How could sexually reproductive organisms have persisted generation after generation if, initially, their reproductive organs were merely a hint of their necessary functional states? Turning Darwin’s argument of purpose on its head, Mivart asked: How could anything but the fully formed version of a feature, particularly one that was essential for sustenance of life or procreation, be beneficial to an organism? Using flatfish (soles, flounder) as one of many examples leading to doubting Darwin’s assumption, Mivart questioned the advantage of selection causing one of the fish’s eyes to be dragged gradually from one side of its head, across the rough sand of the ocean floor, until it reached its present position near the eye that had been on the opposite side of the body. Clearly, on various levels, the notion of gradual change did not make biological sense. But a saltational model for the advent of novelty – especially if functionally integral to the survival of an individual – was not only biologically sensible, it was also compatible with the pattern of life as illustrated in the fossil record. Indeed, Mivart (pp. 129–130) quotes from Fleming Jenkin’s devastating review of the Origin (“It is really strange that vast numbers of perfectly similar [fossil] specimens are so great; but it is also very strange that the specimens should be so exactly alike as they are, if, in fact, they came and vanished by a gradual change”), and then proceeds with his own argument: “The mass of paleontological evidence is indeed overwhelmingly against minute and gradual modification…[H]ad such a slow mode of origin, as Darwinians contend for, operated exclusively in all cases, it is absolutely incredible that birds, bats, and pterodactyles should have left the remains they have, and yet not a single relic be preserved in any one instance of any of these different forms of wing in their incipient and relatively imperfect functional condition!” Even the bird–like fossil reptile, Archaeopteryx, which had been discovered in 1861, and which Darwin would use in the sixth and last edition of the Origin in an attempted refutation of Genesis, did not pose a problem for Mivart (p. 131): “But even supposing all that is asserted or inferred on this subject to be fully proved, it would not approach to a demonstration of specific origin by minute modification. And though it harmonizes well with ‘Natural Selection,’ it is equally consistent with the rapid and sudden development of new specific forms of life.” In contrast to Darwin, who invoked forces external to the organism as the primary provocateurs of change (e.g. as in gradual environmental change, or use–disuse), Mivart hypothesized the source of novelty as lying primarily internally, within the cells of the organism itself. In formulating this theory, Mivart turned to the inert inorganic world for an analogy relevant to the living one. In apparent anticipation of some of Waddington’s (Waddington 1940) ideas many decades later, Mivart (Genesis, p. 114) suggested: Judging the organic world from the inorganic, we might expect, a priori, that each species of the former, like crystallized species, would have an approximate limit of form, and even of si-

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ze, and at the same time that the organic, like the inorganic forms, would present modifications in correspondence with surrounding conditions; but that these modifications would be, not minute and insignificant, but definite and appreciable, equivalent to the shifting of [a] spheroid on to another facet for support.

Mivart (pp. 114–115) then quotes from a Mr. Murphy (“Crystalline formation is also dependent in a very remarkable way on the medium in which it takes place…And [as] the Rev. E. Craig found that [different chemicals affected copper crystal growth]…[t]he changes take place not by the addition of new crystals, but by changing the growth of the original ones.”), after which he comments: “These, however, may be said to be the same species, after all; but recent researches by Dr. H. Charlton–Bastian seem to show that modifications in the conditions may result in the evolution of forms so diverse as to constitute different organic species.” In contrast to Darwin’s efforts to explain away the “gaps” in the fossil record, Mivart (p. 143) expands his model to incorporate them: Now all these difficulties [e.g. the absence of fossils in old strata and of intermediate forms] are avoided if we admit that new forms of animal life of all degrees of complexity appear from time to time with comparative suddenness, being evolved according to laws in part depending on surrounding conditions, in part internal–similar to the way in which crystals (and, perhaps from recent researches, the lowest forms of life) build themselves up according to the internal laws of their component substance, and in harmony and correspondence with all environing influences and conditions. [comment added]

But, what is the internal element of Mivart’s saltational theory? After all, one could claim that Darwin’s theory of pangenesis embodied an internal component to eventual evolutionary change. The difference between the two scholars lies in Mivart’s thinking in terms of novelty emerging as a result of alterations in the regulation of an organism’s development: Altogether, then, it appears that each organism has an innate tendency to develop in a symmetrical manner, and that this tendency is controlled and subordinated by the action of external conditions, and not that this symmetry is superinduced on ab externo. In fact, that each organism has its own internal and special laws of growth and development. If, then, it is still necessary to conceive an internal law or “substantial form,” moulding each organic being, and directing its development as a crystal is built up, only in an indefinitely more complex manner, it is congruous to imagine the existence of some internal law accounting at the same time for specific divergence as well as for specific identity. A principle regulating the successive evolution of different organic forms is not one whit more mysterious than is the mysterious power by which a particle of structureless sarcode develops successively into an egg, a grub, a chrysalis, a butterfly, when all the conditions, cosmical, physical, chemical, and vital, are supplied, which are the requisite accompaniments to determine such evolution. (pp. 186–7) [T]he new forms must be produced by changes taking place in organisms in, after or before their birth, either in their embryonic, or toward or in their adult, condition. (p. 233)

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Reminiscent, at least in spirit, of Wright’s (Wright 1932) “shifting balance theory” (represented by the topographic map of differing gene combinations) and Waddington’s (Waddington 1940) “epigenetic landscape,” Mivart (pp. 228–9) frames his theory of developmental reorganization in terms of a rapid transition from one stable state to another: The conception of such internal and latent capabilities is somewhat like that of Mr. Galton…according to which the organic world consists of entities, each of which is, as it were, a spheroid with many facets on its surface, upon one of which it reposes in stable equilibrium. When by the accumulated action of incident forces this equilibrium is disturbed, the spheroid is supposed to turn over until it settles on an adjacent facet once more in a stable equilibrium. The internal tendency of an organism to certain considerable and definite changes would correspond to the facets on the surface of the spheroid.

Equally interesting is how Mivart’s (Genesis, p. 230) language anticipates Bateson’s (Bateson 1894) “undulating theory” or “theory of repeated parts”: [A]s the atoms of a resonant body may be made to give out sound by the juxtaposition of a vibrating tuning–fork, so it is conceivable that the physiological units of a living organism may be so influenced by surrounding conditions (organic and other) that the accumulation of these conditions may upset the previous rhythm of such units, producing modifications in them–a fresh chord in the harmony of nature–a new species!

Mivart (p. 231) then asks: “Are new species now evolving, as they have been from time to time evolved? If so, in what way and by what conceivable means?” To which he responds: [W]e…saw that minerals become modified suddenly and considerably by the action of incident forces… We have thus a certain antecedent probability that if changes are produced in specific manifestation through incident forces, these changes will be sensible and considerable, not minute and infinitesimal. Consequently, it is probable that new species have appeared from time to time with comparative suddenness, and that they still continue so to arise if all the conditions necessary for specific evolution now obtain. (p. 236) [A]n internal law presides over the actions of every part of every individual, and of every organism as a unit, and of the entire organic world as a whole. It is believed that this conception of an internal innate force will ever remain necessary, however much its subordinate processes and actions may become explicable. That from such a force, from time to time, new species are manifested by ordinary generation just as Pavo nigripennis appeared suddenly, these new forms not being monstrosities but harmonious self–consistent values… (p. 239)

Countering Darwin’s argument against saltation in Variation that only a solitary individual would be the bearer of a novelty, Mivart (p. 236) hypothesizes that “as the same causes produce the same

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effects, several individual parent forms must often have been similarly and simultaneously affected.” As for Darwin’s theory of inheritance, Mivart (p. 216) dismisses it with a quote from Delpino: “Thus, in Pangenesis, everything proceeds by force of unknown elements, and we may ask whether it is more logical to prefer a system which assumes a multitude of unknown elements to a system which assumes only a single one?”

Darwin’s rebuttal Given Mivart’s rejection of virtually all of Darwin’s assumptions – with the major exception of allowing that natural selection plays a role in eliminating monstrosities, rapidly eliminates antecedent species, and “favours and develops useful variations, though it is impotent to originate them or to erect the physiological barrier which seems to exist between species” (Genesis, p. 240) – it is not surprising that Darwin harshly criticized the former scientist the following year in the sixth (and, as it turned out, last) edition of the Origin. In this edition of the Origin, published in 1872, the year after Genesis, Darwin doggedly maintained the theme of “natura non facit saltum” and devoted thirty pages specifically to Mivart’s objections to his theory and his objections to Mivart’s. Mr. Mivart believes that species change through an “internal force or tendency,” about which it is not pretended that anything is known. That species have a capacity for change will be admitted by all evolutionists; but there is no need…to invoke any internal force beyond the tendency to ordinary variability, which through the aid of selection by man has given rise by graduated steps to natural races or species. Mr. Mivart is further inclined to believe, and some naturalists agree with him, that new species manifest themselves “with suddenness and by modifications appearing at once.” For instance, he supposes that the differences between the extinct three–toed Hipparion and the horse arose suddenly. He thinks it difficult to believe that the wing of a bird “was developed in any other way than by a comparatively sudden modification of a marked and important kind…This conclusion, which implies great breaks or discontinuity in the series, appears to me improbable in the highest degree. (Origin, 1872, p. 239)

Although Mivart did hypothesize an “internal force” as a generator of sudden change, his model was at least consistent with the observation of the abrupt appearance of novelty in domesticated plants and animals, the discontinuity (i.e. lack of seamless continuity) between apparent species, and the gap–riddled pattern of life history as recorded in the fossil record. In contrast, Darwin attempted to dismiss the relevance of all three of these observations, appealing, as he stated in the quote above, to a sense of probability (or, in this case, improbability). Also as seen in the quote above, Darwin pushed the envelope of credulity, not only by making it seem that the origin of new breeds of domesticated plants and animals is typically by “graduated steps,” but also by adding the claim that this process has led to the emergence of new species of domesticates. In the former assertion, Darwin neglected his own recognition of the reality of niata cattle and other examples of the sudden origin of novelty, while, in the latter, he clearly entered the realm of the imaginary. Darwin’s (Darwin 1872) most relevant objection to Mivart’s theory is found in his questioning the expectation that more than one individual will emerge with the same novelty. Although this

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suggestion is logically consistent with the argument for change via an internal force that causes developmental reorganization, Darwin (p. 240) makes light of it: “Hence in order that a new species should suddenly appear in the manner supposed by Mr. Mivart, it is almost necessary to believe, in opposition to all analogy, that several wonderfully changed individuals appeared simultaneously within the same district.” Of course, “analogy” is the key word here, inasmuch as Darwin promotes the case of gradual change in domesticates through artificial selection. But, upon reflection, Darwin’s counterargument merely reiterates the essence of the variation–natural selection argument that Fleming Jenkin demolished in his review of the first edition of the Origin (see Schwartz 1999): “This difficulty [the sudden appearance of many individuals with the same novelty]…is avoided on the theory of gradual evolution, through the preservation of a large number of individuals, which varied more or less in any favourable direction, and of the destruction of a large number which varied in an opposite direction” (Darwin 1872, p. 240; comment added). Relying on repetition rather than validation, Darwin states outright: “That many species have been evolved in an extremely gradual manner, there can hardly be a doubt” (p. 240) for “when we look to the special parts of allied species, instead of to distinct species,…numerous and wonderfully fine gradations can be traced, connecting together widely different structures” (p. 241). Again, the question must be raised: What is the basis of these assertions? If Darwin can take issue with Mivart’s hypothesis on the grounds that there is no evidence of an “internal force,” are we then expected to take his word for the existence of “numerous and wonderfully fine gradations…connecting together in widely different structures”? Where is the demonstration of the reality of gemmules, of the effects of use and disuse, of blending inheritance, or of the power of natural selection to produce novel structures? Indeed, it is here we can appreciate Delpino’s objection to pangenesis: Why should we invoke a multitude of unknowns when an alternative theory predicts only one mechanism? But Darwin (Darwin 1872, p. 242) continues his attack on Mivart, concluding with an appeal to embryology: It is notorious that the wings of birds and bats, and the legs of horses or other quadrupeds, are undistinguishable at an early embryonic period, and that they become differentiated by insensibly fine steps. Embryological resemblances of all kinds can be accounted for…by the progenitors of our existing species having varied after early youth, and having transmitted their newly acquired characters to their offspring, at a corresponding age. The embryo is thus left almost unaffected, and serves as a record of the past condition of the species. Hence is it that existing species during the early stages of their development so often resemble ancient and extinct forms belonging to the same class. On this view of the meaning of embryological resemblances…it is incredible that an animal should have undergone…momentous and abrupt transformations; and yet should not bear even a trace in its embryonic condition of any sudden modification; every detail in its structure being developed by insensibly fine steps.

On the face of it, this passage would appear to echo von Baer’s (Baer 1828) laws regarding the commonality of embryonic stages among vertebrates until the point at which each kind of animal veers off onto the ontogenetic path that will mold it into the adult of its taxon, replete with its specifically distinctive features. But it is obvious that Darwin’s invocation of ontogeny is actually

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a foil for gradualism, using the smoothly transitional nature of organismal growth and development as supposed evidence of the gradual nature of evolutionary change. Future suggestions aside as to how novelty could become imbedded early on in ontogeny (e.g. de Beer’s, 1930, theory of clandestine evolution), what is interesting about Darwin’s dismissal of such a possibility and his focus on “structure being developed by insensibly fine steps” is that the only avenue he leaves open along which change can occur is by adding stages to the end of an individual’s ontogeny. This, of course, is precisely the primary interplay between ontogeny and phylogeny that Haeckel (e.g. 1866) envisioned when he formulated the Biogenetic Law (“ontogeny recapitulates phylogeny”). Both Darwin and Haeckel envisioned links between adult individuals, with change occurring at the terminal stage of ontogeny.

Wherefore Thomas Henry Huxley? Darwin’s intensified adherence to notions of gradual transformational change from Variation to the last edition of the Origin, and his assault on Mivart in the latter work, is both interesting and curious given the reviews he received of the first edition of the Origin, not only from Fleming Jenkin, but also and especially from his intellectual defender, Thomas Henry Huxley. For, in his review of the Origin, which was published in 1860 (and reprinted thereafter, e.g. in Huxley 1876, Lay Sermons, Addresses, and Reviews, hereafter referred to as Sermons), Huxley was anything but restrained in his criticism of Darwin’s rejection of rapid morphological change. For example, on p. 257 in Sermons, in the reprinted review of the Origin, Huxley writes: We do not speak jestingly in saying that it is Mr. Darwin’s misfortune to know more about the question he has taken up than any man living. But this superabundance of matter must have been embarrassing to a writer who, for the present, can only forward an abstract of his views; and thence it arises, perhaps, that notwithstanding the clearness of the style, those who attempt fairly to digest the book find much of a sort of intellectual pemmican–a mass of facts crushed and pounded into shape, rather than held together by the ordinary medium of an obvious logical bond: due attention will, without doubt, discover this bond, but it is often hard to find. Again, from the sheer want of room, much has to be taken for granted which might readily enough be proved; and hence, while the adept, who can supply the missing links in the evidence from his own knowledge, discovers fresh proof of the singular thoroughness with which all difficulties have been considered and all unjustifiable suppositions avoided, at every reperusal of Mr. Darwin’s pregnant paragraphs, the novice in biology is apt to complain of the frequency of what he fancies is gratuitous assumption.

In apparent heed of his own criticism of Darwin’s unrestrained use of example, Huxley keeps his reference to individual cases to a bare minimum. Nevertheless, he does refer to one of Darwin’s own citations: the abrupt appearance of a “monstrosity” that became the basis of a new breed of sheep, the Ancon or Otter sheep. He does so (p. 265) to demonstrate not only that it “appears to have arisen in full force, and…per saltum,” but also to argue that “[i]t was no case of what is commonly called adaptation to circumstances; but, to use a conveniently erroneous phrase, that

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variations arose spontaneously.” But, while Darwin would say that such demonstrations in domesticated animals are not applicable to wild species, Huxley takes an opposing view: Varieties then arise we know not why; and it is more than probable that the majority of varieties have arisen in this ‘spontaneous’ manner…But however they may have arisen, what especially interests us at present is, to remark that, once in existence, varieties obey the fundamental law of reproduction that like tends to produce like, and their offspring exemplify it by tending to exhibit the same deviation from the parental stock as themselves. (p. 266) If a variation which approaches the nature of a monstrosity can strive…to reproduce itself, it is not wonderful that less aberrant modifications should tend to be preserved even more strongly; and the history of the Ancon sheep is, in this respect, particularly instructive. (p. 267)

Anticipating Mivart’s Genesis, Huxley used Darwin’s argument for rejecting monstrosities – in this case the Ancon sheep – as a reflection of nature in the “wild” as the basis for coming to a diametrically opposed conclusion: monstrosities are biologically, and therefore evolutionarily, instructive. Further like Mivart, Huxley (e.g. see pp. 266–271) rejected natural selection as playing a role in the emergence of novel features. Unlike Mivart, however, Huxley did not speculate on how novelty is produced “spontaneously” – a fact that seems incongruous given Huxley’s (Huxley 1863) emphasis only a few years later on development and the emergence of differences between taxa. Instead, Huxley took the approach of questioning on philosophical grounds the validity of Darwin’s claims. For example, on pp. 294–295 of Sermons, he writes: Inductively, Mr. Darwin endeavours to prove that species arise in a given way. Deductively, he desires to show that, if they arise in that way, the facts of distribution, development, classification, &c., may be accounted for, i.e. may be deduced from their mode of origin, combined with admitted changes in physical geography and climate, during an indefinite period. And this explanation, or coincidence of observed with deduced facts, is, so far as it extends, a verification of the Darwinian view. There is no fault to be found with Mr. Darwin’s method, then; but it is another question whether he has fulfilled all the conditions imposed by that method. Is it satisfactorily proved, in fact, that species may be originated by selection? that there is such a thing as natural selection? that none of the phaenomena exhibited by species are inconsistent with the origin of species in this way? If these questions can be answered in the affirmative, Mr. Darwin’s view steps out of the ranks of hypotheses into those of proved theories; but, so long as the evidence at present adduced falls short of enforcing that affirmation, so long, to our minds, must the new doctrine be content to remain among the former–an extremely valuable, and in the highest degree probable, doctrine, indeed the only extant hypothesis which is worth anything in a scientific point of view; but still a hypothesis, and not yet the theory of species. After much consideration, and with assuredly no bias against Mr. Darwin’s views, it is our clear conviction that, as the evidence stands, it is not absolutely proven that a group of animals, having all the characters exhibited by species in Nature, has ever been originated by selection, whether artificial or natural.

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And on pp. 297–8, …Mr. Darwin’s position might, we think, have been even stronger than it is if he had not embarrassed himself with the aphorism, “Natura non facit saltum,” which turns up so often in his pages. We believe, as we have said above, that Nature does make jumps now and then, and a recognition of the fact is of no small importance in disposing of many minor objections to the doctrine of transmutation. …Our object has been attained if we have given an intelligible, however, brief, account of the established facts connected with species, and of the relation of the explanation of those facts offered by Mr. Darwin to the theoretical views held by his predecessors and his contemporaries, and, above all, to the requirements of scientific logic. We have ventured to point out that it does not, as yet, satisfy all those requirements; but we do not hesitate to assert that it is as superior to any preceding or contemporary hypothesis, in the extent of observational and experimental basis on which it rests, in its rigorously scientific method, and in its power of explaining biological phaenomena, as was the hypothesis of Copernicus to the speculations of Ptolemy. But the planetary orbits turned out to be not quite circular after all, and, grand as was the service Copernicus rendered to science, Kepler and Newton had to come after him. What if the orbit of Darwinism should be a little too circular? What if species should offer residual phaenomena, here and there, not explicable by natural selection?

But others did come after Darwin. Even though Darwin was a well–known naturalist prior to 1859, if we use the publication of On The Origin of Species as the public emergence of Darwin into the realm of evolutionary theory, we must recognize Huxley and Mivart as subsequent major critics of the credos of natural selection and gradual transformation. Curiously, though, it is Mivart who Darwin publicly attacked, and who was arguably a primary provocation for the 1872 revision of the Origin. Of no less import than Darwin’s determined adherence to gradual transformation in this latter work was his clinging to the theory of pangenesis. Indeed, as he clearly appears to have dug in his heels on gradualism and a rejection of saltationism and the importance of “monstrosities” for understanding evolutionary change in reaction to Mivart, he seems to have done the same with regard to pangenesis and his cousin Galton’s (Galton 1871) experiments that failed to support it. By the late nineteenth century, with no less energy than Darwin brought to bear on his theories, Bateson (Bateson 1894) and de Vries (de Vries 1889) were arguing with force and conviction for the sudden origin of novelty as well as for decoupling the origin of novelty from selectionist scenarios, and relegating the role, if any, of natural selection to the survival of species. Although inspired by Darwin’s theory of pangenesis, de Vries’ theory of intracellular pangenesis was actually a rejection of the former notions of inheritance. And, certainly, in 1903, the capstone year of this workshop, Morgan, a trained embryologist, was as vocal as any scientist in rejecting the hyperbole and circularity of Darwinism as being of any evolutionary import. Why, then, did the questioning of Darwinian explanations of smoothly transformational change become submerged until the recent advent of “evo–devo” thinking? Ironically, it was through the work of Morgan, who in just over a decade went from lambasting Darwinism and Mendelism in 1903 as metaphysical flights of intellectual fancy to melding the two disciplines into

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the population–genetics thinking that informed the evolutionary synthesis (see review in Schwartz 1999).

Conclusion Although Darwin is often lauded for embracing embryology and development in his theory of gradual change via natural selection, his perspective was clearly at odds with Mivart’s. It may be true that both scholars envisioned a source external to the individual as playing some role in the emergence of organismal change. But it is equally obvious that only at this vague level might we seek a favorable comparison between these two scientists. Otherwise, for Darwin, biologically real novelty lies only in the minutiae of individual difference, which, in turn, derives from any number of sources that, often through use or disuse, leave their marks on an individual. The notion that use–disuse can engender change by causing elements of a postnatal individual to become altered, and that this effect can then, via gemmules, be passed on to future generations, fits the envisioned role of natural selection: Both concepts externalize the ultimate source of organismal modification. Blending inheritance is, therefore, the only aspect of Darwin’s theory of pangenesis that might conceivably be regarded as representing an internal component of an individual’s biology. With the substitution of Mendelism and population genetics for blending inheritance and gemmules in early formulations of neo–Darwinism (Morgan, e.g. 1916), Darwin’s theory of evolution by means of natural selection seemed to be unassailable (e.g. Simpson 1962). Although use–disuse arguments were supposedly purged from Darwinism (by way of singling out Lamarck as the lone advocate of such lunacy; see Burkhardt 1977), it is obvious from the language of Darwinian explanation still in vogue that, in essence, they were not (e.g. see examples in Schwartz 2004, 2005). Even though Darwinism today claims a basis in genetics, the emphasis is not only on the incorrect notion of there being “genes for things” – similar to the idea that selection chooses features to serve a purpose – but also and contradictorily on the similarly biologically unreal notion that selection can direct the course of genetic change by selecting behavioral or morphological traits that anticipate their benefit to an individual. One might thus characterize present–day Darwinism as the “vacuum theory of evolution” (Schwartz 2005, in press). In contrast, Mivart’s emphasis on internal reorganization affecting an organism’s development represents an entirely different biological perspective. Although hypothesizing an “external force” as the initiator of a process of change, whatever course organismal change takes is rapid and random with regard to the circumstances in which the altered organism finds itself. Most importantly, Mivart seats organismal change in the context of an internal restructuring of developmental processes. At one point in time, the developmental organization of an organism is in equilibrium, as is the spheroid lying on one of its facets. In order for change to occur, this equilibrium must be disrupted. Ultimately, the spheroid will come to rest in equilibrium on another facet; that is, developmentally, the organism will be in equilibrium in a different or novel state of organismal organization. If the resultant novelty is ill suited to its bearer’s circumstance, the individual will most likely not survive. Even if one chooses to equate the “elimination” of individuals with “natural selection,” this process or phenomenon is involved neither in the

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production of novelty, nor in the differential selection of individuals that are either more fit than others or supposedly fulfilling a particular adaptive trajectory. With the exception of incorrectly predicting that more than one individual will emerge with the same novelty because they will respond similarly to the same provocation, Mivart’s focus on development is unexpectedly compatible with the emphasis in evolutionary developmental genetic theory (“evo–devo”) on novelty resulting via the differential recruitment of regulatory molecules in different signal transduction pathways (e.g. Carroll et al, 2005; Gilbert and Bolker 2001; Raff 1996; Maresca and Schwartz, n.d.; Schwartz 1999). That is, in contrast to the Darwinian population genetics model of continually changing “genes” or “genomes” underlying the emergence of minute variations of a phenotypic trait, modern cell and molecular biology have demonstrated that cell and DNA stability or homeostasis is the rule. The potential for change thus occurs when this “equilibrium” (to use Mivart’s term) is disrupted and new pathways of molecular communication become available. As Gilbert and Bolker (Bolker 2001, p. 451) put it: Embryologists now recognize receptors and signal transducing molecules as components of the competence apparatus that enable certain cells to respond to specific inducers. These signaling pathways are the bases of embryonic induction, which is in turn the core of organogenesis. If macroevolution involves changing morphological features, then the alteration of signal transduction pathways becomes critical for any discussion of large scale evolution.

In a very basic way, then, it might not be inappropriate to delineate the beginning of a “Darwinism”–“evo–devo” debate in the late nineteenth century, between one of Victorian England’s leading comparative anatomists, St. George Mivart, and Darwin himself.

Postscript Given Darwin’s seemingly career–long entrenchment in gradualism and rejection of saltationsim, it is with some surprise to read entry 130 in his Red Notebook, which according to Herbert (Herbert 1980), was written sometime during March, 1837: The same kind of relation that common ostrich bears to (Petisse. {lesser or Darwin’s rhea} & diff kinds of Fourmillier {antbird}): extinct Guanaco {llama} to recent: in former case position, in latter time. (or changes consequent on lapse) being the relation. – As in first cases distinct species inosculate, so must we believe ancient ones: [(] not gradual change or degeneration, from circumstances: if one species does change into another it must be per saltum – or species may perish. = This representation of species important, each its own limit & represented. – Chiloe creeper {thorn–tailed Rayadito}: Furnarius {ovenbird}. Calandria; inosculation alone shows not gradation. {comments added}

Reading this passage and then those as well as other notebooks that followed is a frustrating experience inasmuch as there is no obvious reason for Darwin to have abandoned saltational ideas as completely as he did. Indeed, as the quotes above from Variation make clear, Darwin had before him the basis of a saltational theory that was even supported by evidence of heredity: not only could “monstrosities” interbreed successfully, they could also reproduce with “normal,” parental– type individuals, and thereby perpetuate their novelties. Observations to the contrary, Darwin’s

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constant assertions of a seamless web having existed among living species as well as between descendents and their extinct ancestors might betray a non–biological concern. Namely, were he to embrace saltation, the door would remain open for his religious contemporaries to invoke special creation to explain the abrupt appearance of species in the fossil record as well as the discontinuities between extant taxa. It is, therefore, perhaps a bit ironic that an intellectual enterprise – neo–Darwinism – that went and continues to go out of its way to denegrate and discredit alternative, saltation–like theories for the origin of novelty was built on such an imaginary foundation.

Jeffrey H. Schwartz, University of Pittsburgh, [email protected]

Acknowledgements I thank James Lennox for guiding me to the entry in the Red Notebook.

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References Baer, K. E. von 1828. Über Entwicklungsgeschicte der Thiere: Beobachtung und Reflexion. Königsberg: Bornträger. Bateson, W. 1894. Materials for the Study of Variation, Treated with Especial Regard to Discontinuity in the Origin of Species. New York: Macmillan. Burkardt, R. W., Jr. 1977. The Spirit of System: Lamarck and Evolutionary Biology. Cambridge, MA: Harvard University Press. Carroll, S. B., Grenier, J. K., and Weatherbee, S. D. 2005. From DNA to Diversity: Molecular Genetics and the Evolution of Animal Design (2nd ed.). Malden, MA: Blackwell Publishing. Darwin, C. 1859. On the Origin of Species by Means of Natural Selection or the Preservation of Favoured Races in the Struggle for Life. London: John Murray. ———. 1868. The Variation of Animals and Plants under Domestication (2 vols). London: John Murray. ———. 1871. The Descent of Man and Selection in Relation to Sex. London: John Murray. de Beer, G. R. 1930. Embryology and Evolution. Oxford: Oxford at the Clarendon Press. de Vries, H. 1889. Intracellular Pangenesis. Translated from the German by C. S. Gager (Open Court, Chicago, 1910). Galton, F. 1871. Experiments in pangenesis, by breeding from rabbits of a pure variety, into whose circulation blood taken from other varieties had previously been largely transfused. Proc. Roy. Soc. (Biol.), 19: 393–404. Gilbert, S. F. and Bolker, J. A. 2001. Homologies of process and modular elements of embryonic construction. In The Character Concept in Evolutionary Biology (G. Wagner, ed.). New York: Academic Press, pp. 435–454. Haeckel, E. 1866. Generelle Morphologie der Organismen: Allgemeine Grundzüge der organischen Formen– Wissenschaft, mechanisch begründet druch die von Charles Darwin reformirte Descendenz–Theorie (2 vols). Berlin: Georg Reimer. Herbert, S. (ed.) 1980. The Red Notebook of Charles Darwin. Bull. Brit. Mus. (Nat. Hist.), hist. ser., 7: 1–164. Huxley, T. H. 1860. Review of “The Origin of Species.” The Westminster Review (reprinted in Huxley, 1876). ———. 1863. Man's Place in Nature. New York: D. Appleton. ———. 1876. Lay Sermons, Address, and Reviews. New York: D. Appleton. Maresca, B. and Schwartz, J. H. n.d. Environmental change and stress protein concentration as a source of morphological novelty: sudden origins, a general theory on a mechanism of evolution. Mivart, St. G. 1871. On the Genesis of Species. London: John Murray. Morgan, T. H. 1903. Evolution and Adaptation. New York: Macmillan. ———. 1916. A Critique of the Theory of Evolution. Princeton: University of Princeton Press. Raff, R. A. 1996. The Shape of Life: Genes, Development, and the Evolution of Animal Form. Chicago: The University of Chicago Press. Schwartz, J. H. 1999. Sudden Origins: Fossils, Genes, and the Emergence of Species. New York: John Wiley & Sons. ———. 2004. Trying to make chimpanzees into humans. Hist. Phil. Life Sci., 26: 271–277. ———. 2005. The Red Ape: Orangutans and Human Origins (revised). Boulder, CO: Westview Press. ———. in press. Molecular systematics and evolution. In Encyclopedia of Molecular Cell Biology and Molecular Medicine (EMCBMM), (R. A. Meyer, ed.). Weinheim: Wiley–VCH Verlag. Simpson, G. G. 1962. Preface to The Origin of Species (6th ed.). New York: Macmillan (reprinted 1972). Waddington. C. H. 1940. Organisers and Genes. Cambridge: Cambridge at the University Press. Wright, S. 1932. The roles of mutation, inbreeding, crossbreeding and selection in evolution. Proc. Sixth Internatl Congr. Genetics, 1: 356–366.

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Message in a Bottle: The Business of Vaccines and the Nature of Heredity after 1880 Andrew Mendelsohn

Introduction Before the 1880s there was no consensus on the nature of microorganisms. Some students of bacteria saw constant species. Others saw a flux of form and function. Each saw in the sense of both literally seeing through the microscope and perceiving in relation to rival biological theories or classifications. Differences in view were thus at once descriptive and theoretical. Violent debates raged. Research projects arose to great influence and just as quickly came crashing down. It is usually said that the transformists/“unitarians” lost (the botanist Carl Wilhelm Nägeli, the surgeon Theodor Billroth, many lesser–knowns) and the “Linnaeans” won (the botanist Ferdinand Cohn, the physician Robert Koch, their followers) – and that this made possible a science of bacteriology at all, relieving biology of the notion that, in Nägeli’s oft–ridiculed formula, all microscopic life was a single species in continual flux of form and function, producing sometimes souring of milk, sometimes butyric fermentation in sauerkraut, sometimes the aging of wines, sometimes cholera, and so on. That is the standard story. 1 Yet in fact during the 1880s, as I have argued recently, far from one view winning out over the other, a consensus was rapidly achieved in which neither of these views prevailed. Instead, bacteria were seen to vary within species.2 How could this have happened? How could an international and notably French–German consensus on a supremely controversial and uncertain topic – bacterial species – have been so rapidly forged? This historical problem generates the following essay. My answer is, in a word, vaccines. I shall argue in the first part of this paper that an unusually powerful model of species and variation – a model in the sense of both exemplary case and research object – was inadvertently provided by early vaccines, notably the anthrax vaccine invented by Pasteur in 1880–81 and then mass–produced and distributed, manipulated in laboratories as well as used on farms around the world.3 In this process, as I shall argue in the second part of the paper, heredity (at least in the realm of microscopic life) came to be located within and redefined by an enterprise of control and testing, production to an exact standard and reliable distribution. This enterprise was akin to yet differed in revealing ways from animal and plant breeding, which in this period became all–important to the making of a biological science of heredity. The resultant new meaning of heredity suggests historical change related but not equivalent to the well–known shift from “soft” to “hard” heredity, or from heredity as force to heredity as structure.4 1

2 3

For “unitarians” versus “Linnaeans,” see Mazumdar 1995, which revises the standard story by showing that the two traditions long continued their conflict but in a series of other research domains, such as immunology and serology. Mendelsohn 2002. Comments welcome as to whether the history of science offers earlier examples of a standardized, mass– produced and distributed research object, as distinct from instruments. The answer would be a definite yes if (some?) chemical reagents count as research objects rather than instruments.

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I. Manufacturing Variation In a series of papers in 1880 and 1881, Louis Pasteur reported that he had produced weakened or “attenuated” cultures of the microorganisms of fowl cholera and anthrax, which conferred immunity to these diseases when inoculated in animals. He called these cultures of attenuated virulence “vaccins.” This work is celebrated as the beginning of immunology and artificial immunization. Yet it also had other profound if less remembered implications. Bacteriological workers in many countries began manipulating the virulence of their cultures and soon enough other microbial properties such as morphology, pigment formation, colony form, capacity to liquify gelatine, and sporification, by altering the recipe of the culture medium or the time–span between resowings in fresh medium, or the temperature, or the exposure to air, and so on. They interpreted many of the observed changes in form and function as biological variation. Thus much early medical bacteriology became a science of what the preeminent French veterinary scientist Auguste Chauveau called “experimental variation.” 5 Entire chapters of the standard German and French bacteriological handbooks were soon devoted to variation or “variability.”6 This is remarkable on two counts. First, the received view has been that a dogma of species constancy made bacterial variation unthinkable or at best heretical until the early twentieth century. Second, biology at large could boast nothing comparable. In the early 1880s, when Wilhelm Roux and other academic scientists had barely begun to preach a new biology as experimental laboratory science; when questions of species, variation and inheritance were still being pursued largely through observation in field and museum; when Mendel would not be rediscovered and classical genetics begin for another 20 years, dozens of physicians more or less remote from academic biology and working in mostly medical and public health laboratories around the world, were building a system of sustained cellular–level, in vitro experimental research on what they and their contemporaries saw as biological variation and indeed mechanisms of evolution.7 What was so special about virulence work? How did it allow observers to see change in bacterial properties as variation within species, rather than contamination or “transformism”? There are two answers to this question. Variable virulence modelled variation within species because it was observed to correlate with variable severity of what remained clinically distinct, or specific, diseases: bacteria that were attenuated or augmented in their virulence and morphologically sometimes less than uniform could still be used to provoke a predictable set of clinical effects in animals, varying only in their severity.8 And secondly, fully attenuated, no longer pathogenic organisms did not have to be seen as transformed into different species because they exhibited their original species identity by acting as specific vaccines when injected into animals. 9 (Total loss of pathogenicity was no minor species issue: recall that these organisms were often even named after their associated diseases, as in Bacillus anthracis.) Thus the identity and stability of

4 5 6 7 8 9

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See Gayon 1995. Chauveau 1889b, 789. The most important of these was Kruse 1896b. See Mendelsohn 2002, 18–26. Pasteur 1880c, 324–26. Chauveau 1889a–b.

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biological species was guaranteed not by a botanical classification system and rules and skills of biological observation, but by clinical and vaccinal practices and effects. Microbial variation existed as a biological phenomenon not despite, but because of its medical, even clinical identity. This is not meant as a subtle point. Scientific knowledge took shape through a practical matrix in a way akin to the role of the steam engine in the origins of thermodynamics, or telegraphy in the rise of electromagnetic field theory, a comparison on which I shall elaborate below. All this was accomplished, inadvertently, more by the distribution of vaccines than by the distribution of journal articles reporting methods and results, of instruments and visual representations, or even of skill and tacit knowledge through circulation of laboratory personnel.10 Pasteur’s celebrated announcements and publications of 1880–81 certainly did not convince his rival founder of bacteriology Robert Koch or lead to experimental confirmation. Koch and his disciples were unable to achieve attenuation in their own laboratory: they reported culturing anthrax bacilli on gelatine for fifty generations and on a potato surface for nine months, through over one hundred generations, without observing any alteration in virulence. They charged that so–called “attenuated” cultures were in fact merely contaminated with common non–pathogenic organisms.11 Evidently, experimenting on virulence was neither practically nor conceptually coherent and plausible enough to be replicated, much less to serve as a model case for variation in other bacterial characteristics. On the theoretical side, there was no agreed–upon concept for the mechanism of hereditary change. Bacteriological researchers were soon able to use a wide range of often incompatible concepts: variation, progressive modification, acclimatization, race formation, adaptation, degeneration, transformation, and not least, selection in the Darwinian sense. This smorgasbord of biological concepts and theory could not provide the coherence of the phenomenon. Moreover, “variation” was itself a heterogeneous category at the time. Biologists hotly debated the origin and nature of variations, as well as the kinds of variation (continuous versus discontinuous). Bacteriologists drew on a concept that was itself ambiguous. Thus it was not the case that the virulence and vaccine model allowed bacteriologists to assimilate their phenomena to some settled standard ‘biology’ of their time. On the practical side, there was no standard unit of measure, no “meter” or “ohm” of virulence. 12 Nor was there a uniform method of attenuation: all sorts of things were done to bacterial cultures and over a range of times. The anthrax vaccine, on the other hand, was uniform. You could buy a bottle on the market – “käuflich im Handel” as Koch put it13 – and test it on animals, or try to alter it by passing it through animals in series, or use it as the known starting point for an experimental manipulation in culture. Following the spectacular success of the public trial at Pouilly Le Fort in 1881, requests for vaccine poured into Pasteur’s laboratory.14 His associate Charles Chamberland began large– scale production. Sales and distribution were handled by a commercial agent, Boutroux, in Paris. Tens of thousands of sheep and cattle were vaccinated in France in 1882 alone. Within 10 years 3.3 million French sheep and 438,000 cattle had been vaccinated. In Italy, the government provided 10 11 12 13 14

Cf. Collins 1985; Latour 1987. Koch 1881, 200; Gaffky 1881, 121–126; Loeffler 1881, 134–141. See Kruse 1896a, 299, for one attempt to define a scale of six “Virulenzstufen” by delimiting sets of microscopic pathological changes in animals. Koch, Gaffky and Loeffler 1884, 247. See Pasteur’s published correspondence for July 1881; Valery–Radot, 3:220.

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Pasteur’s vaccines free of charge. Production laboratories were established in places near and far – Austria, Spain, South America, Russia, Australia.15 (See figure 1.)

Fig. 1. Loir, Adrien, Pasteur's Vaccine of Anthrax in Australia: as a preventative against Cumberland Disease in sheep, cattle and horses, published c.1891, Sydney, Back Cover Advertisement. Boutroux distributed the bottles to farmers, but also to scientists. (Researchers also obtained from Boutroux the Pastorian vaccines for fowl cholera and swine erysipelas, though these played a far lesser role in the story told here.) Reference to Boutroux crops up everywhere in the relevant bacteriological, medical, and veterinary journal literature of the 1880s. For the scientists, Boutroux’s agency and all the work by Chamberland and his assistants that went into those bottles – and of course the procedures and skills detailed in instruction manuals and diagrams (see figure 2.) – made the bottles’ contents a model arguably unprecedented in its distribution, stability, and unequivocality as witnessed by the dramatic and repeatable set of effects produced by following the instructions and inoculating from the bottles into animals.

15

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M’Fadyean 1894, 327.

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Fig. 2. Only after attenuated cultures began to be distributed and used as vaccines did phenomena of attenuation become ‘real’ and powerful enough to be recognized and studied by Koch and his associates, rather than dismissed as mere contaminations. Before distribution of the vaccine, even a less sceptical, English bacteriologist who did manage to achieve sporeless anthrax cultures harmless even to mice nonetheless could not confirm that this attenuation was “transmitted” to the next generation.16 As late as 1888, researchers in the laboratory of Koch’s colleague Carl Flügge at Breslau, working on the nature of the attenuation process, were unable to attenuate swine erysipelas bacilli and obtained them instead as vaccines from Boutroux. 17 It is remarkable that Koch and his associates first failed to replicate Pasteur’s results and then succeeded after the vaccine was publicly tested at Pouilly Le Fort and hit the international medical marketplace. Koch and his disciples obtained vaccines from Boutroux in Paris, an especially well thermostatically controlled incubator from the firm of Wiesnegg also in Paris, which could maintain a temperature 16 17

Klein 1883, 9, 64 (paper based on author’s 1881 report to Local Government Board). Smirnow 1888, 244.

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“without the slightest oscillations” for weeks, and proceeded to attenuate fully one of their own anthrax cultures and inoculate it into sheep. Now Koch saw and represented a biological phenomenon of hereditary change. He affirmed that “the high scientific importance” of Pasteur’s discovery of attenuation lay in the fact “that the new properties are also preserved in the progeny [Nachkommen] of the attenuated bacilli.” This was “of the highest interest not only for etiological research, but equally for biological science.”18 Hans Buchner, student of the transformist Nägeli, glimpsed why Pasteur’s experiments had become, even for Koch, real phenomena of hereditary modification of microorganisms, whereas Buchner’s own similar earlier results had been ignored or rejected. The reason was “that Pasteur’s protective vaccinations, which are now being practiced – even if at first only in trial form – in nearly all European countries, made it impossible even for Koch to continue to doubt the existence of an attenuated anthrax bacterium.” 19 Pasteur’s vaccine business distributed not only prophylaxis against disease, but also inadvertently this message in a bottle. Though to the eye nothing but clear liquid in a bottle, the mass–manufactured vaccine culture served as a model in several senses: (1) as an object for demonstration, available virtually off the shelf as needed to show variation within bacterial species; (2) as an exemplary case rendering variation of other microbial characteristics plausible and investigable without casting doubt on the rigor of one’s methods or the existence of bacterial species; (3) as a live research object to be worked upon in vitro or in vivo, whose uniformity, purity, known production method and initial properties made it manipulable in predictable yet exploratory ways. The bottles were not, of course, at sea. They were part of wider human organization. There is an illuminating analogy to the physical sciences. Like the early physical standards laboratories of the same period, vaccine laboratories helped make a world in which local science could become global. They distributed a standardised material that could be tested and manipulated – with predictable outcomes – in other laboratories as well as used on farms. And with this material, a set of techniques and gestures (see figure 2). In this sense, the Pastorians’ vaccine laboratory and their commercial agent and distributor, Boutroux, became like the Cavendish Laboratory at Cambridge, which James Clerk Maxwell did not wish to become a “manufactory of ohms” (the unit of electrical resistance) but which in effect did.20 Yet unlike the calibration and distribution of electrical measurement devices, the manufacture and distribution of vaccines fundamentally recast the nature of the phenomenon. It changed what sort of descriptive and theoretical statements could plausibly be made. Nature had to be rewritten: the harmless “hay bacteria” species which Buchner had so recently transformed out of anthrax bacteria were now – with no admission of error or retraction of previous claims – “my attenuated anthrax bacteria.” 21 Microscopical biology’s mountainous intellectual terrain of theoretical divides and commitments and antagonisms was ironed flat. The explanandum in most history and sociology inspired by Bruno Latour is: How and why does science work? By contrast, my explanandum here is the changing specific content of a science. The successful anthrax vaccine, one of Latour’s main explananda in his studies on Pasteur, is my explanans. In this sense, the distribution, use and monitoring of vaccines, the network of vaccine 18 19 20 21

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Koch 1882, 216–18, 227–28; see also Koch, Gaffky and Loeffler 1884, 233. Buchner 1883, 411. Schaffer 1992; Latour 1987. Buchner 1880; Buchner 1882, 253.

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producers and consumers, functioned more as telegraphy in relation to the rise of electromagnetic field theory than the Cavendish in relation to the ohm. Historian of physics Bruce Hunt has shown field theory to have arisen through telegraphy as it was made to work and put to work in British projects of empire and trade.22 A new biology of microscopic life, I suggest, emerged through vaccines as these were developed and produced in the 1880s and ’90s for animal industry, hygiene, and colonialism. Animal vaccines’ potential relevance to the “success of colonization” was apparent as early as 1881, when the French Association for the Advancement of Science met in Algiers and thus, in the words of its president, followed in the footsteps of “French arms” to “take possession, in its turn, of this land Africa,” a continent which science and the oeuvre civilisatrice were rendering nothing less than “France, extended across the sea.” Somewhat less predictably, the president continued his address with the question: “What is a virus?” Auguste Chauveau, director of the prestigious National Veterinary School at Lyon and the most important researcher on attenuation outside of Pasteur’s group, launched into a detailed account of Pasteur’s discoveries (and those of his own research school) culminating in “the permanent and transmissible attenuation” of pathogenic microorganisms.23 These spheres of activity, by the same token, provided bacteriology with conceptual tools. From colonialism and its zoological and anthropological sciences, bacteriologists took the concepts of acclimatization and race; from medicine, those of “degenerate” and “abnormal”; from agronomy, as we shall see below, the breeder’s concept (and aesthetic–economic ideal) of “fixed” varieties. These terms were transfered from man (and animals) to bacteria, which themselves were already seen as growing in “cultures” and “colonies.”24 At a joint meeting of the five French academies, Henri Bouley explained that in Pasteur’s vaccine cultures, attenuation became “a race character” and thus these microbes are “races degenerated from their original power and become beneficial by their very weakness.”25 Evoking contemporary anthropological stereotype, Emil von Behring, in the year he received the first Nobel Prize in medicine, argued that changes of culture medium and environment yielded “statistically almost more abnormal than normal diphtheria bacilli” and dismissed the idea of conformity to a “normal type” as the equivalent of expecting the human race to conform to the “Apollo Belvedere.”26 Agronomy, medicine, and colonialism were not only the matrix, but became also the content of bacteriology. Using the language of acclimatization, race, degeneracy, and abnormality, bacterial change could be described in terms that left species untouched; all these terms came to be ordered under variation. 27 Neither a celebrated experiment, nor a type–specimen located in one laboratory or museum, nor the application of any biological theory or classification system, but rather an increasingly global scientific and commercial matrix of practice and meaning, of vaccine bottles and bodily 22

23 24 25

26 27

Hunt 1991. With the difference that empire and telegraphy created what Hunt called a “market” for the reception of field theory already created by Faraday, whereas I am arguing that the new biological content of bacteriology and the nature of its phenomena was created through vaccine–making and the vaccine enterprise. Chauveau 1881, 482–83, 490. A good early example of frequent use of the term bacterial “colonies” is Eberth 1872. Bouley 1882, 547; see also Chauveau 1885, 355. The language of race continued to be used routinely in technical publications in the twentieth century; e.g., Pfeiffer 1903, 37: “Rassen des Choleraerregers,” “Cholerarassen.” Behring 1901, 81–82. See Mendelsohn 2002, 18–24.

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effects, sustained the phenomenon of bacterial hereditary change or “variation” and consensus about it. Without that matrix, variation within bacterial species would not have been merely a neglected topic or heterodox viewpoint. It would not have – indeed, had not – existed as an object of inquiry. Students of bacteria had not seen or conceived of such a thing: they had seen either immutable species or protean mutability. Likewise, later students of bacteria, working within a different matrix, would stop seeing it. In the early twentieth century, the unifying vaccine model was displaced by the rise of routine bacteriological diagnosis for endemic diseases and the application to bacteria of the mutation theory of Hugo de Vries.28 Bacterial “variation” began to fragment into myriad phenomena: mutation and dissociation, cyclogeny, phage effects, and transformation of type, and eventually enzyme adaptation and transduction. 29

II. Fixing Heredity In the second part of the paper, I turn from hereditary change to heredity itself. In modeling variation, vaccines modeled hereditary change and thus phenomena of inheritance, at least implicitly. Implicitly, because bacteriologists rarely contributed to the explosion of scientific writing on heredity that characterised the last decades of the nineteenth century. In the absence of such writings, what I offer here is, instead, the intellectual history of practices. The historian can interpret what the hereditary meant and how it changed by looking at practice and its rationales: not how things were done (as in much of the past twenty years’ work on scientific practice, material culture, tacit knowledge, skill, training), but which things were done and why. Why exactly did bacteriologists regard attenuation and return to virulence as hereditary changes rather than merely physiological adaptations? – as in Claude Bernard’s experiments on the physiological adaptability of animals to lack of oxygen under a belljar, which Pasteur liked to cite.30 The simple answer seems to be time – the multiple generations through which microorganisms could be observed to pass and over which a given degree of virulence or attenuation could be found to persist. At first, in Pasteur’s earliest attenuation work on fowl cholera, and among other researchers circa 1880–82, persistence from one cultivation or “generation” to the next was indeed the criterion for inheritedness; it showed that “the bacillus having become modified by time, transmits to its offspring this acquired mitigation.” 31 As late as 1882, reporting his own experiments, Koch gave no other criterion for inheritance than the regular, stepwise (Stufen) character of the attenuation process over the passage of time and thus bacterial generations.32 Soon enough, however, time ceased to be an adequate criterion. Increasingly, modification was more consistently and confidently judged hereditary if it persisted under changed conditions. Why? In 1884, for example, in their major research paper on anthrax vaccination and immunity, Koch and his associates now reported on anthrax cultures attenuated in his laboratory 28 29 30 31

32

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This theory–practice relationship has been explored by Olga Amsterdamska 1987. For an overview, see Brock 1990. Pasteur 1876, 241; Duclaux 1920, pt. 6. Klein 1883, 65 (based on his government report of 1881). See Pasteur 1880d. Modification and persistence over many “generations” was also the criterion for those claiming species transformation; see Buchner 1880. Koch 1882, 217. See also Jahresbericht 1882, 1883, p. 13; Jahresbericht 1883, 1884, p. 15.

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and then cultivated at room temperature for two years, during which time they never regained their virulence. From this he concluded that the attenuation was “passed on from generation to generation [von Generation zu Generation weiter vererbt].”33 Without much ado, Koch made persistence under changed conditions the test for – and thus in effect the nature of – heredity. Others cited him in agreement.34 Now, persistence under changed conditions was in fact a paradoxical criterion: if changing the conditions created the hereditary variety in the first place, changing the conditions again or restoring them, would not test inheritance, since it ought simply to yield a new hereditary variety (or the original); this was in fact something Pasteur had explicitly shown he was able to do, namely return attenuated anthrax or fowl cholera microorganisms to virulence by passaging them through animals, beginning with the weakest possible ones and moving in series to more and more robust creatures.35 But the objection was not made. And no paradox was perceived. Why not? The answer in a word, again: vaccines. What does make sense of this criterion of heredity is to suppose that it was modeled on – or indeed synonymous with – vaccine safety and efficacy.36 Thus the question: Are modifications of virulence hereditary? was in fact increasingly being asked as: Can we produce safe and effective vaccines? Will there be accidents? Might vaccines, injected into animal bodies, regain their virulence and thus become a danger? Sure enough, the criterion of persistence under changed conditions (such as when cultured as usual or when injected into an animal) first appears in Pasteur’s publications at precisely the point at which he asks whether the attenuated fowl cholera microbe could be “a true vaccine, comparable to the cowpox vaccine.” And the criterion was formulated at first as a test not for inheritedness, but simply for useability as a vaccine.37 In practical vaccine research contexts, this criterion was emphasized and, as we saw with Koch and his associates, became the criterion for inheritedness. 38 By the same token, in theoretical contexts, in research projects designed to “explain” the process of attenuation rather than develop vaccines, the criterion of persistence under changed or usual conditions was not emphasized.39 Despite the confidence of Pasteur’s various pronouncements, these life–and–death questions could not be resolved quickly in the laboratory or by public vaccine trials such as the celebrated demonstration at Pouilly Le Fort. They remained the questions of the day in the veterinary societies, journals and government bureaux.40 And these became criteria for judging the merits of various methods of vaccine–making, a hotly contested arena.41 Koch and others reported that 33 34 35 36

37 38 39 40 41

Koch, Gaffky and Loeffler 1884, 236. For example, Beumer 1887, 1–2. Pasteur 1881, 336. The terms vaccine “safety” and “efficacy” are taken from M’Fadyean 1894, 331; for another example, see Flügge 1888, 209: “Gefahrlosigkeit und Sicherheit.” An alternative reason for changing the criteria would be to meet the objection that persistence of a given degree of attenuation could indicate merely that each new generation of organisms was undergoing the same purely physiological change in response to conditions, rather than inheriting the degree of attenuation. But I have not found this argument in the sources. Pasteur 1880a, 299–300. Interestingly, at this early stage, Pasteur refers not to safety but to the “fear,” once had by Jenner concerning cowpox, that one would have to return always to the original preparation. See, for example, Beumer 1887, 1–2. Flügge 1888, 208–15. Börner 1882, 698; Rózsahegyi 1882, 27; Koch, Gaffky and Loeffler 1884, 261. Geison 1995, chap. 6.

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cultures attenuated using Pasteur’s young and hapless rival Toussaint’s method of heating at relatively high temperatures (47 or 50 or 53 degrees celsius) tended eventually to return to virulence. Cultures attenuated using the Pastorians’ method of heating at only 42–43 degrees celsius over longer periods kept (“bewahren”) their attenuated state even “in later generations” and in the spores. The lower the temperature, “the more securely [sicherer] the physiological varieties seem to keep their [new] properties.”42 “More securely,” sicherer: at work here was the traditional conception of heredity as a force, as varying in strength, as having degrees. 43 I shall return to this point below and in the conclusion. These were issues of vaccine production. There were also issues of vaccine distribution. Pasteur’s production and distribution chief, Chamberland, outlined these too in a chapter on “variation in the virulence of the anthrax vaccines.” Might vaccines become too weak and thus be ineffective and unsafe? The biological stability of varieties was synonymous with successful “conservation.”44 The “theory of virulence” one former Koch student presented, years later, to the master on his sixtieth birthday was equally a method of “conserving virulence in vitro” rather than by costly and time–consuming continual animal passage.45 The terminology of conserving and conservation was at once the language of the technical and social entreprise of storage and distribution and a language for describing the bacterial cultures themselves: “they will conserve their own virulence.”46 Chamberland aimed at a system in which bottles of vaccine could be “expedited throughout the world, as far as the farthest countries, retaining [gardant] their preservative properties,” a network of global biological transport. He reported that the Pasteur team studied the conditions “of conservation” and the causes of vaccine instability throughout 1882. They concluded that the vaccine must be “fresh” or “recently prepared,” or if the second vaccine, then veterinarians should put it “in a cool place [au frais], in a cellar for instance,” without opening the tubes and using within 12–14 days. Vaccines could, however, be prepared with “minute” rigor such that they would last two or three months. Ultimately, however, the goal would be to build myriad “little factories” in far–off countries.47 When accidents did occur, it was said, for example at a meeting of the French veterinary society in 1882, that Pasteur’s “vaccines were not fixed as he had hoped.” 48 The pages of Chamberland’s 1883 chapter on “Variation in the virulence of the anthrax vaccines” bore the header “Relative fixity of the vaccines.”49 Fixity, the breeder’s concept and aesthetic, economic ideal of “fixed” varieties, is a keyword for understanding how the vaccine model changed the meaning of heredity in the world of microscopic life. The currency of the term in these discussion may have come from the veterinary context of anthrax vaccination, but presumably also from the Pastorians own prior twenty years’ work in agricultural science and industry, which had made 42 43 44 45 46 47 48 49

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Koch, Gaffky and Loeffler 1884, 250, and see 252 (“Varietäten”). The same terminology occurs in Haeckel’s laws of heredity: “um so sicherer und vollständiger auf alle folgenden Generationen vererbt” (Haeckel 1866, 187). Chamberland 1883, chap. 30. Pfeiffer 1903, 48: “Virulenzkonservierung im Reagenzglase.” Chamberland 1883, 282; see also Pasteur 1880a, 299–300 and 1880c, 327, and reports in veterinary journals, such as Bouley 1881, 405; Rózsahegyi 1882, 24; Jahresbericht, Jahr 1881, 1882, 10. Chamberland 1883, 282, 295–96. Société centrale de médecine vétérinaire, séance 8 juin 1882, “Sur certains accidents consécutifs à la vaccination charbonneuse,” reprinted in Chamberland 1883, 290. Chamberland 1883, 284–295.

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them well aware of the theory and practice of animal and plant breeding, especially concerning grape vines and silkworms. For an attenuated fowl cholera culture to be a vaccine, Pasteur wrote in his first paper on attenuation, “It would be necessary, if I may so speak, that it were fixed in its own variety [fixé dans sa variété propre] and that one would not at all be constrained always to go back to its original preparation when one wishes to use it.”50 Later, anthrax inoculation “accidents” showed that given batches of anthrax vaccine were not “absolutely fixed,” not yet “true races with fixed characters.” With Jenner’s vaccine, one did not have to fear “these atavistic returns to the primitive virulence.” A safe vaccine was “a family of virus in which the attenuation is fixed by heredity.”51 Fixée par l’hérédité, “fixed by heredity”: again, as in Koch’s conception, heredity was a force. Yet it was also capable of transcending its own nature as a phenomenon of degree to achieve fixity, the cessation of degree. Despite bacteriologists’ adoption of the keyword fixed/fixity,52 the relation between the business of vaccines and that of animal or plant breeding was not straightforward. Whereas the bacteriologists came to imply that variety was truly hereditary only when it was “fixed,” for the breeder, to fix a trait was not to make it hereditary but to stabilise hereditary change (or maximize the force of heredity) and thus prevent regression or “atavism.” The term fixed connotes inalterability, but in fact it belonged to the conception of heredity as a force ranging from weak to strong. As Darwin glossed: “fixedness of character, or strength of inheritance.” 53 The French plant breeder and leading seed company family Vilmorin acknowledged that the constancy of a breed depended on continued selection,54 and Hugo de Vries emphasized this as the “universal experience of breeders.”55 Vaccines were different. Unlike the breeder whose pedigree animals or seeds passed into the hands of buyers equally interested in maintaining the product, the vaccine–maker had to let go his product into the wild. His control ceased at the point of inoculation into millions of animal bodies. Thus constancy of characters after cessation of controlled culture was not a happy exception or a distant ideal, but an immediate life–and–death necessity. Thus although productive yield and profitability or economic viability of plant and animal breeds were roughly analogous to safety of vaccines, bacteriologists’ pursuit of vaccine safety and efficacy was turning the question of whether something was hereditary into a yes/no question, rather than a question of degrees of strength, or indeed degrees of “certainty” or “uncertainty.”56 Fixed versus unfixed, rather than fixity as a high degree, Darwin’s “strength of inheritance.” Speaking with science studies or history and sociology of science, we could say at this point that vaccine safety and efficacy constructed, produced, constituted heredity as fixity and in a new, absolute sense. Yet this language would be too socially and morally neutral: the stability in 50

51 52

53 54 55 56

Pasteur 1880a, 299; see also Pasteur and Thuillier 1883, 531: “chacan de ces états [de virulence] est suceptible d’être fixé par la culture”; Smirnow 1888, 242: “Es gelang mir, auf diese Weise drei gut unterscheidbare Virulenzstufen zu fixiren.” Chauveau 1885a, 617, 620–621; see also Cornil and Babès 1890, 242. See the introduction to the most important literature review on “variability”: Kruse 1896, 476: “Befestigen lassen sie [die Varietäten] sich durch Wiederholung der Züchtung in alten Kulturen” (emphasis added). Darwin 1888, 2:47. Gayon and Zallen 1998, 260 De Vries 1906, 787. For “certainty” and “uncertainty,” see Gayon 1995, 64.

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question was not just technical or epistemic stability, but safety. The nature of heredity, at least among the microscopic organisms, was coterminous with securing against accidents, shouldering responsibility for animal welfare and industry, minimising the potential for lawsuits or disputes with angry farmers or governments, ensuring commercial success for a vaccine over its competitors. It was the flip–side of these practical, medical, economic goals and the system of techniques and organisation designed to make those goals realizable. In this sense, the nature of heredity has a social and technical history. To quote from an introductory section in the most important bacteriological handbook around 1900: When “the artificially induced loss of virulence [becomes] a lasting property of the strain, transmitted from generation to generation . . . we call such strains vaccins.”57 Shorn here of its medical meaning, the word vaccins could be equally a biological term. To conclude: Are the contours of this story limited to the history of microbiology, or is there wider significance for the history of heredity? In 1866 phenomena of “fixity” (Befestigung) and methods by which organisms could be “‘pure’ cultured” (“rein” fortgepflanzt) still belonged to but one of nine “laws” of heredity outlined by Ernst Haeckel, and they remained firmly within the conception of heredity as force.58 After 1880 the pursuit of vaccine fixity, of safety and efficacy, exemplified a shift from heredity as a force, a phenomenon essentially of degree, toward heredity as a phenomenon essentially of presence and absence – a world in which Johannsen’s “pure lines” and genotype/phenotype divide, Mendel’s laws, and the search for units of heredity would soon make the most sense and command the chief attention.59 For the history of biology, breeding has most obviously been important in two ways: (1) in the formulation of Darwin’s theory partly through the analogy to artificial selection; (2) in the making of Mendelian genetics as a science whose main practice was the breeding practice of hybridisation – using or making “true” varieties to cross with one another and keeping records of the results. But the vaccine story suggests another relationship of practical breeding–like enterprises to biological science in this period. Here was one important area of science in which heredity was freed from environment and atavism alike and coming literally to mean permanence. Bacteriologists routinely contrasted “temporary” and “permanent” modifications.60 And this permanence was not found through observation of the workings of nature in the wild or even the research laboratory as such, but created through a practical enterprise of making things reliable, context–independent, accident–free. Heredity would be like a safe, effective vaccine. The vaccine story may be an example of how agricultural and other enterprises of biological stabilization and standardization in this period contributed to this shift in the meaning of heredity – a shift that was independent of hybridization and the origins of Mendelism; independent of the “hardening” of heredity through the rejection of inheritance of acquired characteristics by August Weismann and others; independent of cytology and of biological theories of subcellular structures

57 58 59 60

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Wassermann 1903, 248. Haeckel 1866, 187–88: “Gesetz der befestigten Vererbung.” I thank Wolfgang Lefèvre for directing my attention to Haeckel’s heredity laws. This shift is related but not identical to the shift from “force” to “structure” identified by Gayon 1995. See, for example, Smith 1894. For a biologist’s language of “temporary” versus “durable” characters and “permanency,” see de Vries 1906, 774, 786.

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and units of heredity. This would not be merely another example of how scientific phenomena are stabilized at any given historical moment. For here was a historical change that was itself a process of stabilization, a shift away from ambiguity and degree. “True” heredity would seem to be less an ideal or proposition or a perception of nature as such, than something whose existence was warranted or predicted by such enterprises as vaccine–making, animal and plant breeding or domestication, perhaps also human breeding. If the setting into motion of people and objects through natural–historical and agricultural projects, trade, colonialism, industrialisation and urbanisation was a key determinant of the history of heredity in the early modern period, 61 in the 19th century an ever increasingly man–made world of controlled and stabilized variety would seem to be crucial. No wonder it seemed to Wilhelm Johannsen, who displayed his debt to breeders such as Vilmorin, “daß die Verhältnisse der reinen Linien das eigentliche Fundament der Erblichkeitslehre sein müssen, selbst wenn man in den meisten Populationen – vor allem in der menschlichen Gesellschaft – überhaupt nicht mit reinen Linien zu tun haben kann.” 62 His pure line methodology was a scientific expression, a research–program residue, of this man–made world. A similar contrast had impressed itself upon Darwin, writing of inheritance: If animals and plants had never been domesticated, and wild ones alone had been observed, we should probably never had heard the saying, that ‘like begets like.’ The proposition would have been as self–evident as that all the buds on the same tree are alike, though neither proposition is strictly true. For, as has often been remarked, probably no two individuals are identically the same. . . . The saying that ‘like begets like’ has, in fact, arisen from the perfect confidence felt by breeders, that a superior or inferior animal will generally reproduce its kind . . . 63

Darwin continued and nuanced the picture in an important way: “Inheritance is not certain; for if it were, the breeder’s art would be reduced to a certainty, and there would be little scope left for [his] wonderful skill and perserverance.” And yet in the end, amidst the ambiguities of the “wild,” of what Johannsen called “most populations,” one could observe that, “Hard cash paid down [for prize animals], over and over again, is an excellent test of inherited superiority.” 64 Those “excellent test[s]” – hard cash paid again and again, vaccines safely injected by the millions – made clear heredity’s nature for contemporaries in ways that biological theorizing and experimenting alone could hardly do. It was by no means the only nature of heredity being revealed at this time.

61 62 63 64

Müller–Wille and Rheinberger 2004, 13. Johannsen 1903, 9. Cf. Gayon and Zallen 1998, 244, 260, who challenge the similarity between fixed breeds and pure lines. Darwin 1888, 1:531–32. Darwin 1888, 1:534.

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Quite the opposite, de Vriesian mutationism (with its own roots in a different horticulture) was both an argument against the reality of breeders’ stable varieties and an attempt to reestablish biology on the basis of genetic instability.65 Heredity by 1903 bore a Janus face, reliable and mutable.

Andrew Mendelsohn, Imperial College London, [email protected]

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Undermining the breeding analogy to the origin of species was necessarily as important to de Vries as establishing it had been to Darwin; de Vries 1901–03, 1:4–9, 86–87, passim; de Vries 1906, Lecture 27.

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References Amsterdamska O., 1987, “Medical and Biological Constraints: Early Research on Variation in Bacteriology”, Social Studies of Science, 17:657–87. Behring E. von, 1901, Diphtherie (Begriffsbestimmung, Zustandekommen, Erkennung und Verhütung), Berlin: Hirschwald, 1901. Beumer, Otto. 1887. Der derzeitige Standpunkt der Schutzimpfungen. Wiesbaden: J.F. Bergmann. Bouley, Henri. 1881. “Chronique,” Recueil de médecine vétérinaire, 6th ser., 7:401–423. ———. 1882. “La nouvelle vaccination,” Revue scientifique, 3rd ser., 2:546–550. Brock, T.D., 1990, The Emergence of Bacterial Genetics, Cold Spring Harbor: Cold Spring Harbor Laboratory Press. Buchner, Hans. 1880. “Ueber die experimentelle Erzeugung des Milzbrandcontagiums aus den Heupilzen” in Carl Wilhelm Nägeli, Untersuchungen über niedere Pilze aus dem Pflanzenphysiologischen Institut in München. München, 140–177. ———. 1882. “Kritisches und Experimentelles über die Frage der Constanz der pathogenen Spaltpilze” in Carl Wilhelm Nägeli, Untersuchungen über niedere Pilze aus dem Pflanzenphysiologischen Institut in München. München, 231–285. ———. 1883. “Die Umwandlung der Milzbrandbakterien in unschädliche Bakterien und die Entgegnung R. Koch’s an Pasteur,” Virchows Archiv 91:410–422. Chamberland, Charles. 1883. Le charbon et la vaccination charbonneuse d’après les travaux récents de M. Pasteur. Paris: Bernard Tignol. Chauveau A., 1889a, "Sur les propriétés vaccinales de microbes ci–devant pathogènes transformés en microbes que la culture destitua de toutes propriétés virulentes" Archives de médecine expérimentale et d'anatomie pathologique, 1:161–202 ———. 1889b, "Recherches sur le transformisme en microbiologie pathogène: Des limites, des conditions et des conséquences de la variabilité du Bacillus Anthracis," Archives de médecine expérimentale et d'anatomie pathologique, 1:757–97. ———. 1881. “Ferments et virus,” Revue scientifique, 3rd ser., 1:482–492. ———. 1885. “L’inoculation préventative du choléra,” Revue scientifique, 3rd ser., 10:353–360. ———. 1885a. “L’atténuation des virus,” Revue scientifique, 3rd ser., 10:614–623. Collins, Harry M. 1985. Changing Order: Replication and Induction in Scientific Practice. Sage. Darwin, Charles. 1888 [1868]. The Variation of Animals and Plants under Domestication, 2nd edn., 2 vols. London: John Murray. De Vries, Hugo. 1901–03. Die Mutationstheorie: Versuche und Beobachtungen über die Entstehung von Arten im Pflanzenreich. 2 vols. Leipzig: Veit. ———. 1906 [1904]. Species and Varieties: Their Origin by Mutation. Lectures Given at the University of California by Hugo de Vries. Ed. D.T. MacDougal. 2nd edn. Chicago: Open Court. Duclaux E., 1920 [1896], Pasteur: The History of a Mind, tr. E.F. Smith and F. Hedges, Philadelphia: W.B. Saunders. Flügge, Carl. 1888. “Studien über die Abschwächung virulenter Bacterien und die erworbene Immunität,” Zeitschrift für Hygiene 4:208–30. Gaffky G., 1881, “Experimentell erzeugte Septicämie mit Rücksicht auf progressive Virulenz und accomodative Züchtung,” Mittheilungen aus dem kaiserlichen Gesundheitsamte, 1:80–133. Gaffky G., Pfuhl W., Schwalbe J. (eds), 1912, Gesammelte Werke von Robert Koch, 2 vols. in 3, Leipzig: Thieme. Gayon, Jean. 1995. “Entre force et structure: genèse du concept naturaliste de l’hérédité” in Le paradigme de la filiation, ed. J. Gayon and J.–J. Wunenburger. Paris: L’Harmattan, 61–75. Gayon, Jean and Doris T. Zallen. 1998. “The Role of the Vilmorin Company in the Promotion and Diffusion of the Experimental Science of Heredity in France, 1840–1920, Journal of the History of Biology 31:241– 62. Geison G.L., 1995, The Private Science of Louis Pasteur, Princeton: Princeton University Press. Haeckel, Ernst. 1866. Allgemeine Entwickelungsgeschichte der Organismen = vol. 2 of Generelle Morphologie der Organismen. Berlin: Georg Reimer. Hunt, Bruce. 1991. “Michael Faraday, Cable Telegraphy and the Rise of Field Theory,” History of Technology 13. Jahresbericht über die Leistungen auf dem Gebiete der Veterinär–Medicin, Jahr 1881 – Jahr 1883, 1882–1884. Johannsen, Wilhelm L. 1903. Über Erblichkeit in Populationen und “reinen Linien”. Jena: Gustav Fischer. Klein, E. 1883. “On the Relation of Pathogenic to Septic Bacteria, as illustrated by Anthrax Cultivation,”

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Quarterly Journal of Microscopical Science, n.s., 23:1–68. Koch R., 1881, “Zur Ätiologie des Milzbrandes”. In: Gaffky et al. 1912, 1:174–206. ———. 1882, “Über die Milzbrandimpfung: Eine Entgegnung auf den von Pasteur in Genf gehaltenen Vortrag”. In: Gaffky et al. 1912, 1:207–31. Koch R., Gaffky G., Loeffler F., 1884, “Experimentelle Studien über die künstliche Abschwächung der Milzbrandbacillen und Milzbrandinfektion durch Fütterung,” Mittheilungen aus dem kaiserlichen Gesundheitsamte, 2:147–81. Kolle W., Wassermann A. (eds), 1902–04, Handbuch der pathogenen Mikroorganismen, 4 vols. in 5, Jena: Gustav Fischer. Kruse, W. 1896a. “Krankheitserregung” in Die Mikroorganismen, mit besonderer Berücksichtigung der Ätiologie der Inkfektionskrankheiten, ed. Carl Flügge, 3rd edn., 2 vols. Leipzig: Vogel, 1:271–419. ———. 1896b. “Variabilität” in Die Mikroorganismen, ed. Flügge, 1:475–493. Latour B., 1988, The Pasteurization of France, tr. A. Sheridan and J. Law, Cambridge, Mass.: Harvard University Press. ———. 1987. Science in Action: How to Follow Scientists and Engineers through Society. Open University Press. Loeffler F., 1881, “Zur Immunitätsfrage,” Mittheilungen aus dem kaiserlichen Gesundheitsamte, 1:134–87. M’Fadyean, John. 1894. “Vaccination against Anthrax,” Journal of Comparative Pathology and Therapeutics 7:325–332. Mazumdar P.M.H., 1995, Species and Specificity: An Interpretation of the History of Immunology, Cambridge: Cambridge University Press. Mendelsohn, J. Andrew. 2002. “‘Like All That Lives’: Biology, Medicine and Bacteria in the Age of Pasteur and Koch,” History and Philosophy of the Life Sciences 24:1–36. Müller–Wille, Staffan and Hans–Jörg Rheinberger. 2004. “Heredity – The Production of an Epistemic Space.” Max Planck Institute for the History of Science, Preprint 276. Pasteur L., 1876, Études sur la bière. In: Vallery–Radot 1922–39, 5. ———. 1880a, "Sur les maladies virulentes, et en particulier sur la maladie appelée vulgairement choléra des poules". In: Vallery–Radot 1922–39, 6:291–303. ———. 1880b, "Sur le choléra des poules; Etude des conditions de la non–récidive de la maladie et de quelques autres de ses caractères". In: Vallery–Radot 1922–39, 6:303–12. ———. 1880c, "De l'atténuation du virus du choléra des poules". In: Vallery–Radot 1922–39, 6:323–30. ———. 1880d. “Sur la virulence du microbe du choléra des poules,” manuscript note. In: Vallery–Radot 1922–39, 7:52–54. Pasteur L., Chamberland C., Roux E., 1881, "De l'atténuation des virus et de leur retour à la virulence". In: Vallery–Radot 1922–39, 6:332–38. Pasteur, Louis and Louis Thuillier. 1883. “La vaccination du rouget des porcs à l’aide du virus mortel atténué de cette maladie.” In: Vallery–Radot 1922–39, 6:527–34. Pfeiffer, Richard. 1903. “Zur Theorie der Virulenz” in Festschrift zum sechstigsten Geburtstage von Robert Koch. Jena: Fischer, 35–48. Rózsahegyi, Aladár von. 1882. “Versuche mit der Pasteur’schen Schutzimpfung gegen Milzbrand in Ungarn,” Deutsche medizinische Wochenschrift 8:24–27. Schaffer, Simon. 1992. “Late Victorian Metrology and Its Instrumentation: A Manufactory of Ohms,” in Invisible Connections: Instruments, Institutions, and Science, ed. Robert Bud and Susan E. Cozzens. Bellingham, Wash.: SPIE Optical Engineering Press. Smirnow, G. 1888. “Ueber das Wesen der Abschwächung pathogener Bacterien,” Zeitschrift für Hygiene 4:231–60. Smith, Theobald. 1894. “Modification, Temporary and Permanent, of the Physiological Characters of Bacteria in Mixed Cultures,” Transactions of the Association of American Physicians 9:85–109. Vallery–Radot P. (ed.), 1922–39, Oeuvres de Pasteur, 7 vols., Paris: Masson. Wassermann A., 1903, “Wesen der Infektion”. In: Kolle and Wassermann 1902–04, 1:223–87.

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The Chromosomal Theory of Heredity and the Problem of Gender Equality in the Work of Theodor and Marcella Boveri1 Helga Satzinger

In the early years of the twentieth century Theodor Boveri (1862—1915) and Walter Sutton (1877—1916) proposed the chromosomal theory of heredity. In his paper ‘Ergebnisse über die Konstitution der chromatischen Substanz des Zellkerns’, published in 1904, Theodor Boveri summarised the results of two independent fields of research: experimental cytology on the one hand, which was his field of expertise, and Mendelian hybridisation experiments on the other. At the end of his 130-page paper he came to the conclusion: ‘The probability becomes extraordinarily high that the characters traced in Mendelian experiments are actually bound to specific chromosomes’.2 Working independently, Sutton and Boveri correlated the behaviour of chromosomes during cell cleavage, germ cell development, and cell fusion in fertilisation to the recently rediscovered laws of Gregor Mendel.3 In doing so, they laid a cornerstone of modern genetics, as our textbooks on the history of biology tell us. The localisation of the hereditary material in the chromosomes became the precondition of all gene-mapping projects of the twentieth century, the first one starting in 1911 with the famous fruit fly Drosophila melanogaster and a recent one being the HGP with humans.4 Yet, upon a closer examination of how Boveri established the chromosomal theory of heredity, things do appear confusing. First of all, Boveri did not call his theory the ‘chromosomal theory of heredity’, he gave no name for his combination of two fields of research. In 1904 he was proposing the ‘theory of chromosomal individuality’, attributing to each chromosome a specific relevance 1

2

3 4

This paper is part of an extended work on the history of genetics from the perspective of science and gender studies. I want to thank Staffan Müller-Wille, Karin Hausen, Christiane Eifert and my colleagues at the Wellcome Trust Centre for the History of Medicine at UCL for critical and supportive comments. Boveri (1904), p. 117: ‘... so wird die Wahrscheinlichkeit, daß die in den Mendelschen Versuchen verfolgten Merkmale wirklich an bestimmte Chromosomen gebunden sind, ganz außerordentlich hoch.’ [Translation: H.S.] Translating Boveri’s texts into English is difficult, both because the meaning of certain terms changed over the years, and because some German forms of conceptualisation escape an appropriate one-to-one translation. The terminology of genetics was developed in the first decade of the twentieth century, but Boveri did not use it: he used his terms derived from cytology and from early Mendelian work in Germany. Especially for the mathematically defined entities, which were called Mendelian ‘genes’ after 1909, a translation is not possible without changing the meaning in an a-historic way. Boveri most frequently used the term ‘Anlagen’, which I do as well in this paper, adding the English expression ‘disposition’ as an indicatory term in brackets. The only contemporary, but still very late translation of a text of Theodor Boveri was done by his widow Marcella Boveri in 1929: Boveri, Theodor: The Origins of Malignant Tumors. London, Baillière, Tindall & Cox, 1929, German: 1914. Here the term ‘inheritance factors’ is used for the German ‘Erbfaktoren, über die wir durch die Mendelforschung unterrichtet sind’, thus linking this hereditary unit directly to the experimental approach of the Mendelians. Marcella Boveri’s translation, not using the term ‘gene’ but ‘inheritance factors’, indicates a conceptual difference between the ‘heredity’, she and her husband were investigating and the transmission of ‘genes’, the Mendelian geneticists were after. This will become clearer, as I hope, at the end of this paper. The terms ‘mitosis’ for cell fission and ‘meiosis’ for the cell divisions during germ cell development were coined in 1905. See: Churchill (1970), esp. p. 445. Rheinberger and Gaudillière (2004); Gaudillière and Rheinberger (2004).

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for the inheritance of certain characters. This difference in the theory’s name signifies an important difference in the understanding of heredity that Boveri had in mind, compared to that of his contemporaries and even that of recent genetics. Second, well equipped with new experimental results, Boveri called his own theory into question shortly before his untimely death in 1915. Since 1902 he had claimed that the chromosomes imprinted parental properties onto the next generation’s cell or organism, as the substrate of the hereditary material. 5 In 1915 he expressed his doubts: ‘Experience does not teach us what the substrate that contains the paternal [!] ‘Anlagen’ [‘dispositions’] might be.’6 Thus, the cell nucleus probably no longer could be seen as the carrier of the ‘Anlagen’, but as a factor enabling development only [‘Entwicklungsfaktor’]. Boveri came to the surprising conclusion — and he did it explicitly in the face of the rapidly growing science of genetics — that ‘Our knowledge of heredity itself ... amounts to nearly nothing.’7 These doubts of Theodor Boveri reflect a very complicated process in the generation of knowledge in cytological research, which aimed at an understanding of heredity. This research used very sophisticated experimental approaches and microscopic techniques to illuminate the interplay of the cell’s plasma and its elements on the one hand and the chromosomes in the cell’s nucleus on the other. Both plasma and chromosomes took part in the process of heredity — the question was how they interplayed. The creation in 1902/1904 of the scientific fact that bound hereditary ‘Anlagen’ to chromosomes was a decision taken in favour of the chromosomes as the decisive entity in the cell. In the same move the cell’s plasma became the supportive element, active only in the ontogenetic realisation of an organism. This decision had far-reaching consequences for the science of heredity and the understanding of the functions of the cell. It helped to create a split into two disciplines, genetics dealing with the chromosomes, and embryology investigating the plasma and development. It was reinforced in further developments in genetics, social and experimental, which centred the experimental approach on the chromosomes, the genes in the chromosomes, and the DNA as their basic structural unit. Needless to say, the belief in an identifiable, stable, Mendelian hereditary unit was crucial for the belief in the feasibility of eugenics and plant and animal breeding, and it is still alive in the current understanding in medicine of a ‘gene for’ a certain disease. By using the word ‘decision’ I am implying that there were other options available around 1900. Because the chromosomal theory of heredity was not accepted immediately by Boveri’s 5

6

7

‘In diesen väterlichen und mütterlichen Kernelementen [i.e. den Chromosomen] müssen wohl die dirigierenden Kräfte liegen, welche dem neuen Organismus neben den Merkmalen der Species [sic] die individuellen Eigenschaften der beiden Eltern kombiniert aufprägen.‘ Boveri (1902), p. 35. ‘Welches das Substrat ist, das die väterlichen Anlagen enthält, darüber lehrt die Erfahrung nichts. Den Kern läßt sie nur als Entwicklungsfaktor, nicht als Träger der erblichen Anlagen erkennen.’ Boveri † (1918), here p. 467. Published posthumously and in an unfinished state by his widow, Marcella Boveri. The manuscript reached the publishing journal on 10 April 1917; p. 417. - The word ‘Anlage’ in embryology refers to an undifferentiated cellular structure out of which a certain organ or limb may develop. In this sense it also can mean a potential of a given structure, whereas the ‘Entwicklungsfaktor’ refers to a function only, enabling the development. Both notions have their difficulties, as it was not known how at the microscopic and submicroscopic level the development — not growth — of an organism took place. ‘Über die Vererbung selbst ... wissen wir so gut wie nichts.’ Ibid., p. 417.

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contemporaries due to various scientific and some idiosyncratic reasons, we can say that options existed.8 The impression of a certain fragility of the theory is underlined by the fact that, in the years 1914/1915, Boveri himself found it difficult to stick to his own theory, which he had developed ten years earlier and still praised in 1913, a fact overlooked by current historiographers and biologists. Evidence for the importance and activity of cell plasma had been provided by Boveri himself in the years before and after 1902/1904, when he was studying heredity in cell fission and embryonic development in the flatworm Ascaris and in sea urchin embryos. Thus, data on the plasma’s importance were available at the time of the chromosomes being made the site of the ‘Anlagen’; they were there at the beginning of ‘the century of the gene’. The cytoplasm and its activities were not discovered by geneticists at the end of the twentieth century in the form of ‘“postgenomic” metabolic pathways’ or ‘multiple systems of inheritance’. 9 It was there in the founding years of genetics and it is currently rediscovered by developmental geneticists praising Boveri as their long forgotten founding father.10 In this paper I shall argue that the problem of gender equality was one important cultural and social factor in the making and stabilisation of the chromosomal theory of heredity. This social problem was a factor that supported the decision in favour of the chromosomes as the hereditary material and hampered the possibilities to formulate the interaction of cell plasma and chromosomes as a kind of co-operation, most likely non-hierarchical, by using the experimental evidence available in 1915. Historians of biology have shown for the 1920s and later that the relations of cell nucleus/ chromosomes and cell plasma were understood according to highly contested political concepts, such as the monopoly of power exercised by the nucleus upon the plasma, the ‘Kernmonopol’. 11 According to recent science and gender studies, the chromosomal theory of heredity is an example of how the order of the cell’s elements follows a gendered hierarchy. Analysis of the scientific language and the metaphors used reveals the familiar Aristotelian, hierarchical dichotomy between male form and female matter: on the one hand, the (male) active, controlling, imprinting chromosomes or genes, represented by the sperm; on the other, the female, passive, obedient cell plasma, the egg.12 But, in considering the making of the gendered order of the cell before World War I, something else becomes apparent. The decision in favour of the chromosomes as the carriers or place of the ‘Anlagen’ reflects an uneasiness. Cytologists based their understanding of hereditary processes on the investigation of cell fusion in fertilisation and subsequent cell fissions. Since 1875 fertilisation was understood within the paradigm of cell theory, thus entailing a highly alarming connotation. Former theories of fertilisation saw a male force active on female matter, thus providing a hierarchical order of unsurpassable eternal and cosmic dimensions. 13 At the new 8

9 10 11 12 13

Thomas H. Morgan, Hans Driesch, and Oskar Hertwig, just to mention three important contemporary scientists, did not immediately accept the chromosomal theory of heredity in the decade before World War I. Cremer (1985); Gilbert (1978). Keller (2000), p. 9. Moritz (1993); Moritz (1996). Harwood (1993), pp. 315—350; Sapp (1987). The Biology and Gender Study Group (1989); Keller (1995). Keller extends the argument to the ‘gene action talk’, in which the gene even became the only representative of life itself. See Lesky (1951), pp. 125-159, on the canonisation.

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material level of cells only, the male tended to be much smaller than the female, if not irrelevant. In this situation the chromosomes defined as the hereditary substance helped to rescue gender equality for the male. It was in no way a feminist move to claim gender equality at the level of cells and chromosomes; it was a move to regain at least some male influence in the realm of generativity and heredity in a situation in which matter mattered. Contemporaries noticed that a negotiation of the social gender order was taking place in cytology. In 1909 the Prague scientist and historian of biology Emanuel Rádl saw the ‘philosophy of gender’ [die Philosophie des Geschlechts], ‘the subject of deepest thoughts throughout the ages, culminating in the science of the chromosomes’.14 Rádl criticised Boveri, Hertwig and other cytologists for their claim that there was equality and no fundamental difference between men and women. The situation appeared to be even worse: Rádl uttered the fear that man himself was in danger. The research of Jacques Loeb had shown that fertilisation and egg development could be initiated by chemicals only. In Rádl’s words, ‘Some potassium chlorate or everyday salt taken from the kitchen may substitute the male of the Echinide, of the worms, the starfish and other animals, if not the human male himself.’15 Before looking more closely into Boveri’s formulation of the chromosomal theory of heredity, some remarks on his social background are relevant.16 His work is the work of a creative couple in the sciences.17 Theodor Boveri was born the son of a medical doctor in 1862 in a small town of northern Bavaria. He studied in Munich, where he got his doctorate title, beginning with ancient history and philosophy, then changing to anatomy. Again he changed to the Institute for Zoology under the directorship of Richard Hertwig, and continued his cytological work on the cell in fertilisation and development. In 1893 he became professor for zoology and comparative anatomy at the University of Würzburg, primarily teaching medical students. The institute became an internationally renowned place to do the latest research in cytology, and several female scientists of the first generation worked here and completed their doctoral dissertations. Theodor Boveri gained such a high reputation in the German scientific community that he was assigned to become the director of the newly founded Kaiser Wilhelm Institute for Biology in Berlin-Dahlem in 1911/ 1913. The institute and the composition of its staff was planned by Boveri, but in the end he did not take up its directorship. Since 1897 he had been married to the U.S.-American Marcella O’Grady (1863—1950). After their marriage they co-operated scientifically all his life, with Marcella Boveri remaining in the shadows of her husband. The daughter of a Boston architect, she was the first ‘woman to graduate with a concentration in biology’ at the Massachusetts Institute for Technology. She studied comparative zoology and embryology at Bryn Mawr College, carried out research at the Marine Biological Laboratory (MBL) Woods Hole and received an appointment as a teacher of biology at Vassar Women’s College in 1889. Here she became a full professor in 1893 and developed a new curriculum. In 1896 she left for a sabbatical year with 14 15 16 17

Rádl (1909), p. 498. ‘Die Philosophie des Geschlechts, welche zu allen Zeiten den Gegenstand tiefsten Nachdenkens bildete, kulminiert heute in der Lehre von den Chromosomen.’ Ibid., p. 501. ‘Ein wenig Chlorkali oder Küchensalz ersetzt, wenn nicht geradewegs den Mann, so doch das Männchen der Echinide, der Würmer, der Seesterne u.a. Tiere.’ Baltzer (1962); Neumann (1998). Wright (1997). See, for comparison, the case studies of several marital co-operations in: Pycior, Slack, and Pnina (1996).

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Theodor Boveri at the Würzburg Institute in order to do a PhD dissertation, which she published in 1903. She stayed, raised one daughter and returned to the United States in 1927 to work as a biologist, teaching at the Albertus Magnus Women’s College in New Haven. 18 In the life of the Boveris the woman was not restricted to a purely female domain: she took part in his professional work, understood what he was doing, and co-operated in the experiments and other scientific activities. In addition, she was responsible for the running of the house, with a child and servants.19 Marcella and Theodor Boveri lived at a time when women in Germany were struggling hard for regular access to academic training and independent income and for an improved legal standing. Women in Germany faced a severe step back as in the years around 1900 a new German Civil Code was passed, which made the situation for women much worse than before. It gave husbands control over their wives: they had the final decision in any respect, they controlled women’s means and property, they decided on all matters concerning the children, on her professional activities and so on. The women’s campaign for suffrage ended in 1919, with the new Weimar constitution giving women the right to vote.20 The women’s movement was an important social force in the time of the Boveris, and its repercussions can be seen in their professional life. The scientific work of Theodor Boveri has to be regarded as the work of a married couple, thus creating some problems for the historiographer. How to talk of a work that is authored by one person only, but created by two from a certain time, from 1897, onwards? The publications bear his name only. How to escape ‘the Mathew Matilda Effect in Science’, which attributes to the known male scientist the contributions of his collaborator?21 I would like to pay tribute to Marcella Boveri’s contribution to the shared work. She was active in the performance of the experiments, but she did not publish under her own name, with one exception only. 22 Marcella Boveri even did not finish her late husband’s last and incomplete paper, she cannot be viewed as his invisible co-author, and it does not seem legitimate to attribute every sentence that Theodor Boveri authored to her as well.23 For lack of a better solution, I refer to Theodor Boveri when I am discussing the papers he put his name on, thus risking a continuation of her ‘silencing by his pen’.24 I use the phrase ‘the Boveris’ when I am referring to their common work and not explicitly to him as the author of a specific statement in a paper. All the usual elements that take part in the creation of a scientific fact can be found in the making of the chromosomal theory of heredity. There was a good deal of experimental and observational material involved, women’s work, inductive and deductive modes of reasoning, several hypotheses carefully combined, with the most important ones left to be tested further in complicated experiments. Scientific enemies had to be fought in the development of the new theory [e.g. Oskar Hertwig], and scientific authority had to be accumulated by building bridges to promising new fields of research, such as the Mendelian hybridisation experiments.

18 19 20 21 22 23 24

Wright (1997), pp. 629-636. Ibid. pp. 638-643; Boveri (1982), pp. 10-50. See, e.g., Bleker (1998); Hausen (1986); Special Issue ‘Universität — Frauen — Universitäten’, Feministische Studien 20/1, 2002; Gerhard (1990), here pp. 137-324. Rossiter (1993). Boveri (1903). Boveri † (1918). Pycior, Slack, and Abir-Am (1996), p. 6.

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The above-mentioned paper of 1904, ‘Ergebnisse über die Konstitution der chromatischen Substanz des Zellkerns’, presents a combination of the Boveris’ own observations and experimental results and the results of twenty years of cytological research, undertaken by an international group of male and female cytologists and zoologists. Most of them came from Germany, the United States and Belgium. Nearly all of them also had worked for some time at marine laboratories, like the Stazione Zoologica in Naples; they used insects, sea urchins, frogs, Ascaris and other species as animal models.25 The paper starts with the interpretation that the number of chromosomes was a characteristic one for each species and that the chromosomes appeared in a reliable and constant way in every cell division. This ‘persistence of chromosomes’ was a hotly debated issue of the time and not at all agreed upon by all cytologists.26 This finding was combined with the results of two of the sea urchin experiments Theodor Boveri had begun in 1889 and continued with Marcella Boveri. 27 In the so-called Merogonie experiment, egg cells were deprived of their nucleus and fertilised with one or two sperm of another sea urchin species. In the early experiments made before 1902, the resulting larvae did not show maternal properties, thus proving relevance of the sperm’s nucleus. The second experiment used sea urchin eggs that had been fertilised simultaneously by two sperms; it showed abnormal developments due to the wrong number of chromosomes in the various cell lineages derived from the fertilised egg. Both experiments used the variation of the number and quality of chromosomes in fertilised eggs and its effects on the developing embryo. The experimental results allowed the conclusion that each chromosome had its own relevance for the development of the new organism. This interpretation was called the ‘theory of chromosomal individuality’. Attributing a specific quality to each chromosome and describing the regular reduction of chromosomes during germ cell development, it was possible to see a parallel between the behaviour of chromosomes and the Mendelian ‘Anlagen’. Most of the experiments stabilising the ‘theory of chromosomal individuality’ were performed in a sophisticated way in the years to come until the final ones led to some destabilisation.28 The gender problem lay in the apparent size difference of the germ cells, the maternal ones contributing much more material to the offspring than the paternal ones. As parthenogenesis showed, the egg cell made a different and a much greater contribution to heredity than the spermatozoon did.29 Theodor Boveri had elaborated on that explicitly in a short paper of roughly 40 pages, ‘Das Problem der Befruchtung’ [The Problem of Fertilisation]. 30 The paper had been published in 1902 and made the first step to the chromosomal theory of heredity, linking chromosomes to the Mendelian ‘Anlagen’ as the parental properties [Eigenschaften] were situated in the chromosomes. 25

26 27 28 29 30

Some names should be mentioned: Edmund B. Wilson, Thomas H. Morgan and his wife Lillian Morgan, C. E. McClung, Nettie Maria Stevens, Kristine Bonnevie, Oskar Hertwig, Yves Delage, Eduard van Beneden. See, Cremer (1985). The first paper on a series of experiments of 25 years is: Boveri (1889). Boveri (1910); Boveri (1908). Boveri (1904), p. 112. ‘... und es hat ... die Eizelle ... eine andere und ungleich viel größere Bedeutung bei der Vererbung als die Samenzelle.’ Boveri (1902). The paper derived from a talk at the Versammlung Deutscher Naturforscher und Ärzte in 1902. All of the following quotations are from this paper.

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The understanding of fertilisation was the starting point for the understanding of heredity in cytology. The Boveris’ research can be characterised as the ‘investigation into processes, which cause the generation of a new individual with specific properties from the parental procreative substances.’31 Hence, research into heredity had to begin with the germ cells. The paper on the problem of fertilisation began with the proud declaration that the untranslatable ‘uralte menschheitsgeschichtliche Problem’ finally was solved. Everywhere in organic nature, even at the level of protozoons, two sexes co-operated in the creation of their offspring. This, according to Theodor Boveri, was new.32 In his eyes the co-operation occurred in a reciprocal way, as both germ cells were dependent on each other. Both germ cells had a potential for cell cleavage and development, but they were inhibited. Through co-operation they could overcome this inhibition, through co-operation they supplemented each other in their intrinsic urge to procreate and combine different properties, through co-operation they had enough plasmatic and nutritive substances necessary for the building of the embryo.33 This order of the germ cells resembles quite clearly the ideal of a German middle-class/ bourgeois couple around 1900, practising the model of the ‘Arbeitspaar’ [working couple] in a gendered division of labour, like the Boveris themselves. The unquestioned purpose was the raising of children and the common production of its economic basis, enhanced by the dowry of the woman. At the level of biological reasoning, Theodor Boveri derived his concept of the reciprocity of the male and female germ cells from evolutionary thinking. Boveri was no Darwinist in the sense of seeing selection as one important force of evolution. According to Boveri, the evolution of the organisms was a progressive process driven by intrinsic forces, the ‘bildnerische Elementargesetzlichkeit’, leading from the primitive ‘Urzustand’ [primordial state] to utmost complexity.34 Attempting to find a reason for the existence of male and female germ cells, Boveri described a line of development starting with single cells procreating via cell fission. Then copulation developed between two equal cells. At the evolutionary state of colonies of 16 cells, a differentiation of copulating cells developed — and in the case of the flagellate Eudorina elegans, the first egg and sperm cells were to be found, characterised by their difference in size. The ‘Urzustand’ was the self-sufficient cell, procreating through growth and subsequent fission. Bees and other insects procreating partially by parthenogenesis exhibited this property of the egg cell. The egg cell in higher animals still had this self-sufficiency, albeit a bit hampered by inhibition. Boveri compared the egg cell to a perfect clock that was missing the spring. In 1902 he had to give up his earlier idea that the centrosom of the sperm caused the cell cleavage of the (fertilised) egg cell, as this process was not found to be a general one in all organisms. Thus, Boveri abandoned the concept of a male induction of embryonic development, which he saw as being in perfect line with the Aristotelian notion of the female providing the matter and the male giving the activating stimulus for the movement of the matter.35 Having lost this possibility of explaining a 31 32 33

34

Es ging um die ‘Erforschung jener Vorgänge, ... durch die aus den elterlichen Zeugungsstoffen ein neues Individuum mit bestimmten Eigenschaften hervorgeht.’ Baltzer (1962), p. 81. The long tradition of not seeing two sexes or more existing in the plant kingdom had been overcome in the 16th century; now the protozoons had it as well. Theodor Boveri followed August Weismann’s line of argument that amphimixis, the combination of different parental properties in the offspring, was the purpose [Zweck] of fertilisation. Boveri (1902), p. 36. Ibid. p. 38.

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universal biological difference between the male and the female, he tried to create a full reciprocity of male and female germ cells: ‘One could say the spermatozoon is fertilising the egg, one could also say that the spermatozoon is being fertilised by the egg.’36 Both cells needed each other to get procreation started. However, Boveri had to concede that it was always the egg cell that started development: there was parthenogenesis only and no androgenesis. No development of an embryo started from the sperm. Having to admit a generative difference in germ cells, Boveri posed the question: ‘How does it come that the properties of the sperm are not suppressed within the egg? How do they cope with the properties of the egg, which exceeds the sperm in size by the thousands and millions?’37 Boveri sought rescue in the mere statement that the sperm could cope with the egg, though he did not know how. He referred to the experience that ‘generally the father’s influence on the constitution of the child equalled that of the mother’s’. 38 This equal influence was guaranteed by equality at the level of chromosomes. Boveri saw his work in the tradition of others, starting with Carl Naegeli’s postulate of an ‘Idioplasma’. Naegeli had claimed that a substance was present in every cell in a very small quantity, which derived from equivalent substances in the egg and the sperm cell. This substance provided an equal force of heredity [‘gleiche Vererbungskraft’] of both parents, despite the enormous differences in their material contributions for the gestation of a child.39 After 1900 the Mendelian laws provided an experimental approach to prove parental equality in heredity. This claim of equality, however, entailed a specific definition of heredity. In a rather circular reasoning, heredity now only dealt with properties characterised by their binary difference in both parents. In other words, only the inheritable differences between members of a species could be dealt with and localised in the chromosomes. All the general properties of an individual, like the inheritable features of the mammalians, the properties of the genus and higher classificatory groups, were not included in this definition.40 For Boveri, the chromosomes of the sperm incorporated the sperm’s equal influence on the offspring’s properties in this narrow sense of inheritable properties. Describing the behaviour of chromosomes after the sperm’s integration into the egg, he rhapsodised in a pseudo-religious language: ‘Indiscernible the grown nucleus of the sperm stands face to face with the nucleus of the egg; in fullest equality in size, form and number [‘Gleichheit nach Größe, Form und Zahl’] the paternal and maternal nuclear elements [‘Kernelemente’] lie close to each other. They are passed on in the same combination to the daughter cells and, as we may suppose, to all the cells of the new individual. All this happens with unsurpassable, painstaking care. In these paternal and maternal nuclear elements lie the directing forces, which in combination imprint onto the new organism not only the properties of the species but the individual characters of the parents.’ 41

35 36 37 38 39 40

Ibid. p. 23. Ibid. p. 34. Ibid. p. 35. ‘Zahllose Erfahrungen ... lehren, daß der Vater auf die Konstitution des Kindes im allgemeinen ebenso viel Einfluß hat, wie die Mutter.’ Boveri (1902), p. 35. Boveri (1904), pp. 102-103. The paradigmatic example for the species-specific properties lying in the chromosomes was derived from the crossbreeding of a horse and a donkey. It made a considerable difference in the offspring if the maternal animal was the horse or the donkey, thus showing the influence of the maternal cell plasma.

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Boveri was so fascinated by the notion of two equal sets of chromosomes in the fertilised egg that it took him some time in the following years to accept the findings of Nettie Maria Stevens (1861-1912) on chromosomal sex determination. According to this theory, two types of male germ cells existed, which differed in the number of chromosomes, thus determining the sex of the offspring.42 Stevens had worked with Boveri in his institute in Würzburg in 1903, and she was the one who immediately saw the applicability of Boveri’s chromosomal theory of heredity to the problem of sex determination. Boveri did not — he was trapped in the creation of gender equality at the level of chromosomes. As late as 1908 he revised his interpretation that there always were two completely equal nuclei in the germ cells.43 In the following years, Theodor Boveri argued as a convinced Mendelian geneticist. In 1913 he praised the results of the latest research, which enabled the localisation of certain characters onto the sex chromosome: ‘sex and colour blindness — what could be more different? Nonetheless, there is nearly no doubt that the ‘Anlagen’ for both characters are located in the same chromosoma.’44 He thus was on the same track as his scientific enemy Thomas H. Morgan, who, with his group of PhD students had started mapping Drosophila genes onto sex chromosomes.45 The Boveris, however, favoured a different experimental approach: heredity was to be investigated in embryonic development. The qualities of the germ cell’s nucleus were to be analysed here. This approach was called in analogy to the spectral analysis of light in physics: ‘Embryonalanalyse des Zellkerns’ [‘Embryonic analysis of the cell nucleus’]. Heredity was to be investigated in fertilisation and ontogenesis; the methods used were a combination of cytological analysis and hybridisation experiments, including interspecific crossbreeding of sea urchins. In co-operation with his wife, Marcella O’Grady, Theodor Boveri designed experiments to change the chromosomal constitution of the egg cell and fertilised egg to observe and interpret their abnormal development. He planned to use a device that was to remove single chromosomes from a cell, and various other techniques were applied to influence a cell such that the relation of plasma and chromosomes was changed. These experiments aimed at the understanding of the interplay of chromosomes and cytoplasm at various stages of embryonic development. The last Merogonie experiments did not provide the desired result that the chromosome was the site of the ‘Anlagen’. Marcella Boveri published her husband’s last paper in its unfinished form in 1918 under the very unpromising title: ‘Zwei Fehlerquellen bei Merogonieversuchen und die Entwicklungsfähigkeit merogonischer, partiell-merogonischer Seeigelbastarde’. 46 If this paper contained at least some dynamite to shake the chromosomal theory of heredity, then most probably nobody took notice — with one exception perhaps: Richard Goldschmidt (18781958).47 For the geneticists after World War I, there was no longer a need to take notice of the 41

42 43 44

45 46

‘In diesen väterlichen und mütterlichen Kernelementen müssen wohl die dirigierenden Kräfte liegen, welche dem neuen Organismus neben den Merkmalen der Species die individuellen Eigenschaften der beiden Eltern kombiniert aufprägen.’ Boveri (1902), p. 35. Brush (1978). Boveri (1909). ‘Geschlecht und Farbenblindheit, was könnte verschiedener sein. Und doch können wir ... kaum zweifeln, daß die Anlagen für beide Eigenschaften in dem gleichen Chromsoma lokalisiert sind.’ Boveri (1913), p. 16. See, Kohler (1994); Gilbert (1978). ‘Two sources of artefacts in merogonic experiments and the potential for development of merogonic, partially merogonic sea urchin bastards.’

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results of a paper that promised to deal with methodological problems in a very complicated experimental system using sea urchin development, as it no longer was in use in the research into heredity. The main and unexpected results of Boveri’s last paper are that embryonic development is divided into two phases. In the first phase, the plasma alone is active; the chromosomes step in at a later stage. In addition, chromosomes and plasma need to be compatible with each other in order to enable proper development. As mentioned, the results were such that Boveri doubted his own proposal of 1904, that the chromosomes might be the material substrate of heredity. But this last, unfinished paper shows as well that these new and severe doubts were outweighed by the desire to believe, in disregard of the new findings, that the substance within the cell nucleus actually was the hereditary substance [die ‘Vererbungssubstanz’]. 48 Boveri did not reformulate the interaction of plasma and chromosomes in heredity according to his and his wife’s new findings. It makes no sense to speculate what Boveri might have done had he lived longer. He ended with drawing a line between the ‘exakte Vererbungslehre’, the newly developed genetics, on the one hand and the research into the processes of ontogenesis on the other. The one approach used hybridisation experiments, the breeding of pure lines, it applied Mendel’s laws and the notion of a gene; the latter asked how the constellation in the zygote leads to the ‘Erbeffekt’ [Charakter] that the geneticists [‘Vererbungsforscher’] deal with. For Boveri, only the latter equalled ‘heredity itself’, the ‘Vererbung selbst’. In the work of the Boveris the interaction of plasma and chromosomes in the fertilised egg and the developing organism was conceptualised in various ways. In the same move in which gender equality at the level of the chromosomes was introduced, a gendered hierarchy between the chromosomes and the plasma was established. In the years 1902 and 1904, Theodor Boveri appreciated the Aristotelian notion of fertilisation as female matter set in motion by a male activating impulse, but he had to concede that this solution was not a general one and dropped it. The Aristotelian hierarchical dichotomy between form and matter could, however, be applied to the chromosome-plasma relation by interpreting the chromosomes as entities that contained ‘conducting forces to imprint paternal properties onto the egg cell and the organism of the next generation’. Theodor Boveri was criticised by colleagues who saw him as advocating the autocracy of the nucleus within the cell.49 Boveri replied that the cell plasma and the nucleus were dependent on each other and that neither could exist alone; he used his opponents’ metaphor: the huge crowd of workers necessary for the autocrat’s existence was present in the cell plasma and inherited by the plasma.50 His own analogy was that of the brain and the body to exemplify the mutual dependency of chromosomes and plasma, or of the plan of an architect and the construction workers building the house.51 Obviously, mutual dependency was not an equal dependency. It entailed a clear hierarchy following the Aristotelian model, which ensured a gender hierarchy as 47

48 49 50 51

Goldschmidt who followed his own track in conceptualising the genes and the chromosomes became the head of one department at the Kaiser-Wilhelm-Institute in Berlin in 1914. He appreciated Boveri’s work until the end of his own life. Boveri (1918), p. 468. Boveri (1904), p. 103. ‘Vor allem wird eben im Protoplasma das ganze Heer des Arbeitsvolks vererbt, ohne welches selbst ein Alleinherrscher, wenn wir einmal dieses Bild gebrauchen wollen, nicht existiert.’ Ibid, p. 113. Ibid, p. 103.

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well. The order of cell nucleus and plasma followed this double hierarchy. At the level of gender, it parallels the legal situation of a married couple according to the civil code of the time. The man had the legal power to make all decisions, he was the head of the household, and all the negotiations and co-operations made to keep the family going were left invisible between husband and wife. In 1914/15 there was no new conceptualising of the nucleus-plasma relation possible, albeit the possibility had to be considered that the chromosomes were not the decisive forces but enabling the specific ‘Gestaltung’ of the developing organs of an organism only. The cause for its concurrence with the specific ‘Gestalt’ of the parents could lie in other parts of the cell. 52 It is even much more remarkable that the Boveris did not re-conceptualise the plasma-chromosome relationship as Theodor and Marcella Boveri provided important findings on the interplay of the plasma and the cell nucleus, findings Theodor Boveri is praised for nowadays by developmental geneticists.53 1. The centrosoms and spindles organised the distribution of chromosomes during cell cleavage. They organised the positioning of the hereditary material/chromosomes within the cell and thus ‘decided’ on their fate during development. 2. Substances and processes in the plasma were important for the first phase of embryonic development before gastrulation. 3. Even more dramatic: the plasma was able to reorganise the chromosomes; it could change their size and composition. This process, called ‘Chromosomendiminution — chromatin diminution’, was observed for the first time by Boveri in 1887 in Ascaris, and examined in extensive cell lineage studies later on. Boveri showed in 1899 that only the germ cells did not undergo chromatin diminution. For him, this was a necessary finding to support Weismann’s germ line theory and his own theory of chromosomal individuality. The reorganisation of the chromosomes by the plasma during ontogenesis was crucial, as a process was needed which could explain an unequal distribution of hereditary material during ontogenesis so that cell differentiation could occur. It was a process of circular reasoning due to which chromosomal diminution could not happen in the germ line. Let me end with a kind of a-historic thought experiment using a bricolage of the Boveris’ findings of the plasma’s own activities. It would have been a very radical move to claim that the plasma could rearrange the chromosomes of the germ line as well.54 Boveri had postulated that chromatin diminution happened in species other than Ascaris, even if it was not visible under the microscope — so why not in the germ line, where one could not see it as well? Obviously, there was no way of conceptualising an interaction between plasma and chromosomes in heredity which was a co52

53 54

The nucleus (Kern) was only to be recognized ‘als ein Organ ..., welches dazu da ist, die Ausbildung des Larvendarms, des Skeletts usw. zu ermöglichen’ while ‘die spezifische Gestaltung dieser Prozesse, d.h. die Übereinstimmung mit der spezifischen Gestalt der Eltern, in anderen Teilen der Gameten ihre Bedingungen haben werden.’ [emphasis by Boveri]. Boveri (1918), p. 467. Moritz (1993). See for comparison the fate of the contemporary experimental system in hereditary research using the worm Planaria. Planaria did not have a visible germ line thus providing reasons for a completely different understanding of heredity as a process of metabolism, not linking it to a specific structure in the cell. The scientists using Planaria did not gain the social authority to efficiently compete with the Morgan group in the US. Mitman and Fausto-Sterling (1992).

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operative, non-hierarchical one, which would have made the egg cell much more powerful than the chromosomes of the sperm, and which would have resulted in a completely new understanding of ‘heredity’ and its science. For the social gender order there was obvious progress attached to the knowledge of genetics. In 1934 the Norwegian geneticist Otto Mohr enthusiastically praised it, claiming that the times of pure male genealogy were over: ‘one of the most far-reaching achievements of modern biology is the definite establishment of the fact, that men and women are genetically equivalent’. 55 For genetics, however, it might not be seen as a success story.

Helga Satzinger, Wellcome Trust Centre for the History of Medicine, University College London, [email protected]

55

Mohr (1934), p. 207.

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References Baltzer, Fritz. 1962. Theodor Boveri. Leben und Werk eines großen Biologen 1862-1915. Stuttgart: Wissenschaftliche Verlagsgesellschaft. Bleker, Johanna, ed. 1998. Der Eintritt der Frauen in die Gelehrtenrepublik. Zur Geschlechterfrage im akademischen Selbstverständnis und in der wissenschaftlichen Praxis am Anfang des 20. Jahrhunderts. Husum: Matthiesen Verlag. Boveri, Margaret. 1982. Verzweigungen. Eine Autobiographie. Herausgegeben von Uwe Johnson. München: dtv. Boveri, Marcella. 1903. "Über Mitosen bei einseitiger Chromosomenbindung." Jenaische Zeitschrift für Naturwissenschaften 37: 401-446. Boveri, Theodor. 1889. "Ein geschlechtlich erzeugter Organismus ohne mütterliche Eigenschaften." Sitzungsberichte der Gesellschaft für Morphologie und Physiologie München 5: 73-80. ———. 1902. Das Problem der Befruchtung. Jena: Gustav Fischer. ———. 1904. Ergebnisse über die Konstitution der chromatischen Substanz des Zellkerns. Jena: Gustav Fischer. ———. 1908. "Zellen-Studien VI. Die Entwicklung dispermer Seeigel-Eier. Ein Beitrag zur Befruchtungslehre und zur Theorie des Kerns." Jenaische Zeitschrift für Naturwissenschaften 43: 1292. ———. 1909. "Über die Beziehungen des Chromatins zur Geschlechts-Bestimmung." Sitzungsberichte der physikalisch-medicinischen Gesellschaft zu Würzburg: 1-10. ———. 1910. "Die Potenzen der Ascaris-Blastomeren bei abgeänderter Furchung. Zugleich ein Beitrag zur Frage qualitativ ungleicher Chromsomenteilung." Festschrift vom 60. Geburtstag Richard Hertwigs, Vol. 3. München: 133-214, Taf. XI-XVI. ———. 1913. Vortragsmanuskript. Nachlaß Theodor Boveri. Handschriftenabteilung der Universitätsbibliothek Würzburg. ———. [First edition in German: 1914]1929. The Origins of Malignant Tumors. London: Baillière, Tindall & Cox. Boveri, Theodor †. 1918. "Zwei Fehlerquellen bei Merogonieversuchen und die Entwicklungsfähigkeit merogonischer, partiell-merogonischer Seeigelbastarde." Archiv für Entwicklungsmechanik der Organismen 44: 417—471. Brush, Stephen G. 1978. "Nettie M. Stevens and the Discovery of Sex Determination by Chromosomes." Isis 69: 163-172. Churchill, Frederick B. 1970. "Hertwig, Weismann, and the Meaning of Reduction Division circa 1890." Isis 61: 429—457. Cremer, Thomas. 1985. Von der Zellenlehre zur Chromosomentheorie. Naturwissenschaftliche Erkenntnis und Theorienwechsel in der frühen Zell- und Vererbungsforschung. Berlin, Heidelberg et al.: Springer. Gaudillière, Jean-Paul and Hans-Jörg Rheinberger, eds. 2004. From molecular genetics to genomics: the mapping cultures of twentieth-century genetics. London, New York: Routledge. Gerhard, Ute. 1990. Unerhört. Die Geschichte der deutschen Frauenbewegung. Hamburg: Rowohlt. Gilbert, Scott F. 1978. "The Embryological Origins of the Gene Theory." Journal of the History of Biology 11: 307-351. Harwood, Jonathan. 1993. Styles of Scientific Thought. The German Genetics Community 1900—1933. Chicago: Univ. Chicago Press. Hausen, Karin. 1986. "Warum Männer Frauen zur Wissenschaft nicht zulassen wollten." Hausen, Karin and Helga Nowotny (eds.). Wie männlich ist die Wissenschaft? Frankfurt a. M.: Suhrkamp: 31-40. Keller, Evelyn Fox. 1995. Refiguring Life. Metaphors of Twentieth-Century Biology. New York: Columbia Univ. Press. Keller, Evelyn Fox. 2000. The Century of the Gene. Cambridge, London: Harvard Univ. Press. Kohler, Robert E. 1994. Lords of the Fly. Drosophila Genetics and the Experimental Life. Chicago: Univ. Chicago Press. Gilbert, Scott F. 1978. "The Embryological Origins of the Gene Theory." Journal of the History of Biology 11: 307-351. Lesky, Erna. 1951. Die Zeugungs- und Vererbungslehren der Antike und ihr Nachwirken. (Abhandlungen der Geistes- und Naturwissenschaftlichen Klasse Jahrgang 1950, Nr. 19. Akademie der Wissenschaften und der Literatur in Mainz.) Wiesbaden: Franz Steiner Verlag. Mitman, Greg and Anne Fausto-Sterling. 1992. "Whatever happened to Planaria? C.M. Child and the Physiology of Inheritance." Clark, Adele E. and Joan H. Fujimura (eds.). The Right Tools for the Job. At Work in Twentieth-Century Life Science. Princeton Univ. Press: 172-197.

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Mohr, Otto L. 1934. Heredity and Disease. New York: W.W. Norton & Co. Moritz, Karl B. 1993. Theodor Boveri (1862-1915). Pionier der modernen Zell- und Entwicklungsbiologie. Jena: Gustav Fischer. Moritz, Karl B. and Helmut W. Sauer. 1996. "Boveri’s Contributions to Developmental Biology — A Challenge for Today." International Journal for Developmental Biology 40: 27-47. Neumann, Herbert A. 1998. Vom Ascaris zum Tumor. Leben und Werk des Biologen Theodor Boveri (18621915). Berlin, Wien: Blackwell Wissenschafts-Verlag. Pycior, Helena M., Slack, Nancy G., and Pnina G. Abir-Am, eds. 1996. Creative Couples in the Sciences. New Brunswick, New Jersey: Rutgers Univ. Press. Pycior, Helena M., Slack, Nancy G. and Pnina G. Abir-Am, eds. 1996. "Introduction." In: Pycior et al. (eds.). Creative Couples in the Sciences. New Brunswick, New Jersey: Rutgers Univ. Press. 3-35. Rádl, Emanuel. [1909] 1970. Geschichte der biologischen Theorien in der Neuzeit, II. Geschichte der Entwicklungstheorien in der Biologie des XIX. Jahrhunderts. Leipzig: Wilhelm Engelmann. Reprint Hildesheim: Georg Olms. Rheinberger, Hans-Jörg and Jean-Paul Gaudillière, eds. 2004. Classical genetic research and its legacy: the mapping cultures of twentieth-century genetics. London, New York: Routledge. Rossiter, Margaret. 1993. "The Mathew Matilda [sic] Effect in Science." Social Studies of Science 23: 325341. Sapp, Jan. 1987. Beyond the Gene. Cytoplasmic inheritance and the Struggle for Authority in Genetics. New York: Oxford Univ. Press. "Special Issue ‘Universität — Frauen — Universitäten’." Feministische Studien 20/1. The Biology and Gender Study Group. 1989. "The Importance of Feminist Critique for Contemporary Cell Biology." Tuana, Nancy (ed.). Feminism and Science. Bloomington, Indianapolis: Indiana Univ. Press: 172—187. Wright, Margaret R. 1997. "Marcella O’Grady Boveri (1863-1950). Her Three Careers in Biology." Isis 88: 627-652.

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Hugo de Vries's transitions in research interest and method Ida H. Stamhuis1

I said, no- one deserves the title botanist who is not also a physicist and a chemist. (Hugo de Vries looking back to the beginning of his research career around 1870)

Assumptions in which way the hereditary characteristics are determined through the composition of their carriers are not yet made; this elaboration of the theory of heredity is unnecessary for the time being.2 (Hugo de Vries in 1889 about hereditary particles)

To get an impression of the context in which the modern ideas of heredity were embedded, I will concentrate on Hugo de Vries (1848–1935), the person who put his stamp on the emerging science of genetics, yet soon distanced himself from it. I will examine the pre–Mendelian stages in De Vries’s scientific development. ‘Context’ will in this case mean that I discuss Hugo de Vries’s early scientific interest and methodology, which was not in heredity and evolution but in plant physiology, and that I will connect that to his later research interest and methodology. Heredity was not a general concept from the beginning, but would slowly emerge; I will try to make plausible that something parallel was true with respect to the methodology that had to be used to study the concept of heredity. Looking at Hugo de Vries’s early work in plant physiology and comparing that to the – for him – most important later work in heredity and evolution, his Intracellular Pangenesis (Intracellulare Pangenesis) and his Mutation Theory (Die Mutationstheorie), the contrast in research methodology is striking: the first is what I will call ‘reductionist’, because an explanation of biological phenomena is sought with the help of physics, chemistry and quantitative reasoning, whereas the second does not try to do so, and is qualitative.3 In this essay my aim is to discuss the methodological aspects of De Vries’s first choice for plant physiology and then his transition from plant physiology to heredity and evolution. I will try to gain some understanding as to why he chose to work in these two fields in botany with such radically different approaches, and to discuss how he dealt with this contrast. I will address this problem by showing that originally it was not

1

2 3

I am grateful to Erik Zevenhuizen, who is writing a biography (in Dutch) on De Vries, for his critical response to my ideas, especially on the physiological foundation of De Vries’s ideas on heredity, and for his meticulous comments on the whole text. I thank my Free University colleagues (Ab Flipse, Teun Koetsier, Frans van Lunteren and Wijnand Rekers) for their comments during a discussion meeting. Thanks to Daniel Carroll for his correction of the English. See the more elaborate quotations and references in the sections ‘Change of Research Focus” and “Change of Research Method”. Reductionism has been extensively written about. Here I will take reductionism to mean the explanation of biological phenomena with the help of a so–called ‘underlying’ less complex field of knowledge: chemistry, or physics, or not an ‘underlying’ but nevertheless less complex field of knowledge: mathematics. In the case of mathematics a characterization like ‘modelling’ is perhaps better, but to make clear that in all cases the effort is to try to explain biological phenomena with the help of a simpler field of knowledge, I use one concept in all cases (Jongeling 1997).

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obvious that the methods in these two fields would be so different. For De Vries the original methods turned out to be unsuccessful and ultimately he would no longer consider these as useful. To gain deeper insight into De Vries’s research choices, it will also be helpful to sketch his background: the milieu he came from, his secondary school time, and the atmosphere at the university in Leiden, where he studied botany. I will start by dealing with these aspects. After discussing his change of research focus, I will include a short characterization of his theory of heredity and I will discuss a number of criticisms levelled at De Vries’s Intracellular Pangenesis, because he did not explain these ideas within the framework of his plant physiological work. In addition I will show that he tried out a quantitative model of heredity, but had to abandon it due to the resulting inconsistencies. In the next section I will argue that in the 1890s he again tried out a reductionist approach, namely a statistical/probabilistic one. And although this approach resulted in 1896 in the rediscovery of the Mendelian Laws, for De Vries it was not a complete success, since that approach was not compatible with his theory of heredity. I discuss only an aspect of the background of De Vries’s research choices. There is an erudite essay about De Vries’s transition from plant physiology to genetics (written in Dutch) by Bert Theunissen, from which I will now extract a few observations. 4 Theunissen discusses the views of De Vries and other Dutch scientists in that time on the importance of science for the progress of society, because that may have influenced their research choices. Their opinion was that science should have a civilizing influence. Moreover science could help to solve problems of society, both social and economic. According to De Vries, botany could very well serve that goal. Within botany the study of heredity was a possible choice, but other choices were also possible. That De Vries nevertheless chose heredity was for other reasons. Theunissen suggests that for De Vries the reason for leaving the field of plant physiology, the collaboration with his colleague at the University of Amsterdam, the physical chemist Jacobus H. van’t Hoff, will have been a decisive factor. Aided by De Vries’s work on osmosis, Van’t Hoff was able to formulate his theory of dilute solutions for which he and Svante Arrhenius would later receive a Nobel Prize. De Vries subsequently felt that he had to redefine the demarcation between each other’s sphere of activity. Why, then, did De Vries choose to focus on genetics? Theunissen points, just as I will do, to De Vries’s early interest in this field. Theunissen also argues that De Vries’s ultimate aim, to be able to control mutations, was in accordance with the more general idea of Dutch scientists of the ’improvability’ of society with the help of science.5 But this would motivate De Vries later in his life than the period I discuss in this paper. For my discussion of De Vries’s research choices and methods, I will mainly rely on existing scholarship. In addition, this material will be augmented by the rich primary resource represented by the preserved 450 letters De Vries exchanged with his colleague and friend Jan Willem Moll (1851–1933).6 Little, is available in English about De Vries’s youth and student time, but in Dutch a number of biographies have been written, which will be useful for my aims. 7 A contemporary 4 5 6 7

(Theunissen 1992). ‘improvability’ is a translation of the Dutch ‘maakbaarheid’. (Stamhuis 1995). The collection of letters is in the University Library in Groningen. Moll’s letters are copied in three copy books, to which I refer as CB1, CB2 and CB3. (Van der Pas 1970) and (Zevenhuizen 1998a) are biographies in English; (Heimans 1948), (De Veer 1969), (Smit 1980) and (Visser 1992) in Dutch.

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biography was published in 1900 by one of his first pupils, the later professor of botany in Utrecht, Friedrich A.F.C. Went (1863–1935).8 We can use this text, supplemented with modern biographies, to gain deeper insight into his boyhood, student time, the time spent as a researcher in plant physiology and his turn to heredity. Another interesting source is a short biography written by his friend and colleague Jan Willem Moll on the occasion of his seventieth birthday. 9 Moll had been a colleague and friend of De Vries since 1872 and commented upon many concepts of De Vries’ scientific publications. Although Moll’s comments on De Vries’s work were sometimes very critical, to the outside world he always defended and supported him. For the discussion of the significance of the reductionist approach in De Vries’ work on heredity, I draw on two of my own publications.10

Life Sketch Hugo de Vries was born in 1848 into a family of intellectual and political distinction. His father was successively a member of the ‘Raad van State’, an important national advisory council for the government, a minister of Justice and a member of the ‘Tweede Kamer’ (Parliament); Hugo’s uncle was professor of Dutch language at Leiden. His maternal grandfather was an archaeologist. So his family was intellectual, but more in the humanities than in the natural sciences. Notwithstanding this family interest, it is told that even as a primary school pupil Hugo was interested in nature. He lived in Haarlem near dunes, woods and the sea. He regularly took long walks in these woods and dunes and studied the flora native to these natural environments. At the age of twelve he participated in a competition initiated by the Dutch Society of Agriculture to make a collection of hundred dried plant species from the surroundings of Haarlem. 11 He started to compose a herbarium and was awarded an honourable mention. During holidays his family made trips to various parts of the country and it is said that on these occasions his brother made drawings and Hugo looked for plant species. During his time at secondary school he and his family moved to The Hague, the Dutch administrative capital, because of the political career of his father. At secondary school the classics played an important role, and Hugo would later say that he was not happy with the neglect of modern languages and the natural sciences. In 1866 the period of university study came (fig. 1). That he would go to university was obvious given his milieu, but not his wish to study natural sciences, especially not botany; he registered to study natural philosophy (‘philosophia naturalis’). Apparently his interest in natural history was not in accordance with his family’s interest. Moreover, there were no ‘distinguished’ jobs after such a study, the available few in secondary school teaching not being considered ‘distinguished’. Moreover, botany was connected to agriculture and, according to his biographer Went, in this period of the upcoming industrial revolution, industry enjoyed a higher standing in his family than agriculture.12 Ultimately, though, Hugo’s family accepted his choice.

8 9 10 11 12

(Went 1900). (Moll 1918). (Stamhuis et al. 1999); (Stamhuis 2003). ‘Competition’ is a translation of the Dutch ‘prijsvraag’. (Went 1900), 266.

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Fig. 1. Student Hugo de Vries.(Archive Hugo de Vries, University of Amsterdam). In 1859 Darwin had published the Origin of Species. De Vries’s teacher, the professor of botany W.F.R. Suringar, did not agree with Darwin’s ideas on evolution. Hugo de Vries therefore did not hear about it from his teacher but read Darwin’s book for the first time in 1868, his second year of study, in a German translation he had bought at an auction. According to Went, he and his fellow students discussed it, abandoned the idea that species could not change into others, and became convinced adherents of Darwin’s evolutionary theory.13 In the meantime Hugo de Vries encountered another way of practicing botany than he learnt from his teacher Suringar, who was working in the field of systematics and wanted Hugo to study and describe lichens. In the same year as he studied Darwin’s work, Hugo studied the new book 13

(Went 1900), 267–268; (Visser 1992), 160–161.

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Lehrbuch der Botanik by the German Julius Sachs, in which the new plant physiological approach of botany was discussed. De Vries also found an opportunity to apply this new method, because the University of Groningen held a competition entitled “What is known of the effect of warmth on plant roots?” He participated in this contest and did the necessary experiments at home, because at the university in Leiden there was no opportunity to carry out these kinds of investigations. He used his answer in the Groningen Prize Contest, for which he was awarded the Gold Medal in 1869, for his thesis, and was awarded his PhD in 1870. To explain the physiological interest of De Vries notwithstanding his education in systematics by Suringar, his biographer Went informs us that De Vries was not only interested in the outside but also in the inside of the plant: in its anatomy and vital functions. 14 I would add that Hugo might have been an ambitious young man, who will soon have noticed that his teacher’s interest was old fashioned. He may well already have had greater ambitions. The most obvious opportunity to do something which was new and which was considered of real scientific value was the new plant physiology. The practitioners of plant physiology could be found especially in Germany, the country on which Dutch scientists were mainly oriented. Moreover, Sachs had discussed an applicable research program in his book. De Vries would later also refer to the teaching of his physics professor Rijke; this professor will have played a stimulating role in choosing a research field in which physics was essential. After gaining his PhD, Hugo de Vries left for Germany. His first visit to the old Wilhelm Hofmeister in Heidelberg was not successful, but subsequently, in 1871, he went to the young Julius Sachs (1832–1897) in Würzburg. That was the beginning of a fruitful relationship with Sachs and with the new botany that Sachs represented (fig. 2).15 However, De Vries also had to earn a living and he therefore spent the next four years as a secondary school teacher in Amsterdam. During this period he frequently spent his entire summer break in Würzburg. In April 1875 he was commissioned by the Prussian Ministry of Agriculture to write monographs on agricultural plants. He was able give up his job as a teacher and settled in Würzburg. In 1877 he became a ‘Privatdozent’ at the University of Halle, and published his Habitilationsschrift About the mechanical causes of cell stretching (Über die mechanischen Ursachen der Zellstreckung). De Vries noticed that the professor of botany in Halle was not happy with his presence. Moreover, student interest in his lectures was minimal; it is therefore not surprising, that De Vries was not very enthusiastic about the job in Halle.16 He was therefore happy when in the next year, in 1878, he was appointed lecturer at the University of Amsterdam where he became a full professor in 1881. 17 14 15 16

17

(Went 1900), 267. Letter from De Vries to Moll, 24–04–1875 from Würzburg “I am now busy with micro- chemical work” (Ik ben thans druk met microchemischen arbeid bezig); see also the letter of 28–06–1875. (Went 1900), 271; (Visser 1992), 161–162; Letter from De Vries to Moll, 22–05–1877 from Halle “Gradually I start to notice (…) that it will not be easy for me (..). Don’t tell anyone about this.” (Ik begin langzamerhand te merken (…) dat ik het hier niet gemakkelijk zal hebben (…) vertel hiervan maar aan niemand iets). Letter from De Vries to Moll, 09–09–1877 from Den Haag “I have been proposed for the post of lecturer in plant physiology in Amsterdam. Salary 1500 Dutch guilders.” (Ik ben tot lector in de plantenphysiologie te Amsterdam voorgesteld. Tractement Fl. 1500). Letter 19–09–1877 from Würzburg “This morning I received the official announcement of the Municipal Executive of Amsterdam about my appointment” (Heden morgen ontving ik de officieele mededeling van B. en W. van Amsterdam omtrent mijn benoeming).

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In 1896 he also became the director of the Hortus Botanicus of the university. In the Netherlands he became a ‘Man of Significance’, even before the so–called rediscovery of Mendel in 1900 and the publication of his Mutation Theory (in 1901 and 1903).18 Notwithstanding various job offers from Dutch and foreign universities, he remained in Amsterdam until his retirement in 1918.

Fig. 2. Julius Sachs and Hugo de Vries left and right at the front. (Archive Hugo de Vries, University of Amsterdam).

18

Went published in 1900 his biography of De Vries in the series ‘Mannen en Vrouwen van Beteekenis in onze Dagen’ (Men and Women of Significance in our Days).

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Change of research focus I will now discuss his research interest and focus attention on the apparently great transition from plant physiology on the one hand, to heredity, evolution and mutations on the other. 19 It is striking that De Vries’s colleague and friend Moll distinguished three stages in De Vries’ work, of which the first two are in plant physiology and the third in heredity, evolution and mutations. 20 De Vries’s pupil Went did not set De Vries’s work on plasmolyses apart from his other plant physiological work, for example on the movements of plants, as clearly as Moll did. Moll gave De Vries’s work on plant physiology more weight than we would expect. Moll stated that this last work took place during a so–called preparatory period, and his work on plasmolyses during De Vries’s second period, which Moll characterizes as ‘very important for science’. It is strange that Moll more or less distinguished two periods of plant physiological work, since from the list of De Vries’s publications it follows that these two kinds of work cannot be distinguished chronologically. An explanation may be that De Vries’s chemical physiological work on turgor, plasmolyses and isotonical coefficients (fig. 3) made a greater impression than his work on the growth of plants, the movement of plant vines, the causes of tree rings and of transport in the plant, and that for that reason Moll felt the need to clearly distinguish between the two. 21

Fig. 3. An example of De Vries’s reductionist plant physiological work. Cephalaria leucantha. 1) Turgescent young cell; 2) In 4% nitric acid solution; 3) In 6% nitric acid solution; 4) In 10% nitric acid solution. (De Vries 1918-1927) Vol. 1, 396. (Library Free University Amsterdam). Went as well as Moll discuss De Vries’s work on heredity and evolution as something different from plant physiology. Went asserted that as soon as De Vries became acquainted with Darwin’s 19 20 21

See also (Visser 1992) and (Theunissen 1992). (Moll 1918). (Moll 1918), 2. About difference in prestige of different methods in plant physiology see (De Chadarevian 1996).

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ideas on evolution in 1867, they made a strong impression on him. 22 To support this view, Went pointed to the theses that De Vries added to his dissertation on 1870 in which he defended evolutionary theory, although he knew that his supervisor would oppose them. One of these theses was: The hypothesis of Pangenesis [...] cannot explain the variability of the species. 23 Went even stated that “since that time the great questions of variability and heredity absorbed De Vries’s mind and never released him. Although it is true that (…) through the influence of Sachs, for a time he went into an entirely different field of investigations.”24 Before Went started to discuss De Vries’s work on ‘heredity and variability’ he stated25 Certainly few have studied Darwin’s work so thoroughly as De Vries. When he was conducting his investigations in the field of mechanical physiology, (…) his mind also remained engaged in the great questions of the theory of descent, and he soon collected all facts in this field that he encountered in the literature. As soon as he had his own field for experiments at his disposal, directly some of the experiments were dedicated to questions of variability and heredity. We thus see that De Vries was driven more and more in this new direction, especially since during the academic year 1881–82 he gave a course in the theory of descent for students of botany. I believe that all who attended those lectures – not only philosophers, but also medical students from various academic years – will not easily forget the clear way in which such a rich subject matter, which was difficult to gather, was discussed by a convinced adherent of Darwinism.

Went must have attended these lectures. Much later, in an interview published in 1935, De Vries himself commented upon his original views on the role of physics and chemistry for physiology and botany, and his later transition of research focus.26 He said: I felt in that time that physics and physiology were so closely connected that they were only different communications about the same events, and I also said that to Hofmeister. I said, no–one deserves the title botanist, who is not also a physicist and a chemist. Then I had something in my head that had not yet a name. You must realize that Van’t Hoff and Arrhenius were then still schoolchildren. And names like physical chemistry, chemistry, biochemistry, colloids, cell physiology and the like were not used until thirty to forty years hence. (...)When I came back in Amsterdam, I wanted to continue with what is now termed physical chemistry of the cell. I felt it as a kind of responsibility towards my esteemed teacher of physics Rijke. 22 23 24

25

(Went 1900), 167–268; (Visser 1992), 160. “De hypothese van de pangenese kan de variabiliteit der soorten niet verklaren.” (Went 1900), 268. “Sedert dien tijd hebben de groote vraagstukken van erfelijkheid en variabiliteit zich meester gemaakt van den geest van De Vries en hem niet meer losgelaten. Wel is waar betrad hij (…) onder den invloed van Sachs, een tijdlang met zijn onderzoekingen een geheel ander veld.” (Went 1900), 289. “Weinigen hebben zeker Darwin’s werken zoo grondig bestudeerd als De Vries. Ook toen hij de onderzoekingen op het gebied der mechanische physiologie verrichte, (…) bleef zijn geest zich bezig houden met de groote vraagstukken der afstammingsleer en hij verzamelde spoedig alle feiten, hierop betrekking hebbende, die hij in de literatuur aantrof. Zoodra hij beschikken kon over een eigen terrein voor proefnemingen, werden dadelijk eenige van die proeven gewijd aan vragen over erfelijkheid en variabiliteit. Zoo zien wij De Vries meer en meer gedreven in deze nieuwe richting, vooral sedert hij in den cursus 1881–82 een college voor botanici gaf over de afstammingsleer. Ik geloof, dat al degenen die dit college volgden – en het waren niet alleen philosophen, maar ook medici uit verschillende studiejaren – niet licht de heldere wijze zullen vergeten, waarop een zoo rijke stof, die moeilijk bijeen te garen was, door een overtuigd voorstander van het Darwinisme werd behandeld.”

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But nobody knew exactly what I intended; the child had no name, the institute was not equipped. For students of medicine, I had to teach general physiology and theory of heredity, and this confronted me with my old love, evolutionary theory. Only during holidays was I able to do experimental cytology. However Ms. De Vries had three young children and wanted to spend the summer in the country. So I abandoned osmosis (...) and I started with what is now termed genetics.

He suggested that he ceased work on plant physiology after his appointment in Amsterdam in 1878, but that is not correct. Although he had also started with experiments in variability and evolution on a modest scale, he continued to publish in plant physiology until 1889, the year of the publication of his Intracellular Pangenesis. His correspondence with Moll clearly reflects that after 1889 his physiological work abruptly came to a halt. In 1887 and 1888 he wrote about the tonoplast, about permeability of the protoplasm, about isotonical coefficients, plasmolyses and permeability, about the influence of gravitation on protoplasm and the lowering of the freezing point of raffinoses.27 In 1887 he started to write about sunflower seeds, about numbers of rows of seeds in corn ears, about conifer seeds and about monstrosities in plants. 28 In a letter dated May 19th 1888 his interest in heredity and variability became more explicit: he wrote about August Weismann’s work, stating that he appreciated Weismann’s germ plasm theory. 29 In September 1888 he mentioned for the first time his intended booklet on heredity and in October 1888 he sent the first part to Moll for comment.30 Subsequently we find no further references to plant physiological work. I think that it is well founded to conclude that in his youth De Vries loved to walk in nature and to study plants in the countryside. This passion endured throughout his lifetime (fig. 4). He was however not satisfied to carry out investigations in the old fashioned systematics, but was ambitious and eager to work on new and modern botanical questions. Notwithstanding the opinion of his teacher Suringar, he became a convinced adherent of Darwinian thinking. 26

27

28

Quotation from the Prager Presse, June 2, 1935. The German part I copied from (Theunissen 1992), 112; the Dutch part I copied from (De Veer 1969), 18. “Ich fühlte schon damals dass Physik und Physiologie irgendwie nahe zusammenhängen, dass sie nur verschiedene Berichte über ein und dasselbe Geschehen vorstellen und habe es auch Hofmeister gesagt. Niemand sollte Botaniker heissen, der nicht zugleich Physiker und Chemiker ist, sagte ich. Ich hatte damals etwas im Kopfe, das noch keinen Namen trug. Van’t Hoff und Arrhenius waren damals noch Schulbuben, müssen Sie wissen. Und Namen wie Physikalische Chemie, Biochemie, Kollide, Zellphysiologie und dergleichen mehr sind erst nach dreissig bis vierzig Jahren geprägt worden. (...). Als ich nach Amsterdam kam, wollte ich zuerst selbstverständlich in dem, was man heute Physikalische Chemie der Zelle nennt, fortfahren. Ich fühlte es übrigens halb als Gebot meines geliebten verstorbenen Lehrers [P.L. Rijke]. Aber niemand wusste, wass ich eigentlich vorhabe, das Kind hatte keinen Namen, das Institut war nicht eingerichtet, rings herum stand zu den Versuchen nichts Lebendes zur Verfügung:” “Voor de medicijnstudenten moest ik college geven in de algemene fysiology en erfelijkheidsleer, en dit confronteerde mij weer met een oude liefde, de evolutietheorie. Alleen in de vacanties zou ik experimentele cytologie hebben kunnen doen. Maar mevrouw De Vries had drie kleine kinderen en wilde de zomer buiten doorbrengen. Aldus heb ik de osmose verlaten (...) en ben begonnen aan datgene wat men tegenwoordig genetica noemt.” Letters from De Vries to Moll, from Amsterdam, 07–01–1887; from Amsterdam 13–12–1887. Letter from De Vries to Moll, from Amsterdam 12–01–1888; from Amsterdam 24–02–1888, Letter from De Vries to Moll, 04–03–1888 from Amsterdam. De Vries informs Moll that research of Arrhenius affirmed the previous results of De Vries. About sunflower seed and corn: Letter from De Vries to Moll,: from Amsterdam 20–09–1887, 27–10– 1887; Letter from De Vries to Moll, from Amsterdam 27–03–1888.

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However, for the time being he was absorbed in successful plant physiological work in which Sachs became an influential factor and of which the Groningen Prize Contest marked the beginning. He may at that time have had the impression that there was not yet an interesting Darwinian research program. However, when circumstances made it easy to grow plants and his mind became again engaged in heredity and evolution because he lectured in it, and perhaps because it became clearer how to do interesting work in heredity and variability, he did not hesitate to change his research focus and to start to investigate questions of heredity. Theunissen also pointed to the possible roles of the promising applicability of this new field and the need for the redefinition of the demarcation line between him and his colleague, the physical chemist Van’t Hoff. After the publication of his Intracellular Pangenesis his change of research focus was complete.

Fig. 4. Hugo de Vries with students on a botanical excursion. (Archive Hugo de Vries, University of Amsterdam).

29 30

Letter from De Vries to Moll, from Hilversum, 19–05–1888; again on Weismann’s work in a letter dated 07–06–1888 from Hilversum. Letter from De Vries to Moll from Amsterdam 04–10–1888.

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De Vries’s theory of heredity To understand the discussion of the change of De Vries’s research method, it is necessary to know something of the content of his theory of heredity.31 I will sum up the most important aspects. According to Intracellular Pangenesis the properties of visible characters depend on small invisible particles, which De Vries called 'pangenes'. They are either inactive or active, and they are able to grow and multiply in both states. Their activity is dependent on the type of cells in which they are located. There are two kinds of cell lines. There is a line of cells from the fertilized egg cell to the germ cells of the newly formed organism. This cell line is called the germ line; the other lines are called somatic lines. The cells of the somatic lines develop into the cells forming the organs of the organism. De Vries stated that pangenes are usually inactive or latent in the germ lines, and develop their greatest activity in the somatic cells. Differentiation of organs is due to the fact that individual pangenes, or groups of pangenes, develop more strongly than others. In the nucleus of each cell of the organism all types of pangenes of the individual are present, in the cytoplasm only those that will become active. In the nucleus most pangenes remain latent. They become visible and active only in the cytoplasm. Pangenes multiply in the nucleus, partly for the division of the cell nucleus, and partly in order to be transported later to the cytoplasm. There has to be transport from the nucleus to the cytoplasm. Transmission of pangenes is the function of the nucleus, their development the function of the cytoplasm. In the nucleus, most pangenes only have to multiply. In the cytoplasm pangenes will continue to multiply (usually more so than in the nucleus). Sometimes they will change there from active to latent, or sometimes the other way around. Some will become active immediately after their arrival, others later. The development of a character requires a minimum number of active pangenes of the same kind. When the number of pangenes becomes larger, the expression of the character will become stronger. If the numbers of certain pangenes are reduced, the corresponding visible property will be only weakly developed; if their number is strongly reduced, they become latent. Therefore, the number of similar pangenes is very important. In the cytoplasm, this number is decisive for the function of the various organs, in the nucleus for the hereditary force. Pangenes do not represent morphological parts of the organism, or cells, or parts of cells, but specific individual characters. They can vary independently of each other. De Vries stressed the independence of hereditary properties. Hybridisation is performed in order to transfer properties from one variety to another. Therefore the study of hybrids is of great importance. Hybrids show that the nature of a species is not unitary. Properties of hybrids can be as clearly distinguished and are as constant as those of pure species. Heredity must have a material basis, which can be none other than the living protoplasm. In every cell division two new pangenes arise which are identical to the original one. In every cell division all types of pangenes present go, as a rule, to both daughter cells. Exceptions to this rule are the starting point of the emergence of varieties and species. There are two kinds of variability. Firstly, the numbers of pangenes may vary in what is called 'fluctuating variability'. Secondly, during successive cell divisions, pangenes may change their nature slightly, or even considerably. 31

(Stamhuis et al. 1999), 243–244.

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Change of research method In this section I will explore the thesis that Hugo de Vries’s Intracellular Pangenesis can be considered as a transition; in various cases a physiological approach was suggested, but it had to be abandoned, because it was not possible to explain the complicated hereditary phenomena in that framework (fig. 5).

Fig. 5. Drawing of Rubus fruticosus laciniatus from The Mutation Theory (The original German edition, Vol. 2, p. 689) The caption is: ‘The distinguishing feature of the variety, the deep indentation of the leaf edge, expresses itself in the leaves as well as in the petals. Both phenomena are expressions of the same material carrier of the internal character’. (Archive Hugo de Vries, University of Amsterdam). Let me start by making clear that it is not true that for his Intracellular Pangenesis De Vries abandoned a physiological approach, or that he put it aside. Quite the contrary is the case. Throughout the booklet he was very clear that his theory had a physiological basis and that his hereditary carriers, which he called pangenes, were morphological particles. However, since it was mostly not yet possible to connect the hereditary characteristics with the physiological material basis, his opinion was that, to enable progress in the field of heredity, it would be better to put considerations about that basis aside for the time being. I will refer to a few passages of his booklet in which he put this view into words. In the introduction to this booklet he stated that he had to “explore the basic ideas of pangenesis.” His starting point was that “the physiology of heredity” could be investigated during the “microscopic investigation of the cell division and the

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fertilization, and the morphological substrate.” He continued: “One must not try to explain the morphological details of these processes, because our knowledge is still far too restricted.” De Vries’s opinion was that if such an explanation was nevertheless possible, then that should of course be given, but that was only seldom the case.32 In the chapter in which he discussed the hypothetical carriers of species characteristics, he asserted that33 an assumption about the nature of these hypothetical units (…) must be connected to their relation to the hereditary characteristics. Assumptions in which way the hereditary characteristics are determined through the composition of their carriers are not yet made; this elaboration of the theory of heredity is unnecessary for the time being.

That he always realized that his theory had a physiological basis is also clear from the fact that later he talked at various times about the “physiology of heredity”.34 Although he explicitly formulated his opinion in his booklet on heredity of 1889, his readers would not be satisfied with it. They would criticize aspects of his theory that he could not explain physiologically or that were even implausible from a physiological point of view. I will cite a few examples. Moll commented upon a concept version of Intracellular Pangenesis in 25 written pages and an accompanying letter.35 In the accompanying letter he criticized what he called the “hypothesis of intracellular pangenesis itself.”36 Moll's main criticism concerned what he considered to be an inconsistency in De Vries’s hypothesis. De Vries had argued that the pangenes moved from the nucleus to the other parts of the cell in protoplasm streams, the normal route of transport in the plant.37 Moll's comment was38: But now the reader will proceed further on this road. He remembers that H. de V. is himself the man who considers the protoplasm movements as the means of transport for food through the entire plant. He thinks of the ppl [protoplasm] connections between the cells, a fact which is as well established as the many facts you have used, and therefore he feels forced to assume a transport of pangenes through the entire plant and thinks that in this way he acts completely in your spirit.

32

33

34 35 36 37 38

“den Grundgedanken der Pangenesis (…) auszuarbeiten” “die Physiologie der Erblichkeit” “die mikroskopische Erforschung der Zelltheilung und der Befruchtung und das morphologischen Substrat”. “Nicht die morphologischen Einzelheiten jener Vorgänge soll man zu erklären suchen, dazu ist unsere Kenntniss noch viel zu beschränkt.” (De Vries 1889), 6–7. “Eine (…) Annahme über die Natur jener hypothetischen Einheiten (…) bezieht sich auf ihre Beziehung zu den erblichen Eigenschaften. In welcher Weise diese durch den Aufbau der Träger bestimmt werden, darüber werden bis jetzt keine Annahmen gemacht, denn auch dieser Ausarbeitung bedarf die Theorie der Vererbung vorläufig nicht.” (De Vries 1889), 36. e.g. (De Vries 1897), 63. Copy of the letter from Moll to De Vries: CB1, 168 ff., October 15, 1888. See (Stamhuis 2003), 125–130. “de eigenlijke hypothese der intracellulaire pangenesis zelve”: copy of the letter from Moll to De Vries, CB1 168 ff., October 12, 1888. (De Vries 1889), 141. “Maar nu gaat de lezer op dien weg als vanzelf verder. Hij herinnert zich dat H. d. V. zelf de man is, die de protopl–bewegingen als middel van vervoer voor voedsel enz. door de geheele plant beschouwt. Hij denkt aan de ppl verbindingen tusschen de cellen onderling, een feit even vast staande als vele feiten die je gebruikt hebt en hij ziet zich dus genoodzaakt een vervoer van pangenen door de geheele plant heen aan te nemen en meent daardoor zelfs zeer in je geest te handelen.”

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De Vries’ earlier statements about transport in the plant by protoplasm streams implied that pangenes could not only move from the nucleus to the protoplasm, but also from the protoplasm to the nucleus and even from one cell to another. Moll stressed that accepting this consequence would mean that De Vries’s theory of heredity did not essentially differ from Darwin's provisional intercellular theory of pangenes. “He [the reader] will think that it would have been better if you had made it clear from the start that you accept Darwin's pg [pangenesis] entirely and now want to show that the more recent results are entirely in accordance with it.” 39 Moll formulated his opinion rather confrontationally: “There is no reason to assume that pangenes are able to leave, but not to enter the nucleus.” But although this reasoning was in accordance with his physiological work, De Vries did not accept this conclusion. In Moll’s words: “In one word, this closure of the nucleus to its former inhabitants is an auxiliary hypothesis, which you frame because you have said from the start that you don't want to have anything to do with ‘movable pangenesis' and with heredity of acquired characters, and you feel now that without this hypothesis you return to these ideas.” We do not have de Vries’s immediate reaction to this criticism, but in his booklet he discussed the problem briefly.40 In a section entitled “Comparison with Darwin's Transport Hypothesis” (“Vergleichung mit Darwin's Transporthypothese”) he admitted that on the basis of the physiological knowledge available to Darwin there had been no reason to doubt the possibility of transport of a material carrier of hereditary information from one cell to another. He went on to say that, especially thanks to Weismann’s convincing arguments that acquired characters are not hereditary, it had become clear that such transport was very improbable. So he put the physiological approach aside in favour of hereditary evidence. An example of the critical reception of Intracellular Pangenesis is a review in the Botanische Zeitung41 in 1881 written by Georg Albrecht Klebs, who had studied with Julius Sachs in Würzburg.42 The criticism centred around the nature of pangenes. Klebs concluded that pangenes must be a kind of imprint of the cell organs, but he complained that it was not explained how they were able to perform their activity. He wondered what exactly the difference was between active and inactive pangenes and upon what that difference was based. Nor was it explained how it was possible that the functioning of a pangene depended entirely on its location: in the cell nucleus it caused the force of heredity and in the protoplasm it was responsible for the physiological activity. In addition, Klebs found the idea of the latency of pangenes unclear, and remarked that the claim that the level of expression of characters depended on the number of pangenes led to dubious consequences. Klebs criticized De Vries because he had simplified a complex issue and Klebs wrote that it was not difficult to formulate a few questions that would be difficult to answer. For example, De Vries’s theory did not explain the essence of an organism, the connection of all its parts, and their union into one entity. De Vries did not explain much at the cell level either, for example, how 39

40 41 42

“Hij zal dus meenen, dat je beter gedaan hadt met voorop te zetten, dat je Darwin's Pg geheel aanneemt en hier aan wilt toonen, dat de latere resultaten van het onderzoek daarmede geheel in overeenstemming zijn.” “Er is geen enkele reden om aan te nemen, dat pangenen wel uit, maar niet in den kern kunnen gaan.” “In één woord deze geslotenheid van den kern voor zijn gewezen inwoners is een hulphypothese, die je opstelt, omdat je van den beginne al gezegd hebt, dat je met de bewegelijke pangenesis en erfelijke verworven eigenschappen niets te maken wilt hebben en nu voelt dat je er zonder die hypothese weer toe terugkeert.” (De Vries 1889), 142–146. (Klebs 1889). See (Stamhuis 2003), 137–138.

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the cell is generated and how pangenes influence each other in the cell. Because of these deficiencies, Klebs concluded, De Vries’ theory would probably not be generally accepted. The criticism of another commentator, Cornelis Adrianus Pekelharing, professor of physiology at the medical faculty of the University of Utrecht, shows parallels with Klebs’s criticism. De Vries had discussed his ideas with Pekelharing and on that occasion his concept of latent characters had also proved to be rather controversial.43 With reference to that occasion he wrote to Moll:44 I cannot compromise on P.'s objection against latent characters, it makes it even more difficult to say what I mean. I sense that I am usually unsuccessful in this anyway. That the nucleus is the storehouse of all types of pangenes was what I meant to say, for example. However, I didn’t manage to say it. I wish rather that I had not started it.

The basis of the criticism by Pekelharing, Klebs and Moll was that the claims of De Vries’s theory were not explained physiologically. Moll asked how it was possible that pangenes can leave a nucleus but cannot return to it, or move to other cells. It was a well–known physiological fact that transport within the plant takes place by protoplasm streams, which stream into and out of the nucleus and from cell to cell. Movement only from the nucleus to the cell could not be explained. Pekelharing’s and Klebs’s criticisms of the latency of characters can be understood analogously. De Vries needed the concept of latency. He had to argue that all types of pangenes are present in all cells. For the explanation that nevertheless cells develop differently, he needed the concept of latency. However, the physiological questions remained unanswered: how it was possible that in some cells a pangene (or character) did not develop activity while in other cells it became active; how some pangenes in the same cell became active while others remained latent; and also, how it was possible that their activation depended on their position within the cell: in the nucleus or in the protoplasm. De Vries did not explain what physiological processes were responsible for this; the only reaction he could give was that these concepts could explain important hereditary phenomena and that therefore he believed that they were true. De Vries had already come to the conclusion that in some cases he could not use the usual physiological approach to explain hereditary phenomena. In his booklet on heredity he had also put that into words. However, others confronted De Vries with it, because they experienced De Vries’s argumentations as unsatisfying, inconsistent and incomplete.

Calculating offspring character values In Moll’s detailed comment on a concept version of Intracellular Pangenesis there is a passage, which shows that De Vries tried out a quantitative model of heredity, but that he could not

43 44

In his detailed comment on Intracellulare Pangenesis Moll also referred to Pekelharing's criticism of the latency of characters. Copy of comment from Moll to De Vries, especially 170 and 172. “P's bezwaar tegen latente eigenschappen kan ik niet toegeven, ook maakt het het nog veel moeilijker om te zeggen wat ik bedoel. Ik bespeur dat dit toch veelal mislukt. Dat de kern is de bewaarplaats van alle soorten van pangenen, en dat de questie van de inactiviteit bijzaak is, was bv mijn bedoeling. Doch 't was mij niet gelukt dat te zeggen. 'k Wilde wel dat ik het niet begonnen was.” Letter from De Vries to Moll, November 1, 1888.

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maintain it, because it resulted in inconsistencies.45 Unfortunately we do not have De Vries’s concept at out disposal, only Moll’s response. The relevant parts of Moll's comment are: 46

I am not sure whether this calculation contributes to clarity. If 1 on p. 54 is right, then 2 must be right, as I changed it above, and then at 3 p. 55 reinforcement has to take place. If this is not in accordance with the facts, then that is proof that your formula is wrong and then I would rather delete it. On second thoughts it is like this. In case 2 one gets an individual with the character in question above or below the zero point. In case 3 with the character above the zero point. Like this:

I think that these are the results of your calculation. Your further calculation on p. 55 I do not understand. Doing that you forget that by zero point (see p. 54) you mean the mean character if all individuals of the culture are mixed. And if you want to stick to this, I don't see what

45 46

See (Stamhuis et al. 1999), 244–246. “Of deze berekening nu wel tot de duidelijkheid bijdraagt weet ik niet. Als 1 op p. 54 waar is, dan moet 2 waar zijn, zooals ik het boven veranderde, en dan moet bij 3 p. 55. versterking plaats hebben. Als dit nu niet met de feiten overeenkomt dan is het een bewijs dat je formule verkeerd is en dan zou ik die liever weglaten. Bij nader inzien is het aldus. In geval 2 krijgt men een individu met de bedoelde eigenschap meer of minder dan het nulpunt. In geval 3 met de bedoelde eigenschap hooger dan het nulpunt. Aldus: [And then the three cases of crossing schemes follow] Dit zijn dunkt mij de uitkomsten van je som. Your further calculation on p. 55 I do not understand. With that you forget that with the zero point (see p. 54) you mean the mean character if all individuals of the culture are mixed. En wil je je daaraan houden, dan zie ik niet in wat een verdere berekening meer geeft dan het bovenstaande, terwijl de lezer moeite zou hebben je te volgen. ___ (.....) Je doel is uit de feiten der bevruchting te halen dat de erf. eig. onafhankelijke grootheden zijn. (.....) Eindelijk de berekenings–?tirade? is meer een 3e (p. 54) onderwerp om de Anwendung v.d. voorstell. der zelfstandigh.v.erf.eigensch. op het fixeeren duidelijk te maken:” Copy of Comment Moll, CB1, 175– 176.

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more a further calculation will give than the preceding one, whereas the reader will have difficulties in following you. ___ (.....) Your aim is to derive from the facts of fertilization that the hereditary characters are independent entities. (.....) Finally the calculation –?tirade? is rather a 3rd (p. 54) subject to make clear the Anwendung (application) of the notion of the independence of the hereditary characters on the fixation.

In the passages quoted, Moll used elementary arithmetical methods such as adding, subtracting and averaging. First the mean of the character value of the whole culture was calculated and that became the reference point, and was therefore characterized as zero. Individual values of that character became the differences with respect to this mean value. The character value of the individual offspring was the sum of the values for that character of both parents. Therefore, a hybridization of an organism with a character value 3 units above the average and another with a character value 3 units below the average would result in offspring with the mean value. Moll’s interpretation of De Vries's calculations as resulting in an additive hereditary theory has strange consequences: offspring of parents with values higher than the mean value will have an even higher value for that character than both parents, and offspring of parents with values lower than the mean similarly an even lower value. It is clear from the first passages of Moll's comment, in which he wrote that in case 3 reinforcement had to take place, that this consequence was clear to him. Moll knew that this consequence was definitely not in accordance with the facts. He was successful in conveying his objections against such an approach; De Vries took Moll's objections to heart and did not include the quantitative model in his book. The conclusion of the preceding two sections is that De Vries’s publication of his theory of heredity was surrounded by ideas that a reductionist explanatory framework was advisable; physiological or additive. This turned out to be impossible. De Vries’s hereditary ideas could not yet be reduced in this way. Notwithstanding, De Vries did not yet abandon a reductionist approach. In the next section his efforts to introduce statistical and probabilistic thinking in his work will be discussed.

Again a quantitative approach of heredity: rediscovery of the Mendelian laws Klebs’s conclusion of his review of De Vries’s Intracellular Pangenesis was that a general acceptance was not probable because of the large gaps and deficiencies in the hypothesis. Klebs’s prophecy came true; De Vries’s theory was considered too speculative. As I have discussed elsewhere, De Vries responded by developing a sizable research program of hybridization and other experiments with the aim of making his ideas on heredity more convincing.47 The experiments that he performed resulted in a huge amount of data, of which it was not directly clear how to interpret them. During the period of his physiological work De Vries had encountered the efforts of L.A.J.

47

(Stamhuis et al. 1999); (Zevenhuizen 1998b).

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Quetelet, F. Galton and W.F.R.Weldon to apply statistics to biological data and to make the normal distribution play an important role.48

Fig. 6. An example of the statistical interpretation of De Vries´ research results. De Vries called the curves Galton curves and the underlying law: Law of Quetelet and Galton. 1) Curve of the number of fruits of Oenothera Lamarckiana; 2) A. One sided curve of number of petals of Caltha Palustris; B. One sided curve of number of petals of Weigelia Amabilis; 3) One sided curve of number of petals of Ranunculus bulbosus; 4) Ranunculus bulbosus. A) One sided curve of number of flowers; B) Symmetrical curve of the number of flowers. C) Symmetrical curve of number of flowers. (De Vries 1918-1927) vol. V, 505. (Library Free University Amsterdam). De Vries started to try out statistics to interpret the results of his experiments (fig. 6). To that end he had to design his experiments accordingly. Through this approach in 1896 he encountered the 48

(Zevenhuizen 1998b), 431–435.

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laws that after 1900 would be named the Mendelian laws. He became persuaded that these laws were just what he needed, because the independence of distinct hereditary factors was central to them. Therefore, initially the great popularity of Mendel served him very well. Notwithstanding, because he was absorbed by the writing of the first volume of the The Mutation Theory, he paid little attention to these laws in 1900.49 In the course of time the experimental foundation of his pangenes theory was no longer the only aim of these experiments; criticism of Darwin's selection theory became another central point. Although he did not use the terminology of his pangenes theory in The Mutation Theory, it is clear from the way he expressed himself that he always had this theory at the back of his mind and that he tried to combine the two theories. He was more or less successful in bringing his ideas on pangenes and on mutations in line with each other, but it was more difficult to mould the Mendelian laws into the framework of his ideas on heredity and mutations. When his attempts to describe Mendelian crossings in terms of pangenes and mutations are examined precisely, it becomes obvious that he became entangled in a number of contradictions. As discussed elsewhere, he described these crossings in terms of his pangenes theory in two different ways, but in both cases unsolvable inconsistencies emerged. 50 He did not admit this explicitly, but some of his remarks convey the impression that he was more or less aware that Mendel's laws on the one hand and his theories on pangenes and mutations on the other hand could not be reconciled. Moreover, it remains unclear how his “normal” fluctuating variability could be fitted in with the quantitative approach of Mendelism, because he did not confront them with each other. He may not have found Mendelian genetics important enough to feel obliged to bring all kinds of concepts of his theories in line with it. However, it is not easy to imagine how the concept of fluctuating variability, in which numbers of the same pangenes can decrease or increase rather randomly, could be made compatible with paired Mendelian hereditary factors. The incompatibility between De Vries’s ideas on heredity and mutations and the statistical formulated Mendelian laws may have caused De Vries’s need to relativize the importance of the Mendelian laws, as he did in several passages of The Mutation Theory and elsewhere.51 I will give a few examples from The Mutation Theory. De Vries remarked that the splitting of the hereditary factors does not only occur in the case of Mendelian crossings, but also in hybrids where the splitting is not always complete.52 De Vries formulated the opinion that the deeper, the so–called systematically higher, characters are generally transferred to the next generation without changes and therefore without splitting.53 In addition, polyhybrids are possible whose elementary characters are connected so strongly that they cannot be split.54 In general, only the phylogenetically younger characters, or the race characters, obey Mendel's splitting rules, although even this is not the case with all of them.55 De Vries hypothesized that the occurrence of Mendelian crossings pointed to the termination of the mutation capacity within a family. 56 And 49 50 51 52 53 54 55 56

(Stamhuis 1995), 20–22. (Stamhuis et al. 1999), 259–264. (Meijer 1985), 220–223; (Theunissen 1994), 247–248; (Stamhuis 1995), 19–24. (De Vries 1903), 74. Ibid, 79. Ibid, 115. Ibid, 140–141. Ibid, 458.

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he apparently did not think of these kinds of organisms as the most interesting ones. He, moreover, emphasized that the possibility has always to be taken into account that phenomena occur that do not obey these hybridization laws.57 He could also not accept the rule that, when the offspring of Mendelian crossings possess the recessive character, the following generations remain forever constant, but stated that when it was allowed that this constancy was not absolute, an influence of an earlier hybridization could remain present in a latent state. 58 Somewhere in the offspring the dominant character could re–emerge. All of these assertions are more or less incompatible with the Mendelian laws. Although with hindsight we may conclude that he was very successful with this new attempt to use a reductionist framework, it was not the way he ultimately experienced it. The Mendelian framework was not in accordance with his ideas on heredity and evolution. It is plausible that this led him to consider the Mendelian framework as not being of essential importance.

Conclusion In this paper I have mainly restricted myself to a discussion of the methodological aspects of Hugo de Vries’s research choices, and showed that it was not clear from the beginning by which methodology heredity should be investigated. I intended to link Hugo de Vries’s later ideas and methods in heredity and mutations to the context of his early ideas and methods in plant physiology, and see whether a thread can be discerned in his scientific development. A conclusion is that this ambitious young man, who from his early youth showed a focused interest in botany, was from the very beginning of his career not only interested in plant physiology, but also in heredity and evolution. That he chose to work in physiology was because a research program in plant physiology was more easily attainable for him. He was however also steeped in Darwin’s publications on evolution and heredity. When it became clearer to him how to approach the problems of heredity and when the circumstances became favourable, he did not hesitate and started to work in that field. This work was first mainly theoretical, but later he started a sizable experimental program, first in variability and heredity but later – through his work on mutations – also in evolution. We see that on numerous occasions the approach he had become used to in his work in plant physiology was connected to his new work. He himself stated that his ideas on heredity had a physiological basis, although it was not yet known how to explain hereditary phenomena physiologically. Notwithstanding De Vries’s statements on this topic, others criticized his new ideas on heredity because De Vries did not explain them in the way as they were used to in his plant physiological work. He himself tried to apply quantitative methods, first an additive model in his original theory of heredity of 1889 and later, in 1896, a probabilistic approach in the interpretation of his hybridization and other experiments. Although this last effort was very successful in hindsight, in both cases De Vries found these approaches unsatisfactory. In all cases the reductionist argumentations could not satisfactorily answer the questions in which he was interested. The problems of heredity and evolution that he studied could not yet be moulded in such a reductionist way. Therefore he distanced himself from the applicability of these 57 58

Ibid, 486–487. Ibid, 525–526; see also 537–538.

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approaches. In the case of physiology he stated explicitly that for the time being they could not be used for his hereditary theory. In the case of the probabilistic Mendelian laws he relativized their significance, but in the background their incompatibility with his own theories will have been a factor in this diminution. I will close with the following observation. In the discussion of De Vries’s transition from plant physiology to heredity and evolution we can also take into account that for De Vries there is an additional reason why the contrast between the two research approaches will not have been so sizable as we experience it now. During the period of De Vries’ plant physiological investigations, Darwin was also working in the same field. In 1862 he published Fertilisation of Orchids, in 1875 Insectivorous Plants, in 1877 Forms of Flowers and in 1880 The Power of Movement in Plants. Although De Vries’s mentor in plant physiology Julius Sachs and Darwin criticized each other’s approach, they worked on the same kind of plant physiological problems. 59 At the end of the 1870’s Hugo de Vries corresponded with Darwin, and in 1878 visited him. 60 In their correspondence and during their meeting they discussed their similar plant physiological investigations: among others about their work on climbing plants. So De Vries knew that Darwin, who had formulated the theory on evolution for which De Vries strongly admired him, was also engaged in plant physiological problems. It is not improbable that the physiological interest of Darwin was a factor that diminished De Vries’ feeling of the contrast between plant physiology at the one hand and heredity and evolution at the other.

Ida H. Stamhuis, Vrije Universiteit, Amsterdam, [email protected]

59 60

(De Chadarevian 1996), 17–41. (Van der Pas 1970). See also (Theunissen 1992), 101.

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References Soraya de Chadarevian. 1996. ‘Laboratory Science versus Country House Experiments. The Controversy between Julius Sachs and Charles Darwin’, British Journal for the History of Science 29: 17–41. Jo Heimans. 1948. “Hugo de Vries (16 Februari 1848–1948)”. In: J. Heimans and Th. Weevers, Hugo de Vries. Voordrachten ter herdenking van zijn hondertste geboortedag op 16 februari 1949, Amsterdam: 1–9. Bas Jongeling, 1997. Wat is reductionisme? In: Willem Drees (ed.), De mens: meer dan materie? Kampen. Georg A. Klebs, 1889. “Intracellulare Pangenesis”, Botanische Zeitung 47: 734–739. Onno G. Meijer,. 1985. “Hugo de Vries no Mendelian?” Annals of Science 42: 189–232. Jan Willem Moll. 1918. “Hugo de Vries, 1848 – 16 Febr. – 1918”, 5 pp. I found this publication labelled as number 86 in one of the three files with numbered reprints of Moll's publications, which are located in the collection of the library of the University of Groningen coded C 1295, and don’t know where it is published. Peter W. van der Pas. 1970. ’The Correspondence of Hugo de Vries and Charles Darwin’, Janus 57: 173–213. Peter W. van der Pas, 1970. “Hugo de Vries”. In: Charles C. Gillispie (ed.), Dictionary of Scientific Biography vol. 14: 95–105: Peter Smit. 1980. ‘Hugo de Vries (1848–1935). Het veredelen van cultuurplanten’ in: A.J. Cox en M. Chamalaun ed., Van Stevin tot Lorentz. Portretten van Nederlandse natuurwetenschappers, Amsterdam: 163–176. Ida H. Stamhuis. 1995. 'The “rediscovery” of Mendel's laws was not important to Hugo de Vries; evidence from his letters to Jan Willem Moll', Folia Mendeliana 30: 13–30. This journal issue appeared in 1997. Ida H. Stamhuis. 2003. ’ The Reactions on Hugo de Vries’s Intracellular Pangenesis; the Discussion with August Weismann, Journal of the History of Biology 36: 119–152. Ida H. Stamhuis, Onno G. Meijer and Erik J.A. Zevenhuizen. 1999. “Hugo de Vries on Heredity, 1889–1903. Statistics, Mendelian Laws, Pangenes, Mutations” Isis 90: 238–267. Bert Theunissen. 1992. “De Beheersing van Mutaties. Hugo de Vries' Werdegang van Fysioloog tot Geneticus”, Gewina 15: 97–115. Bert Theunissen. 1994. “Closing the door on Hugo de Vries' Mendelism”, Annals of Science 5: 225–248. P. H.W.A.M. de Veer. 1989. Leven en Werk van Hugo de Vries, Groningen. Rob P.W. Visser. 1992. ‘Hugo de Vries (1848–1935). Het begin van de experimentele botanie in Nederland’ in: J.C.H. Blom e.o. ed., Een brandpunt van geleerdheid in de hoofdstad. De Universiteit van Amsterdam rond 1900 in vijftien portretten, Hilversum enz.: 159–178. Hugo de Vries, 1918–1927. Opera E Periodicis Collata.Utrecht: Oosthoek. 7 vols. Hugo de Vries, 1889. Intracellulare Pangenesis. Jena: Fisher. Reprinted in (Hugo de Vries, 1918–1927) Vol. V: 1–149. Hugo de Vries, 1897. ‘Erfelijke monstrositeiten in den ruilhandel der botanische tuinen', Botanisch Jaarboek 9. Hugo de Vries, 1901–1903. Die Mutationstheorie. Versuche und Beobachtungen über die Entstehung von Arten im Pflanzenreichh; zwei Bänder. Leipzig: Veit. [Friedrich A.F.C.Went,]. 1900. “Hugo de Vries.” Mannen en Vrouwen van Beteekenis in onze Dagen 31: 263– 320. Erik Zevenhuizen. 1998a. ‘Hugo de Vries: life and work’, Acta Botanica Neerlandica 47: 409–417. Erik Zevenhuizen. 1998b. ‘The hereditary statistics of Hugo de Vries’, Acta Botanica Neerlandica 47: 426– 463.

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Herbert Spencer’s two editions of the Principles of Psychology: 1855 and 1870/72. Biological heredity and cultural inheritance Snait B. Gissis

Introduction My paper is a report on one aspect of my project that addresses Spencer’s role in the process of explicitly evolutionising the human sciences in the second half of the 19 th century. Introducing an evolutionary component in the human sciences immediately confronts the issues of biological heredity and of cultural inheritance, –– whether clearly distinguished or not, whether intertwined or not, whether specific mechanisms for them can be adduced or not,–– and thus introduces a significant, if not crucial, element into these fields of knowledge. My presentation is in many ways an internalist one. I will not delineate genealogies of ideas nor draw contextual filiations in Spencer’s Principles of Psychology (PP) as much of this work has been done, and redone, rather thoroughly from various aspects during the past three decades, particularly by Robert M. Young, J. D. Y. Peel, Roger Smith, C. U. Smith, Robert Richards and Rick Rylance. My paper falls within the ‘evolution bent’ in reading Spencer, but perhaps with a difference in the significance attributed to the evolutionary tradition within which Spencer was located, as I have given an interpretation which throws light on Spencer‘s position within the human sciences of the second half of the 19th century. In an attempt to combine the various parts of my reading of the two editions of PP from the perspective of biological heredity and cultural inheritance I have used Spencer’s Lamarckism as my needle and thread. Let me first very briefly point to some elements in the context of Spencer’s enterprise. The period between the two editions was eventful. It witnessed several developments that had important political repercussions.1 Similarly, several notable intellectual and cultural controversies took place during the period. Also, a number of books that brought a new perspective to bear on social institutions and on diverging human capabilities were published. 2

1

Eg: 1846 – Repeal of Corn Laws. 1850 – Telegraph cable laid under English Channel. 1851 – Great Exhibition ("Crystal Palace"). Population of United Kingdom at 21 million. 1855 – Livingston discovers Victoria Falls. – Civil Service Commissioners appointed. 1857–8 – The Mutiny in India. 1858 – First Atlantic cable laid. 1860 – Budget–changes to Free trade. 1861 – Albert dies; Victoria retires into mourning. 1861–5 – American Civil War. 1867 – Second Reform Bill: enfranchises many workingmen; adds 938,000 to an electorate of 1,057,000 in England and Wales. (Disraeli's legislation)– Factory Act– extends restrictions on working hours for women and children to all factories– South African diamond fields discovered. – Fenian Rising in Ireland. 1869 – Suez Canal opened. 1870– Forster's Elementary Education Act establishes School Boards starts a system of state elementary schools. 1871 – University Tests Act removes religious tests at Oxford and Cambridge. – Trade unions legalized. Newcastle engineers strike for a nine–hour day. 1872 – Public Health Act– creates sanitary authorities with medical officers. 1873 – Population of the United Kingdom at 26 million (France 36 million). 1875 – Artisans’ dwellings Act encourages slum clearance. 1876 – Victoria named Empress of India. – Compulsory school attendance in Great Britain. 1877 – Transvaal annexed.

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I would suggest that some of these were relevant not only to Spencer’s later writings on sociology ( ‘The Study of Sociology’, ‘Principles of Sociology’) but also to the collectivist aspect of his work on psychology. Being a time of debates and controversies, the period from the 1850’s until the mid 1870’s was rich in newly founded publication organs for the various sides in those debates (over 170 new periodicals were started).3 This meant that both the publication of the 1st and the 2nd edition were not singular events but part of an ongoing activity in which the issues discussed in Spencer’s books formed one thread in a yarn in which things scientific, political, social and biological, not to mention religious, were intimately interwoven. 4

The two editions of the ‘Principles of Psychology’ Spencer published the PP1 in 1855. It consisted of one volume of some 620 pages; He issued the second edition first as sections to subscribers and then as volumes between 1870–72. The three volumes that made up the second edition totalled around 1800 pages. The 3rd edition appeared in 1880 and was identical to the second except for one chapter of some 50 pages set almost at the end, called ‘congruities’. The PP in its first edition was Spencer’s second book. He had published his Social Statics in 1851. Given that his essays just before and after issuing the first edition dealt with aspects of progress and evolution, the question arises why he did decide on psychology as his project. His testimony does not answer that question,5 and thus one is left to guess. I view it as an attempt to lay down the foundational assumptions that would enable him to pursue his enterprise of refashioning the contemporary human sciences. PP 55’ is foundational in the following sense: it dealt with ‘human nature’, ‘human understanding’, and the requirements for humans ‘to associate together’ –– the ever recurring themes of 18th- and early 19th- centuries philosophers –– but with a difference: those parts that dealt with philosophy were based on physiology, and the parts that dealt with psychology or ‘the social’ were based on an evolutionised physiology. It aimed to encompass all the known functioning of the human mind and of the human psyche, but it did so by situating humanity as a component of a natural continuity. The basic analytical units, the mechanisms, the generalisations, the laws, all would have to apply to living organisms at large in 2

3

Eg: Major works in British anthropology written between the appearance of the 1st and the 2nd edition of ‘Principles of Psychology’: Maine – Ancient law 1861; Lubbock – Prehistoric Times 1865; McLennan – Primitive Marriage 1865; Tylor – Researches into the early History of Mankind 1865; Lubbock – The Origin of Civilization and the Primitive Condition of Man 1870; Maine – Village Communities in the East and West 1871; Tylor – Primitive Culture 1871. Among the new periodicals some had been very short–lived, others survived until well into the early 1880’s. I have noted down some of those which either published articles or reviews on Spencer’s work, or dealt with issues similar to those discussed in his books of those years: The Leader 1850, The Saturday Review of Politics, Literature, Science and Art. 1855, The National Review 1855, The Oxford and Cambridge Review 1856, Bentley’s Quarterly Review 1859–60, Macmillan’s Magazine 1859, National Reformer 1860, Cornhill Magazine 1860, Home and Foreign Review 1862, The Reader 1863, The Fortnightly Review 1865, The Contemporary Review 1866, The Academy 1869, Fraser’s Magazine 1870, The New Quarterly Magazine 1873, The Nineteenth Century 1877; The Natural History Review 1854, The Naturalist 1864 , Nature 1869; The Geologist 1858, Geological Magazine 1864, A Quarterly Journal of Science 1871; The Anthropological Review 1863, The Journal of the Anthropological Institute of Great Britain and Ireland 1872; Asylum Journal of Mental Science 1853 continued by Journal of Mental Science 1859–1875, Mind 1876, Brain 1878 , Journal of Physiology 1878.

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order to apply to humans. It ‘naturalised’ human behaviour by putting it on a par with the behaviour of any living organism. The patterns and the biological mechanisms of heredity that would apply to living nature would apply to human perception and consciousness, as well as to unconscious, purposeful, planned and spontaneous human activities; to thought, feelings, desires, and to both individual and social behaviour. Thus, to create and establish a new psychology meant to lay the foundations for sociology, anthropology, politics and ethics. Spencer’s position on heredity when applied to all these fields had far reaching implications. In both editions, in so far as Spencer employed Lamarckian assumptions, there was an intentional effort to blur the distinction between biological heredity and cultural inheritance. The hereditary mechanism resorted to a delineated, alternating–rhythmic pattern between the two. The important feature was in the choice of ‘basic units’, which were neither solely mental nor solely physiological (and their process of recombining, etc was a hybrid one too). He resorted to the method of analysis characteristic of the empiricist–sensational associationist schools– i.e. to choosing basic units which were assumed to be the most elementary, primitive, while all other elements were regarded as combinations and compoundings of these units. In contradistinction to his predecessors, the evolutionary mechanism he had adopted operated on these basic units and their compoundings resulting in processes of complexification. Let me give as an example Spencer’s discussion of the emergence of consciousness in the two editions. Both editions indicated the significance of ‘change’ as a constitutive element in the explanatory framework. In the 1 st edition such a basic unit was what Spencer defined as ‘resistance’ – ‘action by direct contact, being the primary action, and at the same time the simplest and most definite… the sensation of resistance, through which this fundamental action is known.. …(resistance) is the only species of external activity which we are obliged to think of as objectively and subjectively the same’(1 st edition p 269– 270) – a hybrid unit. 6 In the 2nd edition Spencer gave an example of the most elementary physiological–neural unit, once again a hybrid, in discussing consciousness: ‘which state of 4

5

This becomes apparent when one looks at the list of major writings by Darwin, Lyell, Spencer and Wallace between the beginning of the 50’ and mid 70’s: Spencer Social Statics 1850; Spencer A Theory of Population , deduced from the General Law of Animal fertility 1852; SpencerDevelopment hypothesis 1852; SpencerPrinciples of Psychology 1855; SpencerProgress, Its Law and Cause 1857; SpencerTranscendental Physiology 1857; Wallace On the Tendency of Varieties to Depart Indefinitely from the Original Type 1858; Darwin On the Origin of Species by Means of Natural Selection. 1859; Spencer The Social Organism 1860; Lyell The Geological Evidence of the Antiquity of Man 1863; Wallace The Origin of Human Races and the Antiquity of Man 1864; Wallace The Origin of the Human Races and the Antiquity of Man deduced from the Theory of Natural Selection 1864; Spencer Principles of Biology 1864; Darwin The Variation of Animals and Plants under Domestication 1868; Walllace The Malay Archipelago 1869; Wallace Sir Charles Lyell on Geological Climates and the Origin of Species 1869; Wallace Contributions to the Theory of Natural Selection 1870; Darwin The Descent of Man and Selection in relation to Sex 1871; Darwin The Origin of Species by Means of Natural Selection. 6th edn 1872; Spencer Principles of Psychology 2nd edn 1870–72; Darwin The Expression of the Emotions in Man and Animals. 1872; Spencer The Study of Sociology 1873; Spencer Descriptive Sociology 1874; Spencer The Comparative Physiology of Man 1875; Wallace The Geographical Distribution of Animals 1876 Spencer himself, in his Autobiography, noted that ‘Psychology underlies sociology; and there had to be specified a number of those more special truths of psychology which have to be handed in to sociology as part of its data’ (Autobiography, vol ii, p. 240). Similarly he stated ‘After dealing with general psychology, it became requisite to enter upon the special psychology of man in preparation for sociology. Certain traits of human nature are presupposed by the ability to live in the associated state…’ (Duncan, Life and Letters of Herbert Spencer, vol ii , p.355, my italics)

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consciousness caused by a blow, may be taken as the primitive and typical form of the nervous shock… something of the same order as that which we call a nervous shock is the ultimate unit of consciousness;…’(2nd edition vol I p. 151). As Duncan indicated, Spencer in the 2nd edition ‘…ventured the hypothesis that sensations of all kinds and by implication higher feelings of all kinds result from the compounding and re compounding…so that there is an ultimate element of mind…’(Life and letters, ed. Duncan, vol ii, 348) . Human competences (or ‘tendencies’ or ‘latencies’) were neither solely biological nor solely mental–cultural. They were pre–organised moulds, inherited as neural modifications which functioned as neural–psychological patterns for organising experience and behaviour. At the same time they were open to further functional– adaptational modifications by that experiencing, which in its turn would be biologically inherited. Thus biological heredity was cultural inheritance; and heredity was conceptualised on the boundary between the individual and the collectivity (the ‘species’, the ‘race’). The Darwinian mechanism of natural selection played a very subsidiary role. Darwin was read through Lamarckism (which generally agrees with P. Bowler’s argument) and the role of variations in Spencer’s discussion of Darwinian mechanism was almost merely rhetorical. In fact, having both mechanisms helped Spencer draw a distinction between human ‘mental’ evolution and evolution at large. The central role of environment and of adaptation to its changing conditions could, theoretically, be interpreted in two major ways: I. As a facet of a hard–core biological determinism that would overrule whatever effects social changes of environment could have. The capabilities of the individual would then be conceived as merely a reflection of the conditions of existence of the species and thus be bindingly hereditary. Biological determinists could then apply concepts of biological purity to race, to class. II. The role of environment would be expressed through the weight of the formation and the transmission of socially and culturally functional, adaptational patterns such as habits, customs, traditions, and thus it could highlight ‘progress’ as an open ended endeavour. As Spencer conceived of biological evolution as a movement from homogeneity to heterogeneity, and understood it as complexification, it could, to a large extent be identified with ‘progress’ when applied to humans. Even the distinction between white and non–white races was not wholly a determinist one in PP. Lamarckism offered a double perspective on ethics and on society at large. It emphasized the overall importance of the milieu in shaping present and future generations through the inheritance of acquired characteristics, with 'use–repetition–habituation’ as a major explanatory mechanism. This implied the possibility of shaping the future: present changes could be bequeathed as prospective biological traits to be further elaborated in the future. The seeming determination of the present by the past was one consequence of such thinking, the other being the moulding of the present in light of a projected utopian future. But there was a price to be paid. What made possible this seemingly magic alternation between the cultural and the biological, which allowed for hybrid basic units, which made room for psychology as a unique 6

Spencer’s explanations of how space and time, motion and matter , were constituted derive from the analysis and then extrapolation of that hybrid basic unit). In choosing effort–resistance as a basic unit Spencer aligned himself in more than one sense with the psychological programme of the ideologues and its derivatives. In fact, this programme had been one of the principal sources of influence upon Lamarck’s later work on Man in the ‘Systeme analytique’ and in some dictionary items he had written in the second decade of the 19th century.

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science of the individual, which sanctioned the bridging between the solely experienced and the absolutely innate without ascribing to Kant’s transcendentalism and its vagaries, was the role assumed by the collectivity in Spencer’s theory. The Spencerian individuals contained the collective within themselves as their adaptational mode. This was the collective inheritance transmitted as biological heredity. It formed the moulds which fashioned and shaped the individuals’ experiencing, behaviour and thought. The changes were truly feasible only on the level of collectivities because though modifications were inherited by individuals, they were effected and inherited as social habits and psychological patterns (e. g. in the corollaries– cooperation). Thus the ‘future perspective’ of Lamarckism was wholly dependent on its being a social, collective rather than an individual construction. Evolutionising meant accepting the priority of the collectivity at least on the methodological and epistemological levels. In that sense Durkheim made explicit the implicit cost of Lamarckian evolutionising in the human sciences. The later crisis of neo–Lamarckism both in biology and in the human sciences helped turn collectivist assumptions into a methodological anathema. I would further suggest that in the period between the late 1840s and early 1870s, ((in some sense until the implications of Galton’s Hereditary Genius (1869) sank in) evolutionising psychology meant to discuss physiology and heredity in conjunction. In the more general context of ‘progress’ it meant gradual racialisation both in terms of imperial policies and in terms of social class determination on the one hand and of human developmental potentialities from a psychological point of view on the other hand. In this interim period individuals from the British middle class could gain a measure of immortality as their experiencing became that of the collectivity jointly, and the collective patterns became part of each individual singly. Moreover, behavioural patterns– habits– of the individuals were conceived as shapers of the cultural history of the collectivity. But it is doubtful, that it was a way to overcome the self–interested discreteness of individuals in most of political and economical theories of the time. The 1870’s can be viewed as a fairly clear dividing line between those who would investigate the brain only, and those who would discuss categories of thought culturally and philosophically only (e.g. the Neo–Kantians). The mixing of modes of discourse, of metaphors and analogies which Spencer used — philosophical, medical, biological, general–scientific — would not have been possible later on. Thus, in more than one sense, a comparison of Spencer’s two editions affords today’s reader an insider’s gaze into the passage from one phase to the other, and not just in terms of evolutionary models used, or conceptions of heredity. As is well known, from the mid 1860s until the late 1880s Spencer‘s writings were translated and disseminated all over Europe and the USA. They were reviewed, discussed and emulated. His writings had an impact on the human sciences of that era in a way which has not yet been adequately delineated. The period between his 1st and 2nd edition of PP was roughly the time in which anthropological work became more clearly divided into what we now call physical and cultural anthropologies. The last third of the 19th century was also the period where one finds more writings on society which intended to produce a synthesising oeuvre that encompassed an analysis of modern society, its structure, its dynamics, its peculiarities and what was then termed ‘its ills’ or ‘its pathologies’. This usually came together with a genealogical narrative, often couched in evolutionary terms and intended as a causal explanation. I would claim that the underpinning

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of at least some of the then more influential sociologies and cultural anthropologies was a peculiar combination of evolutionised biology with Lamarckian hereditary mechanism. This amalgam was first encountered and put to usage for that purpose in Spencer’s psychology. Quite a few of those who used the rhetoric of Darwinian ‘natural selection’ did in fact read Darwin along the same lines as Spencer, i.e., through some version of Lamarckism. The ‘Lamarckian amalgam’ made it possible to deploy the biological reservoir of problématiques, models, metaphors and analogies both for legitimation and for constitution. It was of great significance to fields of knowledge that were in the process of establishing themselves as autonomous and later as academic disciplines of the human sciences. One of the main assets of this Lamarckian amalgam was its hereditary mechanism which in many extant versions at the time allowed for cultural inheritance to be biologised and vice versa. The ‘Lamarckian amalgam’ catered also for the urgent need to present these theories not only as ‘diagnoses‘ of the past and the present, but also as suggestions for potential cure, ‘prescriptions’ for the future, based on ‘scientific analysis’. By evolutionising psychology Spencer gained hitherto untapped resources for psychological analysis, which provided numerous examples and illustrations. These were the materials authored by social and anthropological thinkers. Just as a Lamarckian evolutionist might use examples for the process of complexification from organic nature, Spencer used examples from ‘social nature’ as described by social thinkers, and particularly by anthropologists. In this connection recall that by the turn of the 18th century the then existing traditional, principally non- European societies had already been presented as earlier phases or stages in human progress (at times dubbed ‘human civilisation’). By the middle of the 19th century this was collapsed with a rather rigid view on the capabilities and potentialities of whites of European origin as compared to non–white groups, ordinarily described as ‘the lower races’. The portrayal and analysis of ‘the poor’, ‘the lower classes’ in general and of women were often conducted using the same rhetoric, and the same categories, attributes and localisation on the scale of progress. Though many of the anthropological and sociological theories of that period appeared to be methodologically individualistic, their implicit assumption was in fact collectivist in the above stated sense. Spencer’s psychology was the first to boldly espouse this mechanism for the purposes of describing ‘human nature’ and ‘human society’, and thus the first to harbour this discrepancy between claimed and de facto methodological assumptions: this had significant implications in his and in other contemporary theories. In a more detailed paper on which this paper is based I go into a rather detailed comparison of the two editions: first their overall plan, and then to issues related to the methodology and to the rhetoric of psychology as a new science. This serves as a background to my discussion of Spencer’s evolutionary mechanisms and their consequences: in terms of evolutionary models used, Lamarckian evolutionism, and the meaning and usage of Darwinian ‘natural selection’. Here, I shall just briefly highlight each of these latter issues. Psychology was constituted by Spencer in the 1870s edition as a ‘relational science’, a science that dealt with the relationship of the relations among outer phenomena with those among inner phenomena, i.e. with the relationship between environment, the organism and its physiological and ‘mental’ reactions to the stimuli of the environment. In both editions, each category used by

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Spencer in the description, analysis and explanation of psychology became constituted as a hybrid one – neither solely mental in the traditional sense nor solely physiological. The main feature of the evolutionary mechanism was perceived as adaptation between organisms and their environment (which for Spencer included ‘social environment’). Inheritance of acquired patterns of action was its principal mechanism. ‘Adaptation’ was then transposed into Spencer’s psychological discourse as the continual process of adjustment, conformation, correspondence, and the fitting of ‘internal and external relations.’ 7 The actual term ‘adaptation’ was used in the 1st edition only in relation to complex organisms ( which, presumably, were fully active in this process). Organisms were graded according to the degree of ‘correspondence’ between the internal and the external environment; life itself was defined in relation to it. In the 1st edition the continual adaptation was to be achieved through processes of differentiation, integration and thus coordination; the transposition of this model unto the domain of psychology was supposed to explain the evolution of distinctions between ‘outer’ and ‘inner’ within the states investigated by psychology, up to and including the emergence of consciousness and the explanation of intelligence. Consequently, various crucial conceptualisations such as those of space and time were presented as developing in an evolutionary manner both in the individual and in culture in general. In the 1870s edition, evolutionary processes at large, and those which were dealt with by psychology in particular, were characterised as a passage or movement from homogeneity to heterogeneity, along the lines suggested in various contemporary essays of Spencer. 8 In the 2nd edition there were two, rather than one, mechanisms of evolution called on to explain how change could be brought to bear on organisms and what the change consisted of. The one often called ‘direct equilibration‘ was the Lamarckian mechanism – in which the organism was also active. The other often called ‘indirect equilibration’ was the Darwinian natural selection which referred both to the survival of the more adjusted or better fitted and to their predominance in terms of reproduction. Spencer thought that the Darwinian mechanism of natural selection could work in evolution in general but would not suffice to explain what he considered the extraordinary development and complexification of the neural structures/the mind in the higher organisms (i.e. in humans). In this, he aligned himself with both evolutionists and non– evolutionists during the 1860’s who had adopted for various reasons somewhat similar positions. However, the main mechanism for solving problems 9 –– methodological, epistemological as well as structural –– was the usage of certain presuppositions of Lamarckism, namely: 01. a graded continuity of the organic world; 02. the evolutionary process was developmental and gradual; 03. a directedness of the evolutionary change towards growing complexification by way of adaptation via increasing differentiation, specification and coordination of these. Spencer 7 8

e. g. PP 1855, pp. 366–375. ‘Transcendental physiology’ (1857, ‘National review,’ under the title ‘The Ultimate Laws of Physiology’) and elaborated in his book ‘First Principles’(1862). Most of Spencer‘s interpreters in the 2nd half of the 20th century related this to his reading of von Baer particularly via Carpenter’s ‘Principles of Physiology ‘ (1851).

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defined this (after von Baer when discussing the development of the embryo) as going from homogeneity to heterogeneity, with increasing definiteness; 04. the environment, whether external, internal, or for Spencer also social 10, affected changes directly through adaptation; 05. a predominant role was assigned to the environment as a process of ‘posing problems and difficulties’ to the actively reacting organisms; 06. the mechanisms of use and disuse and habituation in the organisms ‘process of adaptation; 07. the organic entities discussed were to be considered as differentially self– organising; 08. the results of the process of adaptation of the individual organisms in the form of what Spencer called ‘functional modifications’ (‘acquired characteristics’ or in Lyell – ’peculiarities’) were biologically transmitted inter–generationally; 09. the gradual change in individuals of a species( ‘race’ in PP) occurred in such a fashion that all members of the group, ignoring individual differences, turned out to share the same changes, or more precisely the same results of the gradual process of change. (this assumption was actually at the core of Lamarck’s ‘transformism’); 10. all presuppositions applied equally to the whole organic nature and to humans, as ‘life under all its forms has arisen by a progressive unbroken evolution and through the immediate instrumentality of what we call natural causes.’ Let me exemplify what sort of ‘work’ evolutionary Lamarckism was supposed to perform and at what cost. Already in the 1855 edition Spencer used habituation– habit formed as a result of repetition– as an important explanatory tool. Habituation allowed the performer ( the reasoner, the speaker) to make significant shortcuts, because stages in the ‘mental’ process ‘merged’ and ‘sank to the unconscious,’ and the ‘shorter version’ became the ordinary one. One of his more detailed examples11 was an analysis of how a child learned to classify and to read. Here evolution and development became conflated, and Spencer argued that this process of ‘merging and sinking,’ whereby the activity of those specific parts of the reasoning, classifying, estimating etc. were converted into an ‘automated’ one, started at early infancy and thus became ‘organic.’ Conceptions of space and time were the developmental product of experiencing rather than its condition. Consciousness itself was defined as a consequence, an emergent consequence, of experiencing changes, rather than its precondition. Consequently, an infant’s perception would differ from that of a child, and a child’s from that of an adult. The sense of necessity of seeing the world this way rather than another, was consequent upon the ‘shortcut mechanism’ of ‘merging and sinking’.

9

10 11

As is well known, Spencer came from a dissenter family from Derby, and his father participated in the ‘Derby philosophical society.’ By his own testimony Spencer learned his Lamarckism as a young man by reading Lyell’s presentation of it and its refutation (particularly the first chapters in the second volume of the ‘Principles of Geology’). One would assume that he had also read Chambers’ ‘Vestiges’, he mentioned reading Rymer Jones, William Carpenter, Henri Milne–Edwards etc. From the late 1840s on he lived in London and was close to some of the radical figures of the 1830s. One can surmise that he was familiar with the extant versions of Lamarckism, catastrophism and other in–between contemporary narratives of evolution, as well as with the natural theology arguments against it. e.g. PP 1870-72, p. 381 PP 1855, pp. 205–207.

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For Spencer human adaptation was distinguished by the ability to learn and pass on gathered and organised information to other members of the collectivity. This process was accompanied by a parallel gradual extension of the relevant environment.12 (In certain contexts, human evolutionary adaptation was called ‘progress’ and was perceived as a highly complex hybrid.) In the then contemporary context reflex action and instinct were difficult notions to explain for both those who were committed to innate ideas and those who were not, particularly without assuming divine power. Spencer claimed that if his mechanism of accumulative–repetitive– experience was regarded as developmental and evolutionary, then it could explicate both. Reflex action was thus defined as the basic/primary hybrid unit, as a simple one, the compounding of which would be called ‘instinct’.13 As such the results of its action were to be passed on through hereditary transmission i.e. a biological heredity of habituated psychological patterns of experiencing acquired through the life of an individual. The thrust of Spencer’s argument concerning instinct and reflex action meant that every mental procedure had to be in some sense a learned one . Spencer was fully aware of the fact that this explanation could not suffice, given that it had to provide for the universally necessary and permanent character of both instinct and reflex action, also given individual idiosyncracies, diversity of environments etc. I suggest that Spencer therefore resorted to collectivist assumptions, and these became a major underpinning of the whole system. Experience and behaviour became manifestations of collective hereditary transmission, working equally on all possible generations of individuals of the relevant group. What was transmitted was not solely a psychological–cultural pattern. Rather, in line with the notion of hybrid units, a changed biological pattern, neural in this case, was hereditarily transmitted. This biological pattern, which he called ‘tendency’ (or ‘latency’) collectively provided the members of the collectivity with modified competences through which their experiences would be organised. This ‘tendency’ moulded the physical and psychological behaviour of individuals.14 These competences in their turn were the enabling preconditions for further individual accumulation of experience and thus for learning and communication. They allowed for further minute gradual modifications having had already been organised in specific moulds common to all members of that very general collectivity (humans in our case). 15 Experience and learning processes very gradually formed and fashioned biologicalpsychological competences — the mechanism which operated via a collective biological heredity. 12 13

14 15

PP 1855, pt. 3, ch. 8. One of the hallmarks of this process in humans was the combined ability to forge both specifications and generalisations which mutually generated relevant activities . It seems that Spencer took over some of the features of reflex as propounded by Laycock, e. g. it being a mediator between organism and environment, its automatic character, its pervasiveness at all levels of the neural system. In Laycock it appeared within the context of discussing addiction (alcoholism). See also the two appendices at the end of vol I of Spencer’s 2nd edition: ‘on the actions of anaesthetics and narcotics, Consciousness under chloroform’ pp 631–640. It is worth noting that throughout the 19th century discussions of addiction were a locus of a ‘mixed discourse’ on heredity––biological– psychological. (pp. 526–538). In a footnote which discussed perception of space: ‘….being inherited by the infant in a proximate form, is progressively modified by the daily activities that accompany development, until it reaches complete form.’ Concurrently there were, Spencer argued, modifications that had to do with the overall adjustment of the organism on the basis of the ‘ancestral one’(PP 1870-72, vol II, ch. 1, p. 193)

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In the cases of instinct and reflex action the moulding, which originally might have been more elastic, became inflexible through biological transmission of gradually hardened patterns. Presumably this inflexibility had its adaptational uses. Spencer’s account of instinct was just one instance of the fundamental mechanism of the new psychology, applicable to organic nature at large, to the development of an individual from infancy to maturity, and to the progress of humanity (as response to specific needs which produced that pattern in the first place) . Once this form of hereditary transmission was established as an explanatory tool, it could be used as an explanation for more flexible patterns of behaviour as well. Spencer went into great length in explaining the necessary level of development of the neural system in order for such a pattern to be formed and to be transmitted. He also drew a fine distinction between automated cognition, behaviour which developed over the life span of individuals, and behaviour that was so to say, ‘ready–organised’ at birth. Conversely, (hereditary) instincts could, according to Spencer, become a component of some more complex activity, in which case the isolated repetitive character would become distorted, experience and action less automated, until it lost both its character and its adaptational role. The latter process seems an extrapolation of the concept of reflex as a simple automated unconscious event, stretching it to cover conscious and complex ones. Purposive behaviour could be completely physiologised in this manner. E.g. feelings, which were called ‘inborn’ or ‘innate sentiments’ in the philosophical discourse, were explained by the same mechanism of experience, repetition, habituation, inter-generational, biological, hereditary transmission, moving from the level of individual heredity to the level of collective heredity. Thus Spencer treated ‘will’ as a natural continuation of the same. One can hardly think of more significant concepts than ‘will’ and ‘reason’ in the Victorian ethos and in Victorian culture (for both believers and non–believers). In their diverse interpretations, these encompassed the moral code and the ‘self forming’ ideals of the ascending British middle class.16 The decomposition of sensationalist notions, of a utilitarian world view, of Kantian transcendental epistemology, of certain facets of associationism, of faculty psychology, all depended on a specific mode of biologising and evolutionising. This mode was expressed in the notion of compounding and complexifying of hybrid basic units and the notion of collective hereditary biological, neural transmission of psychological and cultural moulds of experiencing.

A brief note on Spencer’s use of ‘natural selection’ There were three contexts in which ‘natural selection’ was used, with or without mentioning variations: 1. in the ‘biological’ volume– when discussing aspects of development of the neural system. There it was posited as the main mechanism for its primary development, through the production of certain variations. But even there Spencer specified that in higher organisms the causal productive mechanism was the Lamarckian one (e.g. PP 1870-72, vol. I, pp. 526, 614–15). 16

As well as in the medical discourse of that time; In this case Spencer’s analysis undermined the separateness and distinctness of the particular chain called the ‘ego’– but at the same time performed a salvational act by anchoring it, once again, in collective experience and biological heredity. Instead of the traditional ‘will’ and ‘freedom of will’ Specner suggested to his audiences the laws of growing adaptation.

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2. in the chapter on ‘Pleasures and Pains’. There Spencer assumed the hybrid basic units, and the other mechanisms discussed earlier. These accounted for the description of pain and pleasure, but not for their functions as tools for adaptation within the evolutionary psychological context. Utilitarians viewed pleasure and pain as instruments with the help of which to rear/raise and control individuals and lead them towards the ‘right’ modes of behaviour in the social–economic sphere. For Spencer pleasure and pain were more of a total gauge of the organism’s (including the human) state of affairs — how it coped with vicissitudes and perturbations and the continual changes in its environment. As indices of adaptation pleasure and pain would be found to function better in those individual organisms which lived longer, reproduced more, and thus replaced the other ‘less– adjusted’. This was how Spencer viewed natural selection in that context (PP 1870-72, vol. I, pp. 279–281). This was also the kind of context in which he would use the expression ‘survival of the fittest’. ‘Habitat’ and ‘migrations’ also made their first appearance in that context. It is worth noticing that Spencer distinguished there between organisms with foresight (humans and ‘some of the highest allied races’, i.e. apes) and other species. He then went on to produce a narrative of human evolution. This narrative was based on the familiar 18th century division of four general stages: hunting–gathering, pastoral, agricultural, and urban–industrial big societies. The relevant issues concerning natural selection are: Spencer’s emphasis on ‘the social’ as a type of environment which called for adaptation; the failure of successful adaptation(in terms of psychology) of humans in the passage from the second to the third stage and more pronouncedly from stage three to stage four (‘They were thereby cut off from activities like those of the men whose characters they inherited, and were forced into activities in which their inherited characters furnished no incentives’(282)). 3. The third case in the 2nd edition with ‘natural selection’ in the foreground was in the ‘Corollaries’ section, at the end of Vol II–2. The chapter on sympathy and sociality was mentioned earlier and the question here is whether the inclusion of natural selection made a significant difference in Spencer’s analysis. This chapter was one of Spencer’s ideologically laden ones in the PP. It was explicitly stated there that cooperation and mutual aid were the higher aims of social evolution, and this was juxtaposed with a sharply formulated criticism of military societies. According to Spencer, military societies encouraged selective sympathy and sociality. (In some of his later books such as ‘Principles of Ethics’ one finds a ‘reverberation’ of the distinction found in the ‘Corollaries’ between sympathy exercised towards one community and its withdrawal from anyone outside that community.) One can barely regard that as an adoption of a Darwinian group selection, which because of the inner contradictions it created, hampered their own evolution towards the desired ends. We may perhaps call it, anachronistically, ‘cognitive dissonances’. The occurrence of sentiments of sympathy and sociality and their increase was related to other social structural factors (power structure, family structure, societal type), spatial diffusion as well as to relative degree of intelligence. The sentiments were formed and acquired through the Lamarckian mechanism, while the survival of the fittest played the role of an additional factor ensuring the smooth functioning of the sentiment. Clearly, the most important discussion of that in the biological volume of the 2 nd edition was on ‘pleasures and pains’, concluding the second section –the inductions of psychology– in vol. I. Both feelings were explained in terms of the adaptational strategy of the organism. Throughout the

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evolutionary process pain indicated a perturbation to the adaptational equilibrium between the organism and its surroundings, while pleasure indicated a growing adjustment. Spencer did not explain the development of such gauges on the basis of hereditarily transmitted biological transformation. Rather, he presented them as a result of natural selection: the benefit which accrued thereupon would be for the collectivity (‘race’), in terms of its long term survival. 17 The role Spencer assigned to variations, which were central to Darwin’s mechanism, with which he certainly became acquainted shortly after the publication of the Origin, was in my opinion mostly rhetorical. He mentioned them whenever the aspect dealt with was predominantly biological. Furthermore, even there, it was relegated to dealing with simpler structures and organisms. It is not clear whether the variations ‘did’ anything in his employment of ‘natural selection’. There were, however, a couple of clear instances. One was related to the development of the nervous system, and thus clearly related to the biological aspect of the discussion. The other occasion in which natural selection was conjoined with variations was an interesting one. This was when Spencer’s previous dictum on pleasure and pain as indicators of adaptation seemed to be contradicted by phenomena which he defined as ‘… a seemingly abnormal desire to dwell on that which is intrinsically painful.’ This he found in certain modes of pity, defined as a very peculiar mixture of pain with pleasure, – e.g. watching the suffering of another – and he joined them with what he called ‘all the bodily appetites.’ These according to him included the sexual drive and the parental instinct. In that context he said: ‘…(that feeling of pity etc.) is not one which has arisen through the inherited effect of experience, but belongs to a quite different group, traceable to the survival of the fittest simply – to the natural selection of incidental variations.’ (vol. II, p. 623, my emphasis). What did he make of these variations? In a letter to Darwin in 1871 he wrote: ‘though I have endeavoured to show that instinct is compound reflex action, yet I do not intend thereby to negate the belief that instincts of some kinds may arise at all stages of evolution by the selection of advantageous variations. I believe that some instincts do arise: and especially those which are operative in sexual choice’ (Life and Letters, ed. Duncan, vol. I, p. 196). In 1899 he wrote a sort of ‘intellectual biographical sketch’ called ‘The Filiation of Ideas’, in which he referred to psychology, to the inheritance of functional modifications, and argued that ‘Though, nowadays, I see that the natural selection of variations in the nervous systems has been a factor, and in the earliest stages, perhaps the most important factor…’ (Life and Letters, ed. Duncan, vol. II, p. 324, Spencer’s italics) he would still stick on to his Lamarckian view etc. This fits in with the first case. 17

Incidentally the ‘Corollaries’ section was the most political one in the whole of the two editions. There Spencer’s position on policies and economies of war was stated bluntly. All in all, in those chapters and in the chapter ‘On pleasures and pains’ Spencer offered a grim view of modern military industrialised society, and gave a first approximation to what would, twenty years later, turn into two versions of binary sociological typology: that of Durkheim and that of Toennies. Durkheim in ‘De la division du travail social’ and Toennies in ‘Gemeinschaft und Gesellschaft’ suggested a typology of traditional versus modern societies. Schematically, and put somewhat unfairly, Durkheim criticised certain aspects of the traditional societies, particularly their homogeneity and lack of space for individual freedom, and thus called their type of solidarity ‘mechanical.’ While Toennies extolled their community of beliefs and solidarity of ties and support; he characterised them as ‘organic’; Durkheim saw modern societies as complex (differentiation, division of labour etc.), they allow space for individual autonomy and idiosyncracies, and have a solidarity and cooperation which emerges from the webs of interdependencies among their members (thus he called them ‘organic’); Toennies viewed modern societies as devoid of human solidarity, based solely on contractual ties, capital and gain oriented, socially isolating their members and he characterised them as ‘mechanical’.

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In his autobiography, when he mentioned the universal biological transformation in PP, he wrote that at the time he was committed to the Lamarckian mechanism ‘ …and was unconscious that in the absence of indirect adaptation effected by the natural selection of favourable variations…’ the explanation proffered could not suffice (Autobiography vol. I, p. 502): ‘…what Mr Darwin called ‘natural selection’ might more literally be called survival of the fittest…’; Also ‘some individuals in a species are so constituted that their moving equilibria are less easily overthrown than those other individuals, and these are the fittest which survive’ (Autobiography vol. II, p. 100). In the ‘Corollaries’ section of the 2nd edition, where social issues such as socialty, sympathy, egoism and altruism were discussed, Spencer emphasised the inaptness of natural selection in coping with adaptational situations in which complex organisms such as humans were involved. He claimed that Lamarckian mechanisms were much too slow for the fast pace of change in social environments as the alternation between the biological and the cultural needed more time to get usefully established. These were regarded as significant factors in the explanation furnished for the ills of modern civilisation. I would argue that though Spencer did acknowledge and accept that there was another evolutionary mechanism beside the Lamarckian he used, it played a very subsidiary role in his PP. Put more generally, one could perhaps say that Spencer viewed the mechanism of natural selection as one that did a ‘negative work’, rather than the creative one which Darwin attributed to it. This brings me to reiterate my claim that Spencer’s edifice of PP cannot be understood without taking into account his brand of Lamarckism.

To conclude– I have argued that in both editions Spencer’s innovative stance stemmed from a particular evolutionising of an amalgam of tenets from the then current mental and physiological theories of psychology. One of the consequences of this was that it became possible for Spencer to posit hybrid categories– neither solely physiological nor solely mental, and to intentionally blur the demarcation between biological heredity and social, cultural and psychological inheritance. This whole edifice could be coherently sustained if the underpinning of the blurring or hybridising became collectivist, while the rhetoric – political, ideological, and scientific – remained individualistic, and thus could provide legitimation for a new science.

Snait Gissis, Cohn Institute for the History and Philosophy of Science and Ideas, Tel Aviv University [email protected]

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References Spencer’s writings used in the paper Social Statics: or the conditions essential to Human Happiness specified and the First of them developed (1850) *A Theory of Population , deduced from the General Law of Animal fertility (1852) * Development hypothesis (1852) *The Universal Postulate (1853) Principles of Psychology (1855) *Progress, Its Law and Cause (1857) *Transcendental Physiology (1857) *The Social Organism (1860) First Principles (1862) Principles of Biology (1864) *The Classification of the Sciences (1871) Principles of Psychology 2nd edn. (1870–72) The Study of Sociology (1873) Descriptive Sociology (1874) *The Comparative Physiology of Man (1875) Autobiography (1904) The life and letters of Herbert Spencer (ed. David Duncan) (1908) Articles are marked by *

List of secondary sources: The following is the list of major secondary sources used for the actual writing of my paper. General and specific books consulted on social, political and cultural history of Britain, on the history of Darwin, Wallace and Darwinism, on philosophy, the sciences etc. and on the state of the biological sciences and medicine at the time are not included. Boring, E. G. 1929. A History of experimental Psychology: New Yorck. D. Appleton. Bowler, P. 1988. The non Darwinian revolution. Baltimore: John Hopkins. ———. 1984. Evolution: The History of an Idea. Berkeley: U California P. Burrow, J. W. 1961. Evoloution and Society. Cambridge: UP Cambridge Camic, Charles. 1986. ‘A Matter of Habit.’ American Journal of Sociology 91 Danziger, K. 1982. ‘Mid Nineteenth century British Psycho Physiology: a neglected chapter in the History of Psychology.’ In The Problematic Science: Psychology in Nineteenth-Century Thought. Edited by W. Woodward and M. Ash. New York: Praeger. Daston, L. J. 1982. ‘The Theory of Will Vrs. the Science of Mind.’ In The Problematic Science: Psychology in Nineteenth-Century Thought. Edited by W. Woodward and M. Ash. New York: Praeger. Gillispie, C. C. 1959. Genesis and Geology. Cambridge, Ma.: Harvard UP. Greene, J. C. 1981. Science Ideology and World View. Berkeley: U California P. Hearnshaw, L. S. 1964. A Short History of British Psychology 1840–1940. London: Methuen. Jacyna, L. S. 1981. ‘The Physiology of Mind, the Unity opf Nature and th Moral Order in Victorian Thought.’ British Journal For the History of Science 14 Jones G., and R. A. Peel, eds. 2004. Herbert Spencer: The Intellectual Legacy. London: Galton Institute. Lightman, B., ed. 1997. Victorian Science in Context. Chicago: Chicago UP. Lovejoy, A. 1959. ‘The Argument for Organic Evolution before the Origin of Species 1830–1858.’ In Forerunners of Darwin 1745 –1859. Edited by Glass B, O Temkin & W L Straub Jr. Baltimore: John Hopkins. Mandelbaum, Maurice. 1971. History, Man and Reason. Baltimore: John Hopkins UP MacLeod, Roy. 2000. The ‘Creed of Science’ in Victorian England. Aldershot: Ashgate.

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Peel, J. D. Y. 1971. Herbert Spencer. The Evolution of a Sociologist. London: Heineman. Peters, R. S., ed.1953. Brett’s History of Psychology. (1912–21). London: George Allen and Unwin. Reed, E. S. 1997. From Soul to Mind. New Haven: Yale UP. Richards, G. 1992. Mental Machinery, Part 1. London: Athlone Press. Richards, R. J. 1982. ‘Darwin and the Biologizing of Moral Behaviour‘ The Problematic Science: Psychology in Nineteenth-Century Thought. Edited by W. Woodward and M. Ash. New York: Praeger. 43- 64. ———. 1988. ‘The Moral Foundations of the Idea of Evolutionary Progress.’ In Evolutionary Progress. Edited by Nitecki M. H. Chicago: Chicago UP. Richardson, A. 2001. British Romanticism and the Science of the Mind. Cambridge: Cambridge UP. Rylance, Rick. 2000. Victorian Psychology and British Culture 1850–1880. Oxford: Oxford UP. Secord, J. A. 2000. Victorian Sensation. Chicago: Chicago UP. Smith, C. U. 1982. ‘Evolution and the Problem of Mind: Part I. Herbert Spencer ‘ Journal of the History of Biology 15 Smith, R. 1997. The Norton history of The Human Sciences. New Yorck: Norton. ———. 1973. ‘The Background of Physiological Psychology in Natural Philosophy.’ History of Science XI ———. 1977. ‘The Human Significance of Biology : Carpenter, Darwin and the vera causa.’ In Nature and the Victorian Imagination. Edited by Knoepelmacher U. C. & G. B. Tennyson. Berkeley. U California P. Stocking, G. W. 1968. Race, Culture and Evolution. New Yorck. Free Press. ———. 1987. Victorian Anthropology. New Yorck. Free Press. Turner, J. H. 1985. Herbert Spencer –a Renewed Appreciation. Beverly Hills. Sage. Warren, H. C. 1921. A History of the Association Psychology. New Yorck. Scribner & Sons. Young, R. M. 1967. ‘The Development of Herbert Spencer’s concept of Evolution ‘ Actes du Xle Congres International d'Histoire des Sciences Warsaw: Ossolineum, vol. 2, pp. 273–78. http://human–nature.com/rmyoung/papers/ ———. 1970. Mind, brain and adaptation in the nineteenth century : cerebral localization and its biological context from Gall to Ferrier. Oxford: Clarendon. ———. 1985. Darwin's Metaphor : Nature's Place in Victorian Culture. Cambridge: Cambridge UP. ———. 1990. ‘Herbert Spencer and Inevitable Progress.’ In Victorian Values. Edited by G. Marsden. London: Longman.

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Writing Heredity: Emile Zola’s Rougon-Macquart and Thomas Mann’s Buddenbrooks Ulrike Vedder

I. „Writing heredity“: I have bracketed these two terms to explore the relations between two sets of concepts: heredity and its theorisation, on the one hand; and the writing process, respectively literacy, on the other. In doing so, I do not intend to present Zola as a theorist of heredity who assimilated the theories of his age more or less accurately; nor do I intend to present Thomas Mann as a genealogist who ‘applies’ theories of degeneration to his characters. Instead, I shall take Zola as an example to examine how writing and different media operate in theorising heredity; I will focus on the last novel of Zola’s cycle Les Rougon-Macquart, which circles around Docteur Pascal, a physician and scholar of heredity; subsequently, and by way of brief comparison only, I will bring into view the written media that appear in the context of the genealogy of Mann’s Buddenbrooks by considering their role in establishing, respectively abolishing such genealogies. II. Le Docteur Pascal (1893) is the twentieth – and final – novel in Zola’s Les Rougon-Macquart (187193). As is well established, the cycle is subtitled Histoire naturelle et sociale d’une famille sous le Second Empire. The subtitle is striking in several respects, such as its opposition of nature and culture – that is bridged by a plain and simple ‘and’; moreover, we note Zola’s use of the term histoire naturelle to refer not just to the human species, but also to a single genealogical group. His time indication is striking, too, in that histoire naturelle does not envisage any longue durée, but merely the nineteen years of the Second Empire. Zola had already consigned the Second Empire to history with the publication of the first volume in 1871 and brought it to a narrative conclusion in volume 19, La Débâcle (1892), that offers an account of the defeat suffered in the FrancoPrussian War of 1870/71. So if he intends another volume to close the cycle a second time, it functions to render an account of the family’s demise – and of a feeble new beginning – following the downfall of empire – beyond all political coordinates, focusing fully on the family’s ‘histoire naturelle’.

Heredity Whereas Balzac’s Comédie humaine had focused on the synchronic ‘cataloging’ of the social species on the lines of zoological species, Zola’s „histoire naturelle“ can be considered a change within the species, that turns on heredity as its pivot. Given its lawfulness, as attested by science, heredity – as a biological term – has the potential to structure Zola’s voluminous serial novel, quite unlike mythical fate that would amount to little more than a ridiculous organisational principle in the face of the cycle’s expansiveness – and that would lack modernity, moreover. And as Zola describes his plans for the cycle in a preliminary note, „Il ne faut pas user du mot fatalité, qui serait

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ridicule dans dix volumes. Le fatalisme est un vieil outil“. Les Rougon-Macquart – initially conceived as a cycle of ten novels and later amounting to no less than twenty – thus unfolds the question how ancestral disposition takes effect on descendants and which variants emerge in the various constellations. Such structuring by the laws of heredity aims at the underlying, deeper dimensions of a genealogical order, whereas unstructured, unordered diversity reigns on the surface, among the individuals observed, that is; as Zola suggests, it calls for hereditary analysis: „individus, qui paraissent, au premier coup d’œil, profondément dissemblables, mais que l’analyse montre intimement liés les uns aux autres“.1 In the Préface to the entire cycle, Zola formulates his programme to make visible the underlying, submerged ‘liaisons’; he also names the technique for doing so: „Je tâcherai de trouver et de suivre [...] le fil qui conduit mathématiquement d’un homme à un autre homme.“ 2 This thread, which interconnects human beings and marks a path toward cognisance, opens up a broad field of meanings: (1) To begin with, the thread as a plaited connection evokes three concepts: the rhizomatic network or mesh; the hierarchized family order, for instance as a genealogical (that is family) tree; and Wittgenstein’s supposedly linear, yet by all means brittle conception of family ressemblance. (2) The thread as a means of orientation refers to the labyrinth, and (3) the thread as a steering mechanism evokes an author holding all the strings; as Zola notes in the Préface, „je tiendrai tous les fils“.3 Zola thus bases his twenty-novel cycle on the concept of lawful heredity as a prestigious model sanctioned by science. He adopts it as a narrative technique of authorial control, and strives to realise the programmatic verve of the preface throughout the entire cycle, in which heredity proves to be an extraordinarly productive generator of novels: „Je dois le suivre strictement, il est en même temps ma force et mon régulateur.“ 4 The novel concluding the cycle is about the theory and technology of such literary generativity, that converts life into writing and generates life through writing – albeit lives that exist but on paper and whose veins convey but ink. This last novel, which Zola himself referred to as „conclusion scientifique de tout l’ouvrage“, 5 elaborates once more the interrelations between the various family members – over five generations – that have appeared in each of the novels by seeking to systematise and theorise them. The interrelations are organized along hereditary – and not along psychological or juridical lines, for instance – in accordance with a notion of heredity that is conceived particularly in pathological terms.6 A scene set in an asylum where Adelaïde is an inmate makes this point clear. Adelaïde, 105 years old, is the family’s ‘great mother’, „leur mère à tous“ 7; not only do both branches of the family go back to her – the legitimate Rougons and the illegitimate Macquarts – but so does the transmission of hereditary phenomena of degeneration: „la lente succession des accidents nerveux et sanguins“.8 At the asylum, Charles, Adelaïde’s debilitated great-great grandson, sits across from his grandmother, who is paralysed and undead, and has fallen into a 1 2 3 4 5 6 7 8

Zola (1960), p. 3. Ibid. Ibid. Zola (1961), p. 799. Gumbrecht focusses on Zola’s problems to separate the synchronic (social) and the diachronic (biological) perspective (cf. Gumbrecht (1978), p. 41). Ibid., p. 799f. Cf. Föcking (2002). Zola (1967a), p. 1105. Zola (1960), p. 3.

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profound silence over the years; Charles unifies various impairments of previous generations of his family in that its central nervous and celebral deficiency to exert itself in a „débordement des appétits“9 in a compulsive-obsessive manner – reaching as far as décadence and madness, illness and murder – has assumed the physiological features of a haemophiliac. Adelaïde und Charles look remarkably alike, which makes them appear as if they bracketed their lineage – „la chaîne se déroulait, dans son hérédité logique et implacable“10 – and their deaths coincide: Whereas Charles gently bleeds to death, the apathetic aged woman is stricken by shock – because the steady flow of blood reminds her of the bloody deaths of her lover and her grandson – and dies of cerebral apoplexy. In this spectacular scene, the diversification that unfolds in the course of the cycle’s novels thus becomes compressed to absolute closeness [closure], which is accounted for in hereditary terms.11 The fact that the family destroys itself through the continued transmission of hereditary impairment reveals just how close(d) it is: „la famille brûlera comme une matière se dévorant ellemême“, as Zola notes in the Plans préparatoires.12 This closeness takes effect, too, with respect to Pascal Rougon, the principal protagonist, even though he positions himself outside the family: Not only does he not bear his family’s name and is addressed simply as Docteur Pascal, but he acts as his family’s external positivist observer and scientific investigator, studying it „avec l’attention d’un naturaliste surprenant les métamorphoses d’un insecte“.13 He makes use of his family as a generalisable „exemple à la science“,14 that is as material for his work on theories of heredity. He has been observing and recording his family’s hereditary development for thirty years, under implicit and explicit reference to various contemporary theories: : „Il était donc allé des gemmules de Darwin, de sa pangenèse, à la périgenèse de Haeckel, en passant par les stirpes de Galton. Puis, il avait eu l’intuition de la théorie que Weismann devait faire triompher plus tard, il s’était arrêté à l’idée d’une substance extrêmement fine et complexe, le plasma germinatif, dont une partie reste toujours en réserve dans chaque nouvel être, pour qu’elle soit ainsi transmise, invariable, immuable, de génération en génération.“15 Prosper Lucas’ Traité philosophique et physiologique de l’hérédité (1847-50) enters Pascal’s theory of how heritable information [genetic material] determines the human being and shapes his classification of four cases of heredity: „l’hérédité directe“, „l’hérédité indirecte“, „l’hérédité en retour“, „l’hérédité d’influence“. The fifth case, „l’innéité“, completes the schematic that has the structural advantage of allowing the classification of all human beings without fail. Pascal classifies himself as a case of „innéité“, a family member unimpressed by the laws of heredity, thus defining himself as an uninvolved exception, as an objective and dispassionate witness, as a scientific documentarian („un de ces cas fréquents qui font mentir les lois d’hérédité“).16 9 10 11

12 13 14 15 16

Ibid. Zola (1967a), p. 976. But this spectacular hereditarian continuity is irritated by the narrative and its allusions to royal images (Charles is cutting out kings and queens out of paper, he has a royal air, as a haemophiliac he alludes to royal families etc.), that contradict the scientific claim of the novel. Cf. Warning (1999), p. 248. Zola (1967b), p. 1741. Zola (1960), p. 301. Zola (1967a), p. 1015. Ibid., p. 946. Cf. Schmidt (1974). Zola (1967a), p. 66f.

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And yet he is involved in his family and its laws on several levels. Indeed, the fact that he has chosen his own family as the material basis of his research ties him to it. His critical state of health that will lead to his death confronts him with his family affiliation, too, as it also compels him to look for the cause of his impairment in the hereditary ailments of previous generations. As such, this reveals Docteur Pascal as a rather ambiguous figure, whose detached indifference toward his object of study collides with his involvement in „le legs inévitable de sa terrible ascendance“. 17 Yet it is Pascal’s incestuous relationship with his niece Clotilde Rougon – which remains surprisingly uncommented – that manifests the family’s closeness in the most far-reaching manner. The novel offers a euphoric description of their work on Pascal’s research, their great love, the celebration of their union and their life together, resulting in the birth of a new generation – albeit not until after Pascal’s death. Although both characters are demarcated from the Rougon-Macquart – Pascal has brought over Clotilde from the inner circle of her family to live with him and has thus ‘corrected’ her hereditary disposition („tu as corrigé mon hérédité“ 18) – their choice of lover ties them nonetheless to the confines and bonds of their family. Phantasmatic and narrative aspects of their incest are superimposed here. On the one hand, incest does not have the connotations of décadence and hereditary decline, but authorizes itself through health and nature. The greatest possible closeness between Pascal Rougon and Clotilde Rougon therefore aims at making new the Rougon family, which their child personifies. It is not the combination of difference that generates the new, but the doubling of resemblance, of the near-to identical. This comes to a head in that the child born of this incestuous association is elevated and consigned to the realm of myth: It appears in the last chapter (after Pascal’s death) as a redeemer, a saviour of the human species, as a Messiah even. Another narrative function that incest has here is to cover the incompleteness, brittleness, the „fêlure” that the greatest possible family closeness brings to a head, threatening the RougonMacquart family and its underlying theory of heredity time and again. In Zola’s seventeenth novel La Bête humaine, there is mention of the „fêlure héréditaire” – of the cleft, of the fissure, of the void – from which the protagonist, who is losing himself, ensues, „des trous par lesquels son moi lui échappait“. Gilles Deleuze has referred to this “fêlure héréditaire”, this hereditary decomposition as the „grande hérédité“, the proper heritage of the Rougon-Macquart that is, that the account of the everyday hustle and bustle might well conceal, but that nonetheless inscribes death in the novels themselves.19 The „fêlure héréditaire“ undermines both the scientific and aesthetic perfection of the family’s genealogical tree – which I shall turn to in a while – as much as the devoted vitalism with which Pascal and Clotilde evoke „la vie” evermore and factor out death. Insofar, their incest can be read as an attempt to conceal the fatal void with the greatest possible proximity and closeness. Following these comments on the key significance of heredity for the novel, I would like to consider the relevance of writing and of written media to heredity and its theorising.

17 18 19

Ibid., p. 1164. Ibid., p. 1154. Deleuze (1972).

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Writing Pascal develops his theories of heredity in a number of ways: through observation and by keeping records of his observations, by collecting documents in files, and by distilling and visualizing these in a genealogical tree. His filing cabinet contains a wealth of files, manuscripts and notes that require ordering and need to be assigned meaning, respectively pulled together, to make sense. The material calls for a structure that satisfies both scientific and aesthetic requirements. Structuring this material will not result in a definite form, such as a printed scientific treatise, but will rather assume the shape of continually reworked manuscripts striving to grasp new aspects of heredity theory. Such openness to change, for which the file („le dossier“) is an indispensable medium (as I shall explore in more detail below), reveals at once the mobility of the object of writing, namely life, that „vie“ that Zola evokes incessantly in the novel. Designing a book of life by deciphering the book of life: this is not only the objective of Docteur Pascal, physician and scientist, but also that of Docteur Pascal the documentarian and author. Insofar as Pascal’s writing starts out from observation(s) and experience(s), that is from empiricism, it is closer to narrating than to theorising. In this respect, we come across that quintessential scene in which he explains his research to Clotilde, recapitulating his entire family and its heredity interrelations in the process. He does so by tracing his finger along the genealogical tree and offering a detailed and passionate account. The knowledge that the tree formalizes is thus not codified once and for all, but can be transposed into oral accounts, scientific treatises and – last but not least – into novels. This scene amounts to a mise en abyme of the entire cycle, since not only do the titles of all twenty novels occur verbatim here, but so, too, does an abridged version of the novels’ storylines based on their principal characters. Just as the cycle is mirrored in this scene, so too is its personnel – Pascal und Clotilde – mirrored in the family’s history. The literary self-reference that is apparent in this mise en abyme thus produces an overwhelming mirror effect that centres attention in a compelling manner on heredity, conceived as a theorising of self-reference (in that heredity produces family resemblance, for instance, which in turn provides evidence for heredity). Docteur Pascal’s hypothesising, described recurrently, can be transferred onto the novel itself, respectively the entire cycle, which can be conceived as a hypothetical enactment. These are writings that have no place in scientific-academic writing because they are unable to articulate any final knowledge; rather, they ‘test’ thinking and knowing in the novel’s fictional space, that thus assumes the character of a laboratory.20 The openness of this fictional space corresponds with the openness of the question about heredity, which partly explains Pascal’s passionate 21 preoccupation with it, „parce qu’elle restait obscure, vaste et insondable, comme toutes les sciences balbutiantes encore, où l’imagination est maîtresse.“22 The way in which heredity is written conditions the way in which it is conceived. Here, it is „l’imagination“ that marks out the fictional space in which scientists come by their first hypotheses and that both scientists and poets inhabit: „le domaine des poètes autant que des savants“.23 The poets, however, form the „avant-garde“. This is able to discover unknown 20 21 22

Cf. Preiss (1983), p. 126. Pascal, „amant discret de la science“ (Zola (1960), p. 72). Zola (1967a), p. 947.

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territories and new solutions on account of the power of literature to access the world: „ils découvrent les pays vierges, indiquent les solutions prochaines“ (1008). 24 The mise en abymescene has shown that Docteur Pascal is both a scientist and a narrator, respectively an author.25 On the same terrain, poets have other means and technologies than scientists, even if the media they apply in their writing intersect, as the cases of the file and the genealogical tree illustrate, to which I shall now turn.

Files Pascal’s filing cabinet, one of the most essential objects not only of his study, but of the entire novel, has pivotal status. The key to it is a central object of desire. It wanders from pocket to pocket to be kept as safe as possible from view; eventually, it finds its ultimate hiding place – under the dead Docteur Pascal’s pillow. The key functions as a constant pointer to the cabinet, behind whose doors the files of the Rougon-Macquart family have been collected to be deciphered for the invisible laws of heredity. Even if there is a confusion of papers, a „pêle-mêle“, in the depths of the cabinet, the family files have been put into methodical order: „toute une série d’énormes dossiers s’alignaient en bon ordre, classés méthodiquement. C’étaient des documents divers, feuilles manuscrites, pièces sur papier timbré, articles de journaux découpés, réunis dans des chemises de fort papier bleu, qui chacune portait un nom écrit en gros caractères.“ 26 Sundry papers – documents, handwritten sheets, lettered stamping paper and newspaper cuttings – are gathered between strong blue file covers that have been labelled with large letters. Here, a bureaucratic ambition steps alongside the scientific one, emphasizing the materiality of the medium under reference to different qualities of paper and various functions of registration. In a narrative perspective, files and dossiers are specific insofar as they report the history of their own genesis in logging the entries of the documents they contain, for instance. They operate additively, absorbing everything indiscriminately and unresistingly; through the internal mobility of their contents, however, they enable synoptic readings under differentiated keywords, ever new readings that is. This, however, requires that the filing cabinet is not merely a labyrinthian repository, but an item of furniture permitting the ordering, relocation and further use of files, thus making it possible to couple them back to the user. In Docteur Pascal’s case, alphabetic and chronological order compete against each other, depending on whether he is interested in individual specifics or connections relevant to heredity and succession.27

23 24

25

26 27

Ibid., p. 1008. „Ah! ces sciences commençantes, ces sciences où l’hypothèse balbutie et où l’imagination reste maîtresse, elles sont le domaine des poètes autant que des savants! Les poètes vont en pionniers, à l’avant-garde, et souvent ils découvrent les pays vierges, indiquent les solutions prochaines. Il y a là une marge qui leur appartient, entre la vérité conquise, définitive, et l’inconnu, d’où l’on arrachera la vérité de demain...“ (Ibid.) Cf. Borie (1981), p. 140: „[...] en se proclamant docteur, Pascal ne dit pas toute la vérité, dans la mesure où il est aussi l’archiviste et, pourquoi se le dissimuler, le créateur de sa redoutable famille. Des ‘RougonMacquart’ et de leur auteur, ce dernier roman offre un tableau en abîme: l’écrivain y figure au milieu des siens, entouré de ses personnages asservis, comme le chasseur de ses trophées.“ Zola (1967a), p. 919. Cf. ibid., p. 1008.

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The medial conditions of the file just mentioned lead Docteur Pascal to ever new theorising of the laws of heredity, whenever he attends to the files and recombines the documents gathered therein. This results in a correspondence of the media format – the files – and their subject (heredity), because combining is a procedure that is both file-specific and concerns heredity at one and the same time. This becomes apparent when Docteur Pascal is taken ill and asks himself which case of heredity applies to him: „les dossiers répondaient par toutes les combinaisons possibles. Elles se présentaient si nombreuses, qu’il s’y perdait“.28 In his constant rereading and recombining of files he loses his way in different heredity cases, which yield different forecasts for him depending on their combination. After all, files have a retrospective orientation, like the modelling of the laws of heredity; their forecasting power, moreover, is highly doubtful. The filing cabinet is emptied by Pascal’s mother, in the night he passes away. She has been wanting to destroy his records for years to be able to make public her family’s fame above reproach and beyond blemish, to erase the dark stains that have tarnished her family’s history. Now she goes about burning all the papers in the open fireplace. In consigning the documents to the fire, she separates the files out into individual sheets, though not to read them, but because separate sheets burn better than convoluted paper. Destroying the files and therefore family memory thus replaces reading in order to erect another memory in place of the mobile papers, namely the endowment of a home for senior citizens through which she intends to monumentalize – and petrify – family fame. While the ceremonious laying of the foundation stone of the „Asile Rougon“ is underway in the last chapter, Clotilde sits in Pascal’s former study with her baby, as his will has bequeathed his estate to her. She is stowing baby clothes away in what used to be Pascal’s filing cabinet, and comes across some half-burnt remnants of files. Even though these are barely legible, so that the research that had preoccupied Pascal for years is destroyed and science is thrown back by twenty years, the remnants provide Clotilde with an opportunity to furnish an account of her own. Figures, memories, stories – that is living literature – arise from these remnants: „les phrases se complétaient, un commencement de mot évoquait les personnages, les histoires. [...] Et chaque débris s’animait“ (1215).29 The wreckage – „débris“ – reveals that it is not a matter of completeness when it comes to furnishing a narrative from the archive. Every narrative is always only a version of the archive, whereas the archive is an open labyrinth through which a number of paths could lead. Narrating depends on which guiding thread (‘fil conducteur’) is chosen.

Genealogical tree Alongside the illegible remnants of the files, the family’s genealogical tree has survived the autodafé unharmed, although it has become unreadable without the files. Whereas the files furnish an account of how they came into existence – as I have discussed above – the genealogical tree more or less distills their essence, operating as a document that has no knowledge of its genesis – which is even more amazing as this is a matter of a family lineage drawn out in schematic form. Nevertheless, the family’s genealogical tree as it is published in Les Rougon-Macquart can be made visible in its genesis: in the different models of trees that Zola tries out during his working process. 28 29

Ibid., p. 1033f. Ibid., p. 1215.

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Following an outline by Zola, the physician Georges Pouchet drew an “arbre mathématique” 30 (fig. 1) as an attempt to calculate the hereditarian relations (1:1 in the first generation, 1:2 in the second, 1:4 in the third generation etc.). According to such a law of proportion the relationship between the first and the fifth generation, between Adelaïde and Charles should be 1:16 – and not appear as if they bracketed their lineage. Another possibility is not a diagrammatic but a written tree (fig. 2) that can even be read from the left to the right, from the legitimate to the illegitimate branch that is. In 1878, Zola published a genealogical tree with the eighth novel Une page d’amour. But as the cycle progressed the writings overlapped the frame, more persons and more characteristics were added, and what Zola had called “ma force et mon régulateur“, „sans me permettre d'aller ni à droite ni à gauche“,31 gained its own growing dynamic (fig. 3). With the last novel Le Docteur Pascal finally a last genealogical tree – completed on Pascal’s deathbed – was published in 1893 (fig. 4). When the novel refers to the relationship between the files and the genealogical tree – „cet Arbre généalogique [...] dont les volumineux dossiers n’étaient que le commentaire“ 32 – the notion of the commentary brings into view the genealogical tree as a sacred text or a legal text requiring indefinite interpretation to make it meaningful and keep it alive. The genealogical tree functions to authenticate the scientific character of the novels and to exercise authorial control over them. Moreover, the scientific technology involved in the genealogical tree as a medium refers not merely to the function of representing knowledge, but also to that of generating it. That the temporal dimension of genealogy and heredity is translated into a spatial arrangement in the genealogical tree renders it possible to visualize the relations between the living and those long deceased, thus giving postulated heredity a visual structure. Changes in the succession of generations, that are not free from the coincidental, are captured and codified by the coherence and regularity of the genealogical tree. Its representativeness, moreover, implies a mode of thinking and a recording in terms of hierarchical deductions: The trunk explains the branches that in turn explain the leaves,33 so that the complexity-enhancing possibilities of the tree’s ramified crown remain retraceable back down to the tree’s roots within the framework of heredity theory. As a schematic representation of genealogy and lineage, the genealogical tree has narrative functions, too, even though it differs from the family bible or chronicle that are closer to narration, as Mann’s Buddenbrooks illustrates. In Pascal’s genealogical tree, entries are reduced to names, dates and heredity-diagnostic keywords – a compression of knowledge in written and diagrammatic form that has to be unfolded through narrative to be made meaningful, productive and transmittable. Docteur Pascal’s „testament scientifique“34, in which he verbalizes the findings of his research (in the presence of his colleague) on his deathbed, is beyond recovery for science, because only the genealogical tree itself has survived, and not Pascal’s spoken commentary or his files.

30 31 32 33 34

Ibid., p. 1008. Zola (1961), p. 799. Zola (1967a), p. 929. „Le tronc explique les branches qui expliquent les feuilles“ (Ibid., p. 1019). Ibid., p. 1178.

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Fig. 1: Les Manuscripts et les dessins de Zola. Vol. 1: L’Invention des lieux. Edited by Olivier Lumbroso. Paris 2002, p. 537 .

Fig. 2: Ibid., p. 533.

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Fig. 3: Ibid., p. 535.

Fig. 4: Zola (1967a), between p. 912 and 913.

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The genealogical tree, however, has gaps, too, despite Pascal’s efforts to attain its perfect beauty and completeness: „pas un trou“.35 The gaps are of a systematic kind. They include positing Adelaïde as the great mother, without recording her ancestors – and their illnesses – however. As is generally known, every origin, including that of a histoire naturelle, results from cultural norming, as the ‘root’ of the genealogical tree makes particularly clear; here, all hereditary impairments originate in Adelaïde’s „névrose originelle“ that – of all dates – dates back to the revolutionary years around 1790. Another gap opens up when Pascal is able to enter his own position in the tree, but does not know how to locate the path of succession, and is hence unable to classify himself as a heredity case.36 Just as little is he able to already classify the descendants of the youngest generation, which impairs both the explanatory force of the genealogical tree in scientific terms and its aesthetic guise: „Le pis était, pour la beauté de son Arbre, que ces gamins et ces gamines étaient si petits encore, qu’il ne pouvait les classer.“ 37 Pascal updates his genealogical tree to his last breath: When he enters his date of death as he is dying, recording overtakes reality – and it is precisely reality that recording brings into being because the strain that writing involves accelerates Pascal’s death. He also enters the birth of his child for the coming year, although Clotilde is only two months pregnant and her bereavement poses a threat to her pregnancy. Yet she does give birth – not only because she receives the deceased’s last letter in which he summons her home, but also because the child has already been entered in the „arbre généalogique“. A „genealogical imperative” emanates from the genealogical tree – an imperative that conjoins the linear narration in the epic novel with the idea of ancestors and descendants and with thinking in terms of origins and processes 38 and that requires a future that is not void: a narrating, in other words, that glosses over the „fêlure héréditaire“, the fatal void, and attempts to suspend it. The pathos of the novel’s ending ensues from this function in stylizing the child as the forthcoming hero and Messiah. At the same time, this stylization alludes to the notion of immortality. The genealogical tree should not just be a tree of knowledge – in its two meanings as knowledge and as sexual encounter as in the mise en abyme-scene featuring Pascal und Clotilde – , but also that other tree, the one located in paradise that has remained forever beyond human reach (cf. Genesis 3, 22): the tree of life. And thus the unborn child is entered as „l’enfant inconnu“ and becomes a new space in the genealogical tree to be written on, a placeholder of the future: it is entered in order not to have to accept the end of the family, nor the termination of writing at the close of the last novel; its entry, in other words, occurs in order to suspend death. In dealing with death, the files are closed, the last date of death is entered in the genealogical tree, and the chronicle is ruled off. In the novel, however, death is deferred because the debris yields narratives, and because the letters of the dead still reach their addressees, and because the wills that have been drawn up only begin to take effect gradually.

35 36 37 38

Ibid., p. 1006f. „Pourquoi, mon Dieu! l’Arbre ne voulait-il pas lui répondre, lui dire de quel ancêtre il tenait, pour qu’il inscrivît son cas, sur sa feuille à lui“ (Ibid., p. 1034). Ibid., p. 1018f. Cf. Tobin (1978).

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III. VISTA By way of a brief vista, I would like to turn to Thomas Mann’s novel Buddenbrooks. Verfall einer Familie (1901). In the face of Hanno Buddenbrook’s death, the last descendant of a wealthy mercantile family of Lübeck, the belief in immortality is evoked on the last page of this novel, too. Here, however, this is delegated to Sesemi Weichbrodt, a character delineated with unmistakable irony. The „Decline of a family” is reported from a narrative distance, quite unlike the high tones in which Le Docteur Pascal is narrated. The narrative distance in Mann’s novel, which deprives the demise of the Buddenbrooks of all hope for a new beginning, takes effect on how the family’s decline unfolds over four generations. Whereas Zola is intent on establishing the causal nature of the illness, Mann makes no reference to any clearly diagnosable hereditary impairments. Instead, it is rather a decline that is psychologically motivated and delineated in social terms that neither bookkeeping tricks nor legally determined succession can delay: „die Psychologie ermüdenden Lebens“.39 Following the end of Zola’s large-scale naturalist project of reproducing reality in a „roman scientifique“, Thomas Mann places emphasis on a culturally determined, multicausal history of decline, whose ironic depiction allows for distance to the characters, their afflictions and their self-deception through pre-formed discourses. Mann also subjects the family chronicle to this procedure. The chronicle represents the family’s genealogical self-insurance: a representative, gilt-edged booklet in which Jean Buddenbrook makes a verbose entry of the birth of his fourth child. In doing so, „kaufmännische Schnörkel“40 („mercantilistic curlicues”) and sentimental sentences blend with the god-fearing lines over and over. As he turns back the pages, Jean Buddenbrook reads previous entries about himself with „damp eyes” („mit feuchten Augen“).41 The emotion which grips him as he reads these entries time and again is an expression of a circular sentimental preoccupation with himself, a stagnating self-referentiality, which is in turn a symptom of the Buddenbrook’s increasing incapacitation and morbidness. Since the booklet is organised along chronological lines and is hard-backed, no stimulating recombinations are possible – unlike with Pascal’s files and their collection of loose sheets. There is no reference system other than the lapse of time; the genealogy remains codified in its media format, and is only readable in its linearity. Lacking any possibility of being reordered, its end is inevitable, however; thus the final stroke that Hanno dreamfully draws under the chronicle as a child proves to be a clear-sighted forecast, even if his father slaps him the face for this show of disrespect. It might come as a surprise that the media that document and convey reflection on heredity and decline in Zola and Mann – files, genealogical tree and chronicle – are all old writing media. Newer technical media, which would be imaginable too, such as photography, are absent. Which has nothing to do with the authors being out of touch with modernity, but rather casts light on the fact that the assertion of a family and hereditary association is a discursive-symbolic construction. The nature of visual media to furnish evidence, such as photography (employed around 1900 to perform facial anthropometry), proves its value only in an argumentative context – given that it is not merely a question of stating the proliferation of resemblance, which does not 39 40 41

Mann (1981-86), p. 10. Mann (1960), p. 53. Ibid., p. 56.

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necessarily result in a kinship order. Both novels put on view the power of this discursive construction, without which family heredity and decline would be inconceivable: Zola´s does so in a programmatic and identificatory manner, whereas Mann’s is derisive and distanced.

Ulrike Vedder, Zentrum für Literaturforschung, Berlin [email protected] Translation: Mark Kyburz

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References Borie, Jean. 1981. Mythologies de l’hérédité au XIXe siècle. Paris. Deleuze, Gilles. 1972. “La grande hérédité, la fêlure (= Introduction à La Bête humaine, 1967)”. In Les critiques de notre temps et Zola. Edited by Colette Becker. Paris. 44-49. Föcking, Marc. 2002. Pathologia litteralis. Erzählte Wissenschaft und wissenschaftliches Erzählen im französischen 19. Jahrhundert. Tübingen. Gumbrecht, Hans Ulrich. 1978. Zola im historischen Kontext. Für eine neue Lektüre des Rougon-MacquartZyklus. München. Mann, Thomas. 1960. Gesammelte Werke in 13 Bänden. Edited by Peter de Mendelssohn. Bd. 1: Buddenbrooks. Verfall einer Familie. Frankfurt/M. Mann, Thomas. 1981-86. Gesammelte Werke in 20 Bänden. Edited by Peter de Mendelssohn. Bd. 16: Rede und Antwort. Frankfurt/M. Preiss, Axel. 1983. “Pascal, ou la biodicée médicale”. In Les Cahiers Naturalistes, Vol. 29, No. 57. 116-131. Schmidt, Günter. 1974. Die literarische Rezeption des Darwinismus. Das Problem der Vererbung bei Emile Zola und im Drama des deutschen Naturalismus. Berlin. Tobin, Patricia. 1978. Time and the Novel. The Genealogical Imperative. Princeton. Warning, Rainer. 1999. „Kompensatorische Bilder einer ‚wilden Ontologie’: Zolas Les Rougon-Macquart“. In Die Phantasie der Realisten. München. 240-268. Zola, Emile. 1960. “La Fortune des Rougon”. In Les Rougon-Macquart. Histoire naturelle et sociale d’une famille sous le second Empire. Volume 1. Edited by Armand Lanoux. Paris. 1-315. Zola, Emile. 1961. “Une page d’amour”. In Les Rougon-Macquart. Histoire naturelle et sociale d’une famille sous le second Empire. Volume 2. Edited by Armand Lanoux. Paris. 797-1092. Zola, Emile. 1967a. “Le Docteur Pascal”. In Les Rougon-Macquart. Histoire naturelle et sociale d’une famille sous le second Empire. Volume 5. Edited by Armand Lanoux. Paris. 913-1220. Zola, Emile. 1967b. “Documents et plans préparatoires”. In Les Rougon-Macquart. Histoire naturelle et sociale d’une famille sous le second Empire. Volume 5. Edited by Armand Lanoux. Paris. 1667-1781.

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Heritage – Appropriation – Interpretation: The Debate on the Schiller Legacy in 1905 Stefan Willer

This new year sees a jubilee of one of Germany’s most renowned poets and playwrights: Friedrich Schiller, who died 200 years ago, in May 1805. In the first weeks of 2005, there was already much ado about Schiller in German newspapers; there is going to be a significant increase of Schiller stagings in German theaters; German publishers have started new editions of Schiller’s works; in March 2005, the Berlin Academy of Arts will arrange a 24 hours-reading of Schiller’s works with chancellor Schröder as first lecturer; and so on and so forth. Much of this boom reminds of the celebrations one-hundred years ago, when in 1905 Schiller was commemorated for his 100 th obit. That remembrance gave rise to large-scale discussions on the justification of claiming Schiller for various cultural and political interests – discussions which were centred on the notions of cultural property and inheritance. When I, some months ago, proposed to Staffan Müller-Wille and Hans-Jörg Rheinberger to do a paper on this Schiller debate, I mainly thought of it as of a historical matter. Strangely enough, the actuality of this jubilee only occured to me several weeks after I had written down my proposal. The reason for my denseness probably lies in the urgency and the emphasis of the 1905 debate: a kind of articulating the desire for cultural property which seemed to me fairly outdated. I know better now. To name but one example: Two weeks ago, German journalist Burkhard Müller strongly emphasized Schiller being part of our inheritance. In an article in the Süddeutsche Zeitung entitled “Schiller and the Future”, he argues that Schiller’s quest for the ‘ends’ of History in his famous lecture Was heißt und zu welchem Ende studiert man Universalgeschichte? (What is, and to what end do we study, Universal History?) still has an afterlife, an aftermath, that it is thus a legacy and heritage. In his article Müller says explicitly that Schiller bequeathed us this question: “Diese Frage hat uns Schiller vererbt”; and he also asks what it means to be Schiller’s heir, “[wenn] wir ihn heute beerben.”1 It would not have been likely to find formulations like these in a West-German newspaper ten or fifteen years ago. In Germany, ‘Erbe’ as a concept of cultural tradition, in the decades before 1989, was mainly regarded as a domain of Marxist aesthetics and, above all, of the official cultural and educational policy and bureaucracy of the GDR in its proclaimings of the progressive, humanistic, classical and socialist heritage.2 The more these socialist traces of the concept of ‘Kulturerbe’ fall into oblivion, the easier it becomes in German discourses to connect to the semantics of cultural heritage which is well-established in English as well as in French speaking countries – patrimoine culturel – and hence in the globalized canon of cultural values, the ‘world cultural heritage’, or simply ‘world heritage’. But this global concept is far from being self-evident. Its history goes back to the 19 th century in which the ‘invention of tradition’ played a decisive role in the self-conception of nations as 1 2

Müller (2004), 15. Cf. Dautel (1980); Willer (2005).

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ontological entities.3 This political impact of tradition has to do with a monumentalization of culture which is characteristic of the 19th century. This is not only true for sites, buildings, and museums, but also for literary culture with its rather vague material status. The great critical editions of literary lifeworks were often regarded as national affairs and had by 1900 established themselves as monuments of cultural legacy. As for the ‘Weimarer Klassik’ in Germany, it worked as an example for this kind of monumentalization. A wide range of Goethe and Schiller editions were available, the literary estate of both writers was stored in the ‘Goethe and Schiller Archive’ in Weimar since the 1880s. In the Schiller celebrations and discussions of 1905, you can see that the concept of cultural heritage and inheritance is ubiquitous – testifying that in the course of the 19 th century this idea had become more and more prominent. The aim of my paper is therefore to substantiate the importance of the history of cultural heritage in the context of “A Cultural History of Heredity”. Concepts of handing down knowledge and art within a society are closely linked to biological theories about the transmission of traits and, especially, to juridical and economical regulations concerning the transfer of properties. I will concentrate on this latter aspect in the following considerations as it displays the contested and controversial status of heritage. Official jubilees and celebrations of cultural tradition generate a scene on which these controversies can be staged. Speaking of a ‘scene’: Maybe the playwright Friedrich Schiller was such an important figure for the German cultural heritage of the later 19 th century precisely because of his singular ability to create theatrical conflicts which were both rhetorically and philosophically powerful. It was Schiller’s theatrical pathos along with the main subject of his poetry – individual freedom confronted with tyranny or social restraints – that established and renewed his importance for what was claimed by opposite parties as their own tradition. More generally, the importance ascribed to the particular poet Friedrich Schiller shows to which extent cultural tradition – and creation of cultural value – are based on the instance of ‘the author’. The modern copyright as conceived in the late 18th century sees the author not only as the producer of artefacts, but also as the person owning the intellectual property of his work. At the same time – around 1800 –, reformed codes of law like the Code civil or the Allgemeines Preußisches Landrecht granted proprietary rights on a large scale, which rendered necessary, among others, a new configuration of laws of succession, the case of succession being a specific form of transferring and acquiring property in which economy immediately touches the lives of the juridical subjects. The case of succession means death and change of proprietorship at the same time. This is also true for the author subject in his status as originator and owner of his works. If property is ascribed to such things as poems and dramas – but also to letters and diaries, as soon as they were composed by an author in the modern, emphatic meaning of the word –, then, of course, the death of the author4 gains importance for the question of his intellectual property. But what exactly happens when the intellectual proprietor deceases? One of the main questions debated in the beginning of the 19th century is: if, how and when his proprietary right is transferred to the general public. For there seems to be a consent that there is no possibility for the author’s descendants to inherit this proprietary right. In 1819 a copyright commission of the 3 4

Cf. Hobsbawm and Ranger (1983). Cf. the ‘classical’ post-structuralist papers by Barthes and Foucault: Barthes (1967); Foucault (1969).

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German Federal Assembly (‘Bundesversammlung’) formulates that the descendants of an author “cannot inherit the spirit by which a work is produced and by which alone it can be perfected, following its own peculiarity” (”[dass] sie den Geist, aus welchem ein Werk hervorgegangen und durch welchen es nach seiner Eigentümlichkeit allein vervollkommnet werden kann, nicht erben können”)5 – at which it is significant that in German peculiarity, ‘Eigentümlichkeit’, and property, ‘Eigentum’, are etymologically and conceptually very close to one another. 6 Nevertheless there are considerable objections to transfer this ‘Eigentümlichkeit’ of an author into public ‘Eigentum’ as soon as he is dead. For among the characteristics of the modern author there is also the need of doing business with his works. This is where the publishers come into play. They participate in the intellectual property on a contract basis, and they want to safeguard their proprietary rights – especially the protection against reprints – as long as possible. Speaking of Friedrich Schiller, there was a relevant litigation referring to this very problem of publisher’s copyright and public interest in the 1850s. Finally the publisher Cotta managed to have his print privilege prolonged so that just in 1859, the year of Schiller’s 100 th birthday, it was impossible to do free reprints of his works. This was all the more important as the public interest in 1859 was already substantial, Schiller becoming a kind of national hero and the Schiller celebrations having a considerable symbolic impact on the German nation building which was to be put into practice with the ‘Reichsgründung’ of 1871. However, the publisher’s intervention shows that concrete economical interests could decisively interfere in the appropriation of an author and his works by the general public; moreover, it puts into question whether the author, even in his living years, can really be regarded as the owner of his works. On this background, some verses of Goethe, related to Schiller’s posthumous fame, seem rather idealistic and a little too optimistic: “Schon längst verbreitet sich’s in ganze Scharen, / Das Eigenste, was ihm allein gehört.” (Long since, his most personal property is being spread out in crowds, resp. into the crowd.) Goethe himself plays a specific role in inheriting Schiller’s property. Not only did he have a strange affinity to the skull of his late friend which he had exhumed to keep it inside his own house some years after Schiller’s death,7 he also tried (in vain) to complete Schiller’s unfinished last play Demetrius, and he wrote a lengthy Epilogue to Schiller’s famous (and likewise lengthy) poem The Bell, from which the last quotation was taken.8 All of these symbolic actions were part of a technique of mourning9 whose aim was, as Goethe himself puts it in a letter to Cotta from June 1805, to represent not what was lost but what remained: “ich werde in diesem Sinne weniger das, was wir verloren haben, als das, was uns übrigbleibt, darzustellen versuchen.” (MA 6.1, 903) For Goethe, Schiller’s remains, or remainder, were obviously not the dead poet’s eternal belongings, but that which could be appropriated. In Goethe’s Epilogue, this appropriation is even backdated into Schiller’s living years: for several times, the sentence “Denn er war unser” (For he was ours), is repeated as a kind of refrain throughout Goethe’s stanzas. 5 6 7 8

9

Quoted from: Bosse (1981), 127-128. Cf. Plumpe (1979). Cf. Schöne (2002); for further details and connexions regarding the history of cranioscopy and brain research cf. Hagner (2004), 70-78. Goethe (1985-1998), vol. 11.1.1., 297-300, 300. Cf. the poem’s first version, vol. 6.1, 90-92. (Subsequent references to the Münchner Ausgabe are cited in the text using the abbreviation MA with volume and page number.) Cf. Horn (1998), 107-129.

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Some months after Schiller’s death, his poem The Bell and Goethe’s Epilogue were staged for a solemn commemoration which was repeated several times in the following years in Weimar. Goethe promoted these celebrations and repeatedly added new verses and stanzas to his Epilogue. This was the beginning of the bourgeois Schiller remembrance which culminated in the jubilees of 1859 and 1905. The significance of this remembrance for a cultural history of heredity of the 19th century lies in the fact that Schiller not only seemed so likely to be appropriated – by surviving contemporaries as well as by posthumous followers –, but that at the same time his life, works and death altogether were interpreted as a last will and testament for posterity as a cultural community. To quote once again Goethe’s Epilogue – which by the way was one of the mostquoted texts in the 1905 Schiller celebrations –, it is the poet’s fatherland that has to execute his sacred last will: “Oh! möge doch den heil’gen, letzten Willen / Das Vaterland vernehmen und erfüllen.” (MA 6.1, 92) And, according to Goethe, it is especially in the act of celebrating that posterity can somehow effect a balance of values: it reimburses, as it were, the cultural heritage it received by paying tribute to the dead poet. “So feyert ihn! Denn was dem Mann das Leben / Nur halb ertheilt, soll ganz die Nachwelt geben.” (What life only gave to him by halves, posterity shall give in full.) (MA 11.1.1, 299) Most of the official speakers in the dozens of ‘Schillerfeiern’ celebrated in 1905 in many German cities, universities, schools and clubs, see their task in elucidating “in which sense Schiller has to be regarded as a national poet.”10 This formulation comes from the speech of Eugen Kühnemann, a philologist who was at that time director of the Royal Academy in Posen/Poznan (a city in a region which was Polish up to the end of the 18th century and then again after the first World War). The print version of the speech says that the return on sales of the booklet is “to be applied to the erection of a Schiller bust in Posen” – intended as a visible sign for the “spiritual unity” of the “greater Germany” which Kühnemann conjures explicitly at the “eastern boundaries of the fatherland”, as he puts it.11 More drastically, one professor Max Hecht promotes “The Idea of a Schiller Memorial in Königsberg” in Eastern Prussia which in his vision would work as an outpost against Slavian barbarism and a bullwark against the “assailing Polishness”. 12 In spite of this aggressive way of defending national possessions, Kühnemann in his Posen speech declares his fellow citizens the “glücklichen Erben” (lucky heirs) of the national idea of the 19 th century for which Schiller is seen as a forerunner.13 Indeed, the interpretation of the present as a matter of inheritance is at the core of the national-conservative claim of Schiller’s legacy. Many of the speakers see quite clearly that the historical Schiller could not at all be interpreted as a nationalist author; but they affirm that he ought to be read like this today. Kühnemann says, “It is the essence of the great figures of intellectual history that they are appropriated anew, time and again, and that every age has its own vision of them.”14 ‘Appropriation’ is a plainly programmatic concept that is not only valid in the nationalist context. In appropriating Schiller, the issue is neither the proprietary right of the author (so that you would have to respect his own intentions), nor the perpetuation of unalienable and 10 11 12 13 14

Kühnemann (1905), 25. Ibd., 5. Hecht (1905). Kühnemann (1905), 25. Ibd., 6.

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unchangeable cultural values, but a kind of actualization in which these values can be converted and thus valorized anew. It is this very conversion which was understood as an accumulation of cultural capital. To put it in the words of Erich Schmidt, renowned German philologist, in his Schiller speech at the Berlin University: The Germans of the beginning 20 th century shall follow Schiller, but not as “satte Erben” (saturated heirs), but as “Mehrer des Reichs” – a rather strange formulation, meaning literally ‘those who augment the Empire’, but obviously understanding ‘Reich’ also in the sense of ‘Reichtum’ and thus speaking of an augmentation of cultural wealth and value.15 This kind of actualization is the concern of many speakers, as can be seen in titles like Schiller und die Deutschen der Gegenwart (Kühnemann), Schiller und die Gegenwart (Schiller and the Present),16 Schiller und die neue Generation (Schiller and the New Generation),17 Schillers Bedeutung für das Maschinen-Zeitalter (The Meaning of Schiller in the Age of Machines).18 This kind of actualization and appropriation is essentially related to the case of succession, thus, to the death of the author, which in these speeches works as a pathos formula for the vitality of his afterlife. Kühnemann says, “He bequeathed us his own life as an exquisite national possession”; and he emphasizes the meaning of the fact that it is not a birthday but an obit which is celebrated: “Whenever the nation celebrates the anniversaries – birthdays and obits – of its great men, it manifests a feeling of duty to take an interest in them just like in the next of our kin. But we do not think of their infirmity and mortality, but of their work. And in that sense celebrating an obit is even more meaningful than celebrating a birthday. For it is not important to us that the man existed but what he achieved for us.”19 So, in spite of Schiller’s ‘idealism’ often claimed by conservative speakers as a monument against contemporary materialism, the concept of inheritance ultimately effects a materialistic interpretation of intellectual history – that is, it uses the capitalistic logic of ‘achieving’ a performance by making it convertible into cultural value. It seems obvious that the political Left was highly suspicious about any conservative claim of Friedrich Schiller. In fact there was a large debate on the Schiller legacy among socialist and social democratic writers and theorists as well. This was not only a defiance to a national-conservative image of Schiller, but also a response to the inner-socialist discussion on ‘revisionism’, meaning the contested influence of bourgeois thinking on socialism. Falling back on bourgeois cultural traditions had been, ever since the beginning of the socialist movement, a nuisance and a necessity at the same time. It was especially the bourgeois pathos of liberty, with Schiller being the most illustrious protagonist in German literature, that had a great impact on agitational speakers like Ferdinand Lassalle, whose lectures to proletarian audiences, given in the 1860s, sometimes took several hours – and could therefore use a great deal of pathos. For Marx and Engels, on the other hand, Schiller’s pathos was not to be separated from its bourgeois interpretations in the later 19th century, either as a means of escaping reality or as an encouragement of nationalism. So it was the celebrations of 1859 that had the most important impact on Marx’s and Engels’s idea of Schiller – who appears to be one example of “tradition weighing on the brains of the living like a nightmare”, as Marx put it in his famous essay The 18th 15 16 17 18 19

Schmidt (1905), 6-7. Windelband (1905). Fulda (1905). Kammerer (1905). Kühnemann (1905), 10, 7.

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Brumaire of Louis Bonaparte.20 Still Marx and Engels were far from condemning bourgeois tradition as such, but they drew a distinction which proved efficient, namely, a dualism of bourgeois utopism on the one hand, and bourgeois ideology on the other. In Friedrich Engels’s words from 1872, the task was “to conserve what is really worth-while being conserved of the knowledge – science, art, social manners etc. – handed down by history, but not only to conserve it, but to turn it from a monopoly of the ruling class into a common good of the entire society.” 21 To be really sure what is worth-while being conserved, one has to draw the distinction in every single case of cultural tradition. So the important thing here is the value judgement. It is obvious that even here the connection between handing down and evaluating is a matter of inheritance. In one of his late writings from 1888, Engels explicitly uses this notion when he says that the German proletarian movement is “heir to classical German philosophy”. 22 By the first decade of the 20th century, the concept of inheritance was already well-established among socialist writers. It not only worked to distinguish wanted from unwanted cultural relics but also to designate the ‘bequest’ of Marx and Engels, the socialist ‘classics’, as they were soon called. So an important figure of that trend could be characterized by a fellow combatant as the “executor of Marx’s and Engels’s legacy” just because he intended to “bind the German working class to classical poetry, to Lessing, Schiller and Goethe, with untearable ties”. 23 This is what Rosa Luxemburg wrote about the left wing social democrat Franz Mehring who was one of the protagonists in the Schiller debate of 1905. So it is a definitely bourgeois notion of property transfer and accumulation of wealth that is used as an affirmative concept of marxist cultural theory. Nevertheless, the socialist and socialdemocratic contributions of 1905 aimed at bourgeois Schiller remembrance in a very critical way. A social-democratic newspaper speaks of a “Schillerkult”,24 thus referring to the solemnity of most of the celebrations, but also to the industry of Schiller monuments and busts – plus kitsch objects for domestic use like brooches, needles and drinking-glasses. More basically, the cultural technique of celebrating as such is made dubious. The writer and politician Kurt Eisner points at what he calls “Schiller-Baalsdienst” (Schiller idolatry) with a pun: bourgeois Germany, he says, “feiert Schiller, um von Schiller zu feiern.”25 This pun uses the double meaning of ‘feiern’ in German which means ‘celebrating’ as well as ‘resting from work’. So, what Eisner says is that the bourgeoisie celebrates Schiller in order to rest from the effort of really understanding him, whereas the way of the working class should be working on, or, working up Schiller. Franz Mehring’s writings played an important part in popularizing this idea. In 1905 he published several articles on Schiller and, above all, his book Schiller. Ein Lebensbild für deutsche Arbeiter. In this book – a 150 page survey of Schiller’s life and works – Mehring declares his own partiality from the first lines on. He says that he is attempting a reading of Schiller apt to the class struggle of the present, and that he is about to picture Schiller “von der sicheren Warte” (from the certain standpoint) of the working class.26 Of course, as a marxist, Mehring does not think that 20 21 22 23 24 25

Marx [1852] (1969), vol. 8, 115. Engels [1872] (1969), vol. 18, 221. Engels [1888], vol. 21, 307. Luxemburg (1984), 104. [N.N.](1905), 97. Eisner (1905), 24.

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this partiality is in any contradiction to historical objectivity, but that it is the only possibility to analyze and come to terms with historical contradictions as such. In the case of Schiller, Mehring argues that his famous idealism is not in contradiction to reality – as his bourgeois followers claim –, but that it indicates contradictions within reality. According to Mehring, the actualization of Schiller for the class struggle must be based on the finding that Schiller’s actual solutions, namely the invention of an ‘aesthetic state’ and the concept of ‘play’ to overcome the contrarieties of reality, are false, but that the highness of his conviction (”Hoheit der Gesinnung” 27) is right. In this very distinction – or, in this ‘distinguishability’, if the word may be allowed – lies the marxist concept of cultural inheritance. In the end of his “Lebensbild”-book, Mehring summarizes that the working class cannot regard Schiller as an infallible teacher or forerunner, but still has an inviolable right to claim his bequest, and will always pay honour to it: “was ihr [der Arbeiterklasse] von seinem Erbe gebührt, das hält sie in unantastbaren Ehren.”28 In this sentence, the distinction between the ‘inheritable’ and the ‘non-inheritable’ is confirmed: you only have to pay tribute to what is due to you, ‘was einem gebührt’. In his short article “Schiller und die Arbeiter”, written also in 1905, Mehring puts even more clearly that “the modern working class accepts Schiller’s inheritance only with critical reservation” by separating “what in him is still alive and what has died off”. 29 Thus an objection is raised against the concept of “Nachruhm”, posthumous fame, which Mehring himself takes into account in his Schiller-book. When he comes to the end of his “Lebensbild”, he uses heroic words on dying and afterlife not at all dissimilar to the corresponding passages in the official and semi-official ‘Festreden’: “While his body crumbled into dust, his great name lived on.” And Schiller is quoted with two verses articulating that earthly life vanishes, while the dead last forever: “Denn das irdische Leben flieht, / Und die Toten dauern immer.” 30 From a socialist point of view this concept of afterlife had to be judged as insufficient or even false. Since Marx’s early writings, the socialist movement was definitely sceptical about any combinaton of historical memory on the one hand and eternity on the other. In his already cited 18th Brumaire of Louis Bonaparte Marx had suggested to do away will all kinds of what he called “welthistorische Totenbeschwörungen” (historical conjurations of the dead).31 So, the internal separation of Schiller’s bequest into a living and a dead part was essential. It is only in this sense, Mehring continues in his article on “Schiller und die Arbeiter”, that modern workers may utter Goethe’s phrase, “For he was ours.”32 And Mehring adds two more famous lines by Goethe, Faust’s often quoted maxim on inheritance: “Was du ererbt von deinen Vätern hast, / Erwirb es, um es zu besitzen.” (To really possess what you have inherited from your fathers, you have to earn it.) (MA 6.1, 553) This again means appropriation – but in a specific way. Mehring’s use of this concept is not so much based on the desire of having ‘the complete Schiller’ at his disposal, but rather on a selection, which also means: a fragmentation of this cultural 26 27 28 29 30 31 32

Mehring [1905a] (1961), Vol. 10, 91. Ibd., 241. Ibd., 240. Mehring [1905b] (1961), Vol. 10., 280. Mehring [1905a] (1961), 235. Cf. Schiller (1958), 426. Marx [1852] (1969), 115. Mehring [1905b] (1961), 280.

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tradition. Speaking in terms of inheritance: Following Mehring, one does not have to decide whether to accept the inheritance or to waive it altogether, but one can decide which parts of it are acceptable. As a juridical practice, this might seem somewhat arbitrary, and it was not at all undisputed as an aesthetic practice. Marxist aesthetics since the 1930s made strong efforts to conceptualize appropriation in a more dialectical way, without simply recurring to political actualization.33 For Mehring, anyway, the internal separation of the heritage produced a distance between the testator and the heir which was crucial for the heir’s self-legitimation. And it was also for reasons of self-legitimation that the socialist concept of cultural inheritance was at the same time an attempt to disinherit those that had been in possession before. As the journalist Karl Korn put it, the working class was to “enter upon the inheritance of the intellectual heroes that was forfeited by the bourgeoisie.”34 The bourgeoisie of course struck back. Several Schiller speeches of 1905 emphasize that Schiller will never be the hero of the proletariat but will always remain the poet of the middle classes. With a hundred years of historical distance, this contention can be seen as a negotiation of cultural values at a wider range. Moreover, it shows that these values have to be negotiated to be valorized at all. This idea is also displayed in contemporary economical and philosophical value theories. Georg Simmel’s Philosophy of Money for instance (first published in 1900) is based on the perception that values have no causes within the things evaluated but that they are expressed and generated in comparison to other values. Values as such are downright relative; evaluation quite simply is relativity. In Simmel’s words: “Only relativity generates the objects’ value in the objective sense, because it is only in relativity that the things are kept in distance from the subject.” 35 Speaking of the relativity of values in the context of cultural inheritance, I think it is not just a pun to refer to the double sense of relation. By relating yourself to Schiller, you became, as it were, Schiller’s relative and thus were entitled to be his heir. In this perspective, even the most enthusiastic contributions to the debate ultimately do not refer to ‘eternal’ or ‘immaterial’ values, but they operate relatively and economically. Relations of values need “Träger” (carriers), as Georg Simmel puts it. The main carrier substance he deals with is money, “the embodied relation of economic values.” 36 But Simmel regards art as well as a “structure of comparing values.”37 When it is about cultural inheritance, one could say that this structure is generated by a comparison between the old and the new. The creation of new cultural values requires an estimation of how the difference between innovation on the one hand, and tradition, the old, the existing on the other will be evaluated at a certain point of time. Thus you can estimate why and how this difference is likely to be stored in the cultural memory.38 So the idea of converting and augmenting cultural values by appropriating texts written by the dead poet Friedrich Schiller reveals its economical logic. It is by relating oneself to elements of the 33 34 35 36 37 38

Cf. Franz (2000), 190. Korn (1907/08), 414. Simmel [1900] (1989), 135. Ibd., 130. Ibd., 163. Cf. Groys (1992), 47.

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classical tradition that one’s own contribution can be evaluated and valorized in relation to the existing one. All the more, in the threefold ratio of the act of inheritance – someone inherits something from someone – the active part is shifted to the heirs. But still the testator’s perspective has to be taken into consideration. Even though appropriation is affirmed throughout this debate, none of the contributors would have wanted to play the part of the legacy hunter. This is where interpretations and ways of reading intervene. For of course it is in reading Schiller that he is made heritable for the various heirs claiming his legacy. So, which is the way of reading an author that is most likely to legitimate the reader as his heir? Franz Mehring criticizes in his Schiller book that the national-conservative reception is characterized by deliberate misreadings, and especially: misquotings, for instance by using words of the sovereigns in Schiller’s dramas to make statements about current problems of government. In the earlier socialist agitational rhetoric, on the other hand, classical quotations sometimes were even changed in their wording to make them applicable to the appropriator’s political aims. However it is striking that in most of the contributions of 1905 the policy of quoting Schiller does not seek the plain evidence of substantial statements in favour of – or against – certain political, cultural or pedagogical programs, but that they rather look for hints concerning the business of appropriation itself. This is why, I think, Goethe’s Schiller-Epilogue is so often quoted and why so many speakers recur to the Schiller celebrations of 1859 – as if the 1905 celebrations were a kind of frame in which preceding acts of remembrance could be observed. Among the Schiller quotations themselves, quite clearly those are preferred that deal with questions of death and remembrance, of posthumous fame and of aftermath. Eugen Kühnemann’s already mentioned speech closes in an exemplary way by quoting a Schiller epigram that displays the difference between mere physical procreation of the species (the genus), which is said to be the business of “millions”, and the tradition of humanity, which is only ensured by the happy few: “Millionen beschäftigen sich, daß die Gattung bestehe, / Aber durch wenige nur pflanzet die Menschheit sich fort.”39 In conclusion: The meaning ascribed to inheritance in this German cultural-political debate of the early 20th century makes it evident that the history of cultural heritage plays an important role in a cultural history of heredity. The ‘Schiller cult’ of 1905 shows to which extent the culture of inheritance can be regarded as a way of negotiating with the dead. The interesting part of these Schiller commemorations is not that they confirm the fact that there were conflicts between conservative and socialist interpretations of the classical tradition. Beyond this divide I have tried to argue that every definition of cultural heritage is based on appropriation, because it is the very concept of property that permits the translation from juridical and economical to aesthetic transactions of bequest.

Stefan Willer, Zentrum für Literaturforschung, Berlin [email protected] 39

Schiller (1958), vol. 1, 303.

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References [N.N.][1905] 1998. “Schillers Nachruhm.” In Schiller-Debatte 1905. Dokumente zur Literaturtheorie und Literaturkritik der revolutionären deutschen Sozialdemokratie. Edited by Gisela Jonas. Berlin. 97-103. Barthes, Roland. [1967] 1994. “La mort de l’auteur.” In Œuvres complètes. Vol. 2. Edited by Éric Marty. Paris. 491-495. Bosse, Heinrich. 1981. Autorschaft ist Werkherrschaft. Über die Entstehung des Urheberrechts aus dem Geist der Goethezeit. Paderborn. Dautel, Klaus. 1980. Zur Theorie des literarischen Erbes in der “entwickelten sozialistischen Gesellschaft” der DDR. Rezeptionsvorgabe und Identitätsangebot. Stuttgart. Engels, Friedrich. [1872] 1969. “Zur Wohnungsfrage.” In Karl Marx and Friedrich Engels, Werke. Vol. 8. Edited by the Institut für Marxismus-Leninismus beim ZK der SED. Berlin. ———. [1888] 1969. “Ludwig Feuerbach und der Ausgang der klassischen deutschen Philosophie.” In Karl Marx and Friedrich Engels, Werke. Vol. 8. Edited by the Institut für Marxismus-Leninismus beim ZK der SED. Berlin. Eisner, Kurt. [1905] 1998. “Schiller-Baalsdienst.” In Schiller-Debatte 1905. Dokumente zur Literaturtheorie und Literaturkritik der revolutionären deutschen Sozialdemokratie. Edited by Gisela Jonas. Berlin. 17-24. Foucault, Michel. [1969] 1994. “Qu’est-ce qu’un auteur?” In Dits et écrits 1954-1988. Vol. 1. Edited by Daniel Defert and François Ewald. Paris. 789-821. Franz, Michael. 2000. “Aneignung.” Ästhetische Grundbegriffe. Vol. 1. Edited by Karlheinz Barck et al. Stuttgart. 153-193. Fulda, Ludwig. 1905. Schiller und die neue Generation. Stuttgart. Goethe, Johann Wolfgang. 1985-1998. Sämtliche Werke nach Epochen seines Schaffens. Münchner Ausgabe. Edited by Karl Richter. München. Groys, Boris. 1992. Über das Neue. Versuch einer Kulturökonomie. München. Hagner, Michael. 2004. Geniale Gehirne. Zur Geschichte der Elitegehirnforschung. Göttingen. Hecht, Max. 1905. Die Idee eines Schillerdenkmals in Königsberg. Königsberg. Hobsbawm, Eric J., and Terence O. Ranger, eds. 1983. The Invention of Tradition. Cambridge. Horn, Eva. 1998. Trauer schreiben. Die Toten im Text der Goethezeit. München. Kammerer, Otto. 1905. Schillers Bedeutung für das Maschinen-Zeitalter. Berlin. Korn, Karl. 1907/08. “Proletariat und Klassik.” Die neue Zeit. Wochenschrift der Deutschen Sozialdemokratie 26/2: 414. Kühnemann, Eugen. 1905. Schiller und die Deutschen der Gegenwart. Festrede bei der Posener Schillerfeier. Posen. Jonas, Gisela, ed. 1988. Schiller-Debatte 1905. Dokumente zur Literaturtheorie und Literaturkritik der revolutionären deutschen Sozialdemokratie. Berlin. Luxemburg, Rosa. 1984. Gesammelte Briefe. Vol. 5. Edited by Annelies Laschitza and Günter Radczun. Berlin. Marx, Karl. [1852] 1969. “Der achtzehnte Brumaire des Louis Bonaparte.” In Karl Marx and Friedrich Engels, Werke. Vol. 8. Edited by the Institut für Marxismus-Leninismus beim ZK der SED. Berlin.. Mehring, Franz. [1905a] 1961. “Schiller. Ein Lebensbild für deutsche Arbeiter.” In Gesammelte Schriften. Vol. 10. Edited by Thomas Höhle et al. Berlin. 91-241. ———. [1905b] 1961. “Schiller und die Arbeiter.” In Gesammelte Schriften. Vol. 10. Edited by Thomas Höhle et al. Berlin. 278-281. Müller, Burkhard. 2004. “Schiller und die Zukunft. Er stellt eine Frage, die uns bis heute quält”, Süddeutsche Zeitung (December 30th, 2004), 15. Plumpe, Gerhard. 1979. “Eigentum – Eigentümlichkeit. Über den Zusammenhang ästhetischer und juristischer Begriffe im 18. Jahrhundert.” Archiv für Begriffsgeschichte 23: 175-196. Schiller, Friedrich. 1958. Sämtliche Werke. Edited by Gerhard Fricke and Herbert G. Göpfert. München. ———. 1958. “Das Siegesfest.” In Sämtliche Werke. Vol. 1. Edited by Gerhard Fricke and Herbert G. Göpfert. München. ———. 1958. “Die verschiedene Bestimmung.” In Sämtliche Werke. Vol. 1. Edited by Gerhard Fricke and Herbert G. Göpfert. München. Schmidt, Erich. 1905. Rede bei der Schiller-Feier der Königlichen Friedrich-Wilhelms-Universität Berlin am 9. Mai 1905 im Königlichen Opernhause. Berlin. Schöne, Albrecht. 2002. Schillers Schädel. München. Simmel, Georg. [1900] Philosophie des Geldes. In Gesamtausgabe. Vol. 6. Edited by Otthein Rammstedt. Frankfurt a.M. 1989.

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Willer, Stefan. 2005. “Politik der Aneignung. Die ‘Erbetheorie’ in den ‘Weimarer Beiträgen’ der siebziger Jahre.” Weimarer Beiträge 51. Windelband, Wilhelm. 1905. Schiller und die Gegenwart. Heidelberg.

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From pedigree to database. Genealogy and Human Heredity in Germany, 1890–1914 Bernd Gausemeier

The 1911 Dresden Exhibition for Hygiene gave a considerable boost to the German eugenics movement. Only one year after its foundation, the German Society for Racial Hygiene was able to present itself in a highly successful show for the popularization of medical knowledge. 1 The section organized by the Society confronted visitors with an impressive array of illustrations showing the progress of the biological sciences – diagrams on hybridization experiments or on the mechanisms of cell division and fertilization. The eugenicists’ preferred exhibit, however, were pedigrees visualizing the impact of heredity on human life – family trees showing the transmission of night blindness, musicality, tuberculosis or ‘moral insanity’. The use of pedigrees was also a major topic in the international meeting of eugenicists connected with the exhibition. In a keynote talk, the genealogist Hans Breymann expressed that he wasn’t completely happy with what he had seen in the showrooms.2 With mild sarcasm, Breymann stated that the medical profession had led genealogical method into a state of babylonic confusion. While generations of genealogists had worked to standardize and to simplify the method of pedigree–keeping, medical practitioners were excessively creative in inventing new genealogical systems. This situation, Breymann argued, would not only make the collected material incompatible, it would also create a serious problem for the propagation of eugenics: methodological ignorance might repel the hobby genealogists who formed a considerable potential of aides for the case of eugenics. Bringing genealogical method to eugenicists and interesting genealogists in problems of heredity would, in his eyes, provide eugenics both with a solid scientific groundwork and with more social acceptance. The most urgent common task for medical researchers and genealogists, according to Breymann, was the establishment of comprehensive collections for medical family research. He bemoaned that there were excessive public expenditures for “the cure and accomodation of degenerates”, but no support for the scientific study of human heredity that would help to control the spread of mental diseases. Since state institutions showed more interest in “deep sea research and polar expeditions than for this most profound problem of humanity”, he called for concerted action of genealogists and scientists.3 Breymann was neither confronting his audience with unheard ideas nor were his proposals as unpopular as he pretended. Among psychiatrists, there was a vivid discussion on the use of genealogical collections for the study of mental diseases. Around 1910, there were several schemes to reorganize and to standardize the family histories collected in asylums and for setting up regional surveys of ‘burdened’ families. The psychiatrist Robert Sommer suggested to complete such efforts with a national central office for psychiatric family research within the public health administration; other doyens of German psychiatry like Alois Alzheimer and Emil Kraepelin voiced similar ideas.4 1 2 3

von Gruber/ Rüdin (1911); Weindling (1989), p. 230; Weingart et al (1992), p. 206. Breymann (1912 ). Ibid., p. 28.

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The collection of family histories had been a part of psychiatric practice since the early 19th century, but it was not before the 1890s that German genealogists developed interests in this aspect of their occupation. After then, however, genealogical associations quickly built up contacts with the medical profession. This trend was spearheaded by the organization Breymann presided over, the Zentralstelle für deutsche Personen– und Familiengeschichte (German central office for family history). Its establishment in 1903 reflects the boom of family research in late 19th century Germany. Leading genealogists felt the need to coordinate the collecting activities of the countless clubs and societies. Shortly after ist foundation, the Zentralstelle sought to join forces with medical scientists. In 1908, its representatives reached an agreement with psychiatrists and eugenicists (among them the leader of the German Racial Hygiene movement, Alfred Ploetz) that assigned it as a central collecting point for genealogical material that might contribute to the understanding of “heredity, degeneration and regeneration”.5 This scheme exemplifies how dramatically the role of genealogy changed around this time. It was no longer regarded as an aristocratic pastime or as an auxiliary method for political history. In 1913, it was no longer unusual that a manual of genealogy contained contributions on psychiatric and anthropological applications and on the uses of family research in the social sciences.6 The change that is visible here did not only consist in an increasing use of genealogical method by medical researchers interested in hereditary transmission. The field of genealogy as a whole acquired a new meaning. In his History of Sexuality, Michel Foucault has devoted considerable attention to the redefinition of genealogy in the 18th and 19th century. While ancien regime genealogy represented the old descent and the strong alliances of a family, the pedigree of the bourgeoisie contained the ancestors’ diseases and anomalies: “The care for the pedigree became the concern about heredity.”7 Foucault relates this change to an essential shift in the history of modern society: While “classic” genealogy was associated with a historical regime (dispositif d’alliance) under which the distribution of wealth and of social prestige depended on the politics of marriage, the new form corresponded to a regime (dispositif de la sexualité) defined by the values of individual health and the control of reproduction. Thus, the metamorphosis of genealogy reflects profound structural changes in family, kinship and society. The two regimes should be understood as overlapping rather than successive historical formations, and the same applies to the different forms of genealogy. Pedigrees were used to visualize hereditary phenomena already in the 18th century, but this was only a first step in a long process of structural transitions. All notions about the transmission of physical features and diseases are necessarily based on genealogical knowledge. However, the rise of hereditarianist ideas in 18th and 19th century medicine was not necessarily associated with the use of pedigrees. Medical practioners usually treated the hereditary aspect as part of the nosographical description – in other words, the family history was an integral part of a narrative explaining the genesis of a disease. 8 Comprehensive collections of genealogical material emerged primarily in psychiatry. In the first half of the 19th century, asylums began to keep regular records about of their patients’ familial background. 9 In 4 5 6 7 8

Sommer (1913), pp. 394f; Rüdin (1911), p. 571. Zentralstelle (1908), pp. 106f. Heydenreich (1913). Foucault (1983), pp. 149f. Nukaga (2003).

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the same period, criminologists tried to reconstruct the ancestry of delinquents in order to understand the emergence of “moral degeneration”.10 Systematic surveys of alleged hereditary phenomena were closely connected to the rise of medical and penal institutions. Their genealogical collections allowed the definition and registration of menacing sub–populations, the insane and the criminal. The establishment of the Zentralstelle represents a different quality of institutionalized genealogy. The genealogists pursueing this project wanted to incorporate the specific collections of asylums, prisons and hospitals, as well as aristocratic or bourgeois family archives, into a comprehensive genealogical network of the whole population. This idea implied the redefinition of genealogy from a private into a public affair. Moreover, the limits between ‘normal’ and ‘abnormal’ genealogies were to be dissolved. It seems obvious that these plans were essentially a consequence of the growing political influence of the eugenics movement. Yet, the redefinition of genealogy can be related to a whole complex of epistemic shifts in the human sciences.

“Scientific genealogy” Only some years before the establishment of the Zentralstelle, German genealogists were virtually untouched by the discussions about the hereditary transmission of diseases or mental qualities. One of the first scholars to raise the topic in genealogical journals was the lawyer Stephan Kekulé von Stradonitz, son of the eminent chemist August Kekulé, and a co–founder of the Zentralstelle. But the most influential voice calling for a reorientation of the discipline was the historian Ottokar Lorenz (1832–1904), whose ‘Manual of the entire scientific genealogy’ (1898) made a lasting impact both on genealogists and scientists. Lorenz saw his manual as a reaction to the revival of genealogy in large parts of German society – which was, according to his view, unfortunately only taking place in private associations, not in the historical departments of the universities. But it was not his main idea to support the reconstruction of aristocratic customs. The largest parts of his book dealt with new developments in hereditarianist medicine, psychiatry and psychology. Given the mass of material compiled in this field, this is hardly surprising, but Lorenz saw a much deeper need to discuss the biological aspects of family research. He held that the achievements of modern science demanded a complete redefinition of the scope and the methods of genealogy. Genealogy, Lorenz stated, had always been concerned with the phenomenon of human heredity, but “the high ... standard of present natural science ... allows a totally different degree of certainty and comprehension for genealogical research than it would have been possible in earlier times of human observation.” 11 Lorenz was referring to the advances of cytology of the 1880s and 1890s that had brought about the chromosome theory of heredity. Even though the mechanisms of transmission were still obscure, the basic principles of sexual reproduction seemed to be firmly established: “Nothing today can be regarded as more secured by the exact ... investigation of the cell ... than the complete equivalence of the germ–plasms emanating from both sexually different individuals, and accordingly genealogy has, in its own domain, to regard and to evaluate the paternal and the 9 10 11

Jacobi (1834), p. 312. Becker (2002), p. 342. Lorenz (1898), p. 338

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maternal lineage as the basic elements of all examinations of the individual as well as of family, kin, people and species.”12 What he regarded as the pivotal result of modern biology, thus, was the microscopic ‘proof’ for the biological equivalence of the sexes. Lorenz stressed this simple fact because he regarded it as a crucial problem that a majority of genealogists only considered the male lineages of a family. The first step towards a modern genealogy had to be the emancipation from the aristocratic idea of patrimony. However, the appeal to the exact sciences had deeper implications. Lorenz made special reference to August Weismann, though he frankly admitted that he only knew the renowned zoologist’s ideas from secondary sources. Apparently he only used Weismann’s name as a synonym for experimental research on fertilization and cell division. Despite his own recurrent statement that genealogists should never meddle in unsettled scientific discussions, he situated himself in one of the most vivid biological debates. Weismann’s theory of the continuity of the germ plasm, which was most influential in the biological discussion, met considerable resistance in the medical community. The outright rejection of an inheritance of aquired characteristics challenged the prevailing view on the nature of hereditary disease. Weismann’s most eminent opponent Rudolf Virchow regarded the germ plasm doctrine as a symptom for a problematic development: since experimental biology had successfully emancipated itself from mother medicine, it had ceased to listen to the medical practitioner’s experience. 13 Virchow’s critique reflects that the study of heredity was, at the end of the 19th century, in a highly ambivalent situation. On the one hand, the mechanisms of heredity seemed to become a tangible object of exact scientific examination. For pathologists and psychiatrists, on the other hand, the term “heredity” simply stood for the occurence of certain maladies in a family. Distinguishing between “direct”, “collateral” and “atavistic” forms of heredity, they were lacking a clear–cut concept to analyze the mode of transmission itself. Lorenz’s program of “scientific genealogy” implied the promise that sophisticated genealogical methods would help to overcome this problem. However, Lorenz was not primarily dealing with pathological, psychiatric and psychological studies because he wanted to provide scientists with a new understanding of heredity. He considered it necessary to interest historians and sociologists in this field because they usually neglected the impact of heredity on human life. Nevertheless, most of his references to hereditarianist literature were marked by an acerbic criticism. Lorenz especially accused psychiatrists of being too careless in postulating hereditary influences. In his view, this biased perspective corresponded to shortcomings in genealogical methodology: medical family studies were too often based on an inadequate use of pedigrees. Pedigrees represent the descendants of a single ancestor. Medical pedigrees departing from a sick person and tracing the spread of the disease in the following generations unavoidably suggested the multiplication of the evil seed – in other words, they produced a picture of increasing degeneration. Similarly, Lorenz warned to derive any conclusions about a hereditary burden from isolated cases of madness in the ancestry – after all, who would not find “some well developed fools” among his forefathers? 14 The adequate tool of a biological genealogy, Lorenz argued, was not the pedigree, but the ancestral chart 12 13 14

Ibid., p. 347. Virchow (1886), p. 1. Lorenz (1898), p. 387.

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(Ahnentafel), the genealogical system showing the complete ancestry of a living person. The ancestral chart was the proper form to represent the total “germ plasm” of a given individual. On this basis, it would be possible to survey the number of certain diseases in the ancestry and to assess if a hereditary influence could reasonably be assumed. “Scientific genealogy”, thus, was not primarily defined as a method to detect more hereditary diseases, but rather as a way to constrain the growing influence of “hereditarianist superstition” (Vererbungsaberglauben) in the public.15 In the eyes of the conservative historian, the fear of degeneration was a modernist folly stirred up by fashionable authors like Zola and Ibsen. For Lorenz, the idea that mankind was riding a biological avalanche was a complement to the excessive belief in progress. Accordingly, he took a reserved attitude towards eugenics. His sceptical remarks show that it was clearly not his idea to boost the acceptance for genealogy by jumping on the eugenic bandwagon. On the contrary, he was repelled by radical ideas about “race improvement”. He ridiculed psychiatrists who hoped “that the better organized society of the future will, counselled by psychiatry, generally bar burdened persons from marriage”. 16 However, psychiatry was by no means dominated by the careless hereditarianism Lorenz criticized. By the end of the 19th century, heredity was commonly regarded as an important factor in the genesis of mental diseases, but there was a widespread awareness that its influence could hardly ever be measured exactly. The place of heredity in etiology was rather challenged than strenghthened as the advances in bacteriology provided new insights about many diseases. Further, the fear of degeneration did not find unanimous support among psychiatrists. Robert Sommer was an influential voice in the medical discussion raising quite similar objections like Lorenz. He strongly condemned the “pessimistic world–view” of the “doctrine of décadence” that was often propagated by the misleading use of pedigrees.17 Sommer also demanded a strict distinction between the terms Heredität and Vererbung. The occurence of similar symptoms in one family (Heredität) made it probable that hereditary transmission (Vererbung) was involved, but it could not be regarded as a proof in the proper sense of the word. This scepticism, however, did not restrain but rather reinforce the interest in human heredity. Both Lorenz and Sommer called for more methodic family research in psychiatric institutions. Sober work with these genealogical data would support the insight that the powers of heredity were not an inescapable fate but a controllable phenomenon. But what could the genealogist actually contribute to the study of heredity? Lorenz did not propose a clear guideline how to interpret the collected material. He even doubted that heredity followed a definite, intelligible law. Yet, he claimed that proper genealogical method would enable scientists to demonstrate certain regularities of human heredity, particularly for what he considered to be its central question – the problem of latency or “atavism”. The reappearance of hereditary traits after several generations seemed especially important to Lorenz because he regarded it as the main source for the modern “horror” of hereditary diseases. A comprehensible explanation why the forefather’s maladies suddenly struck a family, he reasoned, would help to chase off the “ghosts” of heredity that haunted the bourgeois society. 18 15 16 17 18

Ibid., p. 438. Ibid., p. 437. Sommer (1901), p. 67. Lorenz (1898), p. 438.

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In fact, Lorenz used the term atavism for all irregular forms of transmission. And the only case he discussed in detail was not even about the problem of latency, but about the constancy of a “family type”. As a historian with profound knowledge about European dynasties, he chose the most prominent example of physical characteristics prevailing in royal families – the Habsburg dynasty and its protruding lip. Basically, Lorenz’ approach did not differ from the ideas prevailing in medical family research. Every ancestral generation represented the total germ plasm of an examined person – for example, each grand–grandparent contributed 1/8 of the present generation’s genetic material (this simple fractional arithmetic was, in fact, not in full accord with Weismann’s ideas). Yet, Lorenz stressed a phenomenon that is especially striking in royal genealogies: due to marriages within the family, the actual number of ancestors in higher generations differs from the theoretical one. In other words, almost no ancestral chart (not only in the notoriously incestuous nobility) would display 16 or 32 different individuals in the fourth or fifth generation, but rather an increasing number of “multiple” antecedents. For Lorenz, this “loss of ancestors” was a crucial aspect that was usually disregarded in the study of human heredity. The Habsburg family was an ideal case to demonstrate how a certain trait (the lip) became a constant family characteristic through repeated intermarriage. Lorenz showed that striking cases of the lip appeared whenever marriages between Habsburg royals and women from side lineages of the dynasty occurred.19 In so far, he did not primarily trace an hereditary defect, but rather used a physical trait to describe how a dynasty maintained its biological and social identity. It was surely not only a result of monarchist sentiments that the Habsburg case has been discussed by several other scholars before and after Lorenz. Royal genealogies provided a favorable material for family studies: they could be followed over a long time span, there were excellent family records and, above all, portraits. Medical researchers dealing with a more common kind of people were aware that informations on a patient’s ancestors were for the most part insecure. The crucial question for hereditarianists was how medical data of the living generations could be recorded in a standardized form. But what notions of heredity formed the conceptual basis for such collections? Robert Sommer stated that organized family research was necessary in order to understand the “familial relations of mental diseases and their distribution in the whole country”. 20 Still at a time when he had accepted Mendelism as the theoretical key to human heredity, his primary question was not how certain diseases were transmitted but how they could be classified. Psychiatrists were used to see the nervous diseases as complex system of various clinical patterns. Accordingly, Sommer hoped that exact statistics about the familial and regional distribution of disorders would provide new insights into their “family relations”. Heredity, in this perspective, was rather a method than the object of investigation. Psychiatrists were usually not inclined to regard complex mental disorders as Mendelian units. The first scientist to follow this approach was the ardent racial hygienist Ernst Rüdin, who built up a large psychiatric family register at the German Research Institute for Psychiatry after World War I. Rüdin also introduced a genealogical method that differed distinctly from the ideas of Lorenz or Sommer. Instead of an exhaustive examination 19 20

Ibid., p. 407. Sommer (1913), p. 394.

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of selected family histories, Rüdin used broad samples of “short” genealogies (at most three generations) for the statistical calculation of Mendelian ratios.21 Such a statistical use of family research was clearly beyond Lorenz’s scope. Generally, his approach to heredity was descripitve, not analytical. Despite his references to experimental biology, he did not really understand heredity as a phenomenon that might be explained by a single principle. In so far, he was a typical representative of late 19th century hereditarianism. In Théodule Ribots influential book L’hérédité psychologique, for example, heredity still appeared as closely linked to reproduction – i.e. the circumstances of procreation were discussed as a possible influence on the qualities of the offspring. Unlike Ribot, Lorenz was not inclined to render romantic ideas about the extraordinary talent of children begotten in extramarital love affairs. 22 Yet, he allusively accepted the predominance of the paternal germ plasm. 23 In fact, he had not completely dissociated himself from the patrilineal concept of heredity he pretended to refute. With reference to such inconsistencies Wilhelm Weinberg, a pioneer of Mendelian statistics, later criticized Lorenz’ book as an overrated and insufficient contribution to the study of human heredity.24 But this critique missed both the original aim and the actual impact of ‘scientific genealogy’. Lorenz did not primarily define genealogy as a scientific tool to analyze human heredity, but as a way to understand human society and history.

History Lorenz used pathological and psychiatric examples in order to demonstrate the problems and possibilities of studies about heredity. As an historian, he was more interested in the transmission of “normal” psychological qualities. Yet, even though he claimed it to be unquestionable that individual characters were for the most part shaped by hereditary factors, he was rather unsatisfied with the scientific treatment of this problem. When it came to the inheritance of mental qualities, he was even more sceptical in questions of hereditary diseases. He directed especially harsh critizism against the most influential author on this topic, Francis Galton. Galton’s compilations of families with certain professional or intellectual preferences were, in his eyes, hardly a proof for the hereditary nature of ‘talent’, but a highly redundant affirmation of the well–known fact that a pear–tree doesn’t yield apples.25 Lorenz was especially unhappy with the superficial equation between talent and vocation: after all, the fact that a judge had a number of judges among his relatives gave little evidence if he was particularly good in his job. Lorenz’s limited esteem for Galton does not reflect a distrust in his conclusions, but an aversion against purely statistical method. For Lorenz, understanding human heredity was inextricably linked with detailed genealogical case studies, which were absent in Galton’s work. A convincing proof for the transmission of mental qualities was, in his eyes, a matter of meticulous historical critique. The excessively cited pedigree of the Bach family, for example, appeared to him as a questionable evidence for hereditary artistic genius since it represented a typical form of professional tradition. 21 22 23 24 25

Rüdin (1911). Ribot (1884), p. 175. Lorenz (1898), p. 407. Weinberg (1911). Lorenz (1898), p. 425.

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Lorenz considered it more fruitful to follow cases in which outstanding talent emerged from an unlikely social background. Lorenz did not primarily conceive genealogy as a tool to demonstrate the preponderance of nature over nurture. The influences of education and social customs were an integral part of his concept of family research. He envisaged (royal) family studies combining political history and hereditary psychology into a form of historical characterology – a model study about medieval Saxon royalty was provided by his pupil Ernst Devrient.26 Lorenz himself did not confine himself to the discussion of specific case studies. His program of “scientific genealogy” was about larger political issues. Much more than about the ghosts of degeneration, Lorenz worried about a phantom that scared the majority of German politicians and academics: Socialism. He explicitly defined “scientific genealogy” as a weapon in the struggle against egalitarian ideas. The modern contempt for genealogy itself was, in his words, a typical effect of “the social–democratic doctrine which thinks to be able to disengage itself from the natural foundations of human existence”. However, the “scientific spirit” of the present age and a growing “genealogical awareness” would repel these pernicious influences. The current revival of genealogy, Lorenz emphasized, was not about a return to an aristocratic class society, but would evoke a new consciousness that the specific qualities of families and individuals were the product of “genealogically developed characters.” “Under this banner, scientific genealogy now fights the social doctrines like aristocracy fought democracy in former times.”27 Biological knowledge was to replace the old aristocratic pedigree– consciousness. Lorenz did not only want to use his genealogical arms against Marxism. He first developed his ideas on genealogy in a 1891 book devoted to the celebrated historian Leopold von Ranke. With the master of Historism as his patron, he attacked all idealistic notions of social progress. Contemporary historiography with its preference for ideas and institutions, he complained, displayed an “insurmountable disgust for the marriage–bed and for births” and described historic events “as if they had happened on the moon”.28 Apparently it was time to bring “real life” and “real people” back into history. As human life was a biological process taking place in families, the old auxilliary disciplin of genealogy had to be considered as the starting point for all historical reflections: “Genealogy is the immediate certain fact in terms of natural history, the given element of historical events.”29 But what concept of history resulted from this heraldic motto? Clearly, the genealogic turn implied a highly conservative perspective. Kekulé, usually in full accordance with Lorenz, explicitly referred to Treitschke’s catchword “great men make history” – then, it was up to the genealogist to show how great families made great men.30 Still, Lorenz offered a little more than the prospect of a biologized form of dynastic chronologies. His basic idea was that the succession of generations formed the true basis of all historic change. Thus, generations were the units determining the rhythm of history. Lorenz called it a “natural law” that a generation (in the male 26 27 28 29 30

Devrient (1897). Lorenz (1898), p. 18. Lorenz (1891), p. 188. Ibid., p. 257. Kekulé (1900), p. 114.

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lineage) comprised 33–35 years, and he claimed that this regularity explained the historical identity of a period. In a similar fashion like the influential philosopher Wilhelm Dilthey, he attributed the ideas and political aims of a generation to common juvenile experiences. Lorenz, inevitably, perceived generations primarily as generations of monarchs which formed the character of a period. His concept of generation, thus, was clearly rather a sociological than a biological one, nevertheless it served an anti–sociological objective. According to Lorenz, the generational perspective illustrated that “ideas do not possess men, but men possess ideas, and accordingly inherit them or reject the inherited ones.”31 Generational history was to emphasize the role of deliberately acting people and to reject all notions of a transcendental progress in history. But the concept of generation also implied that historicity was largely determined by biological conditions. While Lorenz stressed that the coherence of generations was shaped by common memory, he attributed the constancy of national histories to biology. He described the existence of nations and classes as a result of inbreeding or, in his own special terminology, “loss of ancestors”. Not only the aristocracy secluded itself by establishing rigid regulations for marriage, also rural societies and premodern urban groups were formed by restricted possibilities for mating. But there were also limits for generative seclusion. Lorenz stressed that every genealogist was aware that even royal family trees showed an increasing number of common elements after some generations. No period in (European) history knew a total, caste–like separation, “there has always been a complete mixture among people, which ascends and descends between the generations, like the waves in the sea.”32 Yet, he regarded the inhibition of connubial mixture as one of the basic laws of human existence: “There is a tendency rooted in human nature to reduce the number of ancestors. The law of attraction between the kindred and the coequal sometimes becomes abandoned ..., but on the whole it is ineradicable, since love prospers best with loss of ancestors and equality of birth (Ebenbürtigkeit).”33 All social grouping, then, was the result of more or less conscious inbreeding. Whenever this law was violated and distinctions between classes and ethnic groups broke down, the inevitable result was decay and revolution. The insights of genealogy, Lorenz proclaimed triumphantly, disproved the idea of complete equality of all human beings as a socialist reverie. The most unthinkable offence against natural order was equality between white and coloured races: A total mixture of races, he warned, would bring about the end of civilization. Lorenz was not simply providing a vindication of racial segregation and class hierarchy. He proposed a perspective on human society which was consequentially based on the idea of selection. Genealogy was the adequate method to visualize how social stratification, i.e. social order, was produced. Developing Lorenz’ ideas further, the historian Armin Tille defined genealogy as a social science in its own right. According to Tille, the basic idea of genealogy was that “the individual man is an intellectual abstraction” while kinship formed the “constant basic element of society.”34 Due to this anti–individualistic perspective, the genealogical approach was closely related to organicist concepts in contemporary sociology. Tille explicitly referred to Ferdinand Tönnies’ influential dichotomy between community and society, claiming that 31 32 33 34

Lorenz (1891), p. 255. Lorenz (1898), p. 316. Ibid., p. 334. Tille (1913), p. 376.

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genealogy provided the appropriate tool to analyze how “organic” communities were generated. He particularly recommended socio–genealogical studies on the formation and transformation of classes and professional groups. There were even more clear–cut ideas about the sociological application of genealogical databases. Robert Sommer proposed a very ambituous agenda for psychological family research. Besides his psychiatric research, Sommer worked in the field of experimental psychology, with a special interest in aptitude tests. The approved methods of individual psychology, he claimed, had to be complemented by an examination of the tested person’s family. The combination of family research and psycholoigcal assessment would generate a new, exact social psychology. Sommer regarded it as the crucial problem of modern society that many people were not choosing the vocation adequate to their inborn ability. He held that it was the “task of social psychology, and of a psychological social policy which has to be derived from it, to guide the natural dispositions of the individuals, as far as possible, into those professions where they can develop the greatest effectivity in the interest of society.”35 Sommer conceded that his science was still far from the diagnostic knowledge needed to accomplish this aim. However, his objectives clearly show that the idea of genealogical databases was closely related to a technocratic vision of society. Sommer hoped that sophisticated collections of family records were to provide a tool for the professional selection of individuals. On the whole, this implied a vision of control over the process of selection that shaped human society. There was also a strong affinity between the idea of “scientific genealogy” and the interest in population development. Fears about stagnating birth rates as a consequence of urban life began to spread in the 1890s. Kekulé strongly suggested that genealogists should turn to demographic problems since “in principle, all questions about population development are genealogical questions”.36 In fact, the genealogical perspective implied a novel concept of population. Rather than just comprising the living inhabitants of a nation, the genealogical concept of population included the past generations. This idea was most clearly expressed in the plans for a national genealogical register propagated by Lorenz’s followers. The genealogical network they envisaged was to represent the complete ancestry of the nation. Rather than as an association of citizens, the population would appear as an organic entity – a virtual body of the people or, as the German eugenicists expressed it, a Volkskörper.

“Genealogical awareness” The plans for a national genealogical register, thus, had a double meaning. On the one hand, it was to serve eugenicists to identify abnormal hereditary dispositions, on the other hand, the reconstruction of common ancestry promised to strenghten the sense of national community. The latter aspect was related to the fact that such large collections could only be accomplished with the cooperation of amateur genealogists. As Breymann had stressed, this mobilization implied the chance to popularize biological thinking. In a similar form, this idea had been expressed by Lorenz. Though he emphatically claimed that genealogy was a science, he decidedly did not want 35 36

Sommer (1907), p. 6. Kekulé (1900), p. 109.

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to turn it into a method for experts. It was up to the “scientific genealogist” to advise laymen how to collect the right material for their descendants.37 Lorenz regarded the interest in family history as a part of the human condition and an indicator of cultural standard. The growing popularity of genealogy, thus, was a sign of true social progress. Lorenz hoped that due to “raising living conditions”, more and more people would develop an interest in their own descent or, as he called it, “genealogical awareness”. Genealogical awareness, he hoped, was to become an effective weapon against all socialist and egalitarian evils. This idea was eagerly taken up by his followers. As one co–founder of the Zentralstelle put it with monarchist zeal, genealogy had a “high moral value” since it formed “a strong bulwark against subversive activities of all kinds” by strenghthening the sense for family and fatherland and therefore supporting a state–loyal attitude.38 Yet, Lorenz had also emphasized that the practice of family research would help to popularize the insights of modern biology. With the gradual integration of genealogical organisations into the eugenics movement, this aspect became more and more predominant. Sommer developed this idea with a remarkable consequence. Like Lorenz, he strictly rejected all proposals for compulsory eugenic measures, labelling them as true expressions of contemporary moral degeneration. All the more, the “racial decay” of modern society had to be fought through eugenic education. And the most effective way to spread the awareness of imminent degeneration was “the general penetration of the people with the ideal of natural aristocracy”. 39 The idea of aristocracy, Sommer demanded, had to be reconstructed in it original sense: as a feeling of responsibility for the own family’s specific qualities. The practice of genealogy, thus, was the best way to make people internalize the ideas of eugenics. After World War I, this aspect gained importance. The position of genealogy as a method for exact studies on human heredity became increasingly questionable. On the one hand, it had turned out that most hereditary phenomena were not explicable in simple Mendelian terms, on the other, twin research emerged as a new standard method of human genetics. In 1930, the eugenicist H.W. Siemens stated that family research had not yielded the scientific results once hoped for – and most likely never would. Nevertheless, he stated that it was indispensible for the cause of eugenics, since the popular practice of genealogy created an emotional connection to family and people, accordingly providing the most “unobstrusive” and continuous way of propaganda for racial biology.40 Genealogy, after all, has never only been a method for the science of heredity. As a practice that is deeply connected with the self–definition of individuals and collectives, it imposes a specific dynamic on the study of human heredity. Genealogy provides the thinking about heredity with a moment of historicity, but it also brought about a biologization of history. Lorenz defined genealogy as “the bridge on which historical and natural science meet”, and he was right to stress the importance of this encounter.41 The emergence of a hybrid field between history, sociology and biology proved to be most momentous for the early 20th century human sciences. The 37 38 39 40 41

Lorenz (1898), p. 139. Ueltzen–Barkhausen (1905), p. 10. Sommer (1907), p. 221. Siemens (1930). Lorenz (1898), p. 26.

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genealogical perspective did not only foster selectionist and organicist concepts of history and society. It also changed the scope of eugenics. Only the idea that population was an organic body and that the hereditary abilities of families, nations and races were the true basis of historical development, provided eugenics with its particular authority and assertiveness.

Epilogue: Total genealogy The high time of genealogical collections was to come after World War I, when eugenics became an integral part of public health policy. Representatives of the Zentralstelle were involved when the Reichsgesundheitsamt discussed the foundation of a National Institute for eugenics and human heredity in 1923.42 These plans eventually led to the establishment of the Kaiser Wilhelm Institute for Anthropology, Human Genetics and Eugenics, an institute not devoted to genealogical collections. Such databases flourished in Ernst Rüdin’s genelogical department at the Deutsche Forschungsanstalt für Psychiatrie, which assembled the largest collection of psychiatric family histories in Germany. With support from the state of Saxony, the eugenicist Rainer Fetscher built up a database of the “inferior” which was designed to comprise all Saxon convicts and their families. The Bavarian ministry of Justice established a Criminal Biological Record Office in 1924, which had the right to access prisoner’s files, but also to school and parish records. 43 The anthropologist Walter Scheidt called for the replacement of classic anthropological race typologies by “population biology”, a method based on a complete genealogical survey of rural populations. Scheidt was also an outstanding figure among those eugenic visionaries who demanded a national genealogical register, as he proposed a central statistical office assembling all medical and juridical records on the whole population and its complete known ancestry. 44 While such plans never materialized, Scheidt’s concept of local population studies became seminal for the program of a “racial survey of the German people” carried out in the early 1930s. But the heyday of “population genealogy” was still to come under National Socialism. The Nazi Peasant’s League (Reichsnährstand) and the Nazi Teacher’s Association launched a project aiming at the total working–up of all German parish registers up to the 17th century. The material was to be issued in “kinship books” for every German village designed to raise the rural population’s awareness of common racial descent. The second – and more important – plan was to create a comprehensive database by using a filing system developed by Scheidt.45 Card indices would allow to trace every individual’s ancestry, including the accessible information about their health. Reichsnährstand officials hoped to link up these files with the material collected in the eugenic “inventory” campaign of the psychiatric asylums. They were also in touch with the Nazi authorities who were trying to detect people of “gypsy” descent. Originally, the Reichsnährstand had started the systematic registration of genealogical sources for similar reasons: it was meant to simplify the procedures connected with the proof of “purely Aryan” descent. The project, thus, combined the

42 43 44 45

Weingart et al (1992), p. 241. Weindling (1989), p. 385. Scheidt (1930). Klenck/Kopf (1937).

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idea of pure descent and the eugenic vision of total control over genetic defects. In Foucauldian terms, the conjunction between the myth of the blood and the control of heredity was carried to the extreme. Total genealogy was both to create a sense of racial aristocracy in all “racially pure” Germans and to defend the racial community against all threats to the “hereditary health”.

Bernd Gausemeier, MPI für Wissenschaftsgeschichte, Berlin, [email protected]

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References Becker, Peter. 2002. Verderbnis und Entartung : eine Geschichte der Kriminologie des 19. Jahrhunderts als Diskurs und Praxis. Göttingen. Breymann, Hans. 1912. Über die Notwendigkeit eines Zusammengehens von Genealogen und Medizinern in der Familienforschung. Archiv für Rassen– und Gesellschaftsbiologie 9: 18–29. Devrient, Ernst. 1897. Die älteren Ernestiner. Eine genealogische Charakteristik. Vierteljahresschrift für Wappen–, Siegel– und Familienkunde 25: 1–133. Foucault, Michel. 1983. Der Wille zum Wissen. Sexualität und Wahrheit. Vol. 1. Frankfurt/M. Gruber, Max von, and Ernst Rüdin, eds. 1911. Fortpflanzung, Vererbung, Rassenhygiene. Illustrierter Führer durch die Gruppe Rassenhygiene der Internationalen Hygiene–Ausstellung 1911 in Dresden. München. Heydenreich, Eduard, ed. 1913. Handbuch der praktischen Genealogie. Leipzig. Jacobi, Maximilian. 1834. Ueber die Anlegung und Einrichtung von Irren–Heilanstalten mit ausführlicher Darstellung der Irren–Heilanstalt zu Siegburg. Berlin. Stradonitz, Stephan Kekulé von. 1905. Ziele und Aufgaben der wissenschaftlichen Genealogie (1900). In: Ausgewählte Aufsätze aus dem Gebiete des Staatsrechts und der Genealogie, edited by Stephan Kekulé von Stradonitz. Berlin. 103–128. Klenck, Willy and Ernst Kopf. 1937. Deutsche Volkssippenkunde. Berlin. Lorenz, Ottokar, and Leopold von Ranke. 1891. Die Generationenlehre und der Geschichtsunterricht. In: Die Geschichtswissenschaft in Hauptrichtungen und Aufgaben. Vol. 2, edited by Ottokar Lorenz. Berlin. ———. 1898. Lehrbuch der gesammten wissenschaftlichen Genealogie. Stammbaum und Ahnentafel in ihrer geschichtlichen, sociologischen und naturwissenschaftlichen Bedeutung. Berlin. Nukaga, Yoshio. 2003. Tracing Genealogical Methods: The Development and Use of Family Trees in the Case of Hereditary Chorea. Paper presented at the MPIWG Berlin. Ribot, Théodule. 1884. L' hérédité psychologique. 2nd ed. Paris. Rohde, Friedrich. 1895. Über den gegenwärtigen Stand der Frage nach der Entstehung und Vererbung individueller Eigenschaften und Krankheiten. Jena. Rüdin, Ernst. 1911. Einige Wege und Ziele der Familienforschung mit Rücksicht auf die Psychiatrie. Zeitschrift für die gesamte Neurologie und Psychiatrie 7: 487–585. Scheidt, Walter. 1930. Ein bevölkerungsbiologisches Reichsarchiv. Allgemeines Statistisches Archiv 22: 561– 568. Siemens, Hermann Werner. 1930. Bedeutung und Methodik der Ahnentafelforschung, Archiv für Rassen– und Gesellschaftsbiologie 24: 185–197. Sommer, Robert. 1901. Diagnostik der Geisteskrankheiten für praktische Ärzte und Studierende. Berlin/ Wien. ———. 1907. Familienforschung und Vererbungslehre. Leipzig. ———. 1913. Familiengeschichtliche Quellenkunde im Gebiete der Psychiatrie und Anthropologie. In: Heydenreich (1913): 388–398. Tille, Armin. 1913. Genealogie und Sozialwissenschaft. In: Heydenreich (1913): 371–388. Ueltzen–Barkhausen. 1905. Bericht über die Entstehung, Gründung und bisherige Tätigkeit der Zentralstelle. Mitteilungen der Zentralstelle 1: 8–16. Virchow, Rudolf. 1886. Deszendenz und Pathologie. Archiv für pathologische Anatomie und Physiologie und für klinische Medizin 103: 1–14, 205–215, 413–436. Weinberg, Wilhelm. 1911. Vererbungsforschung und Genealogie. Eine nachträgliche Kritik des Lorenzschen Lehrbuches. Archiv für Rassen– und Gesellschaftsbiologie 8: 753–760. Weindling, Paul J. 1989. Health, race and German politics between national unification and Nazism, 1870– 1945. Cambridge. Weingart, Peter, Jürgen Kroll, and Kurt Bayertz. 1992. Rasse, Blut und Gene. Geschichte der Eugenik und Rassenhygiene in Deutschland. Frankfurt/Main. Zentralstelle für deutsche Personen– und Familiengeschichte. 1908. Mitteilungen der Zentralstelle 4.

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Bismarck the Tomcat and Other Tales: Heredity and Alcoholism in the Medical Sphere, The Netherlands 1850–1900 Stephen Snelders, Frans J. Meijman and Toine Pieters

1. Introduction A tale Once there lived in Paris a baker who was well known and appreciated in his neighbourhood for his very excellent pies and cakes. His business was therefore thriving. Not only his customers enjoyed the baker’s products, so did various rodents whose habitats overlapped those of the humans. To assert his dominance over these animals the baker had taken in another lodger: an Angora cat. The fierceness of this tomcat earned him, in these days after the catastrophal defeat of the French by the German armies, the name Bismarck. Bismarck justified his name and turned out to be an excellent rat–catcher. Not only rats were to his liking, though. Bismarck also took a fancy to the delicious pastry produced by the baker. This in itself was not a problem. But unfortunately the baker, who had grown up in a culture where the use of alcohol was considered normal and necessary, also made pastry of which this drug was one of the ingredients. Especially his rumcake was (deservedly) famous. It was also appreciated by Bismarck the tomcat. After consuming one of these cakes the people of the neighbourhood would see Bismarck walking and tip–toeing in a pleasantly drunk condition. They did not realize that this was the first step in a fast deterioration of Bismarck’s physical and psychological condition. Having taken a liking to the rumcakes, Bismarck became, in short, a dipsomaniac. And where in the beginning his reaction to the alcohol had been one of pleasant drowsiness, now he would react less favourably to the drug. He became ill–tempered, more and more wild, and finally mad until he started to attack, not the rodents, but the humans, and had to be put out of his misery.

This late nineteenth century story reads as the reverse of Egar Allan Poe’s The Black Cat, the story of the decline and fall of an alcoholic who ends up attacking (and is revenged upon by) his cat. Admittedly Poe’s story is far more chilling. But then his story was literature and published in a literary journal. However, this originally German tale of the tomcat Bismarck was meant to serve the progress of science. It was used instrumentally in the March 1901 edition of the Dutch medical weekly, the Geneeskundige Courant voor het Koninkrijk der Nederlanden (‘Medical Journal for the Kingdom of the Netherlands’) to support the claim that alcoholism presented a most serious health problem that deserved imminent medical attention.1 In an allegoric sense the story warns against the dangers of alcohol abuse, and against the low threshold between an ‘innocuous’ use of alcohol in daily life and the degeneration into madness. Around 1900 it was generally accepted that chronic alcoholism could be inherited or transmitted to descendents as morbid nervous predispositions. Together with tuberculosis and syphilis, alcoholism was regarded as a major cause of degeneration and as such defined as a public 1

GC 55 (1901), no. 7.

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threat that should be curbed by public health measures. In a number of European and American countries support was mounting for ‘hard–line’ policies of eugenics, such as marriage restrictions and involuntary sterilizations. However, as we will discuss in this paper, other scenarios of fighting the ‘alcoholism peril’ were also enacted. To understand how these scenarios took shape in Dutch medicine in the second half of the nineteenth century, we need to take into account the diversity and fluidity of medical and public debates around degeneration and heredity.

2. Research focus and questions This paper reports on preliminary findings of the on–going research project on ‘Heredity and Concepts of Illness and Health in the Netherlands in Modern History’. 2 The focus here is on the second half of the 19th century. The paper investigates the roles played by notions of heredity in Dutch medicine, by analysing discourses on prevention and treatment of alcoholism and alcohol abuse in the Dutch medical literature. It is commonplace in the historical literature to relate medical positions on alcoholism in the 19th century to evolving concepts of degeneration and heredity.3 Two recent volumes are exemplary of the general sort of accounts of nineteenth century approaches to degeneration and hereditary diseases such as alcoholism. Craig Heron, in his wonderful history of alcoholism in Canada, writes about the last decades of the century: The growing numbers of physicians who believed inebriates had inherited their “craving” for alcohol shared the environmentalist concern that these degenerates be confined under medical care at an early stage, in the hope of weaning them from their destructive habit. Yet most of those in the medical profession who believed in the crucial importance of heredity saw little hope for drunkards in the end and took limited interest in them. Doctor’s optimism about institutional treatment for any kind of mental–health problem was waning by the end of the nineteenth century. Prevention became more important than cure.4

The same position is held by John W. Crowley and William L. White in their history of the first U.S. asylum for alcoholics, the New York State Inebriate Asylum in Binghamton, which opened in 1864. They write: “Whereas ‘made’ drunkards were capable of responding to the care of moralists and reformers, ‘born’ drunkards should be placed in the hands of professional authorities who would control them by legal or medical means. The appropriate object of such control was the congenital dipsomaniac, whose chances of full recovery were deemed to be small but whose threat to society was deemed to be large.”5 In both case studies alcoholism shows double faces: cause and product of degeneration, vice and malady.6 The readings above are in line with the idea, that medical hereditarianism in the 19th century transforms from ‘soft’ into ‘hard’, i.e. that it became increasingly deterministic and fatalistic with concomitant implications in medical and public domains. According to some 2 3 4 5 6

Earlier findings are reported in Snelders (2003); Snelders and Pieters (2003); Idem (in press). For an overview of the rise of this relationship between alcoholism and degeneration, cf. Bynum(1984); Sournia (1990). Heron (2003), 143. Crowley and White (2004), 77. White (2003).

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authors this happened because this hardening conveniently explained the failure of medicine to find cures for the problem of alcoholism and other diseases.7 But the thesis of a ‘hardening’ hereditarianism has also been explained by what has been called ‘the hardening face of nature’ in scientific and cultural understandings. “Medical hereditarianism assumed much stronger form in the 19th century, when nature turned from a benevolent, purposeful entity to assume the harsh face of physical necessity.”8 These claims are supported by evidence from the medical literature of the period. Within this perspective, the tale of Bismarck the tomcat can be read as representing the serious social and biological problem of the incurable, degenerative hereditary disease alcoholism, as represented in one of the higher mammals. We consider alcoholism to be an exemplary case to study the dynamics of medical hereditarianism, since alcoholism was considered to be an important cause and consequence of degeneration, shown in mental diseases. As Gianna Pomata has written: “It is significant that most of the 19th century doctors’ interest in hereditary diseases shifted from gout, the patrician malady, to insanity – a disease considered to be endemic at the other end of the social ladder.” 9 We therefore focus on the questions how, to what extent, and why knowledge of inheritance was anchored in medical concepts and practices around alcoholism. By studying five leading Dutch medical journals, this paper looks for changes in the medical conceptualization and practical approaches around alcoholism. The main goal is the closer inspection of the fore–mentioned assumptions regarding the transformation of hereditarianism, in 19th century Dutch medicine. The paper makes a distinction between two analytical levels. First there is the level of theory and conceptualization within medicine; second, we distinguish the level of application and practical approaches. Medicine is a field in which a logical coherence between these two levels often seems to be lacking: it is a science as well as an art.10 In medical practice ‘elastic’ approaches dominate, given the unruly nature of medicine. The semiotic status of hereditarianism, that we label ‘plastic’, involves flexible boundaries between the respective influences of ‘nature’ and ‘nurture’. Connecting concepts of hereditary disease to that of individual constitutions did not necessarily have to entail a desire on the part of physicians to rationalize and to excuse their inability to treat a range of persistent chronic maladies, including alcoholism. It could also justify an elastic approach in treatment, undisturbed by hereditary determinism. Discourses of conceptualization as well as practices of treatment and prevention have both to be contextualized. On both these levels more broader cultural themes and social beliefs, in this case especially that of degeneration, become apparent. In this paper a contribution to the historical assessment of the connections between heredity and alcoholism in the medical domain between 1850 and 1900 will be made by a focus on an unexplored source: medical journals and publications from the Netherlands. The Dutch are not regarded as pioneers of medical and political activities in this field. It might therefore be that a focus on the Netherlands shows developments more representative of overall developments in the Western hemisphere. In following and analyzing the evolution of ideas and views of Dutch medical doctors on heredity and alcoholism we will particularly focus on diffusion patterns as well 7 8 9 10

Weiss (1987), 19; Dowbiggin (1997), ix–x; Waller (2002); Idem (2003). Pomata (2003), 151. Ibid., 150. Weatherall (1995).

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as differentiation and transformation of: 1. medical thought about hereditary transmission; 2. concurrent medical treatment and prevention practices regarding alcoholism. All the five major Dutch medical journals were selected for research. These are: 1. The Nederlandsch Tijdschrift voor Geneeskunde (‘Dutch Journal for Medicine’; in this paper abbreviated as NTvG). This journal appeared weekly and was (and still is) the major journal for Dutch physicians, in the nineteenth century being the journal of the Dutch Society for the Advancement of Medicine. It was created in 1857 after a fusion of nearly all the existing medical journals, with one significant exception (see no. 2). NTvG was politically oriented to the liberals; it counted leading hygienists who advocated public health policies among its contributors. 2. The Geneeskundige Courant voor het Koninkrijk der Nederlanden (‘Medical Journal for the Kingdom of the Netherlands’, in this paper abbreviated as GC) was the one medical journal of importance that remained outside the fusion to NTvG. First published in 1847 and appearing weekly as well, it was politically more oriented to the conservatives. 11 3. The Psychiatrische Bladen (Psychiatric Papers’ in this paper abbreviated as PB) started publication in 1883 and were devoted to subjects of psychiatry and neurology, as reflected in the change of its name in Psychiatrische en Neurologische Bladen (‘Psychiatric and Neurological Papers’, in this paper abbreviated as PNB) in 1897. 4. The Geneeskundige Bladen uit Kliniek en Laboratorium voor de praktijk (‘Papers from Clinic and Laboratory for Medical Practice’, in this paper abbreviated as GB) appeared for the first time in 1894. They aimed to present the contemporary discussions in clinical and laboratory research. 5. Finally, the Tijdschrift voor Sociale Hygiëne en Openbare Gezondheidsleraar (‘Magazine for Social Hygiene and Public Health’, in this paper abbreviated as TSH) was started in 1899 as the ‘mouthpiece’ of the hygienist movement. Its first two volumes can only be taken into account in our analysis.

3. Concepts At the start of the 20th century, Dutch doctors in general took the concept of hereditary predisposition to alcoholism for granted. Research reports in the Dutch medical journals translated from the German, French, and British literature, that appeared over the course of the 1890s, and were based on family–tree research and statistical studies of the inmates of asylums, seemed to establish beyond doubt certain facts. Alcoholism (or dipsomania) had a strong tendency to be hereditary. However in most cases it changed form into other mental diseases. Inmates of asylums were to a large, but debated degree (42,6 % according to one French study) hereditary insane due to the influence of alcoholism. The hereditary influence of alcoholism expressed itself in different forms: directly as delirium tremens and periodic bouts of alcohol 11

On the Dutch medical press until 1857, see Delprat (1927). On NTvG: Idem (1932). Unless mentioned otherwise, references to NTvG are to the second series, starting in 1865. On the politics of NTvG and GC: Houwaart (1991), 224.

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abuse; indirectly in other psychoses; and biologically in ‘deprivation of the progeny’ and extinction in the third and fourth generation. This pathological form of heredity could be transmitted by inheritance due to chronic alcohol abuse of one or both of the parents, or due to alcohol intoxication of the germ–plasm during sexual intercourse, or due to what we would call foetal conditioning by an alcoholic mother.12 The following excerpt from an article by the physician J. Kat, an active member of the temperance movement, on the influence of alcohol on posterity was fairly typical of the articles in medical journals around 1900. ‘Das grauige Kapital [the drab capital, i.e. the influence of alcohol abuse] shows on the basis of health statistics produced by researchers in all parts of the world, that retardation, imbecility, idiocy, epilepsy, neurasthenia, criminality, are for the greatest part grounded in alcohol use of parents before and during the conception.’13

A similar but in terms of degenerative heredity more articulate view can be found in the writings of Amsterdam gynaecologist Hector Treub. Treub´s 1900 New Year speech to Amsterdam doctors, subsequently printed in GB, has been regarded as an early example of eugenic thought in the Netherlands.14 According to Treub, there existed five ‘laws of heredity’: 1. direct inheritance from the parents; 2. inheritance of traits from an earlier generation, i.e. atavism. 3. indirect or collateral inheritance from a collateral family line. 4. ‘initial inheritance’ from the condition of the parents during cohabitation. For example a drunk parent increased the chance for idiocy in the child. 5. inheritance of influence, or telegony: for instance a white woman cohabitating with a negro would give birth to a mulat. If she later cohabitated with a white man, the child would again be a mulat. As far as this fifth law is concerned Treub was rather sceptical. In his speech he mainly engaged himself with the first three laws and he came to some far–reaching conclusions, as we will see. In fact, his expositions were meant to stress the necessity of medical investigation and advice on heredity before marriage. This might indeed seem to point to a hard form of hereditarianism, emphasizing genetic determinism and fatalism. But when we study these ‘laws’ more thoroughly on the level of conceptualization then the fourth law appears to be deviating from the others. According to this law hereditary characteristics could be acquired during lifetime. Treub’s example can be read as a plastic and more ‘soft’ form of hereditarianism. 15 Before further discussing the nature of Treub´s hereditarianism and whether or not Treub’s concept of heredity was typical of Dutch medical thought around 1900, we will explore the transformations of medical thought about hereditary transmission since the 1850s. The underpinning of Treub’s laws of inheritance was based on the results of a research method that had been popular for a long time: family–tree research. For instance, Treub used a family–tree taken from the most influential medical authority on heredity in the second half of the 19th century, French psychiatrist Bénédict–Augustin Morel. Treub used this tree to show how chronic 12 13 14 15

For example: NTvG 27 (1891) II, 328–330; Ibid. 31 (1898) II, 554; Ibid. 34 (1898) II, 496; Treub (1900). NTvG 40, I (1904), 108. Noordman (1989). Treub (1900).

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alcoholism could lead in the following generations to idiocy and madness. 16 Treub’s exposition was neither new as far as the methodology was concerned nor with regard to the research data on display. The investigation of family trees had been pioneered by Franz Wilhelm Lippich in Laibach. In a study published in 1834 Lippich had given a statistical analysis of two hundred alcoholic patients, and found their offspring to be generally more unhealthy than the general population of Laibach.17 Morel’s treatises on degeneration (1857) and mental illness (1860) stimulated similar studies. His own family–trees famously represented the hereditary degeneration caused by alcoholism: in the first generation moral stupefaction, brutalisation, sapping of the body; in the second: hereditary drunkenness, mania and palsy; in the third: hypochondria, suicidal tendencies; and in the fourth finally: mental defections, idiocy, and premature decease, ultimately leading to extinction. In Morel alcoholism shows its double faces as vice and malady, as cause and consequence of hereditary predispositions and vicious environments. 18 But did Morel offer anything sensationally new in the conceptualization of alcoholism? The impact of his work may have been more due to the elaboration of existing notions than to a revolutionary new approach. Even before Morel published his volumes, leading Dutch psychiatrist J.N. Ramaer emphasized in 1852 that inebriety was the cause of hereditary mental diseases, primarily idiocy. Ramaer referred to Lippich’s evidence, but also to the knowledge of this hereditary degeneration among the ancient Greeks.19 What seems to happen over the course of the next half a century is that this theory of degeneration is repeated again and again, occasionally confirmed by new family studies. We did not find any divergent opinions. In Dutch medical literature from the 1850s until Treub’s lecture Morel was again and again cited as authority and evidence for the double faces of alcoholism. In his public health manual of 1872 the leading hygienist Ali Cohen incorporated the views of Morel.20 So did twenty years later psychiatrist Pierre F. Spaink in his 1892 monograph on alcoholism.21 Jan Broers in his M.D.–thesis on alcoholism, morphinism and chloralism, the first Dutch medical treatise on the broader subject of addiction, lamented that especially chronic alcoholism had the ‘important disadvantage’ that posterity had to suffer for the sins of its ancestors (in Morelian degeneration vice becomes the disease of the descendants, and disease becomes the vice).22 GC regularly referred in its columns to the effects of a ‘Morelian degeneration’, for instance in 1870 by publishing the research results and conclusions of Morel’s pupil Doutrebente on hereditary madness, or three case–studies of another French physician, Taguet, in 1877. 23 Neither GC nor NTvG ever doubted the scientific truth of Morelian degeneration, nor did any of the other magazines. The discussions about evolution and heredity that arose after the publication of The origin of species in 1859 did not exert any influence on this. Family–tree research done after Morel was 16 17 18 19 20 21 22 23

Ibid., 34. Bynum (1968), 175–176. Morel (1857); Idem (1860). On Morel: Coffin (2003). Ramaer (1852), 97–100. Ali Cohen (1872), 156. Spaink (1892), 19–20. Broers (1886), 125. GC 24 (1870), no. 10; Ibid. 31 (1877), no. 49, 50, 51.

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regularly cited and always confirmed his conclusions: only the exact percentage of the expression of predisposition wavered, but this was not taken in any sense as invalidating the theory. GC reported in 1884 on German research, in which among idiots in c. 10 % of the patients evidence was found of chronic alcohol abuse or dipsomania.24 In 1899 the news was brought that a research in Bonn had uncovered 709 descendants of a ‘well–known’ alcoholic woman who had died in 1800. Of these descendants 462 had become murderers, criminals, beggars, or prostitutes, and had cost the German government six million francs.25 Leading articles once more explained the degenerative effects of alcoholism on succeeding generations in 1895 and 1900. 26 Psychiatrist A. Tellegen did not doubt in PB in 1884 the hereditary consequences of alcoholism, but recognized that figures on the hereditary etiology of madness differed extremely, from 4% to 90%! But research had one severe problem: persons could have the predisposition to madness, but die before this predisposition was expressed.27 NTvG also endorsed the relationship between hereditary degeneration and alcoholism by citing family studies. R. Demme from the University of Bern saw Morel confirmed in 1890 in his study of twenty families, in which one or both of the parents were alcoholics, over a period of 12 years. Of 57 children, only 10 (17,5 %) had had a normal development. 25 had died in the first weeks after birth, and 22 showed congenital defects: defective physical development, chorea, epilepsy, idiocy. In a group of 10 families with moderate drinkers as parents, 50 of the 61 children had grown up healthy (81,9 %). 28 In 1895 NTvG reported the conclusions of French psychiatrist Legrain: in his asylum 42,6 % of the inmates were hereditary insane due to the influence of alcoholism.29 This research was in 1898 confirmed by the study of 1200 cases of hereditary insanity by Farquharson in Britain. Dipsomania had according to Farquharson a strong tendency to be hereditary, although in most cases it changed form into other mental diseases.30 To the reader of GC in 1901 then, the story of Bismarck the tomcat might have had implications that went beyond the moral decline of one individual cat. Most likely, Bismarck’s decline symbolized the fate of a whole race of dipsomanic cats. It would have been as recognizable to a reader of GC in the 1850s. Which makes us wonder, how medical thought about alcoholism and degeneration relate to the dynamic and controversial discussions about biological evolution in this period. Connecting degeneration theory and evolutionism started in the Dutch medical literature in the early 1880s. This means that the impact of Darwin on the medical framing of alcoholism and degeneration was almost non–existent in the 1860s and 1870s. Darwin seems not to be relevant in any way to this framing, for which Morel was sufficient authority. The theory of degeneration was as acceptable to those physicians who showed a positive stand towards the new evolution theories, 24 25 26 27 28 29 30

GC 38 (1884), no. 6, 7. Z., ‘De afstammelingen van een alcoholist’, GC 40 (1899), no. 40. Niermeijer, ‘Alcohol en alcoholisme’, GC 49 (1895), no. 31; A.N.J. Hazedoes van Almkerk, ‘Alcoholisme en de houding van medici te dien opzichte’, GC 54 (1900), no. 10. A.O.H. Tellegen, ‘Eenige beschouwingen over krankzinnigheid, hare oorzaken en hare behandeling’, PB 11 (1884), 5–46. S.K. Hulshoff, review of R. Demme, ‘Ueber den Einfluss des Alkohols auf den Organismus des Kindes’, NTvG 27 (1891) II, 328–330. NTvG 31 (1895) II, 554. NTvG 34 (1898) II, 496.

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as to their more traditional and religiously minded colleagues. We must remember that Morel himself was not a Darwinist in any sense. He was a Catholic who framed degeneration in terms of the ‘Fall of Man’.31 The great authority for the physician on these matters in the 1880s was not any (Neo–) Darwinian biologist, but one of their own, the German pathologist Rudolph Virchow. Virchow was not only the dominant figure in establishing and transforming German laboratory medicine in the second half of the 19th century. He was a physician profoundly interested in relating clinical research to medical practice. Virchow was moreover one of the leaders of the German liberals and his political activities almost led him into a duel with Bismarck – the statesman, not the cat.32 NTvG discussed in 1882 a lecture of Virchow on Darwin and anthropology, in which he cautioned against a rushed generalization and transmission of Darwin’s theories from animals to humans.33 The great challenge to conceptualizations of alcoholism and heredity in the medical domain was provided by August Weismann´s new concept of the ‘germ plasm’. Weissman´s theory had a major impact on the redefinition of the concepts of heredity at the end of the 19th century. Starting from the question of how the germ plasm with the inherited characteristics could reproduce itself, Weissman conceptualized the soma, the body, as a mere transport vehicle for the germ plasm. In doing so he separated the problem of heredity and the problem of growth and differentiation. Changes in the soma were not transmittable to the germ plasm. Weismann ‘proved’ in 1883 that traits required during one’s lifetime could not be inherited by descendants. However this proof did not have the impact in the medical domain historians of biology have often accorded to it.34 W. Koster discussed the matter in 1886 in NTvG. He called Weismann’s Bedeutung der sexuellen Fortpflanzung für die Selektionstheorie a ‘phantastic–speculative evolution theory’, that seemed to contradict established pathological and clinical knowledge. Virchow gave acclimatisation as an example of the inheritance of acquired characteristics, while Weismann was of the opinion that the acclimatised individual was already, by chance, adapted to his new environment. Koster thought that Weismann’s idea of the continuity of the germ plasm explained much, for example the inheritance of the ‘Jewish type’. But if Weismann was right there could not exist ‘infectious and hereditary’ diseases. However, syphilis and tuberculosis were regarded as ample proof for the claim of the hereditability of these common diseases. 35 A year later, a leading article in GC clearly stated that under specific conditions acquired characteristics could be inherited. Why this happened was still a mystery; that it happened, beyond doubt. Weismann notwithstanding, the article informed its readers that: “The acquired character [developed by education, environment, etc, the ‘envelope’ of the true, inherited character] later changes again into the inherited character, because it is inherited by the descendants”. GC specified that not the characteristics themselves, but the predisposition to develop them was inherited. Expression depends on circumstances. The more often characteristics occurred in 31 32 33 34 35

Huertas (1992). Ackerknecht (1953). NTvG 18 (1882) I, 823–824. On the discutable idea that Weismann disproved the inheritance of acquired characteristics: Bowler (1988). W. Koster, ‘Ontwikkelingsleer en ziektekunde’, NTvG 22 (1886) I, 341–349.

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family–trees, the more chance that they would return in later generations. A person with a powerful imagination could transfer his acquired characteristics more easily to his descendants: the example given was that of an alcoholic father. Inheritance of equal characteristics occurred, but more often polymorphism, the unequal distribution of dispositions. Expression of these predispositions was dependent on circumstances: for example shock, misery, strain. Under the right circumstances the predisposition could even express itself as genius, as in the case of Schopenhauer. Morel’s family–trees once again figured prominently to demonstrate the mechanisms and patterns of hereditary transmission.36 This perspective of plastic expression was compatible with existing medical traditions. It did not conflict either with the new theory of evolution as expressed by Darwin. It was recognized that Darwin himself believed in the inheritance of acquired characteristics. 37 But even to Weismann’s ideas the medical community was not unfavourably disposed. In 1891 Zwaardmaker, one of the editors of NTvG, called the theory of Weismann a hypothesis of great value, a progress in the direction of a mechanical explanation of nature. There was much that seemed to support Weismann, such as the occurrence of atavisms and of morphological characteristics that were insurmountable for the individual, but not for the species as a whole. Still, the theory was held far from proven.38 Koster in turn undertook in 1893 a review of Weismann. There is no evidence for the inheritance of acquired characteristics in a positive sense, he wrote, but Weismann did not deny inheritance in a negative sense: agents as alcohol or virus syphiliticum could damage sperm cells and lead, when sperm and egg mingled, to a spontaneous poisoning of the germ plasm. 39 This ‘inheritance in a negative sense’ became in the course of the 1890s an equally satisfactory and convenient explanation for the hereditary degeneration caused by alcoholism as the inheritance of acquired characteristics had been. What we have here is a typical form of medical eclecticism, producing a workable explanatory tool that met a need, based on doctor’s experiences and seemingly proven by empirical family–tree studies. Possible inconsistencies between the fore– mentioned biological concepts were noted but amended and adjusted to produce a medical argument consistent with a perspective of plastic expression of hereditary predisposition. 40 It is of importance to look at the adjustment and amendments of knowledges of heredity by doctors from a functionalist perspective, and not from the perspective of consistency with developments in the scientific sphere. At the end of the century there was consensus about the three possible mechanisms by which alcoholism could be inherited and which explained both Treub’s ‘laws of inheritance’ and the onset of the degeneration process, i.e. the transformation of a metabolic disorder into morbid nervous dispositions. As the authoritative Swiss researcher and temperance activist Auguste– Henri Forel explained in 1892 on the Fourth International Congress against alcohol abuse, alcohol itself was a toxic agent that led to degeneration of progeny.41 Alcohol abuse of the mother could 36 37 38 39 40 41

‘De overerving van zenuw– en zielsziekten’, GC 41 (1887), no. 22, 23, 24. ‘De overerving van verworven eigenschappen’, GC 43 (1889), no. 43. NTvG 27, (1891) I, 418–420. NTvG 29 (1893) II, 293–311 For similar developments in France: Pinell (2001). Le Rütte Jr., ‘Het vierde internationale congres tot wering van het misbruik van sterken drank’, PB 11 (1892), 220–221.

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lead to poisoning of the foetus in utero. Blastotoxie occurred when a child was conceived while one or both of the partners in the sexual act were drunk. A third mechanism, Blastophtorie, functioned when one of the parents was a chronic alcoholic whose germ plasm was seriously poisoned by the alcohol.42 While in accordance with neo–Darwinism, this plastic perspective on hereditary transmission also fit into Morelian degeneration schemes. Even the positive implications of the inheritance of acquired characteristics were saved: in the course of generations healthy living could restore the vitality of the germ plasma.43 J. van Rees, physician and leading prohibitionist, explained in 1902 in a propaganda brochure against alcohol use that the germ plasm could regenerate in the third or fourth generation, on the condition that it was mixed with undamaged plasm. 44 Forel furthermore considered, as NTvG explained its readers in 1895, that man had a ‘plastic’ disposition: the expression of the hereditary disposition could take different forms, depending on opportunity and exercise, and did not have to take a pathological form per se. 45 Given the plastic nature of attempts by Dutch doctors at adjusting theories on heredity to medical problems this raises the question to what conclusions regarding alcoholism treatment policies the transforming thoughts about hereditary transmission led.

4. Approaches: public health Dutch doctors regarded alcohol abuse and alcoholism both as an individual and a public health problem. In particular the consumption of strong liquor, especially the Dutch jenever (geneva), was high on their agenda. Quite a number of physicians, however, considered fermented drinks such as beer and wine as nutritious and stimulating health. The concerns about the consumption of jenever were not a new feature of the 1850s, but had been around since at least the 18th century.46 N.B. Donkersloot, psychiatrist and editor of GC, in 1854 put the abuse of strong liquor on a par with the state lottery as one of the two great disasters that had overcome the Netherlands. Not only was the use of strong liquor an individual health problem, since it badly affected physical health and was the cause of mental and moral aberrations. It was also a public problem, since it meant economic misfortune for consumers (who spend their money on drink) and destroyed religion and higher morality. Donkersloot pleaded for jenever prohibition, a position he would keep advocating in GC until his death in 1890.47 It is easy to see that the Morelian twist to the degeneration story fitted Donkersloot like a glove, and that it continued to be acknowledged in his articles in GC. It is also remarkable that Donkersloot’s views on jenever were exactly the same as those of one of the leading hygienists, L. Ali Cohen, almost two decades later. In his public health manual of 1872 Ali Cohen named the use of jenever as the chief source of the misery of the Dutch people. He expanded the usual description of jenever’s health hazards with the consequences for posterity as described by Morel. 48 Just as 42 43 44 45 46 47 48

On these views: Finzen (1977), 31. Ibid., 33. van Rees (1902). NTvG 31 (1895) I, 324. For analyses of the British ‘gin craze’, one of the first modern drug scares: Warner (2002); Dillon (2003). Donkersloot (1854). Ali Cohen (1872), 155–160.

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Donkersloot, Ali Cohen was unlike most of his colleagues also concerned about the physiological effects of fermented alcohol: already in 1863 he referred in NTvG to French research that had shown that beer and wine were ‘false’ nutritients, since they diminished the use of real nutrition. Its only value lay in its therapeutic use in medicine, as a stimulant. 49 The question of the nutritious and therapeutic value of alcohol was one of the few questions regarding the effects of the drug that physicians continued to discuss in the medical press. By the 1890s these discussions had become tied up with the discussions whether physicians should advocate moderation, as the majority obviously did, or whether they should preach abstinence. These discussions could take fierce forms, for instance at the 4th International Congress against alcohol abuse in The Hague in 1893, with Forel as foremost spokesman for the extremists.50 Two years later at the next international congress the issue was even more hotly debated. W.P. Ruijsch reported that the moderates (such as himself) were treated as traitors by their opponents.51 The psychiatrist Spaink was saddened by the exaggerations of the advocates of prohibition. Alcohol had its use as medication, he thought, although he prohibited its use in his own asylum in Apeldoorn.52 Hardly anyone questioned the public health hazards of chronic alcohol abuse. Ali Cohen was a contributing editor to NTvG, and the hygienists allied themselves politically to the liberals, while Donkersloot was the editor of the more conservative GC. But their position on the alcohol problem was the same, and Ali Cohen too advocated preventive and repressive measures in this ‘war on alcohol’. These strong opinions of prominent members of the medical profession contributed to the enactment of the first law on alcohol use (Drankwet) of 1881. This law regulated to some extent the trade in distilled liquor by preventive and repressive measures (such as limitation of the number of pubs and a prohibition of liquor sale to children younger than 16) and made public inebriety an offence. Until the First World War more than 4000 men and women would end up doing forced labour in state prisons for this offence. 53 To the temperance movement this law was totally inadequate, but this does not concern us here. What is relevant is that the whole spectrum of sides within the medical profession, that is the sides that expressed themselves in the medical press, were of the same opinion concerning the necessity of state regulation in the fight against alcohol abuse. However NTvG never got around to discussing the exact nature and desirability of the ‘preventive and repressive’ measures. Prohibition was certainly not one of them, partly because Dutch physicians were clearly too fond of their own alcohol intake. In 1900 the diatribe of Hazedoes van Almkerk against any use of alcohol in GC was coincidentally illustrated with an advertisement for a wine seller. 54 In 1901 on the general meeting of the Dutch Society for the Advancement of Medicine the representative of the city of Dordrecht argued that it was not the task of the Society to prohibit alcohol use. He evoked laughter when he added: “This representative has at least not noticed anything of this these 49 50 51 52 53 54

NTvG First series, 7 (1863), 664–665; Ibid. Second series 1 (1865) I, 523–533. On 19th century therapeutic use of alcohol: Paul (2001). NTvG 29 (1893) II, 321–323. NTvG 31 (1895) II, 551–552. NTvG 31 (1895) I, 92–93, 326. Van der Stel (1995), 156. A.N.J. Hanedoes van Almkerk, ‘Alcoholisme en de houding van medici te dien opzichte’, GC 54 (1900), no. 10.

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days [at the social gatherings surrounding the meetings].”55 The majority of Dutch physicians did not intend to confuse the fight against alcoholism with their personal enjoyment of life. GC took more stringent positions. To its editors, the hereditary degeneration caused by alcohol abuse was the ‘damnation of the human race’, and of more importance to the medical profession than bacteria research, sources of malaria, adulterated food, or miasmas. 56 Measures against public inebriety, as would become law in 1881, were not the right method, although Donkersloot did advocate fines and, on second offence, loss of citizen’s rights.57 Donkersloot consistently advocated that the only radical measure would be to make alcohol a prescription drug. Other, less radical measures, would be a state monopoly on the sale of alcohol and the close of the jenever pubs on Sunday and Monday morning.58 Donkersloot and his collaborators advocated now and then in GC forced abstinence for alcoholics.59 After his death his successors continued in the 1890s to warn against the dangers of alcohol and alcoholism. They advised physicians to give a good example and make propaganda against alcohol abuse. They advocated alcohol as a prescription drug, warned against the degeneration of progeny, and against the introduction of absinth. 60 The tale of Bismarck the tomcat was only another example of the tradition of the fight against alcoholism upheld by GC. But as with the degeneration theories this tradition had already been around in the 1850s. Did the link between degeneration and the advocacy of public health measures show any new transformations in the second half of the 19th century? We can indeed pinpoint changes in health policy strategies. In the 1880s and 1890s some voices are heard in the medical literature that seem to point to more support for eugenic policies based on genetic determinism and fatalism. These voices run parallel to the ‘hardening’ of hereditarianism in the biological sciences. But as we have seen, this parallel development should not seduce us into making any causal connections. Possible policy implications of the various biological theories are discussed only once, by Koster in 1886 in NTvG. According to him, the inheritance of acquired characteristics provided some hope that by a policy of (social) hygiene a better species could be created, whereas Weismann´s theories only led to prospect of a Spartan State, in which the inferior had to be eliminated. Koster´s line of argument here shows rather close similarities with a neo–Lamarckian perspective on positive eugenics. 61 But on the not the hereditarianism hardens, but the conclusions of some doctors concerning public health and prophylaxis. However, before the 1900s these voices remain a minority with no political impact. A first eugenic voice (although not in any way connected to a ‘eugenic movement’) can be heard in PB in 1884. It is based, not on hard hereditarianism, but on the ‘proven’ inheritance of 55 56 57 58 59 60 61

NTvG 37 (1901) II, 153. ‘Erfelijke dronkenschap’, GC 31 (1877), no. 49, 50, 51. GC 24 (1870) no. 23; ‘Beteugeling der dronkenschap’, ibid. 34 (1880), no. 40, 41, 42, 43, 44. ‘Het alkoholisme, zijn verspreiding, werking op het persoonlijk en maatschappelijk organisme, en de middelen om het te bestrijden’, GC 33 (1879), no. 5, 6, 7. N.B. Donkersloot, review of B.W. Richardson, Volksonderwijs over alcohol, GC 33 (1879), no. 26; Ibid. 42 (1888), no. 5, 49. C.W. Bollaan, review of Spaink, Over alcoholismus, GC 46 (1892), no. 38; Ibid. 49 (1895), no.2; Niermeijer, ‘Alcohol en alcoholisme’, ibid. no. 31; absinth: ibid. 48 (1894), no. 12. Koster, ‘Ontwikkelingsleer en ziektekunde’, NTvG 22 (1886) I, 341–349.

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acquired characteristics. Psychiatrist Tellegen draws a provisional conclusion from this theory. Physicians should advise before marriage on the heritability of certain diseases in the family–trees of the couple. Significantly, he adds, this is only done seldom.62 Actually the only thing Tellegen seems to do is to point out and advocate a practice that is current among some physicians, a practice that was regarded as consistent and normal for those familiar with Morelian degeneration and the inheritance of acquired characteristics: advice before marriage. 63 It is new that it is mentioned, but far from new as part of the private world of medical practice. When Spaink in 1892 suggests that confirmed drunks should not be allowed to marry this seems a continuation of what physicians (or anybody else with common sense) had always thought. 64 Treub has in 1900 basically the same position as Tellegen, since he asks for a medical advice before marriage that is not legally binding.65 There is some echo of this idea, when GC reports on the French idea of having a heredity report in the military passport.66 But that seems to be all, apart from one voice, which we would like to argue speaks the language of ‘modern eugenics’. G. Jelgersma, psychiatrist and editor of PNB, in 1897 reviewed the magazine of the temperance movement in an editorial. He is sympathetic and recommends the magazine. But he also has his doubts about the goals of the temperance movement. Is prohibition of strong liquor indeed the adequate method to significantly reduce the number of mentally insane (with as much as one– third, as the temperance activists claim) and of criminals and beggars (for the greater part)? The figures from those ‘dry’ American states where prohibition rules suggest otherwise. This should not be surprising, since alcohol abuse is from a scientific point of view as much consequence as cause of insanity. Where the ‘weak’ are not able to get alcohol, they ruin themselves by other means: in ‘dry’ Iowa the use of opium has increased. And now we hear the first and only voice in the medical press before 1900 that sounds distincty ‘social–darwinist’ in our popular sense. Prohibition would be counterproductive, since it would only keep the weak alive and allow them to reproduce themselves. It would interfere with natural selection. Jelgersma therefore feels that it is the état maladif, the pathogenic predisposition that is the cause of alcohol abuse and insanity. This pathological heredity should be controlled – by legislation of reproduction. For sure, Jelgersma does not deny the efficacy of anti–alcohol propaganda since there is also another form of alcohol abuse with a non–hereditary etiology: the great number of people who suffer from the increased demands and stress of modern society, and who seek escape and relaxation in drink. For those the activities of the temperance movement are very well suited. But as for the hereditary predisposed: they should not be allowed to reproduce. 67 As said before, Jelgersma’s voice here sounds as a first trumpet blast of eugenic thought. It is however interesting that his discourse can also easily be constructed as a ‘modern’ fin–de–siècle remake of more traditional views on the necessity of medical advice before marriage, just as Morelian degeneration could for its purposes in the medical domain as well be explained by the inheritance of acquired characteristics as by negative inheritance. Personal distaste of a physician 62 63 64 65 66 67

Tellegen ‘Eenige beschouwingen’, PB 11 (1884), 5–46: 18–25. On the survival of this practice in the United States: Pernick (1996). Spaink (1892), 26–27. Treub, ‘Huwelijk en ziekte’, GB 7: 29–48: 44–45. GC 56 (1902), no. 28. G. Jelgersma, review of De Wegwijzer. Maandblad voor geheel–onthouding, PNB 1 (1897), 287–294.

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for an alcoholic or an addict in general could as well be justified by his immoral behaviour and danger for the public community in the 1850s as for his Darwinian unfitness and danger for the public community in the 1890s. Of course, what is new is Jelgersma’s plea for legislation, but in the Dutch context this is a very solitary position before 1900 – even Treub does not want to go that far. What is of primary interest here is that ‘eugenic’ ideas of regulation of reproduction (by regulating marriage) are actually already around and even well–accepted by physicians in the second half of the 19th century. It remains however ‘invisible’ because it does not, or only now and then, extend to demands for public legislation. We must remember here that eugenic thought in various countries (but not the Netherlands) primarily made its essential impact on state legislation in the political, social and economic crises of the Interbellum.68

5. Approaches: individual health In practice, doctors would have to deal more with individual patients and their treatment options than with public health strategies. What can we say about the dynamics in conceptualizations around heredity and alcoholism and individual health approaches? First of all, we must discount one important historiographical notion about the treatment of alcoholism in the 19th century: the myth of therapeutic pessimism. If anything, the case of alcoholism and heredity shows that the idea of a ‘hardening’ hereditarianism as explanation of therapeutic failure can not be generalized. Estimations of recovery percentages under the right therapeutic regime ran as high as 40% (and as low as 25–30%) at the end of the century, a figure given by not extremely optimistic authors, and a figure that in the year 2004 is unsurpassed by modern addiction treatment methods.69 Of course we should not take these figures at their face value, but they clearly indicate something else than therapeutic pessimism. This does not mean to say that we cannot find any such pessimism in the medical press. In 1888, P. Wellenbergh reported in PB on his visit to the psychiatric hospital in Graz, Austria, run by the eminent degeneration specialist Richard von Krafft–Ebing. According to the latter, psychiatrists should make a distinction between madness that is not hereditary and can be cured in 70 to 80 % of the cases, and paranoia, that is the expression of a hereditary predisposition and can only rarely be cured. The implications for the alcoholic seemed evident. 70 Spaink was of the opinion that the symptoms of alcoholism were more severe in hereditary alcoholics. This was logical, because they were already insane or alcoholic before they had had their first drink; after their first drink, they had to become alcoholic.71 However this did not lead Spaink to therapeutic pessimism concerning his patients with a hereditary predisposition. A.N.J. Hanedoes van Almkerk, medical supervisor of the first Dutch asylum for inebriates in Hoog–Hullen, described his patients in the most pessimistic and abhorrent terms: ‘inferior’, ‘incongruency of the brain parts’, ‘Aztec skulls’ in a denigratory sense, predisposed children with the attitude of wild animals. 68 69 70 71

This argument, based on a comparative literature review on Britain, the United States, Germany, Sweden, Russia, and the Netherlands, is developed more fully in Snelders and Pieters (2003). NTvG 25 (1889) I, 563, 636; Spaink, Over alcoholismus, 72. PB 6 (1888), 96–103. Spaink (1892), 16–17, 26.

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But at the end of his article he claimed to cure 75% of his patients that stayed for more than one year of treatment, and 25% of those that stayed for a shorter duration. 72 His abhorrence for degenerates did not led him to pessimism in his work, although we must add that not all alcoholics in Hoog–Hullen were of course thought to be hereditary predisposed to their condition. It was reported from Hoog–Hullen that, because 60% of its patients were in their thirties, it was not probable that hereditary conditions were the main cause of their alcoholism. 73 Our thesis is that discovering hereditary antecedents in the family–tree of an alcoholic did not lead into pessimism because physicians generally thought that the expression of the predisposition was plastic, or fluid, dependent on environment and circumstances. These fluctuations in individual predisposition could be combined with the idea that treatment should be “a psychiatric treatment in accordance with the individuality of the patient”.74 It also combined with the possibilities of prophylaxis. The physician could, according to Ruijsch, cure where possible, and prevent by his influence in the families of his patients: by calmly and quietly explaining that alcohol is not necessary for everyone, by banishing it from children’s diets, by pointing to alcohol abuse in cases of illness, and to its negative consequences especially for those predisposed to alcoholism.75 In general, explained PB in 1888, a hereditary predisposition to insanity in individuals could be countered with the provision of a healthy wet–nurse for young children (in case of insanity or alcoholism in the mother), caution against the appearing of ‘brain congestions’, and a sensible education.76 Plasticity of genetic expression and elasticity of treatment theoretically opened the way for, not therapeutic pessimism, but the possibility of the reversal of degeneration. The readers of GC could read in 1870 about the research results of Morel’s pupil Doutrebente, one of whose conclusions was: “It is beyond doubt that races can regenerate themselves [emphasis added by GC], i.e., that through the influence of a harmless factor at least part of the descendants can climb to a higher position.”77 Thirty years later the same position was phrased by Van Rees in the terminology of negative inheritance: the damaged germ plasm could regenerate in the third or fourth generation if mixed with new, undamaged plasm.78 We can therefore subscribe to some extent to the conclusion of the study of Claus Finzen on alcoholism and degeneration in the German language scientific literature around 1900: many psychiatrists gave precisely their attention to the prophylaxis and treatment of alcoholism because alcoholism was seen as the main cause of mental diseases, and work in this field seemed to hold great promises of success. 79 Apart from the propaganda effect, what did treatment approaches consist of? Basically, alcohol abuse, delirium tremens, and dipsomania were treated with the same approach, consisting of abstinence, diet, and exercise.80 Once again, this was nothing new. Sir Walter Scott described 72 73 74 75 76 77 78 79 80

Hanedoes van Almkerk, ‘Alcoholisme en de houding van medici’, GC 54 (1900), no. 10. NTvG 37 (1901) I, 1333. Broers (1886), 30. NTvG 31 (1895) II, 714. Van Deventer, ‘Eenige opmerkingen over de psychiatrische behandeling van krankzinnigen’, PB 6 (1888), 27–28. ‘Erfelijke krankzinnigheid’, GC 24 (1870), no. 10. Van Rees (1902). Finzen (1977), 31. Broers (1886), 30; Spaink (1892), 71.

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in 1830 a medical treatment of a patient with alcoholism resulting in delirium tremens: “a gentle course of medicine (...) retire to [the patient’s] own house in the country, observe a temperate diet and early hours, practicing regular exercise, (...) avoiding fatigue.” 81 We can safely assume that this method would have been in use for centuries. The historian W.F. Bynum has concluded on 19th century medical approaches: “(…) alcoholism and alcohol–related problems could be treated by relatively simple measures like a wholesome diet and complete abstinence from alcoholic beverages. The alcoholic on occasion could be reformed and returned to society, hence the prognosis, even if often perceived to be bleak, was not so grave as that of many [other] asylum patients.”82 Furthermore, the second half of the 19th century witnessed extensive experiments and interventions with psychopharmacological medication. In these fifty years an astounding collection of drugs were tried out, especially to fight the effects of delirium tremens. In the 1850s, opium is still the chief medication in cases of delirium tremens and withdrawal symptoms. The idea is to artificially induce sleep, during which the “anomalies of the brain and nervous system” will automatically disappear.83 Opium therefore assisted the body in curing itself. Another drug in use was digitalis. Later in the century chloral hydrate, morphine, lupuline (in the 1870s), strychnine (in the 1880s), and other medication are in use, including purgatives and nausea cures (in which all food and drink are dosed with alcohol). Continuously more or less favourable reports are published in the medical journals.84 The great problem for physicians was not a lack of (more or less) effective treatment methods. The great problem was that the generally accepted best treatment method required total abstinence and a retreat from daily life to recover and cure. But most actual or potential patients did not have their own country house to effect their cure. In 1891 a special asylum for inebriates, Hoog–Hullen in Eelde, was opened, directed by a medical supervisor. 85 But neither this initiative nor changing thought about heredity led to significant changes in the overall medical approaches to alcoholism before 1900. Far from being helpless against alcoholism in its various forms, the physician of the second half of the 19th century had an impressive armoury of methods to help and cure his patients. Of course there are no reliable figures on the efficacy of these methods, but there was no cause for a priori therapeutic pessimism, nor did conceptualizations around heredity lead to that consequence.

6. Conclusions The Dutch medical journals of the second half of the 19th century show that doctors in the Netherlands integrated transforming knowledge around heredity in a flexible and fluid way in their conceptualizations and approaches of alcoholism. On a conceptual level, their goal seems not to have been theoretical consistency with scientific knowledge production, but its amendment and adjustment in the construction of workable explanatory tools. Morelian degeneration, neo– 81 82 83 84 85

Scott (2001), 19. On British medical approaches to alcoholism in the 18th and early 19th centuries: Porter, ‘The drinking man’s disease’. Bynum (1985), 63. NTvG 1 (1857), 355, 594. E.g., GC 24 (1870), no. 3; GC 32 (1878), no. 1; GC 33 (1879), no. 26; GC 34 (1880), no. 41; NTvG 20 (1884) I, 203. Van der Stel (1995), 189–194.

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Lamarckian inheritance of acquired characteristics, Darwinian evolution, or Weissmannian poisoning of the germ plasm could and were all used to produce these tools. They could be fitted to explain doctor’s experiences of phenomena of hereditary degeneration, of plastic expression of predispositions, and even of the possibilities for hereditary regeneration. In short, the transfer of knowledge from the scientific to the medical sphere was by no means hierarchically downward. On the level of approaches, knowledges of heredity did not connect to a therapeutic pessimism. The fight against alcoholism, based on a plastic concept of heredity, and incorporating elastic and pragmatic treatment and prevention practices, was offering hope for individual cures, as well as being an instrument in the long–term regeneration of the population. We must be hesitant to characterize public health strategies connected to heredity around 1900 as primarily concerned with collectives or the ‘race’, and not with individuals. Although not static, the medical domain in the Netherlands before 1900 does not come under the grip of a hardening hereditarianism. How Bismarck the tomcat would have fared under medical supervision in the subsequent decades will have to become clear in follow–up research.

Stephen Snelders, Frans J. Meijman and Toine Pieters, Vrije Universiteit, Medisch Centrum, Afdeling Metamedica, Amsterdam, [email protected]

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References Abbreviations in footnotes: GB: Geneeskundige Bladen uit Kliniek en Laboratorium voor de praktijk GC: Geneeskundige Courant voor het Koninkrijk der Nederlanden NTvG: Nederlandsch Tijdschrift voor Geneeskunde PB: Psychiatrische Bladen PNB: Psychiatrische en Neurologische Bladen TSH: Tijdschrift voor Sociale Hygiëne en Openbare Gezondheidsleraar Ackerknecht, Erwin H. 1953. Rudolf Virchow: Doctor, statesman, anthropologist. Madison: University of Wisconsin Press. Ali Cohen, L. 1872. Handboek der openbare gezondheidsregeling en der geneeskundige politie met het oog op de behoeften en de wetgeving van Nederland. Vol. I. Groningen: J.B. Wolters. Bowler, Peter J. 1988. The non–darwinian revolution: Reinterpretations of a historical myth. Baltimore: Johns Hopkins University Press. Broers, Jan. 1886. Alcoholisme, morphinisme, chloralisme, op zich zelf en in verband met elkaar beschouwd. Beverwijk: D.S. Slotboom. Bynum, W.F. 1968. ‘Chronic alcoholism in the first half of the 19th century.’ Bulletin of the History of Medicine 5: 160–185. Bynum, W.F. 1984. ‘Alcoholism and degeneration in 19th century European medicine and psychiatry.’ British Journal of Addiction 79: 59–70. Coffin, Jean–Christophe. 2003. ‘Heredity, milieu and sin: The works of Bénédict Augustin Morel (1809– 1873).’ In: A cultural history of heredity II. 18th and 19th centuries. Max–Planck Institut für Wissenschaftsgeschichte, Preprint 247: 153–164. Crowley, John W. and William L. White. 2004. Drunkard’s refuge: The lessons of the New York State Inebriate Asylum. Amherst: University of Massachusetts Press. Delprat, C.C. 1927. ‘De geschiedenis der Nederlandsche geneeskundige tijdschriften van 1680 tot 1857.’ NTvG 71 I: 3–116, 1711–1824, II: 13–86. Delprat, C.C. 1932. De wording en geschiedenis van de eerste 50 jaren van het Nederlandsch Tijdschrift voor Geneeskunde 1857–1907. Haarlem: De Erven F. Bohn. Dillon, Patrick. 2003. Gin: The much–lamented death of Madam Geneva. Boston: Justin, Charles & Co. Donkersloot, N.B. 1854. Loterij en jenever. Twee rampen over Nederland. Tiel: H.C.A. Campagne. Dowbiggin, Ian Robert. 1997. Keeping America sane: Psychiatry and eugenics in the United States, 1880–1940. Ithaca: Cornell University Press. Finzen, Claus. 1977. Der Alkoholismus als Problem der Degeneration um die Jahrhundertwende. M.D.–thesis, University of Kiel. Heron, Craig. 2003. Booze: A distilled history. Between the Lines: Toronto. Houwaart, E.S. 1991. De hygiënisten. Artsen, staat & volksgezondheid in Nederland 1840–1890. Groningen: Historische Uitgeverij. Huertas, Rafael. 1992. ‘Madness and degeneration, I: From “fallen angel” to mentally ill.’ History of Psychiatry 3: 391–411. Morel, Bénédict–Augustin. 1857. Traité des dégénérescences physiques, intellectuelles et morales de l’éspèce humaine. Paris: Ballière. Morel, Bénédict–Augustin. 1860. Traité des maladies mentales. Paris: Masson. Noordman, Jan. 1989. Om de kwaliteit van het nageslacht. Eugenetica in Nederland 1900–1950. Nijmegen: SUN. Paul, Harry W. 2001. Bacchic medicine: Wine and alcohol therapies from Napoleon to the French paradox. Amsterdam: Rodopi. Pernick, Martin S. 1996. The Black Stork: Eugenics and the death of “defective” babies in American medicine and motion pictures since 1915. New York: Oxford University Press. Pinell, Patrice. 2001. ‘Degeneration theory and heredity patterns between 1850 and 1900.’ in: Jean–Paul Gaudillère and Ilana Löwy, eds. Heredity and infection: The history of disease transmission. Routledge: London: 245–259.

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Pomata, Gianna. 2003. ‘Comments.’ In: A cultural history of heredity II. 18th and 19th centuries. Max–Planck Institut für Wissenschaftsgeschichte, Preprint 247: 145–151. Porter, Roy. 1985. ‘The drinking man’s disease: The ‘pre–history’ of alcoholism in Georgian Britain.’ British Journal of Addiction 80: 385–396. Ramaer, J.N. 1852. Dronkenschap en krankzinnigheid. Eene voorlezing. Tiel: Gebr. Campagne. Rees, J. van. 1902. ‘De invloed van de alkohol op het kind vóór de geboorte (erfelijkheid).’ In: Idem et al. Het kind en de alcohol. Amsterdam: Hoofdbestuur der Ned. Onderw. Propaganda–Club voor drankbestrijding: 1–8. Scott, Walter. 2001. Letters on demonology and witchcraft. Ware: Wordsworth Editions. Snelders, Stephen. 2003. ‘Changing concepts of heredity in the 20th century: A new perspective.’ Paper read at conference, Anglo–American medical relations: Historical insights, 19–21 june, The Wellcome Trust Centre for the History of Medicine at University College, London. Snelders, Stephen and Toine Pieters. 2003. ‘Van degeneratie tot individuele gezondheidsopties. Het maatschappelijk gebruik van erfelijkheidsconcepten in de twintigste eeuw.’ Gewina 26: 203–215. Snelders, Stephen and Toine Pieters. in press. ‘Alcoholism and degeneration in Dutch medicine around 1900.’ In: P. Dassen and M. Kemperink, eds. The many faces of evolution in Europe, c. 1860–1914. Groningen: Groningen University Press: 87–100. Sournia, Jean–Charles. 1990. A history of alcoholism. Oxford: Basil Blackwell: 98–114. Spaink, Pierre F. 1892. Over alcoholismus. Amsterdam: J.H.&G. van Heteren. Stel, Jacob Carel van der. 1995. Drinken, drank en dronkenschap. Vijf eeuwen drankbestrijding en alcoholhupverlening in Nederland. Hilversum: Verloren. Waller, John C. 2002. ‘“The illusion of an explanation”: The concept of hereditary disease’. Journal of the History of Medicine and Allied Sciences 57: 410–448. Waller, John C. 2003. ‘Poor old ancestors: The popularity of medical hereditarianism, 1770–1870’, in: A cultural history of heredity II. 18th and 19th centuries. Max–Planck Institut für Wissenschaftsgeschichte, Preprint 247: 131–144. Warner, Jessica. 2002. Craze: Gin and debauchery in an age of reason. New York: Random House. Weatherall, D.J. 1995. Science and the quiet art: Medical research and patient care. Oxford: Oxford University Press. Weiss, Sheila Faith. 1987. Race hygiene and national efficiency: The eugenics of Wilhelm Schallmayer. Berkeley: University of California Press. White, Paul. 2003. ‘Acquired characteristics: The (pre–genetic) material of the ‘self–made man’.’ In: A cultural history of heredity II. 18th and 19th centuries. Max–Planck Institut für Wissenschaftsgeschichte, Preprint 247: 67–82.

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A Changing Landscape in the Medical Geography of ‘Hereditary’ Disease: Syphilis, Leprosy, and Tuberculosis in Hawai‘i (1863-1903) Philip K. Wilson

It has been argued that during the second half of the 19th century a ‘hardening’ occurred in beliefs about the nature of heredity.1 ‘Inherited’ disease had long been viewed as the result of both nature and nurture. Appropriately nurturing against the hereditary predisposition to a disease could, it was argued, diminish the likelihood of the disease ever being expressed. This longstanding ‘soft hereditarian’ view of disease was later eclipsed by more ‘hard hereditarian’ claims that inherited disease was solely dependent upon a non-malleable nature. The current quest for better understanding the hereditary nature of particular diseases is hardly new.2 Considerable attention has been directed towards the meaning of heredity in relation to disease within the context of Europe, England, and the United States. However, the extent to which these beliefs changed elsewhere around the globe and whether this change was concurrent with that in Euro-America has been little explored. This paper begins to uncover some nineteenthcentury viewpoints about ‘hereditary’ disease in a different location – Hawai‘i. Remote though it is, Hawai‘i’s potential as a critical colonial outpost garnered the attention of Germany, England, France, and the United States during the nineteenth century. European explorers had opened new vistas to the eyes and to the mind, portraying their newly found ‘specimens’ as savages in primitive cultures. Ethnographers busily concocted labels of new races and varieties of humans within newly updated classification schemes. Missionaries focused upon all of the Polynesian Islanders as prime targets for Christianization. Governments envisioned these islands as colonial outposts which, when acquired, would strengthen their fortunes and futures. Euro-American Imperialism consequently provoked many native population groups to reinforce their respective cultural beliefs and traditions, including their beliefs and practices related to health and disease. Of the many Pacific islands ethnographically grouped at the Polynesian Islands within the triangulated zone of Hawai‘i, New Zealand, and Easter Island, a cultural heredity-mindedness was particular apparent in three of them: Tahiti, Tonga, and Hawai‘i. With particular interests in the disease patterns and healing traditions from the Native Hawaiian cultural perspective, I am interested in examining notions of heredity in relation to disease within this particular island group. My particular goal is to ascertain the extent to which views of the heritability of disease in Hawai‘i conformed with the general ‘hardening’ of hereditarian views expressed in the EuroAmerican medical writings during the second half of the nineteenth century. Adding the perspective of heredity from earlier Hawaiian perspectives will add a further dimension into our worldview of beliefs about the inheritance of disease.

1

2

Carlos López-Beltrán (1994) described that a malleable view existed in the “soft hereditarianism” beliefs of the early nineteenth century in contrast to the more objective qualifications of a nature-based, “hard hereditarianism” later in the century. For example, Wilson (2003a and 2002a).

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To begin reassessing just what natives and newcomers may have viewed as “hereditary” regarding both native and foreign disorders suffered by people in Hawai‘i during the nineteenth century, the existing literature must be questioned anew. Primary source material is crucial. The nineteenth-century published medical and public health writings, ethnographical accounts, scientific expedition reports, and travelogue memoirs help to capture contemporary anthropological perspectives of the Hawaiian people. These writings supplement the native perspectives from cultural luminaries including David Malo, Samuel Manaiakalani Kamakau, and John Papa Ii that were compiled at the time and are now available in English translation.

Introduction Hawai‘i faced a number of significant disease outbreaks during the 1800s. Venereal disease began devastating the native population soon after it arrived via James Cook’s sailors and continued, despite missionary attempts of Christian cleansing, to ravage Hawai‘i for many years. The Okuu (probably what Westerners’ viewed as Asiatic cholera) ravaged Hawai‘i beginning in 1804. 3 Measles was introduced to the islands in 1848. Smallpox arrived in Oahu in 1853 claiming at least 5000 lives, then returned in force in the early 1880s.4 Tuberculosis became endemic among the Hawaiian island populations during the nineteenth century, just as it did across the globe. And leprosy, possibly first identified in Hawai‘i as early as 1819 by members of a French scientific expedition, later became a disorder that troubled not only the afflicted, but also their families as it precipitated a legal segregation and isolation of the diseased to Kalawao (and later Kalaupapa) on Moloka‘i. Those who manifested signs of the leprous taint were gathered and hauled off to the settlement at Kalaupapa in the name of quarantine – for life. Finally, what has been characterized as the bubonic plague swept through the islands in 1899, taking a further toll upon the inhabitants at the close of the century. The “infectious” diseases mentioned above were frequently described in the 19 th century by Euro-American physicians, missionary settlers, expeditionary voyagers, and itinerant travelers – all haole (white foreigners), hapa (part) haole, or “Hawaiianized haole” – who provided critical insights into these diseases from various perspectives.5 However, most of the scholarship to date has yet to cover the entire spectrum of illness and disease as experienced in nineteenth century Hawai‘i. Indeed, I have noticed little mention of “inherited” or “hereditary” disease in recent writings about the Hawaiian kânaka (people) of the 1800s.6 This paucity of information is puzzling. For when one peruses nineteenth-century EuroAmerican medical writings and patient narratives, one finds that disease – or at least the propensity to disease – was commonly viewed as being hereditarily transmitted. This claim is true

3 4 5

6

Schmitt (1970). Greer (1969). We owe much of our historical contextualization of these emerging infectious diseases of nineteenthcentury Hawai‘i to the scrupulous research and widely read writings of O.A. Bushnell and C. S. Judd. Their devotion to advancing our understanding and treatment of disease in Hawai‘i, both past and present, was unprecedented. Their intellectual approach to explaining the birth and significance of new diseases has been replicated in the work of others, including Diamond (1996) and Igler (2004). See, for example, Green and Beckwith (1926), Larson (1962/63), and Blaisdell (1989).

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even for the so-called “infectious” diseases mentioned above. It is also puzzling in that among the Polynesians, the Hawaiian islanders were a hereditary conscious group. Pedigrees, in the form of epic genealogical chants have long been a part of Hawaiian culture. Kinship was, in the nineteenth century, an important concern for the kânaka maoli or ‘ôiwi (people of the bone) as well as for the ali‘i (ruling families). Hawaiians had established families of ruling chiefs as a class distinct from common islanders.7 But the Hawaiian kânaka (people) became increasingly worried throughout the nineteenth century about the repeated threats to the hereditary rule of the islands. Ascribing to a belief that the “nation lived while his [Kamehameha’s] descendants lived and ruled,” both natives and rulers remained focused upon the struggle to maintain this hereditary rule. Thus, thinking about the possible hereditary nature of diseases was, to some extent, a natural extension of the hereditary-mindedness among the kânaka maoli. A complete reassessment of hereditary disease within Hawaiian heritage is, however, beyond the scope of this paper. For now, I want to focus upon some ideas about heredity in Hawaiian history regarding three particular diseases common among Hawaiians and Euro-Americans during this period: syphilis, leprosy, and tuberculosis. Particular attention will be devoted to the timing and the extent to which hereditary conceptualizations of these diseases may have been modified by the Germ Theory, a new explanation of disease that gained considerable support in at least some parts of Europe and the United States during the second half of the nineteenth century. Prior to turning to each of these diseases, it may be helpful to identify, using a few examples, several traditional Hawaiian cultural beliefs concerning heredity, healing, and disease.

Traditional Hawaiian beliefs concerning heredity and disease Inheritance has been a traditional concern among native Hawaiians in regards to health and disease. In Hawai‘i, ancestry appears to have been as important for the pedigree of the healer as for that of the sufferer. For example, Kamakau recounted that only if the kahuna lâ‘au lapa‘au (medical healer) is “an upright person” – one who is pure and clean of person and deed, who “obeys the laws of that land as well as those of the akua (god or goddess),” – only then will he be “guided properly by true revelations of his spirit guides” in that the “secret things of his ancestors” will be revealed to him.8 Knowledge that certain illnesses were “inherited in a family through generations,” was a common belief to all kânaka maoli (native Hawaiians). The “secret things” that medical kâhuna received from their ancestors included wisdom about treating diseases that the an ‘aumakua (deified ancestor) or a kumupa‘a (ancestral deity) passed along hereditarily. These diseases were known to be “very resistant.” They did not “respond to [regular] medicines” and there were “not many kâhuna who are able to work on [them].” Healing “could not be done by outsiders or strangers because their voices and appeals would not be heeded by the ‘aumakua.” Since these diseases had been introduced through familial lines, it was thought to be more “the work of the family” than of the kâhuna in overcoming them.9

7 8 9

Goldman (1970), p. 234. Kamakau (1964), p. 95. Kamakau (1964), p. 97.

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Newborn babies were generally inspected by a “kahuna of predispositions” (kahuna pa‘ao‘ao). If a baby was found to be ill, it was imperative for the kahuna pa‘ao‘ao to ascertain whether or not it was “an ailment that afflicted the ancestors, coming down from them and afflicting the parents and then the child.” For if such ailments were not promptly and properly treated, the child was likely to suffer from “a more severe ailment” in adulthood.10 Untreated hereditary predispositions were believed to hold the capacity for changing into adult diseases including generalized states of emaciation, localized swellings, or specific disorders including tuberculosis and dysentery. Hereditary diseases represented a particular grouping of disorders that were commonly understood around the world as being capable of manifesting themselves throughout many generations. However, within Hawaiian cultural belief, a hereditary disease could be manifested for all future generations. Since a “relationship” had been forged “between ‘aumakua and descendants,” the Hawaiian people became “actual children (keiki ponoi) of the gods.” Therefore, a hereditary influence over illness and disease held the potential to last forever. 11 However, the meaning of “forever” in regards to inheritance in Hawai‘i was challenged during the later nineteenth century as islanders faced repeated threats to the hereditary rule of the islands. After the death of Hawai‘i’s ruler, Kamehameha IV (Liholiho), in 1863 until the defeat of the Home Rule Party in 1902, Hawai’i encountered turbulent confrontations by outsiders regarding the hereditary succession of its rulers. It was also during this period that many new claims were being tested in Hawai’i as to the inheritance of particular diseases. Therefore, focusing upon this time frame of threat upon the Hawaiian belief of hereditary rule, I will look briefly at the meaning of heredity in regard to what have been deemed as Hawai‘i’s “destructive trinity of diseases”: syphilis, leprosy, and tuberculosis.12

Syphilis It could be easily assumed from standard historical accounts that describe syphilis arriving de novo on the Sandwich Islands with Captain James Cook’s men in 1778, and its quick and wide-spread devastation, that contemporaries viewed syphilis as a contagious, infectious disease. Such a statement, however, is only part of the story. Taking a broader perspective on this disease and the Hawaiian people offers new insights into the history of Hawai‘i during the 19 th century in terms of island rule, the new-found wealth of sugar, immigration related to agricultural needs, concerns of ‘the other’ consequent to immigration, the changing demography of the island populations, and – most critical to our interests – changes in how people in Hawai‘i understood the etiology of this island scourge. William Ellis, the surgeon’s second mate aboard Cook’s ship, HMS Discovery, conjectured that it was only since the arrival of his ship that the island natives had become diseased “in a violent degree.” He identified European voyagers as the underlying “cause of this irreparable injury.” 13 Alonzo Chapin, one of a series of nine missionary physicians sent to serve the medical needs of the

10 11 12

Kamakau (1964), pp. 101-102. Kamakau (1964), p. 99. Mouritz (1916), p. 58.

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Hawaiians, claimed in 1838 – a half-century after Cook’s men established contact between the Hawaiians and “the other” – that Those, who have the credit of the discovery of the islands, and of exhibiting first to the astonished gaze of the simple and ignorant natives, some of the ingenious and useful implements and commodities of enlightened lands, and who sailed ships so enormous in size as to have been regarded as floating islands, inhabited by supernatural beings, must also receive credit of having introduced among these islanders . . . the vilest and most loathsome disease ever sent as a punishment for transgression.14

Chapin, in remarks of 1850, cited syphilis as “the most prominent cause of the decrease in population” of Hawaiian natives.15 Though census tabulations of this era leave room for speculation as to the precise population counts, there is general agreement among scholars that the number of native Hawaiian declined sharply between 1778 and the second half of the nineteenth century.16 While the Hawaiians “died off,” what one literary scholar described as a “strange cosmopolitan society of Chinese, Japanese, Portuguese, half-breed and Americans sprouted in their stead.”17 Many native Hawaiians hurled scorn against the largest immigrant population at mid-century, Chinese men in particular, as being “unchaste” and “whose cohabitation with Hawaiian women was doing nothing but contributing to their [the Hawaiian] continued infertility.”18 This so-called “sliding way of death” prompted considerable reaction. 19 On one level, the government expressed concern to “cease or at least limit” the importation of Chinese men. 20 Such efforts, however, were also seen by many authoritative figures as counterproductive to the need of new plantation labor as sugar increasingly became the unofficial, corporate King of the Islands. Another King, Kamehameha IV, worked on another level to develop public health measures against syphilis. “Our first duty is that of self-preservation,” the King claimed. “Our acts are in vain 13

14

15

16

17 18 19 20

Ellis (1782), pp. 73-74, as cited by Stannard (1990), p. 329. For other broad analyses of Cook’s impact upon the island from an infectious diseases perspective, see Stannard (1989), A. Bushnell (1993), and O.A. Bushnell (1993). Ralston (1984) also provides a helpful overview of key cultural changes within this time period. Chapin (1838), reprinted excerpts, Halford (1954), p. 291. These nine physicians were Thomas Holman, Abraham Bletchley, Gerrit Judd, Dwight Baldwin, Alonzo Chapin, Thomas Laton, Seth Andrews, James Smith and C.H. Wetmore. Chapin (1850), p. 93. Views expressed by many foreigners who traveled to or were at least temporarily relocated in the islands was consistent with that of many activists promoting Hawai‘i’s reclamation of sovereignty. This view is also apparent in blanket statements by historians who claim, the “white man’s presence was killing the Hawaiians.” Daws (1973), p. 74. State statistician Robert C. Schmitt (1968) provided a descriptive analysis of census taking. His numbers show a decline of the percentage of “native” Hawaiians among the people on the Hawaiian islands from 95.8 % in 1853 to 28.5 % in 1896. In identifying what contemporaries as well as later scholars viewed to be the likely causes for this decline, Schmitt (1968), p.159, that the “role of syphilis has been mentioned frequently.” Haunani-Kay Trask (1993), University of Hawai‘i Professor of Hawaiian Studies and an outspoken advocate for Hawaiian sovereignty, argues that an even more drastic decline occurred beginning from a considerably larger initial native Hawaiian population, pp. 6-7. A revised edition was published in 1999 by the University of Hawaii Press. Carré (1930), p. 237. Osorio (2002), p. 174. Judd (1977) attributed the use of this term to Ozzie Bushnell, p. 593. Osorio (2002), p. 174.

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unless we can stay the wasting hand that is destroying our people.” Consequently, the Queen’s Hospital was opened in 1859, devoted primarily to care “for sick and indigent Hawaiians,” in which a syphilitic ward was opened within its first year.21 But what of heredity and syphilis? Infection, regardless of its source, was not the only cause associated with the spread of syphilis.22 On one level, discussion of hereditary syphilis was propagated among islanders through the medical textbooks that Western-trained missionary physicians added to their collection while working on the islands. 23 Physicians during this era diagnosed syphilis based upon the visualization of signs. Hereditary syphilis often appeared in newborns as a particular constellation of signs. 24 It was also these visible manifestations that foreign visitors “inspected” and noted in their written characterizations of syphilis amongst the Hawaiians. But to the Hawaiians, the signs of syphilis held meanings beyond that routinely noted in the medical literature. As noted earlier, syphilis had been particularly tied to the arrival of Captain Cook. According to Hawaiian cultural belief, Cook personified the white god Lono. Lono had once been present among the Hawaiians, but had gone away. Cook’s appearance was commonly touted as Lono’s return.25 Upon Cook’s arrival, Hawaiian’s claimed, “Now our bones shall live” because “our ‘aumakua (ancestral spirit) has returned.”26 Within the genealogical tradition of Hawaiian gods and priests, Cook’s own death had been a foretold event. According to some contemporary accounts, Cook’s (Lono’s) revenge or wrath for his death would be delivered via later personifications of Lono who, in turn, were connected to Cook through their godly or priestly inheritance. Accordingly, some Hawaiians perceived that the syphilis spreading amongst them was the physical manifestation of Cook’s revenge being delivered unto them through the hereditary lines of the god Lono. Some Hawaiians explained this hereditary transference or transmission of disease in terms of mana. Mana, according to one leading structural anthropologist of Hawaiian culture, was “the creative power” that Hawaiians describe as “making visible what is invisible, causing things to be seen, which is the same as making them known or giving them form.”27 Views attributing disease to supernatural, spiritual causes were consistent with the aims of the kahuna lâ‘au lapa‘au (medical healer) and the Hawaiian missionary. The spirits underlying disease would, Hawaiian 21 22 23

24 25

26 27

Halford (1954), p. 238. Wilson (2003b), pp. 18-21. Such nineteenth-century textbooks, according to Halford (1954), pp. 131-132, included Benjamin Bell’s Treatise on the Venereal Disease, Alphée Cazenave’s On Cutaneous Diseases, René Theophile Hyacinthe Laennecs’ On Diseases of the Chest, M-F. Xavier Bichat’s Epitome of Physiology, Anatomy, and Pathology, Samuel Solomon’s Guide to Health, John Armstrong’s On Fevers, François Magendie’s Physiology, and John Mason Good’s Study of Medicine. It may be helpful to note that the distinctions twenty-first-century medical writers draw between the terms “hereditary” and “congenital” as applied to disease were not so clear in the nineteenth century. See Wilson (2003b), pp. 18-19. For an over view of the importance of “reading” marked children during this era, see Wilson (2002b). In How Natives Think: About Captain Cook, For Example, cultural anthropologist Marshall Sahlins (1995) elaborately analyzed this view of Cook as Lono, defending his interpretation against that postulated by Gananath Obeyesekere (1992). Ozzie Bushnell has also incorporated the Hawaiian view of Cook as Lono in his well received fictionalized historical account, The Return of Lono: A Novel of Captain Cook's Last Voyage, a work that that originally appeared as Peril in Paradise (1956). Kamakau (1961), as cited by Sahlins (1981), p. 7. Sahlins (1981), p. 31.

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healers claimed, remain active for generations and could return along specific lines of inheritance to inflict successive members in a family’s lineage. Christian missionaries also kept the hereditary thinking about syphilis alive as they spread their word to the peoples of Hawai‘i. Underlying disease, promiscuously-spread disease in particular, lay original sin. This disease, an William Bliss, one nineteenth-century traveler to the Islands noted, represented the “iniquity of the parents . . . visited upon the children, even to the third and fourth generations.”28 Christianizing the islands would, the missionaries hoped, cleanse the people by blocking this inheritance of original sin. Such reform would, it was argued, sway the trend away from the demise of the native population. With this frame of mind, missionaries – and missionary physicians – focused particularly on Christianizing the kua‘âina (rural folk in the hills and valleys) whom they perceived had been “beyond [the] reach of procurers and poxed foreigners,” who had “refused to tolerate intimacies with foreigners,” and who had “no taint of venereal diseases. For it was the kua‘âina, the missionaries claimed, that held great potential for the “preservation of the [native] race.”29 Alcoholism has, as I have argued elsewhere, been one of the “bad habits” frequently associated with the spread of syphilis.30 Alcoholism was also a disorder that, throughout the nineteenth century, was often claimed to have a hereditary etiology.31 This view, common in Euro-America, also gained support among the Hawaiians. Lunalilo, while prince and king, was known to be “constantly tempted to indulge his inborn craving for drink,” in that he was “lacking the constitution” to resist.32 The apparent high incidence of alcoholism in Hawai‘i was also exacerbated by environmental factors. For instance, many kânaka maoli in the nineteenth century who grew increasingly concerned about the depopulation of the native population and the threat of the complete disappearance of the “Hawaiian race” appear to have frequently turned to alcohol to palliate their concerns. By the early twentieth century, heredity was still supported in etiological thinking about syphilis in prominent Western writings. Even the Don of medical authorities, Sir William Osler, argued that cleansing “parents before marriage” remained the “most certain prophylaxis” against the hereditary spread of syphilis – that “most tragic form of the disease.” 33 Missionaries, physicians and native Hawaiians agreed that, uncontrolled syphilis would continue its wrath through its hereditary spread to future generations. Uncontrolled, syphilis was also thought to be able to transform or metastasize into another disease – leprosy. 34 28 29 30 31

32 33 34

Bliss (1873), p. 76. Halford (1954), p. 198. Wilson (2003b). Bynum (1984). In Wilson (2003a), I have demonstrated that Dr. Erasmus Darwin, grandfather of the naturalist Charles Darwin, was a leading advocate for temperance and documented many pedigrees of intemperance (i.e., alcoholism) in the early 1800s that ran through English families – including his own. Gordon-Cumming (1883), vol. 2, p. 224. Osler and McCrae (1915), vol. 2, p. 200. George L. Fitch, physician to the Kalaupapa leper settlement from 1882 to 1884 widely endorsed this view. Other physicians who spent time in Hawai’i – as did others across the globe – noted the great difficulty in differentiating signs of leprosy from those of syphilis. Samuel Kneeland (1873), p. 405, depicted leprosy as “inseparably mixed with syphilis.” Frank Enders (1877), p. 719, viewed leprosy “not as a disease sui generis but [as] an offspring of syphilis.” He reaffirmed this view, p. 719, citing the beliefs of other physicians that “the eradication of syphilis from these [Hawaiian] Islands, would eventually cause the disappearance of leprosy.”

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Leprosy Leprosy, a disease known for centuries throughout Europe, became a major health concern closely intertwined with Hawaiians during the mid 19th century. As Nathaniel Emerson, the physician son of a Hawaiian missionary, observed, the “roots” of leprosy are “very deep and are intricately interwoven with the whole fabric of the community.”35 Native Hawaiians came to believe that they held some special susceptibility to leprosy that was not seen in the haoles living on the islands. According to one government physician, George Fitch, the haole has “acquired a kind of hereditary immunity to leprosy as a result of centuries of exposure in Europe.”36 Even the licentious haole, who readily contracted syphilis were deemed to be “protected” from what Fitch viewed as the “fourth stage of its consequence” – leprosy. Missionary physician, Dwight Baldwin, noted its presence on the Hawaiian island of Maui as early as 1840. However, the Berlin-educated physician, William Hillebrand is typically credited with having noted its first appearance in Honolulu in 1848. The introduction of this “fresh item of the infinite curse which has come upon this [Hawaiian] race,” was blamed, by many, upon the Chinese.37 Indeed, it soon became colloquially knows as ma’i Pâkç (the Chinese disease). The afflictions of leprosy became well known to people on all the Hawaiian islands. 38 Its manifestations were remarkable. One well known piece of historical fiction recounts an 1893 episode on meeting lepers. The lepers were monsters – in face and form grotesque caricatures of everything human. They were hideously maimed and distorted, and had the seeming of creatures that had been racked in millenniums of hell. Their hands, when they possessed them, were like harpy-claws. Their faces were the misfits and slips, crushed and bruised by some mad god at play in the machinery of life. Here and there were features which the mad god had smeared half away, and one woman wept scalding tears from twin pits of horror, where her eyes had once been. Some were in pain and groaned from their chests. Others coughed, making sounds like the tearing of tissue. [Some] were idiots, more like huge apes marred in the making, until even the ape were an angel. They mowed and gibbered in the moonlight, under crowns of drooping, golden blossoms. One, whose bloated ear-lobe flapped like a fan upon his shoulder, caught up a gorgeous flower of orange and scarlet and with it decorated the monstrous ear that flip-flapped with his every movement.39

For a decade after its first description in Honolulu, leprosy was hardly mentioned in the growing number of widely read travel accounts of Hawai‘i. Mark Twain, for example, omitted any reference to leprosy in his well known Letters From Hawai‘i because, according to historian A. 35

36 37 38 39

Nathaniel Emerson to Rudolph Meyer, January 3, 1889, as cited by Daws (1973), p. 78. Insightful overviews of leprosy within the Hawaiian peoples are found in many works, including Stoddard (1895), Thompson (1897), Mouritz (1916), Weymouth (1938), esp. pp. 153-183, Farrow (1954), Wellman (1968), O.A. Bushnell (1963, 1968), Kalisch (1973), esp. pp. 500-501, Judd (1984), Gussow (1989), Veith (1992), pp. 300, 302, 304, Merwin (1998), Castillo (1992), and Inglis (Forthcoming). Daws (1973), p. 133. Bishop (1966), p. 223. Kalisch (1973), p. 501. London (1914 edition), pp. 50-51. Although the word “leper” connotes strong stigmatization and disrespect today, I have maintained its usage throughout this paper when I believe that it is important to contextualize nineteenth-century vernacular.

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Grove Day, Twain “did not wish to frighten off the business men who would be his most important readers.”40 However, among the people of Hawai‘i, both native and haole, leprosy hardly held a threat to visiting foreigners. Although the mode of its transmission was unclear, most initially viewed that it did not spread in a contagious manner. Writings of the period strongly suggest that physicians and Hawaiian deemed that leprosy was “somehow transmitted along hereditary lines,” 41 for the “leprous taint” was observed to run “strong in many families.”42 Such a view was consistent with the concurrent claims half-way across the globe in the leprosy asylums in Norway. There, Daniel Cornelius Danielssen and Carl Wilhelm Boeck had concluded from their studies that leprosy was passed along familial lines.43 This view was also consistent with the Hawaiian missionaries’ Biblical account of inheriting leprosy as a curse. The curse of Elisha seemed to be “irrevocably fixed” upon the Hawaiian just as the leprosy of Naaman was cursed to “cleave unto thee and unto thy seed forever.”44 Yet within this framework of thinking, another Biblical message resonated within the Hawaiian population. For similar to the treatment of lepers in the Biblical era, measures were soon undertaken in Hawai‘i to segregate the lepers from the “clean.” To briefly review this action, we turn again to Dr. Hillebrand. William Hillebrand remains a complex figure in the Hawaiian history of the 1860s. Regarding leprosy, he seems to have served several masters. It was Hillebrand who initially claimed that the leprosy in Hawai‘i owed its importation to the Chinese “coolie” labor force. Such medical concerns, however, did not prevent him, in 1865, from acting, as the King’s appointed Commissioner, to procure additional coolie plantation labor. That same year, Hillebrand prompted the Legislative Assembly to establish a leprosy Hospital and Detention Center in Kalihi that would help to better identify and contain the lepers in Honolulu. Once detected, the lepers were, according to the 1865 segregation “Act to Prevent the Spread of Leprosy,” sent off to the newly acquired governmental land at Kalawao, on the eastern side of the Hawaiian island of Moloka‘i. The history and lore of the lepers at Kalawao, and later after the transfer of this settlement to Kalaupapa on the northern side of Moloka‘i has been retold – frequently embellished – in much Hawaiian history and literature. Most common is the tale of Father Damien, a Belgian Catholic priest of the Sacred Hearts, who arrived at Kalaupapa in 1873 to offer spiritual and physical care for several hundred lepers secluded there, and who, eleven years later, was diagnosed with and subsequently died from leprosy.45 However, Kalaupapa also represented a cornerstone for the study of a reputed hereditary disease. Leprosy was, as noted above, known to afflict members of different generations within one family, but curiously this pattern of transmission appeared only in some families – even when looking at just the native Hawaiian community. Such transmission could, of course, have been explained by contagious spread among members of a household that often included several 40 41 42 43 44 45

Day (1977), p. 65. Daws (1973), p. 6. Bishop (1966), p. 223. Rokstad (1964), p. 65. Bliss (1873), p. 97. Emmett Cahill (1990) offers a photo journalistic overview of the history of this settlement.

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generations of family members living closely together. However, more often than not, leprosy appeared only in a few individuals of one family’s generation, sparing most of the family members, all of whom had been in relatively similar contact with each other. Moreover, many kôkua (caregivers) voluntarily worked amongst the lepers in the isolated community at Kalaupapa, yet they never contracted leprosy. Was some form of protection from this disease hereditarily transmitted? Or, was it a weakened physical constitution among the afflicted that had been passed along family lines? It initially startled hereditarian thinking when news reached Hawai‘i from Norway regarding Gerhard Henrik Armauer Hansen’s 1873 isolation of rod-shaped bodies routinely observed microscopically in the mass of bacteria present in nodules of leprous patients. 46 However, neither Hansen nor his contemporaries could demonstrate a clear causal relationship by inoculating “clean” animals or humans with material taken from a leprous nodule. Thus, Hansen’s findings prompted little change of thinking in Hawai‘i – at least initially. Leprosy was becoming a “national blight” for Hawai‘i. Forty-eight Protestant ministers, both Hawaiian and haole, who gathered at the 1873 Hawaiian Evangelical Association meeting conjectured that “our Hawaiian people will become in a very few years, a nation of lepers.” 47 Indeed, even members of a later generation within a leprous family were at times among those sequestered and sent off to Kalaupapa. The Kalaupapa containment had not reduced leprosy’s spread on the other islands. Based upon this information, the Hawai‘i Legislative Assembly concluded in 1873 that leprosy was not a contagious concern, and they attempted to reverse their earlier decision and vowed to send no more lepers to Moloka‘i.48 However, commercial interests prevailed as other authoritative figures within the Hawaiian community wished to keep lepers out of sight from the malihini (foreign observers) visiting Honolulu. These malihini were, after all, the observers whose future financial investments would continue to bolster the economic growth of the islands. In 1882, according to Harvard Professor of Dermatology, James C. White, the medical world had “almost universally” adopted the opinion that “leprosy is not contagious, and that it is endemic mostly because it is hereditary.”49 Although most medical authorities agreed with White’s claim, emphasizing that it continued to “appear in the descendents of . . . [particular] families,” others argued that this finding alone “proves nothing a priori, for the . . . continuance [of a disease] among relations may [also] be used . . . as the best evidence of its communicability by contagion.”50 Alas, this “unquestionable doctrine” continued to be questioned. The “important point to be determined,” they argued, is “the proof of [contagion], not the dis[proof] of [heredity].” Maintaining this secluded population of lepers allowed physicians during the last two decades of the nineteenth century to use Kalaupapa as an experimental study site from which they hoped to 46

47 48 49 50

Hansen’s initial article (1874) is available for English readers as “”Causes of Leprosy” (1955). See also Feldman (1965), pp. 412-416. In the twentieth century, leprosy was officially renamed “Hansen’s Disease.” Daws (1973), p. 63. Osoro (2002), p. 177. White (1882), p. 435. White (1882), p. 435.

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ascertain the precise hereditary underpinning of leprosy. Hawai‘i provided the case study site whereby the “recent introduction of leprosy into an insular nation” and the sequestration of most of those with the diseases into a further isolated community provided “that virgin field for observation so essential for the proper study” of the transmission of leprosy. 51 Much was at stake over the Kalaupapa findings regarding leprosy. On one level, marriageability between lepers as well as between individuals from ‘clean’ and ‘unclean’ families depended upon these observers’ conclusions. On another level, the entire usefulness of the settlement was in question. One figure at the center of this question was George Fitch. It was Fitch who, in 1882, had become Hawai‘i’s greatest proponent of drawing the interconnections between leprosy and syphilis. Leprosy, he argued, was the fourth stage of syphilis. His view, according to one observer, became “quite a popular topic of discussion amongst the laity and the medical fraternity in Honolulu.”52 It also exacerbated the already present diagnostical dilemma that existed between these two disorders. Fitch drew upon the findings at Kalaupapa as well as authoritative reports from around the globe for his 1884 Report to the Board of Health. He construed the information in this report as thoroughly supportive of a hereditary transmission of leprosy. “There appears no more need . . . for restricting the liberty of lepers” by sequestering them in perpetual quarantine at Kalaupapa than for “restricting the liberty of those with the gout.” Segregation, he continued, “except in so far as it prevents [the] hereditary transmission of the disease has absolutely no effect towards checking it.”53 Going against the tide, though unsupported in his belief, was Harvard dermatologist James White. His view was based upon the observation that just as in the case of the “syphilization” of Hawai‘i, leprosy appeared far too quickly for it to be exclusively caused by heredity. For if heredity was acting as the only factor, he argued, “it would have required several generations to have accomplished such results.”54 Additional support that this was certainly not the case in Hawai‘i, some surmised, came from the finding that no word existed in the Hawaiian language for this disease. Nor could, as Morell Mackenzie argued, “ancestral proclivity” alone explain the “sudden outbreak of the diseases [across the globe] in races [that were previously] altogether free of it.” 55 Also going against the tide were the Hawai‘i-based physicians, Eduard Christian Arning and Arthur Mouritz. Arning, a highly skilled bacteriologist, arrived in Honolulu in late 1883 from Germany. Mouritz, an Oxford-educated physician, came to Hawai‘i and served as a resident physician and Medical Superintendent to the Leper Settlement at Kalaupapa from 1884 to 1887. Arning acknowledged that a “disposition” to leprosy, a “certain weakness to resist its attacks,” “may possibly be transmitted by heredity,” but he remained firm that “leprosy itself was not congenital.”56 In 1884, he designed an experiment that would, he argued, prove once and for all, that leprosy could also spread via contagion. This involved the inoculation of a condemned 51 52 53 54 55 56

White (1882), p. 435. Mouritz (1916), p. 54. Cited in Mouritz (1916), pp. 384, 396. White (1882), p. 438. Mackenzie (1889), p. 938. For an elaborate contemporary description of specific global sites affected by the recurrence of leprosy, see Tebb (1893). Arning 14 November 1885 report to Gibson, cited in Mouritz (1916), p. 326.

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prisoner, Keanu, whose sentence the Hawai’i Privy Council commuted from hanging to life imprisonment so that he might serve as a subject for Arning’s experiment. Keanu, a strong man who showed no signs of leprosy or other illnesses, had the leproma (a leprous filled swelling) that had been removed from a young leper colony resident and sutured over an incised site on his forearm. Arning removed material from Keanu’s body near the inoculation site for a number of months. To his great disappointment, no bacterial signs of leprosy were found nor did Keanu develop any visible signs of leprosy before Arning returned to Europe in 1886. The resident Kalaupapa physician, Mourtiz, watched over Keanu after Arning’s departure and, a year later, diagnosed him as a “confirmed leper.” Arning presented this case at the First Congress of the Society of German Dermatologists held in Berlin in 1889. Shortly thereafter, however, doubts over Arning’s findings surfaced. Leprosy was, it was noted, endemic in Hawai‘i, and, as Mouritz argued, some of Keanu’s relatives were also known to be inflicted. 57 Moreover, Mouritz had not succeeded in any of over a hundred similar attempts to inoculate healthy Hawaiians with the disease.58 Arning’s later writings show more than disappointment. Although he had once viewed Hawai’i as a natural laboratory for the study of leprosy, he now saw it much differently. For in Hawai‘i, “the intense feeling which everything connected with leprosy necessarily evokes in so small and terribly afflicted community, cannot favour the slow and tedious process of purely scientific work.” And, no small obstacle in and of itself, there one also had to contend with the “character of the natives.”59 Concern mounted on all sides of leprosy’s etiology for the remainder of the century. Foreign dignitaries and visitors, such as M.G. Bosseront d’Anglade, Commissioner and Consul of France, noted in 1893 that there are “very few native families who are not affected” with leprosy – both the disease and its stigmatization – as well as by the separation of their family members to Moloka‘i. In his travelogue, he admitted his inability to absolutely ascertain whether leprosy was contagious. The “combination of two conditions appears necessary for leprosy to develop,” he concluded. First, “prolonged contact with other affected persons,” and second, “individual susceptibility to disease.” The susceptibility, he argues, is seemingly inherited. He cited an example of the inherited passage of this susceptibility in the case of a “healthy Kanaka man who during ten years at the [leprosy] settlement had successively married four women patients, [and] begotten leprous children by each of his wives, and yet never gave any indication that he himself had become a leper.”60 Thus, for many, as for Bosseront d’Anglade, the ambivalence over what precisely guided leprosy’s spread kept the matter controversial for some time.

57 58 59

60

Keanu’s familiar linkage to this disease was also raised by another resident physician at Moloka‘i, Sidney Bourne Swift. Tebb (1893), pp. 127-128. Daws (1973), pp. 142, 234-235. For an overview of the attempts to inoculate humans with leprosy, see Dubois (1952). Arning to Nathaniel Emerson, 10 May 1886, as cited by Daws (1973), p. 278. Keanu was retained at Moloka‘i, a “punishment,” according to California University pathologist, D.W. Montgomery, that was “ten times more severe than the death penalty. Physician William Jelly concurred, noting “had [Keanu] known what leprosy is, [he] would without hesitation, have preferred the guillotine, the garrote, or the hangman’s noose” to his fate in Kalaupapa. Tebb (1893), pp. 124-135. Bosseront d’Anglade (1987).

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It should be noted that by the century’s end, we do find heredity being eclipsed by contagion as the chief explanation for Hawai‘i’s leprous population. However, this turnabout resulted more from politics than from medicine. New immigration channels would be opened if Hawai‘i was formally annexed to the United States. Fear of exporting leprosy together with coolie labor loomed, reaching unprecedented levels in California in 1897.61 When, as New York public health authority Prince A. Morrow argued, it is considered that “more than ten percent of the Hawaiian race are affected with leprosy,” he conjectured that “it becomes a serious question as to what will be the effect of the absorption of this tainted population upon the health interests” of the United States if the islands are annexed to them.62 The Hawaiians themselves are not the real problem, he argued, for each of them is “essentially insular in his tastes and habits and shows little disposition to leave his native shores.” Rather, it is the labor forces who, having interbred with the native Hawaiians, are the likely ones to carry the “seeds of . . . [this] deadly contagion [i.e., leprosy]” with them when imported to the Western shores of the United States. 63 “All of these facts,” he concluded, “should be carefully considered and their importance from a sanitary point of view carefully weighed by our legislative authorities before deciding upon the [U.S.] annexation of Hawaii with its leprous population.”64 Hawaiian sovereignty came to a close soon after the U.S. Congress’s passage of an act to Annex the Hawaiian Islands to the United States on July 7, 1898. Political turmoil had begun with the overthrow of the Hawaiian Constitutional Monarchy in 1893 and the self-designated provisional government to oversee the new U.S. protectorate. It remained strong through the time that proexpansionist U.S. President, William McKinley ushered in the annexation bill. “Among the early results of annexation,” the physician Burnside Foster argued, will “undoubtedly be a largely increased immigration to as well as emigration from the islands.” Many who “either know or suspect that they have [leprosy] will undoubtedly escape to this country while those from this country who settle in Hawaii will be thrown into more or less intimate relations with the already infected but unrecognized lepers.” Thus, it would “certainly seem worthwhile,” Foster continued, “for the United States to take this question immediately at hand, and to appoint a commission of bacteriologists properly equipped and with every facility for the study of the leprous problem. England has gained the eternal gratitude of humanity for her Jenner and her Lister, France for her Pasteur, Germany for her Robert Koch. Shall not America . . . gain further glory by striking leprosy from the calendar of human afflictions?”65

Tuberculosis The rise of bacteriological thinking in the nineteenth century, promulgated primarily via the work of Robert Koch, diverted the medical gaze, focusing it more upon germs than germ cells. Tuberculosis, one of the most common diseases of the nineteenth century, is among the most critical emerging infectious diseases across the globe in our era. To combat medicine’s diminished 61 62 63 64 65

Gussow and Tracey (1970), p. 439. Morrow (1897), p. 582. Morrow (1897), p. 588. Morrow (1897), p. 590. Foster (1898), p. 305.

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stronghold over tuberculosis, current investigations focus upon uncovering changes in the genetic evolutionary processes between the tubercular microbe and anti-microbial agents as well as identifying inheritable alterations that have influenced the germ’s pathogenicity. Curiously enough, in the nineteenth century, the heredity of tuberculosis was also being investigated. But unlike the present focus upon the etiological agent itself, earlier genetics studies were inferred from tuberculosis’ transmission along family lines. Divergent thoughts existed surrounding the hereditary nature of tuberculosis during the 1800s. Medical authorities who purported that this “white plague” was hereditary offered as evidence its frequent appearance in multiple members and successive generations of the same family. Some argued that “from the beginning of medical time it was considered that tuberculosis ran in families; that heredity had much to do with the occurrence of the disease.”66 Within the early nineteenth century, we find a number of prominent European, British, and American physicians endorsing its hereditary basis. Antoine Portal “maintained categorically” that tuberculosis was “hereditary and had nothing to do with contagion.” Gaspard Laurent Bayle argued that heredity was the “chief aetiological factor” underlying this disease. René-Théophile-Hyacinthe Laënnec, whose innovation of the stethoscope was targeted towards the study of respiratory diseases acknowledged that although tuberculosis “appeared to be contagious is some countries, this did not seem to be the case in France” – his homeland. Gabriel Andral, William Cullen, Sir John Forbes and Sir James Clark also supported this theory.67 Prominent Boston medical authority, Henry I. Bowditch wrote to a general reading audience in 1869 that “Undoubtedly it is true that public opinion considers consumption [i.e., tuberculosis] as hereditary, and medical experience seems to support this view. We presume that there is scarcely a physician anywhere who would not admit the truth of this belief.” Yet, he adds, “no physician would dare to say that . . . consumption would necessarily be transmitted from parent to child.”68 However, Robert Koch’s 1882 discovery of a bacillary germ as what he deemed the etiological agent responsible for tuberculosis provoked serious rethinking about this disease among the medical community and the general populace. According to biomathematician and eugenicist Karl Pearson’s later reflections, following Koch’s discovery, the “idea of infection dominated” and consequently, an “immediate neglect” arose regarding any reputed “hereditary factor” underlying this disease.69 Although it is true that Koch’s discovery “immediately” spread throughout the globe, we are only beginning to fully appreciate that his pronouncement did not immediately convert all ‘hereditarians’ into supporters of the germ theory.70 Indeed, many physicians, especially older physicians, remained skeptical of any germ-based etiology, and they continued to practice believing tuberculosis to be hereditarily transmitted.71 When confronted with findings suggesting 66 67

68 69 70 71

Lewis (1923), vol. 1, p. 178. Keers (1978), p. 53, reviewed these proponents’ beliefs. Additional supporters of a hereditary passage of tuberculosis are found in Castiglioni (1933). For a helpful outline of the changing perceptions of this disease, see Burke (1955). Bowditch (1869) as reprinted in Rosenkrantz (1994), p. 63. Pearson (1912), p. 3. Worboys (2001) and Carter (2003), p. 196. Ott (1996), p. 128. For more on the persistence of hereditarian thoughts, see Wilson (Forthcoming).

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that they disregard heredity as the primary cause of tuberculosis, they typically replied with something like the following argument. How can we account for the cases where the parents, having died of Consumption, the children are necessarily attacked on arriving at a certain age, with a severe type of the disease? And, moreover, there are several instances . . . where the children, who happened to be scattered in various parts of the world, were yet attacked and succumbed to the fell disease at almost the same age?72

However, the hereditarians were divided over precisely what they deemed to be heritable regarding tuberculosis. Whereas a large faction had once argued that “the disease” itself was inherited, many physicians began to assert during the late 1800s that it was something about this disease that was inherited. In other words, a hereditary tendency, predisposition, or diathesis to disease existed.73 Diathesis, the term commonly used at this time, referred to the “inherent liability to consumption which ‘ran in families’ and was handed down from one generation to another.”74 Indeed, the extent to which the germ theory predominated medical thought regarding tuberculosis during this period remains unresolved. In Hawai‘i, we find it present in successive generations of missionaries, natives, and chiefs. 75 There, tuberculosis “vied with syphilis” as the “singularly most remarked upon disease . . . during the first half of the nineteenth century.”76 However, as the century progressed, tuberculosis, part of what Mouritz characterized as Hawai‘i’s “destructive trinity of diseases” (together with syphilis and leprosy), had come to overshadow the other two such that it remained in “almost full possession of the field.”77 When we look to Hawai‘i during the two decades following Koch’s discovery, we find the most vocal support for a hereditary transmission as being the primary explanation of tuberculosis among the island population. Most typically, the discussions closely followed descriptions of leprosy. The connection between these two diseases was not merely coincidental, for considerable clinical conundrums arose because leprosy, at least in its early forms, mimicked the signs of tuberculosis. Some physicians considered the “tuberculosa” or “tuberculous condition” that formed on the face, legs, arms, and trunk of lepers as affiliated with an inward pulmonary tuberculosis.78 However, over time, the manifestations of leprosy became more distinguishable, and it did not involve the lungs.79 Tuberculosis also held a particular relationship with syphilis on the islands. It was widely held that an individual demonstrating “defects of development” or having parents whose diseases 72 73 74 75 76 77 78 79

Williams (1882), p. 618. Ackerknecht (1982). “Tuberculosis” (1911), vol. 27, p. 357. Middleton (1971), p. 452. Stannard (1990), p. 342. Mouritz (1916), p. 58. This finding is remarkable considering that in the mid-1800s, several physicians noted the Hawaiian’s “entire exemption from pulmonary tuberculosis.” Gulick (1855), p. 197. Kneeland (1873), pp. 404-405, and Tryon (1883), p. 447. Although sanitaria were developed on four of Hawaiian Islands, tubercular patients admitted themselves voluntarily. They never suffered the stigmatization commonly felt among the leprous patients.

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“stamp[ed] sufficient weakness upon the[ir] offspring” frequently gave rise to children with a diminished resistance to tuberculosis. Prominent among such parental-derived diseases was syphilis.80 And again, as seen in beliefs about syphilis and leprosy, those determined to Christianize the Hawaiian islands integrated Biblical wisdom with medical reports. Dr. Bowditch provided Hawai‘i’s missionaries considerable support through popularizing tuberculosis as follows. It is often seen that the “diseased conditions of the parents, sometimes, alas! Due to their own or to their ancestors’ previous excesses, [create] tender bodies of [newborn] children . . . so tainted that life becomes a burden. We have often seen in such [tuberculosis] cases the terrible vindication of the power of the old Mosaic law, ‘For the sins of the fathers are visited upon the children . . . unto the third and fourth generations.’”81

One additional connection may be gleaned regarding tuberculosis and inheritance in the Hawaiian islands. King Lunalilo’s tuberculosis brought an end to his reign in 1874, only thirteen months after he had been elected King by the Hawaiians upon the death of Kamehameha IV. Between the time of his death and the U.S.’s takeover of control of the Hawaiian islands, the hereditary lineage of rulers successively lost ground. Interestingly, it was over this same time that the germ theory fought to overtake hereditary beliefs in the transmission of diseases. Perhaps the end of hereditary rule in Hawai‘i parallels, in ways that have yet to be uncovered, the transference of power from the hereditarians to the germ theorists regarding disease.

Hawaiian highlights expanding our perspective of hereditary disease Perhaps Hawai‘i is marginalized and remote to most cultural worldviews, both then and now. Still, this reflection upon a seemingly remote Hawaiian culture during the last half of the nineteenth century reminds us that in the views of some folk and physicians, “heredity” and “inheritance” as related to disease were, at times, quite distinct from concurrent views expressed in Euro-Ameican writings. For example, long before Westerners sought to identify the hereditary substance within humans, Hawaiians already envisioned a hereditary substance, mana, (spiritual power), as a contributory factor to health and disease. If, for example, the flow of mana was disrupted, disease would likely follow. Spirits, ghosts, wraiths and astral bodies are ancestral, and thus, as Hawaiian heritage reminds us, their influence must be considered in discerning a culture’s view of heredity as an etiological factor of disease. Similar beliefs are found and may be contextualized within other Polynesian Island cultures. Anne E. Becker analyzed Fijian culture, noting that the spirit, often ancestral, is often attributed as the cause of an illness.82 Although Fiji lies just outside of the typical island range denoted as the Polynesian group, it has long been populated with a large percentage of Polynesians whose belief systems have come to influence Fijian culture. If visual detection was important for a culture to identify the presence of a hereditary disorder, then the sufferer must be visible. If, however, we follow the suggestions presented in some early 80 81 82

Osler and McCrae (1915), vol. 1, p. 317. Bowditch ([1869] 1994), p. 65. Becker (1995).

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nineteenth-century writings, we are left uncertain as to whether these diseased individuals were actually kept alive for long. “Deformed, sluggish, or ill-tempered babies were unceremoniously put out of the way,” it has been argued. Children with “deformities and trying illnesses of one sort or another,” many of which could have been brought on through inheritance, were frequently disposed of quickly or “placed out of sight and sound . . . and allowed to languish and waste away.”83 Despite what some modern readers may think of such practices, if indeed they were as common as nineteenth-century authors lead us to believe, we should also appreciate that such practices were condoned within the moral and religious framework of native Hawai‘i at that time and discuss them within that historical context.84 Expanding research into these and related areas will allow us to recapture more about the entire spectrum of illness and health as viewed by Hawai‘i’s ka po‘e kahiko (the people of old). Such views can complement and strengthen the Western-centered narratives that have dominated decades of the telling of the Hawaiian past were applied to states of disease within different cultural perspectives across the globe. Such efforts will allow us to better contextualize the significant prior contributions regarding “infectious” and “contagious” diseases within Polynesia with an even broader worldview of disease and history. This focus upon Hawai‘i exemplifies our need to be aware that specific cultural contexts drive specific and precise meanings into such polysemic words as “inheritance” and “heredity.” It also helps us appreciate the need to consider comparative cultural views as we conceptualize the meaning that heredity held during earlier periods. It remains to be determined just how contemporaries of the nineteenth century viewed immigration and racial mixing as altering, perhaps exacerbating, hereditary influences upon disease within these new mixes of peoples in Hawai‘i. This area of inquiry will be covered in my next foray into heredity, disease, and Hawai‘i as I look at Hawai‘i through the eyes and minds of early 20th-century Anglo-American eugenicists.

Philip K. Wilson, Penn State College of Medicine, Hershey, PA [email protected]

83 84

Lee (1966), pp. 172, 178. Tobin (1994). Cultural anthropologist George M. Foster (1976) provided timeless reminders to historians of the need to be ever cognizant of the comparative ethnographical contexts when pursuing historical work regarding “other” cultures.

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References Ackerknecht, Erwin H. 1982. “Diathesis: The Word and the Concept.” Bulletin of the History of Medicine 56: 317-325. Becker, Anne E. 1995. “The Body as a Community Forum: Spirit Possession and Social Repossession.” In Body, Self, and Society: The View From Fiji. Philadelphia: University of Pennsylvania Press, 104-126. Bishop, Isabella Bird. 1966. Six Months in the Sandwich Islands. Honolulu, HI: University of Hawaii Press for Friends of the Library of Hawaii. Blaisdell, Kekuni. 1989. “Historical and Cultural Aspects of Native Hawaiian Health.” Social Process in Hawaii 32: 1-21. Bliss, William R. 1873. Paradise in the Pacific: A Book of Travel, Adventure, and Facts in the Sandwich Islands. New York: Sheldon and Company. Bosseront d’Anglade, M.G. 1987. A Tree in Bud: The Hawaiian Kingdom 1889-1893. Honolulu, HI: University of Hawaii Press. Bowditch, Henry I. [1869] 1994. “Consumption in America.” In From Consumption to Tuberculosis: A Documentary History. Edited by Barbara Gutmann Rosenkrantz. New York: Garland Publishing, Inc. Burke, Richard M. 1955. An Historical Chronology of Tuberculosis. 2nd ed. Springfield, IL: Charles C. Thomas. Bushnell, Andrew. 1993. “‘The Horror’ Reconsidered: An Evaluation of the Historical Evidence for Population Decline in Hawai‘i, 1778-1803.” Pacific Studies 16: 115-161 Bushnell, O.A. 1956. Peril in Paradise. New York: Ace. ———. 1963. Molokai. Cleveland: Cleveland World Pub. Co. ———. 1968. “The United States Leprosy Investigation Station at Kalawao.” History of Science in Hawaii 2: 76-94. ———. 1993. The Gifts of Civilization: Germs and Genocide in Hawai‘i. Honolulu, HI: University of Hawaii Press. Bynum, W.F. 1984. “Alcoholism and Degeneration in 19th-Century European Medicine and Psychiatry.” British Journal of Addiction 79: 59–70. Cahill, Emmett. 1990. Yesterday at Kalaupapa. Honolulu, HI: Mutual Publishing. Carré, Jean Marie. 1930. Robert Louis Stevenson: The Frail Warrior. Translated by Eleanor Hard. New York: Coward-McCann. Carter, K. Codell. 2003. The Rise of Causal Concepts of Disease: Case Histories. Aldershot: Ashgate. Castiglioni, Arturo. 1933. History of Tuberculosis. New York: Froben Press. Castillo, Stephanie J. 1992. Simple Courage: The Story of Hawaii’s Leprosy Epidemic: A Historical Portrait for the Age of AIDS. Co-produced by Olena Productions and Hawaii Public Television. Chapin, Alonzo. ([1838], 1954). “Remarks on the Sandwich Islands, their Situation, Climate, Diseases, and their Suitableness as a Resort for Individuals Afflicted with or Predisposed to Pulmonary Diseases.” The Hawaiian Spectator 1: 248-267. ———. 1850. “Remarks on the Venereal Disease at the Sandwich Islands.” Boston Medical & Surgical Journal 42: 89-93. Daws, Gavan. 1973. Holy Man: Father Damien of Moloka. New York: Harper & Row. Day, A. Grove. 1977. Books About Hawaii: Fifty Basic Authors. Honolulu, HI: University of Hawaii Press. Diamond, Jared. 1996. Guns, Germs, and Steel: The Fates of Human Societies. New York: WW Norton. Dubois, A. 1952. “Quelques inoculations volontaires ou accidentelles de la lèpre aux êtres humains.” Royal Colonial Belge Bulletin des Séances 23: 199-213. Ellis, William. 1782. An Authentic Narrative of a Voyage Performed by Captain Cook and Captain Clerke, 1776-1780. London: G. Robinson. Enders, Frank N. 1877. “Leprosy as Observed in the Sandwich Islands.” Transactions of the International Medical Congress 1876. Philadelphia, 717-722. Farrow, John. 1954. Damien the Leper. Garden City, NY: Image Books, Doubleday. Feldman, William H. 1965. “Gerhard Henrik Armauer Hansen: What Did He See and When?” International Journal of Leprosy 33: 412-416. Foster, Burnside. 1898. “Leprosy and the Hawaiian Annexation.” North American Review 167: 300-305. Foster, George M. 1976. “Disease Etiologies in Non-Western Medical Systems.” American Anthropologist 78: 773-782. Goldman, Irving. 1970. Ancient Polynesian Society. Chicago: University of Chicago Press.

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Gordon-Cumming, C.F. 1883. Fire Fountains: The Kingdoms of Hawaii, Its Volcanoes and the History of the Missions. London: Blackwood. Green, Laura C. and Martha Warren Beckwith. 1926. “Hawaiian Customs and Beliefs Relating to Sickness and Death” American Anthropologist 28: 176-208. Greer, Richard A. 1969. “Oahu’s Ordeal: The Smallpox Epidemic of 1853.” In Hawaii Historical Review: Selected Readings edited by Richard A. Greer. Honolulu, HI: Hawaiian Historical Society, 37-88. Gulick, Luther H. 1855. “On the Climate, Diseases, and Materia Medica of the Sandwich (Hawaiian) Islands.” The New York Journal of Medicine 14: 169-211. Gussow, Zachary. 1989. Leprosy, Racism and Public Health Social Policy in Chronic Disease Control. Boulder, CO: Westview Press. ——— and George S. Tracey. 1970. “Stigma and the Leprosy Phenomenon: The Social History of a Disease in the Nineteenth and Twentieth Centuries.” Bulletin of the History of Medicine 44: 425-449. Halford, Francis John. 1954. 9 Doctors & God. Honolulu, HI: University of Hawaii Press. Hansen, G. Henrik Armauer. 1874. “Spedalskhedens Arsager.” Norsk Magazin for Laegevidenskaben 4: 7679. ———. 1955. “Causes of Leprosy.” International Journal of Leprosy 23: 307-309. Igler, David. 2004. “Diseased Goods: Global Exchanges in the Eastern Pacific Basin, 1770-1850.” American Historical Review 109: 693-719 Inglis, Kerri A. Forthcoming. “Criminalizing the Victims of Disease: Leprosy in Hawai‘i, 1865-1969.” In Engendering Health in the Pacific: Colonial and Contemporary Perspectives, edited by Vicki Lukere and Margaret Jolly. Judd, Charles S. 1977. “Depopulation in Polynesia.” Bulletin of the History of Medicine 51: 585-593. ———. 1984. “Leprosy in Hawaii, 1889-1976.” Hawaii Medical Journal 43: 328-329, 333-334. Kalisch, Philip A. 1973. “Lepers, Anachronisms, and the Progressives: A Study in Stigma, 1889-1920.” Louisiana Studies 12: 489-531. Kamakau, Samuel M. 1961. Ruling Chiefs of Hawaii. Honolulu, HI: Kamehameha Schools Press. ———. 1964. Ka Po‘e Kahiki (The People of Old). Honolulu, HI: Bishop Museum Press. Keers, R.Y. 1978. Pulmonary Tuberculosis: A Journey Down Through the Centuries. London: Baillière Tindall. Kneeland, Samuel. 1873. “On Leprosy, as it Exists in the Sandwich Islands.” The Richmond and Louisville Medical Journal 15: 404-411. Larson, Nils P. 1962/63. “The Highly Developed Art of Medicine in Old Hawaii.” Ciba Journal 24: 29-33. Lee, W.S. 1966. The Islands. New York: Holt, Rinehart and Winston. Lewis, Paul A. 1923. “The Relation of Heredity to Tuberculosis.” In Eugenics Genetics and the Family. Scientific Papers of the Second International Congress on Eugenics, edited by C.B. Davenport, et al. Baltimore, MD: Williams & Wilkins. London, Jack. 1914. “Koolau the Leper.” In The House of Pride and Other Tales of Hawai‘i. New York: Grosset & Dunlop, 47-94. López-Beltrán, Carlos. 1994. “Forging Heredity: From Metaphor to Cause, a Reification Story.” Studies in the History and Philosophy of Science 25: 211-235. Mackenzie, Morell. 1889. “The Dreadful Revival of Leprosy.” Nineteenth Century 26: 925-941. Merwin, W.S. 1998. The Folding Cliffs: A Narrative of 19th-Century Hawaii. New York: Alfred A. Knopf. Middleton, William Shainline. 1971. “Early Medical Experiences in Hawaii.” Bulletin of the History of Medicine 45: 444-460. Morrow, Prince A. 1897. “Leprosy and Hawaiian Annexation.” North American Review 16: 582-590. Mouritz, A[rthur]A[lbert] St. M[aur]. 1916. ‘The Path of the Destroyer’: A History of Leprosy in the Hawaiian Islands and Thirty Years Research into the Means by which it has been Spread. Honolulu, HI: Honolulu Star-Bulletin. Obeyesekere, Gananath. 1992. The Apotheosis of Captain Cook: European Mythmaking in the Pacific. Princeton, NJ: Princeton University Press. Osler, William and Thomas McCrae. 1915. Modern Medicine: Its Treatment and Practice. 2nd ed. Philadelphia: Lea & Febiger. Osorio, J.K.K. 2002. Dismembering Lâhui: A History of the Hawaiian Nation to 1887. Honolulu, HI: University of Hawaii Press. Ott, Katherine. 1996. Fevered Lives: Tuberculosis in American Culture since 1870. Cambridge, MA: Harvard University Press. Pearson, Karl. 1912. Tuberculosis, Heredity and Environment. London: Dunlau and Company. Ralston, Caroline. 1984. “Hawaii 1778-1854: Some Aspects of Maka’ainana Response to Rapid Cultural Change.” Journal of Pacific History 19: 21-40.

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Rokstad, Ingvald. 1964. “Gerhard Henrich Armauer Hansen.” International Journal of Leprosy 32: 64-70. Sahlins, Marshall. 1981. Historical Metaphors and Mythical Realities: Structure in the Early History of the Sandwich Island Kingdoms. Ann Arbor, MI: University of Michigan Press. ———. 1995. How Natives Think: About Captain Cook, For Example. Chicago: University of Chicago Press. Schmitt, Robert C. 1968. Demographic Statistics of Hawaii 1778-1965. Honolulu, HI: University of Hawaii Press. ———. 1970. “The Okuu – Hawaii’s Greatest Epidemic.” Hawaii Medical Journal 29: 359-364. Stannard, David E. 1989. Before the Horror: The Population of Hawai‘i on the Eve of Western Contact. Honolulu, HI: University of Hawaii Press. ———. 1990. “Disease and Infertility: A New Look at the Demographic Collapse of Native Populations in the Wake of Western Contact.” Journal of American Studies 24: 325-350. Stoddard, Charles Warren. 1895. The Lepers of Molokai. Notre Dame, IN: Ave Maria Press. Tebb, William. 1893. The Recrudescence of Leprosy and Its Causation. London: Swan Sonnenschein. Thompson, J. Ashburton. 1897. “Leprosy in Hawaii: A Critical Enquiry.” In Mittheilungen und Verhandlungen der Internationalen Wissenschaftlichen Lepra – Konferenz zu Berlin im October 1897. Berlin: August Hirschwald. Tobin, Jeffrey. 1994. “Savages, the Poor and the Discourse of Hawaiian Infanticide.” Journal of Polynesian Society 106: 65-92. Trask, Haunani-Kay. 1993. From A Native Daughter: Colonialism and Sovereignty in Hawai‘i. Monroe, ME: Common Courage Press. Tryon, J.R. 1883. “Leprosy in the Hawaiian Islands.” American Journal of the Medical Sciences 85: 443-450. “Tuberculosis.” 1911. In Encyclopaedia Britannica, 11th ed. Cambridge: Cambridge University Press. Veith, Ilza. 1992. “Parallels Between AIDS, Leprosy, and Syphilis.” Hawaii Medical Journal 51: 300, 302, 304. Wellman, Klaus F. 1968. “Notizen zur Geschichte des Aussatzes im Königreich Hawaii.” Sudhoffs Archive: Vierteljahrsschrift für Geschichte der Medizin und der Naturwissenschaften, der Pharmazie und der Mathematik. 52: 221-256. Weymouth, Anthony [pseudonym Ivo Gieke Cobb]. 1938. Through the Leper-Squint: A Study of Leprosy from Pre-Christian Times to the Present Day. London: Selwyn Blount. White, James C. 1882. “The Question of Contagion in Leprosy.” American Journal of the Medical Sciences 84: 433-454. Williams, C.T. 1882. “The Contagion of Phthisis.” British Medical Journal. 2: 618-621. Wilson, Philip K. 2002a. “Eugenicist Harry Laughlin’s Crusade to Classify and Control the ‘Socially Inadequate’ in Progressive Era America.” Patterns of Prejudice 36: 49-67. ———. 2002b. “Eighteenth-Century ‘Monsters’ and Nineteenth-Century ‘Freaks’: Reading the MaternallyMarked Child.” Literature and Medicine 21: 1-25 ———. 2003a. “Erasmus Darwin on Hereditary Disease: Conceptualizing Heredity in Enlightenment English Medical Writings.” In Cultural History of Heredity II: Eighteenth and Nineteenth Centuries, edited by S. Müller-Wille and H-J Rheinberger. Berlin: Max-Plank-Institut für Wissenschaftsgeschichte Preprint 247, 109-122. ———. 2003b. “Bad Habits from Bad Genes: 19th & 20th-Century Eugenics Attempts to Eliminate Syphilis and Associated ‘Defects’ from the U.S.” Canadian Bulletin of Medical History 20: 11-41. ———. Forthcoming. “Confronting ‘Hereditary’ Disease: Eugenic Attempts to Eliminate Tuberculosis in Progressive Era America.” Journal of Medical Humanities. Worboys, Michael. 2001. “From Heredity to Infection: Tuberculosis, 1870-1890.” In: Heredity and Infection: The History of Disease Transmission, edited by Jean-Paul Gaudillière and Ilana Löwy. London: Routledge, 81-100.

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How Cultural Is Heritage? Humanity’s Black Sheep from Charles Darwin to Jack London Marianne Sommer

Charles Darwin’s black sheep of my title was one of several near-synonyms of what was frequently referred to as atavism. Other related concepts were reversion (to type) or throwback, and arrest of development or stagnation. It seems that reversion was an important concept within plant breeding to refer to the reappearance of traits of the wild type.1 Used by evolutionists such as Herbert Spencer (1820-1903), Charles Darwin (1809-82), and Ernst Haeckel (1834-1919), atavism meant the complete or partial reversion of a plant or animal to an ancestral state. However, at a time when the mechanisms of heredity were unknown, concepts such as atavism had to remain equally diffuse. In addition, when applied to humans they became colored by socio-cultural attitudes of the time. Clearly, atavism was about questions of heredity from the start. Yet, possibly indicative of a general indeterminacy, the concepts of reversion and stagnation were not restricted to any particular theory of heredity or human evolution, associated with an emphasis on the so-called Lamarckian mechanism of the inheritance of acquired characteristics or Darwinian spontaneous variation and natural selection. Rather, atavisms regarded as caused by either of the two processes are often difficult to tell apart. For example, in a ‘Lamarckian’ framework, it was possible to see an impoverished environment as the cause of regression or stagnation, which in effect looked similar to the ‘Darwinian’ notion that an environment in which there were no selective pressures, or the wrong ones, might lead to regression or stagnation. In fact, within non-scientific discourses, the latter was sometimes seen as being possible within one individual.2 However, in my talk I have argued that reversion and arrest were intimately related to recapitulation theory, the belief that the individual in its embryonic development repeats stages its species passed during evolution. Michael Hagner has therefore asked the fundamental question of why atavism did not come to preoccupy non-biological discourses earlier, at the time of the recapitulationism of Naturphilosophen and transcendentalists such as Lorenz Oken, J. F. Meckel, and Etienne Serres.3 Indeed, as has been pointed out by Jonathan Harwood, the concept of atavism works particularly well within a linear-developmental view of evolution. At this point, I may only offer a few tentative remarks on the subject, deferring a more detailed discussion to a future opportunity. 1

2

3

The plant-breeding origin of the concept of reversion was stressed by Roger Wood. Where atavism is concerned, an article in Scientific Gardening of 1833 referred to Joseph Duchesne (1549-1609), when explaining atavism as designating the phenomenon that children often resemble one of their grandparents more than their parents (OED online, etymology). Even so, the difference between ‘hereditary’ and ‘acquired’ atavism became increasingly important for example in connection with criminality and homosexuality (e.g. Nye, 1976, who deals with acquired and inborn criminality; Williams, 1990, who discusses acquired versus congenital homosexuality; Pizer, 1961, who sees Frank Norris’ Vandover as acquired and McTeague as hereditary atavism (see below)). On these see for example Gould, 1977, pp. 39-52; Ospovat, 1976.

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Obviously, the notion of atavism was bound up with evolutionism, which in the Anglophone world mainly under concern here did not come to dominate either scientific or popular discourses until after Darwin’s On the Origin of Species (1859). At that time, a new kind of recapitulationism, in accordance with Darwinian evolutionism, became widespread. However, as I discuss elsewhere, the tree-like structures that came to be proposed as representations of phylogenies did not in all cases completely turn away from a linear sequence. This seems to have been particularly true for anthropology, where the acceptance of human antiquity and the turn towards evolutionism added the parameters of time and development to a racial hierarchy already in place in physical anthropology. Incorporating the new insights from comparative ethnology and prehistoric archaeology, an inevitable series of ever higher cultural and anatomical stages came to be seen as mandatory passages for all human races and civilizations in their evolution. This progression from savage to civilized, or primitive to modern, was analogized to individual embryonic development. As a consequence, fossil and living ‘non-white races’, children, the insane, etc., were interpreted as occupying stages of the human evolution of mind and body which Western man had passed in his ascent to the apex of nature and culture. The ‘savage races’ were understood as both simultaneously, offshoots of the line leading to the modern ‘civilized races’ and stages through which the latter had passed in their evolution. They were projected back in time, so that a scala naturae structure was essentially maintained.4 Thus, evolutionary progressionism had been accompanied by the counter-concepts of reversion and stagnation from the beginning. However, towards the end of the century, the general self-assuredness was increasingly troubled by a fear of regression and degeneration, both of which could signify the return to a primitive state.5 This was motivated among other things by the notion that struggle was essential to progress, combined with the observation that growing sections of Western societies were excluded from the fight for survival – a threat epitomized in such dandy and effeminate figures as Oscar Wilde.6 In the latter half of the nineteenth century until well into the twentieth, atavisms and their close relatives came to haunt philosophical, literary, sociological, and psychological discourses as the shady side of the progressionist paradigm. The wider cultural appeal of the biological concepts might exactly have been enhanced by their insufficient differentiation and imprecise definition within an evolutionary biology uncertain about the underlying mechanisms of hereditary continuity and change. They may have been symptomatic of a growing uncertainty, their pervasiveness signifying an attempt at ostracism and control over what was perceived as the nonor anti-progressive. While in this sense they seem to have functioned as an instrument for securing the status quo, atavism was also employed as a way of conceptualizing the duality of the human being – heir to animal instincts and cultured mind.

4 5

6

Sommer, 2005. Within a cyclic model of history, in which senility signified a return to an infantile condition, both degeneration and regression could indeed indicate a return to an ontological or phylogenetic earlier and lower stage. On progressionism and fears of degeneration see for example Bowler, 1989, 1993. For one of the most instructive documents of the fin-de-siècle culture and its concerns see Nordau, 1968 (1895).

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The atavistic disposition could therefore also be perceived as lurking within everyone. Somewhat paradoxically, the belief that anyone of us may fall victim to partial or full atavism was sometimes accompanied by a more positive interpretation of the phenomenon. Indeed, atavistic traits or events could become instruments of self-discovery, a means of integrating the individual with the common racial heritage, or of integrating the altruistic and civilized with the more selfish and animal aspects of personhood; a trend that would culminate in the new psychology at the dawn of the twentieth century. Although this is already a longer version of the talk I gave in Berlin, the paper offers no thorough discussion of atavism and similar concepts. Most of the questions that have been raised during the workshop have to remain unanswered for the time being. A closer analysis of the relation between degeneration and regression, as well as of the development of the concept of atavism over time in the contexts of progressionism, imperialism, theories of racial supremacy, growing nationalism, fear of miscegenation, eugenics, Mendelism and the new genetics, etc., will have to be put aside. Indeed, as Manfred Laublicher has pointed out, within a framework of hard heredity and Mendelian laws of inheritance, the meaning of atavistic traits and individuals must have changed considerably. However, these new insights were not incorporated into evolutionary anthropology until well into the twentieth century and neo-Lamarckian use-inheritance remained a favorite explanation for change. Also popular discourses in America as well as Britain drew from diverse expressions of evolutionism.7 The paper represents work-in-progress and the above attempt at a rough contextualization is meant as an invitation to continue the discussion initiated in Berlin. The paper aims at reconstructing hypothetical passages along which the concepts under concern might have traveled for one particular example – the American fiction writer, journalist, and adventurer Jack London (1876-1916). With the birth of modernism, old and new political, artistic, philosophical, and scientific ideas clashed. London felt caught in-between and struggled against paradoxes in life and writing. He appears to have seen in the notion of atavism a conceptual tool for overcoming opposites of old and new; first and foremost within the human being.

1. Books as tools of the trade: The production of atavisms in Jack London’s laboratory To identify some of the ideas that converged in the notion of atavism, I will approach the subject through London’s prehistoric fiction novel Before Adam, and discuss some of the possible scientific and non-scientific influences on its creation. This approach lends itself to an attempt at a wider cultural understanding of atavism for two reasons. The first is due to London’s writing practices. London referred to his farm at Glen Ellen, California, where he lived and worked, as a “writing laboratory”, in which his more than 15’000 books served as “tools”. London read and wrote according to a strict schedule. He had a sophisticated ordering system for his notes, references, newspaper clippings, correspondence, etc. However, the books he bought and that

7

E.g. Bowler, 1986, pp. 190-198; Oldroyd, 1980. For a critique of the biological notion of atavism see Montagu, 1938, 1945.

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were sent to him by far overextended the space of his library and colonized the house, including the barn. London’s laboratory, like those of so many of the scientists he read, functioned as a hub, where objects and information from all over the world were accumulated, analyzed, and synthesized into something new. London himself was an excessive reader, but also an adventurer and explorer, undertaking ‘field research’ at the places where his stories would be situated. It must be ascribed to London’s passion for sailing that he used the following analogy when explaining the function of his books as tools of his trade: I regard books in my library in much the same way that a sea captain regards the charts in his chart-room. It is manifestly impossible for a sea captain to carry in his head the memory of all the reefs, rocks, [...] harbors, points, lighthouses […] of all the coasts of all the world; and no sea captain ever endeavors to store his head with such a mass of knowledge. What he does is to know his way about in the chart-room, and when he picks up a new coast, he takes out the proper chart and has immediate access to all information about that new coast. So it should be with books.8

And London, in good white-suprematist fashion, conquered many new intellectual coasts, such as evolution, psychology, political economy, travel, navigation, philosophy, drama, poetry, and fiction. His claim to factuality, one suspects, was influenced by the American and French naturalists he studied, first and foremost Émile Zola (1840-1902). Indeed, when accused of plagiarism from The Story of Ab (1897) by Stanley Waterloo, London self-confidently replied that Before Adam presented rather something of a reply, as The Story of Ab was unscientific.9 It is therefore possible to ask what kinds of tools London could have used in his laboratory in the creation of his supposedly scientific Before Adam. In doing so, I will limit myself to three of the coasts London explored, evolutionary biology, literature, and psychology. The book’s preoccupation with atavisms and arrests of development constitutes the second reason for choosing this approach. The novel was first published as a series in Everybody’s Magazine from October 1906 to February 1907 (Figure 1). The main story evolves around three stages of hominization present at the same time in the Mid-Pleistocene. The Tree People are closest to modern anthropoid apes and as their name suggests, they live in trees. The Folk are physically and culturally more advanced than the Tree People. They are in the process of descending from the trees, have less body hair, are capable of rudimentary communication, and have tools. While the members of the Folk are more intelligent but less strong than the Tree People, they also lack the foresight and purpose of the Fire People, who are anatomically modern humans.

8

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Letter to The North American, 30 March 1913, reproduced in Hamilton, 1986, pp. 1-2. David Mike Hamilton, 1986, has investigated and catalogued London’s library and the annotations found in books. While many of the books I will discuss in this paper are explicitly mentioned by Hamilton, in other cases it is only known for certain that London was acquainted with the author. Hamilton, 1986, p. 26.

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Figure 1: The Structure of Jack London’s Before Adam (1906-1907) The three stages of human evolution are no truly independent side-branches of the human family tree; rather, the less progressive groups are cases of arrest of development, or stagnation. However, the fact that London had three different stages of human evolution live simultaneously would not have looked so odd to the contemporary reader. Evolutionists in general conceptualized ‘the lower human races’ as having stagnated at points which represented evolutionary stages in the white man’s prehistory.10 Similarly, more primitive fossil hominids were often interpreted as arrests of development in undemanding environments at the peripheries of the earth. 11 Like the contemporary ‘savages’ of London’s time, the Tree People and Folk are ‘living fossils’. 12 10 11 12

Stocking, 1982, Ch. 6. Sommer, 2005. London, 2000 (1906-1907), e.g. pp. 171, 224-225. In the story of Before Adam, London also described the prehistoric equivalent to the colonization and marginalization of ‘savages’ by ‘civilized Europeans’ underway at his time. The Folk are gradually being displaced and extinguished by the Fire People, who due to their increase in numbers have to gain new territories.

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Furthermore, the three groups are interconnected by missing links such as “Red-Eye, the atavism”. Although Red-Eye is one of the Folk, he represents a throwback to the earlier stage of human evolution occupied by the Tree People.13 Finally, the world of the Pleistocene hominids is accessed through the dreams of a man of London’s time. The narrator’s dreams about the life of his prehistoric ancestor Big-Tooth are resurfacing chunks of racial memory. In a way then, the narrator, too, is an atavism; one might say a psychological atavism: “[…] some strains of germplasm carry an excessive freightage of memories – are, to be scientific, more atavistic than other strains; and such a strain is mine.”14

2. Tools from biology: The concepts of atavism/regression/reversion and arrest/stagnation There can be no doubt that books on evolution played a central role in the making of London’s atavisms and evolutionary stagnations. By 1900, London had studied Spencer, Darwin, Thomas Henry Huxley (1825-95), and Haeckel among others.15 In the evolutionary theory of the Prussian biologist Haeckel, the so-called Biogenetic Law took center stage, according to which an individual’s ontogeny was essentially the shortened and accelerated repetition of its phylogeny. 16 Thus, in its embryonic development, the individual recapitulated the paleontological history of its phylum. This is of relevance, because as I will show the notions of atavism and stagnation were intimately related to the belief in recapitulation theory. With the spread of the Darwinian theory, the view of ontogeny of the Prussian-Estonian embryologist Carl Ernst von Baer (1792-1876) as a process of differentiation and individuation was applied to the view of evolution as another system of divergent development. 17 Thereby, the ideal (Naturphilosophie) and/or non-evolutionary (Cuvier, Owen, von Baer) notion of archetypes of taxonomic groups such as fish, reptiles, birds, and mammals were turned into real common progenitors, even if their fossil bones had not yet been found.18 Darwin, too, integrated von Baerian embryology and a view of evolution as a process of divergence into a recapitulationist framework. Already in the notebooks of the late 1830s, Darwin embraced recapitulation theory; in On the Origin of Species (1859) he argued: As the embryonic state of each species and group of species partially shows us the structure of their less modified ancient progenitors, we can clearly see why ancient and extinct forms of life should resemble the embryos of their descendants, – our existing species […] Embryology rises greatly in interest, when we thus look at the embryo as a picture, more or less obscured, of the common parent-form of each great class of animals.19 13

14 15 16 17 18

Another example of a connecting link is Big-Tooth’s partner the Swift One, who is related to the Fire People. Her father or mother might belong to that ‘higher race’. In general, there is a certain exchange between the three stages of development (London, 2000 (1906-1907), p. 135). London, 2000 (1906-1907), p. 20. E.g. Hamilton, 1986, pp. 8, 14. E.g. Haeckel, 1898 (1868), p. 190. Baer, Carl Ernst von. Über Entwicklungsgeschichte der Thiere. Beobachtung und Reflexion. Königsberg, Teil 1: 1828, Teil 2: 1827. Haeckel, 1898 (1868), Vortrag 13, pp. 314-315 on the three-fold parallelism between comparative embryology, the classification system, and comparative phylogeny, all of which resulted in a tree. Spencer, too, argued for the threefold parallelism (e.g. Spencer, 1897 (1866), Part II, Ch. 2, and Part III, Ch. 5).

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Thus, for the ‘Darwinian bulldog’ Huxley, the pronounced similarity of human and ape embryology was the strongest evidence in favor of their close kinship – too close, as we will see, for humans to be quite safe from the ape within: Startling as the last assertion [that it is only in the last stages of development that the young human being presents marked differences from the young ape] may appear to be, it is demonstrably true, and it alone appears to me sufficient to place beyond all doubt the structural unity of man with the rest of the animal world, and more particularly and closely with the apes.20

To turn now to the notion of atavism, which Haeckel defined in the Natürliche Schöpfungsgeschichte as an organism that represents a throwback to the state of a long lost ancestral generation,21 we find a possible explanation for the phenomenon in Darwin’s The Variation of Animals and Plants Under Domestication. At the outset of the chapter on “Reversion or Atavism”, Darwin gave several terms for the phenomenon besides atavism and reversion, such as throwback, pas-en-arrière, Rückschlag, and Rückschritt.22 Here, Darwin conjectured that in sexual reproduction, the characters of both parents would exist in the offspring in a double state, once blended and once kept apart. The usual case of character expression would be through blending of the parent variants.23 However, there were cases where certain traits were predominant, as he called it, in which the plant or animal offspring may revert to a character thousands of generations remote: “From these several facts it must be admitted that certain characters, capacities, and instincts, may lie latent in an individual, and even in a succession of individuals, without our being able to detect the least sign of their presence.”24 The reversion of characters could be catalyzed by a change in living conditions or by hybridization, or simply happen for no apparent reason. With regards to human races, Darwin speculated that “[…] the degraded state of so many half-castes is in part due to reversion to a primitive and savage condition, induced by the act of crossing [between two races, both low in the scale], even if mainly due to the unfavourable moral conditions under which they are generally reared.”25 Already from these considerations it seems that, particularly where human atavisms were concerned, there was neither a precise definition of reversion, nor were partial and full reversions clearly distinguished. While so far there seems to be no necessary connection between atavism and recapitulation theory, there was such a connection for the concept of arrest of development. Darwin was well aware of the fact when he wrote in The Descent of Man: “[…] it is hardly credible that a complex part, arrested at an early phase of embryonic development, should go on growing so as ultimately 19 20

21 22 23 24

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Darwin, 1964 (1859), pp. 449-450. London, 2000 (1906-1907), p. 172, also followed the von Baerian model. Man’s Place in Nature (1863), in Huxley, 1894, p. 92. On recapitulation theory see Gould, 1977; Ospovat, 1976; Russell, 1916; on Darwin’s recapitulationist ideas see for example Darwin, 1964 (1859), pp. 449450, and Richards, 2002, pp. 526-533. E.g. Haeckel, 1898 (1868), p. 186. Darwin, 1998 (1883), p. 1. Darwin spoke of characters “blending together” (e.g. Darwin, 1998 (1883), p. 11). Darwin, 1998 (1883), p. 29. This notion seems similar to Haeckel, 1898 (1868), pp. 184-186, “unterbrochene oder latente Vererbung”. Like Darwin, Haeckel thought that a return to the wild often caused an animal or plant species to revert to earlier generations. Darwin, 1998 (1883), p. 21.

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to perform its proper function, unless it had acquired such power during some earlier state of existence, when the present exceptional or arrested structure was normal.” 26 But how could such an arrest of development then unambiguously be distinguished from an atavism? In many cases it could not: “When a structure is arrested in its development, but still continues growing, until it closely resembles a corresponding structure in some lower and adult member of the same group, it may in one sense be considered as a case of reversion.” 27 As an ontological model of evolution suggests, a more primitive phylogenetic stage might be reached either through reversion, or by stagnating in one’s embryonic development. Thus, Darwin conceded that microcephalous idiots in as far as their brains resembled those of apes, like many other cases of arrest of development, might just as well be classified as reversions. Thereby both arrest of development and atavism were placed within a recapitulationist framework, which turned them into catchalls for the ape-like, the primitive, the ‘savage’, the childish, and the ‘idiotic’: Arrests of Development.– […] It will suffice for our purpose to refer to the arrested brain-development of microcephalous idiots […] Their skulls are smaller, and the convolutions of the brain are less complex than in normal men. The frontal sinus, or the projection over the eyebrows, is largely developed, and the jaws are prognathous to an “effrayant” degree; so that these idiots somewhat resemble the lower types of mankind. Their intelligence, and most of their mental faculties, are extremely feeble […] They often ascend stairs on all-fours; and are curiously fond of climbing up furniture or trees. We are thus reminded of the delight shewn by almost all boys in climbing trees […] and several cases have been published of their bodies being remarkably hairy.28

Obviously, in this discussion of the phenomenon of arrest of development, supposedly phylogenetically and ontogenetically earlier and lower stages, such as ‘non-white races’, children, and apes (and animals in general), were brought in to characterize the arrested or reverted state of microcephalous idiots. Again, it seems as though the atavistic trait of a microcephalous brain rendered the affected individuals atavistic in general morphology and behavior. London’s atavism in Before Adam, called Red-Eye, closely corresponds to the concept within evolutionary theory (Figure 2). Although Red-Eye lives among the Folk, he represents a regression to the evolutionary lower and more apish type of the Tree People: “He [Red-Eye] was a monster in all ways. Physically he was a giant […] He was abominably hairy […] Furthermore, it was a rare individual among us who balanced himself with his knuckles when walking. Such an individual was an atavism, and Red-Eye was an even greater atavism.”29 As becomes clear from both London’s description and Charles Livingston Bull’s (1874-1932) illustration of Red Eye in 26 27 28

29

Darwin, 1874 (1871), p. 54. Darwin, 1874 (1871), p. 54. Darwin, 1874 (1871), pp. 52-54 (since Darwin used italics, I have marked keywords bold). Darwin based his speculations on the microcephalous condition on Carl Vogt (1817-95), who argued that fossil hominids, ‘lower extant races’ such as ‘the Negro’, and microcephalous idiots represented missing links between the living white races and the recent great apes. The microcephalous idiot was interpreted as arrest of development (Entwicklungshemmung), “[…] welche eine der Stationen bezeichnet, die der menschliche Embryo nothwendig durchlaufen muss und welche durch die Mischung menschlicher und afflicher Charaktere jetzt noch in ihrer Abnormität die Zwischenbildung bezeichnet, die früher in normaler Bildung bestanden haben kann” (Vogt, 1863, p. 278). London, 2000 (1906-1907), pp. 56-59.

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aggressive bearing, he is marked by the primitive morphological traits enumerated by Darwin; but he also betrays a primitive mind.

Figure 2: “It was Red-Eye”; Illustration by Charles Livingston Bull, American wildlife artist, chief taxidermist at the National Museum in Washington D.C. This is further emphasized by the introduction of yet another term, that of the monster, which reconnects the notion of atavism to the older view of monstrosities as the outcome of embryology gone awry, and which brings the very negative connotations to the fore. In The Expression of the Emotions in Man and Animals, where Darwin strongly relied on scales of mental capacity of animal-primitive/savage/insane/child-white adult male, he included lengthy opinions that expressed the belief that recapitulation theory could explain the insane. The ‘idiot’ and the insane were linked to a brute state of humankind dominated by lower instincts:

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Dr Maudsley, after detailing various strange animal-like traits in idiots, asks whether these are not due to the reappearance of primitive instincts – ‘a faint echo from a far-distant past, testifying to a kinship which man has almost outgrown’. He adds, that as every human brain passes, in the course of its development, through the same stages as those occurring in the lower vertebrate animals, and as the brain of an idiot is in arrested condition, we may presume that it ‘will manifest its most primitive functions, and no higher functions’. Dr Maudsley […] asks, whence come ‘the savage snarl, the destructive disposition, the obscene language, the wild howl, the offensive habits, displayed by some of the insane? Why should a human being, deprived of his reason, ever become so brutal in character, as some do, unless he has the brute nature within him?’ The question must, as it would appear, be answered in the affirmative.30

As in the scientific literature, where the stagnations at or reversions to earlier evolutionary stages might affect an individual in its entirety, Red-Eye’s regression to ape anatomy in Before Adam coincides with a reversion to ape mind and behavior. Red Eye is driven by animal instincts: Red-Eye was an atavism. He was the great discordant element in our horde. He was more primitive than any of us. He did not belong with us, yet we were still so primitive ourselves that we were incapable of a cooperative effort strong enough to kill him or cast him out. Rude as was our social organization, he was, nevertheless, too rude to live in it. He tended always to destroy the horde by his unsocial acts. He was really a reversion to an earlier type, and his place was with the Tree People rather than with us who were in the process of becoming men.31

The atavism appears as the epitome of all threats to progress, and the term came to encompass all kinds of undesirables. In analogy to animal breeding, Darwin recognized the need of a society to dispose of such destructive elements as the insane and the criminal through confinement or execution. Other problems were seen as to a certain degree self-regulatory since the violent or intemperate, the restless, or the profligate tended to die prematurely, to emigrate, or to bear few children. In any case, they were not to hand on their dispositions to the next generation: In the breeding of domestic animals, the elimination of those individuals, though few in number, which are in any marked manner inferior, is by no means an unimportant element towards success. This especially holds good with injurious characters which tend to reappear through reversion, such as blackness in sheep; and with mankind some of the worst dispositions, which occasionally without any assignable cause make their appearance in families, may perhaps be reversions to a savage state, from which we are not removed by very many generations. This view seems indeed recognised in the common expression that such men are the black sheep of the family.32

Black sheep as a designation for throwbacks, reversions, atavisms, arrests, or stagnations seems more than any other to convey the multiple meanings of the phenomenon referred to. Like black sheep, atavisms were not only about physical appearance, but also alluded to mental and behavioral traits. They were the unwanted elements in society at large. As indicated by the quote from Before Adam above, the Folk cannot heed Darwin’s advice of isolating atavistic individuals, 30 31 32

Darwin, 1998 (1872), p. 241. London, 2000 (1906-1907), p. 111. Darwin, 1874 (1871), p. 211.

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because they have not yet developed the high degree of sociality needed for such a common effort. As the examples from Darwin and London show, the atavism was generally driven by (even) lower instincts than his fellow creatures. Among civilized people, the atavism particularly lacked the latter’s highly developed social and moral instincts. Judged by contemporary readers, Red-Eye’s unsocial behavior would amount to the criminal – he even kills his own wives. The link to the criminal anthropology of the Italian Cesare Lombroso (1838-1909), who had even reviewed Zola’s La bête humaine (1890, The Human Beast) for its ‘scientific accuracy’ with regards to criminal types,33 seems thus an obvious one. As is well known, for Lombroso, the stigmata of the born criminal represented atavistic features of an evolutionary past. Thus originated the belief that what was regarded as criminal in civilized societies might be normal behavior among ‘savages’, animals, and for children, a belief instantiated by Red-Eye, who ends up completely at ease among the apish Tree People.34 Not surprisingly then, London kept a file on ‘crime and criminals’ and read Enrico Ferri’s Criminal Sociology (1899).35

3. Tools from literature: Instincts as missing links between biology and culture From the above observations it appears that for the atavism as social-misfit and criminal, instinct theory played a central role. Darwin and Spencer, particularly in the editions of The Principles of Psychology published after The Descent of Man, provided similar explanations for the evolution of social instincts and a moral sense.36 They saw the origin of the complex social and moral instincts that distinguish humanity in lower and more common ones. They agreed that the lower instincts, which represented compound reflex actions, were the result of habitual (stimuli and associated) behaviors and thoughts that had impacted brain anatomy. Through the mechanism of the inheritance of acquired characteristics, the modifications in brain structures were passed on to the next generation. Darwin and Spencer regarded sympathy as the basis of altruistic behaviors. Through the capacity to empathize, the individual would learn to suffer from the community’s condemnation and feel happy when praised. With the possibility to reflect on one’s actions conscience arose, which would strengthen the social over the selfish instincts through habit. They also agreed that group selection had catalyzed the development of the highest instincts of sociality and morality. While humans would have obeyed the social imperatives with regards to their own tribe, in a primitive state, they would not have done so towards other tribes – a quality which would not have been judged as amoral. Where they differed was in the way they thought these higher instincts had arisen in the first place. While Darwin in The Descent of Man saw the main factor in the gradual addition of small and spontaneous but hereditary variations to the lower instincts, Spencer’s 33 34 35

36

Lombroso, 1891. London, 2000 (1906-1907), p. 242. See also Lombroso, 1895. Lombroso’s L’Uomo delinquente (1876) was not translated into English until 1911; but his ideas were made known through Havelock Ellis’ The Criminal (1892) and Arthur MacDonald’s Criminology (1893) (Pizer, 1961). In On the Origin of Species (Darwin, 1964 (1859), Ch. 7 – case of neuter insects), Darwin seems to have ascribed more power to spontaneous variations and natural selection in the development of instincts than in The Descent of Man. On the other hand, Spencer increased the role of selection after the first edition of Principles of Psychology, so that their instinct theories converged over time (Young, 1990 (1970), pp. 186-190).

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instinct theory in The Principles of Psychology stressed the impact of the individual’s will and effort through the inheritance of acquired characteristics.37 Generally, that mechanism lay at the heart of Spencer’s Synthetic Philosophy that advocated for a universal law of progress guiding biological, psychological, and social evolution. 38 It ensured that individual or cultural achievements would lead to the corresponding advance in anatomy and mind. Also here, an integrating factor was the von Baerian theory of embryology as a process of increasing differentiation, which Spencer regarded as analogous to an increasing distribution of labor and overall complexity in the organism and in society. Spencer’s emphasis on the individual’s self-determination made the existing social hierarchy inevitably a just one. By analogy, so was the hierarchy of races, since the stage of development of each race reflected its individuals’ combined effort or lack thereof.39 The ‘Spencerian’ combination of the individual’s self-responsibility and a ruthless struggle for survival found wide entry into literary creations. Clearly, Spencerian philosophy was part of the bedrock of London’s worldview. Among London’s favorite authors whose books functioned as tools of his trade were Frank Norris (1870-1902), O. Henry (William Sydney Porter 1862-1910), and Robert Louis Stevenson (1850-94), all of whom dealt with atavistic phenomena in their writings, and might have fought with the threat of regression in their real lives. 40 As in Before Adam, the reversions to lower types they expounded were primarily due to a loss of the particularly human instincts. The Jekyll-Hyde case can clearly be interpreted as atavistic (Dr. Jekyll and Mr. Hyde 1886). Hyde is described as ape-like, of small and muscular stature, hairy, and as a troglodyte. Jekyll represents the normal state of the human being who has risen above his ape ancestry through the acquisition of among other things highly developed moral and social instincts that work as checkers of the selfish animal drives. Similar to London’s Red Eye, the violent, uncontrolled, wanton, and above all criminal, Hyde is the disrupting element among the civilized circle of London gentlemen. In accordance with his animal instincts, Hyde has no conscience: “his every act and thought centered on self; drinking pleasure with bestial avidity from any degree of torture to another; relentless like a man of stone.”41

37

38

39

40

41

Spencer thought that the more complex the organism, the smaller the effect of natural selection. In humans, changes in morphology and behavior were therefore seen as nearly exclusively the result of useinheritance, as natural selection was hindered from eliminating the unfit. However, between human societies, group selection was considered to be still at work (Spencer, 1897 (1866), Vol. I, Part III, Ch. 13; also note that Spencer, 1897 (1866), Vol. I, Part II, Ch. 9, explained ‘spontaneous variation’ as the effect of the recombination of individual direct adaptations). On instinct theory see Spencer, 1895 (1855), Vol. I, Part IV, and Vol. II, Part IX, Chs. 5-8; Darwin, 1874 (1871), Chs. 3-5. Both Spencer and Haeckel criticized August Weismann (1834-1919) for his rejection of the inheritance of acquired characteristics. Haeckel saw particularly with regards to the evolution of instincts no other satisfactory explanation than that they were psychic habits turned heritable (Spencer, 1893, 1893; Haeckel, 1898 (1868), p. 192). On Spencer see also Bowler, 1989, pp. 37-39, 153-154, 157-158, 194-195; 1993, pp. 65-69; Richards, 1987, Chs. 6-7; on social Darwinism in particular see Bowler, 1989 (1983), pp. 285-291, and Bowler, 1990, Ch. 10. Hamilton, 1986, pp. 11-12, 256-258. Darwin himself experienced something akin to a regression to a stage of a lower aesthetic sense when he lost his capacity to enjoy Shakespeare and poetry and replaced them by easily digestible novels (see Levine, 2003, pp. 46-47). Stevenson, 1981 (1886), p. 87.

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Jekyll, in contrast, yearns for the approval and fears the disapproval of his peers. The JekyllHyde problem seems like an aggravation of the condition of ordinary civilized man, still vessel of selfish animal instincts imperfectly controlled by a higher moral sense: “But I had voluntarily stripped myself of all those balancing instincts by which even the worst of us continues to walk with some degree of steadiness among temptations; and in my case, to be tempted, however slightly, was to fall.”42 In accordance with an evolutionary understanding, the potion Jekyll takes in the attempt to bring about a complete dissociation of the two natures, produces a Hydeexistence rid of Jekyll, but there is no Jekyll free of Hyde.43 Jekyll is an inherently composite being, and the more he gives in to Hyde, the less there is left of the non-Hyde part of Jekyll’s body as well as mind. Norris’s tragic hero in Vandover and the Brute (1895, 1914), too, fails in his effort to live up to the moral standard. Like London, Norris held a mix of ‘Darwinian’ and ‘Spencerian’ ideas that made him agree with the anthropological doctrine that without selection, effort, and struggle there would be stagnation or even regression. Western civilization was seen as threatened by growing subcultures that doted on comfort. Whether it was the undemanding environment in which ‘the savages’ stagnated or the ‘lazy and effeminate’ fin-de-siècle dandy cultures, the detrimental effect on mind and body was perceived as analogous.44 Norris’s Vandover, when choosing the company of prostitutes, freaks, gamblers and alcoholics over that of respectable society, automatically adapts to his new surroundings. Rather than finding himself under different selective pressures, the atmosphere of vice is portrayed as a world bare of struggle, where men and women self-indulgently succumb to their animal instincts, giving in to regression: “His [Vandover’s] intellectual parts dropped away one by one, leaving only the instincts, the blind, unreasoning impulses of the animal.” 45 It is only when re-entering the productive world of business and industry that Vandover finds himself subjected to a fierce struggle demanding an effort that he is increasingly unable to live up to. The habits acquired in the late hours of the night have reduced him to the brute and have finally brought him beyond the reach of those checking agents, the attitude of society, conscience, and remorse. Through the figure of Vandover’s friend, Charlie Geary, however, the reader learns that for Norris our atavistic sides were not entirely negative. The rudiments of the brute within were necessary to make one’s way in the struggle for survival and success, albeit under the guidance of an intellect sharpened in competition. But Vandover succumbs, gliding into madness and idiocy, and in the end like Jekyll literally reverts to the animal. He falls to the floor on all fours and in any other way turns into a wolf – his physical and psychic reversion are always in step. Even Vandover’s environment seems to regress 42 43 44

45

Stevenson, 1981 (1886), p. 93. E.g. Stevenson, 1981 (1886), p. 85. For Spencer, for example, women, especially those of inferior rank, ‘savages’, and primitives possessed lower intellectual faculties due to limited experiences in an impoverished environment. However, their faculties were also physically underdeveloped as a consequence of continued under-use over many generations. On the basis of the mechanism of the inheritance of acquired characteristics, the stage of an individual’s or group’s mental (including intellect as well as emotions) and anatomical stage were at corresponding levels (e.g. Spencer, 1895 (1855), Vol. II, Part IX, Ch. 3). The related argument that the cessation of the cause for a certain adaptation leads to the regression of the organ or individual to the state before the cause came into effect can be found in Spencer, 1897 (1866), Vol. I, pp. 199-200. Norris, 1986 (1914), p. 228.

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to primeval times: “The room was small, and at some long-forgotten, almost prehistoric period had been covered with a yellowish paper, stamped with a huge pattern of flowers that looked like the flora of carboniferous strata, a pattern repeated to infinity wherever the eye turned.” 46 The monotonous repetitiveness of the wallpaper seems to mirror the infinite repetition of phylogeny in each individual’s ontogeny and the associated struggle against reversion. As a last possible literary tool of London’s trade, I will turn to a story by O. Henry of the title The Atavism of John Tom Little Bear, which was published July 1903 in the same serial in which London published Before Adam. In it, one encounters a more deterministic kind of atavism. In nineteenth-century evolutionism, the environment was still seen as formative for the development of a human race, but the strong adaptationism of eighteenth-century environmentalism had given way to a less flexible notion of racial variety. The differences created in the evolutionary long-run were perceived as having become hard-wired.47 Similarly, the narrator in the story’s frame claims that the ‘lower races’ are prone to regress to ‘their primitive state’ despite all attempts at civilization. Like Red-Eye, the tropical man “[wi]ll be happiest in his own way.” 48 To convince his guest of the futility of education in the case of ‘the savage’, the narrator tells the story of the educated Cherokee John Tom Little Bear, who had apparently completely assimilated white American culture, but who in a moment of crisis relapsed to his original state of primitive Indian. When a boy and his mother, for both of whom Little Bear deeply cares, are threatened, Little Bear, aided by the brutalizing force of liquor, regresses to ‘the subhuman ways of the savage’, becomes criminal, by murdering the dangerous man and taking his scalp as a trophy. At this turning-point, this atavistic moment, he re-assumes the behavior as well as looks of the Cherokee. He even looses his polished English and reverts to the Creole. When the narrator and Little Bear wake up the following morning, the narrator is happy to see “the nineteenth century” in the Indian’s eyes again. Although firewater may have played its part, Little Bear adds: “Combined […] with the interesting little physiological shake-up known as reversion to type. I remember now.” 49 O. Henry’s deeply cynical farce renders it dramatically obvious how the fall back to a lower stage of development was perceived as a fall back in time. As indicated in this last quote, it was therefore associated with the process of remembering, just as progress was associated with the process of learning. This leads on to London’s last atavism in Before Adam, the psychological peculiarity of the narrator.

4. Tools from psychology: Dreams as psychological regression to racial memory In “A Biographical Sketch of an Infant” (1877), Darwin introduced the notion of a phylogenetic memory on the basis of a recapitulationist interpretation of mental development. At the experience that his son was afraid of the animals of prey at the zoo without ever having had the chance to learn that fear, Darwin speculated: “May we not suspect that the vague but very real fears of children, which are quite independent of experience, are the inherited effects of real dangers and abject superstitions during ancient savage times?”50 The notion of traces of experiences 46 47 48 49

Norris, 1986 (1914), pp. 234-235. Stocking, 1982, pp. 42-68, 110-132; see also footnote 44. O’Henry, 1903. O’Henry, 1903.

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transferred from a prehistoric past was central for the structure of Before Adam, and London described an almost identical instant in the childhood of the narrator, when he is taken by his father to the circus and instinctively fears the lion, even though London took the concept one step further from instinct-like and unconscious memory to conscious remembrance: Nevertheless, it was with fear and trembling, and with much encouragement on his [the father’s] part, that I at last approached the lion’s cage. Ah, I knew him on the instant. The beast! The terrible one! And on my inner vision flashed the memories of my dreams, – the midday sun shining on tall grass, the wild bull grazing quietly, the sudden parting of the grass before the swift rush of the tawny one, his leap to the bull’s back […].51

In Before Adam, more has been handed down to the narrator than an irrational fear of predators. He has access to a memory that in the average person remains hidden in the unconscious – the race memory; it provides the view back through time into our Pleistocene ancestors’ lives. Through his dreams, the narrator, who carries “an excessive freightage of memories”, who is more “atavistic than other strains”, establishes continuity between the individual conscious and a racial unconscious. The notion of instincts as unconscious memory was present in both Darwin and Spencer,52 and London’s explanation of racial memory seems an amalgam of the current biological and psychological theories: Now a fall [of our arboreal prehistoric ancestors from the trees], averted in such fashion [by clutching branches], was productive of shock. Such shock was productive of molecular changes in the cerebral cells. These molecular changes were transmitted to the cerebral cells of progeny, became, in short, racial memories […] There is nothing strange in this [the heritability of memory], any more than there is anything strange in an instinct. An instinct is merely a habit that is stamped into the stuff of our heredity, that is all.53

The way in which London worked with instinct theory, racial memory, and dreams brings to mind the development of these notions in Sigmund Freud (1856-1939). Indeed, London, who had a great interest in psychology, processed texts by Freud in his laboratory. 54 In Totem and Taboo (1913), the way in which individual child experiences might produce neurosis in the adult, was seen as analogous to the fact that experiences accumulated in the childhood of the race, in its primitive state, might haunt a race in its mature, civilized state. Freud meant to bridge the gap between the non-analytic folk-psychology of Wilhelm Wundt (183250

51 52

53 54

Darwin, 1877, p. 288. Spencer, 1895 (1855), e.g. Vol. I, pp. 461-462, too, conceptualized mental evolution as analogous to mental development, which allowed him to use the minds of ‘savages’ and children as evidence of the evolution of the civilized mind from animal mind. London, 2000 (1906-1907), pp. 7-8. See particularly Spencer, 1895 (1855), Vol. I, p. 452, where the transformation of conscious memory into unconscious, organic, and heritable memory is likened to the process of learning where, too, the effort gets less with each repetition, until the habit becomes automatic and unconscious. The heredity-memory analogy guided many evolutionary theories of mind of the last quarter of the nineteenth century (see particularly Samuel Butler, e.g. Unconscious Memory 1880; but also Karl Ewald Hering, “Über Gedächtnis…” 1870 [trans. in Butler]; Richard Semon, Die Mneme 1904; Edward Dinker Cope, The Primary Factors of Organic Evolution 1896). London, 2000 (1906-1907), p. 14. Hamilton, 1986, pp. 129-130.

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1920) and the psychoanalysis of the Zurich school.55 In the small book which contained four articles first published in Freud’s Imago (1912/3), one finds ample expression of the recapitulationist theory and its tenet that ‘primitive human races’ represent frozen ancestral states of western civilized man. ‘Savages’ were prehistoric humans still living amongst us. For psychology, they were of particular interest as illustrations of an earlier stage of mental evolution. Since the neurotic was a kind of regression or atavism, his or her psychology was expected to show similarities to the psychology of ‘the savage’.56 Freud assumed that the Darwinian primeval horde had really existed, and that actual events had led to the initial acquisition of the Oedipus complex that then became part of our phylogenetic heritage.57 In the prehistorically reified Oedipus complex, Freud saw the explanation for all neuroses.58 Without the assumption of a collective psyche that would guarantee continuity in the emotional life of humans, that could bridge the gaps created in the stream of souls through the death of individuals, folk-psychology could not exist.59 Most importantly in this context, Freud not only thought that the individual’s memory spanned beyond his or her life, but saw dreams as the means through which experiences that reached back into prehistory were brought to the surface. Dreams represented regressions to the earliest conditions of the dreamer, a revival of his infancy. Beyond that, the dream might lead to the mental infancy of the race. 60 Even more than Freud’s, Carl Jung’s (1875-1961) psychology meant a revelation to the London who was approaching the end of his life. In fact, London read Psychology of the Unconscious (1916) in his last year.61 Here, Jung presented myths as the collective dreams of young humanity, resurfacing in the dream of the modern individual, which was a form of historical regression to the infantile stage of the human race. In parallel to recapitulation theory based on evidence from comparative anatomy and embryology, Jung analogized the way of thinking of ‘the lower human races’, to the myths of the ancients, to the fantasies of children, and to the dream of the modern civilized adult: “[…] ontogenesis corresponds in psychology to phylogenesis.” 62 This turned the dream into a re-echo of the ancient and prehistoric.63 In the earliest blossoms of psychoanalysis, London finally found a solution to a paradox that had troubled his life as well as his literary work: How were the individual (the hero, the superman,

55 56 57 58 59 60

61 62

Freud, 1913, p. 5. Freud, 1913, p. 7. In Totem and Taboo, Freud already described the neurotic constitution as atavistic. Freud, 1913, pp. 158-161. On Darwin’s influence on Freud in general see Ritvo, 1990. Freud, 1913, p. 174. Freud, 1913, pp. 175-176. On Freud’s concept of racial memory see also Heyman, 1977; Slavet, In progress; Stewart, 1976. E.g. Freud, 1991 (1900), pp. 539-540. Note that in Studies on Hysteria (1895), Freud theorized hysteric traits as rudimentary organs; in Before Adam, on the other hand, the narrator, whose atavistic psychology gives him conscious access to the race memory, is described as hysterical (London, 2000 (1906-1907), p. 9). Not only in his developmental conception of mind, Freud had other precedents than Spencer, but also his dream theory, that is the notion of dreams as access to primeval mental states of humankind, had been anticipated by such figures as the neurologist John Hughlings Jackson (1835-1911), the psychiatrist James Sully (e.g. “The Dream as a Revelation,” Fortnightly 59 (1893), pp. 354-365), and Havelock Ellis (1859-1939). This was true for Britain as well for the U.S. and France, where the notions of mental regression, dissolution, and involution were taken on before 1900 (Sulloway, 1979, pp. 257-275, 321327). Jung, 2002 (1916). See also McClintock, 1970. Jung, 2002 (1916), p. 28.

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the adventurer, the colonizer) and the social (altruism, universal love, brotherhood) to be reconciled? In his literary atavisms such as Red-Eye, London problematized these opposing demands, which were linked to the animal and moral aspects of human nature. 64 Jung emphasized that it was this dual human nature that lay at the bottom of pathological as well as nonpathological psychological problems. Jung saw the process of civilization in a progressive submission and domestication of the animal in humans, which could not be carried out without rebellion from the part of the freedom-loving vestiges of the primordial soul. The neurotic therefore experienced an aggravated condition of a universal human conflict; being at odds with oneself was seen as characteristic of the cultured human being (Kulturmensch).65 The ways in which the novels by Stevenson and Norris foreshadowed the explanation the new psychoanalysis provided for the inner human struggle was thus made strikingly obvious by Jung, who defined neurosis as the symptom of an unsuccessful attempt to synthesize the opposing demands of the conscious moral ideal and the according to contemporary opinion immoral ideal of the unconscious.66 London’s narrator in Before Adam might be seen as approaching the psychoanalytic understanding particularly close, since for Jung and Freud the unfulfilled wish of the unconscious was of infantile, if not animal or prehistoric origin; it had been born in childhood or beyond and thus no longer fit the contemporary adult moral world. In Before Adam, one also encounters the struggle of the individual to distinguish the self from the other, which for Jung was synonymous with the difference between personal (un-)conscious and collective unconscious, and to leverage his or her dual nature. According to Jung, the recognition of the psychic reality of the other within the self would allow to use the energy before bound up in the confusion of the two for their synthesis.67 However, failure of recognition opens up the possibility of interpreting the notion of atavism in psychoanalytic terms as a projected archetype. The atavism would then not so much represent an aspect of those who are thus called, but rather the repressed irrationality of society at large that in a powerful resurfacing is projected on those that are feared.68 In Before Adam London described the dissociation into a wake-a-day personality with a memory of the experiences of the individual and a dream personality with a memory of past-day

63

64

65 66 67 68

Jung, 1971 (1916), pp. 71, 78-79. This interpretation of dreams London met with in Freud, Jung, and also Friedrich Nietzsche (1844-1900) (e.g. 1879. Human, All Too Human: A Book for Free Spirits, trans. Marion Faber with Stephen Lehmann. Lincoln: University of Nebraska Press, 1984, e.g. Vol. II, p. 27, where Nietzsche explained that in the dream an atavistic relic of humanity manifested itself, since the dream represented the way of thinking of prehistoric man). See also Mills, 1955. Note that this tension can be seen within non-atavistic individuals, too. Lop-Ear, for example, stays with the wounded Big-Tooth even though this is dangerous and “he is anxious to be gone”, and the narrator takes this as “a foreshadowing of the altruism and comradeship that have helped make man the mightiest of the animals”. He “often meditates upon this scene – the two of [them], half-grown cubs, in the childhood of the race, and the one mastering his fear, beating down his selfish impulse of light, in order to stand by and succor the other” (London, 2000 (1906-1907), p. 91). Jung, 1971 (1916), p. 22. Jung, 1971 (1916), p. 23. Jung, 1971 (1916), pp. 74-75. Jung, 1971 (1916), p. 97, explained for example the devil in this way, as an avatar of the shadow archetype, that is of the dangerous aspect of the unrecognized dark half of the human being. Besides the devil, Jung identified the demon as another main archetype, which often appears in the guise of the medicine man, old, dark-skinned and of the mongoloid type.

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race experiences.69 In this sense as in Freud’s and Jung’s, the racial memory really was the memory of an other, a memory that may lead back to the very beginning: “For Big-Tooth also had an otherself, and when he slept that other-self dreamed back into the past, back to the winged reptiles and the clash and the onset of dragons, and beyond that to the scurrying, rodent-like life of the tiny mammals, and far remoter still, the shore-slime of the primeval sea.” 70 In Jekyll and Hyde, too, Hyde first exists in the daydreams of Jekyll, and Vandover at one point, close to hysteria, seems to gain a glimpse into the memory of all times – both Hyde, (homophone of hide,) and the wolf into which Vandover turns, are the prehistoric other.71 The dual-characters Jekyll-Hyde, Vandover-Brute, and Narrator-Big-Tooth fail at a synthesis; through the attempt of suppression or persistent segregation, they constantly feed and are finally swallowed by the residues of an evolutionary past. The state of dissociation between individual and collective psyche is maintained if not sharpened. Thus, as suggested by Jungian psychology, the modern self continues to find itself opposite a kind of ‘Negro culture’ (Negerkultur), opposite a primitive state. This is in Jung’s appraisal the general state of his culture – a thin crust of civilization covering a dark-skinned beast (eine dunkelhäutige Bestie).72 In the end, the atavistic sides of the literary personalities win, and the regression in the psychoanalytic sense is complete; at this point, one may recognize the condition of the neurotic who is unable to see the archetypes of the unconscious as psychic realities and instead takes them for concrete realities. In other words, the individual identifies with aspects of the common prehistoric past of humankind. 73 This is where the dream comes in. Jung ascribed Freud the primacy of the insight that the analysis of dreams was the most important method for unraveling the unconscious. 74 London’s narrator calls himself an “atavistic nightmare”, which links the notion of atavism with that of dreams as windows into prehistory – which open particularly wide for him, because his psyche or his brain is atavistic.75 In psychoanalysis, the dream, the myth, and other manifestations of the other in the self gained remedial impact in the healing process of the split human being. The wordless events that might be awakened through a regression into pre-infantile time have their origin in the lives, sufferings, and joys of our ancestors, and they must be idiosyncratically reembodied by the individual. Due to their inherent contradiction to the conscious, they cannot, however, be translated immediately – the dream may function as mediator between the reality of the unconscious and that of the conscious.76 Through the interpretation of dreams, an intense dialogue between unconscious and conscious is initiated. The analysis of dreams fulfils a transcendental function that might enable a personality to realize his or her original potential for wholeness, completeness, and perfection (Individuationsprozess).77

69 70 71 72 73 74 75 76 77

London, 2000 (1906-1907), p. 15. London, 2000 (1906-1907), p. 139. Norris, 1986 (1914), p. 162. Jung, 1971 (1916) pp. 97-98, see also pp. 74-75. Jung, 1971 (1916), p. 97. Jung thought of archetypes as having lives of their own (Partialseelen, pp. 6768). Jung, 1971 (1916), p. 24. On Jung’s reception of Freud see also Jung, 1972 (1906-1916). London, 2000 (1906-1907), “this atavistic brain of mine”, p. 21, also p. 20. Jung, 1971 (1916), pp. 81, 103. Jung, 1971 (1916), pp. 99, 110-111.

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In Before Adam as well as in other writings of London and his contemporaries such as Norris and Stevenson, the problems associated with the duality of human nature were expounded, and ways were sought to overcome opposing demands in the authors’ own lives. 78 The potion, alcohol, madness appear as unsuccessful remedies. In the dream, however, a solution was foreshadowed that would find full expression in Freud’s and Jung’s theories. London immediately recognized the new psychology as the means for synthesizing the racial heritage with the modern self he had been searching for. In the child, the criminal, the insane, the primitive, ‘the savage’, and the atavistic reversion to such states, the selfish animal instincts were set loose. But the same ruthless instincts drove the average Western male on to conquer ever new coasts – to return to London’s metaphor. However, in civilized man, our primordial nature was in conflict with the higher social and moral instincts. In the perfect case, the constructive sides of our animal origin were brought to use, while the destructive aspects were subdued by the moral sense. London ascribed dreams a synthesizing function in the individual’s attempt to gain such a balance. They were atavistic devices in the understanding of the individual as part of the community. Like London’s books, they were memory aids; they enabled an understanding of self as part of the prehistory of the race.

Marianne Sommer, Eidgenössische Technische Hochschule, Zürich [email protected]

78

On a possible influence of Jung on the earlier fiction of London, such as The Call of the Wild (1903), Wild Fang (1906), and Before Adam, see Crow, 1966; on such an influence in the late South Pacific fiction see Reesman, 1988; on Jungian psychology in “The Red One” (1918) see Berkove, 1966. West, 1998, identifies several unresolved contradictions against which Norris struggled – such as effete east and rough west, artist and businessman, romance and realism, nineteenth and twentieth century, morality and strength, egalitarianism and racial suprematism, communism and capitalism, sensualism and intellectualism, primitive and civilized, determinism and freedom – and connects these to the unconscious/conscious dualism. In fact, both Norris and London suffered from similar paradoxes that marked their generation as a whole (on paradoxical stances in the racial question see for example Furer, 1966).

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Introduction Late nineteenth–century anthropological research was to a large extent preoccupied with questions of inheritance, but it was so with regard to two sets of phenomena. On the one hand, cultural and physical differences among humans were subjected to description and comparison. Inheritance was of interest here, as the present distribution of the varieties of mankind was interpreted as bearing some relation to the stages and bifurcations through which human evolution had occurred in the past. On the other hand, societies other than contemporary European ones were scrutinized for the institutions that made them work. Classification of kinship and organization into clans, moieties, castes, etc., in short: the structures along which names, social status, and property were inherited raised the interest of anthropologists here for their relation to the temporal stability of the social order. From historical literature on late nineteenth–century anthropology one could get the impression that it was the first aspect, that of racial descent, which dominated research agendas. Yet some of the most influential works of the period were dedicated to what we today would call questions of social anthropology: Johann Jakob Bachofen (1815–1887), Swiss lawyer and philologist, argued in Das Mutterrecht (1861) for a primeval stage of promiscuity, followed by gynæcocracy, which again, though only in select cultures, yielded to patriarchy. Henry James Sumner Maine (1882–1888) directed attention to the complex relations between kinship and territorial bonds in his Ancient Law (1861), and John F. McLennan (1827–1881) coined the analytical categories “exogamy” and “endogamy” in his first major essay Primitive Marriage (1865). Like Bachofen, both Maine and McLennan were lawyers. Edward B. Tylor in his Researches into the Early History of Mankind and the Development of Civilization (1865), and Henry Lubbock, in his The Origin of Civilisation and the Primitive Condition of Man (1870) laid the foundations for understanding the interdependence of cross–cousin marriage and exogamy, mother–in–law avoidance and matrilocal residence. Finally, Lewis H. Morgan, again a lawyer by training, laid bare the intricacies of kinship terminologies world wide in his Systems of Consanguinity and Affinity of the Human Family (1871) and demonstrated their correlation with forms of marriage and rules of descent, among other things drawing the distinction between descriptive and classificatory kinship systems. Through their reception by Charles Darwin, in his The Descent of Man (1871),1 as well as by Friedrich Engels in Der Ursprung der Familie, des Privateigentums, und des Staates (1884),2 all these works fed into the main–stream ideologies of late nineteenth–century, no less so than evolutionary anthropology did. Was there a common ground for this twofold preoccupation with inheritance? At first sight it may well seem so: Descent, affinity, and genealogy seem to afford a common, and what is more, 1 2

Kuper (1997). Krader (1973): ch. 2.

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universally shared metaphorical framework for the study of both racial descent and kinship. And indeed, as George Stocking has pointed out: Before the twentieth century, “‘[b]lood’ —and by extension ‘race’— included numerous elements that we today would call cultural; there was not a clear line between cultural and physical elements or between social and biological heredity.” 3 Modern European racial discourse, as argued by Renato Mazzolini, had itself grown out of the extension of a caste system of European making with corresponding interdictions and regulations regarding marriage and inheritance.4 Eugenicists and racial hygienists relied on what may clearly be addressed as rules of exogamy (avoidance of incest) and endogamy (segregation by race and class, exclusion of the ‘unfit’) in their programs for social reform. 5 The study of race and kinship seem to be nothing but the two sides of one and the same coin, a widespread, and deeply rooted fear of the consequences of promiscuity and incest, and a corresponding obsession with the rationalization of even the most intimate human relations. As soon as one tries to move beyond analogies, the relation of kinship and racial descent becomes curiously unclear however. The main reason for this is, that the categories used to analyze kinship and racial descent respectively do not map onto each other. Racial categories, in the nineteenth century at least, obeyed the logic of the Linnaean hierarchy, constituting a system of mutually exclusive classes contained within more extensive and again mutually exclusive classes. Ideally, an explanation of their formation would lead back to one or several singular origins. The history of racial descent is one of progressive differentiation, usually represented as the growth of a tree-like structure. Kinship, on the other hand, employs categories that overlap systematically: one individual can be a daughter, a mother, an aunt, a cousin at once, and these categories do not form a hierarchy of nested classes. Their explanation, in the case of humans at least, does not lead back to singular origins, but to bonds of marriage. Reducing kinship to a singular “ancestor” is the task of genealogy, quite a different matter, and one that systematically has to discard certain kin relations. Finally, the history of kinship is one of alliances and separations, of a continually woven ‘fabric […] in which warp and filling yarn correspond to localities and tribes’, as Claude Lévi– Strauss formulated it.6 Darwin, famously, looked for sexual selection or “long-continued intercrossing” as a means to explain the “protean or polymorphic” nature of mankind. 7 My aim in this paper is largely descriptive. I will look at the way in which the discourses of racial descent and kinship related to each other in a selection of texts, with particular emphasis on Morgan’s Systems of Consanguinity and Affinity of the Human Family (1871) and Franz Boas’s (1858–1942) anthropological papers from the late nineteenth century. In particular, I will be interested in how the analysis of kinship, establishing definite relations among individuals, was used by anthropologists of the late nineteenth century as a tool to reach a complete analysis of the varieties of mankind – the “races”, “families”, or “nations” constituting mankind.

3 4 5 6 7

Stocking (1994): p. 6. Mazzolini (in press). See, e.g.: Richardson (2002). Lévi–Strauss (1962) (cited from German edition, Frankfurt a. M. 1969: p. 97). Darwin (1871): p. 550; cf. Endersby (2003).

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Morgan and the analysis of kinship When Lewis Henry Morgan carried out his first interviews with Iroquois informants in Albany 1844, his immediate motive for understanding the workings of their social life was a curious one: He had joined a secret fraternal organization, the Grand Order of Iroquois, and wanted to duplicate the structure of Iroquois society in this organization. After the renewal of his ethnographic interests in 1857, he began to collect more extensively, by setting up a circular containing a recent paper on the ‘Laws of Descent of the Iroquois’ and a set of questions regarding kinship and clan organization, in particular a long list of questions asking for the counterpart of kin terms to the fifth collateral degree in other languages. Morgan had set up this list of questions or “schedule”, as he also called it, with “considerable labor” to be “sufficiently full to describe every known relationship, and yet arranged upon such a method as to be simple and intelligible.” 8 Morgan called his schedule a “new instrument in ethnology.”9 According to the introductory chapter to Systems of Consanguinity and Affinity, he first sent it out, in printed form, to “the several Indian missions in the United Strates, to the commanders of the several military posts in the Indian country, and to the government Indian agents.” Unsatisfied with the answers he thus received –one district officer had answered to Morgan’s questionnaire of the “customs and manners” of natives: “Manners none, customs beastly”–10 Morgan began to undertake “annual explorations among the Indian nations” himself to gather kinship terms. To get at similar information outside of North America, Morgan addressed both the Smithsonian Institution and the U.S. government for assistance. Assistance was granted, and Morgan could rely on a worldwide network of diplomatic staff, missionaries, and scientists to procure relevant data for him.11 What the questions contained in the schedule must have looked like – a copy is preserved in the Smithsonian collection,12 but I have not had a chance to have a look at it– can be derived from the tabular presentation of results in Morgan’s Systems. The data is presented for the “Seneca– Iroquois” and the “Tamil People of South Asia”, both supposedly representative of what Morgan called the “classificatory system of relationship”, i. e. a system that classifies collateral cousins of various degrees together with siblings under one and the same term. The respective Iroquois and Tamil kin terms are tabulated against their translations into English and a long list of terms headed “Description of persons.” Despite its affinities with what Morgan called, in a sweeping generalization, the “descriptive systems” of relationship among the Aryan and Semitic languages, which address each relative by a definite, mostly composite term, the latter “descriptions of persons” do not stem from any natural language. Morgan himself called them “numerical in character”, “resting upon an ordinance of nature,” and both “universal and unchangeable.” 13 The principle they build upon is simple: each possible kin relation between an assumed “ego” and another person, up to the fifth collateral degree, is expressed in a combination of primitive terms:

8 9 10 11 12 13

Tooker (1992). Morgan (1871): p.9. Fox (1877): p. xxxii. Tooker (1992): p. 9. Ibid. Morgan (1877): p. 11.

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“mother”, “father” “son”, “daughter”, “sister”, “brother”, and, for affine relatives, “husband” and “wife”. The list of terms under “Description of person” reaches from simple terms as “my son” to complex ones like “my mother’s mother’s mother’s brother’s son’s son’s son.” All in all, the table lists 237 terms like that. What is the purpose of this exercise? The introductory chapter to Morgan’s Systems of Consanguinty and Affinity contains the following discussion of the difficulties Morgan encountered when asking correspondents to answer his questionnaire: A large number of schedules, when returned, where found to be imperfectly filled out. Misapprehension of the nature and object of the investigation was the principal cause. The most usual form of mistake was the translation of the questions into the native language which simply reproduced the questions and left them unanswered. […]. As our own system is descriptive essentially, a correct answer to most of the questions would describe a person very much in the form of the question itself, if the system of the nation was descriptive. But, on the contrary, if it was classificatory, such answers would not only be incorrect in fact, but fail to show the system. […]. Every system of relationship is intrinsically difficult until it has been carefully studied. […]. It is easy, therefore, to perceive that when a person was requested to work out, in detail, the system of a foreign people he would find it necessary, in the first instance, to master his own, and after that to meet and overcome another, and perhaps, radically different form.14

These observations are highly revealing. They demonstrate that the terms contained under the rubrique “Description of the Persons” are not merely descriptive, but in fact provide a complete analysis of kinship. They break down kinship into each and every possible kin relation, thus serving the translation of terms between different systems by defining their respective extensions. Without this device, the systems of “foreign people” would remain largely opaque to their observers, and, vice versa, informants would be unable to give “correct” answers to the questions posed to them. A peculiar property of the schedule betrays its analytic character: It is, in Morgan’s words “necessarily self-corrective […], since the position of Ego and his or her correlative person is reversed in different questions.” Morgan himself, it is true, systematically blurred the distinction between the schedule of his own making and the “descriptive systems” he saw at work in the Aryan and Semitic language families. His contemporary James McLennan saw that already, and in Ancient Society Morgan felt himself forced too meet McLennan’s criticisms in a 12 page long “note”. 15 But analyticity remains a decisive character of his schedule, one that distinguishes it from kinship terminologies used in everyday contexts.16 It betrays the extent to which his concerns were not philological in the first place. The preface to Systems refers to “philology”, but as a mere “instrument for the classification of nations into families.” And adds, that “[i]t was with special reference to the bearing which the systems of consanguinity and affinity of the several families of mankind might have upon this vital question, that the research, the results of which are ascertained in this volume, was undertaken.” 17 14 15 16

Ibid.: p. 6. Morgan (1877): pp. 509–521. Tooker (1992, p. 9) emphasizes that there is no vernacular kinship terminology that does not provide the possibility to describe each relative by terms as the ones employed in Morgan’s schedule and adds: “Morgan’s schedule rests on this fact.”

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Systems of consanguinity, Morgan believed, were particularly well–suited for the task of classifying humankind: “The family relationships are as ancient as the family. They exist in virtue of the law of derivation, which is expressed by the perpetuation of the species through the marriage relation. A system of consanguinity, which is founded upon a community of blood, is but the formal expression and recognition of these relationships.”18 And in his 1877 Ancient Society he should fomulate: “[Modern institutions] have had a lineal descent through the ages, with the streams of the blood, as well as a logical development.”19 To Morgan the analysis of kinship reveals, one might say, the organization of mankind, and differences in that organization immediately reflect the different stages and branches of its evolution.

Boas and the statistics of culture George Stocking has assigned Boas the role of a founding father of the “modern anthropological culture concept” characterized by “historicity, plurality, behavioral determinism, integration, and relativism.”20 On the other hand, Stocking has portrayed Boas’s work in physical anthropology as instrumental in the “passing of a romantic conception of race – of the ideas of racial ‘essence,’ of racial ‘genius,’ of racial ‘soul,’ of race as a supraindividual organic identity”. In particular it was Boas’s statistical approach that was, as Stocking put it, “subversive of traditional racial assumptions.”21 Boas, as is probably too little known, was an ardent practitioner of physical anthropology and biometry, designing instruments for craniometric measurements, 22 carrying out large–scale anthropometric studies on native populations of North–America, 23 and creating mathematical tools for the study of correlation.24 In the following, I want to argue that Boas advocated a similar statistical approach in his studies of primitive culture as well, and that kinship was integral to that approach. In 1887, Boas became involved in a debate about museum displays. 25 Otis Tufton Mason (1838–1908), curator of ethnology at the Smithsonian institution, had suggested to arrange ethnological displays at the United States National Museum according to a classification of the objects displayed: exemplars of different kinds of artifacts were arranged in series, each representing a stage in the evolution of its kind; the rationale on which this presentation rested was borrowed from natural history. As Boas quoted Mason: [Human inventions] may be divided into families, genera, and species. They may be studied in their several ontogenies (that is we may watch the unfolding of each individual thing from its raw material to its finished production). They may be regarded as the products of specific evolution out of natural objects serving human wants and up to the most delicate machine performing the same function. They may be modified by their relationship, one to another, 17 18 19 20 21 22 23 24 25

Morgan (1877): p. v–vi. Ibid.: p. 10. Ibid: p. 4. Stocking (1983a): p. 230. Stocking (1983b): pp. 192–194. Boas (1890). Boas (1891b). Boas (1894). Jacknis (1985); Jenkins (1994).

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in sets, outfits, apparatus, just as the insect and flower are co–ordinately transformed. They observe the law of change under environment and geographical distribution.26

The alternative Boas proposed was to arrange collections “according to tribes, in order to teach the peculiar style of each group.” The reasons he adduced for this position were epistemological: In regarding the technological phenomenon as a biological specimen, and trying to classify it, [Mason] introduces the rigid abstractions species, genus, and family into ethnology, the true meaning of which it took so long to understand. It is only since the development of the evolutional [sic] theory that it became clear that the object of study is the individual, not abstractions from the individual under observation. We have to study each ethnological specimen individually in its history and in its medium […]. Our objection to Mason’s idea is, that classification is not explanation.27

This seems to be a strange reasoning: First of all, “studying each ethnological specimen individually in its history and in its medium” would, taken literally, be an endless task, and both Mason as well as other participants in the debate pointed out the practical difficulties that an arrangement by tribes would imply.28 Secondly, an arrangement according to tribes seems to involve as much classification as that proposed by Mason, namely a classification by “tribes.” Also this criticism was raised in the debate,29 and the point has only recently been reiterated by a post– modern epigone.30 To this, Boas only had a short, categorical reply: “Such groups [i.e. tribes, and groups of tribes] are not at all intended to be classifications.”31 Boas’s studies of native myths along the North–Pacific coast can serve as an example to elucidate what he had in mind with this strange assertion. In these studies, Boas broke down the myths into constituent “elements” and recorded their distribution within a group of geographically contiguous “tribes.” “We can in this manner,” as Boas explained, “trace what we might call a dwindling down of an elaborate cyclus of myths to mere adventures, or even to incidents of adventures, and we can follow the process step by step.” 32 In more detail, he described this method as follows: If we have a full collection of the tales and myths of all the tribes of a certain region, and then tabulate the number of incidents which all the collections from each tribe have in common with any selected tribe, the number of common incidents will be larger the more intimate the relation of the two tribes and the nearer they live together. This is what we observe in a tabulation of the material collected at the North Pacific Coast. On the whole, the nearer the people, the greater the number of common elements; the farther apart, the less the number.33

26 27 28 29 30 31 32 33

Quoted according to Franz Boas (1887b): p. 485. Boas does not give a reference, and I have not been able to identify his source. Ibid. Dall (1887): p. 587; Powell (1887). Powell (1887). Jenkins (1994). Boas (1887a): p.614. Boas (1896a): p. 2. This paper is a summary of Boas (1895a). Ibid.: p. 3.

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It should be noted, that it is the unequal distribution of elements of myths that defines them as their constituent elements in the first place. Such a “statistical inquiry”, as Boas called it, 34 rested on a “fundamental condition”, which “differentiates our method from other investigators […], who see a proof of dissemination or even blood relationship in each similarity that is found between a certain tribe and any other tribe of the globe” – namely, that the material, on which it was based, was “collected in contiguous areas.”35 This contiguity was not necessarily, Boas emphasized, a geographical one, and it is here were marriage, kinship, and social structure enter the picture. “The social customs of the Kwakiutl,” the ethnic group most intensely studied by Boas during several field trips, are, as he maintains, “based entirely upon the division into clans and the ranking of each individual is the higher –at least to a certain extent– the more important the legend of the clan.” Moreover, “the customs of the tribe are such that by means of a marriage the young husband acquires the clan legends of his wife, and the warrior who slays an enemy those of the person whom he has slain. By this means a large number of traditions of the neighboring tribes have been incorporated in the mythology of the Kwakiutl.”36 It is by relating the distribution of mythical elements to a space whose contiguity can be ascertained in terms of social relations among individuals –alliances as well as antagonisms– that Boas wants to circumvent the pitfalls of analogical reasoning. In the analysis of these relations, Boas clearly relied on the categories that others before him, most prominently Morgan, had developed to account for social structure in terms of kinship. I cannot discuss this in detail here, but let me state, that the clan system that Boas detected among the Kwakiutl was actually more complex than he described it in the quote I gave you above: through marriage it was not the husband personally who acquired the clan status of his wife, but he acquired it “for his son.” 37 Ironically, the grand picture that Boas came up with on the basis of such studies was strongly opposed to that of evolutionists like Morgan: A great many [...] important legends prove to be of foreign origin, being grafted upon mythologies of various tribes. This being the case, I draw the conclusion that the mythologies of the various tribes as we can find them now are not organic growths, but have gradually developed and obtained their present form by accretion of foreign material. Much of this material must have been adopted ready–made […]. We are, therefore, led to the conclusion that from mythologies in their present form it is impossible to derive the conclusion that they are mythological explanations of phenomena of nature […], but that many of them, at the place where we find them now, never had such a meaning. If we acknowledge this conclusion as correct, we must […] admit that, also, explanations given by the Indians themselves are often secondary, and do not reflect the true origin of the myths.38

34 35 36 37 38

Ibid. Ibid.: p. 6. Ibid.: p. 8–9. See Boas (1895b): pp. 334–335. Ibid.: p. 5.

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Conclusion Boas rejection of an “organic growth” of culture, and the model of a “complex growth” 39 by accretion which he advocated instead reveals that his famous environmentalism was far from being Lamarckian in character. The elements of culture are transmitted without bearing the mark of the context in which they originated, and their combination is a matter of historical accident, not of influence or the weight of tradition. The similarities that this model exhibits with respect to the distinction that biologists and biometricians began to draw between the transmission of germinal elements and the development of somatic characters is remarkable. In 1901 Boas wrote: “[… T]he development of culture must not be confounded with the development of mind. Culture […] shows the cumulative effects of the activities of many minds. But it is not an expression of the organization of the minds constituting the community.”40 And when Johannsen canonized the distinction of genotype and phenotype in 1911, Boas whole–heartedly endorsed it. 41 Boas cultural relativism did not develop in opposition to rival claims from physical anthropologists in the explanation of cultural phenomena. His ambition was to base the whole of anthropology – somatology, ethnology, and linguistics alike– on an inductive basis that proceeded not by analogies but by connecting individual phenomena with the help of statistical procedures. 42 Is there something to be said about the motivation to do this? George Stocking has identified the central concern of Boas, related to the Kulturkampf of his native country, with the “search for ‘the psychological origin of the implicit belief in the authority of tradition.’” 43 For Morgan, tradition had been largely inescapable: “[T]he primary institutions of a people are necessarily permanent from age to age […]. It is only by the entire and absolute transmutation of a race from the hunter to the civilized condition, that such institutions can be eradicated.” Likewise Boas admitted, that “[e]ach individual must be influenced to a greater or less extent by the mass of traditional material present in his mind.” But to Boas this influence was not inescapable: “There is an undoubted tendency in the advance of civilization to eliminate traditional elements, and to gain a clearer and clearer insight into the hypothetical basis of our reasoning.” The method to achieve this was “to carry the analysis of any given phenomenon to completion.” I have tried to show that the analysis of kinship, as instituted by Morgan, served this aim, and that it actually succeeded in revealing and overcoming cultural preconceptions. And it is telling, in this respect, that Morgan took “descriptive” systems of kinship terminology to be the hallmark of “civic” societies as opposed to “gentile” ones, that were “founded upon relations purely personal.” 44

Staffan Müller- Wille, Research Center for Genomics in Society, University of Exeter, S.E.W.Mueller–[email protected] 39 40 41 42 43 44

Boas (1891a): p. 20. Boas (1901). Boas (1911). Boas (1896b). Stocking (1992): p. 97. Morgan (1877): pp. 393–395.

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References Boas, Franz. 1887a. ‘Museums of Ethnology and their classification’, Science 9(229): 614. ———. 1887b. ‘The occurrence of similar inventions in areas widely apart’, Science 9(224): 485– 486. ———. 1890. ‘A Modification of Broca's Stereograph’, American Anthropologist 3(3): 292–293. ———. 1891a. ‘Dissemination of Tales among the Natives of North America’, Journal of American Folklore, 4(12): 13 – 21. ———. 1891b. ‘Physical Characteristics of the Indians of the North Pacific Coast’, American Anthropologist 4(1): 25–32. ———. 1894. ‘The Correlation of Anatomical or Physiological Measurements’, American Anthropologist 7(3): 313–324. ———. 1895a. Indianische Sagen von der Nord–Pacifischen Küste Amerikas, Berlin: A. Asher. ———. 1895b. The Social Organization and the Secret Societies of the Kwakiutl Indians, Washington: Government Print Office. ———. 1896a. ‘The Growth of Indian Mythologies. A Study Based upon the Growth of the Mythologies of the North Pacific’, Journal of American Folklore 9(32), 1 –11. ———. 1896b. ‘The Limitations of the Comparative Method of Anthropology’, Science. New Series, 4(103): 901–908. ———. 1901. ‘The mind of primitive man’, Journal of American Folklore, 14(52): 1–11. ———. 1911. The mind of primitive man; a course of lectures delivered before the Lowell Institute, Boston, Mass., and the National University of Mexico, 1910– 1911, Boston: MacMillan. Dall, William H. 1887 ‘Museums of Ethnology and their classification’, Science 9(228): 587 – 589. Darwin, Charles. [1871] 1989. The Descent of Man and Selection in Relation to Sex, New York: Modern Library Reprint. Endersby, Jim. 2003. ‘Darwin on generation, pangenesis, and sexual selection’, in: Cambridge Companion to Darwin, ed. by J. Hodge & G. Radick, Cambridge, Engl.: 69–91. Fox, Robin. 1877. ‘Introduction’, in: Morgan (1877). Jacknis, Ira. 1985. ‘Franz Boas and exhibits: On the limitations of the museum method of anthropology’, in Objects and others: Essays on museums and material culture, ed. by George W. Stocking, Jr. Madison: 75– 111 Jenkins, David. 1994. ‘Object lessons and ethnographic displays: Museum exhibitions and the making of American anthropology’, Comparative Studies in Society and History 36: 242–270. Krader, Lawrence. 1973. Ethnologie und Anthropologie bei Marx, München. Kuper, Adam. 1997. ‘On human nature: Darwin and the anthropologists’, in Nature and society in historical context, ed. by M. Teich et al, Cambridge, Engl.: 274–290. Lévi–Strauss, Claude. 1962. Le Totemisme aujourd´hui. Paris: PUF. Morgan, Lewis H. 1877. Ancient Society, New York: Henry Holt. ———. 1871. Systems of Consanguinity and Affinity of the Human Family, City of Washington (Smithsonian Contributions to Knowledge, vol. xvii). Powell, J. W. 1887. ‘Museums of Ethnology and their classification’, Science 9(229): 612–613. Richardson, Angelique. 2002. Love and Eugenics in the Late Nineteenth Century: Rational Reproduction and the New Women, Oxford: Oxford University Press. Stocking, George. 1983. Race, Culture, and Evolution: Essays in the History of Anthropology, Chicago: University of Chicago Press. ———. 1983a. ‘Franz Boas and the culture concept’, in: Stocking (1983): 195– 234. ———. 1983b. ‘The critique of racial formalism’, in: Stocking (1983): 161–194. ———. 1992. ‘Anthropology as Kulturkampf. Science and politics in the career of Franz Boas’, in: George Stocking, The Ethnographer’s Magic and Other Essays in the History of Anthropology, Madison: 92 – 113. ———. 1994. ‘The turn–of –the–century concept of race’, in: Modernity/Modernism 1: 4– 16. Tooker, Elisabeth. 1992. ‘Lewis H. Morgan and His Contemporaries’, American Anthropologist 94: 357– 375.

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M a x P la n c k I n s titu te fo r th e His to r y o f Sc ie n c e

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Co nf e re nc e

A Cultural History of Heredity IV: Heredity in the Century of the Gene

Table of Contents

Introduction Staffan Müller-Wille, Hans-Jörg Rheinberger and John Dupré

3

Leaving Inheritance behind: Wilhelm Johannsen and the Politics of Mendelism Staffan Müller-Wille

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Mendelian Factors and Human Disease: A Conversation Jean Paul Gaudillière & Ilana Löwy

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Heredity without Mendelism: Theory and Practice of Dairy Cattle Breeding in the Netherlands 1900-1950 Bert Theunissen

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Innovation and Ownership in Living Products: Animals and Fruits in the United States, the 1870s to 1930 Daniel J. Kevles

51

Coalition and Opposition: Heredity, Culture, and the Boundaries of Anthropology in the Work of Alfred L. Kroeber Maria E. Kronfeldner

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Comments on Daniel Kevles and Maria Kronfeldner’s Papers Edna Suárez

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Producing Identity, Industrializing Purity: Elements for a Cultural History of Genetics Christophe Bonneuil

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Mendelism and Agriculture in the First Decades of the XXth Century in Mexico Ana Barahona

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Herbert Spencer Jennings, Heredity, and Protozoa as Model Organisms, 1908-1918 Judy Johns Schloegel

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Clones, Pure Lines, and Heredity: The Work of Victor Jollos Christina Brandt

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Pedigree vs. Mendelism. Concepts of Heredity in Psychiatry before and after 1900 Bernd Gausemeier

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Pedigree Charts as Tools to Visualize Inherited Disease in Progressive Era America Philip Wilson

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Biohistorical Narratives of Jewish history. Contextualizing the Studies of Wilhelm Nussbaum (1896-1985) Veronika Lipphardt

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William Bateson’s Pre- and Post-Mendelian Research Program in ‘Heredity and Development’ Marsha L. Richmond

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Genetics Without Genes: Blakeslee, Datura, and “Chromosomal Mutations” Luis Campos

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Seeing, Breeding and the Organisation of Variation: Erwin Baur and the Culture of Mutations in the 1920s Alexander von Schwerin

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Heredity and the Century of the Gene Raphael Falk

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Introduction This volume contains contributions to a workshop that was the fourth in a series of workshops dedicated to the objects, the cultural practices and the institutions in which the knowledge of heredity became materially entrenched and in which it unfolded its effects in various epochs and social arenas.1 It was organized collaboratively by the Max Planck Institute for the History of Science, Berlin, and the ESRC Research Centre for Genomics in Society, Exeter. Funds from the Academic Research Collaboration programme of the British Council and the German Academic Exchange Service allowed to prepare the workshop in two one-day-meetings of scholars from Berlin and Exeter. The workshop itself was funded by the British Academy and by the Government of the Principality of Liechtenstein. The last workshop in the series had dealt with the period up until the very end of the nineteenth century when heredity had become a central problem for biologists and a wide variety of approaches to attack that problem had begun to flourish.2 The fourth workshop was specifically designed to address a historiographic problem that had arisen within the wider context of our project ‘A Cultural History of Heredity’. Up to the late nineteenth century, the knowledge of heredity took shape by a step-by-step aggregation and integration of discourses from various knowledge domains; while from 1900 onwards it began to condense into and to be shaped by a highly-specialized discipline, the discipline of genetics. This has resulted in a preoccupation of the historical literature with genetics. One of the basic assumptions of our project, however, has been that heredity was always and remained to be more than genetics as a discipline, and that wider notions of inheritance persisted in areas like practical breeding, medical counselling and therapy, eugenics, and anthropology (including cultural anthropology). To widen the scope of inquiry, we therefore decided to focus the workshop on the tools of dealing with inheritance, that is genealogical records and model organisms, and to follow their provenances, metamorphoses, and trajectories.3 The major results, as documented in this volume, can be summarized as follows. 1. During the era of classical genetics (1900-1940) a number of important concepts with respect to heredity were defined, or re-defined, in strictly genealogical terms. These concepts included: clones, pure lines, bloodlines (in animal breeding), family lines (in anthropology), generation, and mutations as instances of change in these lines. The operational definition of these terms allowed researchers like William Bateson, Wilhelm Johannsen, Erwin Baur, Victor Jollos, Herbert Spencer Jennings, and Albert F. Blakeslee to create what some of them referred to as ‘synthetic species’: breeding systems fine-tuned by systematic in- and out-breeding to instantiate particular cases of evolution, as Bateson once put it. The significance of these constructs for the cultural history of heredity is twofold. First, they all had a life prior to and outside biological laboratories or experimental gardens. Pure lines, for example, had been developed by plant breeders, especially 1 2 3

For more information on the project see http://www.mpiwg-berlin.mpg.de/en/research/projects/ DeptIII_Cultural_History_Heredity/index_html. See Conference: A Cultural History of Heredity III: 19th and Early 20th Centuries, Berlin: Max Planck Institute for the History of Science (Preprint, vol. 294). See original call for papers at http://www.mpiwg-berlin.mpg.de/workshops/en/HEREDITY/ announcement4.html, 2008/01/07

3

Introduction

the French seed company Vilmorin, in the late nineteenth century already. And genealogical concepts used in the analysis of human populations had an even longer prehistory in clinical records. Second, the construction of ‘synthetic species’ was largely independent of any specific theoretical assumptions about mechanisms of transmission. Clones of single-celled organisms like Paramecium, for example, could be regarded as ‘naked germ-lines’, thus opening the possibility to study germinal transmission in its own right, without any prior commitment to particular theoretical assumptions about the relationship of soma and germ-plasm. And mutation researchers like Blakeslee or Baur could rely on their systems producing novelties without having to make prior decisions about the nature of mutations. It was in this sense that Jennings insisted against Johannsen that genotypes had to be considered as ‘things’, rather than hypothetical entities. 2. Much research into heredity in the late nineteenth and early twentieth century took place in applied contexts like seed production, breeding yeast and cereals for large-scale beer production, mass-production of vaccines, efforts to further public health, or administration of psychiatric hospitals. Increasing levels of division of labour and bureaucratic control in these areas – the seed company Vilmorin in France had 400 employees around 1900 – led to the establishment of a culture of expertise and scientificity. In these contexts, however, Mendelism featured as only one among many methodologies to realize values that were endorsed by this culture, like analyticity, exactitude, calculability and predictability. Breeders and eugenicists in particular, whether they declared themselves Mendelists or not, shared a combinatorial approach that held a promise for the transparent and reliable production of intergenerational effects. Synthetic chemistry, not physics, provided the model science in this context. 3. An important property of this culture of expertise was its obsession with purity. Purity connects a number of issues that were at stake. It was an instrument of control, as results could be ‘checked’ against the corresponding inputs. It enabled practitioners to ‘fix’ characters and create identifiable and specifiable products. It created a set of discrete and stable life forms, rather than an uncontrolled continuum of variations. And it held a promise to divorce practices from the vagaries of history. Once entities could be held ‘pure’, they could be recombined without being subject to the unpredictable manifold of interactions that ‘impure’ entities like the so-called ‘land races’ in traditional agriculture elicited. Heritability rather than inheritance, prospect rather than retrospect thus became one of the chief criteria for assessing the quality of life forms. In order to advertise, trade-mark, or patent agricultural or microbiological innovations, production methods had to be made transparent and reliable reproduction guaranteed. Heredity was commodified to become heritability, a marketable quality. 4. Mendelism entailed conditions and costs that precluded many areas from adopting it. To do Mendelian experiments, organisms had to be first inbred, then cross-bred, and finally raised in large numbers, to be able to ascertain Mendelian ratios. Asexual organisms and humans, but also many agriculturally significant animals, like cows, could not be subjected to such a practice. This is one of the main reasons why animal breeding and clinical medicine became ‘geneticized’ only well after WWII, and why statistical approaches, developed by the so-called biometrical school

4

Introduction

long before the advent of Mendelism already, persisted in these areas to finally merge with population and quantitative genetics. It was with respect to human populations, in psychiatry, medicine and anthropology, in particular that sophisticated genealogical and statistical techniques –trait pedigrees of various cut, statistical and combinatory tables – were developed and applied to populations by researchers like Wilhelm Weinberg, Ernst Rüdin, or Wilhelm Nussbaum. Originating in administrative record-keeping practices of mental asylums in the late nineteenth century, these techniques retained their bureaucratic character, with the result that key categories and concepts, like ‘race’ or ‘heritability’, were emptied of their biological content and became formal, purely classificatory or statistical notions, although constant slippage from statistical results to presumed ‘genetic’ and thus biological (as opposed to ‘environmental’ or ‘epigenetic’) causes occurred regularly. Such slippages could be productive in terms of posing new research problems, but they were also mobilized to justify oppressive and outright murderous bio-policies. 5. The era of classical genetics was marked by a close, yet conflict-ridden relationship of heredity and history. Prominent biologists like Wilhelm Johannsen saw Mendelism as a way to free technology and society from the weight of tradition. Mendelism’s reductionist view of the organism as composed of modular and largely independent, to some degree even autonomous entities, was prefigured by the debates about cell theory in the nineteenth century and resonated with an industrial culture that placed value on the specificity and reproducibility of innovations. If one were able to atomize life to the degree that its elements would not be affected substantially by the combinations they entered in the course of history, then there would be virtually no limit to the future production of innovations through combination. The future could be made, or constructed, eliminating the power that history and tradition used to have over life. It was in this sense that Alfred L. Kroeber spoke of culture as the ‘superorganic’: Weismann’s separation of soma and germ-line also, in a way, divorced culture from the organic, leaving culture behind as subject to its own ‘systems of inheritance’, systems that varied from culture to culture, and could be studied in their own right by anthropologists. But histories of nations and peoples were also recast as ‘bio-historical narratives’ whose dramatic turning points consisted in race mixtures, migrations, phases of strong selection or isolation, resumed intermarriage, in short: events entirely human, and not enforced by some higher law of history. Knowledge of heredity had evolved into an instrument, not only to analyze the past, but also to shape the future. Staffan Müller-Wille (University of Exeter) Hans-Jörg Rheinberger (Max-Planck-Institute for the History of Science) John Dupré (University of Exeter)

5

Leaving Inheritance behind: Wilhelm Johannsen and the Politics of Mendelism Staffan Müller-Wille

Abstract In 1912 Wilhelm Johannsen codified the distinction of genotype and phenotype to distinguish a space of heredity with an independent logic and metrics from another, physiological and developmental space represented by the cytoplasm and standing for the organism. In addition, for the elements of the genotype, he proposed the notion of the gene. This terminology was gradually taken up by the genetics community. Johannsen’s codification, which was based on breeders’ practices of separating “pure lines,” has profoundly marked all of twentieth century genetics. What has largely escaped the attention of historians of science, however, is the polemical context in which Johannsen made these distinctions. In introducing his neologisms, Johannsen explicitly turned against “historical” notions of inheritance prevalent in eugenics and breeding. Yes, he even denounced the terms “heredity” and “inheritance,” taken in their everyday sense, as inadequate to capture the “modern view of heredity.” “Ancestry by itself is irrelevant; dispositions are decisive,” as he put it in his 1905 textbook Arvelighedslærens elementer. In making such statements, Johannsen was far from denouncing eugenics and breeding as “unscientific” as such. He rather wanted to put these applied sciences on a thoroughly instrumental and constructive basis, with chemistry as a paradigm. In my contribution I will discuss Johannsen’s roots in industrial research and how his view of Mendelism resonated with certain political ideas. Science, for Johannsen, was a modernizing force in as much as it was able to cut ties with tradition.

There is some kind of link, some kinship, among burdocks and beggars, singing in the fields, electricity, a locomotive and its whistle, and earthquakes—there is the same birthmark on all of them and some other things too … Growing grass and working steam engines take the same kind of mechanics. (Andrei Platonov, 1922)1

Looking at the cultural history of heredity in the “century of the gene” poses a number of historiographical problems as explained in the introduction to this volume. I would like to add an additional layer of complication: Current biological research is steering away from genecentrism.2 Heredity, as a consequence, is again supposed to involve much more than the transmission of genes. Such contemporary developments are not neutral with respect to how historians of science conceive of their object, that is, the history of a concept, theory, or discipline. While historians of science, for their own explanatory purposes, must be symmetrical, the historical object they deal with rarely, if ever, is. More often than not, science is radically asymmetrical, by claiming to “progress” and overcome what then turns out to have been “error”

1 2

From a letter to a publisher, quoted in Tolstaya (2000), p. xvi. Keller (2005).

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or “prejudice.” If it were not, it would simply not have a history.3 So if gene-centrism now turns out to have been mistaken, how are we to asses its history in twentieth century biology? There are several ways to answer this question. One is straightforwardly whiggish: Gene centrism was always mistaken, and the century of the gene was simply a century of error. A less whiggish, but still anachronistic answer, is that gene-centrism was a fiction necessary for heuristic reasons, a “stage” that biology had to go through to reach its present state of art. There are finally two reflexive answers, which turn on the presuppositions that have informed the histories written so far. One may want to question that genetics was central to twentieth century biology at all, and argue that a lot more than genetics was going on all along, but has been unduly overlooked by historians. Or one may want to question the meaning of genetics, and argue that genetics was about something entirely different than historians have so far told us. I will follow the latter line of argument in this paper. My aim in this is to open up classical genetics in order to see how it might fit into a cultural history of heredity. A lot of what I am going to say is derived from looking at the life and work of Wilhelm Johannsen, a key figure of early genetics. Yet the structure of my paper will be neither narrative, nor discursive, but rather aphoristic. I want to present a series of observations, all of which, in one way or other, turn around heredity, and all of which, in one way or other, turn around the notion of progress—a fundamentally political notion, as I see it. My overarching claim will be that classical genetics was the expression of an industrial culture that valued the future over the past, progress over tradition, autonomy over authority, and parts over wholes. 1. The first observation I want to make is trivial, but essential, I believe. Classical genetics emerged during a time when Europe was undergoing demographic, economic, and social changes on a massive scale, a development that has become known as “the demographic transition.” Some of the main parameters of the demographic transition are: fall of death-rate and fall of birth-rate, the latter occurring with a characteristic time lag, resulting in rapid population growth; rise in agricultural and industrial productivity; migration from rural communities to urban centres; and an overall rise in living standards, including nutrition and health care. To provide an illustration of the scale of these developments from a small European nation: In Denmark, between 1801 and 1901, 360,000 individuals migrated from agricultural areas to cities—or rather one city, Copenhagen. 143,000 of these went between 1881 and 1890 alone. From 1901 to 1950, the net-loss for agriculture was over 850,000 people. The population as a whole grew from 929,000 to 4,281,000 during the same time.4 The term “demographic transition” was coined in 1929 by the American population scientist Warren S. Thompson, director of the Scripps Foundation for Research in Population Problems. 5 The changes that the demographic transition encompassed, that is, had been under close scientific scrutiny—as is well-known, at least since Malthus. Moreover, the demographic transition associated a number of knowledge domains that arguably had some import for the knowledge of 3 4 5

8

Canguilhem ([1994] 2002), p. 16–20. Andersen (1979), p. 102. Thompson (1929).

Leaving Inheritance behind: Wilhelm Johannsen and the Politics of Mendelism

heredity: eugenics and reproductive medicine, of course; agricultural science, in particular breeding research; and—probably less obviously—microbiology in its medical, agricultural, and industrial applications.6 2. Much of nineteenth-century research into heredity was clearly motivated by a concern, or even fear of degeneration, and this remained true even for much of classical genetics. The five last chapters (out of twelve) of Ronald A. Fisher’s The Genetical Theory of Natural Selection dealt with the “decay of civilizations.”7 As a phenomenon, however, degeneration was unwieldy. In a crude manner, it simply reflected the negative consequences of the demographic transition. Any number of causes—from racial dispositions to alcoholism, from economic depravation to contagious diseases—could be held responsible for degeneration. And any number of measures—from positive eugenics to temperance, from birth control to public hygiene—could reasonably be suggested to counter or even reverse its negative effects. In the eugenic movement, therefore, all of these concerns lay very close. Alcoholism, for example, was thought to be both an effect and a cause of “bad” inheritance, simply by “poisoning” the germ line. 8 The metaphor of hereditary transmission covered the whole spectrum of causes and measures just mentioned, relying on the assumption that the “indefinitely numerous small causes” that generated variation in one generation would have some “average effect upon the offspring” of that generation.9 “Nature” and “nurture,” “heredity” and “environment,” under this perspective, did not separate two organic systems, one responsible for transmission, the other for development, but two sets of causes, one acting from the past, through pedigrees, onto the present, the other acting more or less instantaneously, at a given point in time.10 This perspective becomes particularly clear when Karl Pearson, in his 1910 presidential address to the annual meeting of the Social and Political Education League, titled “Nature and Nurture: The Problem of the Future,” reaches the conclusion that “[t]here is no real comparison between nature and nurture”—a surprising conclusion, if one takes into account that the bulk of the address was actually devoted to a meticulous comparison of the statistical effects of “nature” vs. “nurture.” As Pearson went on to explain his paradoxical conclusion, it is essentially the man who makes his environment, and not the environment which makes the man. That race will progress fastest where consciously or unconsciously success in life, power to reproduce its kind, lies with native worth. Hard environment may be the salvation of a race, easy environment its destruction.11

Hereditary superiority, that is, lies with the innate ability to shape one’s environment, even under adverse conditions, in order to reproduce one’s kind. Nurture, in this case, is not reduced to a mere accident, able to modulate nature only within strictly set limits. Nurture would rather be the 6 7 8 9 10 11

On microbiology see Bos and Theunissen (1995); Mendelsohn (2005); Müller-Wille (2007). Fisher (1930), p. xii. Snelders, Meijman, and Pieters (2005). Pearson (1896), p. 255. Pearson (1913), p. 11–12. Ibid., p. 27.

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very expression of nature, if the latter were only left alone to exert its positive influence. In trying to avert degeneration, the eugenic movement endorsed a view of inheritance as a natural force which had the potential to preserve and promote positive characters, but whose positive effects would only supervene, if society invested a lot to create conditions that would actually foster, and not inhibit, that potential. Nature was glorified as a powerful source of progress and at the same time deeply mistrusted in its ability to bring about progress simply on its own. 3. There was another, more specific, more precisely delineated phenomenon than degeneration, which excited research into heredity. This was reversion, regression, or atavism, the fact that ancestral characters sometimes reappear in more distant, descendant generations, while being absent from intermediate generations. Several points are interesting about this phenomenon. First, it depended on a temporal structuring of populations by generations, on an analysis of descent, that is. 12 Second, it encompassed a wider scope than procreation, the immediate production of offspring by parents, by looking at three generations at least. And third, it connected with an important issue raised in the context of contemporary cell theory, the issue of the relative autonomy of the living, elementary units of which organized beings were supposed to be composed. In the last chapter of The Variation of Plants and Animals under Domestication, presenting his notorious theory of pangenesis, Charles Darwin highlighted two conclusions that attention to reversion suggested with respect to the issue of autonomy. First, he reasoned that the “principle of Reversion [sic],” that “most wonderful of all attributes of Inheritance [sic] ... proves to us that the transmission of a character and its development, which ordinarily go together and thus escape discrimination, are distinct powers.”13 Transmission, that is, and development of a character are independent phenomena. The second conclusion that Darwin drew from the “principle of Reversion” was that “[o]vules and the male element, before they become united, have, like buds, an independent existence. Both have the power of transmitting every single character possessed by the parent form. We see this clearly when hybrids are paired inter se, for the characters of either grandparent often reappear, either perfectly or by segments, in the progeny.” 14 Individual characters, that is, may be transmitted independently of each other. 4. The second conclusion quoted in the previous section is as close as Darwin ever should get at formulating what Mendel, just a few years earlier, had called the “law of development of the progeny of hybrids,” and which we today would call the law of segregation. 15 It also allows us to understand why it was that Darwin thought that the “principle of Reversion” was the “most wonderful of all attributes of Inheritance.” Drawing on the work of Claude Bernard and Rudolf Virchow, Darwin subscribed to the view that “[e]ach organ has its proper life, its autonomy,” and 12 13 14 15

10

Parnes (2007). Darwin (1868), vol. ii, p. 372. Ibid., p. 360. On Mendel’s formulation of this “law” see Müller-Wille and Orel (2007).

Leaving Inheritance behind: Wilhelm Johannsen and the Politics of Mendelism

that each element of the living body “even though it derives its stimulus to activity from other parts, yet alone effects the actual performance of its duties.”16 Now, reversion, according to Darwin, was just the phenomenon that provided decisive evidence for this view, because it could only be explained under the assumption that “every character which occasionally reappears is present in a latent form in each generation ..., ready to be evolved under proper conditions.” 17 To put it differently: the potential to develop a character “under proper conditions” was apparently retained by each transmitted unit—or gemmule, as Darwin called it—independently of the particular bodies it passed through, and independently, in particular, from its combination with other such units in the fertilized egg. This was a relative autonomy only, to be sure. The development of characters clearly depended on “proper conditions,” more specifically, on the “union” of gemmules “with other partially developed cells or gemmules.” 18 But once these conditions were realized, development would always ensue in the same way according to Darwin. The developmental potential of gemmules was supposed to remain unaffected—or untainted, so to speak—by the various organic systems that they became part of while being transmitted. If that were not the case, if gemmules were somehow “tainted” along their way, parents would always leave a trace in their children—which is clearly contradicted by cases of reversion, where grandparental traits reappear without having reappeared in the parents. Darwin thus broke down the organism into two levels of organisation in his theory of pangenesis: a level consisting of the “completely passive or ‘formed material’” of the body, composed by fully developed cells; and a level consisting of the gemmules, thrown off by cells throughout their development, “circulat[ing] freely” through the body, and “multiplying by selfdivision” when supplied with “proper nutriment.”19 Gemmules, that is, represented the body in all its parts, but they did so in a manner that allowed them to circulate, recombine, and develop freely without changing their essential nature. It is in this sense that Darwin, towards the end of his Pangenesis-chapter, insisted that “[t]he child, strictly speaking, does not grow into the man, but includes germs which slowly and successively become developed and form a man.” 20 5. Cell theory, and the associated issue of the autonomy of the elements of organisms, also lay at the ground of Johannsen’s distinction of genotype and phenotype. This becomes especially clear in his contributions to a textbook in general botany that he co-authored with his teacher Eugen Warming in 1900. Among the chapters he contributed was one on the topic of the “periodicity in the life of plants.” Johannsen started this chapter with a thought experiment: If we think about a bacterium in a continuously renewed nutritive fluid of unchanged composition, at constant temperature, in permanent darkness etc., in short, under constant living conditions, we can assume that the bacterium will divide after a certain amount of time, the daughter cells will grow up and also divide etc. Cell division is obviously a consequence of 16 17 18 19 20

Darwin (1868), vol. ii, p. 368–369. Ibid., p. 373. Ibid., p. 374. Ibid. Ibid., p. 404. Darwin is playing here on a famous quote from a poem by William Wordsworth: “The child is the father of the man” (My heart leaps up when I behold, 1802).

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growth and nutrition, as well as of changes in the inner states of the cell provoked by growth and nutrition. And these periodic phenomena appear without the slightest change in external conditions. The nature and order of phases in the life history of bacteria is thus not dependent on a periodicity of external factors.21

Johannsen took this to show that “the course of development [of an organism], its ‘Grundplan’, is independent of external factors or at least not immediately dependent on them.” The reasoning behind this is obvious: If the course of the life of organisms depended entirely on external conditions, conditions held constant should keep them from developing. But organisms do develop, even in a constant environment, in the simplest case undergoing cell division after a certain amount of time. Johannsen believed that the source of this “independent and autonomous periodicity was still entirely mysterious.” So much could be said however: “In many cases there must be properties within the organism, which determine, that a certain activity, e.g. a growth process, causes a state after some time, which contravenes the continuation of that process, may be in an analogous manner as chemical processes ‘cease by themselves’ with the accumulation of their product.”22 Johannsen’s thought experiment, as well as the chemical analogy he used to explain the mysterious ability of organisms to develop, were not simply plucked out of thin air. In interesting ways, both show connections with another of Johannsen’s contributions to Warming’s textbook, a chapter that dealt with “fermentation and putrefaction” and the “occurrence and role of microorganisms in nature.” Here, Johannsen gave credit to Emil Christian Hansen, head of the bacteriology department at the Carlsberg Laboratory in Copenhagen, for the discovery “that there are whole series of yeast species or races that are of very different practical value” for the brewing industry.23 In order to develop methods that could prevent beer from turning sour occasionally, Hansen had adopted the pure culture approach from Louis Pasteur and Robert Koch in 1883. Isolating single yeast cells by repeated dilution and under the microscope, and cultivating them under sterile and constant conditions, allowed him to produce yeast consisting of beneficial strains of brewer’s yeast only. The strains were marketed successfully as Carslberg Bottom-Yeast No. 1 in the same year, and soon spread over breweries world-wide.24 6. Johannsen had himself started his research career at the Carlsberg Laboratory , a private research laboratory in Copenhagen associated with, but largely independent of, the famous beer brewery. In 1881, he entered its chemistry section as a research assistant with the task of applying analytic methods to determine the involvement of organic nitrogen in metabolic processes connected with the ripening and germination of plants, especially barley. In 1887 Johannsen left the Carlsberg Laboratory to take up a lectureship at the Royal Veterinary and Agricultural College in Copenhagen, but continued his collaboration with the Laboratory, now turning to experiments in breeding high quality strains of barley.25 Both projects were intimately connected, because the 21 22 23 24

12

Warming and Johannsen ([1900] 1909), p. 580–581. Ibid., p. 619. Ibid., p. 355. Teich (1983).

Leaving Inheritance behind: Wilhelm Johannsen and the Politics of Mendelism

nitrogen-content of barley was an important variable in the brewing process, and the quality of barley, in consequence, could be assessed, among other things, by measuring its protein content. Variation in plant form was thus reduced to variation in a single, measurable chemical variable. 26 It is in this context that Johannsen must have picked up the “pedigree” method from plant breeders like the French Louis de Vilmorin. Just like pure cultures, pedigrees—or “pure lines” as Johannsen should later call them—were genealogical constructs. A pedigree, or pure line, consisted of all descendants derived from a single individual through self-fertilization. There was therefore, as Johannsen used to put it, “no doubt about the father” in pure lines. 27 The vagaries of ancestry were reduced to a minimum, just as in pure cultures of asexually reproducing organisms, and pure lines could thus be expected to always react in the same way to given environments. In 1903, Johannsen should use pure lines of beans to draw the distinction of genotype and phenotype by demonstrating that selection was ineffective in genetically homogenous populations. 28 7. Johannsen was clearly aware that the use of pure cultures and pure lines did not relieve practitioners from attending to environmental conditions. The quality of beer, as he put it in the Warming textbook—and as he well knew as a Danish bonvivant—would always depend on “the locality in which fermentation went on.”29 Pure cultures and pure lines were therefore not of immediate practical use in local contexts—as Johannsen stated in his 1905 textbook Arvelighedens elementer (Elements of Heredity)—but of use in the circulation of plant material among such contexts only. “Use and propagation (i.e. breeding) both pose their own demands which should never be confused. Use counts on the individual and exploits the most advantageous conditions that can be created for its development to meet the purpose. Continued propagation must count on life-types, and different conditions of life show us here, what potential lies in a respective race.”30 Pure lines did not exist locally, and they could not be intuited based on localized experience. “There are so many who have made experiences,” as Johannsen polemisized against breeders, “starting from their experiences, [they] have formed notions of inheritance in which they believe like charburners.”31 Johannsen had similar views on eugenics, expounded in an article he wrote in 1927 for the Danish journal Naturens verden (The World of Nature) while serving as a member of the Danish state commission that was to draw up one of the first sterilization laws world-wide.32 Johannsen was sceptical about the prospects of eugenic policies, arguing that individual phenotypes were the expression of equations with two, or rather two sets of, unknowns: elements of disposition and environmental factors. “This causes the varied and not always happily concluding lottery in the life and fate of generations,” as the article concluded. 33 25 26 27 28 29 30 31 32 33

Roll-Hansen (2005). Johannsen (1899) summarizes the results of these projects, discussing mutations and the correlation of morphological and chemical variables. See Bonneuil, this volume, for more details. Quoted according to Roll-Hansen (2005), p. 47. Johannsen (1903). Warming and Johannsen ([1900] 1909), p. 356. Johannsen (1905), p. 177. Johannsen (1913), p. 4. On Johannsen’s engagement with eugenic policies of his time, see Koch (1996), p. 57–67. Johannsen (1927), p. 235.

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8. We meet the same kind of contempt for popular conceptions of heredity in Johannsen’s 1911 paper in the American Naturalist, that introduced the expressions genotype and phenotype to the English speaking world. The paper started out with a veritable diatribe against what Johannsen conceived as “the most naïve and oldest conception of heredity,” namely the “view of natural inheritance as realized by an act of transmission, viz. the transmission of the parent’s (or ancestor’s) personal qualities to the progeny.” “The personal qualities of any individual organism,” as Johannsen went on after sketching out the history of the transmission conception of heredity, do not at all cause the qualities of its offspring; but the qualities of both ancestor and descendent are in quite the same manner determined by the nature of the sexual substances—i.e., the gametes—from which they have developed. Personal qualities are then the reactions of the gametes joining to form the zygote; but the nature of the gametes is not determined by the personal qualities of the parents or ancestors in question. This is the modern view of heredity.34

This short passage contains the distinction of genotype and phenotype in a nutshell. It must be seen against the background of what Jean Gayon has called the “Mendelian break,” seperating biometricians from early geneticists, William Bateson in particular. In Darwinism, specifically in the version endorsed by the biometricians, inheritance was conceived of as a force or tendency that could be measured by the statistical “effects” that ancestors had on their descendents. Heredity became synonymous with the descent, lineage, or “pedigree” of an individual. A major characteristic of this approach was its descriptive character, in other words, its independence from any hypothesis about the mechanism of hereditary transmission. Mendelians, in contrast to that, regarded pedigrees not as objects, but as tools to uncover the genetic constitution, understood as an organic structure, of a given parental generation. To quote Jean Gayon: Heredity was not the sum total of ancestral influences; it was a question of structure in a given generation. What happened to the progeny did not depend on what happened to the ancestors of its parents, but only on the genetic makeup of its parents.”35 For Johannsen ancestral inheritance was a “mystical expression for a fiction,” and his contempt for biologists endorsing such views, especially Ernst Haeckel, was profound. 36 The genotype conception represented an “‘ahistoric’ view of the reactions of living beings” that was analogous to a “chemical view.” “Chemical compounds have no compromising ante-act, H 2O is always H2O, and reacts always in the same manner, whatsoever may be the history of its formation or the earlier states of its elements. I suggest that it is useful to emphasize this ‘radical’ ahistoric genotype conception in its strict antagonism to the transmission- or phenotype view.” 37 “Ancestry by itself is irrelevant; dispositions are decisive,” as Johannsen put it provocatively in his 1905 Textbook Arvelighedens elementer.38

34 35 36 37 38

14

Johannsen (1911), p. 130. Gayon (2000), p. 77. See Johannsen ([1914] 1917), p. 20, where Haeckel is referred to as the “high priest” of German Darwinism. Johannsen (1911), p. 139. Johannsen (1905), p. 216.

Leaving Inheritance behind: Wilhelm Johannsen and the Politics of Mendelism

9. What was modern about Johannsen’s “modern view of heredity”? First of all, I would like to maintain, the uncompromising willingness to detach the production of knowledge from tradition, from experience accumulated in the past. Tradition, in Johannsen, appears as a reservoir of myths and prejudices only, which mathematics in conjunction with experiment allows to overcome. Johannsen’s theory of heredity was an exact match of his epistemology. Johannsen was always proud of coming as a largely self-taught outsider to biology. “I am, and always will be, a free-lancer in science,” as he once stated.39 Johannsen’s father, a corporal in the Danish army, had been unable to pay his son a university education, and Wilhelm therefore had to seek his way into academia through an apothecary apprenticeship. Another aspect of Johannsen’s modernity is much more difficult to pin down. Nils RollHansen has remarked, that Johannsen “successfully bridged the gap” between early Mendelians and biometricians.40 Providing this bridge was not, however, only a matter of conceptual innovation. Johannsen’s emphasis on the importance of pure cultures and pure lines demonstrates a technological dimension, the creation of new entities, genes, that allowed to view and to deal with organisms as if they were constituted, quasi-mechanically, of independent elements, which could be transposed and combined at will, while retaining their propensity to produce definite effects under given circumstances. As James Griesemer has argued, genetics did not divorce transmission from development. Far from it, genetics, in its search for “developmental invariants,” was in a sense all about development.41 10. It needs to be emphasized that Mendelism comprised an impossible view of the organism, which is probably no better expressed than by William Bateson’s stunningly self-confident formula: “We can pull out the yellowness and plug in greenness, pull out tallness and plug in dwarfness.” Such statements contain an attractive promise of control over life, and many Mendelians were enthusiastic eugenicists. Not so Johannsen and Bateson. In the latter’s case, as Mackenzie and Barnes have argued, this correlated with conservative political convictions and a deeply rooted aversion against any “ideas of interventionist reformers [like Karl Pearson] riding the tide of advanced industrialism.”42 With Johannsen, the matter seems to have been a bit more complicated. In 1914, Johannsen published a booklet titled “False analogies with respect to similarity, kinship, inheritance, tradition, and development.” It was a sweeping attack against all attempts to analogize the social and the biological world. In particular, Johannsen directed biting comments against the assumption that “an active historical moment” was involved in the formation of individual organic beings, that heredity consisted in a kind of “memory” of the past that played a causal role in individual development.43 There was another assumption, however, that drove 39 40 41 42 43

Quoted in Winge (1958), p. 87. Roll-Hansen (1980), p. 512; cf. Gayon (1998), p. 512. Griesemer (2000). MacKenzie and Barnes (1979), p. 205. Johannsen ([1914] 1917), p. 38.

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Johannsen mad, and that he saw instantiated by Darwin’s theory of pangenesis: the assumption, namely, that the organism is an ensemble of independently reproducing parts. 44 This is surprising, given Johannsen’s own reliance on the tenets of cell theory. And did not his own conceptual creation, the gene, constitute an element of the organism that reproduced itself independently of circumstance? Well, as Lenny Moss and Rafi Falk have shown, Johannsen always remained critical of “gene for” talk, of “genes” being and representing parts of the organism. 45 He consistently resisted the temptation to follow the Morgan school in locating genes on chromosomes. 46 It thus seems that Johannsen, in some sense, wanted to have it both ways. His analytical skills had made his career, initially in an industrial context, where advancing over traditional procedures was what counted to achieve success, and where “atomizing” life into constituent units that could be moved around and recombined at will was a good strategy to make such advances. It was here, indeed, where ancestry was irrelevant, and where dispositions were decisive. Faced with the prospects and projects of eugenicists eager to avert “degeneration,” on the other hand, Johannsen tended to emphasize the haphazard nature of genetic recombination, and the inability to effectively control the future on the basis of genetics. Here, life essentially became a gamble. Johannsen’s writings thus exhibit a deep ambiguity. “Personally,” he wrote in 1923, “I believe in a great central ‘something’ as yet not divisible into separate factors”, and he identified this “something” with the “specific or generic nature of the organism. The pomace-flies in Morgan’s splendid experiments continue to be pomace-flies even if they loose all ‘good’ genes necessary for a normal fly-life, or if they be possessed with all the ‘bad’ genes, detrimental to the welfare of this little friend of the geneticist.”47 Genetic analysis, on one level, was a safeguard of personal autonomy, as it allowed to escape the weight of history, and to literally make one’s own life. On another level, however, it constituted a threat to personal autonomy, by dissolving the life of individuals into ponderable elements and putting this life at the mercy of powers beyond their control. Johannsen, it seems, was acutely aware of this political dilemma that the science of heredity posed. Staffan Müller-Wille ESRC Research Centre for Genomics in Society, University of Exeter, [email protected]

44 45 46 47

16

Ibid., p. 40. Falk (1986), 135–141; Moss (2003), p. 28–44. Churchill (1974). Johannsen (1923), quoted in Moss (2003), p. 38.

Leaving Inheritance behind: Wilhelm Johannsen and the Politics of Mendelism

References Andersen, Otto. 1979. “Denmark.” In William Robert Lee (ed.), European Demography and Economic Growth. London: Croom Helm. 79–122. Bos, Pieter, and Bert Theunissen. 1995. Beijrinck and the Delft School of Microbiology. Delft: Delft University Press. Canguilhem, Georges. [1994] 2002. Etudes d’histoire et de philosophie des sciences concernant les vivants et la vie, reprint of the 7th edition. Paris: Vrin. Churchill, Frederick. 1974. “William [sic] Johannsen and the genotype concept.” Journal of the History of Biology 7: 5–30. Darwin, Charles [1868] 1988. Variation of animals and plants under domestication, 2 vols., Vol. 20 and 21 of The Works of Charles Darwin, edited by P.H. Barrett and P.B. Freeman,. New York, NY: New York University Press. Falk, Raphael. 1986. “What is a gene?” Studies in the History and Philosophy of Science 17: 133–173. Fisher, Ronald A. 1930. The Genetical Theory of Selection. Oxford: Clarendon Press. Gayon, Jean. 1998. Darwinism’s Struggle for Survival. Heredity and the Hypothesis of Natural Selection. Cambridge: Cambridge University Press. ————. 2000. “From measurement to organization: a philosophical scheme for the history of the concept of heredity.” In Peter Beurton, Raphael Falk, and Hans-Jörg Rheinberger (eds.), The Concept of the Gene in Development and Evolution: Historical and Epistemological Perspectives. Cambridge: Cambridge University Press. 69–90. Griesemer, James R. 2000. “Reproduction and the Reduction of Genetics.” In Peter Beurton, Raphael Falk, and Hans-Jörg Rheinberger (eds.), The concept of the gene in development and evolution: historical and epistemological perspectives. Cambridge: Cambridge University Press. 240–285. Johannsen, Wilhelm. 1899. “Fortsatte Studier over Kornsorterne I. Om Variabiliteten med særligt Hensyn til Forholdet mellem Kornvægt og Kvæfstof-Procent i Byg.” Meddelelser fra Carlsberg Laboratoriet 4: 228–313. ————. 1903. Über Erblichkeit in Populationen und in reinen Linien. Jena: Gustav Fischer. ————. 1905. Arvelighedslærens elementer. Forelæsninger holdte ved Københavns Universitet. København and Kristiania: Gyldendal. ————. 1911. “The genotype conception of heredity.” American Naturalist 45: 129–159. ————. 1913. Elemente der exakten Erblichkeitslehre. Mit Grundzügen der biologischen Variationsstatistik. In dreißig Vorlesungen, 2nd edition. Jena: Gustav Fischer. ————. [1914] 1917. Falska Analogier med hänsyn till likhet, släktskap, arv, tradition och utveckling, transl. R. Larsson. Stockholm: Hugo Geber. ————. 1923. “Some remarks about units in heredity.” Hereditas 4: 133–141. ————. 1927. “Racehygiejniske problemer.” Naturens Verden 10: 214–235. Keller, Evelyn Fox. 2005. “The century beyond the gene.” Journal of Bioscience 30: 3–10. Koch, Lene. 1996. Racehygiejne i Danmark, 1920-1956. Københamn: Gyldendal. MacKenzie, Donald, and Barry Barnes. 1979. “Scientific judgment: The Biometry-Mendelism controversy.” In Barry Barnes and Steven Shapin (eds.), Natural order: Historical studies of scientific culture. Beverly Hills, Calif.: Sage. 191–210. Mendelsohn, Andrew. 2005. Message in a Bottle: The Business of Vaccines and the Nature of Heredity after 1880.” In A Cultural History of Heredity III: 19th and Early 20th Centuries. Preprint 294. Berlin: MaxPlanck-Institute for the History of Science. 85–100. Moss, Lenny. 2003. What Genes Can’t Do. Cambridge, MA: MIT Press. Müller-Wille, Staffan. 2007. “Hybrids, pure cultures, and pure lines: From nineteenth-century biology to twentieth century genetics.” Studies in History and Philosophy of the Biological and Biomedical Sciences 38: 796-806. Müller-Wille, Staffan and Vitezslav Orel. 2007. “From Linnaean species to Mendelian factors: Elements of hybridism, 1751-1870.” Annals of Science 64: 171–215. Nilsson-Ehle, Nils Hermann. 1927. “W. Johannsen.” Nordisk Jordbrugsforskning 1927, 361–364. Parnes, Ohad. 2007. “On the shoulders of generations: The new epistemology of heredity in the nineteenth century.” In Staffan Müller-Wille and Hans-Jörg Rheinberger (eds.). Heredity Produced: At the Crossroads of Biology, Politics, and Culture, 1500-1870. Cambridge, MA: MIT Press. 315–346. Pearson, Karl. 1896. “Mathematical contributions to the theory of evolution III: regression, heredity and panmixia.” Proceedings of the Royal Society of London, Series A, 187: 253–318. ————. 1913. Nature and Nurture: The Problem of the Future, Vol. 6 of Eugenics Laboratory Lecture Series.

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London: Cambridge University Press. Roll-Hansen, Nils. 1980. “The controversy between biometricians and Mendelians: A test case for the sociology of scientific knowledge.” Social Science Information 19: 501–517. ————. 2005. “Sources of Johannsen’s genotype theory.” In A Cultural History of Heredity III: 19th and Early 20th Centuries. Preprint 294. Berlin: Max-Planck-Institute for the History of Science, 43–52. Snelders, Stephen, Frans J. Meijman and Toine Pieters. 2005. “Heredity and alcoholism in the medical sphere: The Netherlands, 1850-1900.” In A Cultural History of Heredity III: 19th and Early 20th Centuries. Preprint 294. Berlin: Max-Planck-Institute for the History of Science. 193–211. Teich, Mikulás. 1983. “Fermentation theory and practice: The beginnings of pure yeast cultivation and English brewing, 1883-1913.” Technology and Culture 8: 117–133. Thompson, Warren. 1929. “Population.” American Journal of Sociology 34: 959–975. Tolstaya, Tatyana. 2000. “Introduction.” In Andrei Platonov, The Fierce and the Beautiful World: Stories, translated by Joseph Barnes. New York: New York Review Books. ix–xviii. Warming, Eugen, and Wilhelm Johannsen. [1900] 1909. Lehrbuch der allgemeinen Botanik, 4th edition, transl. E. P. Meinecke. Berlin: Bornträger. Winge, Øjvind. 1958. “Wilhelm Johannsen. The creator of the terms gene, genotype, phenotype and pure line.” Journal of Heredity 49: 83–89.

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Mendelian Factors and Human Disease: A Conversation Jean Paul Gaudillière & Ilana Löwy

A fictitious conversation, in 1930s, between

—Miss Mina Mauser, an indefatigable laboratory worker and enthusiastic follower of the new theories on heredity, who has dedicated all her life to studying hereditary phenomena in inbred mice. Miss Mauser is a typical spinster: austere and slightly fanatical; and —Professor Adolph Influence, a brilliant clinician, who has dabbled in multiple domains of medical investigation, is a pro-natalist, believes in the superiority of “intransmissible” clinical knowledge. Professor Influence is a highly successful doctor and has the charm and polished manners of his profession.

MM: Doctors are always talking about “heredity,” “hereditary predisposition,” and “hereditary conditions.” But if you listen to them carefully, you can see that the term “heredity,” as they use it, is far from clear. They confuse true hereditary conditions with vague “parental influence,” and they include under the same heading the “degeneration” induced by diseases such as syphilis or tuberculosis and behavioural traits such as alcoholism, and even the effects of the poor health of mothers during pregnancy on the newborn child. They’ve never really understood what the new science of heredity is all about.1 Take for example Charles Richet. He is a great scientist, he has won the Nobel prize for his investigations in anaphylaxis, but he is also an enthusiastic supporter of eugenics.2 Richet has suggested that interracial marriages should be prohibited, that men found unfit for military service should not be allowed to marry, that the marriages of sick and mentally handicapped people and of those with identified criminal inclinations should be regulated, and that the sterilization of recidivist criminals should be mandatory.3 He has supported these rather 1 2

3

See, e.g., papers by Andrew Mendelsohn and Patrick Zylberman. In Jean Paul Gaudillière and Ilana Löwy (eds.). Heredity and Infection. The History of Disease Transmission. London: Routledge 2001. On Richet, see Charles Richet. Souvenirs d’un physiologiste. Paris; J. Peyronnet & Cie 1933; André Mayer. “Notice Nécrologique sur M. Charles Richet.” Académie de Médecine. Session of 14 January 1936; CR Académie de Médecine. 1936. pp. 51-64; Gustave Roussy. “Éloge de Charles Richet.” Académie de Médicine. Session of 18 December 1945. CR Académie de Médecine. 1945. pp. 725-731; Stuart Woolf. Brain, Mind and Medicine. Charles Richet and the Origins of Physiological Psychology. New Brunswick and London: Transaction Publishers 1992; Pierrette Estingoy. DEA thesis. “Charles Richet, 1850-1935. Esquisse biographique et bibliographie.” PhD in History. University of Lyon III. Jean Moulin 1993; Pierrette Estingoy. “Charles Richet et la découverte de l’anaphylaxie. Histoire d’un prix Nobel de médecine.” Thesis in Medicine. University of Lyon I. Claude Bernard 1996. Charles Richet. “La sélection humaine.” Eugénisme, Organe de la société française d’eugénique 1923, 3(1). This study was written in 1912, first published in 1919, and in 1921 a revised version of this essay was published by the French Eugenic society.

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extreme measures because he believes in invariable hereditary traits. On the other hand, he also believes that heredity could be modulated by the environment. For him, not all transmissible traits are fixed. Some are, but others are plastic and can be changed by the appropriate manipulation of external conditions. Really, according to these people, anything goes! Richet has even declared that Darwinism should blend with Lamarckism.4 What kind of scientific approach is that? AI: That is precisely the point. We are not dealing with the artificial conditions of a laboratory but with real human beings. We are not breeding homogeneous mice, but trying to help parents have healthy children and provide healthy citizens. You seem to believe that eugenic measures are in contradiction with efforts to improve the health of expectant mothers—but where is the contradiction? We all know that some people—those suffering from advanced tuberculosis or syphilis, those with important physical handicaps, those with a family history of mental diseases— are unfit to be parents.5 Charles Richet wants such people to be formally prohibited from marrying. Most physicians will not go that far but do think that people with hereditary handicaps should be persuaded to refrain from having children. Responsible family doctors are expected to advise their patients on these issues. On the other hand, even individuals without hereditary handicaps can produce sickly and unfit children. The French paediatrician Professor Pinard’s studies have shown that a child’s weight at birth and its subsequent health strongly depend on the health status of the mother during pregnancy. Pinard has also demonstrated that women who work under difficult conditions while pregnant have higher rates of spontaneous abortion, more frequent pregnancy complications and can give birth to sick children. Helping poor women to have healthy children is important for the nation. Paid maternity leave for women who are forced to work for economic reasons and free medical service for all pregnant women are also excellent investments, because a small expenditure during the mother’s pregnancy will save much larger amounts of money spent on an ailing child and, if the child survives, on a sickly adult.6 Charles Richet is a great experimental physiologist, but above all, he is a clinician. He is familiar with the complexity of physiological and pathological conditions and, for this reason, does not treat animals—or humans—like test tubes with legs. His understanding of anaphylaxis— a violent reaction to a “sensitizing” protein—has led him to a more complex view of the interactions between hereditary and acquired conditions. Anaphylaxis, argues Richet, illustrates the impossibility of separating innate physiological reactions from acquired components. Some anaphylactic reactions, such as sensitization by injection of horse serum—the so-called “serum sickness”—are induced by a repeated contact with an external antigen. Other “anaphylactic type” reactions, such as allergies to foodstuffs or drugs, occur with no previous contact with the allergen and are therefore probably induced by innate mechanisms. Nonetheless, both kinds of 4

5 6

20

On Richet’s concept of occasional inheritance of acquired traits, see “Autobiographie de professeur Charles Richet recueillie par le Dr. Pierre Maurel.” Les Biographies Médicales – Revue Mensuelle Illustrée. 1932, 6(8), Part II, pp. 173- 188. See, e.g., William Schneider. Quality and Quantity. The Quest for Biological Regeneration in Twentieth Century France. Cambridge: Cambridge University Press 1990. William Schneider. “Puericulture and Style of French Eugenics.” History and Philosophy of Life Sciences. 1986, 8, pp. 265-277; Charles Richet. “La protection de la maternité.” Bulletin de l’Académie de Médecine. 1917, 77 (3), pp. 605-634.

Mendelian Factors and Human Disease: A Conversation

anaphylaxis are rigorously identical from a physiopathological point of view. 7 And the first concern of a doctor treating a patient with severe allergic manifestations is not to find out if the patient is suffering because he was born sensitized to an allergen, acquired such sensitization sometime in the past, or from a mix of both of these mechanisms. What a doctor really needs to do is find a way to alleviate the patient’s suffering and prevent future incidents. MM: But if we really want to understand disease, not just to try to provide symptomatic treatment of bothersome symptoms, we need to develop a rigorous scientific approach, tested under wellcontrolled experimental conditions. We need to study diseases that run in families and find the Mendelian distribution of diseases known to be hereditary, such as haemophilia. AI: Haemophilia is precisely a very good example. Some scientists believe that this is a true “Mendelian disease,” but the French paediatrician Eugène Apert, who published the genealogical tree of a family with haemophilia, explained that in this case we were dealing with a modification of Mendelian transmission through “maternal inheritance.” Professor Apert provided a good explanation of the complexity of human heredity. Allow me to quote him: Identical transmission of a disease from one generation to the next, which may be called inheritance of the same, is rare. It is, however, the rule for a few illnesses, known as familial diseases. Although these are exceptional, the study of hereditary disorders must begin with them, because they represent the least complex form of morbid inheritance. It should be borne in mind that they are just a small corner of a vast field. If, for instance, the father is a drunkard, he will produce a son differing from the normal type within his lineage. Within this family, there will be a tendency to degenerate, since the father’s sperm, or rather the cell from which it derives, has been the target of obnoxious effects originating in the bad condition of the paternal organism.8

Professor Apert understood that normal inheritance is Mendelian. But he also made it clear that pathological inheritance is more complex than normal inheritance. He also pointed to the fact that familial disorders are rare and that these pathologies are marginal when viewed from a medical and public-health perspective. Geneticists love to draw pedigrees. Pedigrees, to be sure, are very useful, but only if we take them, not as a demonstration of the exclusive role of Mendelian factors but as a means for revealing how these factors are modified by other influences, whether they are environmental, social or physiological.9 Another French doctor, Professor Raymond Turpin, has studied “mongolism.” Other researchers had previously shown that although the condition was inborn, there was no familiar 7

8

9

Charles Richet. “Humorisme ancien et humorisme moderne.” La Presse Médicale. 1 October 1910. English version. Charles Richet. “Ancient Humorists and Modern Humorism.” British Medical Journal. 1910. ii. pp. 921-926; Charles Richet. L’Anaphylaxie. Paris: Felix Alcan. 1911. English translation. Charles Richet. Anaphylaxis. (translated by J. Murray Blight). London: Concable & Company 1913; Charles Richet, “Anaphylaxis.” Nobel Lecture, 11 December 1913. Nobel Lectures. Physiology or Medicine. Amsterdam and London: Elsevier Publishing Company 1967. Vol. 1, pp. 473-492. Eugène Apert. “Traité des maladies familiales et des maladies congénitales.” Paris: Baillière 1907. In Jean Paul Gaudilliere. “Mendelism and Medicine. Controlling Human Inheritance in Local Contexts. 19201960.” CRAS 2000, 323: pp. 1117-1126. Raymond Turpin. “L’avenir des caractères acquis.” Le Progrès Médical. 16 April 1932.

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clustering, and they had decided that the condition was not inherited. Turpin has a different view. “Mongol” children have a furrowed tongue. Turpin claims that this trait is often displayed by their parents as well, which might be an indication of simple Mendelian transmission. On the other hand, Turpin and his co-workers have shown that the birth of a “mongol” child is often associated with late pregnancy; the older the mother, the higher the frequency of mongolism. Their conclusion is that there is no simple Mendelian transmission, but a twofold “familial imprint,” genetic and physiological, that favours mongolism.10 We can perhaps demonstrate the inheritance of a simple trait, but complex traits are, well … complex. MM: I disagree! Suppose we look at cancer. Cancer is without doubt a very complicated disease. Nevertheless, there are important differences in the distribution of malignant tumours in populations. So-called “savages” rarely suffer from cancer: the disease is much more frequent in “civilized” countries. The distribution of types of cancer in specific populations is also very different: people in Northern Europe suffer more from different malignant tumours than people in central Asia, and Negroes in the United States have a much higher incidence of cancer than their African ancestors.11 In addition, we all know about “cancer families.” In some families, every single member seems to die of cancer, often at a relatively young age. This is even known by insurance firms, as they are reluctant to sell life insurance to people who have several relatives who have died young from a malignancy.12 This is something your fellow physicians refuse to admit: they want to persuade the general public that cancer is not hereditary so the disease is seen as curable and people are encouraged to consult a doctor as soon as they detect a symptom that could indicate the presence of malignancy.13 They have therefore opted to avoid the question of the heritability of cancer and hide behind vague terms such as “hereditary influence.” For example, an editorial in the Journal of the American Medical Association stated in 1932: Cancer is a disease of such protracted development and course, so variable in its manifestations and duration, often so difficult of diagnosis and differentiation, that satisfactory study of many fundamental problems on the basis of clinical observations is almost or quite impossible. At the present time it seems safe to maintain that the existence of an hereditary influence on the susceptibility and resistance to cancer has been established for both man and animals. The exact mechanism of the hereditary influence has yet to be determined. The evidence offered by human material is conflicting and inadequate both in amount and character to permit satisfactory analysis.14

10

11 12 13

14

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Raymond Turpin. “Le mongolisme, étude clinique et fonctionnelle.” Semaine des Hôpitaux de Paris. 31 December 1931; R. Turpin, A. Caratzali. “Conclusion d’une étude génétique de la langue plicaturée.” CRAS 1933, 196: 2040-2045; R. Turpin, A. Caratzali.“Remarques sur les ascendants et les collatéraux des sujets atteints de mongolisme.” La Presse Médicale. 25 July 1934, pp. 1186-1190. Federick L. Hoffman. “Cancer in the North American Negro.” American Journal of Surgery and Gynecology. 1931, pp. 229-263, quotation p. 241. J. Paterson MacLaren. Medical Insurance Examination. Modern Methods and Rating of Lives. New York: William Wood and Company 1943. pp. 530-531. E.E. Bashford. The Influence of Heredity on Disease. London: Longmans, Green & Co. 1909. pp. 63-66; James Ewing. “Heredity and Cancer.” Bulletin of American Society for the Control of Cancer. August 1942, 24(8); pp. 4-7. Editorial. “The Influence of Heredity on Cancer.” JAMA. 7 May 1932, reproduced in Bulletin of ASCC, 1932, 4 (8): pp. 3-4.

Mendelian Factors and Human Disease: A Conversation

Fortunately, some researchers have not been discouraged by this muddled argumentation and have set up specific experimental systems to study inheritance in cancer. In the 1910s, the 1920s and now in the 1930s, researchers such as Clara Lynch, Maud Slye, Nathalia Dobrovolskaia Zavadzkaia and Clarence Cook Little have developed “cancer prone” lines of mice and attempted to display the genetic mechanism that made these mice specially susceptible to malignancies. This has not, to be sure, been an easy task, and these researchers have not always agreed among themselves. For example, Maud Slye investigated spontaneous tumours surfacing in mice “families” that were kept under constant investigation and systematically autopsied to identify and document the cause of each death. This strategy has mainly produced pedigrees showing cancer families and non-cancer families of mice, which can be compared with human pedigrees but display clearer patterns of transmission, and has permitted the computation of Mendelian ratios. On the basis of several thousand necropsies and hundreds of charts, Slye claimed in the 1920s that cancer was not only a transmissible factor, but that a common recessive gene was involved in the appearance of tumours in utterly different locations. The geneticist Clarence Little strongly opposed this form of experimental practice, because it did not involve true “pure lines” of mice. Only models based on such lines, Little argued, would allow the development of reliable knowledge on the input of genetic factors to the genesis of cancer. Little opposed the “messiness” and semi-qualitative nature of Slye’s work, while the latter considered Little’s inbred mice as by-products of an artificial selection process that made comparison with humans impossible. This controversy notwithstanding, the breast-cancer mice Little and his co-workers have developed lately have become widely-circulated research tools. Thanks to this and similar animal models such as the “cancer lines” of mice developed at the Curie Institute in France, we will soon be able to understand the true input of heredity to cancer. 15 AI: Of heredity to cancer in mice! Look what has happened to studies that have attempted to investigate more precisely the links between heredity and cancer in human beings. About 10 years ago, the Sub-Committee on Statistics of the Cancer Committee of the League of Nations funded an extensive investigation on the racial determinants of cancer.16 The study on the “relationships between cancer and race” conducted by Professors Alfredo Nicoforo and Eugène Pittard relied above all on anthropological measures to define “race.” The study was limited to European populations. These populations were divided into three main racial types: Homo europeus, Homo alpinus and Homo mediterraneus, according to the distribution of physical traits such as eye and hair colour, height and build, the shape of the nose and the form of the skull. Nicoforo and Pittard painstakingly mapped racial traits on one side and the distribution of tumours on the other, and tried to correlate the two maps. And what did they find? Not much, really. At best, some vague 15

16

Ilana Löwy and Jean Paul Gaudillière. “Disciplining Cancer: Mice and the Practice of Genetic Purity.” In J.P. Gaudillière and I. Löwy (eds.). The Invisible Industrialist. Manufactures and the Production of Scientific Knowledge. London: Macmillan 1998. pp. 209-249; Jean Paul Gaudillière. “Circulating Mice and Viruses: The Jackson Memorial Laboratory, the National Cancer Institute and the Genetics of Breast Cancer.” In Michael Frotrun and Everett Mendelsohn (eds.). The Practices of Human Genetics. Dordrecht: Kluwer 1999. pp. 89-124. The commission was headed by Major Greenwood, and its members were Professor H.T. Delman, Dr. Janet Lane-Claypon, Dr. Henri Methorst, Professor Alfred Niceforo, Professor Eugene Pittard, Major P.G. Edge and Professor Goustave Roussy.

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indication that the Homo mediterraneus seemed to be more cancer-prone than the Homo alpinus. They blamed their relatively unconvincing work on the inadequacy of data on cancer mortality and on the distribution of racial traits in Europe, and they recommended that collection of this type of data should be reinforced, but one just wonders if better data could provide more interesting results.17 And when the British statistician Janet Lane-Claypon investigated correlations between family antecedents and the prevalence of breast cancer, she also failed to show statistically meaningful data.18 The questionnaire for cases of breast cancer and control cases developed by Lane-Claypon included the question, “Is there any information as to other forms of tumour in the family?” For practical reasons—given that people seldom know the cause of death of their grandparents and sometimes lose contact with their siblings—Lane-Claypon concentrated on the causes of the parents’ death. The results were inconclusive. They seemed to indicate that there were more deaths from malignancies among the mothers of women with breast cancer than among their fathers, but the difference was not significant and its meaning was unclear. Lane-Claypon notes that “this may, perhaps, be in part accounted for by the fact that there are fewer unknown causes of death among mothers than among fathers. It may be also a small though real difference.” She is also careful to point out that the limitations of her data are such, that these results, “are not of nature either to prove or to disprove the inheritability of cancer.”19 Well, this is not much different from what is stated in the editorial that you dislike so much in the Journal of the American Medical Association: there is perhaps some “hereditary influence” in cancer, but we are unable to determine what it is exactly and how to measure it. MM: What about FAP, Familiar Adenomous Polyposis? Is this also a vague “hereditary influence” on cancer? FAP is a condition in which people have numerous polyps of the rectum. As you probably know, in 1925, Mr. Percy Lockhart-Mummery, the surgeon at St Mark’s Hospital in London, published three pedigrees of patients with extensive polyposis, which showed that this condition runs in families, and he suggested that the presence of multiple polyps invariably leads to the development of colon cancer.20 Unlike many of his colleagues, Lockhart-Mummery does not believe that cancer develops as a result of “irritation” or “trauma.” He affirms that cancer is always the result of a mutation. His studies of FAP also led him to the conclusion that a second mutation may be needed to transform a simple tumour—in the case of FAP, an adenomatous polyp—into a malignant one.21 In the end, he established an FAP registry, which records pedigrees of families with this disease.22 Here we have, beyond any possible doubt, true Mendelian inheritance of cancer! 17

18

19 20 21

24

Alfredo Niceforo and Eugène Pittard. Considerations Regarding the Possible Relationships of Cancer to Race, Based on Study of Anthropological and Medical Statistics of Certain European Countries. Geneva: Publications of the League of Nations 1928. Janet Lane-Claypon. A Further Report on the Cancer of the Breast with Special Reference to Its Associated Antecedent Conditions. Reports on Public Health and Medical Subjects. N°32. London: Ministry of Health 1926. Lane-Claypon. A Further Report on the Cancer of the Breast. pp. 61-62. John Percy Lockhart- Mummery. “Cancer and Heredity.” Lancet. 1925. i. pp. 427-429. John Percy Lockhart-Mummery. Leaflet. The Origin of Tumours. London: John Bale Sons & Danielson 1932; John Percey Lockhart-Mummery, C.E. Dukes & M.D. Edin. “Familiar Adenomatosis of Colon and Rectum.” The Lancet. 1939. ii. pp. 586-587.

Mendelian Factors and Human Disease: A Conversation

AI: I do not deny that FAP is a truly “Mendelian” condition. But how many people have FAP, compared with the number of people who suffer from non-hereditary cancer of the colon? The FAP story reminds me of Lionel Penrose’s investigation of the role of heredity in mental disease. His study, the Colchester Survey, aimed at a quantitative evaluation of the incidence of inherited mental deficiency among the inmate population at the Colchester asylum. 23 The study comprised two parts. First, Penrose examined the inmates, reporting on both their clinical and psychiatric status. Second, Penrose and his co-workers visited the inmates’ families, supplementing, whenever possible, the Colchester data with clinical and intelligence testing of the parents. The IQ testing provided a continuous scale, from mild to severe deficiency, while the clinical examination provided data on the prevalence of specific pathologies such as mongolism, Huntington’s chorea, dystrophies of endocrine origins, etc. In the final report of the study, Penrose stressed the fact that very few mental disorders, including “mongolism,” followed a Mendelian pattern of transmission.24 Only one condition, phenylketonuria, followed this pattern, but Penrose was aware of the fact that this was a rare disease, a medical curiosity rather than a public-health problem. 25 Even a dedicated student of Mendelian heredity like Penrose has had to acknowledge that while a “pure” Mendelian transmission of pathological traits can be observed in a handful of rare pathologies, for the majority of diseases these traits are hopelessly intermingled with other variables. In short, we clinicians are not opposed to the idea that some diseases may be transmitted according to a Mendelian pattern. We simply do not believe that the study of such patterns is very relevant to our daily practice. Human diseases are very complex phenomena, and so are the vertical transmission of parental traits and the distribution of diseases among populations. We are quite pleased that scientists study the role of Mendelian factors in laboratory animals and in experimental diseases. It is surely a worthy endeavour that will enhance human knowledge, as do astronomy, botany or Egyptology, but we are not convinced that these studies are very relevant to what we are trying to accomplish at the bedside. MM: My dear Professor Influence! You are beautifully eloquent, as the members of your profession often are. Your success as a clinician depends as much, and sometimes more, on your rhetoric and bedside manners than on sound science. But, alas, you belong to a dying breed. Medicine is inexorably being transformed into an experimental science. In half a century or so, all diseases will be defined in genetic terms and medical science will be grounded on studies of Mendelian factors. AI: My dear Miss Mauser! You are entirely wrong. It is your approach that is condemned to become irrelevant, at least in medicine. You and your colleagues will perhaps be able to breed 22

23

24 25

Paolo Palladino. “Speculations on Cancer-free Babies: Surgery and Genetics at St Mark’s Hospital, 19241995.” In Jean Paul Gaudillière and Ilana Löwy (eds.). Heredity and Infection. Historical Essays on the Transmission of Human Diseases. London and New York: Routledge 2001. pp. 285-310. Daniel Kevles. In the Name of Eugenics. Berkeley: University of California Press 1985; J-P Gaudillière. “Le syndrome nataliste. Eugénisme, médecine et hérédité en France et en Grande-Bretagne.” In J. Gayon (ed.). L’éternel retour de l’eugénisme. Paris: PUF 2006. Lionel Penrose. A Clinical and Genetic Study of 1280 Cases of Mental Defect. Penrose Papers. University College London 59/2. Lionel Penrose. Lancet. 1935. vol. 1, p. 23 and vol. 2, p. 192.

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thousands of mice, test their susceptibility to numerous diseases and perhaps even cure some of these animals – but this will not bring you any closer to efficient cures for human diseases. Some day, scientists will move beyond the simplistic concept of Mendelian factors and will find more sophisticated ways of understanding biological and pathological phenomena. MM: I wish I could live to 2006, when it will surely be obvious which one of us is right today! AI: I wish I could live to 2006, when it will surely be obvious which one of us is right today!

Jean Paul Gaudillière & Ilana Löwy Centre de recherche médicine, science, santé et societé, Paris [email protected] [email protected]

26

Heredity without Mendelism: Theory and Practice of Dairy Cattle Breeding in the Netherlands 1900-19501 Bert Theunissen

Introduction In the 1940s and 1950s, Dutch geneticists and animal husbandry specialists repeatedly criticised the practices of dairy cattle breeders. The romantic idea that breeding was an art rather than a science seemed ineradicable, the experts lamented. Some breeders might even be accused of breeding for fancy rather than for utility. Particularly the top breeders seemed virtually oblivious of the fact that dairying was an economic activity and that the productivity of dairy cattle should come first. To bolster their claims, the experts pointed to the dominant role that conformation shows still played in assessing the value of breeding stock, despite the availability of more scientific methods for evaluating the animals’ qualities. Herd-book inspectors, breeders and farmers alike still judged the hereditary potential of a young bull on the basis of phenotypic characteristics. Now in pig breeding, for instance, where the objective was the production of pork, judging a boar on the basis of its weight and conformation made sense, for the animal’s outward appearance might indeed provide an indication of its economically valuable hereditary qualities. Breeding dairy cows however was a different matter. Although there was a clear difference in conformation between dairy cows and cows bred for beef production, correlation studies had shown that most individual details of conformation in dairy cows were unrelated to their milk yield. In the case of bulls, the breeders’ preoccupation with their phenotype was even more questionable. It was not their looks that mattered, but their daughters’ milk yield. An objective verdict on the qualities of breeding stock could only be obtained by progeny testing: the best animals, bulls as well as cows, were those that produced the most productive offspring. What was even worse, the scientists continued, was that many breeders seemed to be on the wrong track altogether in their choice of sires and dams. Despite their claims of being able to ‘see’ an animal’s qualities in its conformation, objective data from the milk recording services showed that milk yields had hardly increased over the last ten to fifteen years. The bulls and cows themselves, however, as the records of the herd-books showed, had definitely changed: they had become smaller, deeper, more short-legged and beefier. True, the Dutch dairy cows were a double purpose breed, producing milk and meat, but the greater part of the profit came from the milk, and the breeders were clearly overemphasizing their animals’ disposition for meat production. Considering the many prizes such stocky animals were awarded at shows, it seemed that breeders were unwittingly turning the Dutch dairy cow into a fancy breed unfit for its main economic purpose. It was high time, the scientists concluded, that fashion and fancy gave way to utility and rationality. Particularly the selection of bulls had to change, since the bull was half the herd, as the 1

A more extensive version of this paper will be published in the Journal of the History of Biology.

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saying went. Bulls should be subjected to progeny testing, and only proven bulls should be widely used as sires. Only then would breeding become a rational practice. 2 Frustrated as they may have been about what they perceived as the conservatism of the breeders, the scientific experts knew that, in the 1950s, the tide was already turning and that a reform of breeding in the sense they envisaged was on the way. In the following decades scientists were to acquire a key role in the business of cattle breeding. Progeny testing did indeed become standard practice, and the influence of conformation shows dwindled steadily. Bull shows, once the culminating points of the perpetual competition among the breeders, were eventually even abolished. While breeders began to lose their influence, the involvement of scientists increased. They worked out a system for progeny testing and helped making the plans for its implementation, which involved a drastic reorganisation of the plethora of organisations and institutions in the field of dairy farming and stock breeding. They also developed the statistical means to judge and rank bulls according to merit, and came up with an index for the exact economic profit to be expected from using a given bull as a sire. 3 The story of how scientists acquired a leading role in dairy cattle breeding is a fascinating one, but my intention in evoking their views of traditional breeding methods was to put a different set of questions in perspective that I would like to explore in this paper. To begin with, Dutch scientists had been claiming already since the beginning of the twentieth century that only progeny testing provided a rational basis for breeding. Why was it that this seemingly simple and sound advice was apparently not heeded by breeders for so long? And what breeding methods did they use then? As indicated, scientists in the 1940s and 1950s disputed the economic effectiveness of breeders’ practices. What were the breeders’ views in this matter? Why, for instance, would they prefer animals whose conformation seemed to have an adverse effect on their productivity? Finally, Mendelism had been around for more than half a century before Dutch geneticists became seriously involved in practical cattle breeding, which raises the question of what the relation between geneticists and breeders had been in the pre-war period. And what was it that conditioned the change in this relationship after the war? Questions of this kind, that broadly speaking concern the relation between theory and practice in agriculture, are receiving increasing attention from historians of plant breeding, as a recent thematic issue of the Journal of the History of Biology has underlined.4 Studies by, among others, Barbara Kimmelman, Paolo Palladino, Jonathan Harwood, Christoph Bonneuil and Thomas Wieland have shown that the role of hereditary theory in plant breeding practices in the early twentieth century was much more complex than was suggested by an earlier historiography that described the reception of Mendelism by practical workers in terms of either ‘successful 2

3 4

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Already in 1927 and 1928, dairy adviser C. Zwagerman had published a series of articles that foreshadowed parts of the later critique of breeders’ practices. Several other articles appeared in the 1930s, but the criticism only gathered steam in the 1940s. I will mention only a few characteristic examples: Zwagerman (1927, 1928); Hagedoorn (1928); Zwagerman (1934); Bosman (1935); Hagedoorn (1939); Hagedoorn (1941); de Jong (1943); de Jong (1947); van der Plank (1948); van der Plank and Hirschfeld (1950); Hoekstra (1957); de Groot and Bekedam (1957); Hoekstra (1958). General histories of dairy cattle breeding in the Netherlands that describe this development are Strikwerda (1998) and Bieleman (2000). ‘Special Issue on Biology and Agriculture,’ Journal of the History of Biology 39/2 (2006), with contributions by Jonathan Harwood, Barbara Kimmelman, Christoph Bonneuil, Thomas Wieland, Karin Matchett and Lloyd T. Ackert.

Heredity without Mendelism

application’ or ‘failed assimilation’ of its principles.5 Historical studies of animal breeding are still very scarce.6 Yet investigations of livestock breeding provide opportunities for instructive comparison, as I hope my analysis of cattle breeding will demonstrate. One of my conclusions will be that Mendelism was, for all practical purposes, irrelevant for pre-war Dutch cattle breeders, and that even scientists in this period agreed with this assessment.7 My example here will be the breeding of Friesian black and white dairy cows in the Netherlands. In the course of the twentieth century Dutch Friesians became the principal type of dairy cattle worldwide. The foundational stock of the American Holsteins, some 7500 animals, were imported from The Netherlands in the late nineteenth century as Dutch Friesians. 8 (Their having become known as Holsteins, soon after their arrival, seems to have been due to an inattentive American government official). In the United States and in Canada, the Friesians were valued for their high milk yield, and they were bred as a pure dairy type, mainly producing milk for consumption. In most European countries however the double purpose cow was preferred, producing meat and milk, as farmland was too expensive in Europe for the extensive land use required for raising beef cattle. After the breeding of Friesians in America had gathered steam, the U.S. and Canada on the one hand and several western European countries on the other became competitors on the world market for Friesians. In the end, the post-war trend towards specialisation would give the American pure dairy type a decisive edge: from the late 1960s onwards, a worldwide ‘Holsteinisation’ took place. Ironically, Dutch farmers nowadays also call their black and whites ‘Holsteins.’ Until the middle of the twentieth century, Dutch breeders of Friesians were among the leading promoters of the European double purpose type. They unabashedly marketed their animals as the world’s best dairy cows—a somewhat arbitrary qualification since even farmers in different European countries not unusually preferred a slightly different balance between meat and milk production. Still, Dutch dairy cows had an excellent reputation. They had the highest average milk yield in Europe and were valued for their harmonious and uniform conformation as well as for their adaptability to different climates and management regimes. Black and white breeding stock found its way to many countries in Europe, America, Asia, Africa and Australia. 9 Dutch farmers also did well in terms of the organisation of breeding: participation in herdbook registration and in breeding and milk recording associations was exceptionally high. 10 Developments in other countries were scrutinised in journals issued by the herd-books and in agricultural and dairy industry periodicals. Germany had more scientists working on breeding problems before the war, Scandinavian scientists were quicker to get a grip on practical breeding, and the quantitative genetics underlying the reform of progeny testing after the war was mainly 5 6 7

8 9 10

For references, see the contributions mentioned in note 3. See for instance Russell (1986); Cooke (1997); Wood and Orel (2001); Derry (2003); Orland (2003); Wood and Orel (2005). It should be added, however, that quite different circumstances conditioned poultry and pig breeding practices, for instance, so that more work is required to obtain a general understanding of the impact of Mendelism on animal breeding. For the history of the Dutch black and whites in America, see for instance Prescott and Price (1930); Mansfield (1985). Grothe (1993). Strikwerda (1998: 192).

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worked out by American scientists, Jay Lush prominent among them. 11 Yet all in all, it can safely be said that the Dutch case provides an illustrative example of European breeding practices before the advent of systematic progeny testing.

Type and tuberculosis Returning to the criticism leveled at the Dutch breeders by agricultural experts in the 1940s and 1950s, the first issue I shall address is the change in type that scientists worried about. While it seemed obvious to them that smallness and beefiness were undesirable in a dairy breed, most experts, then and in later years, seemed to have all but forgotten why Friesians had become so small and stocky over the years. For instance it was suggested (without supporting evidence) that smaller cows, while producing less, had been easier to maintain in the years of crisis before the war.12 Wieger de Jong, professor of animal husbandry at Wageningen Agricultural College (the only institution of its kind in the Netherlands) argued more plausibly that the decrease in size was a side effect of breeding for shows. In terms of procreation, the fate of a bull was decided on at an early age. Animals that matured early, i.e. acquired adult proportions rapidly and fattened easily, were preferred by herd-book inspectors and judges at bull shows, according to de Jong. As it happened, such qualities were more often found in relatively small bulls than in larger ones, which looked rather gawky in their younger years. Since small bulls won the prizes at shows, de Jong concluded, they had been systematically preferred as sires, which in the long run had resulted in a decrease in size of the breed as a whole.13 But then, one might ask, why should inspectors and judges prefer stocky animals in the first place, instead of the tall and lean dairy type that had characterised the Friesian breed in the late nineteenth century? Most scientists entertained no doubts about the answer to this question: fashion and fancy breeding must have been responsible.14 Yet pre-war records show that there was more to the change in type than scientists in the 1950s seemed to remember. Some background information is needed here. Until the middle of the nineteenth century, most regions in the Netherlands had had their own local type of dairy cows. Friesian black and whites were mainly to be found in the sea provinces in the north and west. Yet by the 1890s, black and whites began to replace the local breeds in regions in the south and east. The main reason for this was an increase in profitability of animal husbandry, which had started after the liberalisation of the export markets in many European countries around the middle of the century. The trend of focussing on animal husbandry was facilitated by the improving means of transportation, and it was reinforced by the sharp drop of grain prices in the 1880s, when American grains flooded the world market. Towards the end of the century, farmers on the many small mixed farms in the east and south of the Netherlands by and large concentrated their activities on the production of milk, meat (beef and pork) and eggs. Their 11 12 13 14

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In the Netherlands, facilities for scientific breeding experiments with cattle would become available only after the war. See de Boer and Strikwerda (1990: 11). Stapel (1988: 42, 67). De Jong (1943: 116); de Jong (1947: 8-10). See note 1.

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arable land was mainly used to produce fodder for their animals. Concentrates also became very cheap and were being fed in increasing quantities. Another major stimulus to dairy farming was the establishment of cooperative dairy factories. The creameries lifted a major restraint on the growth of dairy farms in that they solved the farmers’ problem of finding an outlet for their milk. While the number of dairy cows had been more or less stationary until the 1880s, their number rose from some 900.000 in 1890 to about 1.3 million in 1930. 15 Meanwhile, partly as a consequence of the crisis of the 1880s, the government gave up its nineteenth-century laissez-faire attitude with regard to agriculture and began to stimulate and support the improvement of breeding practices. Local and regional milk recording and breeding associations were established in quick succession. Engineers from Wageningen Agricultural College were appointed to act as advisers of these associations and to develop educational programs for the farmers. The herd-books expanded their activities from the mere registration of true-bred animals to the improvement of breeding practices. Finally, more and more creameries, following the example set by dairy factories in Friesland, provided an incentive by paying the farmers for their milk on the basis of its butterfat content. The percentage of butterfat turned out to be heritable and less dependent on environmental circumstances than milk yield, thus providing an excellent opportunity for selection.16 The pages of agricultural newspapers and weeklies such as Het Friesch Landbouwblad, Het Landbouw Nieuwsblad, De Veldbode and De Veldpost testify to the growing importance attached to dairy cattle breeding after 1900.17 Agricultural journalists, academic scientists and government breeding advisers regularly exchanged views on the aims and methods of breeding in such journals, and more and more reports appeared on conformation shows and on the accomplishments of breeding associations and individual breeders. The keen interest taken in the subject by the dairy farmers themselves can be gleaned from the exchanges in the questions and answers section of weeklies such as De Veldbode and De Veldpost. As a consequence of these developments, good breeding stock and particularly good bulls were in high demand in the early twentieth century. The most productive black and whites were traditionally to be found on the specialised dairy farms that exploited the vast natural pastures of the northern and western clay provinces, mainly Friesland and North-Holland, so one would expect the breeders in these regions to have experienced a golden age. They did not do quite as well as expected, however. In the late nineteenth century, the black and whites in Friesland and North Holland were big, gaunt, leggy, sharp-backed, narrow-chested and ewe-necked animals (fig. 1). ‘All milk, skin and bone,’ as a British commentator put it.18 They were milking machines indeed and it was precisely for this reason that thousands of Friesians from these provinces were exported as breeding stock in these decades. Most experts and farmers in the Netherlands however were agreed that Friesians 15 16 17

18

Broekema (1913); Knibbe (1993); van Zanden (1985). For the development of cattle breeding organisations in the Netherlands see for instance Löhnis (1911); Tukker (1924); van Adrichem Boogaert (1970). I will refer mainly to De Veldbode, a widely read weekly established in 1903 that continued to appeared during the whole period under investigation and that reported on all important events and dicussions related to cattle breeding. Its full title was De Veldbode, Geïllustreerd Weekblad voor Land- en Tuinbouw, Pluimvee- en Konijnenfokkerij en Bijenteelt. Stanford (1956: 61).

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could only be profitably exploited on the fertile soils in the sea provinces. These highly productive cows were delicate and demanded high quality food and careful management. This was no new insight. Already in the seventeenth and eighteenth century German buyers, for instance, had learned the hard way that Dutch dairy cattle were difficult to maintain. In the Berlin area the Friesians were for this reason taken care of by Dutch immigrant farmers. 19

Figure 1. Friesian black and whites of the pure dairy type in 1900. Source: K.N. Kuperus & Zonen, Eenige mededeelingen over den uitvoer van Frieschstamboekvee (Leeuwarden 1912), p. 30.

In the Netherlands, it was contended, the mixed farms on the sandy soils in the east and south provided an environment that was no less ‘foreign’ to Friesians. Farmers in these regions could not provide the quality foodstuffs required, and they had neither the means nor the time to provide the care the animals needed. Under such less than optimal circumstances Friesian black and whites were said to become weedy. After a few generations, they were no longer better milk producers than the local breed. Friesians had been bred exclusively for production, wrote herd-bookinspector Iman van den Bosch, one of the most respected experts of the time. This had affected their constitution, and thus they demonstrated the wrong-headedness of the much discussed ‘Zucht nach Leistung’ (selection for production), propagated by the German agriculturist Emil Pott.20 H.M. Kroon, zootechnical expert of Utrecht Veterinary College, agreed that Friesians ran 19

32

Orland (2003: 173-174). Eighteenth-century sheep breeders were also familiar with the problems involved in maintaining foreign breeds; see Wood and Orel (2001: 45-46, passim).

Heredity without Mendelism

the risk of becoming so ‘overbred’ that their functionality was jeopardised. A government report on the improvement of animal breeding similarly warned against the risks of one-sided breeding for production.21 The most damaging allegation of all was that Friesians, if not taken proper care of, were highly susceptible to tuberculosis, a widespread disease among dairy cattle at the time, but one that particularly affected the reputation of the Friesians. A German visitor at a national show remarked that if the conformation of Friesian cattle was not enough to make one suspicious, the constant coughing that could be heard in their stables would not fail to do so. 22 According to veterinarian A. van Leeuwen, German experts even considered Dutch cattle to be the most severely afflicted with tuberculosis word-wide, and Belgian buyers also complained that Friesians were unfit for their soils and often fell victim to the disease.23 In many regions of the Netherlands black and whites from Friesland came to be held in bad repute too. Seemingly healthy Friesian breeding stock was claimed to ‘degenerate’ in other provinces and then to succumb to T.B. Veterinarians compared the fine-skinned and weedy dairy type to the tuberculosis-prone ‘habitus phthisicus’ in humans, characterised by a weak frame and an almost translucent complexion. 24 There was wide agreement that the delicacy and extreme level of performance of the Friesians made them particularly vulnerable.25 A considerable group of farmers on the sandy soils therefore preferred dairying with the Dutch red and white cow, the traditional cattle of the regions along the major rivers, the Meuse, Rhine and IJssel, for short called MRIJ-cattle. These red and whites were more robust and sober, and thus better suited for the circumstances on small mixed farms. Their milk yield was not as high as that of the Friesians, yet they were better meat producers: they could be fattened more easily and the quality of their meat was better. Last but not least, they were claimed to be less susceptible to tuberculosis.26 A second alternative preferred by farmers outside the MRIJ-regions, was a more robust type of black and whites, to be found in the provinces of Groningen and South Holland, where thanks to the availability of agricultural waste products fattening had traditionally been more important than in Friesland, resulting in a preference for heavier animals. Like MRIJ-cattle, cows of this type were believed to be less susceptible to tuberculosis than Friesians. In the 1910s and 1920s, a group 20

21

22 23 24 25

26

Van den Bosch (1906). Pott developed his views in reaction to what was then called ‘Formalismus,’ i.e., selection for phenotypic traits with no demonstrable relation to production; see Pott (1899); Comberg (1984: 122, 336-339). Kroon (1913: 95-99); Löhnis (1911: 28, 46). In the decades after 1900, animal husbandry specialists A.A. ter Haar, A. van Leeuwen and E. van Muilwijk constantly warned readers of De Veldbode not to be misled by the high milk yields that Friesian farmers were able to obtain on their rich soils. Wageningen experts concurred with this view; see for instance de Jong and Koenen (1923). Animal husbandry textbooks contained the same message; see for instance Broekema (1913: 16); Kok (1919: 76); Dommerhold (1927: 10, 14-17). Ter Haar (1913). Van Leeuwen (1905); van Leeuwen (1923). Abbo-Tilstra (2002: 27, 146-147, 201). As in the case of their shortcomings in conformation, the susceptibility of Friesians to tuberculosis was pointed out time and again in agricultural journals and handbooks in the early decades of the twentieth century; see for instance Kroon (1913: 97); Bakker c.s. (1914: 133); Timmermans (1923: 12); van Leeuwen (1924); Dommerhold (1927: 10); van Leeuwen (1931). See for instance ter Haar (1919); ter Haar (1923); Kroon (1913: 107).

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of breeders of this type in Hoornaar in South Holland offered serious competition to the Friesian breeders of black and whites. The provincial rivalry sparked by this competition can be gleaned from the articles that one of the type’s promoters, agricultural journalist E. van Muilwijk, published in De Veldbode. He wrote, for example, that breeders in South Holland should beware of breeding with the effeminate aristocratic bulls from Friesland. For within a few generations, tuberculosis-prone, spiky offspring with a miserable constitution would be the result. 27 Animal husbandry adviser Jacq. Timmermans dared his readers to name a single Friesian bull that had done well in the southern province of Limburg. Imported Hoornaar bulls, on the other hand, had almost without exception improved the local breed in this province. 28

Figure 2. Dirk 4, representative of a more robust type of black and whites. Source: E. van Muilwijk, De preferente zwartbonte N.R.S.-stieren (Den Haag 1937), p. 83.

Figure 2 shows the most famous bull of the Hoornaar type: Dirk IV. For years, particularly in the late 1910s and 1920s, he and his male offspring were considered to represent the ideal type of sire for dairy farmers on the lighter soils. The fact that milk yields were lower than in Friesians was acknowledged but accepted as the price to be paid for a healthy breed. On a more general level herd-book-inspector Iman van den Bosch had argued already in 1906 that it was better to aim for reasonable milk yields with a high butterfat percentage than to strive for record yields of blue milk. Foreigners, he said, preferred milky cows with a sound conformation; cows that literally produced milk like water were undesirable.29 He had a point: English farmers in the 1910s and 1920s 27 28 29

34

Van Muilwijk (1919); van Muilwijk (1925). Together with veterinarian A. van Leeuwen, van Muilwijk promoted the Hoornaar type in De Veldbode for years. Timmermans (1919). Van den Bosch (1906: 597-598).

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described the Friesians as an ‘irrigation breed’ and as ‘mere water carts.’ 30 And most German breeders, according to C. Kronacher, a leading German expert, preferred animals that were more solidly built than the Friesian black and whites.31 Nevertheless, the ‘Dirk IV’ bloodline became less popular in the 1930s, probably because farmers became dissatisfied with the - by then - even less than mediocre milk yield of this type of cow. The most important reason for its dwindling popularity however was that the Friesian breeders of black and whites made a spectacular comeback in these years. They had taken the criticism of their type to heart and had been working to improve it since the 1900s. In the 1920s, Friesian farmers also began a vigorous campaign to eradicate tuberculosis among their animals. With the help of the Friesian black and white herd-book, the cooperative creameries and other provincial organisations, the first provincial animal health service in the Netherlands was established in Friesland in 1919. Other provinces would follow suit only after World War Two. As a result, the black and whites in Friesland would be the first to be declared free from tuberculosis in 1950. 32 A culmination point of the Friesian breeders’ efforts to improve their black and whites was the bull Adema 197, born in 1934 and bred by the reputed Knol brothers in the hamlet of Hartwerd. In the eyes of the cognoscenti this animal was the most glorious Friesian bull ever bred until then. Adema 197 was claimed to represent a type that adapted more easily to varying circumstances than the original Friesians. He was heavier, deeper, broad- and flat-backed, and more short-legged than his late nineteenth century forebears. Yet contrary to the rather crude Dirk IV, he retained the Friesian dairy type in his more elegant lines, supple skin and fine hair. Moreover, Adema 197 exuded ‘nobility,’ as the breeders called it, a term borrowed from horse-breeding of which no straightforward definition can be given. It was used by breeders to denote the extra phenotypic quality or beauty, to be appreciated only by the connoisseur, that distinguished the pick of the breed from animals that were merely phenotypically correct according to the standard of the breed. In the fifties, the significance of the term would give rise to extensive discussions in the herd-book journals. Yet whether it was a useful notion or not, no conformation expert would deny that Adema 197 was an icon of nobility. As to his production qualities, it turned out that Adema 197’s daughters’ milk yield was ‘satisfactory,’ while the milk had a high butterfat percentage. All in all, he thus represented the almost perfect bull according to pre-war criteria. As a foundational bull for what came to be called the ‘modern Friesian,’ he was the most influential Friesian sire for several decades. In the 1950s, there were few true-bred Friesian sires who did not have Adema 197 in their pedigree at least once.33 The modern Friesians, with Adema 197 as their harbinger, restored the breeders in Friesland to the leading position they had had at the end of the nineteenth century. In the fifties they even experienced the heyday of their fame, nationally and internationally. The Friesian herd-book flourished, the small circle of top breeders enjoyed an enormous prestige and their animals were much sought after and were sold for high prices. The five-yearly jubilee shows organised by the herd-book were events that attracted an international audience.34 Foreign buyers were 30 31 32 33

Stanford (1956: 50). Anon. (1928). Abbo-Tilstra (2002: 330). Strikwerda (1998: 96); Strikwerda (1979: 317-333).

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particularly impressed by the uniformity of the Friesian black and whites. They knew that the breeders gave priority to quality of conformation: in Holland, ‘high milk yields [are] not sought,’ a British herd-book official even stated categorically.35 Two other factors contributed to the success of the Friesian black and whites. After Friesland had set the example, bovine tuberculosis was eradicated in all Dutch provinces by the mid1950s.36 Secondly, the differences in fertility between the heavier and lighter soils in the Netherlands no longer obtained in the 1950s as a consequence of the use of artificial fertilisers and improved pasture management techniques.37 Thus, the major obstacles to the spread of the Friesian type had been removed. Still, the constitution of their animals continued to be among the breeders’ central concerns in the 1950s. It was their job, they argued, to safeguard the health and adaptability of the breed. To be sure, productivity came first for run-of-the-mill dairy farmers’ cows, but a different standard was needed for the breeding stock from which these cows were bred. Trade-offs between milk yield and conformation were inevitable, and to strive for uniform and harmoniously built animals was no mere luxury or fancy. A well-bodied cow was an economic cow, and the nobility that distinguished the best breeding stock was to be seen as an extra guarantee for the quality of their progeny.38 Against this background it will come as no surprise that breeders and herd-book officials in these years qualified the growing scientific criticism of their allegedly excessive preoccupation with the phenotypical qualities of their animals as out of place. In the breeders’ opinion, the commercial success of their modern Friesians alone may have sufficed to justify their view of the matter. For it was precisely for their conformation and uniformity that the Friesians were sought after. Moreover, details of conformation that lent nobility to an animal, whether functionally significant or not, were definitely important financially speaking. An animal’s ranking at shows and its score in the herd-book’s point system for conformation had direct consequences for its commercial value.39 For the breeders, beauty was the hallmark of health and adaptability as well as of marketability. And why change a breed that was so obviously successful? ‘This can’t go on forever,’ one of the Knol brothers is said to have admitted, only to add, however, that ‘it is what the farmers want.’40 The animal husbandry director of the ministry of agriculture remarked that while it was impossible to say whether the modern Friesian represented an advance in terms of efficiency of production, it was an undeniable fact that it sold better than the old type. 41

34 35

36 37

38 39 40 41

36

Strikwerda (1979: 64, 96-97, 253-257). Stanford (1956: 186-187). This is not to say that the ‘modern Friesian’ was accepted uncritically by foreign breeders. The hereditary qualities of a group of 57 bulls exported to Great Britain in 1950 gave rise to heated discussions among British breeders; see Anon., 1955. Hofman (1996). Accordingly the N.R.S. decided in 1954 that it was no longer necessary to judge female cattle from different soils in separate categories at shows, as had been customary until then; see Anema and Jepma (196010: 116). See for instance Jepma (1954); Jepma (1957); Anon. (1955); Anon. (1957a); Anon. (1957b); Jepma (1962). Strikwerda (1979: 255). Kroon (1997: 82). Rijssenbeek (1956).

Heredity without Mendelism

A middle ground in the discussion over breeding practices was taken by Wieger de Jong of Wageningen Agricultural College. De Jong had risen from the ranks in both practical and scientific circles. The son of a dairy farmer and a Wageningen graduate, he had worked as a regional breeding adviser and herd-book inspector before being appointed as director of the Dutch national cattle herd-book, the N.R.S., which serviced all Dutch provinces except Friesland. In 1947 he became Wageningen professor of animal breeding and chairman of the N.R.S. 42 Thus representing different constituencies, de Jong weighed the arguments from both sides against each other. De Jong pointed to the difficulties inherent in the notion of constitution. 43 Unquestionably, a healthy constitution was important, yet how were constitution and conformation related? Were short legs stronger, did heaviness indicate longevity, were sturdy animals really healthier? Only empirical studies could decide on such matters, he argued, and these had yet to be undertaken. Nevertheless, de Jong sympathised with breeders who strove for beauty in their animals. Breeding for fancy should not be simply condemned. For many farmers, the joy of breeding and even their happiness in life were bound up with their competitive efforts to create the perfect animal. Conformation shows provided the sporting ground to assess the level of their achievements, and not much would remain of the popularity of breeding without such incentives. While de Jong was undoubtedly correct that, for many farmers, breeding was a labour of love, we might add to his observations that, as Margaret Derry has demonstrated, it is quite impossible to separate the breeders’ esthetic ideals and their commercial considerations in shaping the perfect animal. Breeding for perfection and breeding for the market went hand in hand. 44 While sympathetic towards the breeders’ concerns, de Jong was no less worried about the productivity of the black and whites than his scientific colleagues. Already in 1943 he had shown that there was no correlation between the scores for conformation that animals were allotted by herd-book-inspectors and their milk yield.45 Apparently, the methods that the breeders and the herd-books used to evaluate and improve their animals were not conducive to the improvement of the productivity of the breed. Thus the question arises of what the breeders’ methods actually consisted in and what the underlying assumptions were.

Bloodlines and purity In the late nineteenth century, a growing number of dairy farmers in the Netherlands considered themselves to be not merely dairymen, but also breeders. (While not all dairy farmers were breeders, all breeders were also dairymen. Cows were too costly to raise and maintain merely for breeding purposes; their milk constituted an important part of the breeders’ income —the greater part, for most of them.46) In western Europe, Dutch cows had been well-known for their dairy

42 43 44 45 46

Dekker and Stapel (1976: 315-316). De Jong (1943); de Jong (1947); de Jong (1957). Derry (2003: 156-161). De Jong (1943). Minderhoud (1935: 126).

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qualities for centuries, yet breeding became particularly lucrative in the second half of the nineteenth century, when the fame of the black and whites spread world-wide. The American cattle traders who in the 1870s and 1880s bought thousands of Friesians provided an incentive for organised breeding in the Netherlands in that they stimulated the establishment of herd-books. An American importer, Holstein pioneer Solomon Hoxie, even acted as adviser of the Friesian herd-book founders, and he and several other American buyers became herd-book members.47 There is a pattern here, as Derry has shown: the establishment of herd-books indicates a rising international market.48 The guarantees on paper provided by herdbooks were especially important for American buyers. Whereas a Dutch farmer would never buy a cow that he had not inspected himself, New World geographical distances necessitated American farmers to rely on catalogues and certified pedigrees. What pedigrees had to prove, was ‘purity’ (zuiverheid, in Dutch). In the case of Arabian horses, for instance, purity was ascribed only to animals all of whose ancestors descended from horses that had been bred, literally, ‘in the desert.’ In Shorthorn cattle, purity referred to descent from the breeding stock of a very limited number of British breeders.49 Likewise, a pedigreed Friesian could be trusted to have descended from black and white ancestors bred in the Netherlands. The Dutch national herd-book (the N.R.S.) was established in 1874 and it registered animals belonging to what was then called the Dutch lowland breed, mainly comprising the black and white Friesian, the red and white MRIJ and the Groningen whitehead (the blaarkop).50 To enhance the exclusivity of their black and whites, breeders in the province of Friesland established their own herd-book in 1879, the F.R.S. (Friesch Rundvee-Stamboek). From then on, black and whites from bloodlines in other provinces were no longer accepted for registration in the Friesian herd-book, irrespective of their characteristics or qualities.51 The concept of purity was an ambiguous and contested one.52 For instance, the nineteenth century notion of constancy of a pure race (‘Konstanztheorie’) as propagated by the German horse expert Johann Justinus was based on the conviction that purity resided in an inborn potential of a true race to consistently and unchangingly pass on its defining characteristics through the generations, irrespective of the circumstances under which the animals were kept. 53 Many nineteenth-century practical breeders however knew from experience that the purity of their breeds could not be defined in such a strict sense, and Friesian breeders are a case in point. They were only too well aware that the ‘purity’ of their black and whites needed maintenance even under stable environmental circumstances. This was convincingly demonstrated by the irregular occurrence of red and white calves born from black and white parents. In its early years the Friesian herd-book made no bones about registering such calves and other off-coloured animals. 47 48 49 50 51

52 53

38

Strikwerda (1979: 81-86); see also van der Wiel and Zijlstra (2001: 32-35). Derry (2003: 156-161, passim). Conversely, around 1900, when the German, English and American markets were being closed for live cattle , the herd-books experienced a serious crisis; see Löhnis (1901). Derry (2003), chapter 2. For a history of the N.R.S., see Dekker and Stapel (1976). Still, although this was later denied by F.R.S. officials, there were a few isolated cases of Friesians born outside Friesland that had, in the early years of the F.R.S., been registered by the herd-book (Strikwerda 1979, 144). For histories of the F.R.S., see Zwart (1960); Strikwerda (1979). Derry (2003: 9, passim). Berge (1961: 131-134); Comberg (1984: 106ff); Wood and Orel (2001: 244-246, 264-266).

Heredity without Mendelism

They would soon be relegated to separate registers, however, to please the American buyers for whom a pure Friesian should be black and white.54 Meanwhile, Friesian breeders did believe that their black and whites represented a very old race that had been native to Friesland since prehistoric times.55 Crossing of different breeds, which was still common in other provinces at the time, was supposed to have been rather the exception in Friesland, and a ‘pure’ core of Friesians was supposed to have been preserved through the ages. Accordingly, the most stringent requirement for a Friesian to be accepted for registration concerned geographical provenance: the animal should descend from ancestors bred by Friesian breeders. In this way, the notion of purity functioned exactly as intended, namely to protect the interests of breeders in Friesland and their buyers. As we shall see, geneticists would translate purity into Mendelian terms after 1900, yet the purity concept had connotations of exclusivity and quality that Mendelism could not capture. An example is provided by a veritable cause célèbre in Dutch cattle breeding, the so called coloured spots question (vlekjeskwestie). In the middle decades of the nineteenth century a number of Shorthorn bulls were imported in the Netherlands from the U.K. Some agricultural experts believed that they might improve the beef quality of Dutch cows. The experiment was soon terminated however, because the milk yields of cross-breds turned out to be disappointingly low. Traces of Shorthorn influence remained visible for some time in the colouration of some herds, but these progressively disappeared when the cross-breds were bred up to the original Dutch type for several generations. The idea took hold, however, among breeders and their customers alike, that isolated coloured spots on the lower legs were an indication of lingering Shorthorn influence and for this reason highly undesirable. Animals with spots on the phalanges were in the 1920s even excluded from registration by the F.R.S.56 Until far into the twentieth century agricultural experts and scientists would spill gallons of ink trying to convince the herd-books that excellent animals were for no good reason kept out of breeding programmes in this way. In their view there was no evidence whatsoever that the spots derived from Shorthorns, while, more importantly, a cow’s productivity was in no way affected by the presence of such spots.57 It was to no avail, however. In 1912 the well-known breeder A.D. Groneman conceded that the experts might well be right, yet that breeders had different concerns: buyers, especially foreign traders, wanted pure black and whites, and pure black and whites were not supposed to have spots.58 The herd-books acted accordingly and did not change their policy. Clearly, purity referred to a breed standard that could not be compromised, irrespective of whether a deviation from the standard was genetically or functionally significant or not. At issue here was not a genotype but a commercial ‘brand.’59

54 55

56 57 58 59

Strikwerda (1979: 31-36). See also Dekker and Stapel (1976: 256-267). Bakker (1909) contested this view and argued that the original Friesians had been red and whites, the black and whites having been imported from Denmark after the onslaughts of the rinderpest in the eighteenth century. See for instance Strikwerda (1979: 109-116). A. von Leeuwen, for instance, campaigned for years on end against the exclusion of animals with spotted legs in De Veldbode. See for instance van Leeuwen (1914). Groneman (1912). Dog and horse breeding provide similar examples; see Derry (2003: 158, passim).

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In order to maintain the desirable qualities in their herds, breeders employed methods that, as Roger Wood and Víte£lav Orel have recently underlined, had been common practice among experienced breeders since the eighteenth century.60 Breeders knew that the best strategy to maintain the desirable qualities in their stock was to breed the animals among themselves. In its most strict form, this amounted to inbreeding, which was indeed practised intensively by all experienced breeders of Friesians.61 Even parent-offspring and sibling matings were not shunned. Adema 197, to give but one example, was the product of a mating between siblings; he had only one grandfather while his grandmothers were related as aunt and niece. 62 Meanwhile, the risks of inbreeding were well-known. Offspring had to be selected carefully and some outbreeding might be unavoidable once in a while. Still, the ideal of uniformity, in the breeders’ opinion, could only be reached by close inbreeding. The best breeders created their own ‘bloodlines’ in this way, and these were considered to buttress the quality of the breed as a whole. 63 An additional advantage of breeding in bloodlines was that the herds of the top breeders were not only very uniform, but also slightly different between them, as a consequence of minor variations in conformation in each herd that were consolidated through inbreeding. This subtle distinctiveness was a commercial asset, as it enabled the breeders and their customers to recognise the animals from the better herds. For instance, buyers knew that breeders in North Holland produced black and whites of a slightly larger and milkier type than those in Friesland, while breeders in North Holland liked some of the Friesian bloodlines but not others. 64 An new method for assessing the quality of dairy cows was introduced in the 1890s, after the example of Danish dairy farmers, namely the systematic recording of milk production. Friesland led the way and would remain the province with the highest participation in milk recording. By carefully weighing a cow’s milk yield on a regular basis its yearly production could be estimated, and the figures thus obtained could also be used to assess the hereditary quality of the cows’ sires. Milk recording included measuring the milk’s butterfat content. After Friesian creameries had, again, set the example, farmers in more and more regions of the Netherlands were paid for their milk on the basis of butterfat content. Selection for butterfat content became the primary focus of selection for the breeders in Friesland in particular.65 An instrument to raise the interest in the improvement of breeding methods that had been introduced around the middle of the nineteenth century yet that acquired a much more prominent role in the twentieth century, was the organisation of agricultural exhibitions and conformation shows. The increasing number of local, regional and national shows that breeders associations, agricultural organisations and the herd-books organised after 1900 testify to the growing popularity of the breeding of purebreds as well as to the commercial interests behind it. For as already indicated, show prizes earned breeders money: after a successful show, sales of their stock would immediately pick up.

60 61 62 63 64 65

40

Wood and Orel (2001) chapters 3 and 4. See for instance Hoogland (1921). Strikwerda (1979: 317). Some famous bloodlines were described in monographs; see for instance van Muilwijk (1935). For the history of cattle breeding in North Holland see Kroon (1979); van der Wiel and Zijlstra (2001). Strikwerda (1979: 65-80).

Heredity without Mendelism

There were several other tools that were employed by breeding organisations and the government to rationalise the farmers’ methods to improve their stock. We shall take these in stride in the analysis of the impact of Mendelism on breeding practices.

Mendelism For Amsterdam botanist Hugo de Vries, one of the ‘rediscovers’ of the Mendelian theory of heredity, the improvement of plant breeding and agriculture had been the principal motive for investigating hereditary phenomena, and he considered Mendel’s laws to be directly relevant for the breeding of agricultural varieties.66 While the possible implications of Mendelism for agriculture were thus pointed out from the start, Dutch animal husbandry experts were more hesitant in confronting Mendelian genetics with their field. The subject began to receive serious consideration only in the 1910s, when Mendel’s rules were explained in several monographs and articles.67 Even then, the authors took most of their examples from botany. Examples from livestock breeding only involved very simple Mendelian phenomena, mainly relating to coat colour in farm animals. For example, veterinarian A. van Leeuwen, the stock breeding expert of De Veldbode, after having expressed his reservations about the general validity of the theory, inquired among his readership whether anyone had ever bred a black-and-white cow from red-and-white parents. A group of farmers responded that they had never come across such a combination; only a single farmer believed that he had. Van Leeuwen concluded that alternative explanations could not be ruled out, yet that there was indeed support for interpreting the red colour as a Mendelian recessive trait.68 The presence or absence of horns appeared to fall into the same category, and before long, the more difficult example of coat colour in horses also turned out to be amenable to a Mendelian explanation. In 1910 geneticist Arend L. Hagedoorn, a pupil of Hugo de Vries and Jacques Loeb, was invited by the Dutch Agricultural Society to assist in designing breeding strategies for the improvement of Texel sheep.69 Breeders had been hybridising this breed with English races such as the Lincoln and the Wensleydale for several decades. Aiming for a uniform new type, they were struggling to get rid of unwanted fleece, head and nose colours. Hagedoorn helped them by devising breeding schemes along Mendelian lines. Although his efforts were not unsuccessful, the project was discontinued after some time because of the complexity - and consequently the rising costs - of test-mating and culling.70

66 67 68 69

70

On de Vries, see for instance Stamhuis, Meijer and Zevenhuizen (1999). For the motives underlying his research, see Theunissen (1994). See for instance Hagedoorn (1912); Waardenburg (1913); Giltay (1914); Lotsy (1915); Reimers (1916). Van Leeuwen (1912). Arend Lourens Hagedoorn (1885-1953), animal geneticist and evolutionary theorist, deserves more attention from historians than he has received until now. Basic information on his life and work (in Dutch) can be found in a commemorative issue, published shortly after his death, of the journal of the Dutch Genetics Society, Erfelijkheid in Praktijk (1954). Hagedoorn (1911); Kroon (1917: 43).

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This example illustrates the problems inherent in the application of a Mendelian approach to livestock breeding as opposed to plant breeding. As Wageningen animal husbandry specialist J. Reimers pointed out in 1916, experimenting with plants was easier because they could be selffertilised, and seeds and plants were cheap and could easily be obtained as well as dispensed with in large quantities. Individual animals, on the other hand, especially the larger farm animals, represented a significant economic value and produced far less progeny, and therefore the costs of experiments with animals quickly became prohibitive.71 Deliberately trying to produce even a single—undesirable—red and white calf, for instance, was not something a breeder of Friesians would readily do for experimental reasons, let alone that he could be induced to experiment with several detrimental recessive factors. Moreover, we have only been discussing qualitative characters so far. The economically most important characters of livestock however, such as milk and meat production, are of the quantitative kind. Even experts who were convinced that such characters could also be explained in Mendelian terms, had to admit that in this case the practical application of Mendelian theory was virtually impossible. According to Reimers a quantitative character such as milk yield might be accounted for by assuming that a group of similar Mendelian factors was responsible for the trait. Yet even if a Mendelian breeding scheme, based on this assumption, could be devised to improve milk yields, the complexity and costs of such a programme presented great difficulties. Hagedoorn remarked that breeders of farm animals, contrary to plant breeders, would learn nothing of practical use from a visit to the Svalöf experiment station in Sweden. 72 Accordingly, while Hagedoorn would become a well-known expert in animal genetics, he conducted his experiments with small laboratory animals such as mice. As to the economically important animals, he confined his investigations to animals that were inexpensive, could be kept in relatively large numbers and produced reasonable numbers of offspring, such as chicken and, occasionally, rabbits. Hagedoorn entertained no doubts that the rationality or irrationality of traditional cattle breeding methods could be decided on in Mendelian terms. He was well aware however that a Mendelian reform of breeding strategies was an entirely different matter. Little was known about the genetics of quantitative characters, but there were definitely too many genes involved to be handled by simple Mendelian crossing procedures. Consequently, traditional breeding methods would be indispensable for a long time to come. In 1927 Hagedoorn stated that the influence of genetic theory on cattle breeding practices had been negligible, and in his wellknown Animal Breeding of 1939 he even wrote that the influence had rather been the other way round: geneticists had learned a lot from the best breeders. What geneticists had to offer to the breeders of large farm animals was of a different nature: ‘The geneticists’ main contribution to animal breeding is not an analysis of genes, but an analysis of breeding methods. 73 This view was widely shared among Dutch animal husbandry experts at the time. 74

71 72 73 74

42

Reimers (1916: 2, 27); see also Hagedoorn (1912: 5-6). Reimers (1916: 27, 37-38, 78); Hagedoorn (1912: 83). Hagedoorn (1927); Hagedoorn (1939: 19). Broekema (1913); van Leeuwen (1923); Compte-rendu (1923: 53-58); van Muilwijk (1928); Overbosch and van der Plank (1931). See Derry (2003: 12-13) for a similar assessment with respect to the role of classical genetics in horse and dog breeding.

Heredity without Mendelism

What did the assistance that geneticists might give according to Hagedoorn consist in? To begin with, geneticists and agricultural experts concurred with the breeders that inbreeding was a rational strategy. The haphazard crossing of breeds that had been customary among small farmers until the late nineteenth century had resulted in motley collections of animals with unpredictable and widely varying qualities.75 No improvement was possible in this way, and the national herdbook had been right, in 1906, to have formally subdivided the ‘Dutch lowland race’ into three clearly delineated breeds, the black and white Friesian, the red and white MRIJ and the Groningen whitehead.76 But even stock improvement within clearly defined breeds remained something of a lottery as long as bulls of different provenance were used every few years. It was much better, the scientists agreed with the top breeders, to start from a group of excellent animals and to consolidate their qualities in a closely inbred herd. Purity, translated into Mendelian terms, meant homozygosity, and inbreeding increased the degree of homozygosity. Therefore, inbreeding was a rational strategy of breed improvement, provided it was accompanied by scrupulous selection against unwanted recessive traits. Experts explicitly advised against needless outbreeding with unrelated animals. Animals imported from other regions might not adapt well to local circumstances —as the example of Friesians deteriorating on poor soil discussed above illustrated. Moreover, a bull from a unrelated herd with a long history of its own was bound to be different, genetically speaking, in many characters. Recombination would bring these differences to the surface in the second generation, and thus the achievements of years of careful inbreeding and selecting might be undone.77 At the same time, however, experts also warned breeders not to overestimate the value of pedigrees. Obviously, the productivity of his ancestors should play a role in the choice of a bull. Yet it was of little use to study more than a few generations of an animal’s ancestry. From a Mendelian perspective, it was more instructive to look at a bull’s brothers and sisters, since they provided more reliable insights into his genetic strengths and weaknesses than remote ancestors whose contributions to his genes was insignificant.78 It is difficult to say whether practical breeders took heed of this advice. Yet a cursory survey of herd-book journals and histories of cattle breeding suffices to conclude that where the market for breeding stock was concerned, the preoccupation with pedigrees continued at least as long as inbreeding remained the principal breeding method and as long as a breeder’s reputation was inseparably bound up with the reputation of his bloodlines. For instance, until well after World War II items on individual breeders in herd-book-journals invariably included detailed information on the pedigrees of the foundational animals of their herds. The national herd-book published several illustrated genealogies of the most prestigious bloodlines, and a detailed description of bloodlines constituted the pièce de résistance of herd-book histories.79 Knol Bros. even had the history of their stock-farm and bloodlines privately published. 80 Again, pedigree, like 75 76 77 78 79 80

Kroon (1913: 71, 121); Kroon (1917: 24). Van den Bosch (1906). Kroon (1913: 102); Hagedoorn (1912: 57-64); Lotsy (1915: 15-17, 33); Reimers (1916: 95); Bakker (1926); Hagedoorn (1927: 54, 87-95). Reimers (1916: 89); Hagedoorn (1912: 47-48); van der Plank (1940). See notes 46, 49 and 50. Van Popta (1965).

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purity, was not merely about genes. Famous ancestors, however remote, continued to lend prestige to their bloodline. In a herd’s history resided its quality and distinction. There is an obvious contrast here with the Mendelian interpretation of purity: as soon as a breed becomes pure in Mendelian terms, i.e. homozygous, its history becomes irrelevant. Mendelian breeders may in a sense be said to aim for the elimination of a herd’s history. Another scientific critique of breeders’ practices that could frequently be heard, was that the herd-books were more attuned to the breeders’ commercial interests than to cattle improvement. Herd-books, it was argued, might serve as invaluable tools. Much might be learned, for instance, about hereditary diseases, if only the herd-books would register all descendants of pedigreed animals and would also record their genetic peculiarities.81 It takes no stretch of the imagination, however, to realise that breeders could muster up no enthusiasm for such suggestions. Firstly, they were charged for registration of their animals, so they offered only the best ones for inspection. 82 Secondly, for obvious reasons most breeders preferred malformed progeny from their prizewinning animals to disappear without a trace. The herd-books would lose all their members if complete registration became compulsory, an expert admitted. 83 Thus in the pre-war period at least, the-herd books did not develop into the instruments for rational breeding that the scientists would have liked them to become. By far the most often-heard advice that scientific experts tried to press upon farmers was that rational breeding should be based on progeny testing. Conformation and pedigrees were helpful to find a promising young bull, yet ultimately, it was the performance of his daughters as dairy cows that determined the true value of a sire. Therefore, rational breed improvement required the systematic use of older, tested bulls. From the early years of the twentieth century onwards, experts and animal husbandry advisers tried to drive this message home to the readers of agricultural weeklies and farmers’ handbooks. Hagedoorn relentlessly repeated the message in his scientific and popular publications. Ideally, he added, promising bulls should be tested on a limited number of cows first, and only the best ones should then be widely used as sires. 84 In this case, there is no evidence that the breeders disagreed in principle. Yet again, meeting this demand for rational breeding in practice was a different matter. The ideal situation as sketched by Hagedoorn was impracticable in every respect in the pre-war period. Farms in the Netherlands were small and few farmers milked more than ten cows. For instance, in 1920 the 953 organised farmers in the province of Limburg owned 2990 cows; in the Netherlands as a whole, an average number of ten cows per farm would only be reached in the 1950s. 85 Bulls were costly to maintain, and bull-calves only increased in value until their second or third year. Therefore farmers who could afford a bull of their own as a rule bought a young bull-calf, used it for a year or two and then sold it for slaughtering.86 Thus by the time their daughters began to give milk and their real worth became apparent, most bulls were dead. 81 82 83 84 85 86

44

Reimers (1916: 81, 93); Hagedoorn (1912: 48); Hagedoorn (1927: 130-137). In 1940, for instance, the registration costs of an animal were five guilders; a farm hand at the time earned about fifteen guilders a week (Kroon, 1998: 118). Compte-rendu (1923: 51-53). See also Wibbens (1923: 306-330). Hagedoorn (1912: 47-48; 86, 88); Hagedoorn (1927: 63). For the early decades of the twentieth century see also, for instance, Reimers (1916: 79, 92-93); Kroon (1913: 99); van Krimpen (1905: 13). Timmermans (1920: 615-616); Strikwerda (1998: 67). See for instance van Leeuwen (1904); Wibbens (1907); Löhnis (1911: 46).

Heredity without Mendelism

Farmers might also use a sire owned by a breeder for their cows. Yet if distance and the difficulty of transportation did not preclude such an option, it was, more often than not, the prices that breeders charged for insemination that put farmers off. Around 1910, prices varied between 25 cents and 20 guilders, and the reputed breeder F.A.F. Groneman experienced that small farmers were not prepared to pay the 2 guilders he charged for an insemination by his service-bull. 87 Since the late eighteenth century small farmers in many regions of the Netherlands had traditionally shared a bull purchased with municipal support. There were fine animals among them, yet many poor ones too.88 After 1890, more and more farmers began to organise themselves in breeding associations which enabled them to buy better bulls. Government premiums helped them to keep the good ones for a longer period.89 While some of these associations managed to improve their stock in this way, others fared less well and were discontinued after a number of years. There were indeed many obstacles to be overcome: farmers had to agree on the type of bull to be purchased; after several years of use, father-daughter inbreeding became unavoidable; a shared bull might spread venereal diseases; older bulls might become dangerous or too fat to perform; and the progeny of even an expensive bull could turn out to be disappointing. 90 On the other hand, once a breeding association had acquired a certain reputation, farmers might be tempted to buy their own bull and start their own stock-farm to get a share in the breeding market.91 Besides such complications, the number of cows serviced by a bull exploited by a breeding association was still relatively small, and a reliable assessment of his productive qualities was in most cases only possible after his death. Most bulls that, after a thorough investigation of their offspring, earned the much-coveted title of preferentschap, indicating proven hereditary excellence, were no longer around to receive the honours or were at best near the end of their period of service.92 The ideal situation as envisaged by Hagedoorn and his scientific colleagues, in which a number of young bulls was tested before the best of them—by that time having reached the age of at least five or six—seriously began their tour of duty, was beyond the means, financially and practically, of even the most prosperous breeding associations. In 1941, after Hagedoorn had in a lecture once again underlined the importance of systematic progeny testing, N.R.S. chairman H.W. Kuhn responded that Hagedoorn was apparently ignorant of practical cattle breeding: breeders could not possibly implement such a system, for both practical and economic reasons. 93 And animal husbandry adviser R.G. Anema predicted that current practices would probably not change for a hundred years to come.94 Kuhn’s was a correct assessment of the pre-war situation, yet as to the future he and Anema soon turned out to have been too pessimistic. Progeny testing would become feasible within a few years, after the introduction of artificial insemination in the early 1940s. Interestingly, AI was initially developed to fight venereal diseases causing infertility and spontaneous abortions, which 87 88 89 90 91 92 93 94

Löhnis (1911: 46); Groneman (1956: 37). Van der Wiel en Zijlstra (2001: 57-61). Löhnis (1911: 3); van Adrichem Boogaert (1970: 303-305). Nobel (1912: 10-11); van der Wiel en Zijlstra (2001: 99-109, 145-146). This happened in North Holland, for instance; see Van der Wiel en Zijlstra (2001: 146). Strikwerda (1998: 114). Hagedoorn (1941); Kuhn (1941). Anon. (1941).

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were rampant in the inter-war years, but scientists were quick to realize the potential of AI as an enabling technology for progeny testing. AI opened up the possibility to test a young bull on hundreds of cows at the same time. Years were thus taken off the time formerly needed to assess his hereditary qualities. Further, AI enabled the use of proven sires on an unprecedented scale, and consequently far less bulls were needed than before. This implied a switch from breeding in bloodlines to breeding in populations, which in turn required drastic changes in the organisational structure of dairy cattle breeding. In the process, scientists were to take the lead in breeding matters, while the breeders were slowly but surely relegated to the side-line. 95 This transformation did not take place overnight—it took several decades. In the 1950s, when the ‘modern Friesian’ was at the height of its popularity, the breeders, particularly those in Friesland, held on to their breeding methods and to their convictions about the ideal Friesian type. This was not merely because of the obvious threat that AI posed to the market for bulls, but also, as indicated, because breeders opposed the exclusive focus on milk yield that in their view was part and parcel of the scientists’ pleas for systematic progeny testing. In due course, however, postwar economic pressures forced farmers to scale up, intensify and specialise their farms, and as a consequence the traditional double purpose cow lost more and more ground to the specialised dairy type.96 This played into the scientists’ hands, since the changeover to the pure dairy type favoured bulls that had been tested for high yields. Eventually, in the 1970s and 1980s, the trend towards specialisation would even lead to the demise of the Friesian black and whites and their replacement by their American relatives, the Holsteins, which had been bred purely for production since the 1880s and had left their European ancestors far behind in terms of milk yield. The details of these postwar developments are beyond the scope of this paper. Here I have merely mentioned them to indicate the context of the scientists’ criticism of breeding practices in the 1950s. Circumstances were changing rapidly in those years, and in their campaign for a new approach to cattle breeding scientists all but ignored the conditions on which breeding methods up till then had been predicated. We can now draw some conclusions with respect to what these pre-war methods entailed and how they related to Mendelian theory.

Discussion and conclusions By and large, breeders and animal husbandry experts in the pre-war years entertained comparable views on the best methods for breeding dairy cattle, even though some experts became increasingly critical about the relative weight breeders attached to conformation and pedigrees. Where their opinions diverged, commercial considerations on the breeders’ part were often involved: market demands were not always ‘rational’ from the experts’ point of view. For instance, while some details of conformation might not be demonstrably relevant for milk yield, they did make a difference on the market for breeding stock. Breeders and experts further agreed that selecting animals purely for production was unwise, since Friesians of the extreme dairy type were difficult to maintain and less resistant to disease, particularly tuberculosis. 95 96

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On the development of AI in the Netherlands, see Strikwerda (2007). On these postwar economic pressures, see for instance van der Molen and Douw (1975: 9-35).

Heredity without Mendelism

The breeders’ principal method, which consisted in striving for purity by mains of inbreeding and breeding in bloodlines, was deemed perfectly rational by the experts. Progeny testing and the use of proven sires were propagated widely as indispensable for rational breeding, yet most experts were well aware of the limitations that practical realities set to the implementation of this advice. Geneticists generally considered the breeders’ methods to be consistent with Mendelian theory. The theory confirmed the rationality of aiming for purity and of inbreeding. At the same time however the experts readily admitted that the Mendelian insights into heredity added little of practical value to what breeders already knew. The importance of progeny testing was recognised long before the war. It might be added that the basic idea of progeny testing was not a new insight. It was hinted at already in the biblical phrase that ‘the tree is known from its fruit,’ and some breeders in antiquity were definitely aware that the value of breeding stock was to be gauged from its offspring, as Nicholas Russell has shown.97 The work of eighteenth century sheep breeders such as Bakewell also reflects this principle. This is not to say that systematic and controlled tests were developed already before the twentieth century; there is no convincing evidence for this, not even in the case of Bakewell and his followers.98 On a wide scale, systematic progeny testing became feasible only after the introduction of AI. Mendelism had nothing to do with this. In fact, progeny testing is not predicated on any specific theory of heredity. Besides the availability of AI, its successful implementation after the war rather asked for sophisticated statistical methods than for specific theoretical knowledge about genes or chromosomes and their behaviour. From the late 1950s onwards, breeding methods began to change from breeding in bloodlines to breeding in populations. Accordingly, breeding experts in the Netherlands began to call themselves population geneticists. They should rather have called themselves quantitative geneticists however, since nothing was as yet known of the genes involved in milk production, nor was such knowledge needed for their statistical analyses of milk production through the generations. While these scientists could rightly claim they had taken over the leading role in breeding from the breeders and had turned breeding into a scientific enterprise, the history of classical genetics is of little help in explaining how this came about.

Bert Theunissen Institute for the History and Foundations of Science Utrecht University [email protected]

97 98

Russell (1986: 25). Russell (1986: 204-205, 211).

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References Abbo-Tilstra, Bartje. 2002. Om de sûnens fan it Fryske folk. Tuberculose en haar bestrijding bij bevolking en veestapel in Fryslân, 1890-1940. Leeuwarden: Fryske Akademie. Adrichem Boogaert, D.H. van. 1970. De ontwikkeling van de Nederlandse rundveehouderij in deze eeuw. Een historisch overzicht over de periode tot 1970. s.l.: Ministerie van Landbouw en Visserij. Anema, R.G. and J. Jepma. 1960. Veeteelt-I. Rundvee, varken, schaap, geit en fokleer. Zwolle: Tjeenk Willink10. Anon. 1928. “De nationale tentoonstelling IV. Het oordeel van de Duitsche deskundigen.” De Veldbode 26: 63-865. Anon. 1941. “Over verandering van principe bij de rundveestamboeken.” De Nieuwe Veldbode 8: 8-10. Anon. 1955. “De fokwaardebepaling van stieren van de praktische kant gezien.” De Keurstamboeker 3: 204205. Anon. 1955. “Uit de Britse zwartbontfokkerij.” De Keurstamboeker 3: 521-522, 547-549. Anon. 1957a. “Het keuren van rundvee en wat daarmee samenhangt.” De Keurstamboeker 5: 79-80. Anon. 1957b. Naschrift. De Keurstamboeker 5: 580-581. Bakker, D.L. 1909. Die Niederländische Rindviehzucht. Maastricht. ———— c.s. 1914. Gedenkboek der nationale en internationale landbouwtentoonstelling te ’s-Gravenhage in 1913. Maastricht. ————. 1926. “Over het wezen en de betekenis van inteelt.” De Veldbode 24: 766-767. Berge, S. 1961. “Die geschichtliche Entwicklung der Tierzüchtung.” In E. Schilling (ed.). Wissenschaftliche Grundlagen der Leistungsermittlung und Zuchtverfahren bei landwirtschaftlichen Nutztieren. Schriftenreihe des MPI für Tierzucht und Tierernährung. Sonderband 1961: 127-148. Bieleman, Jan. 2000. “De georganiseerde rundveeverbetering.” In J.W. Schot c.s. Techniek in Nederland in de twintigste eeuw, vol. 3, Landbouw, voeding. Zutphen: Walburg pers: 131-153. Boer, H. de and R. Strikwerda. 1990. “Zestig jaar Nederlandse Zoötechnische Vereniging.” In R.D. Politiek (ed.). Markante ontwikkelingen. Nederlandse Zoötechnische vereniging 1930-1990. Wageningen: Nederlandse Zoötechnische Vereniging: 11-26. Bosch, Iman G.J. van den. 1906. “Stierhouderijen en fokvereenigingen.” De Veldbode 4: 585-588, 597-598, 633-634. ————. 1906. Kort overzicht met bijlagen van de resultaten van het in de jaren 1905 en 1906 ingesteld onderzoek omtrent de Nederlandsche rundveefokkerij in verband met de reorganisatie van de Vereeniging. Het Nederlandsch Rundvee-Stamboek. Amsterdam. Bosman, K. 1935. Onderzoekingen omtrent de melkproductierichting in de fokkerij van het Nederlandsche zwartbonte vee. Wageningen. Broekema, C. 1913. Nederlandsche rundveeteelt. ’s-Gravenhage. ————. 1913. “De gehouden wedstrijd betreffende de organisatie en werking van fokvereenigingen.” De Veldbode 11: 963-965. Broekema, L. 1913. “Rundveeteelt. In Directie van de Landbouw.” De Nederlandsche Landbouw 1813-1913. ’s-Gravenhage: 346-379. Comberg, Gustav. 1984. Die Deutsche Tierzucht im 19. und 20. Jahrhundert. Stuttgart: Ulmer. Compte-rendu des travaux du Congrès international pour l’élevage de l’espèce bovine ( …) (s.l. 1923). Cooke, Kathy J. 1997. “From Science to Practice, or Practice to Science? Chickens and Eggs in Raymond Pearl’s Agricultural Breeding Research, 1907-1916.” Isis 88: 62-86. Dekker, H.W.J. and K.P. Stapel. 1976. 100 Jaar Koninklijke Vereniging het Nederlandsch Rundvee-Stamboek, 1874-1974. s.l., s.n. Derry, Margaret E. 2003. Bred for Perfection. Shorthorn Cattle, Collies, and Arabian Horses since 1800. Baltimore & London: Johns Hopkins University Press. Dommerhold, E.J. 1927. Veeteelt. Zwolle. Erfelijkheid in Praktijk 15 (November 1954) nr 4/5. Giltay, E. 1914. Het principiële uit de Mendel-leer en haar konsequenties. Voordracht voor de Groninger Landbouwbond. Wageningen. Groneman, A.D. 1912. “De vlekken-kwestie.” De Veldbode 10: 805-806. Groot, Th. de and M. Bekedam. 1957. “De melkproductievererving van Friese stieren.” De Keurstamboeker 5: 138. Grothe, P. 1993. Holstein-Friesian, een wereldras. Doetinchem: C. Misset. Haar, A.A. ter. 1913. “Het oordeel van een Duitscher over ons vee op de Haagsche tentoonstelling.” De

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Veldbode 11: 997. ————. 1919. “Veeverbetering op lichten bodem.” De Veldbode 17: 489-490. ————. 1923. “Roodbont vee.” De Veldbode 21: 875. Hagedoorn, A.L. 1912. Oordeelkundige zaadteelt en fokkerij. Middelharnis. ————. 1927. Handboek voor fokkerij en plantenteelt voor Nederland en koloniën. Maastricht. ————. 1927. “De toepassing van wetenschap, in het bijzonder van erfelijkheidswetenschap.” De Veldbode 25. unpaginated Christmas issue. ————. 1928. “De richting van de rundveefokkerij.” De Veldbode 26: 370-372. ————. 1939. Animal Breeding. London. ————. 1941. “Rationeele fokkerij.” De Nieuwe Veldbode 8: 10-13. Hoekstra, P. 1957. Levensduur en rundveeteelt. s.l. ————. 1958. “De waarde der exterieurkeuring voor de Nederlandse rundveefokkerij.” De Keurstamboeker 6: 342-343, 370-371. Hofman, J. 1996. “Het succes van de tuberculose-bestrijding bij rundvee in Nederland, in het bijzonder in de periode na 1945.” Argos 14: 143-152. Hoogland, D.M. 1921. Een studie over familieteelt in de rundveefokkerij. Rotterdam. Jepma, J. 1954. “Enkele gedachten over de veefokkerij.” De Keurstamboeker 2: 274-275. ————. 1957. “Gedachten over veefokkerij.” De Keurstamboeker 5: 314. ————. 1962. “Gedachten over veefokkerij. Nogmaals: de fokrichting.” De Keurstamboeker 10: 251-253. Jong, W. de. 1943. “Productie en exterieur van ons rundvee.” Cultivator: 114-124. ————. 1947. Een halve eeuw productieteelt. Enkele beschouwingen over de toepassing en de resultaten van de melkcontrole bij de rundveefokkerij in Nederand. Wageningen. ————. 1957. “Geluk, ervaring, wetenschap.” De Keurstamboeker 5: 109-111. ————. and S. Koenen. 1923. “Welke eigenschappen dient het vee te bezitten om onder bepaalde bedrijfsomstandigheden te voldoen?” In Compte-rendu des travaux du Congrès international pour l’élevage de l’espèce bovine. ’s-Gravenhage: 599-612. Knibbe, Marijn. 1993. Agriculture in the Netherlands 1851-1950. Production and Institutional Change. Amsterdam: NEHA. Kok, J. 1919. Handleiding bij het onderwijs aan land- en tuinbouwwinterkursussen, vol. VIII, Veelteelt. Groningen. Krimpen, B. van. 1905. Verbetering van het rundvee (fokvereenigingen). Lochem. Kroon, C. 1997. Onze zwartbonte. 100 jaar rundveefokkerij in Noord-Holland. Oostwoud: Kroon. Kroon, H.M. 1913. De tegenwoordige richtingen in de fokkerij der landbouw-huisdieren in Nederland. Maastricht. ————. 1917. De kruisingen in de huisdierteelt. Assen. Kuhn, H.W. 1941. “Rationeele fokkerij. Een wederwoord.” De Nieuwe Veldbode 8: 11-12. Leeuwen, A. van. 1904. “Verplichte stierenkeuring.” De Veldbode 2: 804-805. ————. 1905. “Invoer van levend vee in Duitsland.” De Veldbode 3: 261. ————. 1912. “De standvastigheid van het roode haar.” De Veldbode 10: 98, 121. ————. 1923. “Rundveefokkerij in België.” De Veldbode 21: 461-464. ————. 1923. “Veeteeltkundige snufjes.” De Veldbode 21: 801-802. ————. 1924. “Tuberculosebestrijding.” De Veldbode 22: 465-467; 1931. Tuberculose bij het rund en haar bestrijding. Tijdschrift voor Diergeneeskunde 58: 17-32, 65-79. Löhnis, F.B. 1901. “De veefokkerij in Nederland en het Nederlandsch Rundvee-Stamboek.” Landbouwkundig Tijdschrift: 1-19. ————. 1911. Veefokkerij. Verslagen en mededeelingen van de Directie van den Landbouw nr 6. ’sGravenhage. Lotsy, J.P. 1915. Grondbeginselen van oordeelkundig fokken en telen. Alkmaar: 15-17, 33. Mansfield, Richard H. 1985. Progress of the Breed : the History of U.S. Holsteins. Sandy Creek, N.Y: HolsteinFriesian World. Molen, H. van der and L. Douw. 1978. “Een bedrijfstak in ontwikkeling. De Nederlandsch landbouw in de jaren 1950-1975.” In H. van der Molen c.s. Omstreden landbouw: bijdragen over technische en ecologische aspecten van de Nederlandse landbouw in de naoorlogse tijd. Utrecht: Het Spectrum. Muilwijk, E. van. 1919. “De Friesche richting een gevaar voor de Zuidhollandsche fokkerij.” De Veldbode 17: 881-882, 924. ————. 1925. “De Dirk IV-stam.” De Veldbode 23: 312-314. ————. 1928. “De moderne erfelijkheidsleer en de praktische fokkerij.” De Veldbode 26: 1025-1027.

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————. 1935. De Jacob-Ruiter-Frans-Max-Stam (van 1870-1935): een verhandeling over de oudste adellijke familie in de Nederlandsche rundveefokkerij. Den Haag. Nobel, C. 1912. Veefokkerij. Schagen. Orland, Barbara. 2003. “Turbo Cows. Producing a Competitive Animal in the Nineteenth and Early Twentieth Century.” In Susan Schrepfer and Philip Scranton (eds.). Industrializing Organisms: Introducing Evolutionary History. London and New York: Routledge: 167-189. Overbosch, H.W. and G.M. van der Plank. 1931. “Een bijdrage tot de kennis van het productievermogen van Friesche stamboekkoeien.” Tijdschrift voor Diergeneeskunde 58: 569-582. Plank, G.M. van der. 1940. “Stamboomstudie.” Tijdschrift voor Diergeneeskunde 67: 848-851. ————. 1948. “Fokwaardebepaling van melkvee in de U.S.A. en in ons land.” Tijdschrift voor Diergeneeskunde 73: 185-192. ———— and W.K. Hirschfeld. 1950. “Veeteelt en K.I..” Tijdschrift voor Diergeneeskunde 75: 250-258. Popta, IJ.A. van. 1965. Geschiedenis fokstal Knol Hartwerd 1854-1965. Sneek: Friese Veefokkerij.. Pott, Emil. 1899. Der Formalismus in der landwirtschaftlichen Tierzucht. Stuttgart. Prescott, M.S. and F.T. Price. 1930. Holstein-Friesian History. Syracuse, N.Y. Reimers, J.H.W.Th. 1916. Die Bedeutung des Mendelismus für die landwirtschaftliche Tierzucht. ’sGravenhage. Rijssenbeek, Th.C.J.M. 1956. “25 Jaar geleden werd de Nederlandsche Zoötechnische Vereniging opgericht.” De Keurstamboeker 4: 6-8. Russell, Nicholas. 1986. Like Engend’ring Like. Heredity and Animal Breeding in Early Modern England. Cambridge etc.: Cambridge University Press. Special Issue on Biology and Agriculture. 2006. Journal of the History of Biology 39/2. Stamhuis, Ida H. , Onno G. Meijer and Erik J.A. Zevenhuizen. 1999. “Hugo de Vries on Heredity, 18891903: Statistics, Mendelian Laws, Pangenes, Mutations.” Isis 90: 238-267. Stanford, J.K. 1956. British Friesians. A History of the Breed. London: Max Parrish. Stapel, K.P. 1988. Rundveefokkerij. Zutphen: Terra. Strikwerda, Reimer. 1979. Een eeuw Fries stamboekvee. Leeuwarden: Friesch Rundvee-Stamboek. ————. 1998. Melkweg 2000. s.l.: CR Delta. ————. 2007. Revolutie in het dierenrijk. Theunissen, B. 1994. “Knowledge is Power: Hugo de Vries on Science, Heredity and Social Progress.” British Journal for the History of Science 27: 291-311. Timmermans, Jacq. 1919. “Zuiden vooruit.” De Veldbode 17: 379-380, 608. ————. 1920. “Naar aanleiding van de centrale stierenkeuring te Roermond.” De Veldbode 18: 615-616. ————. 1923. Gedenkboek van het internationale congres voor rundveeteelt (...) te ’s-Gravenhage op 5 en 6 September 1923. ’s-Hertogenbosch. Tukker, J.G. 1924. Rundveehouderij en -fokkerij in Nederland. Verslagen en mededeelingen van de Directie van den Landbouw nr 3. ’s-Gravenhage. Waardenburg, P.J. 1913. Het mendelisme. Elementen der erfelijkheidsleer. Utrecht. Wibbens, H. 1923. “Welke gegevens dienen in het stamboek te worden vermeld (...).” Compte-rendu des travaux du Congrès international pour l’élevage de l’espèce bovine. ’s-Gravenhage: 306-330. ————. 1907. “Eenige koeien van het roodbonte Maas-Rijn-IJssel-veeras.” De Veldbode 5: 243-245. Wiel, Kees van der and Jan Zijlstra. 2001. Paradijs der runderen. Geschiedenis van de rundveeverbetering in Noord-Holland. Wormerveer: RON. Wood, Roger J. and Víte¡lav Orel. 2001. Genetic Prehistory in Selective Breeding: a Prelude to Mendel . Oxford: Oxford University Press. ————. and Víte¡lav Orel. 2005. “Scientific Breeding in Central Europe during the Early Nineenth Century: Background to Mendel’s Later Work.” Journal of the History of Biology 38: 239-272. Zanden, J.L. van. 1985. “De economische ontwikkeling van de Nederlandse landbouw in de negentiende eeuw, 1800-1914.” AAG-Bijdragen 25. Wageningen. Zwagerman, C. 1927. “Opbrengstverschillen en vraagstukken die daarmede verband houden.” Officieel Orgaan van den Algemeenen Nederlandschen Zuivelbond (F.N.Z.) 22: 124-125, 137-139, 152-153, 170171, 178-188, 201-203, 216-217, 244-246, 259-260, 290-291, 681-684; 23. 1928. 7-9, 25-26, 40-41, 59-61. ————. 1934. De wenschelijkheid en mogelijkheid om fokkerij-problemen in ons land te centraliseeren. s.l. Zwart, W.Th. 1960. Tachtig jaar: een speurtocht door het Friesch Rundvee-Stamboek en aanverwante organisaties. Drachten: Laverman.

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Innovation and Ownership in Living Products: Animals and Fruits in the United States, the 1870s to 1930 Daniel J. Kevles

The best known form of intellectual property (IP) protection is the common utility patent, whose requirements include the stipulations that the invention must be man-made and useful. What is patentable in the United States according to statute dates back to the patent law of 1793, which declared, in language written by Thomas Jefferson, that patents could be obtained for “any new and useful art, machine, manufacture, or composition of matter, or any new or useful improvement thereof.”1 The code said nothing about whether an innovation’s being alive or not has any bearing on its patentability. However, in the nineteenth century living organisms were taken to be unpatentable. In the United States, only one living organism was patented —a form of yeast that Louis Pasteur claimed as an “article of manufacture.”2 That was the exception that proved the rule. Plants and animals were not machines or manufactures. Improvements upon them were not identifiable new compositions of matter. And how could one define the utility of an ornamental plant—say, a rose exhibiting a new fragrance or hue? Under the circumstances, through most of the nineteenth century plant and animal improvers did not seek patents on their products, but this is not to say that they were indifferent to intellectual property protection. While they did not speak of “intellectual property” —the phrase was coined in a Massachusetts court case, in 18453—they were alive to the concept. Indeed, plant and animal improvers were no less profit-minded and imaginative than contemporary biotechnologists, and they devised a variety of property-protection arrangements outside the patent system to achieve protection of the IP in their living innovations. The story of their efforts lies at a rich and relatively unexplored historiographical site—the intersection of pre-Mendelian craft knowledge of plant and animal improvement with law and economics. In establishing their arrangements, the improvers recognized, at least tacitly, that they had to deal with several difficulties. No property right is worth the paper it is written on that can not be enforced. The requirements for enforcement of any property right include the ability to specify and warrant the identity of the property. This was easily accomplished with a tract of land by

1

2 3

Jefferson’s phrasing remains at the core of the U.S. patent code, except for the eighteenth-century word “art,” which was replaced in a 1952 Congressional overhaul of patent law by the word “process.” Bruce W. Bugbee. Genesis of American Patent and Copyright Law. Washington, D.C.: Public Affairs Press 1967. p. 152; Fritz Machlup. “Patents.” International Encyclopedia of the social Sciences. David L. Sills (ed.). New York: Macmillan 1968. XI, 461-64. I am grateful to the Andrew W. Mellon Foundation for support of the research from which this article is drawn and to Karin Matchett for indispensable assistance. Pasteur’s patent, no. 141,072, was issued in 1873. Graham Dutfield Intellectual Property Rights and the Life Science Industries. A Twentieth Century History. London: Ashgate Publishing Co. 2003. p. 151. The judge declared, upholding the broad patent of an inventor of cotton spinning machinery: “Only . . . in this way can we protect intellectual property, the labors of the mind, productions and interests as much a man’s own, and as much the fruit of his honest industry, as the wheat he cultivates, or the flocks he rears.” Catherine Fisk. “The History of Intellectual Property Comes of Age.” Key Note Address, Wisconsin Legal History Symposium, University of Wisconsin Law School, November 13, 2004. p. 6, unpublished, copy in author’s possession.

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surveying and recording its meets and bounds. In contrast, specifying the identity of a living organism—for example, a Shorthorn bull or a Concord grape—was problematic, given that defining biological knowledge such as blood types or DNA sequences was unavailable. The establishment of intellectual property also entails reliable reproduction of the product, including its valuable characters. Absent such reproduction, the IP would be worthless. Faithful reproduction of an organism depends on practical and/or theoretical knowledge of heredity. But the achievement of reproductive fidelity posed a problem for plant and animal improvers that the innovators of, say, mechanical reapers did not face. Unlike reapers, living organisms reproduce themselves. If an improved plant or animal reproduced itself faithfully —or could be made so to reproduce itself—the original improver potentially faced competition from the purchaser that in the absence of patents could not be easily forestalled. In the nineteenth century, identification of a living organism could take the form of a written description, a drawing, or a photograph, but such descriptions were by no means exact or adequate for the purposes of IP disputes. The ability to identify and reproduce a plant or animal depended on the improver’s craft knowledge of biology, heredity, and breeding practices. 4 The history of IP in living organisms during the nineteenth century—and, indeed, even long after the rediscovery of Mendel’s laws, in 1900—thus concerns the interplay among such craft knowledge on the one side and the arrangements at that this body of knowledge and skills permitted. In the United States, IP protection in law for living products found its way onto the agenda of plant and animal improvers during the latter third of the nineteenth century. Before then, markets in agricultural stock were largely local, and the seed, nursery, and animal breeding industries were only incipient.5 It is likely that the warrant for the identity and character of what was offered for sale rested on the purchaser’s knowledge of the purveyor and his reputation. How subsequent competition from buyers was handled is largely unknown, but it may not have been an issue if only because in this period the large majority of new animal breeds as well as plant species and varieties were not the product of effort and investment by improvers. They were imported to the United States, usually at the cost and with the encouragement of the federal government. 6 If breeders did invest in improvements, they likely commanded the local market enough to disregard or shame copycat competitors or they may have considered their efforts a pro bono service to the community, finding reward enough in the admiration of the local agricultural society. Attention to IP protection for plants and animals loomed larger after the Civil War, for several likely reasons. Regional and national agricultural markets emerged with the construction of the railroads and amid increasing urban demand for meats, fruits, and vegetables, as well as ornamental plants, trees, flowers, and shrubs.7 The number of animal breeders, orchardists, and 4

5 6

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The locus classicus for information on craft knowledge and practices in plant and animal breeding in the later nineteenth century is, of course, Charles Darwin. Variation in Plants and Animals under Domestication. 2 vols.; London: John Murray 1868. Available on January 29, 2006 at The Writings of Charles Darwin on the Web (http://pages.britishlibrary.net/charles.darwin/texts/variation/ variation_fm1.html). Cary Fowler. “The Plant Patent Act of 1930: A Sociological History of Its Creation.” Journal of the Patent and Trademark Office Society 82 (September 2000): pp. 622-23. Ibid.; Jack R. Kloppenburg, Jr.. First the Seed. The Political Economy of Plant Biotechnology. 2nd ed.; Madison: University of Wisconsin Press 2004. pp. 50-57; Margaret Derry. Bred for Perfection. Shorthorn Cattle, Collies, and Arabian Horses since 1800. Baltimore: The Johns Hopkins University Press 2003. passim.

Innovation and Ownership in Living Products

nurserymen was growing. Eager to be competitive, the proprietors of these enterprises felt the need to offer new and superior breeds or varieties as often as possible. While many such products continued to come from importation, an increasing number were generated by breeders and, in the plant world, by variants and sports found in the fields. Whether generated by hybridization or chance finds, the improved variants usually required effort and investment to turn them into marketable products, a condition that made improvers more attentive to capturing financial returns on the IP they had created. But doing business across vast, impersonal distances, animal and plant improvers could rely much less on reputation to warrant the identity and quality of their products. And the distance as well as the impersonality of the buyer-seller relationships made it all the easier for purchasers to propagate an improver’s innovation and sell it as her own. During the nineteenth century, breeders of pure-bred Shorthorn cattle devised a system for protecting the IP in their animals responsive to these circumstances. Drawing on the methods pioneered by the English breeder Robert Bakewell in the late eighteenth century, they bred through pedigree, selecting for valuable characters and intensifying their embodiment in the animals through inbreeding.8 The resulting purebreds likely tended to possess an essential feature of IP licensing—intergenerational reliability, which is to say that the products of their stud service were likely to resemble them. Warranting the identity of the animals was achieved by registering the pedigrees in publicly available stud-books. The books, originally imported from England, along with the breed, were developed by private entrepreneurs in different states, and by the late nineteenth century they were increasingly characterized by non-uniformity in standards, sloppiness in the records, and general unreliability. As warrants of identity, they left a good deal to be desired. To solve that problem, the Shorthorn breeders moved in 1876 to regulate their market to a degree by forming the American Shorthorn Association. The Association bought the existing registry books and amalgamated them into one. The arrangement thus advantaged genuine Shorthorn breeders and protected buyers against fraudulent sellers—that is, purveyors of putative Shorthorns whose animals were unregistered with the Association.9 This system for the protection of IP in Shorthorns likely exemplified the systems developed for other farm animals and, with some variation, for pets and race horses. 10 It did not firmly protect the IP developed by individual breeders, but it protected very well the collective IP of the cartel of breeders represented by the breed association. In 1891, Liberty Hyde Bailey, the prominent plant scientist and a professor at Cornell University, noted the value of the system: “There is no law to compel one to register an animal, but every breeder knows that it is only through registration that he can advertise, sell and protect blooded stock. And there is no intelligent purchaser who would 7 8

9

10

Fowler. “The Plant Patent Act.” pp. 623-24. On Bakewell, see Harriet Ritvo. “Possessing Mother Nature. Genetic Capital in Eighteenth Century Britain.” In John Brewer and Susan Staves. (eds.) Early Modern Conceptions of Property. London and New York: Routledge 1995. pp. 413-26. See also H. Cecil Pawson. Robert Bakewell. Pioneer Livestock Breeder London: Crosby Lockwoon & Son 1957. Derry. Bred for Perfection. pp. 15, 20-29, 34-36. For a more extensive discussion of the history of IP in animal breeding, see Daniel J. Kevles. “Breeding, Biotechnology, and Agriculture. The Establishment and Protection of Intellectual Property in Animals Since the Late Eighteenth Century.” In Preprint 310, Workshop, History and Epistemology of Molecular Biology and Beyond: Problems and Perspectives. Berlin: Max-Planck-Institut für Wissenschaftsgeschichte 2006. pp. 69-80. Ibid., passim.

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think of negotiating for such stock without having obtained the testimony of the herd-book.” 11 In all, the breed association-stud book system provided the degree and type of protection consistent with what could be done in the then-current state of biological and breeding knowledge to specify the animal’s hereditary essence, warrant its hereditary prowess, and transmit that hereditary essence to succeeding generations. Still, it remains an open historical question whether and how the purveyors of pure-bred animals or stud services managed to discourage purchasers from competing against them with the offspring of their animals. The principal IP-related problem for improvers of plants that were reproduced sexually —for example, corn, the grains, most vegetables, and flowers—was that they did not ordinarily breed true. Sellers of their seed thus could not guarantee the quality and character of any given crop. J. M. Thorburn & Co., a prominent nursery in New York, warned buyers that they gave “no warranty, express or implied, as to description, quality, productiveness, or any other matter of any seeds, bulbs or plants they send out.” Among the reasons was “the well-known tendency of many vegetables to revert to their original types, notwithstanding the care of the seed-grower.”12 Then, too, farmers could save seed from their crops, and then either plant them, sell them, or both, thus undercutting the improver’s control of his IP in the plant. Under the circumstances, the nascent private seed industry paid little attention to IP protection. It was content to rely for new varieties on importation and on the innovations produced by the state agricultural experiment stations established by the Hatch Act, in 1887. Of far greater concern than IP was the competition the seed trade faced from the federal government. Beginning in the 1830s, the U.S. Patent Office and then the U.S. Department of Agriculture distributed seed gratis to farmers —more than ten million packets annually in the 1890s—via members of Congress and their franking privilege. What the seed industry wanted from the policy arena was not IP protection but an end to the seed distribution program, a campaign that succeeded in 1924.13 Innovations and improvements in asexually reproducible plants and trees —the foundation of the horticultural industry—came partly from the hybridizing work of breeders like Luther Burbank but in the overwhelming main from chance finds in the field and orchard. 14 The finds arose from bud sports or fortunate sexual pollinations, but once found they could be reproduced virtually identically by the nurturing of grafts or cuttings. Commercial nurseries acquired such finds, tested them for such characteristics as sturdiness and fruit-bearing qualities, then put them on the market. Stark Brothers Nursery and Orchards, based in Louisiana, Missouri, was one of the oldest and perhaps the largest such enterprise in the country. It sponsored an annual fair that encouraged farmers to submit their good fruits, including those of chance finds, for competitive judgment. In 1893, through this means, the firm learned about an apple tree that produced a luscious red fruit. The next year, it brought the tree with all propagation rights —which is to say all

11

12 13 14

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Liberty Hyde Bailey. Report. “Protection to the Originator of Varieties.” read at the meeting of the American Association of Nurserymen, June 4, 1891. In Transactions of the American Association [of Nurserymen]. June 3-13, 1891. pp. 88-91. J. M. Thorburn & Co.. Catalogue [1908]. Copy in New York Botanical Gardens Archives, Catalogue Collection, Box. 538. Italics in the original. Fowler. “The Plant Patent Act.” pp. 622-23; Kloppenburg. First the Seed. pp. 61-65. Bailey. Report. “Protection to the Originator of Varieties.” pp. 88-89.

Innovation and Ownership in Living Products

its IP—from its owner, a farmer in Iowa, named the fruit the “Delicious” apple, and proceeded to market the tree to the world.15 Nurserymen and orchardists could be confident that the young trees they sold would bear fruit very much like the fruit on the trees from which they had been derived. The question for the purchaser was whether the quality of the tree—for example, its bearing abundance—and of its fruit would live up to its billing. The reputation of a well-known seller —for example, Burbank or Stark Brothers—counted for something, but in the impersonalized setting of the national market nurserymen relied increasingly on advertising, placing ads in horticultural and gardening journals and distributing catalogues across the country. The ads tended to include what amounted to warrants of their products’ quality and identity in the form of farmers’ testimonials about the merits of their fruit trees, shrubs, and flowers. Yet the ease with which, say, valuable fruit trees could be easily reproduced virtually identically, through grafting, and thus numerously multiplied facilitated theft of the developer’s IP. Competitors could purchase the trees, or take cuttings of them from someone’s nursery in the dead of night, then propagate and sell them. Burbank tried to protect himself against such thieves by telling buyers that the way to judge novel fruits was to “look to their source, and also if possible purchase direct from the originator.” He also charged high prices for his innovations – say, $3,000 for a new plum tree, including all “stock and control”—thus attempting to gain in the first sale revenue that would cover his costs and return a reasonable profit. 16 The pricing strategy was intended to capture what economists call all the downstream revenues of which thieves might deprive him, since he would be unable to control the reproduction of the tree once he had sold it. Trouble was that the high first-sale pricing did not work very effectively to compensate horticultural innovators for the loss of IP in their new fruit trees. Nurserymen repeatedly complained that they failed to receive just returns for all their investment of time and money. Burbank fulminated to the readers of Green’s Fruit Grower that he had “been robbed and swindled out of my best work by name thieves, plant thieves and in various ways too well known to the originator. . . . A plant which has cost thousands of dollars in coin and years of intensest [sic] labor and care and which is of priceless value to humanity may now be stolen with perfect impunity by any sneaking rascal. . . . Many times have I named a new fruit or flower and before a stock could be produced some horticultural pirate had either appropriated the name, using it on some old, well-known or inferior variety or stealing the plant and introducing it as their own, or offering a big stock as soon as the originator commences to advertise the new variety.” 17 Burbank as an innovator was largely in the business of selling to nurseries and orchardists, middlemen who would propagate his trees and sell them to gardeners, farmers, and other 15 Dickson Terry. The Stark Story: Stark Nurseries’ 150th Anniversary. Columbus, Mo: Missouri Historical Society 16

17

1966. pp. 38-40.

Catalogue. New Creations in Fruits and Flowers, June 1893. Santa Rosa, CA: Burbank’s Experimental Grounds 1893. p. 12; Catalogue. Twentieth Century Fruits, 1911-1912. Santa Rosa, CA: Burbanks’ Experiment Farms. 1911. p. 1. Copies in Luther Burbank Papers, Library of Congress, Box 14. Bolded print in the source. Burbank to Jacob Moore. May 4, 1898. In Green’s Fruit Grower. June 1898, clipping in Luther Burbank Papers, Luther Burbank Home and Gardens, Archives, Santa Rosa, CA. Scrapbooks. Vol. 2. p. 45. See also Jacob Moore to Chas. A. Green, April 20, 1898; “Protection for Fruit Evolvers.”Editorial. California Fruit Grower. n.d.; and Moore to Peter Gideon. n.d.. Green’s Fruit Grower. ibid. Scrapbooks. Vol. 2. pp. 44, 47, 115.

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consumers. Stark Brothers, which did not breed new fruit trees but only acquired them, was in the business of mass marketing. Realizing the value of their IP by charging high prices would have been counterproductive to their business plan. To protect the IP in their fruit trees, the Starks trademarked them, using the trademark law that Congress passed in 1881. In the 1890s, the Stark catalogues included gorgeous paintings of their named fruits with a small banner beneath each declaring that it was covered by a Trade Mark or, in the case of the Gold Plum, that it was “Trademark Pat[ente]d. Feb. 25, 1895.” 18 The trademark, however, would not prevent someone from obtaining the tree or cuttings from it, propagating the wood, and then selling the tree under a different name. Under the circumstances, beginning in the 1880s and with mounting insistence in the 1890s, American nurserymen began urging the establishment of legal protection for what they called the rights of “originators.” Some of the agitation recommended the expansion of the patent system to include coverage for innovations in plants and trees. Mindful of their exclusion from the patent system, nurserymen wondered why, as the California Fruit Grower put it, “the writer of a book, the composer of a song, the designer of a drawing or the originator of a mechanical device should be protected in their productions, while the originator of an improved flower or fruit is denied the same privilege.” 19 The move to patentability was blocked, however, when, in 1889, in Ex parte Latimer, the U.S. Commissioner of Patents rejected an application for a patent to cover a fiber identified in the needles of a pine tree, declaring that it would be “unreasonable and impossible” to allow patents upon the trees of the forest and the plants of the earth.”20 The commissioner’s ruling formed the basis for what came to be known as the “product-of-nature” doctrine —that while processes devised to extract what is found in nature can be patented, objects discovered there or bred from there can not be patented. In a report to the American Association of Nurserymen in 1891, Liberty Hyde Bailey rejected the horticulturalists’ patenting initiative as in any case unwarranted. New varieties were not inventions, he noted, precisely because they were accidents found in the fields, adding, “When the time comes that men breed plants upon definite laws, and produce new and valuable kinds with the certainty and forethought with which the inventor constructs a new machine, or an author writes a book, plant patents may possibly become practicable.” 21 Bailey held that plant originators should nevertheless be protected, though he doubted that any new legislation would do the job. “It is evident that after a variety is put upon the retail trade it becomes public property, and no statute could further protect it,” he observed. He proposed that the nurserymen draw on existing trademark law to obtain protection through a national register of plants administered by the Department of Agriculture. The originator would send the department “a specimen, description and perhaps picture of his novelty,” and the department would issue a certificate, a type of trademark insuring him “inviolable rights” in his innovation. 18 19

20 21

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Copy in Scrapbooks. Vol. 1, p. 141. Burbank Home and Gardens, Archives. “Protection for Fruit Evolvers.” Editorial. California Fruit Grower. quoted in Luther Burbank, Burbank’s Experiment Farms. The 1899 Supplement to New Creations in Fruits and Flowers. Luther Burbank Papers, Library of Congress, Washington, D.C., Box 14. Daniel J. Kevles. “Ananda Chakrabarty Wins a Patent. Biotechnology, Law, and Society, 1972-1980.” HSPS. Historical Studies in the Physical and Biological Sciences 25: 1 (1994), 111. Bailey. Report. “Protection to the Originator of Varieties.” pp. 88-89.

Innovation and Ownership in Living Products

He acknowledged that thieves could sell the variety under a different name, but he thought that “tricksters” would be discovered and in consequence commercially disadvantaged. The public would soon learn to buy only from originators who possessed a registration certificate, just as they had learned to purchase only animals registered with the breed associations. 22 Bailey, like the Stark Brothers with their trademarked fruits, tacitly assumed that the certificates would not only protect the name of the innovation but also secure to the originator the exclusive right to the plant or tree and to its propagation. But that assumption was severely undercut in 1895 by the ruling of a federal appeals court in the case of Hoyt et al v. J. T. Lovett Co. James Hoyt and Edwin Hoyt, nurserymen in Connecticut, had sued the J. T. Lovett Nursery, in New Jersey, for selling a grape that had been found in the Green Mountains in Vermont. The Hoyts believed they had bought the grape wood with exclusive rights and they had trademarked it as the “Green Mountain Grape.” The court found against the Hoyts partly on grounds that certain facts in the case contradicted the tenets of trademark law as it had been judicially interpreted. But its decision also addressed the scope of trademark protection for living products. 23 Apparently Lovett’s lawyer had raised the issue, contending, in the words of the court, “that the protection of a trade-mark cannot be obtained for an organic article which, by the law of its nature, is reproductive, and derives its chief value from its innate vital powers, independently of the care, management, or ingenuity of man.” The court, while noting that the question was “novel and unprecedented,” agreed, writing: “The Hoyts did not make the Green Mountain vine, nor, strictly speaking, did they produce it. It grew out of the earth, was fashioned by nature, and endowed with powers and qualities which no human ingenuity or skill could create or imitate. If such protection as that now claimed by the complainants was allowed, a breeder of cattle could with equal propriety and reason demand like protection for the natural increase of his herd. In every aspect such claims would seem to be impracticable and inequitable.” 24 Meanwhile, Liberty Hyde Bailey had evidently persuaded the nation’s nurserymen to discard their ambitions for patent protection in favor of establishing trademark protection through a national registry. But the ruling in Hoyt et al had nullified Bailey’s contention that to accomplish their purpose the nurserymen needed only administrative action by the Department of Agriculture. Legislation was necessary, and during the next decade the leading nurseries, including Burbank and Stark Brothers, moved to obtain it, engaging a lawyer in Washington, D.C. named F.T.F. Johnson. In 1906, a bill, perhaps drafted by Johnson, was introduced in the House that would amend the trademark act by authorizing the commissioner of patents to register an originator’s new variety of plant, bush, shrub, tree, or vine. Registration of the name would constitute a trade mark and would include for twenty years the “exclusive right to propagate for sale and vend such variety of horticultural product under the same so registered.” 25 The bill enjoyed broad support from nurserymen, a number of whom wrote letters to Johnson that were introduced at hearings on the bill before the House Committee on Patents, in March 22 23 24 25

Ibid. pp. 89-90. Hoyt et al v. J.T. Lovett Co., Circuit Court of Appeals, Third Circuit, 71 F. 173; Dec. 3, 1895. Ibid. U.S. Congress, House of Representatives, Committee on Patents, Arguments before the Committee . . . on H.R.113570, Authorizing the Registration of the Names of Horticultural Products and to Protect the Same, March 28, 1906, 59th Cong., Washington, D.C.:GPO 1906. pp. 3-5, 12-13.

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1906. M. Crawford, who lived in Cuyahoga Falls, Ohio and who had given the world the Crawford peach, explained the principal reason behind the desire for protection: “An originator may work ten or twenty years to produce a variety worth naming and introducing. If he attempts to introduce it himself he will hardly get enough out of it the first year —the only year he controls it— to pay the printer. The second year he is undersold by competitors, many of whom never saw the real thing.”26 Several committee members expressed sympathy for protecting the rights of the originators, but the committee leadership found the bill before it constitutionally dubious. For one thing, by trying to protect rights in the product by protecting rights to its name, it sought to combine the exclusivity of a patent with the coverage of a trademark. More important, constitutional authority for the granting of federal trademarks rested on congress’ power to regulate interstate commerce. The bill allowed for trademark protection of plants even if they were not sold in interstate commerce, and under prevailing interpretation of the commerce clause it was unconstitutional for congress to regulate intrastate trade. Congressman Frank D. Currier, of New Hampshire, the chairman of the Patents Committee, summarily declared: “The proposition is as clearly unconstitutional as anything can be.”27 Although an immediate failure, the 1906 venture did lead to the formation of a lobbying group, the National Committee on Plant Patents under the American Association of Nurserymen. In 1929, Paul Stark, of Stark Brothers, became chair of the committee. Along with other nurseries, Stark Brothers had been trying to protect its propagation rights in new fruits by imposing contractual obligations upon the purchaser—for example, an agreement that he would neither sell nor give away scions, cuttings, or buds. Liberty Hyde Bailey had suggested in 1891 that nurserymen use such contractual arrangements and the court in Hoyt et al had in passing noted their acceptability. However, the contracts were some times difficult to enforce, which helped energize Stark’s eagerness for the stronger IP protection that a patent would provide, and in 1930, not least because of Stark’s lobbying effort, Congress passed the Plant Patent Act. 28 The act covered only asexually reproduced organisms, and it authorized a patent to anyone who “has invented or discovered and asexually reproduced any distinct and new variety of plant, other than a tuber-propagated plant. . . .”29 Given its requirement of distinctiveness rather than usefulness, it was not a utility patent law. Moreover, it did not establish the conventional legal bargain that granted the inventor a monopoly right in exchange for public knowledge of how the invention was produced so that others could innovate beyond it. In most cases, there was no such knowledge to be disclosed. Liberty Hyde Bailey may have predicted that patent protection would accompany the discovery of the laws of inheritance, but the rise of Mendelian genetics played little or no role in the work of the nurserymen who were the Act’s principal advocates. Even in 1930,

26 27 28

29

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Crawford to Johnson, March 19, 1906, in ibid., p. 10. Ibid., pp. 4-5, 9. Bailey. Report. “Protection to the Originator of Varieties.” p. 90; Hoyt et al v. J.T. Lovett Co., 71 F. 173; Dec. 3, 1895; Fowler. “The Plant Patent Act.” pp. 630-42; Glen E. Bugos and Daniel J. Kevles. “Plants as Intellectual Property. American Practice, Law, and Policy in World Context.” Osiris. 2nd Series. Vol. 7, Science After ’40 (1992): 81-88. Quoted in Fowler. “The Plant Patent Act.” p. 641.

Innovation and Ownership in Living Products

the innovations in fruits that were their stock in trade continued to arise from chance variations in the field rather than breeding on Mendelian principles.30 In all, the Plant Patent Act harkened back to the seventeenth century, when patents were granted as privileges in the market—royal dispensations to encourage commerce in new technologies, often from abroad, or to reward favorites. Indeed, the Plant Patent Act might well have been called the Stark Horticultural Privilege Act, not only because of Stark’s role in its passage but because it granted a privilege of intellectual property protection that was tailored to the practices and needs of horticultural innovators.31 Still, for all its simultaneous restrictiveness and looseness, the act was the first statute passed anywhere in the world that extended patent coverage to living organisms. It helped pave the way for the legal protection of IP in sexually reproducing plants, which Congress authorized in 1970, and for the extension of utility patents to all living organisms other than human beings after 1980, when in the emerging age of biotechnology the U.S. Supreme Court ruled that whether an innovation is alive or not is irrelevant to its patentability.32 Daniel J. Kevles Yale University [email protected]

30

31

32

The rise of Mendelian genetics similarly changed breeding practices and the system of IP protection for animals very little. No doubt one of the reasons was the small number of offspring produced by animals, which makes difficult conventional genetic analysis. In 1925, one farm expert noted, “Up to the present time, the new knowledge of genetics has contributed little” to advances in animal breeding, adding, “Animal breeding proceeds in much the same way as it [did] four thousand years ago.” Derry. Bred for Perfection. pp. 12-14. I am indebted to Mario Biagioli for the analogy of the Plant Patent Act to the earlier practice of awarding patents as privileges. On patents as privileges, see Miller and Davis. Intellectual Property. p.5; and Jessica He. “‘Hail to the Patents!’ The Ethics, Politics, and Economics of the Early Modern Patent System in England.” Senior Essay, Ethics, Politics, and Economics, Yale University 2005. pp. 2-27. For these developments, see Bugos and Kevles. “Plants as Intellectual Property.”; Kevles. “Ananda Chakrabarty Wins a Patent.”; and Daniel J. Kevles. “The Advent of Animal Patents: Innovation and Controversy in the Engineeering And Ownership of Life.” In Scott Newman and Max Rothschild. (eds.) Intellectual Property Rights and Patenting in Animal Breeding and Genetics. New York: CABI Publishing 2002. pp. 18-30.

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Coalition and Opposition: Heredity, Culture, and the Boundaries of Anthropology in the Work of Alfred L. Kroeber Maria E. Kronfeldner

Introduction “If there is nothing beyond the organic, let us quit our false and vain business and turn biologists….”1 This is what anthropologist Alfred L. Kroeber (1876-1960) said in 1916—a time when ideas about heredity changed a lot, when genetics established itself as an experimental science, when hereditarian thinking was gaining wide acceptance in the US, and—last but not least—when American anthropology emancipated itself from being a museum-based profession and became an academic discipline. In face of all this, Kroeber, student of Franz Boas (1858-1942), was fighting for the boundaries and the autonomy of the new academic discipline. And this struggle included a severe opposition to certain kinds of hereditarian thinking. Kroeber tried to accomplish his boundary work by focusing on a concept of culture that not only saves man from being ‘just another animal’ but gives anthropology a distinctive phenomenon for study. According to him, culture is defined as not only opposed but also analogous to biological heredity. In addition, he stressed that the rise of a Weismannian, non-Lamarckian concept of inheritance, today often called ‘hard inheritance,’ and the correspondent denial of ‘soft inheritance’ was necessary for the historical development of such a concept of culture.2 Some historians have acknowledged Kroeber’s point about inheritance of acquired characteristics.3 Yet, they did not concentrate on the consequences of his case for an historical account of the impact of the concept of hard inheritance.4 In part 1, I will say a little bit more on the shifting boundaries of anthropology at the beginning of the 20th century. This makes clear why Kroeber needed an opposition to hereditarian thinking. 1 2

3 4

Kroeber (1916b: 296). Kroeber himself did not use the terms ‘hard’ or ‘soft’ inheritance and today these terms are not necessarily used in the same way by different authors. I will use them in the following sense: hard inheritance is what Weismann’s concept of inheritance implied, which will be specified later in this paper. Soft inheritance is the exact opposite, implying that the hereditary material is malleable at any time, as for instance in Lamarckian inheritance. The term ‘Lamarckian’ is today used in unison for the idea of inheritance of acquired characteristics, even though Lamarck was by far not the only one referring to this kind of inheritance. It was common knowledge of his time and even Darwin believed in it. See Zirkle (1946) on the history of the idea from the Greeks to Darwin. Ernst Mayr (1982) is often quoted as the one who introduced the terms of ‘soft’ and ‘hard’ inheritance. Cyril Darlington, however, used the term “hard heredity” already in 1959, as I learned from Jonathan Hodge during the workshop. Yet, Darlington used the terms with a slightly different meaning (see Darlington 1959: 14, 54-56, Appendix, and compare Mayr 1982: 687). See for instance Stocking (1968: 250-269); Harris (1968: 121); Peel (1971: 143-146); Freeman (1983: 3450); Degler (1991: 96-100). Thus, it is not surprising that Kroeber has been ignored in accounts of the impact of the concept of hard inheritance (as for instance in Paul 1995: 40-49) or in historical accounts of the history of hereditarian thoughts in general, as for instance in Ludmerer (1972). He is briefly mentioned by Kevles (1985). In turn, it is not surprising that a standard history of anthropology, such as Patterson (2001), can ignore Kroeber’s reference to the concept of hard inheritance as important for his concept of culture.

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I will then analyze in part 2 how Kroeber used a Weismannian or non-Lamarckian concept of hard inheritance to secure the boundaries of anthropology. This shows why he wanted geneticists to enter into a coalition with him. I will end, in part 3, with some systematic notes on the concept of culture, with remarks on why Kroeber’s case is important for contemporary debates on evolution, and on why his case is important for writing a cultural history of heredity.

1. The shifting boundaries of anthropology at the beginning of the 20th century That sciences are organized into disciplines means that conceptual boundaries are constantly built and rebuilt: the space of ideas gets delineated into areas of autonomy and exclusive authority over problems. Since ages, anthropology has conventionally been defined as ‘the science of man.’ At the beginning of the 20th century in the US, anthropology was thought to comprise four parts: physical anthropology, ethnology (which was later called cultural anthropology), linguistics, and archaeology. At the same time, it stopped being a mere museum based profession and became an academic discipline, with the usual outward signs this has: curricula, degrees, journals, disciplinary associations etc.5 Naturally, there was a need to define the boundaries of anthropology in the face of other academic disciplines and areas of research, such as psychology, biology in general, and genetics in particular. And this need was also a need to define the internal relationship between physical and cultural anthropology. MARGINALIZING PHYSICAL ANTHROPOLOGY Franz Boas regarded physical anthropology as central to understanding the behavioural differences between groups of people: heredity, a phenomenon considered as part of physical anthropology, was for him one of several factors an anthropologist has to take into account in order to understand the development and behaviour of individuals. His student Alfred L. Kroeber was more radical. He tried to marginalize the field of physical anthropology. Kroeber grew up in a German-Jewish-American intellectual context in New York and received Columbia’s first PhD in anthropology in 1901, the ninth in the whole US. Immediately afterwards, he got a permanent position. His job was to build up a department of anthropology at the University of California, Berkeley. By 1907 he was an important figure in the discipline and counts until today as the most influential figure in the establishment of American anthropology after Boas.6

5 6

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For more on the history of anthropology before it became a scientific discipline and how it developed since then in general see: Hinsley (1981), Darnell (1998), Patterson (2001). See Bidney (1965) for a short review of his life and work, Steward (1973) for a book length one, containing a summary of the biography written by Kroeber’s wife Theodora Kroeber (1970); see also Thoresen (1975) on the establishment, financing, and development of academic anthropology in California.

Heredity, Culture, and the Boundaries of Anthropology in the Work of Alfred L. Kroeber

Figure 1. Anthropologist Alfred L. Kroeber, who strongly believed in culture as important to explain similarities and differences, and Ishi, famous last member of the Yahi tribe, 1911. Photo credit: UC Berkeley, Phoebe Hearst Museum of Anthropology.

For Kroeber, cultural anthropology was at the centre of the discipline, while “the other parts were secondary and marginal and owed their significance to their contribution towards an understanding of cultural history,” as the anthropologist David Bidney says in a review on Kroeber’s impact.7 Consequently, Kroeber never contributed anything to physical anthropology. At the same time, others, geneticists such as Davenport and other anthropologists, pulled in the exact opposite direction: they tried to marginalize cultural anthropology. The following example from the arena of the politics of science will show that anthropologists such as Kroeber had quite concrete reasons to be afraid of losing their jobs. In other words, there was a practical or pragmatic pressure to secure the boundaries of anthropology by marginalizing physical anthropology and by opposing hereditarianism. REPRESENTATION IN THE SCIENTIFIC BUREAUCRACY Between 1916 and 1918, Boas and his students fought for their representation in America’s scientific bureaucracy. At issue were the posts for the National Research Council’s Committee on Anthropology. For historians of anthropology the story is well known. George E. Hale, the Director of the National Research Council, asked William H. Holmes (1846-1933), important figure in American anthropology and defender of a racial interpretation of cultural differences, to organize the Committee on Anthropology. He chose Ale¡ Hrdliçka (1869-1943), who was a defender of physical anthropology as an independent discipline, to take the lead. The goal was to prevent that Boas and his students get control over the committee, i.e. to prevent cultural anthropology from becoming too influential. Yet they could not totally ignore Boas. Holmes thus put Hrdliçka, Boas, and Charles B. Davenport (1866-1944), a geneticist and the leader of the

7

Bidney (1965: 268).

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American eugenicist movement, on the list for the committee. Yet, Hale dropped Boas from the committee because of Boas’ anti-war activism. In April 1917, Madison Grant (1865-1938), a wealthy racist propagandist, who published his best-selling book on the “Passing of the Great Race” in 1916, offered money for the committee’s work in exchange of membership in it. In the end, the committee consisted of Holmes, Hrdli™ka, Grant, and Davenport. And it was Davenport, who had been selected by Hale in February 1918, who was to represent the interests of the Committee on Anthropology to the National Research Council’s Division of Medicine and Related Sciences. In a nutshell, a geneticist, who defended eugenic doctrines, came to represent anthropology in the scientific bureaucracy of the National Research Council. And this was at a time when there were already trained anthropologists to do so. 8 Besides this struggle for representation, there was the emancipation from the older generation of anthropologists such as Holmes, which were not trained as anthropologists and predominantly oriented towards a racial hereditarianism, and the general dominance of racism and eugenics in the US at that time. These are the three main contexts in which cultural anthropologists in the US formed a clear professional identity as cultural anthropologists. That Kroeber perceived a danger (and wanted others to perceive such a danger) in the various developments just mentioned is also evident from the language of war and territory that he used: according to him, biology is a discipline that “forged its weapons, taught itself their use, conquered a territory, and stands forth a young giant of prowess,” in order to “annex the antiquated realm of history that lay adjacent.”9 Now, it was Kroeber who used the biologist’s own concept of hard inheritance to keep up the two oppositions: against the institutional hegemony of biologists and against the scientific hegemony of hereditarianism. According to Stocking, he was the only one among social scientists, who realized that the concept of soft inheritance (i.e. inheritance of acquired characteristics) prevented the autonomy of anthropology and other social sciences.10

2. Alfred L. Kroeber’s boundary work: culture and/as inheritance Kroeber’s boundary work for anthropology found its first peak with a couple of papers between 1915 and 1917, ending with his famous article on “The superorganic” (1917), which established cultural determinism as his major doctrine. THE PSYCHIC UNITY OF MAN AND THE SUPERORGANIC NATURE OF CULTURE Already in 1910, Kroeber laid down the basic frame of his point of view on culture, heredity, and anthropology. His example was morality: according to him, morality is governed by an innate, instinctual moral sense. Yet, variations in moral behaviour between “civilized” and “uncivilized people” are due to different cultural influences and not due to innate differences in the alleged moral sense. In other words, behavioural differences do not imply that there are essential inborn 8 9 10

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For more on this and the history of the Committee on Anthropology see in particular: Stocking (1968: 270-308), Cravens (1978: 89-120); Patterson (2001: 55-60). Kroeber (1916a: 34). Stocking (1968: 259).

Heredity, Culture, and the Boundaries of Anthropology in the Work of Alfred L. Kroeber

mental differences between groups of people: to the contrary, one should assume a psychic unity of mankind and explain the behavioural differences by the influence of what Kroeber called civilization, history, or culture. 11 From this assumption, Kroeber went on to describe culture as “outside of race and independent of the human body.”12 This means that culture influences culture (via behaviour), but it does not influence the body, at least not the innate racial basis of the respective behaviour, and vice versa. That culture influences and thus explains culture means that culture is for Kroeber a system or process sui generis. Culture is “superorganic”13—‘on top’, so to say, of organic matters, relying on “social inheritance or cultural transmission” instead of biological inheritance. 14 To understand his position clearly, the following points have to be taken into account: in his 1915 paper “Eighteen professions,” arguing for the autonomy of anthropology as a distinct discipline, Kroeber assured that the psychic unity of man is not a proven or disproven fact, but a necessary assumption for the “historian,” i.e., the anthropologist, since otherwise “his work becomes a vitiated mixture of history and biology.”15 Yet, at the same time, he acknowledges that history and biology are intertwined and that the degree of their contribution in the development of individuals cannot be tested.16 Yet, the two statements are not contradictory. On the contrary, the argument that culture is a process in its own right is compatible with Kroeber’s claim that the behaviour of individuals and their development is caused by multiple factors, culture being merely one of them. If, however, we look at culture itself, then we see that culture is independent of nature, i.e., a phenomenon that can only be explained by reference to pre-existing culture. It is from this inter-individual phylogenetic perspective so to say, that culture always derives from previous culture, as a cell always derives from previous cells. The last issue that might cause misunderstanding is the issue about holism: Kroeber’s paper on culture as superorganic is often treated as defending a strong holistic conception of culture. 17 Even though I cannot decide this issue here, the following two points should be taken into account. (i) Although Kroeber believes that culture is maintained via individual mental states or individual actions, he also believes that “[c]ivilization is not mental action itself,” but rather “a body or stream of products of mental exercise.”18 This is not pointing to an ontologically dubious whole; it is pointing to a causal inter-individual lineage of the effects of mental acts. (ii) In addition, although he sometimes wrote in 1917 and in 1919 as if individuals are mere passive bearers of culture (implying that their properties do not determine culture and vice versa, i.e., culture does 11 12 13

14 15 16 17

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This psychic unity does not exclude individual differences. It is an “identity of average” as he makes most clear in Kroeber (1917: 194-203). Kroeber (1910: 446). Kroeber (1916b, 1917). Kroeber took the term superorganic from Spencer, who used it in the sense of exo-somatic or artificial as secondary environment of organisms, as Kroeber makes clear in the re-edition of papers from him (Kroeber 1952: 4). Kroeber, on the other hand, uses it for an autonomous system of change and stability, i.e. inheritance. Kroeber (1916c: 368). Kroeber (1915: 285). Kroeber (1915: 285). For a critique of the concept of the superorganic understood in a holistic manner, see Bidney (1944), Herskovits (1948) and the discussion of the issue in Kaplan (1965) that shows that the actual issue is methodological and epistemological, but not ontological; it is an issue about the distinctive subjectmatter of anthropology. Kroeber (1917: 189 and 192).

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not determine the properties of individuals) and as if culture is a special ontological substance, he recanted from this in 1952: he admitted that culture as a whole is not a peculiar emergent entity or substance and that individuals are more important than he put it in 1917. His goal in 1917, he himself says in 1952, was to establish the recognition of culture as an “autonomous” system, independent of “biological explanation.”19 Consequently, if the context of an opposition to hereditarianism is ignored, Kroeber’s claim about the superorganic nature (and its genesis) cannot properly be understood.20 And it was this opposition that correlates with a denial of the Lamarckian principle of the inheritance of acquired characteristics. INHERITANCE OF ACQUIRED CHARACTERISTICS In 1916, in a paper called “Inheritance by magic,” published in the American Anthropologist, Kroeber moved the denial of inheritance of acquired characteristics to the centre of his account. In order to do so, he referred to three important aspects of August Weismann’s (1834-1914) ideas on inheritance: first, that experiments failed to produce positive evidence for the inheritance of acquired characteristics; second, that all cases of evolution are explainable without reference to inheritance of acquired characteristics; third, that inheritance is ‘hard’: that the hereditary material is not produced by the organism, but present from the start, continuously existing, and protected against changes that occur in the somatic tissue. Acquired changes, i.e., changes to the somatic tissue of the organisms, are not heritable on this basis. In Kroeber’s words, Weismann’s “basic idea” was “that the hereditary substance is totally distinct from the organic body, and that therefore the fate of the individual cannot affect the race.”21 In addition, Weismann’s concept of heredity meant that the germ plasm exists over time independently of individuals. The germ plasm is thus sub-individual and inter-individual at the same time—almost as superorganic, i.e., independent of individuals, as Kroeber assumed culture to be. Kroeber also referred to Mendelism, the “new branch of biological science,” as providing a corroboration of this concept of hard inheritance. Kroeber states that “if Mendelism rests on anything at all, it rests on the doctrine of the utter separateness of what it calls gamete and zygote. This separateness may be purely conceptual, but it is the only concept which it has yet been possible for anyone to think out that will explain and hold together the looming mass of facts heaped up by genetic observation and experiment.” Kroeber also mentions that although Mendelians perceive themselves as opposed to

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Kroeber (1952: 7, 22-3). Compare Kroeber & Kluckhohn (1952: 49), but without reference to Kroeber’s papers between 1916-7, or Bidney (1965: 273). A point I originally took from Kuklick (2004). I thus depart from the conclusions drawn by anthropologists such as Bidney (1965), who derive from the failure of a total independence of culture from individuals that the concept of the superorganic did not make any sense. It did make sense, but only in a very specific way: namely, in the sense of a separate system of change and inheritance. Note that I use the term “system” or “process” to follow Kroeber with his late assertion that he does not regard culture as a “substance” (1952: 4, 22). With this I do not want to decide whether the ontological status of ‘culture’ has to be interpreted in a realistic or nominalistic manner. Do beauty or culture exist in themselves or do they merely exist in concrete beautiful things and culture bearing individuals? Either way one can ascertain that culture exists as an inter-individual process or system, a system of change and heredity. In a similar sense, we can say that evolution is a process or system of change that exists, even though individual organisms vanish, without regarding evolution as a specific substance, an extra entity existing in addition to and in the same sense as the evolving entities. Kroeber (1916a: 26).

Heredity, Culture, and the Boundaries of Anthropology in the Work of Alfred L. Kroeber

Darwinism, “one of their fundamental achievements has been the involuntary confirmation by real knowledge of an idea first clearly grasped by a Darwinian theorist.” 22 Despite Weismann and despite Mendelism, the principle of the inheritance of acquired characteristics was still quite popular in the first 20 years of the 20 th century. Because of this, Kroeber called his paper “Inheritance by magic,” since “if found in the minds of uncivilized people,” the belief in the inheritance of acquired characteristics “would be described as belief in sympathetic magic.”23 As one might expect from a trained cultural anthropologist of that time, Kroeber wants to know why the people had such an ‘irrational’ belief—‘irrational’ from his point of view. He cites two motivations for the belief in the inheritance of acquired characters: first, Lamarckian palaeontologists (as well as Mendelians) maintain that Darwinism cannot explain the origin of variation. Thus, in order to account for the origin of variation some scientists call the inheritance of acquired characteristics to the rescue. Yet, Kroeber believes that this is not a viable route for Mendelians, since if they move back to Lamarckian inheritance, they run into a severe tension: the “absolute distinction between gamete and zygote which is the kernel and essence of all the new unit heredity seems contradictory of any possible understanding of use inheritance as a process, and leaves it an empty name.”24 Second, the general public and the social scientists stick to inheritance of acquired characteristics for another reason, as Kroeber states: they stick to it since they still do not distinguish between culture and race (synchronic perspective) and between cultural change and biological evolution (diachronic perspective) in a “consistent” manner. They confuse culture and nature. 25 According to Kroeber, this confusion is caused by the assumption that cultural change, i.e. civilization, is evidence for and is causally linked to biological evolution. In Kroeber’s words, it arises from the assumption that “the acquisition of greater wealth or learning or skill by one group is evidence of a superior faculty for such acquisition inborn in that group through organic heredity.”26 This is what Kroeber calls the “fallacy that the social is organic.” 27 Those who “nominally” employ culture but regard it nonetheless as “ultimately, and in general directly, resolvable into organic factors,” are subject to this fallacy.28 And the cause for this fallacy is the belief in Lamarckian inheritance. And it is true that for instance Herbert Spencer (1855), the most influential Lamarckian with respect to mental traits, assumed that civilization is correlated with biological evolution. According to Spencer, civilization can only be explained by reference to Lamarckian inheritance, where ‘nurture’ becomes ‘nature’ in each generation, leading to innate differences between races.29 New behavioural patterns become habits, which become instincts —via inheritance of 22 23 24 25 26 27 28

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Kroeber (1916a: 27). Kroeber (1916a: 38). Kroeber (1916a: 30). Kroeber (1916a: 31). Compare Kroeber (1916b: 295; 1916c: 370; 1917: 163). Kroeber (1916a: 33). Kroeber (1916a: 36). Kroeber (1916a: 37). —The influence of Boas is evident, since it was Boas who first stressed that culture, language, and race (i.e., the genetic endowment of people) do not covary. See Boas (1894), or Boas (1911). See Richards (1987) and Gissis (2005) on Spencer.

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acquired characteristics; these then play a role in the genesis of new behavioural patterns, which become habits, which then in turn become instincts, and so on. The explanation of the evolution of such mental abilities like intelligence, moral sense, or musical sense, is one of the reasons why Spencer opposed Weismann’s claims about the all-sufficiency of natural selection, which totally excluded inheritance of acquired characteristics. Yet, as part of the well-known debate about the all-sufficiency of selection, Weismann had already answered that Spencer ignores that tradition is an alternative to his Lamarckian explanation. In an essay on music in animals and man, Weismann (1892) claimed that we do not need Lamarckian inheritance to explain the evolution of man’s capacities and achievements, if we admit that there is tradition. According to Weismann, Spencer and others confused achievements (culture or cultural change) with innate abilities (nature or biological evolution). They thus ignore that the first can change without the latter. Weismann illustrated his point with the following thought example: is it possible that there was a Mozart in Samoa, a person with a musical sense or innate ability equal to Mozart’s? According to Weismann, it would indeed be possible. But since the hypothetical “Samoaner Mozart” could not build on already accumulated musical traditions and the corresponding culturally transmitted abilities, it was not possible that the Samoaner Mozart expressed his high musical sense the way the real Mozart did. Kroeber acknowledged Weismann’s essay and heavily relied on it, but regarded it as “a brilliant miss,” since in the end, Weismann “hastened to the inconsequent conclusion that faculties are probably different after all.”30 THE RELATIONSHIP BETWEEN INORGANIC, ORGANIC AND SUPERORGANIC CHANGE It follows from Kroeber’s account that cultural evolution can proceed independently from biological evolution. Kroeber expressed this claim most clearly in the following figure 2:

Figure 2. The relationship between inorganic, organic and superorganic change. Source: Kroeber (1917) p. 211. The continuous line denotes the inorganic, the broken line the organic, and the dotted line the superorganic.

Kroeber presents the graph in order to stress that the lines, representing the three different systems of change (inorganic, organic, and superorganic) develop independently from each other. The important point is B, the first human that was able to learn socially from others; C would be the ‘primitive’ man and D the present moment.31

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Kroeber (1916a: 37).

Heredity, Culture, and the Boundaries of Anthropology in the Work of Alfred L. Kroeber

With this, Kroeber opposed what I would like to call racist hereditarianism.32 The latter regarded the synchronic and diachronic behavioural differences between groups of people as being correlated with and mainly caused by innate differences in ability to produce these cultural differences. Thus, greater wealth and power of one group of people is due to higher innate intelligence. In a diachronic perspective, every cultural change (civilization) is then accompanied by a change in innate endowment. This is what Kroeber denies. 33 But note that, by assuming an inborn faculty of man for civilization and by assuming innate individual differences, Kroeber accepted the hereditarianism of his time. He merely rejected its racist version. 34 In addition, by looking at culture in this manner, cultural inheritance —symbolized by the dotted line—emerges as the very process that makes culture ‘superorganic’. If civilization and biological evolution are as decoupled as Kroeber assumes, then culture becomes clearly visible as a separate, second system of inheritance and change. In the end, culture is conceived as being opposed to biological heredity (culture as superorganic) and, at the same time, it is conceived as heredity of another sort. THE JOINING OF HANDS ACROSS THE GULF Given that we can replace Lamarckian inheritance of acquired characteristics with social or cultural inheritance, Kroeber assumes that “[b]iology and history can join hands in alliance across the gulf that separates them.”35 From a close intertwining interaction of culture and nature in the concept of Lamarckian inheritance of acquired characteristics, we moved with Kroeber to a strict separation of nature and culture on the basis of the concept of hard inheritance. For Kroeber, this conceptual separation is linked to a disciplinary one: biologists should limit their study to biological heredity and the respective organic mental faculties and should leave the explanation of the superorganic culture to the historically working anthropologists. To return, where we started: 31

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Kroeber defined “[h]eight above the base” as “degree of advancement, whether that be complexity, heterogeneity, degree of coordination, or anything else” (Kroeber 1917: 211; Emph. added). A page later, he refers to the increase in number of cultural items and complexity of social organization as the things that distinguish us from the Neandertal people as example of the primitive man. The terms advancement or progress pop up here and there in 1917 and also in other papers. Despite these progressivistic wording, Kroeber tries to distance himself from progressivism by stressing: that “[n]othing is more erroneous than the wide-spread idea and oft-repeated statement that the savage is only a child” (Kroeber 1910: 445), a statement that directly leads to a critique of Darwin and like-minded thinkers who claim that the “savage is in a stage intermediate between the higher animals and ourselves.” (ibid.) Kroeber also stresses that “[a]ll men are totally civilized” (Kroeber 1915: 286) and that he does not use the term civilization for “high” civilization, since for him it makes perfectly sense to talk about “Apache civilization” (Kroeber 1918: 355), which includes their language, their kinship systems, habits, religion, diet etc.—The just cited examples give a mixed message for the question whether Kroeber was still progressivist and less radical in terms of cultural relativism than his teacher Boas. A precise answer is, however, not central for the issues raised here, even if it is important in its own right; it has to wait for another occasion. Kroeber also opposed eugenics, for instance, in Kroeber (1916a: 34-37; 1916c: 370; 1917: 188-9). If eugenics is understood as Kroeber did, that is, in a narrow way as assuming that progress cannot be achieved by social reform (hereditarian eugenics), then it also ignores the possibility of long-term human betterment by cultural inheritance. If eugenics is understood to include Lamarckian points of view, then it reduces culture to environmental influence that is projected into the next generation via biological inheritance. Cooke (1998) suggests that eugenics was Lamarckian (kind of soft eugenics) before 1915 and was predominantly hereditarian afterwards. Kroeber does not say that he can empirically prove that he is right. He merely states that the others cannot prove that they are right. See for instance Kroeber (1916a: 34).

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if there is something superorganic, anthropologists do not have to turn into biologists. Instead, biologists are invited to “join them in a cooperative effort to establish the exact nature and the precise limits of the organic and the superorganic.”36 In the end, Kroeber’s plea for a coalition became true, for instance, when Thomas H. Morgan indeed joined in. Already in 1924, in a paper called “Human Inheritance,” and again in his “The Scientific Basis of Evolution,” he presents social evolution and its peculiar process of “inheritance” as Ersatz for Lamarckian inheritance of acquired characteristics, for which he sees no experimental evidence.37 Social inheritance can be such an Ersatz precisely because it leads to the same effects, i.e. because it is functionally equivalent: efforts to change or to learn during one’s lifetime are heritable and thus not pointless from an inter-generational, evolutionary point of view. Consequently, Morgan advocates the same interdisciplinary division of labour between geneticists and anthropologists Kroeber asked for.

3. Consequences for the concept of culture and the history of hereditarian thinking Even if the concept of culture is still subject to controversial debates, not much has changed with respect to Kroeber’s claim that culture is a system of change that is maintained via a distinctive inter-individual, trans-generational process of cultural inheritance. In this sense culture is even today conceived as a thing sui generis, as autonomous. Let me illustrate this last point in a systematic way by distinguishing between three theoretical roles the concept of culture has played up to the time of Kroeber’s boundary work. THREE THEORETICAL ROLES OF THE CONCEPT OF CULTURE Without much further historical argument, I want to claim that up to the 1920s, with respect to the dichotomy between culture and nature, there have been three major theoretical roles the anthropological concept of culture played in the explanation of behaviour: (C 1) Culture has often been understood as an explanandum: something that is to be explained, by nature or nurture or both of them. I count Tylor’s classic anthropological definition of culture as an exemplar of this category: “Culture or Civilization, taken in its wide ethnographic

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Kroeber (1916a: 35). In Kroeber (1916a: 36 and 1917: 189-192) he therefore refers to Galton as being right in claiming that “between individuals mental faculties are inherited in the same ration and degree, and therefore presumably in the same manner, as physical traits […] But it is an entirely uncompelling inference when he then proceeds to explain the diversity between the attainments of social groups such as ancient Athenians, modern Englishmen, Africans, and Australian natives, as due to differences between the average inherited faculties of the bodies of men carrying the civilizations of these social groups” (Kroeber 1916a: 35). “That heredity operates in the domain of mind as well as that of the body, is one thing; that therefore heredity is the mainspring of civilization, is an entirely different proposition, without any necessary connection, and certainly without any established connection, with the former conclusion” (Kroeber 1917: 192). Kroeber (1916a: 39). Kroeber (1916b: 295). Morgan (1932: 187-217)

Heredity, Culture, and the Boundaries of Anthropology in the Work of Alfred L. Kroeber

sense, is that complex whole which includes knowledge, belief, art, morals, law, custom, and other capabilities and habits acquired by man as a member of a society. ”38 (C 2) Franz Boas is well known to have initiated a change to culture as a more-or-less important factor in the generation of behaviour of individuals. Culture has become an explanans: culture helps explaining behaviour, but has to be distinguished from other factors, like race, in such an explanation of behaviour. 39 (C 3) Kroeber went a decisive step further. He explicitly took culture as a system of change and inheritance in its own right, one that relies on social heritage. Culture thus became again an explanandum, but a new one. And although the early Kroeber thought that culture is also the only explanans for culture as explanandum (only culture explains culture), the late Kroeber admitted that many factors are involved in bringing about the inter-individual system of change and inheritance he called culture.40 It is this last step that I wanted to stress, since it is usually ignored, e.g. even by Stocking, who is well-known for his work on the history of anthropology (especially on Boas, Kroeber, Lamarckism in social science, and the culture concept) and by Cravens and Degler, who are well known for their work on the history of hereditarian thought. Although Stocking, for instance, realizes that Kroeber radicalised Boas approach and further developed the concept of culture, he looks at the concept of culture through a Boasian lens and does not clearly distinguish the second and third way of using the concept of culture. He writes for instance that Boas’ and Kroeber’s concept of culture provided “a functionally equivalent substitute for the older idea of ‘race temperament’. It explained all the same phenomena, but it did so in strictly non-biological terms, and indeed its full efficacy as an explanatory concept depended on the rejection of the inheritance of acquired characters.”41 This is misleading. Boas and Kroeber, first of all, did not have the same concept of culture, since in Kroeber’s hands culture became a system of change and inheritance in its own right. Secondly, Kroeber’s concept did not simply explain the same phenomena, since the concept of culture changed its theoretical role—from an explanans to an explanandum.42 38

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Tylor (1871: 1). As far as I know, the ‘acquired’ in Tylor’s definition bears no systematic role in his account and it is not evident—without further analysis—that it means cultural inheritance in the sense Kroeber means it. Nonetheless, it should be mentioned that Tylor, as does Kroeber, says that he considers questions of race as practically irrelevant for his goals (Tylor 1871: 7). See Stocking (1968: 212-220). My point holds even if Boas sometimes pointed to social learning as part of culture, since he also did not put an emphasis on it as a central aspect. He predominantly regards culture as a special kind of environment, a social environment that influences individual development. This might be the reason why Stocking uses the term “cultural determinism” synonymous to “behavioural determinism” or the “cultural determination of behavior,” i.e., in the sense of ‘culture explains behavior’ and not in the more radical sense ‘culture explains culture’. Nonetheless, Kroeber uses culture in the other two ways vis-a-vis the one he added. This is most evident in Kroeber (1918). Stocking (1968: 265). Cravens (1978) and Degler (1991) also use the term culture mainly for an environmental factor in the development of individuals, even though Degler comes close to my point, when he writes that Kroeber demanded “more than a mere change in assumptions as Boas had done; he was insisting upon a new mode of explanation for human behavior” (Degler 1991: 94). Freeman (1983) probably comes closest to my point of view, but without distinguishing between different roles of the culture concept.

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CONTEMPORARY DEBATES AND A FOURTH ROLE FOR CULTURE The distinction between the three theoretical roles of the culture concept is not only helpful to revise the history of the concept of culture. It is even helpful to understand contemporary debates about the relationship between culture and nature. First, Boas concept is the one that still dominates nature-nurture-debates in psychology and behavioural genetics. Kroeber’s concept, however, is the one that is used in debates about man’s place in nature and in those about the interaction between biological evolution and cultural change. In other words, the first is used in developmental contexts, the second in evolutionary contexts. Both contexts involve different questions of interactions. In addition, there are people who still ignore Kroeber’s concept. Evolutionary psychologists, for instance, reduce culture to a mere triggering condition of innately specified behavioural patterns. Cosmides and Tooby, thus, define culture as “any mental, behavioural, or material commonalities shared across individuals […] regardless of why these commonalities exist.” 43 Culture is here the explanandum, the specific attributes of a group of a people. It is not a factor in the explanation of what people do; it is the explanandum, the phenomenon to explain. At the same time, it is not an explanandum in Kroeber’s sense. On the contrary, it is considered as merely ‘evoked’ through experience in the world. Thus, culture (mental, behavioural, or material commonalities) is basically innate. It can be reduced to the decisive influence of innately specified characteristics of mind. The influences of the natural and social environment are mere triggering conditions. Yet, the social environment is what others, dual-or-multiple-inheritance-theorists 44 as well as ‘standard social scientists,’ as evolutionary psychologists like to say, call culture: a distinctive factor in the explanation of behaviour, that is, an explanans, and a special explanandum at the same time, namely a separate second system of inheritance of ideational units that can and needs to be studied in its own right. And this is exactly what Kroeber wanted to say —with the help of Weismann’s concept of hard inheritance. And this is why I regard his case as historically and systematically important. In a way, Kroeber’s case and the three different usages of the concept of culture offered above show that evolutionary psychology falls back to the 19th century concept of culture: used by Tylor and long ago abandoned in anthropology. Note that what I have said so far holds even though—from our contemporary perspective—we might question whether a psychic unity of man is justified, since mind (or mental abilities) is itself a developmental product of nature and nurture.45 No child is born with a ready-made mind. Culture would then start off from a mere genetic unity of mankind. 42

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That Kroeber wants to distinguish his concept of culture from Boas’ is also evident from Kroeber & Kluckhohn (1952), a review of various definitions and concepts of culture, for which they became famous later on. In this they put Boas together with Tylor into the category of “descriptive” definitions using enumerations and Kroeber into this and into a second category of “historical” definitions: definitions with “emphasis on social heritage,” even though the early papers of Kroeber at centre here are ignored in this review. Cosmides & Tooby (1992: 117). Such as Cavalli-Sforza & Feldman (1981), Boyd & Richerson (1985), Durham (1991), Richerson & Boyd (2005), or Jablonka & Lamb (2005) and the niche construction theory (Odling-Smee, Laland & Feldman 2003). This has been stressed by John Dupré (2004, 1993).

Heredity, Culture, and the Boundaries of Anthropology in the Work of Alfred L. Kroeber

With dual-or-multiple-inheritance-theorists the last argument (that evolutionary psychologists ignore that culture is a second system of inheritance) can even be taken further since these approaches claim that culture interacts with the biological system of inheritance, at an ontogenetic as well as at a phylogenetic level, influencing thereby the distribution of genes in subsequent generations. With this, these approaches actually introduce a fourth theoretical role of the culture concept: (C 4) Culture becomes a factor not only in the ontogenetic development of individuals but a factor in the phylogenetic process of culture and nature interacting in the evolution of organisms that have a body, a mind and a culture. James M. Baldwin (1861-1934), and others at the beginning of the 20th century, made a similar usage of culture as a factor in the evolution of organisms. 46

THE HISTORICAL IMPACT OF THE CONCEPT OF HARD INHERITANCE I will now explicitly drive home the main point of this essay with respect to the cultural history of heredity. What was the historical impact of Weismann’s concept of hard inheritance on how the relationship between nature and culture was conceived? I want to defend the following three claims: (H 1) First, inheritance of acquired characteristics or soft inheritance in general allowed for soft hereditarianism. On the basis of soft inheritance, one could be a hereditarian and give culture a significant role to play in the evolutionary process, since the hereditary material itself was considered as being soft, that is, malleable by cultural or environmental influences. Culture, and that includes education and social reform, could play a role without the need to refer to social or cultural inheritance. (H 2) Given Weismann’s concept of hard inheritance, this possibility was gone. As long as cultural inheritance is ignored, hard inheritance leads to a hard hereditarianism, a picture where cultural and environmental influences cannot exert any influence on the evolutionary process. One could reduce everything to biological inheritance by combining the continuity of the germ plasm with the view that the germ plasm is the sole hereditary material transferred down the generations of individuals. (It was the latter, that has often wrongly been attributed to Weismann as I have shown above). Both of these claims are more or less part of the received view on the impact of soft and hard inheritance. Yet the received view also takes it for granted that the concept of hard inheritance was therefore partly responsible for the vogue of hard hereditarianism—a view where nurture (natural environment and culture) does not play any explanatory role anymore. And indeed, at least until the end of World War I, geneticists as well as the general public predominantly believed in the 46

The relationship between Baldwin, Boas and Kroeber would deserve close investigation here but has to wait for another occasion. Consult Simpson (1953) or Weber & Depew (2003) for more on the ‘Baldwin effect’.

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power of biological inheritance to explain behavioural differences (within and between groups). At least, they usually did not say anything to the contrary.47 This is why Bowler, for instance, writes that the “social consequences of biological determinism” are not a product of social Darwinism or Darwinism as such, but a product of the rise of genetics, which “represents the collapse of a preDarwinian ‘developmental’ view of nature with consequences that were at least as profound as those associated with the initial conversion to evolutionism.”48 I depart from this received view by claiming that: (H 3) Since nothing in the concept of hard inheritance prevented one from acknowledging cultural inheritance, the connection between the concept of hard inheritance and biological determinism (or hard hereditarianism, choose your term) is neither necessary nor historically true, as the examples of Weismann and Kroeber show. The concept of hard inheritance was thus not exclusively linked to hereditarianism, or, to put it in other words, the concept of hard inheritance did not have an unambiguous, unidirectional historical influence. To the contrary, it had an important historical impact on the rise of the concept of culture as a superorganic, separate system of change and inheritance: a concept of culture that led to the break of the hereditarian consensus in the US of the early 20 th century, and that thereby helped establish the boundaries of anthropology. This culture concept flourishes until today, at least in anthropology. Maria E. Kronfeldner Max Planck Institute for the History of Science, Berlin [email protected]

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See Ludmerer (1972), Kevles (1985), Barker (1989), Paul (1995: 40-49). The concept of hard inheritance surely was not the only reason for the dominance of hereditarianism, but it is usually taken as one of the reasons. Part of the hereditarian bias might have been due to the growing scientific success of genetics as an experimental science in explaining biological heredity. Part of it might have been due to sociopolitical views, part of it due to institutional developments, as Cravens (1978) suggests. Usually, Johannsen is cited as an early exception to the rule of ‘geneticists were hereditarians’, and Morgan and Jennings as exceptions of the 20s, e.g. in Paul (1995: 115-117). Bowler (2003: 24).

Heredity, Culture, and the Boundaries of Anthropology in the Work of Alfred L. Kroeber

References Barker, D. 1989. “The biology of stupidity: Genetics, eugenics, and mental deficiency in the inter-war years.” British Journal for the History of Science 22(1): 347-375. Bidney, D. 1944. “On the concept of culture and some cultural fallacies.” American Anthropologist 46: 30-44. ————. 1965. “The contribution of A. L. Kroeber to contemporary anthropology.” International Journal of Comparative Sociology 6: 266-277. Boas, F. 1894. “Human faculty as determined by race.” Proceedings of the American Association for the Advancement of Science for the 43. Meeting: 301-327. ————. 1911. The Mind of Primitive Man. New York: Macmillian. Bowler, P. J. 1989. The Mendelian Revolution: The Emergence of Hereditarian Concepts in Modern Science and Society. Baltimore: John Hopkins Press. Bowler, P. J. 2003. Evolution: The History of an Idea. Berkeley: University of California Press. Boyd, R., & Richerson, P. J. 1985. Culture and the Evolutionary Process. Chicago: University of Chicago Press. Cavalli-Sforza, L., & Feldman, M. 1981. Cultural Transmission and Evolution: A Quantitative Approach. Princeton: Princeton University Press. Cooke, K. J. 1998. “The limits of heredity: Nature and nurture in American eugenics before 1915.” Journal of the History of Biology 31: 263-278. Cosmides, L., and J. Tooby. 1992. “The psychological foundations of culture.” In J. H. Barkow, L. Cosmides and J. Tooby (eds.), The Adapted Mind: Evolutionary Psychology and the Generation of Culture. Oxford: Oxford University Press. 19-136. Cravens, H. 1978. The Triumph of Evolution: American Scientists and the Heredity-Environment Controversy 1900-1941. Baltimore: University of Pennsylvania Press. Darlington, C. B. 1959. Darwin’s Place in History. Oxford: Basil Blackwell. Darnell, R. D. 1998. And Along Came Boas: Continuity and Revolution in Americanist Anthropology. Amsterdam: John Benjamins Publ. Company. Degler, C. N. 1991. In Search of Human Nature: The Decline and Revival of Darwinism in American Social Thought. New York and Oxford: Oxford University Press. Dupré, J. 2001. Human Nature and the Limits of Science. Oxford: Oxford University Press. ————. 2004. “On human nature.” Journal of the Slovakian Academy of Sciences 13: 109-122. Durham, W. H. 1991. Coevolution: Genes, Culture, and Human Diversity. Stanford: Stanford University Press. Freeman, D. 1983. Margaret Mead and Samoa: The Making and Unmaking of an Anthropological Myth. Canberra: Australian National University Press. Gissis, S. B. 2005. “Herbert Spencer’s two editions of the Principles of Psychology 1855 and 1870/72: Biological heredity and cultural inheritance.” In S. Mueller-Wille and H.-J. Rheinberger (eds.), A Cultural History of Heredity III: 19th century and Early 20th century. Max Planck Institute’s preprint series no. 294. 137-151. Harris, M. 1968. The Rise of Anthropological Theory. A History of Theories of Culture. London: Routledge. Herskovits, M. J. 1948. Man and His Works. The Science of Cultural Anthropology. New York: Knopf. Hinsley, C. 1981. Savages and Scientists. The Smithsonian Institution and the Development of American Anthropology, 1846-1910. Washington, D. C.: Smithsonian Institution Press. Jablonka, E. & Lamb, M. 2005. Evolution in Four Dimensions. Genetic, Epigenetic, Behavioral, and Symbolic Variatin in the History of Life. Cambridge, MA: MIT Press. Kaplan, D. 1965. “The superorganic: Science or metaphysics.” American Anthropologist 67: 958-976. Kevles, D. J. 1985. In the Name of Eugenics. Genetics and the Uses of Human Heredity. New York: Knopf. Kroeber, A. L. 1910. “The morals of uncivilized people.” American Anthropologist 12(3): 437-447. ————. 1915. “Eighteen professions.” American Anthropologist 17(3): 283-288. ————. 1916a. “Inheritance by magic.” American Anthropologist 18(1): 19-40. ————. 1916b. “Heredity without magic.” American Anthropologist 18(2): 294-296. ————. 1916c. “The cause of the belief in use inheritance.” The American Naturalist 50(594): 367-370. ————. 1917. “The Superorganic.” American Anthropologist 19(2): 163-213. ————. 1918. “Heredity, environment, and civilization: Attitude of the anthropologist toward race; work in his own special field traces back the history of man culturally and psychologically as response to civilization.” The American Museum Journal 28(5): 351-9. ————. 1919. “On the principle of order in civilization as exemplified by changes of fashion.” American Anthropologist 21(3): 235-263.

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————. 1952. The Nature of Culture. Chicago and London: The University of Chicago Press. ———— and C. Kluckhohn. 1952. Culture. A Critical Review of Concepts and Definitions (Vol. XLVII). Cambridge, MA: Peabody Museum. Kroeber, T. 1970. Alfred Kroeber. A Personal Configuration. Berkeley: University of California Press. Kuklick, H. 2004. “Who owns the past?” Current Anthropology 45: 292-293. Ludmerer, K. M. 1972. Genetics and American Society. Baltimore and London: Johns Hopkins University Press. Mayr, E. 1982. The Growth of Biological Thought. Diversity, Evolution, and Inheritance. Cambridge, MA: Harvard University Press. Morgan, T. H. 1924. “Human Inheritance.” The American Naturalist 58: 385-409. ————. 1932. The Scientific Basis of Evolution. London: Faber & Faber Limited. Odling-Smee, F.J., K. Laland and M. Feldman. 2003. Niche Construction. The Neglected Process in Evolution. Princeton: Princeton University Press. Patterson, T. C. 2001. A Social History of Anthropology in the United States. Oxford and New York: Berg. Paul, D. 1995. Controlling Human Heredity. 1865 to the Present. Atlantic Highlands: New Jersey: Humanities Press. Peel, J. D. Y. 1971. Herbert Spencer. The Evolution of a Sociologist. London: Heinemann. Richards, R. J. 1987. Darwin and the Emergence of Evolutionary Theories of Mind and Behavior. Chicago and London: University of Chicago Press. Richerson, P. J. and R. Boyd. 2005. Not by Genes Alone. How Culture Transformed Human Evolution. Chicago: University of Chicago Press. Simpson, G. G. 1953. “The Baldwin effect.” Evolution 7: 110-117. Spencer, H. 1855. Principles of Psychology (1st ed.). London: Williams & Norgate. Steward, J. H. 1973. Alfred Kroeber. New York and London: Columbia University Press. Stocking, G. W. 1968. Race, Culture and Evolution. Essays in the History of Anthropology. New York: Free Press. Thoresen, T. H. 1975. “Paying the piper and calling the tune. The beginnings of academic anthropology in California.” Journal of the History of the Behavioral Sciences 11: 257-275. Tylor, E. B. 1871. Primitive Culture. London: Murray. Weber, B. H. and D. J. Depew. 2003. Evolution and Learning. The Baldwin Effect Reconsidered. Cambridge, MA: MIT Press. Weismann, A. 1892. Gedanken über Musik bei Thieren und beim Menschen. Aufsätze über Vererbung und verwandte biologische Fragen. Jena: Gustav Fischer. Zirkle, C. 1946. “The early history of the idea of the inheritance of acquired characters and of pangenesis.” Transactions of the American Philosophical Society 35: 91-151.

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Comments on Daniel Kevles’ and Maria Kronfeldner’s papers Edna Suárez

Within the context of our workshop, the papers of Dan Kevles and Maria Kronfeldner could not be more different than they are. One deals with plants, the other with man; one deals with property rights and the other with the institutionalization of disciplines and the history of concepts. One extends along a Braudelian mid-term durée (the realm of general economic trends), and the other along the trajectory of an individual and an academic discipline. Thus, I hope you are not expecting me to find relations among them, though the title of this session, “Contexts of Heredity” may provide me, at the end, with some common questions to both exponents. I am going to begin with Dan Kevles’ paper. His reconstruction of the many ways in which animal and plant improvers sought protection to what they saw as their “intellectual property,” offers us an explanatory framework for the pre-history of the patents of living beings in the United States. As Chakrabarty won the Supreme Court case in 1980, to be warranted a patent for his genetically modified bacterium, many in the United States and around the world wondered not only if the US Congress should have been involved in a new patent legislation, but even more, what historical, economic and even ethical or moral transformations had allowed the Supreme Court judges to take that narrow decision. As the Century of the Gene closed with a mixture of good and bad news for the genetic engineering industry, the growing number of patents of living beings or their parts, or even the patent of bio-macromolecules by US firms, universities and governmental agencies, stands still as a highly controversial issue around the world. What I would like to stress is an outsider’s view of the history of the US, that is, its exceptionality regarding the patentability of life and, more generally, the importance of granting intellectual property rights since more than a century ago. Trademarks and some kinds of property rights developed in different ways across the industrialized countries during the XIXth Century. When I speak of the exceptionality of the US condition regarding the patentability of life and, more broadly, the history of intellectual property rights, I attempt to redirect our attention to what might be seen as an almost “natural” search for financial return by part of the breeders and improvers of animals and plants in XIXth century United States. Clearly, it was not a natural right, as is manifested in the confrontation of values implicated in the “products of nature doctrine.” It is an historical phenomenon, a general economic trend, but also a question of values that came together with the transformation of the American market, and which needs an explanatory framework. I think Daniel Kevles is particularly well situated to give us that framework and that is what I would like to ask of his recent research in what is part of the pre-history of patents. Dan gives us some clues to the exceptionality of the US condition. He argues that before the Civil War “markets in agricultural stock were largely local, and the seed, nursery, and animal breeding industries were only incipient” (p. 3), thus the protection of intellectual property found its way only during the latter third part of the century, as “(r)egional and national agricultural

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markets emerged with the construction of railroads and amid increasing urban demands for meats, fruits, and vegetables, as well as ornamental plants” (p. 4). It is in the context of the competition for markets after the Civil War, of the need to capture financial returns and do business at long distances, that the animal and plant breeders, orchardists and nurserymen, began to seek the “protection of their rights.” The old-regime system of prestige, reliability and admiration of products by the local community, were not sufficient any more. But the succeeding of the advocates of property, of course, did not take place without enforcement measures and even casualties. Dan refers an important triumph of the seed industry in 1924, when they succeeded in their campaign to ban the distribution of free seed to farmers by the US Department of Agriculture and the US Patent Office. My questions to Dan do not have the intention to deviate our analysis to the economic history of the US. Rather, they point to the need of awareness of the historical specificity of these changes. Moreover, I think this issue has something important to tell us about the cultural history of heredity. It is convincing that the development in regulations of intellectual property of plants had almost nothing to do with the rise of Mendelism, since many of the claims of the agricultural industry had to do more with crafts and with the chance finding of mutations and sports in the American fields. But, along the history of the reification of heredity, as Carlos Lopez has called it, a central phenomenon seems to have taken place when heredity began to be considered a commodity, a tendency that has not expired today. Could we say more about the “commodification” of heredity as part of the long-durée history of heredity? I hope that the nature of my comments will make clear that we are not expecting for general answers, but more modest attempts to deal with the specificity of cultural—including of course values—and economic contexts. Maria Kronfeldner’s paper looks in another direction. The context, although also referring to the United States, points to the academic scene and the construction of concepts as part of the boundary work developed by cultural anthropologists in the first part of the XXth century. Here, however, we have the heavy load of hereditarianism of several kinds (including of course eugenics and racism) as a more explicit factor of her historical and conceptual analysis. I find Maria’s distinction of three theoretical roles of the concept of culture illuminating for understanding recent—and not so recent—developments in the intersection of evolution and development. The connection between the concept of culture used by evolutionary psychology and the early concept of Tylor, accounts for the impression that a somewhat old-fashioned concept of culture is at play in this approach. Even more fruitful is her critique of the received view, namely, that hard heredity—that is, the rise of genetics—is responsible for the collapse of the developmental view of nature and for the expulsion of nurture. Maria’s argument, instead, tries to direct our attention to the fact that linked to the idea of hard heredity, in Kroeber’s later work, cultural or social inheritance were certainly expelled from evolutionary accounts. On the developmental level, however, hard inheritance provided the framework of coalition and opposition that anthropology needed in order to distinguish itself from biology and in particular genetics. Cultural inheritance, even if it is embodied in biological individuals, is autonomous from hard genetic inheritance. Heredity was

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always more than genetics. In this particular case the “hardening” of inheritance and its restriction to genetics dialectically provided a space for cultural anthropology to flourish in the academic scene. Maria’s analysis is convincing, in part because it give us a framework to account for the differences between so many versions of hereditarianism in the XXth century, and to clarify some terms in the nature-nurture debate. Though I should confess that I am deeply ignorant of the history of anthropology and I imagine her views might be contested by those who have interpreted the superorganic nature of culture in different terms. My questions, instead, refer again to the context of these developments in American Anthropology. The drawing of disciplinary boundaries, and their changes, can tell us a lot about the actor´s views of themselves. But I long for a more detailed account of the relations of Kroeber and his contemporaries and, more importantly for a comprehension of the century of the gene, I would like to have more details of the connections between the rise of cultural anthropology in the United States and the prevalence, for instance, of dual-inheritance-theorists in contemporary debates. Edna Suárez Universidad Nacional Autónoma de México/ Max Planck Institut für Wissenschaftsgeschichte [email protected]

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Producing Identity, Industrializing Purity: Elements for a Cultural History of Genetics Christophe Bonneuil

In December 1910, just one year after the coining of the terms “gene,” “genotype” and “phenotype” by Wilhelm Johannsen in his Elemente der exakten Erblichkeitslehre (Johannsen, 1909), a symposium on the “Genotype Hypothesis” was one of the key attractions of the meeting of the American Society of Naturalists at Cornell. Leading figures of genetics and plant breeding 1 discussed the various aspects of Johannsen’s “principle of pure lines, as a true analytical implement, as an indispensable method of research in heredity” (Johannsen, 1911, 143). Even more so than “unit factors” or “genes,” “types,” “permanency,” “stability” and “purity” were the buzzwords of this meeting. Almost everybody endorsed then the idea proposed a few years earlier that “the study of the behavior of pure lines is the basis of the science of heredity” (Johannsen, 1903, 9). The papers of this session, published in American Naturalist were continued by follow up discussions in Science (Jennings, 1911b; Shull, 1912a and 1912b). 2 This stream of papers enthusiastically used, promoted and extended Johannsen’s concepts of pure line and genotype. Jennings, who had experimented on the effects of selection on pure cultures of Paramecium,3 explained that “we need badly a term that will include ‘genotypically identical’ series of forms” (Jennings, 1911b, 842). This is why he turned Johannsen’s ‘structural’ genotype concept round to a ‘populational’ one (“a concrete, visible group of organisms” having “the same combination of hereditary characters” and proposed an extended definition of pure lines: (1) in case of vegetative reproduction [of unicellular or pluricellular organisms], (2) in at least some cases of parthenogenesis (where no reduction division occurs), (3) in case of self-fertilization of homozygotic organisms [pure lines stricto sensu in Johannsen’s sense], (4) in case of inbreeding of a group of genotypically identical homozygotic organisms (Jennings 1911b, 841). As early as 1904, Shull had also extended Johannsen’s stricto sensu definition of “pure lines” to any “population relating through budding or other method of vegetative reproduction” (Shull, 1905 quoted in Shull, 1912a, 27), for which the USDA agricultural scientists O. F. Cook and Herbert J. Webber had recently coined the term “clon” soon to become “clone” (Webber, 1903). As a young scientist willing to play a role at the frontier of genetics, Shull verified Johannsen’s claim that F1 hybrids from homozygotic parents show no more variability than pure lines do. As a breeder, he also hoped to command and control the homogeneity and vigour of F1 hybrids (as compared to populations) and to capture heterozygosis in stable, mass-produced and profitable 1 2 3

The speakers were: W. Johannsen, Herbert S. Jennings, George H. Shull, Edward East, Raymond Pearl, J. Arthur Harris and Thomas Hunt Morgan. I thank Christina Brandt for making me aware of several of these papers. See J. Schloegel, this volume.

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life forms. For these reasons, Shull was no less attracted than Jennings by the search for general conceptualisations for genetically identical groups and he joined in spreading the gospel of “clone,” “biotype” and “genotype” concepts. He undertook to extend the validity of Johannsen’s genotype concept to cross-breeding populations (Shull, 1908 and 1911). He also introduced the notion of “clonal varieties” to label under one and the same category not only potatoes and Paramecium, but also perfectly standard and homogeneous heterozygots such as his F1 hybrid corns from two pure lines, since “in the ‘clone’, it is possible to retain as a permanent feature of the group any purely heterozygous character, as for instance the vigorous constitution dependent upon the stimulation of heterozygosis” (Shull, 1912a, 28). The practices of grafting, budding and other vegetative reproduction techniques were common practices, some being as old as agriculture. What was then the rationale behind the creation of new scientific terms in the first years of the XXth century, that related “pure lines,” “clones” and heterozygous F1 hybrid clone-like varieties? And how can we account for the considerable amount of efforts displayed by geneticists, through hard experimental and statistical work on beans, protozoa or chicken, to separate “fluctuation” as what is caused by the environment from what lies in “genotypical constitution?” Why so much work to construct an “intrinsic” genetic identity of organisms, that could be separated from the influence of time and place and could circulate unaltered in new kinds of scientific and economic networks. In other words, what were the specific historical conditions of this period to call for so huge efforts to engineer and conceptualize genetic sameness, genetic stability and genetic purity? In Purity and Danger, an essay dedicated to cultural attitudes toward “impurities” and “pollutions,” Mary Douglas stated: “I consider as partial any explanation of ritual pollution that would limit itself to only one kind of impurity or only one kind of context” (Douglas, 2001 [1966], 21). Following Douglas’ methodological commandment, this article sketches a cultural history of early XXth century genetics that relates new conceptualizations of the identity and connectivity of organisms that postulated the stability of “types” and of hereditary constituents, the rise of quantity production of new life forms (in biological laboratories, agricultural experiment, fields, hospitals and markets), and new cultural attitudes towards time, space, purity, efficiency and fairness. I will argue that the new framings of heredity and these new life forms were designed in and for a new space of flows, a new matrix of practice and meaning that structured both apparently esoteric scientific investigations on Paramecium or Drosophila, and industrial culture of rationalisation and control. The first section will discuss some of the concepts and analytic tools of this paper: Müller-Wille and Rheinberger’s “epistemic space” and Phillip Thurtle “space of flow,” and Boltanski and Thévenot’s “orders of worth” and how I relate them in my attempts towards a cultural history. The second section analyses the shift from a Darwinian space-time of organic fluxes to an experimental-industrial space-time. The third section documents the quest for purity that pervaded late 19th century and early 20th century biological research.

1. “Epistemic space,” “space of flow” and “worlds of worth” The advent of modern genetics has often been described as the advent of the gene as the unit of explanation and the victory of hard heredity over soft heredity conceptions (Mayr, 1982; Fox-

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Keller, 2000). Its heroic advent is still often narrated as the victory of Mendelians over Biometricians, old school breeders, and Neo-Lamarckists (Provine, 1971; MacKenzie and Barnes, 1979; Buican, 1984). But this focus on Mendelism has obscured the complex relations between mendelian hybridisation research programs on the one side, and “pure line research” programs— associated with a more typological than combinatory thought—developed by De Vries and Johannsen. More generally, the Mendelism-centered view of the sciences of heredity in the first half of the 20th century generated a great numbers of anomalies and generated stories of reluctances, exceptions and intellectual speciations: “rediscoverers” like De Vries seeing isolation and mutation as more important driving forces for evolution than hybridization and more powerful tools for plant breeding (Meijer, 1985), Johannsen having little interest in mendelian crosses as a research strategy (Müller-Wille, 2007), interest in cytoplasmic inheritance and nonMendelian heredity (Sapp, 1987), reluctance to Mendelism based on a physiological and Pasteurian thinking that would turn to be very productive a few decades later in the birth of molecular biology in France (Burian and al., 1988), plurality of breeding strategies (beyond the traditional view of breeding revolutionized by Mendelism, or being an application of it) until the mid 20th century, all of them being perfectly rational when considered as technological paradigms which co-evolved with differentiated bio-socio-economic contexts (Palladino, 1994; Harwood, 1997; Wieland, 2005; Bonneuil, 2006). This paper does not aim at analysing the whole zoo of non-Mendelian (or superficially Mendelian) research schools in early 20th century genetics and breeding and weighting the contribution of each in the progress of genetic knowledge (which clearly did not proceed along a single line). What I rather want to explore, is what the various traditions of research on heredity after 1900 (which profoundly disagreed on the role of genes, cytoplasm and the environment in heredity, on the role of mutations and hybridization in evolution, on the value of Mendel’s laws for practical breeding, etc.) deeply shared at more profound levels. If there is a revolution in the study and mastering of heredity at the turn of the 19th and 20th century, and if this revolution cannot be reduced to the diffusion of Mendelism, I will frame this revolution, on the basis of numerous studies from many scholars, as a shift in the “epistemic space,” the “knowledge regime” and the “worlds of worth” in which heredity was thought and manipulated. Staffan Müller-Wille and Hans-Jörg Rheinberger (2004) have called for the need of a cultural approach to understand wide shifts in “knowledge regimes” on heredity. Such shifts, they argue, cannot be reduced to “‘epistemic things’—in the sense of being determined within individual experimental settings”—or to a paradigm, but rather “depended on a vast, spatial configuration of distributed technologies and institutions connected by a system of exchange: botanical gardens, hospitals, chemical and physiological laboratories, genealogical and statistical archives” (MüllerWille and Rheinberger, 2004, 23). To make sense of such wide shifts, Müller-Wille and Rheinberger have coined the term epistemic space, a more “regional” concept than Foucault’s episteme. This concept seems very fruitful because it provides a lens to map the continent drifts between, say, common traits exhibited by the various “epistemic cultures” 4 of heredity in the early 19th century and common traits exhibited by the various epistemic cultures of heredity in the mid 20th century. These authors have exemplified their “epistemic space” approach with the shift they 4

On “epistemic cultures,” see Knorr-Cetina, 1999.

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see around the middle of the XIXth century when heredity became not only a genealogical (vertical) notion but also a spatial (horizontal) one, i.e the cytological space of hereditary germs in the gametes and fertilized egg. Such a shift was made possible by the development of cell theory, by the assumption of an essential fluidity between all organisation levels of organic life from hereditary elements to species and of the existence of similar mechanisms to account for their transformations, and by a focus of the biological gaze towards the search for “patterns and processes that structure life on the intra-specific level” (Müller-Wille and Rheinberger, 2004, 13). They also argue that the epistemic space of heredity “resided in the heart of capitalist institutions from its very inception” (ibid., 23) and that “the emergence of heredity as a research attractor, as a discursive center, occurred in a knowledge regime that started to unfold when people, objects, and relationships among them were set into motion” (ibid., 13). From a different starting point, Philip Thurtle has come to see a similar connection between changing rationality in thinking about heredity and late 19th century’s wide scale circulation of goods and people. His work understands “the history of the science of heredity as a mutation in cultural practices for dealing with space and time” (Thurtle, 2008). 5 From this perspective, Genetics is a science of mass culture in much the same way that the modern newspaper is a communication medium of mass culture or the urban train station is a transportation medium of mass culture. They all are reliant on the same technologies of transportation and communication, they all create a new conceptual space that denies the importance of place in human interactions, and they all support new ways of folding experiences that will lead to modern conceptions of information. Some would even claim they all privilege a lowest common denominator in order to describe human connectedness over new geographic distances and over radically long periods of time. (Thurtle, 2008)

More precisely, Thurtle explores the cultural conditions of possibilities of what he calls “genetic rationality,” one major aspect of which is the quest for a “genetic identity, the unchanging core of heritable material sealed off from the influence of time and place” (Thurtle, 1996, 2007, 2008). Thurtle goes further in showing how different research programs in genetics and breeding, such as Burbank’s and De Vries’, inhabit (and contribute to make emerge) different types of spaces, and document the emergence of a “new type of space at the turn of the century, a space built on the exchange of manufactured commodities, managed by a host of new informational practices” (Thurtle, 2007). The second industrial revolution allowed for increased profits by exploiting economies of scale and increased circulation of goods, and called for a host of new innovations the way information was collected, stored and processed. This, according to James Beniger was the hallmark of the “control revolution,” which took place from the last decades of the 19th century on, in order to increase production efficiency, ensure the safe distribution of standard goods and raise product awareness among consumers (Beniger, 1986). These developments, argues Thurtle, opened up a new type of space, which he calls (after Manuel Castells) the “space of flows,” a space in which objects circulate intensely from one location to another and are designed to be used in a wide number of places; a space in which the values upheld by exchange came to be promoted at the expense of the values of the specific locations. (Thurtle, 2007). Seen from this wide cultural 5

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perspective, the shift documented by Jean Gayon (2000), from force to organisation in views about heredity at the turn of the century shows striking homologies with a global shift form energy and speed (the industrial revolution) to control (the control revolution) in the same period. A third strand of scholarly work that seems to me particularly useful for a cultural history of genetics is provided by Luc Boltanski and Laurent Thevenot “sociology of worth” (Boltanski & Thévenot, 1999 and 2006; Boltanski & Chiapello, 2006). From a wide range of situations where people are led to justify their actions, these social theorists have abstracted a plurality of logics of justifications (“worlds of worth” or, in French “cité”), each being exemplified by a classic author: civic (Rousseau), market (Smith), industrial (Saint-Simon), domestic (Bossuet), inspiration (Augustine), fame (Hobbes), and connexionism (a new order of worth they found in late capitalism’s neomanagement discourse). These seven “orders of worth” are based upon a “convention of equivalence” that brings together different sets of people and objects and creates a certain kind of commensurability that allows judging them and weighting their worth, but the underlying principles of order differ from one another. For instance in the domestic world of worth (cité domestique), “worth depends on a hierarchy of trust based on a chain of personal dependencies. The political link between beings is seen as a generalization of kinship (…). The person, cannot, in this world, be separated from his/her belonging to a body, a family, a lineage, an estate” (Boltanski & Thévenot, 1999, 370). On the other side, in the industrial world of worth (cité industrielle), worth is based on efficiency for a specific function and the relations between persons (and objects) “can be said to be harmonious when organized, measurable, functional, standardized” (ibid, 373). Notwithstanding its weakness of seeming quite fixed and a-historical grammar, the “worlds of worth” perspective can provide a fresh standpoint on many findings of scholars in the history of heredity. As we shall see, the call for purity, pervading the longue durée history of ideas on heredity, can take quite different forms in different worlds of worth. In a cité domestique, purity might refer to kinship, to a trustworthy keeping of pedigree books whereas in a cité industrielle, purity might be redefined by a particular assay (a measurement of performance or a back-cross). Rather than a structural property (say, a homozygotic constitution) whose value is associated with the possibility of its replication, of its mass-production in a stable state, purity was in the mid 19th century dominated by a mix of domestic, inspiration and fame worlds of worth, and hence valued as something particularly rare and unstable, that needs constant care to be maintained.

2. Out of Darwinism’s space-time: erasing time and space, disciplining organic fluxes, controlling variation Peter Bowler has written about “The eclipse of Darwinism” (Bowler, 1983). More than a mere shift in evolutionary thinking around 1900, this “eclipse” also refers to a wider change in biology’s knowledge regime, from an evolutionary space-time to an experimental-combinatory space-time. Mid 19th century biology, eroding the previous dichotomy of individuals and species, saw life as a property extending both downwards (to the cells and molecules) and upwards (to the species, societies and colonies). Biologists of the generation of Darwin, Haeckel, Galton or Weissmann imagined extensive organic traffics, linking the macro and the micro levels of organisation, linking

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organisms and the environment and linking organisms from different species (cell theory, pangene theory, evolution, interspecific hybridization, acquired heredity, symbiosis, etc.). Natural processes and human activities were seen as similar in nature and interconnected so that the observation of one realm would greatly improve the understanding of the other. 19th century biology’s emphasis was on continuous change, exchange and admixture—rather than on stability, fixity, isolation and purity—as fundamendal properties of life and as driving forces of evolution. Darwin’s pangenesis theory, Quatrefage’s “tourbillon vital,” for instance, “nicely demonstrate the degree to which, by the late nineteenth century, individuals had been resolved in an underlying system of circulating, sub-microscopic entities only to re-emerge as ephemeral and contingent results from the interaction of such entities, both with one another, and with their respective environments” (Müller-Wille, 2007). Early 20th century biologists, on the contrary, put the emphasis on isolation as the driving force of speciation (synthetic theory of evolution) and ceased to view naturally occurring hybridisation and gene flow as a major research object, sought for new typological units reinforcing stability and fixity as an underlying principle of life and turned organisms into purified reagents that experimental strategies put in reaction with one another. While 19th century had framed reproduction as a system of circulating and ever changing elementary entities such as “gemmules,” “pangenes,” “organic units,” and so on (Müller-Wille, 2007), it seems as if 20th century biology had—symbolically and practically—disciplined these circulations, fixed these entities into invariant units (stable genotypes and immutable genes, safe at low mutation frequencies,), and had grasped and redefined heredity, as well as many other biological functions, in terms of predictability of effect in controlled biological reactions. OUT OF HISTORY In mid 19th century, Coleman has shown, “embryology, natural history, evolution theory, even cellular anatomy, were historical disciplines,” and one may add plant and animal breeding and human biometry to this list (Coleman, 1977, 162). But in a few decades, the rise of an experimentalist way of knowing in late 19th century (Pickstone 2000; Rheinberger and Hagner, 1993) was associated with a leaving behind of the ideal of historical explanation in biology. This move was particularly strong in the understanding of heredity, from time and vertical transmission to a timeless combinatory structure keeping its permanency though space. Even if they chronologically overlapped, refracted by the diversity of biologists’ individual trajectories, we may distinguish analytically a few steps in this move. A first step, the shift from heredity as historical force to heredity as structure has nicely been analysed by Jean Gayon. “Heredity was not the sum total of ancestral influences; it was a question of structure in a given generation. What happened to the progeny did not depend on what happened to the ancestors of its parents, but only on the genetic makeup of its parents” (Gayon, 2000, p. 77). Indeed, most mid 19th century breeders, physicians and biologists saw heredity as a kind of force whose effect would be stronger and more robust when accumulated over many generations (“atavism”). This view could explain the sudden apparition at one generation of ancestral characters which had ‘skipped’ several earlier generations. Although such a view was still held around 1900 by most breeders and was behind Biometricians’ law of ancestral heredity, Jean

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Gayon has shown that this move to think heredity as a particular constellation of particulate elements rather than a force was taken in the 1860’s and 1870’s by Darwin (1868) and by Galton (1876) himself. Darwin’s pangenesis theory (1868) postulated material particules (the gemmules) gathering in the gametes from throughout the body of the parents so as to be passed to the next generation. This idea of independent organic particles of heredity 6 stemmed from the more general view of the “independent life of each element of the body” (Darwin, 1868 [1990], 119) promoted by Claude Bernard and Rudolf Virchow. In 1876, Francis Galton, in “A theory of heredity,” argued that the basic locus of heredity was not so much a line of transmission from parents to descendants, but rather a cytological space, “the newly fertilized ovum” filled with the “germs or gemmules, or whatever they may be called.” He compared this structural space to a “post office” where mail bags full of letters (the “organic units” of heredity) are processed to be distributed to their recipients: Ova and their content are, to biologist looking at them through microscopes, much what mail bags and the heaps of letters poured into them are to those who gaze through the glass window of a post office. Such persons may draw various valuable conclusions as to the postal communications generally, but they cannot read a single word of what the letters contain. (Galton 1876, 331)

This metaphor is one of the earliest occurrences of the idea of heredity as text and of the idea (if not the word) of heredity as information that has to be stored, processed and redistributed. It is striking that such new conceptions of heredity had been drawn from a comparison with the processing of information in postal services, one of the key technological and bureaucratic activities that boomed during the control revolution of late 19th century. Another metaphor used by Galton to describe the hereditary units of the “stirp,” is the political metaphor of the “nation,” within which some individuals compete to be elected and serve as representatives in the fully developed body (Galton 1876). Whereas Darwin had developed analogies with breeder’s practices (which were still cottage industry and craftmen practices connoting a kind of stewardship over animals and crops that was essentially similar to the shepherd in Plato’s Republic, hence situating heredity in a domestic world of worth), Galton preferred industrial/bureaucratic and political metaphors to make sense of heredity. A second step was the ‘sanctuarisation’ of the particles of heredity into a specific place, “deeply buried in the body” as François Jacob put it, separated from the experience of the organism in its environment.7 Heredity shifted from infinite universe to closed world. Heredity became a matter of inwardness rather than of interactivity. The great American fruit breeder Luther Burbank viewed heredity as “stored environment” and he saw hybridization as a powerful tool precisely because it put into contact two plants originating from different places (Thurtle, 2007). Heredity was bonded to a place. But this bond was cut by new visions of the units of heredity being isolated 6

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Each particle or “gemmule” was for Darwin a precursor for a cell, rather than for a character, as assumed by De Vries in his 1889 intracellular pangenesis. But Darwin did himself a move in this direction when he wrote that “a certain number of gemmules is necessary for the development of each character…” (Darwin, 1868 [1990], 157). “A higher order structure has to exist, still more hidden, more deeply buried in the body. It is in a third order structure that the memory of heredity is located” (Jacob, 1993, 207).

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from both the environment and the particular experience of the organism. By the turn of the century, the organic units of heredity were not anymore going out in the whole body and then back to the gametes: their circulation was disciplined and they came to be confined in the “stirp” (Galton, 1876), the “germinal plasma” (Weissmann, 1883, 1892) and/or in the nucleus (De Vries, 1889). One could think about a specific place of storage separated from the contingencies of the organism. A division of labor, a specialisation of function had been sought for the storage of hereditary information.8 Therefore, as Galton (1889) and Johannsen stressed it, no transmission of hereditary traits did occur from parents to children; but from germinal lines to gametes and from gametes to somatic and germinal lines : “Heredity may then be defined as the presence or absence of identical genes in ancestors and their descendants” (Johannsen, 1911, 159). This was a major reordering of how organisms connected with each other in time and place: the space-time of origins and bonds was replaced by the “deeply buried” cytological space-time. 9 Much related to this second move, the third step may be named a “devitalization” or a “stabilisation” of the units of heredity. For Darwin and for Galton until the 1870’s, as well as for De Vries in the 1880’s, the gemmules (or pangenes) were much more than discrete, independent and particulate bearers: they had their own and rich organoid-like life. Their life history was strongly impacted by their past and was full of events of encounters, repulsion, competition for being “representative.” The hereditary units grew, ate and reproduced; they were viewed as assimilating materials, changing from latent to active state and back, from being prolific to being exhausted and sterile and vice versa, etc. Darwin’s, Galton’s, Weisman’s, Spencer’s and De Vries’ hereditary units had their own family tree, their particular and ever changing history. The micro world of gemmules being analogous to the macro world of organisms in evolution, the particles of heredity were viewed as capable of experiencing differentiation and transformations, increasing in complexity and radically changing in number, state, and fertility. Galton, for instance postulated in the 1870’s that “patent elements” [i.e. the organic units of heredity that were expressed in the parent organisms] were less likely to be transmitted precisely because, having been developed into cells in the parent organism, they were somehow exhausted and less active (less “fertile”) or less numerous in the gametes and could not compete with the “latent elements” (Galton, 1876, 339-340). This accounted elegantly for the law of regression, this “steady tendency to deterioration in exceptional peculiarities” that he had observed in the patterns of inheritance of human genius and that was corroborated by “the avowed difficulty, among breeders, of maintaining the high character of any variety that has been produced by accident” (Galton, 1876, 340). For Galton, “existing races are only kept at their present level by the severe action of selection”: performance was not encoded in hard heredity but constantly maintained through a designed environment of selection. This “nothing is fixed for ever” view of heredity widely 8 9

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Althought the term “information” in anachronical here, I use it having in mind Galton’s metaphor of the post office. This move away from interactivity and bondedness in the definition of the biological self was ultimately extended to man by Sigmund Freud. Whereas early 20th century Biologists searched for the unchanging core of living beings sealed off from any bond in time and place, Freud relocated human identity from a priviledged bond with the environment but rather inward into the ego, “what seems us autonomous, unitary, well separated from anything else” (Freud, 1995, 7). He disqualified what Romain Rolland had called “oceanic feeling,” a feeling in which the individual feels bonded with the entire world, as an expansion of the ego typical of infantile narcissism.

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held by late 19th century biologists and breeders, even when they started to think of heredity as cytological space rather than a force, was very close to the fin de siècle notions of entropy and fatigue described in Anson Rabinbach’s cultural history of the sciences of energy and work in the same period. It is interesting to note that Galton himself explained mental fatigue in quite the same way as heredity. In effect, he concluded that while brilliant and active minds are more subjected by the pathologies of “excess of work,” “les personnes à esprit mou protègent leur propre santé cérébrale” (Galton 1889b, 103). This view strongly echoes to his views on heredity and regression, when he hypothesized that elite traits bearing gemmules are less fertile (hence less inherited) precisely because they are extraordinarily so expressed in an individual. The recurrent theme here is the fragility and fatigue of the elite. As Rabinbach showed, the Darwinist’s move from “nature’s design” to an undecided future made by chance and by endless struggles fuelled a multiform fin de siècle sensitivity for energy dissipation, race degeneration and national decline (Rabinbach, 2004, 49). Rabinbach has shown how much “Helmoltz’s cosmos was a cosmos at work” (Rabinbach, 2004, 123). In a similar manner, late 19th century biologists’ genome, to use an anachronistic term, was a genome at work, a dynamic space of organic activity and competition rather than a typological concept (“genotype”) or a program it would later become in the “century of the gene.” Galton himself made an important move from organoid-like swarming units to stable units of heredity. In Natural Inheritance, he stated that “the stability of type, about which we as yet know very little, must be an important factor in the theory of heredity” (Galton, 1889a, 31). Unlike in his writings from the 1870’s, he now viewed the hereditary units as rather immutable entities, and compared gametes formation with a deal in a cards: heredity was more about lottery than about development (Bulmer, 2003, 129). De Vries followed this move and incorporated the statistical approach for the first time in 1894, referring to Quételet’s urn with white and black balls to explain the 1:2:1 ratio he would later rediscover in Mendel’s work. Finally, in 1903 De Vries, had totally changed his views on pangenes: even if he kept the latent/active dichotomy, he abandoned his earlier view that they could change in their state, number and nature, and he acknowledged that their state was almost invariant (Stamhuis et al, 1999, 247-259). Mendel and neo-Mendelians, thanks to their use of the simplest symbolism to depict complex physiological characters, led to a further drastic reduction of the relevant properties of a hereditary unit, from a whole array of states (numerous/few, fertile/sterile, fast growing or not, well nourished or not, latent or active, circulating more of less actively in the body, etc.), to only “presence/absence” and “dominant/recessive”: the “history” of the units was pointless, their combination was everything. Finally, Wilhelm Johannsen, who coined the terms gene, genotype, and phenotype (RollHansen, 1989), finished the job of disciplining and stabilizing the units of heredity: The concept of the gene as organoid, as small body with its own life and whatnot, is not any more to be taken into consideration by research. The conditions for such a concept are totally absent. A horse nesting in the locomotive as cause of the movement is no less a ‘scientific’ hypothesis than the doctrine of organoids to ‘explain’ heredity.” (Johannsen, 1909, 485)10

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In the course of this devitalization of the “hereditary units,” they became immutable bearers of elementary traits. The range of possible states they could go through (latent/active, attracted/ repulsed, actively growing or not, numerous/few or fertile/sterile) were drastically reduced to being present or absent, dominant or recessive. The production of heredity (“like engenders like”) ceased to be a matter of competition, repulsion or growth but became a more deterministic combinatory game in sets of immutable units. Heredity ceased to be the product of organic history (not fully predictable from the nature of elementary units in their initial state) and became the predictable output of a particular constitution. Charles Lenay catches this shift in a vivid way in a thought experiment to imagine how Mendel’s contemporaries might have read his work: Its reflections on constant differential characters, which like abstract and unalterable qualities, could be distributed in the descendants without being modified or even interact between each other caught the biologists on the wrong foot, who where looking for mechanisms of continuous transformation, materially comprehensible, of characters of the species. (Lenay, 2000, 1058)

Indeed, much more than with mid nineteenth century biology, the new understanding of heredity of the turn of the century, postulating a tuned and predictable machinery of hereditary units, was very much in line with the rising tide of the industrial world of worth. The stable gene concept, whether viewed as a kind of chemical reaction (Johannsen) or as a material particle (De Vries, Morgan…), was the keystone of the new combinatory, structural and a-historical view of heredity. Once hereditary units had no relevant life history, (natural) historical explanations became irrelevant in the understanding of heredity. For Johannsen “ancestral inheritance” was a mere fiction (Johannsen, 1911, 138). “Ancestry by itself is irrelevant; dispositions are decisive” he stated in his 1905 Textbook (Johannsen, 1905, p. 216, quoted by Müller-Wille, 2007). For W. Johannsen, the genotypic constitution of a gamete or a zygote may be parallelized with a complicated chemico-physical structure. This reacts exclusively in consequence of its realized state, but not in consequence of the history of its creation.(…) The genotype conception is thus an ‘ahistoric’ view of the reactions of living beings – of course only as far as true heredity is concerned. This view is an analog to the chemical view, as already pointed out; chemical compounds have no compromising ante-act, H2O is always H2O, and reacts always in the same manner, whatsoever may be the ‘history’ of its formation or the earlier state of its elements (…). A special genotypical constitution always react in the same manner under identical conditions—as chemical or physical structures must do. (Johannsen 1911, 139 and 146)

Such a “life out of history” view of heredity dominated of course the 20th century, or “century of the gene” as Evelyn Fox Keller puts it. For instance, Salvador Luria opposed two levels in life and two realms in biological research: “life in action” and “life in history” (Luria, 1973). While the latter refers to the understanding of evolution, the former encompassed all other fields of biology, 10

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“Die Auffassung der Gene als Organoide u.dergl. ist aber nicht mehr von der Forschung zu berücksichtigen. Voraussetzungen, welche eine solche Auffassung nötig machen sollten, fehlen gänzlich. Ein Pferd in der Lokomotive steckend als Ursache der Bewegung (…) ist eine ebenso ‘wissenschaftliche’ Hypothese als die Organoidslehre zur ‘Erklärung’ der Erblichkeit.” (Johannsen 1909, 485).

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placed under the leadership of molecular biology, in which biological functions were framed as a programmed machine sealed off from the contingencies of history and stochasticity. In the early 20th century, most plant, animal, microbe and human geneticists, no matter whether they embraced Mendelism or not (Bonneuil, 2006), developed a new view of heredity that had lost most of its temporal weight, of its historic meaning. This changing relation to time echoed the larger drive to rationalize plant breeding and agriculture as an industrial enterprise, from a cottage industry based on a kind of stewardship on living populations, to quantity production of elite races and varieties. Willet M. Hays, scientist at the USDA, first secretary of the American Breeders Association (ABA) and soon to become undersecretary of agriculture, gospelled this drive: The work of breed and variety improvements and of breed and variety formation is now going forward, but at a pace too slow for these times when the world is advancing with accelerated speed all along the line. As science, inventive genius, constructive skill, business organization, and great market demands at home and abroad have pushed forward things mechanical, so should ways be found of improving these living things which serve as machines for transforming the substance of soil and air and the force of the sun’s rays into valuable commodities.... The energy of the generative cell, and its development into the mature plant or animal, is more abstruse and more profound than the mechanisms of the mightiest locomotive.... As one machine is more efficient than another, so the blood of one generative, or of a small group of generative cells combined into an efficient variety or breed unit, is more valuable than another. (Hays, 1905, my emphasis)

Gaining time was the leitmotiv. This ideal of getting faster elite cultivars and races attracted both the Mendelians and those, like De Vries, Johannsen, Nilsson and Blaringhem, who believed that isolation and mutations would yield stable improved types even faster than hybridisation. De Vries, praised Hays isolation of distinct elementary species of wheat, similar to the one systematized at the Swedish Svalöf station, which he opposed to the slower Darwinian method of population breeding of the German breeder Rimpau: The American breeder by one single choice isolated the very best strains and observed them to be constant and pure. The German breeder, on the other hand, by selecting a number of ears, must have gotten an impure race, and needed a long succession of years and a constantly repeated selection to attain, in the end, the same result. (De Vries, 1907, 102)

In other words, breeding methods inspired by Darwin’s evolutionary theory were both rejected as scientifically unsound and too time consuming for rational practical breeding. The slow time of evolution had no place in the modern world. As Hays recalled, That association [the ABA] (…) has suggested that the scientists in biological lines turn for a time from the interesting problems of historical evolution to the needs of artificial revolution. (…) It has thus recognized that the wonderful potencies in what we are wont to call heredity may in greater part be placed under the control and direction of man, as are the greater physical forces of nature. (Hays, 1903, opening address of the ABA, quoted by Castle, 1951, 62, my emphasis)

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From “historical evolution” to “artificial revolution,” the time had come for living organisms being redesigned along the needs of agro-food world markets and the time-space matrix of the industrial revolution. Emile Schribaux, the first professor at the Paris Institut National Agronomique to teach genetics, gospelled in a similar way for “the improvement of the plant machine” (Schribaux, 1911, 17). William Bateson went even further in the machine-like conception of life when he boasted: “We can pull out the yellowness and plug in greenness, pull out tallness and plug in dwarfness” (quoted by Müller-Wille, this volume). This new industrial spirit affected the space-time in which living beings came to be understood and manipulated. What connected organisms in time and space was re-attributed to combinations of hereditary units that were a-historical and extracted from the influence of locality. In this new framing, A pure race, for a given character, is not, as previously believed, a race that possesses a long lineage of ancestors having this character; it is simply a race in which the character is produced from the union of two gametes of the same kind. (Meunissier, 1910, 13)

Most breeders and geneticists saw the genesis of races, not anymore as the produce of time and vertical transmission but instead of their own proper combinatory engineering. Such an expropriation of the past in the production of relevant life forms opened a wide space for transforming the future with a forward looking, manipulatory, bio-political attitude. Much in line with English and American geneticists and eugenists, Philippe de Vilmorin, head of a major European seed company acknowledged that this new conception of time, space and artifice in the optimisation of living beings, “can have a crucial influence on the future improvement of our species” (Vilmorin, 1910, 12). EXPERIMENTALIZING VARIATION Closely linked to the conjuring away of history, the changing framing of variation is another postDarwinian turn of early 20th century genetics. Variation was for Darwin a fundamental property of life, a continuous and ongoing process. This view implied that breeders had to continue the selection from generation to generation so as to counter this permanent drift if they wanted to maintain their varieties at the best level. Many historians of early Mendelism and of the Biometricians-Mendelian controversy have shown that the concept of the fixity of the pure line (Johannsen, 1903) came to be a weapon against biometricians’ view of heredity as continuous (Roll-Hansen, 1978 and 1989; Provine, 1971). This controversy was only the tip of the iceberg of late 19th-century “new biology”’s systematic attempts to produce—or freeze or orient —variation by all kinds of experimental means. Trained in a more professionalized and experimentalist context of late 19th century biology, many young biologists departed from Darwinian amateur and “panoramic regime” 11 of knowledge making, and from Darwin’s views on biology and on variation. Some, like Hugo De Vries, one of the three “rediscoverers” of Mendel’s law in 1900, developed an experimental program to show that big variations, which he called “mutations,” rather than small continuous 11

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On panoramic knowledge regimes, see Thurtle, 2008.

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ones, could account for speciation and evolution. This idea was held by many scholars of evolution and embryology, including T.H. Morgan. They developed a large array of experimental strategies to produce and track variations in controlled “milieux de culture.” The hundreds of mutant strains of Drosophila in Morgan’s laboratory by 1914 are only one example here (Kohler, 1994). In the course of this experimentalization of biology (Rheinberger and Hagner, 1993), a bunch of new life forms were created and grown up in the laboratories, and variation came to be seen as a process amenable to experimental command and control rather than a “natural” process. As it was possible to “fix” the environment (standard and constant milieu), it was also possible (and sought for) to fix the organism itself through generation by repeated inbreeding or cloning (see discussion below on this new culture of purity and repeatability of life). The reframing of variation from a natural, historic and continuous phenomena to something that could be experimental, discontinuous, artificially and systematically engineered happened in a period when standards and standardisation were a deep concern (Schaffer, 1994; Wise, 1995). The experimentalisation of life went hand in hand with the industrialisation of life, in the same matrix of practice and meaning. Variation and fixity of biological entities came into existence as biological phenomena because they were a central issue in agricultural, medical and industrial practices in the “control revolution” of the turn of the century. Andrew Mendelsohn has nicely shown how a shift from “soft” to “hard” heredity in microbiology occurred in the context of stabilisation of mass produced vaccines, in which “vaccine safety and efficacy constructed, produced, constituted heredity as fixity and in a new, absolute sense” (Mendelsohn, 2005, 95). For “heredity (at least at the realm of microscopic life) came to be located within and redefined by an enterprise of control and testing, production to an exact standard and reliable distribution” (Mendelsohn, 2005, 85) In the first decade of the 20th century, it became a commonplace among breeders and geneticists to contempt Darwinian’s view on variation for having missed the basic fact that elementary species, pure sorts or pure lines are fixed and show no more variation in which selection can work on. A Canadian scientist visiting Svalöf’s station reported that : “the Darwinian idea of the omnipresence of hereditary variation in all life was still held by Nilsson who regarded it as necessary to continue the selection from generation to generation to effect a complete fixation of the characters (…) this idea came to be abandoned” (Newman, 1912, 28). Johannsen as soon as 1898 (Roll-Hansen, 2005) as well as De Vries (1907) condemned Darwin-inspired breeding methods and called attention for pre-Darwinian techniques and concepts such as Vilmorin’s “pedigree selection” and the “elementary species,” a concept that had been promoted in the 1850’s-1870’s by the French creationist naturalist Alexis Jordan. For Jordan the species boundaries could be determined with certainty only through experiment, i. e. through cultivation side by side of various forms over several generation, so as to see if their differences bred true. But Jordan’s elementary species concept, was discarded as splitting hair by the leading naturalists, such as Darwin’s friend Joseph D. Hooker, Georges Bentham and Asa Gray. These leading figures, who headed the great herbaria in Europe and the United States, maintained the armchair study of dried plants fragments, rather than experiment, as the cornerstone of proof-making in taxonomy. They also imposed successfully the broad species concept through the imperial power of standardizing enterprises such as the colonial Floras, Bentham’s Genera Plantarum and the Index Kewensis

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(Bonneuil, 2002). Against “species mongers” Hooker and his allies argued that the broad species was more in line both with the study of biogeography and evolution (framing life as a constant flow of variations interconnecting all taxonomic categories) as well as with the imperial project of a commercial unification of the world. It was for instance necessary to attribute a unique technological and commercial value to a species of rubber plant, whether it was collected in different places and named under different (“bad”) species names by various travellers. In sharp contrast with this natural historical and imperial metrology, experimental biologists and geneticists of early 20th century rehabilitated “Jordan’s classical work” (Johannsen, 1913, 389) and small species concept. While the first metrology was part of a larger drive to rationalize extractive mercantile enterprises, the second kind of metrology, allowing the creation and circulation of new stable and pure forms of life at the subspecific level, which were turned into mass commodities, was constitutive of the space of flow of agro-industrial goods. More than just a finer grained taxonomy, what was rehabilitated in this move was a typological view and a search for a stable biological type as “the most biological concept in the science of heredity” (Johannsen, 1905 quoted by Roll-Hansen, 1978; but see also Theunissen, 1994 on De Vries typological views, and Mayr, 1973 for a general analysis). Shull, one father of the hybrid corn, posited very vividly this “modern view of heredity” (Johannsen, 1911, 130), in the history of biological ideas: The doctrine of evolution had to overthrow the [creationist] conception of permanency of specific types (…). It was Darwin’s great triumph (…) to convince the scientific world – and through the scientific world, ultimately the whole world—that everything is in a state of flux, and that there is no such thing as permanency among living things. Owing to the work of De Vries and the other early students of modern genetics, permanency of type again demands serious scientific consideration (…). The old idea of the immutability of specific types was based upon almost total ignorance of genetics, as was likewise the Darwinian conception of fluidity and gradual change (…). The critical work of the past few years has brought a great change and the new idea of permanency is gaining ground with the growth of experimental knowledge. (…) we can definitely say that types are absolutely permanent and do not, at least in some cases, gradually change into something new. (Shull, 1911, 234-35)

If the analysis and production of stable forms of life constituted the heart of the young genetics, at least two strands of approaches of stability were in competition. Some found the basic immutable entities in the “unit factor” or gene (or in the linkage group localised in the chromosome), whereas others, including Johannsen, opposed factorial genetics and saw the genotype 12 as a whole, the elementary species, or its most radical form, the pure line, as the key permanent biological type (Churchill, 1974). This contrast hints to discontinuity between Mendelism and “pure line research” that has been neglected by historians. Indeed, although the two lines of research were convergent, Johannsen (1909, 1911) as well as Shull (1911) and East (1911) presented the “pure line principle” or “pure line theory” as a research programme that was clearly distinct from “hybridisation studies.” Although they diverged about the level that displayed at best stability and immutability and where hard heredity could be located, theses two lines of research strongly 12

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converged in rejecting 19th-century biology’s dissolution of the type in an infinite variability of ever circulating beings (species at the macro level and hereditary units at the micro level), modifying themselves mutually when encountering one another. They agreed on the promotion of authenticity, fixity of the biological self, of a “genetic identity” deeply buried and sealed off from the effects of time and place… but manipulatble by the experimenter. These timeless, experimental/manipulatory and typological views of life reinforced or created boundaries in the biological world, in the social world, as well as between the natural, the cultural and the social realms: — It established a great divide between the past and the future, between time consuming empiricism to planned improvement, between cottage industry and mass production of elite organisms. — Its focus on stability, predictability and standardisation reinforced a great modernist divide between the natural world of landraces and the optimized world of pure strains and high yielding cultivars devised by human scientific genius. Till late in the 20th century, the discourse on crop biodiversity was a discourse about the past and the “origins.” Vavilov, for instance, promoting a systematic exploration of “the whole initial varietal potentialities of the world” (my emphasis), stated: The vast resources of wild species, especially in the tropics, have been practically untouched by investigation (…) An actual mastery of the processes of evolution (…) can be accomplished only through the combined efforts of a strong international association and through the removal of barriers impeding research in those most remarkable regions of the world (1932, 331 and 342).

This is a typical colonial and modernist discourse of discovery and scientific use by civilized/ scientific Man, of untapped resources: gene flow was framed as a resource from the past (for the breeding science and industry), rather than an ongoing process in which farmers’ knowledge and agency make a difference in future conservation and innovation, as in the recent ‘participationist’ and ‘connexionist’ discourse of in situ crop diversity conservation (Bonneuil and Demeulenaere, 2007). In this modernist framing, interspecific and intervarietal crosses were seen as a specialized undertaking of the professional geneticists (breeder, cytogeneticist…), gate keepers of the boundaries between (elementary) species. Landraces were redefined as “ecotypes, derived from populations upon which natural selection operated during very numerous generations in the same environment” and hence conjuring away any role of the farmers (Bustarret, 1944, 346). — Hence a stronger boundary and division of work between farmers and breeders (whose social status was based on their mastering of artificial crosses, large scale screening and pure line breeding). In the following decades, the seed was constituted into an object of public policy that promoted quality standards that reinforced the professionalisation of plant breeding and the making of seed into a commercial “input” (Bonneuil and Thomas, in press).

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— Finally a stronger boundary was drawn between academics and practical breeders: in sharp contrast with Darwin’s reliance on breeders’ testimony, leading figures of the new “genetics,” like Johannsen and Shull, considered breeders as not trustworthy in matters of heredity (Johannsen, 1911; Glass, 1980). Müller-Wille and Rheinberger (2004) have argued that , in the new epistemic space of heredity, biological transmission and cultural transmission, that were in the mid 19th century thought with the same concepts, became strongly separated. I believe that the four modernist divides mentioned above were key elements of this general drive.

3. Industrializing purity As a prelude to a cultural history of purity in the rise of genetics, this section tells two stories of discovery and two stories of justification of purity. Together, these stories document how purity and stability became both a norm of industrial production, a norm of scientificity in experimental biology and a norm of fairness in social and economic relations. VILMORIN’S PURE LINES AND THE PROBLEM OF CONTROL IN THE SEED CHAIN Both Nilsson, De Vries and Johannsen acknowledged the French breeder Louis de Vilmorin’s work as a crucial step toward the notion of pure line. He pioneered the “pedigree breeding” technique of selecting individuals (rather than groups of plants) and documenting “a perfectly correct genealogy of all plants from the beginning of the experiment” (Vilmorin, 1859, 44). In a 1856 work on sugar beet breeding for sugar content, he also noted that the progeny of some individuals was sometimes homogenous and sometimes highly variable, and he proposed that breeding should not only search for high performance types but also for lines with minimum variability (Gayon and Zallen, 1998). Reducing variability, and increasing the predictability of a standard agronomic or technological performance of the seeds was a major goal as the seed trade from improved varieties developed, not only for vegetables and flowers, but also for major crops such as beet, wheat and barley. As grain and seed markets extended, trust networks and standards had to be extended. Inspired by similar regulations on fertilizers trade, seed regulations emerged in different industrializing countries setting minimal purity standards and minimum germination rate. A standard, rather than variable output was expected from improved seeds by the advanced farmers that pioneered their use. Homogeneity was also valued to facilitate the use of harvest machines. The Vilmorin Company employed about 400 employees in 1889 and, as one of the leading seed companies in the World, had to meet the challenge of quality standardization and control to maintain its position. Even before seed trade regulation imposed national standards, Vilmorin conducted both cultivar performance assessment at a multi local scale before release and, in the 1880’s, routine seed quality testing (Flavien 1889, 17). To ensure quality downwards on the seed chain, it was necessary to control quality upward at the seed multiplication level. Ideally, this could be done at Vilmorin’s estate near Paris by skilled, disciplined and carefully managed waged manpower. But this method of production of commercial seeds was costly. In order to decrease production costs, seed multiplication was

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subcontracted to farmers near the Loire, under the supervision “of a special team of inspectors who checked all farming operations” (Flavien 1889, 14). The assessment of seed purity delivered by these farmers was made easier if the initial purity of the seeds delivered to them for multiplication was perfect, so as to detect adventitious mixtures more easily. A visiting engineer specialised in scientific management of work concluded with admiration: In seed production, the division of work adopted at the Company Vilmorin aims at producing, with all the necessary care, seeds of extremely pure races [“de races extrêmement pures”], and multiply them widely in such conditions that, without loosing their purity [“franchise de race”], they can be delivered at the lowest possible price. (Flavien, 1889)

The Vilmorin Company hence combined pedigree selection technique to produce pure lines with a particular quality control for seed production where purity was a key element to ensure standard quality at the lowest production cost. As a perfect example of the “control revolution” documented by James Beniger (1986), purity was here a powerful tool for the Company to keep control along a seed production chain where the work was subcontracted. STABILIZING BEER, BARLEY AND BEANS : FROM PASTEUR TO JOHANNSEN Andrew Mendelsohn has argued that the mass production of attenuated strains of Anthrax as vaccine by Pasteur’s laboratory in the 1880’s was one moment of shift from heredity as force to heredity as presence or absence of a permanent and transmissible trait (Mendelsohn, 2005, 91). Microbiology and its new zoo of small “corpuscules,” each having its agency and its particular chemical or medical action is indeed certainly one of the roots of the particulate and structural view of heredity. Microbiology was also a field of practices and discourses of purity. Pasteur, for instance, in his works on the spontaneous generation, the vinegar, wine and beer fermentations on the 1860’s and 1870’s, sought for purity not as a quantitative magnitude (“more or less pure”) but as “an absolute absence, mathematical if I may say” of any germ other than the studied one (Pasteur 1876, 218). It was a question of yes or no. Pasteur even wondered whether the beer industry could have attained its actual state without having followed his principles of pure culture (Pasteur 1876, 216-17). He noted that the commercial beer contained not only the beer yeast but also undesirable other yeast species (such as Saccharomyces pastorianus) and “disease germs” (vinegar and milk bacteria, and various fungi) which were responsible for turning the beer spoiled after some time, and discarded as undrinkable. This problem of conservation of the beer was only partly, and at high cost, solved by the large scale use of refrigeration techniques (ice machines) from 1880 on. The beer demand was highest in warm months but it was easier to produce and keep unspoiled in cold months. While steam boiling, cooling compressors, new energy sources and bottling machinery made mass production possible, “keeping quality” remained the bottle neck in the mass scale production. J. C. Jacobsen, the founder of the large Carlsberg Breweries in Copenhagen and a great admirer of France came across Pasteur’s work just when he was creating a laboratory, then to become a world centre for biochemical research. Emil Christian Hansen head of the physiology department of the Carlsberg Laboratory deepened Pasteur’s taxonomy of the micro-organisms present in the beer leaven, and greatly improved its method of “pure culture.” To make sure that a strain derived

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from a single cell, he used in 1882 the gelatine-substrate technique he had learned during a visit to Robert Koch in Berlin. A first beer production trial was held in 1883 at Carlsberg Brewery, with different pure lines from the good yeast species he had named Saccharomyces carlsbergensis, including the one he called “Carlsberg bottom yeast n°1.” Large scale production of pure yeast bottom-fermented beer was reached in 1885, and an apparatus for the continuous production of pure culture yeasts was devised (Glamann, 1988). Jacobsen wrote in 1884 that “from now on fermentation in my brewery will wholly be carried out by means of this pure yeast, produced from a single cell! Truly a triumph of scientific research!” (quoted from Teich, 1983, 121). By then Carlsberg controlled almost half of the Danish lager beer market (Boje and Johansen, 1998, 60). Abroad, the “pure yeast” method was heralded among German brewers by beer scientists like M. Delbrück who stated that “the yeast is a working machine” transforming sugar into alcohol, hence amenable to industrial rationalisation (Delbrück 1884 quoted in Sibum, 1998, 48; see also Teich, 2000). Finally in the 1890’s, except in Britain, most of the large Breweries in the industrialized world turned to pure yeast technology, an innovation that together with ice machines, train transportation systems, urbanization and changes in alcohol consumption patterns, transformed the brewing industry into one of the most advanced, global, concentrated, capital-intensive and mass-scaled food industry of the time. Stabilizing beer so that it could flow along the global networks of an expanding beer market, while remaining immutable, had implied a thorough disentanglement of the quality problems that resulted from the production and transportation conditions from those which could be fixed by controlling the “intrinsic” nature of yeasts and standardizing them into pure lines. Barley posed similar issues: its germination kinetics, sugar and protein content were key properties whose optimization and standardisation were required to rationalize the production process and massproduce standard quality beers. From 1881 to 1887, just by the time when pure yeast was introduced, Wilhelm Johannsen was research assistant at Carlsberg Laboratory and explored Barley’s ripening and germination’s chemistry and physiology (Teich, 1983). Later, when Johannsen took a position at the Royal Veterinary and Agricultural College in Copenhagen, he engaged in barley breeding for brewing quality in collaboration with Carlsberg. In this research he combined Galton’s biometrical and statistical methods and Vilmorin’s and Hansen’s pure line principle to work with descendents from single individuals through self-fertilization (RollHansen, 2005, 43-47, Johannsen 1899). It is in this context that Johannsen picked up the creed of purity as a norm of proof, efficiency and fairness. The vagaries of uncontrolled and changing environmental conditions and uncertain ancestry had to be erased so as to harness at large scale new forms of life engineered to react in the same way to given conditions. “The study of the behaviour of pure lines is the basis of the science of heredity, even if populations—especially human populations—are not made of pure lines” wrote Johannsen in 1903 in his study on the ineffectiveness of selection in genetically homogenous populations of beans (Johannsen, 1903, 9). Pure lines, and the particular kind of typological thinking that was associated with them, were the cornerstone of the production of both sound knowledge and of large agro-food markets and industries. Louis Blaringhem, lecturer at the Sorbonne Faculty of Science engaged in a Svalöf-like barley breeding program for the brewing industry, and an ardent promoter of De Vries’ and Johannsen’s views in France, stressed:

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The ideal thing for industry is to operate on products whose nature is well defined and always identical. There exist excellent methods for the purification of inert matter, such as fractioned distillation (…) [but] living matter is complex and the grower, unaware of the value of these methods, cannot provide the guarantee expected by the industrialist, hence a difficulty in economic exchanges. (Blaringhem, 1905, 362)

As pure yeast culture had transformed the beer industry, he heralded that stable and pure lines of crops would revolutionize agriculture and boasted that “the future belongs to pedigree pure sorts” (Blaringhem, 1913, 87). PURITY AS FAIRNESS IN THE INDUSTRIAL WORLD OF WORTH There was actually nothing new in using “types” or traits transcending space and time, stability and purity, to maintain social boundaries (Douglas, 1966). As pointed out by Nicholas Russel in his exploration of early modern animal breeding, “the parallels between the human obsession with title, hereditary position and social caste and animal pedigrees, are too obvious to need emphasis.” (Russel, 1986, 19). Russell argues further that the careful keeping of pedigrees, for lack of disentanglement of the effects of environment and heredity, acted mainly as a political tool to legitimate the political power of the aristocracy. What was new then in the understanding and valuing of purity at the turn of the 19th and 20th century? My argument is that, in the post-Darwinian space-time that we have documented in the preceding section, the search for purity was basically reframed within the “industrial world of worth” (Boltanski and Thevenot, 2006). From 17th to mid 19th century, purity was framed in a mix of “fame,” “inspiration” and “domestic” world of worth. Breeding was a search for fashionable conformations (cf Bakewell, “fame” world of worth) and aesthetic criteria (“inspiration” world of worth) rather than purely economic performance; personal knowledge and “breeder’s eye” (“inspiration” world of worth) deeply mattered as well as ancestry, stewardship and interpersonal relations (“domestic” world of worth). Purity was viewed as something highly valuable because it was particularly rare and unstable, and needed constant care to be maintained. The key trait was rarity. Genealogies (General Stud Book) acted as tools to organise trust around the genealogical value of English Thoroughbred from Arabian origins in a context of scarcity since new importations of Eastern bloodstock was banned (Russel, 1986, 99). In the same way, the “degeneration” of elite cultivars (including Major Hallet’s “pedigree seeds” of late 19th century) acted as incentives for farmers to buy new seeds to the plant breeder. In a context when breeding was a cottage industry lacking of powerful information and control technologies (including intellectual property rights) necessary to control large trade networks, scarcity and instability of elite breeds was consubstantial with the economics of breeding (Berlan, 2001). There was no need to extend trust in elite breeds further than intellectual property could extend. Purity was hence “sticky.” It was altered when races and varieties moved in time and space, and remained dependant on continuous and distinctive practices of the breeder, rather than warranted into an inward timeless constitution of the organisms. As we have seen, this was also much in line with fin de siècle’s visions of entropy and fatigue, with Darwin’s idea of continuous variation as well as with Galton’s view of the “stirp” as a space of (inter)activity and competition.

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In a few decades at the turn of the 19th and 20th century, purity was drastically reconceptualized. Of course it remained dependant on adequate skills and practices (from avoiding contaminations in microbiology to careful avoidance of cross breeding to maintain pure lines), but it was made more robust because these practices could be codified and routinized and because living populations were now seen as basically made of intrinsically stable and pure types, which just needed to be sorted out. Purity was not anymore the produce of history, but rather a structural property (homozygocy) that scientists could master across time and space. 13 Purity lost its domestic dimension. Purity was assessed not only “vertically” (through checked genealogies), but also “horizontally” as predictable functional performance (whether a Gauss curve, a replicable biological effect, a safe vaccine or a high strength wheat). Purity lost in the same movement its association with rarity. It became valuable not because it was an unstable and rare state of living being, but on the contrary because it was amenable to quantity production and to replication across time and space. In the industrial world of worth, fairness in the commercial exchange was associated with the purity and stability of the product. Darwin-inspired population breeding methods and their impure/unstable products, which were previously valued because they created a special role for elite social groups to intervene actively so that the world would not fall apart in chaos, entropy and degeneration, became the stigma of unfair exchange: [If continuous selection would be the right way] it means that all the seed destined for sowing should be produced directly [by the breeder] (…) it is easy to see that the gain made by the breeder of a new variety depends, for a large part, on the acceptance of this proposition. (De Vries, 1907, 43)14 Almost all cultivated varieties have been obtained by selection. Only [elementary] species born by mutation deserve agronomists’ consideration because they alone are genuinely stable (…)But [with Darwinian selection methods] once the variety is put to the market, (…) it quickly loses its value (…) the fast degeneration of the improved seeds he sells ensures the renewing of his order books (…) [hence] the focus on breeding by selection and the present neglect of cultivating forms born by mutation. Only these are stable from their very birth. (Blaringhem, 1905, 377)15

Purer was not only more efficient, it was also fairer. The reframing of purity practices and meaning in the “industrial world of worth” of the control revolution that accompanied the industrialization of the agro-food sector in the last decades of the 19th century, informed not only laboratory and

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Paradoxically, genealogical techniques (book keeping and other inscription devices) became even more important as history was expelled for the framing of heredity. Parentship was important only in so far as it was taken into manipulation, isolation and traceability techniques, but it was not anymore central. Constitution rather than connection was the hallmark of heredity. As Rheinberger would put it, in plant genetics, genealogy became a “technical object” rather than an “epistemic thing.” I thank Jean-Pierre Berlan for attracting my attention towards this book from Hugo de Vries. See Berlan, 2001. Blaringhem was then a young left wing intellectual, and as a son of a medium farmer, he shared the third Republic’s ideological project to free the farmers from the domination of merchants and notables, a project that combined the industrial world of worth and the civic world of worth.

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economic practices but also judgement about fairness and about the kind of social order that was desirable. PURITY AS NORM OF PROOF AND THE MAKING OF BIOLOGY AS AN EXACT SCIENCE Der wichtigste Probierstein hier ist aber das kritische Experiment mit genotypisch ‘reinem’ Material. (Johannsen 1913, iii) Es ist in der Geschichte der neueren Biologie auffallend, daß zu einer Zeit, wo man, in Bezug auf Mikroorganismen, durch ‘Reinkultur’ (durch Kultur mit einer einzigen Zelle als Ausgangspunkt) äußerst wichtige Resultate erhielt, in der Erblichkeitsforschung die Verhältnisse der höheren Organismen fortwährend in weit gröberer, summarischer oder statistischer Weise studiert wurden. Was die Arbeitsmethoden eines Koch oder eines Hansen für das exacte Studium der Mikroorganismen bedeutet haben, dasselbe bedeutet auch für die Erblichkeitsforschung die Reinkultur, d. h. die individuelle Nachkommenbeurteilung, wie sie Vilmorin und Mendel präzisiert haben: Ohne Reinkultur keine klare Einsicht, sonder Konfusion und Irrtum! (Johannsen, 1913, 196)

This new industrial framing of purity informed not only laboratory and economic practices and normative views about the kind of social order to be reached in the 20th century but also the very meaning of how biology should be an exact science. Exactness differed from precision as Bateson stated in 1902: “We have been told of late, more than once, that Biology must become an exact science (…). But exactness is not always attainable by numerical precision” (quoted by par Provine, 1971, 71)16. The focus on exactness was not in the title of Johannsen 1905 book in Danish, but was introduced in his 1909 Elemente der exakten Erblichkeitslehre, and already discussed in his 1906 talk at the International Conference of Genetics. His creed for exactness was a weapon against biometricians who, Johannsen thought, measured precisely, but measured the wrong thing. “Exact biological analysis” meant indeed for him “the fundamental distinction of true type differences and fluctuations” (Johannsen, 1907, 110). Ignoring the existence of biotypes, biometricians worked with ill-defined categories. Interestingly, Johannsen illustrated this point with an analogy taken from the industrial world of the second industrial revolution: If anybody makes a study as of the speed of the railway-cars, the botanist noted, he will of course regard every train or type of train separately: express train, local trains, goods trains, and so on. (…) But what would be said of an enquirer who, for solving the problem, collected statistics as to the speed of the different carriage-classes (…) and by this method found out that the average speed of the first-class car was much greater than the average speed of the third-class car—for in the express trains (in the continent at least) there are only (…) first and second-class cars, while in the local trains the third-class cars is the majority (…) I must confess that the main part of biometrical work in questions of heredity somewhat resembles such preposterous statistics. (Johannsen, 1907, 99) 16

This argument soon became a topos. For instance, after Bateson and Johannsen, Jennings stated that “the men who (…) have lectured on the necessity of becoming exact are the strongest opponent of exact experimental and biological analysis—seeming to feel that mathematical treatment renders other kinds of exactness undesirable” (Jennings 1910, 143). See also Johannsen 1913, 154.

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In other words, the metaphor stated, biometricians developed a pre-control revolution kind of knowledge, which could not help anybody to understand and manage the modern world. Pure lines and biotypes offered as well the promise to get out of the vagaries of species delimitation in taxonomy. As H. C. Watson had written to Darwin, “The short truth is, that we have no real proof or test of a species in botany. We may occasionally disprove an alleged species by seeing its descendants become another such species, —or we may unite two by finding a full series of intermediate links.” As I showed, the chaos in taxonomy had been avoided by imposing the rather conventional “large species” norm to the naturalists community through the botanical tools of imperial power (Bonneuil, 2002). But the epistemological weakness of such a closure remained. Systematics seemed unable to cope with observer-dependent knowledge and to achieve communicability and accumulation. So the rehabilitation of the small species concept by early 20th century geneticists also offered the promise to establish a “new botany” as an exact experimental science working with new taxonomical units. Hansen and Pasteur were seen as the founding father of this exact biology because they achieved an “exact analysis of yeast population” (Johannsen, 1907, 104). “Heredity can only be studied in an exact manner by breeding experiments” and, Johannsen added, there are two ways to do it: the “analytical experiment” with pure lines, and the synthetic experiments of “hybridology” (Johannsen, 1907, 103). As proponents of the spontaneous generation in Pasteurs’ mouth half a century earlier, Neo-Lamarckians were, for Johannsen, committing the sin of impurity “Most of the “neo-Lamarckian” literature demonstrates the necessity of exact experiments” (Johannsen, 1907, 104). “Contamination,” once the stigma of the unskilled microbiologist, became the sin of the bad student of heredity. As Jenning phrased it, Castle’s experiment showing effects of selection in rats and guinea pig were of poor value because they “dealt with races of complicated descent; they plunge us at once into all the difficulties due to interweaving, blending and transfer of characters from one genotype to another” (Jennings, 1910, 140). On the contrary, purity was required if one was to turn living forms into elements of experimental systems and measure replicable and constant effects under similar conditions (Rader, 1999; Löwy and Gaudillière, 1998): The ideal material for any genetic, biological or agronomical research is of course the pure line; because of its intrinsic stability, in space and time, it makes possible to control the factor ‘heterogeneity of the plant material’ in the experiments. (Bustarret, 1944, 353)

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Conclusion While biometry inferred a norm from heterogeneous life forms, exact genetics materially produced normal life forms, in creating populations made of strictly standard, identical and highly performant organisms: “pure sorts,” “pure lines,” “clones,” F1 hybrids corns, “inbred lines,” all being “isogene individuals” (Jennings, 1910, 152). Singling out “the ‘Shakespeares’ of the species” was the motto of both sound science of heredity and rationalized industrial mass-production: As electrical energy must be harnessed, so these investigations are showing that the peculiar breeding potencies of the rare plant or animal must be singled out and given opportunity to work. Both in practical breeding and in evolutionary studies the individual with exceptional breeding power is gaining respect. . . . The world is learning to seek the ‘Shakespeares’ of the species with the same avidity that it seeks gold mines. (Hays, 1906 quoted in Boyd, 2001, 654)

Indeed, in the early 1900’s, “the climate of biological opinion was favorable to the pure line theory” (Provine, 1971, 108). In his pioneering reading of the communications of the 1910 “Genotype Hypothesis” Cornell symposium mentioned in the introduction of this paper, William Provine depicted how Shull, East, Jennings and Pearl and many geneticists were then riding the tide of “the modern view of heredity.” Jennings and Pearl argued, on the basis of the pure line theory, that selection in cross-breeding population was incapable of changing a character beyond the existing limit of variation. East and Shull, deeply sharing the typological belief in “the discreteness, uniformity and permanence of the types” (Shull, 1911, 237), sought to extend the genotype concept to open-pollinated crops and rushed to sort out and amplify the very best genotype of open-pollinated crops such as maize in the same way as this had been done for selffertilizing crops: A (…) demonstration that populations of cross-breeding plants and animals are composed of fundamentally distinct types, intermingled but not changed by panmixia, and capable of being separated by appropriate means and of being shown to possess the discreteness, uniformity and permanence already demonstrated for the genotypes of self-fertilized and clonal races, will add greatly to the importance of the fundamental conception of permanency of types involved in the work of De Vries and Johannsen. (Shull, 1911, 238)

The innovative step was to infer that, if all plants of a corn field resulted from the cross between two parents, or, in other words, from the combinations of two among “numerous elementary species,” then “the fundamental problem in breeding this plant is the development and maintenance of that hybrid combination which possesses the greatest vigor,” i.e. to mass produce the one best cross (Shull, 1908, 300, my emphasis). A difficulty was the depression caused by inbreeding to get fixed pure lines as parents to be crossed. But Shull did not loose hope because, together with this depression, he obtained in his pure lines “the gradual lessening of variability,” which Vilmorin had searched for half a century earlier and which Johannsen had predicted on the basis of his “pure line theory” (Shull, 1911, 244; Johannsen 1907). William Provine argued that, at the 1910 symposium, Johannsen’s “pure line theory seemed so obvious that most outstanding geneticists accepted it without adequate proofs” (Provine, 1971, 108). Similarly, Ernst Mayr has criticised the limitations of the typological view of the species

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promoted by De Vries’ and Johannsen’s work. For instance, De Vries’ typological views led him to postulate the genetic homogeneity of natural populations due to strong natural selection pressure. For him, the normal situation in nature was that interbreeding individuals were of identical genetic composition (hence the need for mutations to make a difference in evolution) (Theunissen 1994, 243-44). He also, as we have seen, dismissed mass selection as leading less efficiently (longer time, more impurities) “in the end, [to] the same result” than pedigree pure line selection (De Vries, 1907, 102). While reasoning in this way however, De Vries overlooked the possible productivity of gene recombination in interbreeding individuals under lower selection pressure, a productivity that was acknowledged later in the evolutionary synthesis, and harnessed in “recurrent selection” breeding schemes from the 1940’s on.17 Similarly, when Jennings dismissed selection in cross-breeding populations for being incapable of changing a character beyond the existing limits of variation or when East and Shull discarded the value of breeding strategies based on pure lines rather than on populations, they conjured away the importance of recombination that was key to the practical success of breeding and that would later become central in quantitative genetics and population genetics (Provine, 1971, 122). 18 The point in mentioning the discontents of genetic purity is not here to blame geneticists, from a “histoire jugée” perspective, for having missed or delayed new and fruitful scientific avenues in the 1900’s. My point is rather twofold. A first observation is that Hardy-Weinberg’s 1908 law and the study of allele frequencies in mixed populations can be seen as more “Mendelian” than the hybrid corn innovation. In many ways, East and Shull’s work towards hybrid corn, rather than a Mendelian innovation in plant breeding, as often depicted by geneticists and historians of genetics, resulted from the kind of typological view of the species that E. Mayr and W. Provine criticized. This can help historians to avoid the plot of the hybrid corn as a Mendelian success story (Bonneuil, 2006) and sharpens the analytical distinction made earlier in this paper between two ways of stressing stability and permanency in biology and heredity by 1900: one taking the hereditary unit or gene as the immutable unit, and one taking the biotype as the immutable unit. These were in fact two “stabilisation” strategies that both emerged from the wider drive to reshape life in a new industrial time-space of flows. We have developed this argument in detail in this paper as far as the “biotype” or “clone” strategy is concerned. But the second strategy of singling out and stabilising immutable genes for valuable traits (disease resistance, productivity, chemical composition adapted to industrial transformation, etc.) has done a similar and complementary job: it has allowed to put “hereditary units” in circulation within a global scientific-economic network of plant breeding, where they were reassembled into

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Population and quantitative geneticists have later argued that in fact the two strategies presented by De Vries in 1907 do not only differ in their rapidity, but also that they don’t lead to the same result on the long run. The isolation or pedigree method, corresponding to a maximum of selection pressure, impedes any further recombinations and leads to a plateau in genetic improvement. This plateau can be overcome only by an alternance of slow selection pressure cycles allowing recombinations to happen (close to mass breeding) and pure line breeding cycles to extract parent lines foir the hybrids. This principle is the basis of “recurrent selection” breeding techniques (Jenkins, 1940). Two years before the Cornell meeting, G. H. Hardy and W. Weinberg came in 1908 to new ways of understanding genes flows in natural populations, that concluded for the existence of regularities in the proportions of alleles in mixed populations without supposing that theses populations would quickly tend to homogeneity.

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new placeless products, such as the Green Revolution cultivars that were made insensitive to photoperiods and grown worldwide (Bonneuil and Demeulenaere, 2007). Hence my second point: to understand why “the climate of biological opinion was favourable to the pure line theory,” and why natural selection seemed so unimportant as compared to stability of the type,19 historians of genetics need to acknowledge how deeply the quest for purity and stability in early genetics and plant breeding has been shaped within a larger drive to rationalize the agro-food sector and mass-produce the one best. As Mary Douglas demonstrates in her cultural understanding of attitudes toward “impurities” and “pollutions,” we cannot understand early genetics’ obsession for purity and stability by just looking at genetics: Defilement is never an isolated event. It cannot occur except in view of a systematic ordering of ideas. . . . The only way in which pollution ideas make sense is in reference to a total structure of thought whose key-stone, boundaries, margins and internal lines are held in relation by rituals of separation. (Douglas, 1966, 42)

From the few rituals of separation documented in this paper, it clearly appears that early 20th century genetics emerged in a larger scientific/economic/cultural matrix of practice and meaning that reframed how organisms were connected together in time and space and with their environment (Thurtle 2007). In this wide cultural shift, a deep and intrinsic genetic identity was constructed for living organisms, separated from the influence of the place and the environment. New “pure” and stable life forms were mass-produced in laboratories and industries, which could circulate without alterations through extending “space of flows,” be they inter-laboratory networks (the most famous being the circulation of strains within the Drosophila community) or larger scientific/economic/medical/cultural hybrid networks of the control revolution. Finally, I must confess a major hole in this article: the material practices have only been superficially discussed here, even though they should be documented in any comprehensive cultural history of the birth of genetics. Although the geneticists of the turn of the century promoted stability and purity as a constitutional and intrinsic property of life (typological conception of biotypes and structural view of purity as homozygoty), they knew, as well as historians know, that the production and maintenance of these pure forms of life necessitated hard work, industrial scale observation and treatment of minute differences, and standardisation activities. As shown by Kohler (1994) with the production of the standard drosophila, with a stable rate of crossing over in every part of the chromosomes, the coming into being of pure life forms rested upon labor-intensive and capital intensive “networks of purity,” being elements of the control revolution. So an entire aspect that should have been addressed in a more comprehensive essay on the cultural history of early 20th century genetics is the question of the transformations of the material practices of observation, recording, book-keeping, processing and manipulating that were associated with the shifts we have described. Although we have a few good pioneering works (Kohler, 1994; Rader, 1999; Löwy and Gaudillière, 1998; Müller-Wille, 2005), at hand, 19

De Vries, for instance, argued in his Mutationstheorie that natural populations were genetically homogenous, because selection would quickly wipe out any new form that was less vigorous or replace the whole population by a beneficial new mutant. So the normal situation in nature was that interbreeding individuals were of identical genetic composition and only mutations made a difference in evolution (Theunissen 1994, 243-44).

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much remains to be done to explore the “industrial” scale and organisation of the work and use of sorting machines in a breeding station like Svalöf (that established the standards for many others), the role of bureaucratic microtechniques (standardized forms for observations; inscription devices such as maps, registers, fieldbooks; management of information flows, etc.), the disciplining of bodies associated with the production of pure life forms and controlled experimental environments. Only with this additional work on the mundane microtechniques of genetics, will it be possible to grasp fully the intimate relations between norms of objectivity, exactness and precision, visions of life, bureaucratisation, mass-production and large-markets at the turn of the century. Christophe BONNEUIL Centre Koyré d’Histoire des Sciences et des Techniques (CNRS) and INR-TSV [email protected]

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References Blaringhem L. 1905. “La notion d’espèce, application aux progrès de l’agriculture et de l’industrie des notions nouvelles sur l’espèce.” Reprint from Revue des Idées 17, 15 mai 1905. ————. 1913. Le perfectionnement des plantes. Paris: Flammarion. Beniger, James. 1986. The Control Revolution: Technological and Economic Origins of the Information Society. Camebridge, MA: Havard University Press. Berlan, Jean-Pierre and R. C. Lewontin. 1986. “The political economy of hybrid corn.” Monthly Review 38(3). Science, technology and capitalism: 35-47. Berlan, Jean-Pierre. 2001. “Political economy of agricultural genetics.” In R. Singh, K. Krimbas, D. Paul and P. Beatty (eds.). Thinking about evolution: Historical, philosophical and political perspectives. Vol. 2. Cambridge: Cambridge University Press: 510-528. Boje, P and H. C. Johansen. 1998. “The Danish brewing industry after 1880.” In R.G. Wilson and T. R. Gourvish (eds.). The dynamics of the international brewing industry since 1800. London: Routledge. 5974. Boltanski, Luc and Ève Chiapello. 2006. The New Spirit of Capitalism. London-New York: Verso. ———— and Laurent Thévenot. 1999. “The Sociology of Critical Capacity.” European Journal of Social Theory 2: 359-77. ———— and Laurent Thévenot. 2006. On Justification. The Economies of Worth. Princeton: Princeton University Press. Bonneuil, Christophe. 2002. “The Manufacture of Species: Kew Gardens, the Empire and the Standardisation of Taxonomic Practices in late 19th century Botany.” In M.-N. Bourguet, C. Licoppe et O. Sibum (dir.). Instruments, Travel and Science. Itineraries of precision from the 17th to the 20th century. Routledge. 189-215. ————. 2006. “Mendelism, plant breeding and experimental cultures: Agriculture and the development of genetics in France.” Journal of the History of Biology 39(2): 281-308. ———— and E. Demeulenaere. 2007. “Une génétique de pair à pair? L’émergence de la sélection participative.” In F. Charvolin. (dir.). Les sciences citoyennes. Vigilance collective et rapport entre profane et scientifique dans les sciences naturalistes. Ed. de l’Aube. 122-147. [german version accepted in the Berliner Journal für Soziologie, to appear in 2008]. ———— and F. Thomas. In Press. Du maïs hybride aux OGM. Histoire de la génétique et l’amélioration des plantes à l’Inra. Quae, to appear late 2007. Bowler, Peter. 1983. The eclipse of Darwinism. Boyd, William. 2001. “Making Meat. Science, Technology and American Poultry Production.” Technology and Culture 42: 631-664. Buican, Denis. 1984. Histoire de la génétique et de l’évolutionnisme en France. Paris: PUF. Bulmer, Michael. 2003. Francis Galton: Pioneer of Heredity and Biometry. Baltimore: The John Hopkins University Press. Burian, R.M., J. Gayon and D. T. Zallen. 1988. “The singular fate of genetics in the history of French Biology, 1900-1940.” Journal of the History of Biology 21: 357-402. Bustarret J. 1944. “Variétés et variations.” Annales agronomiques 14: 336-362. Castle. William E. 1951. “The Beginnings of Mendelism.” In L. C. Dunn (ed.). American Genetics in the 20th Century. Essays on the Progress of Genetics During its First 50 Years. New York: The Macmillan Company. Chadarevian, Soraya de. 1996. “Laboratory science versus country-house experiments. The controversy between Julius Sachs and Charles Darwin.” British Journal for the History of Science 29: 17-41. Churchill, Frederick. 1974. “William Johannsen and the genotype concept.” Journal of the History of Biology 7: 5–30. Coleman, W. 1977 [1971]. Biology in the Nineteenth Century. Problems of Form, Function, and Transformation. Cambridge: Cambridge University Press. Darwin, Charles. 1868 [1990]. Variation of animals and plants under domestication, 2 vols. [The quotes’ pagination are from La découverte de l’hérédité. Une anthologie. Paris: Press Pocket. Douglas, Mary. 1966. Purity and Danger. An analysis of concepts of pollution and taboo. London [the quotes’ paginations are from the French translation from 2001. Paris: La Découverte & Syros]. East E. M. 1911. “The Genotype Hypothesis and Hybridization.” The American Naturalist 45(531): 160-174. Falk, Raphael. 1986. “What is a gene?” Studies in the History and Philosophy of Science 17: 133–173. Flavien, E. 1889. “La culture des graines, bulbes et plants reproducteurs. Maison Vimorin-Andrieux.” Les grandes usines de Turgan, revue périodique. Août 1889. (reprint consulted in the documentation services and archives from Vilmorin-Limagrain, La Ménitré).

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Fox-Keller, Evelyn. 2000. The century of the gene. Cambridge, MA: Harvard University Press. Freud, Sigmund. 1995 [1930]. Le malaise dans la culture [Civilization and Its Discontents]. Paris: PUF. Galton, Francis. 1876. “A Theory of Heredity.” The Journal of the Anthropological Institute of Great Britain and Ireland 5: 329-348. ————. 1889a. Natural Inheritance. Macmillan. ————. 1889b. “Recherches sur la fatigue mentale.” Revue Scientifique 43: 98-103. Gayon, Jean. 2000. “From measurement to organization. Aphilosophical scheme for the history of the concept of heredity.” In Peter Beurton, Raphael Falk, and Hans-Jörg Rheinberger (eds.). The concept of the gene in development and evolution: historical and epistemological perspectives. Cambridge: Cambridge University Press: 69–90. ———— and D. T. Zallen. 1998. “The role of the Vilmorin Company in Promotion and Diffusion of The Experimental Science of Heredity in France, 1840-1920.” Journal of the History of Biology 3: 241-262. Gayon, Jean. 1998. Darwinism’s Struggle for Survival. Heredity and the Hypothesis of Natural Selection. Cambridge: Cambridge University Press. Glamann, Kristof. 1988. Louis Pasteur and Carlsberg. A contribution to the history of pure yeast cultivation. Copenhagen: Carlsberg Foundation. Glass (Bentley). 1980. “The Strange Encounter of Luther Burbank and George Harrison Shull.” Proceedings of the American Philosophical Society 124(2): 133-153. Harwood, Jonathan. 1997. “The reception of genetic theory among academic plant-breeders in Germany, 1900-1930.” Sveriges Utsädesförenings Tidskrift 107 (4): 187-195. Hays Willet. 1905. “Breeding Problems.” Proceedings of the American Breeders’ Association 1, pt.2: 197. Quoted in Barbara Ann Kimmelman. A Progressive Era Discipline: Genetics at American Agricultural Colleges and Experiment Stations, 1900-1920 (Ph.D. dissertation, University of Pennsylvania, 1987). 379. Jacob, F. (1993). The logic of life. A history of heredity. Princeton, N. J.: Princeton Univ. Pr. Jenkins M. T. 1940. “The Segregation of Genes Affecting Yield of Grain in Maize.” Journal of the American Society of Agronomy 32 (1): 55-63. Jennings H. S. 1910. “Experimental Evidence on the Effectiveness of Selection.” The American Naturalist 44, No. 519: 136-145. ————. 1911a. “Pure Lines in the Study of Genetics in the Lower Organisms.” American Naturalist 45: 79-89. ————. 1911b. “‘Genotype’ and ‘pure line’.” Science 34 (885): 841-842. Johannsen, W. 1899. “Fortsatte Studier over Kornsorterne I. Om Variabiliteten med særligt Hensyn til Forholdet mellem Kornvægt og Kvæfstof-Procent i Byg.” Meddelelser fra Carlsberg Laboratoriet 4: 228313. ————. 1903. Über Erblichkeit in Populationen und in reinen Linien. Jena: Gustaf Fischer. ————.1905. Arvelighedslærens elementer. Førelæsninger holdte ved Københavns Universitet. Copenhagen: Gyldendalske bokhandel. ————. 1907. “Does Hybridization Increase Fluctuating Variability?” Report on the Third International Conference 1906 on Genetics. London. 98-113. ————. 1911. “The genotype conception of heredity.” American Naturalist 45 n•531: 129-159. ————. 1913. Elemente der exakten Erblichkeitslehre. Mit Grundzügen der biologischen Variationsstatistik. In dreißig Vorlesungen. 2nd edition. Jena: Gustav Fischer [1st ed. 1909]. Keller, Evelyn Fox. 2005. “The century beyond the gene.” Journal of Bioscience 30: 3–10. Knorr-Cetina, K. 1999. Epistemic Cultures: How the Science Makes Knowledge. Cambridge (UK): Harvard University Press. Kohler R. E. 1994. Lords of the Fly: Drosophila Genetics and the Experimental Life. Chicago Univ. Press. Lenay, Charles. 2000. “Hugo De Vries: From the Theory of Intracellular Pangenesis to the Rediscovery of Mendel.” Comptes Rendus de l’Académie des Sciences. Ser. 3, Sciences de la vie 323: 1053–60. Löwy, Ilana and Jean-Paul Gaudillière. 1998. “Disciplining cancer: mice and the practice of genetic purity.” In Jean-Paul Gaudillière and Ilana Löwy (eds.) The Invisible Industrialist. Manufactures and the Production of Scientific Knowledge. London: Macmillan. 209–249. Luria, S. 1973. Life, the unfinished experiment. New York: C. Scribner’s sons. MacKenzie, Donald, and Barry Barnes. 1979. “Scientific judgment: The Biometry-Mendelism controversy.” In Barry Barnes and Steven Shapin (eds.). Natural order: Historical studies of scientific culture. Beverly Hills, Calif.: Sage. 191–210. Mayr, Ernst. 1982. The Growth of Biological Thought. Diversity, Evolution and Inheritance. Cambridge, Mass. and London. Meijer, O. G. 1985. “Hugo de Vries no Mendelian?” Annals of Science 42: 189-232.

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Mendelsohn, Andrew. 2005. “Message in a Bottle: The Business of Vaccines and the Nature of Heredity after 1880.” In A Cultural History of Heredity III: 19th and Early 20th Centuries. Preprint 294. Berlin: MaxPlanck-Institute for the History of Science. 85-100. Meunissier A.. 1910. La loi de Mendel et ses applications, Versailles, Aubert et Cie. (tiré à part extr. du Bull des anciens élèves de l’ENHV). Müller-Wille, Staffan. 2005. “Early Mendelism and the subversion of taxonomy: Epistemological obstacles as institutions.” Studies in History and Philosophy of Biological and Biomedical Sciences 36(3): 465-487. ————. 2007. “Hybrids, pure cultures, and pure lines: From nineteenth-century biology to twentieth century genetics.” Studies in History and Philosophy of the Biological and Biomedical Sciences (In press). ———— and Vitezslav Orel. 2007. “From Linnaean species to Mendelian factors: Elements of hybridism, 1751-1870.” Annals of Science 64: 171–215. ———— and H-J. Rheinberger. 2007. (eds.). Heredity Produced. At the Crossroads of Biology, Politics and Culture, 1500-1870. Cambridge, Mass: MIT Press. ———— and H-J. Rheinberger. 2004. Heredity—The Production of an Epistemic Space. MPIWG. Preprint 276. Newman, L H. 1912. Plant Breeding in Scandinavia. Ottawa: The Canadian Seed Growers’ Association. Olby, Robert. C. 1979. “Mendel no Mendelian?” History of Science 17: 53-72. ————. 1985. Origins of Mendelism (2nd ed.). Chicago: Univ. of Chicago Pr. ————. 1987. “William Bateson’s introduction of Mendelism to England: a reassessment.” British Journal for the History of Science 20: 399-420. ————. 1990. “The emergence of genetics.” In R.C. Olby et al.. Companion to the history of modern science. New York & London: Routledge. Palladino, Paolo. 1990. “The political economy of applied research: Plant breeding in Great Britain, 19101940.” Minerva 28: 446–468. ————. 1993. “Between craft and science: plant breeding, Mendelian genetics, and British universities, 1900-1920.” Technology and Culture 34, n• 2: 300-323. ————. 1994. “Wizards and devotees: on the mendelian theory of inheritance and the professionalization of agricultural sciences in Great Britain and the United States, 1880-1930.” History of Science xxxii: 409444. Pickstone, J. V. 2000. Ways of knowing: a new history of science, technology and medicine. Manchester: Manchester University Press. Pasteur, Louis. 1876. Etudes sur la Bière. Paris: Gauthier-Villars. Paul, Diane B. and Kimmelman, Barbara A. 1988. “Mendel in America: Theory and practice, 1900-1919.” In Rainger et al. (eds.). American development of biology. Philadelphia: Univ. of Pennsylvania Press. 281310. Rabinbach, Anson. 1990. The Human Motor: Energy, Fatigue, and the Origins of Modernity. New York: Basic Books. [quoted pages are from the French translation. Paris: La Fabrique 2004]. Rader, K. 1999. “Of Mice, Medicine, and Genetics: C.C. Little’s creation of the inbred Laboratory Mouse, 1909-1918.” Studies in the History and Philosophy of Biological and Biomedical Sciences 30 (3): 319-343. Rheinberger, Hans-Joerg and Michael Hagner (eds.). 1993. Die Experimentalisierung des Lebens: Experimentalsysteme in den biologischen Wissenschaften, 1850-1950. Berlin: Akademie Verlag. Rheinberger, Hans-Jörg. 1997. Towards a history of epistemic things. Synthetizing proteins in the test tube. Stanford Univ. Press. ———— and Staffan Mueller-Wille. 2004. “Gene.” In The Stanford Encyclopedia of Philosophy,(http:// plato.stanford.edu/entries/gene/). Roll-Hansen, Nils. 1978. “The genotype theory of Wilhelm Johannsen and its relation to plant breeding and the study of evolution.” Centaurus 22: 201-235. ————. 1989. “The crucial experiment of Wilhelm Johannsen.” Biology and Philosophy 4: 303-329. ————. 1997. “The role of genetic theory in the success of the Svalöf plant breeding program.” Journal of the Swedish Seed Association 4: 196-207. ————. 2005. “Sources of Johannsen’s genotype theory.” In A Cultural History of heredity III, 19th and Early 20th Centuries. Preprint 294. Berlin: Max-Planck-Institute for the History of Science. 43–52. Russell, Nicholas. 1986. Like engend’ring like, Heredity and animal breeding in early modern England, Cambridge: Cambridge University Press. Sapp, Jan. 1987. Beyond the Gene: Cytoplasmic Inheritance and the Struggle for Authority in Genetics. New York: Oxford University Press. ————. 2003. Genesis. The evolution of biology. Oxford: Oxford University Press. Schaffer, Simon. 1994. “Empires of physics.” In R. Staley (ed.). The Physics of Empire. Cambridge: Whiple Museum of History of Science.

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Schribaux E. 1911. Notice sur les travaux scientifiques de M. E. Schribaux. Paris: Impr. de A. Tournon. Shull, G. H. 1908. “The composition of a field of maize.” Am. Breeders Assoc. Rep. 4: 296-301. (Report of the meeting held at Columbia, January 6-8 1908). ————. 1911. “The genotypes of maize.” American Naturalist 45(532): 234-252. ————. 1912a. “‘Genotypes’, ‘biotypes’, ‘pure lines’ and ‘clones’.” Science 35 (888): 27-29. ————. 1912b. “‘Phenotype’ and ‘clone’.” Science 35 (892): 182-183. Sibum, H. Otto. 1998. “An old hand in a new system.” In Jean-Paul Gaudillière and Illana Löwy (eds.). The invisible industrialist : manufactures and the production of scientific knowledge. London: Macmillan. Stamhuis Ida H., Onno G. Meijer and Erik J.A. Zevenhuizen. 1999. “Hugo de Vries on heredity, 1889-1903: Statistics, Mendelian laws, pangenes, mutations.” Isis 90: 238-267. Teich, Mikulás. 1983. “Fermentation theory and practice: The beginnings of pure yeast cultivation and English brewing, 1883-1913.” Technology and Culture 8: 117–133. ————. 2000. Bier, Wissenschaft und Wirtschaft in Deutschland, 1800-1914. Ein Beitrag zur deutschen Industrialisierungsgeschichte. Vienna: Böhlau Verlag. Theunissen, Bert. 1994. “Closing the door on Hugo de Vries’ Mendelism.” Annals of Science 51(3): 225-248. Thurtle Phillip. 1996. “The creation of genetic identity. The implications for the biological control of society.” Stanford Electronic Humanities Review, volume 5, (Supplement: Cultural and Technological Incubations of Fascism). http://www.stanford.edu/group/SHR/5-supp/text/thurtle.html. ————. 2002. “Harnessing Heredity in Gilded Age America: Middle Class Mores and Industrial Breeding in a Cultural Context.” Journal of the History of Biology 35(1): 43-78. ————. 2007. “The Poetics of Life: Luther Burbank, Horticultural Novelties, and the Spaces of Heredity.” Literature and Medicine, soon to appear. ————. 2008. Breeding True: Information Processing and the Rise of Genetic Rationality (University of Washington Press, In Vivo: Cultural Mediations of Biomedicine series, in press, kindly sent to me as personal communication). Vilmorin, Louis de. 1859. Notices sur l’amélioration des plantes par le semis et considérations sur l’hérédité dans les végétaux, précédées d’un Mémoire sur l’amélioration de la carotte sauvage. Paris : Librairie agricole. Vilmorin P. de. 1910. La Génétique et la quatrième Conférence internationale de génétique. Paris: Imprimerie de Duruy. Vries, Hugo de. 1907. Plant-breeding. Chicago: Open Court Publishing. Webber Herbert J. 1903. “New horticultural and horticultural terms.” Science 18 (459): 501-503. Wieland, Thomas. 2004. Wir beherrschen den pflanzlichen Organismus besser...: Wissenschaftliche Pflanzenzüchtung in Deutschland, 1889-1945. München: Deutsches Museum. Wise, Norton (ed.). 1995. The values of precision. Princeton Univ. Press.

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Mendelism and Agriculture in the First Decades of the XXth Century in Mexico Ana Barahona

Introduction History and philosophy of science have played a fundamental role in the comprehension of science in modern culture and society. Studies about history of science in Latin America began during the last four decades of the 20th century, developed under the model of European sciences and their influence. Many of these projects were based upon the diffusion model proposed by George Basalla in 1967,1 since it offered a historic, comparative and transcultural analysis, including epistemological and sociological considerations. However, despite being one of the first contributions to the field of social studies of science, the application of this model meant paying too much attention to the development of science within metropolitan areas without considering the local complexities, i.e. without considering the local characters of the so called “peripheral” countries, such as Mexico.2 Current studies in the sociology of science, philosophy of science and scientific literature have validated the comparative and local vision of historical work. These studies have identified central elements in the process of diffusion and have developed more precise ways to deal with its complexity.3 It is necessary to carry out historical studies that take in consideration the generated interactions after the first contact between imported scientific novelties and their result in local contexts. For example, how the introduction of scientific disciplines or techniques in different countries in Latin America has had different impacts in scientists’ status and their interaction in local political structures.4 This new vision demands the study of local organizations and scientific institutions focusing on the scientific and technical elites, which at different times and in different countries, have identified problems and offered solutions at the same time that they have given a series of beliefs, objectives and ideals to the scientific community. In this sense, the study of the “periphery” becomes a local study, ant its narrative depends in contextual aspects and not on general standars. Therefore, the introduction of genetics in Mexico will be treated as a social history of science and practices in a local context, and not as a result of “difusion” or imperialist/colonial imposition. This does not mean that the role played by the import of practices and techniques, resources and ideas, can be ignored; these elements will be treated as part of the conditions that allow us to explain the particular manner in which genetics was introduced in Mexico. During the 19th century plant and animal breeding relied basically on hybridization, massive selection and individual selection techniques, which where continuously modified according to 1 2 3 4

Basalla, 1967. Chambers, 1993. Latour, 1987, and Vessuri, 1994. Home and Kohlstedt, 1991; Petitjean, 1992; McClellan, 1992, Palladino and Worboys, 1993, and Vessuri, 1994.

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the place and organism in which they where utilized.5 In México, as in general in Mesoamerica, this kind of practices had deep cultural roots in the selection of maize varieties by peasants, that lead to the “creation” of the vast amount of biodiversity that characterizes the region. New theoretical knowledge, general theories such as Mendelism, and concrete knowledge of species and individuals radically changed the planning and execution of the breeder’s work at the beginning of the 20th century. The introduction of Mendelism was a practical asset insofar it changed the idea and purpose of plant breeders and hybridizers. 6 The acceptance and use of Mendelian laws of inheritance were connected to agriculture not only in Mexico but also in other countries such as the United States and England. However, because of its geographic proximity and the type of academic and technical exchange between Mexico and the United States at the beginning of the 20th century, the development of genetics in Mexico can be seen as parallel to that of the United States though there are significant differences. I will try to analyze the scientific conditions and social relations that allowed the introduction and establishment of genetics in Mexico in the early 20th century, which was consolidated and institutionalized during the second half of the century. I will examine the effect that small communities had during the introduction of genetics in Mexico, making an emphasis on the main role played by two groups, that of engineer Edmundo Taboada and that of the Rockefeller Foundation, during the period between the 1930’s up until the late 1950’s. In the first section I will try to briefly explain the first mention of Mendel's laws by Alfonso L. Herrera, despite the fact that it had no effect in the creation of institutions dedicated to the problems of inheritance. Regardless of the development of ideas and postures towards inheritance by the medical Mexican community in the 19th century, genetic principles did not contribute until the 1930’s through agricultural programs that intended to carry out plant breeding and that responded, in great measure, to economic necessities derived from political postures of Mexican governments after the Mexican Revolution that took place between 1910-1921. This subject will be treated in the second section. In the final part of the present work, the subject to be treated will be genetics applied to plant breeding which began during the government of General Lázaro Cárdenas del Río (1934-1940) under the guidance of agronomic engineer Edmundo Taboada Ramírez (1906-1983). Nonetheless, genetics applied to plant breeding was developed in two trends: the one introduced by Taboada in the Office of Experiment Stations (OCE, Oficina de Campos Experimentales, later called Agricultural Research Institute, IIA, Instituto de Investigaciones Agrícolas) and the one introduced by the Rockefeller Foundation. These two trends focused in solving problems of different strata of Mexican agricultural population.

5 6

During the twentieth century other technics such as the creation of mutations were introduced, and more recently genetic technologies, like the ones used in genetically modified organisms. Methodologically, there are two ways to approach the problems of inheritance. One is the study of pedigrees, as was the case in the 18th century when some human characteristics such as polydactyly, hemophilia and color blindness, were studied and recorded. The other method is by breeding that was employed by two schools in the 19th century, the species hybridizers and the animal and plant breeders, which had very different interests and objetives. See, Mayr, 1982.

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1. The first mention of Mendel in Mexico The years between 1810 and 1869 stand out in Mexican history as a period in which the country was immersed in a series of terrible internal conflicts and continued foreign interventions generated by capitalistic interests of European powers and the United States. During these years Mexico faced an attempt of reinvasion (Spain, 1829), several violent mutilations of its territory in the north caused by the United States and two wars with France (1838 and 1864). There were economic, administrative, political, social and cultural internal problems. In the field of science the immediate repercussion was translated into a certain impoverishment in comparison with the advances that were achieved during the time of the Borbonic reforms (last decade of the 18th century). The armed struggles originated by the Movement of Independence (1810-1821) caused the departure of the majority of Spanish and German scientists that had come from Europe following the Borbonic Reforms, scientists that had carried out a great labor within Novo Hispanic science. Nonetheless, as documented by Guevara Fefers scientific activity did not completely disappear and despite adverse condition some areas where developed.7 During the second half of the 19th century in Mexico, in good part due to French influence, the medical community had developed the notion of “inheritance” in the sense of understanding certain diseases that appeared recurrently in family lines or those that presented themselves in certain age-ranges, and that until that moment were incurable. The ideas of inheritance during the 19th century in Mexico suffered an important transformation; there was a transition from vitalism to reductionism, a change that was inplemented in the 1870’s with the introduction of positivist thought in Mexican intellectual circles. There was an impulse of experimentation for hypothesis verification and explanations were connected with material entities. Towards the end of the 19 th century Mexican scientists like other scientists around the world were searching for the general principles underlying the “laws of heredity.” At the beginning of the 20th century the Mexican medical community kept these ideas and it was not until 1904 that the first explicit reference to “Mendel’s law of dominance” appeared in the writings of Alfonso L. Herrera (1868-1942), but more in an evolutionary context than in a practical or applied one.8 Whether the medical community was not convinced of the truth and implications of Mendel’s theory or whether its most urgent interests were far from the theoretical problems involved in the transmission of hereditary diseases, in conjunction with the limitations that the economy applied to research budgets during revolutionary times, the fact is that programs about genetic investigation were not initiated in medicine nor in the incipient biology. 9 7 8

9

Guevara Fefer, 2002. Herrera was son of the notable Mexican naturalist Alfonso Herrera (1838-1901), who enjoyed many privileges from the government of President Porfirio Díaz (1877-1911). Alfonso L. Herrera obtained a degree in pharmacy in 1889, and was immediately appointed to the zoology and botany chair at the Teachers College (Escuela Normal para Maestros) and as an assistant naturalist at the National Museum (Museo Nacional) both in México City. In 1890 he was also appointed assistant in the Natural History Section of the National Medical Institute (Instituto Médico Nacional) in México City. In 1902, Herrera established the first general biology course in Mexico at the Escuela Normal and, in 1904, published the textbook Concepts of Biology (Nociones de Biología) to be used in the course. Herrera’s book and his teaching represent the first serious introduction of modern biology and Darwinism in Mexico. Barahona and Gaona, 2001.

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Without any doubt, Herrera is the most important Mexican biologist of the late 19 th and the early 20th centuries.10 He was a great connoisseur of Lamarck, Darwin and Haeckel, Trevinarius and Humboldt, Cuvier and Lyell as well as Hugo de Vries and Mendel. His two most important works Biología y Plasmogenia11 and Recueil des Lois de Biologie Generale12 are recognized as his most important scientific contributions to biology.13 In these two works Herrera speaks of variation in the context of his evolutionary conception (plamogeny) as produced by use and disuse and the direct influences of the environment. For Herrera there is an innate tendency to variation, but he blurs the line between variation and selection, in accordance to his Lamarckian vision of inheritance. For Herrera morphological and functional variations are caused by mutations. The mutagenic action is carried out by means of determinant influences of cellular physico-chemical factors. It may be seen that Herrera’s thought is linked more to the polemic between soft versus hard inheritance of the late 19th century in other countries.14 The work of Herrera constitutes an important bastion for Mexican biology. 15 However, in the field of genetics, Herrera’s work had no impact due to the lack of development of research lines on genetics and/or the lack of institutions dedicated to genetics research. Nevertheless, by 1900 Herrera participated actively in the Commission of Agricultural Parasitology (Comisión de Parasitología Agrícola), where agricultural research in Mexico was pioneered at the beginning of the 20th century. Established in 1900 by the Ministry of Development, its work was centered on fighting plant pests, specially the extermination of the orange fruit fly. According to Olea Franco it was created in the right moment because in 1899, the Horticultural Council of California had forbidden the import of Mexican oranges, as the fly that infested them was considered dangerous to the orange industry in California. In this way in 1900 Herrera was the head of a team created to research the orange plantation in the state of Morelos. Their first work was to start a campaign for the extermination of the pest. Herrera and collaborators doubted that the fly could get established in California because the climate was very different from Morelos, and because there were no pests in other states in the country. Despite these investigations the final disclosure was that the Mexican orange was forbidden in California, and in the following years research in the fruit fly larvae were

10 11 12 13 14 15

Beltran, 1951. Herrera, 1924. Herrera, 1897. Beltrán, 1968, and Beltrán, 1982. Mayr, 1982. Herrera was criticized because his approach was at odds with the urgent social needs of the time, which called for improved economic development and basic health, instead of advancing theories about the origin and evolution of life. His opponents insisted that practical studies should have priority over theoretical concerns. Moreover, Herrera’s evolutionary ideas were in conflict with religious and social prejudices held by political and religious sectors that had considerable social influence. The institutionalization of biology in Mexico was a complex process, closely related to the establishment of a biological community and the beginning of this discipline in Mexico, and thus leading to the formation of a specific discourse. This process was also influenced by the political environment of the time in which the revolutionary conflict (1910-1917) and then, the institutionalization of the Revolution (1929), motivated academic groups to look for better places to develop their activities. Control of biology returned to a community which had been previously consolidated. It was impossible to think of an autonomous biology that shifted away from medical control. See, Ledesma and Barahona. 1999; Barahona and Ledesma. 2002, and, Ledesma and Barahona. 2003.

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going to be one of the most important research projects of the Commission. It was dissolved in late 1907 and Herrera went back to his evolutionary studies.16

2. Political backgound and agriculture in Mexico Mendelian genetics was introduced in the United States and other countries through agriculture in the late 19th century and the early 20th century.17 In the United States animal and plant breeders that searched practical results, incorporated Mendelism more quickly than other academic groups. Genetics applied to plant breeding began to be used almost immediately after the “rediscovery” of Mendel’s laws by E. M. East and C. H. Shull in the Agricultural Experimental Station of Connecticut and in Cold Spring Harbor respectively in the year 1905. These first studies of inbreeding and crossbreeding were carried out in maize. Since then, this type of research spread to other universities of agriculture in the United States, such as the universities of Minnesota, California, Washington, Ohio, and Illinois among others. One of the most important achievements of the studies carried out in the United States was the production of double hybrid maize by Gorge H. Shull, Edward M. East and Donal F. Jones, in the late 1910’s when they were looking for the inheritance patterns of quantitative characters. 18 This development was marked by the economic interest that its introduction to agriculture meant for the large enterprises, therefore programs were developed that included among their purposes the introduction of hybrid maize in other countries like Mexico and Colombia, where the native varieties of maize with open pollination basically competed against the idea that hybrids were responsible for the increase in crops in the United States. This was an example of agricultural techniques and genetic knowledge exported from their point of origin, the United States, to peripheral countries like Mexico; however it took its own direction in Mexico in order to adapt to local conditions, necessities, and political circumstances. Scientific agricultural investigation, coordinated by government institutions has its origin in the Porfiriato (1877-1911). Porfirio Díaz’s government showed a great interest in encouraging agricultural exportation since this generated foreign currency and it helped to get the equilibrium in the balance of payments.19 The support for the generation of agricultural products of exportation included not only the legal facilities for producers, national as well as foreign, 20 but also the introduction of machinery and modern agricultural implements, as well as the application of medical and biological sciences in the care of crops and animals (mostly cattle). Díaz’ dictatorship favored higher education and scientific research in accordance with the French model, together with the positivist tradition, introduced to Mexico by Gabino Barreda during the regime of President Benito Juárez (1858-1861; 1865-1867; 1871-1872). Francisco I. 16 17 18 19 20

Olea Franco, A. 2002. The Comission was disolved by the Minister of Development Olegrio Molina, who founded the Estación Agrícola Central one year later and annexed it to ENA at San Jacinto. See Paul and Kimmelman, 1988, and Palladino, 1993. See East, 1936, and Shull, 1946. Webster, 1992. For example, for the acquisition of “uncultivated” properties (that in many cases belonged to indigenous communities) and the use of federal waters.

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Madero’s call for universal suffrage and the prohibition of reelection gave rise to an armed uprising (November 20, 1910) that marks the start of the Mexican Revolution. After Díaz’ resignation, Madero assumed the presidency on November 6, 1911, but he was assassinated in February 1913 by Mexican Army General Victoriano Huerta, who remained in power until 1914, as the war against the usurping government continued. After taking the capital city in 1915, Venustiano Carranza, one of the revolutionary leaders, headed a new government. Carranza promulgated a new political constitution in 1917, but was assassinated in 1920. Political instability prevailed through the 1920s, because the right wing forces continued the struggle and formed governments alternating with those of the Revolution until the late 1920s. During the administration of Porfirio Díaz, legal frameworks and agricultural tools were developed; agricultural research was stimulated by the creation of the first experimental stations, professional level agricultural education, and the modernization in 1907 of the National Agricultural College (Escuela Nacional de Agricultura),21 giving a more technical orientation to its careers. The modernization of the ENA was very important since it allowed the development and consolidation of agricultural instruction in Mexico, and became the cradle of technicians capable of connecting with agricultural communities, offering the benefits of scientific knowledge, taking advantage of the local farmers’ knowledge and providing orientation to agricultural politics. 22 After the revolution, new curricula and titles were created, such as “agronomic engineer,” “veterinarian,” and “technician in agricultural mechanics and agronomy.” 23 By the 1920s there were already programs for improvement of cotton cultivars, the study, introduction, and improvement of new and cultivated agricultural varieties, and the cataloging of hybrids and their possible uses. Since 1929, during the administration of Emilio Portes Gil (1928-1932), the Department of Agriculture and Promotion (Secretaría de Agricultura y Fomento) developed a plan to improve land redistribution and reorganize the production of the raw materials that the country needed. 24 Both activities were ideals that emanated from the Mexican Revolution. The “ejido”—a form of communal land-holding and social organization—was revived (its historical roots date from prehispanic and colonial times) under the slogan: “the land belongs to him who cultivates it;” it 21

22

23 24

The creation of the ENA goes back to the 19th century. In 1832 the Ospicio de Santo Tomás in the Federal District was transformed into the first School of Agriculture by government decree, imparting courses in botany, practical agriculture and applied chemistry, but it was closed because of political problems. Later in 1843 there was a second attempt for its creation but this one also did not work. It was not until 1850 that the school was created again in the Colegio de San Gregorio in the Federal District, and in 1853 it was fused with the Veterinary School, creating the National School of Agriculture and Veterinary. In 1854 it was moved to the Convent of San Jacinto, in the Federal District, as the ENA. In 1861 by decree of president Benito Juárez, the first “Organic Law of Public Instruction” was established, which made it dependant of the Ministry of Public Instruction, but in 907 it became dependent on the Ministry of Agriculture and Promotion. Between 1914 and 1917, the ENA was closed due to the armed conflicts and moved to the ex-hacienda de Chapingo in the State of Mexico in 1923, where it is found today as the Autonomous University of Chapingo. It is in that moment where it adopted the motto: “Teaching the exploitation of the land, not of man.” According to Olea Franco, enrollment at ENA was smaller in the middle 1920s than in the years 19081910. This means that enrollment grew more in the years prior to the revolution than under the revolutionary government of President Plutarco Elías Calles. After the 1920s the ENA population estimated was between 1 and 2.5 thousand students. Olea Franco, 2002. Reyes, 1981, p. 127. Portes Gil, 1929.

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could not be taxed or mortgaged because it was a family good transmitted only in a hereditary manner. In 1932, during president Abelardo L. Rodríguez government (1932-1934), the National Agronomic Commission (Comisión Nacional Agraria) was created within the Department of Agriculture and Promotion, with the following objectives: guaranteeing that the national plant and animal products would satisfy, totally and foremost, the needs of the whole population, and establishing the regulatory norms needed by public agencies, within the principles of an economy directed towards a social organization of agriculture based on the ejido. There was a great effort towards improving the teaching of technical agriculture, through the creation of the “agricultural engineer” major at the ENA, which had a totally practical approach. During the 1930s and 1940s, two political tendencies can be distinguished in Mexico’s power circles—with roots dating back to the Porfiriato which influenced research in plant genetics. On one side were those who, as heirs of the Mexican Revolution, believed that farmer agriculture, based on a tradition of communal land-holding, had priority over the creation of a successful agriculture; on the other side were those who thought that Mexican agriculture could only improve by becoming a large-scale private enterprise, far from socialist agrarianism. 25 During the administration of the General Lázaro Cárdenas del Río (1934-1940), research was started seeking to increase large scale food production, whereas during the Porfiriato a primary objective had been the exportation of grains. A main objective of General Cárdenas—a convinced agrarian—was to transform the organization of agriculture and to grant credit and technical support to farmers. The first agronomists trained in the new agricultural techniques shared the “Cardenist” philosophy and focused on solving problems affecting the average farmer. On March 18, 1938 General Cárdenas nationalized oil, provoking a dispute that lasted until the early 1940s. The foreign corporatations operating in Mexico rejected the right of the Mexican government to nationalize oil and not receiving any economic compensation. The American and British-Dutch oil companies and their governments, imposed economic sanctions on Mexico, and many Mexican imports such as silver, were at stake, because suspending them would have been an important economic blow to Mexico. The economic pressure imposed on Mexico by the US government and the oil companies took place in the midst of an economic crisis that made Mexico’s position unstable.26 The WWII context allowed the relative settlement of this dispute, with repayment of foreign loans being a never-ending task. The presence of the Vice-Presidentelect Henry A. Wallace at the inauguration of President Manuel Avila Camacho on December first, 1940, was a good sing of the changing times in bilateral relations. A crucial number of agreements for the wartime alliance were accelerated, and the US committed itself to loan money, give technical assistance, export technology, and help introducing in Mexico modern agricultural technologies.27 The scenario of a farmer policy based on the ejido changed drastically with the government of General Manuel Ávila Camacho (1940-1946); the capitalist tendency reappeared, supported by the private sector, favoring levels of production that would surpass the family needs of the ejido25 26 27

Hewitt de Alcántara, 1985. Meyer, 1977. Mexico provided laborers, petroleum, strategic mineral, henequen and other imprtant plants fibers. See Olea Franco, 2002.

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based farmer, so as to meet the food needs of the greatly expanding cities and, above all, the needs of the developing industries.28

3. The Office of Experimental Stations and the Rockefeller Foundation, two trends in Mexican agriculture. 3.1. THE WORK OF EDMUNDO TABOADA AND THE OFFICE OF EXPERIMENTAL STATIONS (OCE) Concern for the improvement of agricultural technology was an intermittent part of Mexico’s official politics since the beginning of the 20th century,29 however researches intended for the augmentation of food production did not begin until the 30’s, mainly during Cárdenas’ administration, and even with more strength during the 40’s, with the active participation of agronomic engineer Edmundo Taboada Ramírez. Edmundo Taboada Ramírez was born in Ciudad Guzmán, Jalisco, on December 12, 1906. He studied elementary school in Ciudad Guzmán and entered the ENA in 1922, where he graduated as an agronomic engineer with a master in irrigation in 1929. From 1928 to 1929, still as a student, Taboada worked as a topographer in the National Commission of Irrigation (Comisión Nacional de Riego) and later worked in the Sistema de Riego del Mezquital in Tula, Hidalgo, as a planning and calculation assistant. In 1930 he entered the Ministry of Agriculture in the Department of Chemistry and Soils and in 1931 became the head of the Special Analysis Section of the Central Chemistry and Soils Laboratory of the Agricultural Direction. None of Taboada’s teachers knew about genetics, so he read his first genetics book as a request of Waldo Soberón, the head of ENA and director of the Section of Experimental Stations when Taboada joined it, in order to explain certain principles to him. He had great mathematical skills, so he found quantitative genetics very appealing. He was self-taught and became an expert on genetics. From this came the idea of sending Taboada to study abroad and learn genetics formally. In 1932 Taboada traveled to Washington, D.C. for two months in order to study the analysis of soils carried out by the Bureau of Soils of the United States’ Agricultural Department. He was appointed agronomical attaché in the Mexican embassy in Washington, D.C., so that he could enter in the United States to study. In 1932 he studied genetics, applied plant genetics, cytology, mycology, physiology, and wheat, bean and maize improvement at Cornell University in New York under the direction of R. A. Emerson and H. K. Hayes, but he was not formally admitted into the graduate program or given official recognition for the courses he took. He would later receive an invitation by geneticist H. K. Hayes at the University of Minnesota, USA, to study plant parasitology, especially in wheat rust with Dr. E. C. Stakman. With the purpose of studying grinding methods and experimental planning with relation to the improvement of wheat varieties, Taboada traveled in 1933 to the Experimental Central Farm

28 29

Hewitt de Alcántara, 1985. As I have said, there were two opposed visions of rural development, one that emphasized the importance of adapting modern technologies in agriculture while slowing down agrarian reforms, and the other that urged for a deep reform in Mexican agriculture.

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in Ottawa, to study the organization of agricultural research programs that were used by the net of Canada’s Agronomic Experimental Stations. Upon his return to Mexico in 1934, Taboada was appointed Head of the Agronomic Experimental Station at El Yaqui, Sonora, a position from which he soon resigned due to medical reasons. At this station Taboada initiated his first genetic investigations, which consisted in selecting between different varieties of sesame those which were better adapted to Sonora’s environmental conditions. Since 1936 Taboada was a professor at ENA, where he imparted courses in general genetics, plants genetics and agronomic experimentation and investigation. He wrote the first book in genetics text in Mexico, Apuntes de Genética in 1938, with the intention of imparting his courses.30 This book encompasses the history of genetics, Mendelism, the chromosome theory of inheritance, cytogenetics, mutation, gene interaction, and population genetics. There are many references to Charles Darwin and natural selection, explaining how selection works on natural populations and the evolutionary process. Taboada describes with admiration the work of Thomas Hunt Morgan, the leader of the Drosophila research group at Columbia University and CalTech, and the work of Emerson, who was the leader of the Maize Genetics Group at Cornell University. For Taboada, the works of Darwin, Morgan, and Emerson were the cornerstone of biology. His textbook treated the various topics in a simple manner, with emphasis on the basic genetic principles. Taboada’s Apuntes de Genética became very important for teaching genetics and also for the popularization of genetics in Mexico. 31 With the creation of the Section of Experiment Stations in 1928 under the Ministry of Agriculture and Development, national programs of scientific research were promoted, especially for the genetic improvement of wheat and maize or phytotechnic, paving the way for the creation of an experimental station netwok. Taboada was appointed Head of the OCE in 1940, Director of the IIA from 1947 to 1960, Secretary of the National Council of Research and Superior Agronomic Instruction at the Ministry of Agriculture and Livestock from 1960 to 1964, and Consultant for the Ministry of Agriculture and Livestock from 1965 to 1970. As I have pointed out, the philosophy of economic development during Cárdenas administration was an agronomist one; Cárdenas believed that high productivity in the Mexican countryside was intimately linked to social structure changes that would transform the great capitalistic agronomic properties in cooperatives of peasants and farmer workers. In this manner, the first Mexican agronomists instructed in the application of new agronomic technologies shared Cárdenas philosophy and were more concerned with finding solutions to the practical and social problems faced by peasants than with importing foreign technology. 32 The Cardenist group headed by Taboada33 was formed and carried out its researches within the experimental stations where, for the first time, the more usual hybridization techniques that Taboada had imported from the United States and Canada were implemented. 34 The OCE was created in 1940 and Taboada was its first director. 35 At the beginning, this office coordinated ten experimental stations segregated in the entire country. During its first six years of life, in these stations different varieties of maize adapted to the ecological and economical 30

31

Taboada first taught mathematics and immediately afterwards genetics. When another geneticist, the spanish emigré José Luis de la Loma y de Oteyza, arrived at ENA in 1938, he passed down to him the genetic teaching reponsabilities. See, INIA, 1985. Taboada, 1938, and Barahona et al, 2003.

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conditions of producers from different states were selected. For example, the first variety of improved maize was obtained in 1940 and was called Celaya. It was an open pollinated variety obtained through selection, superior to the local varieties then cultivated and tested by farmers in the state of Guanajuato. Regarding wheat production, a collection of varieties from farmer’s fields were initiated, performance tests of the best agronomic qualities were carried out, and the first crossbreeding between high producing Mexican varieties, susceptible to the stem’s chahuixtle, and American varieties with low adaptation qualities but resistant to this disease, was accomplished. Researches with rice, sesame, sugar cane, rubber, beans, potato, cotton, olive, figs, hemp fiber and guayule were also carried out. For Taboada, the implementation of an experimental method was necessary since it could give an impulse to Mexican “scientifc” agriculture.36 The best way to carry out experimental work consisted in experimenting inside laboratories provided with special equipment (besides greenhouses, instruments for maize emasculation and controlled crossbreeding, among others) to obtain control of all possible varieties, installed in the experimental agronomic stations. Genetic improvement, always linked to Taboada’s experimental method, can be expressed in this way: it has been observed in certain species of plants like maize, which have a process of natural open pollination, that in the course of long successive self-fertilization, productivity is reduced to half of the previous generation. In this way, if the process is long enough, successive generations diminish to a very low state of productivity, due to the fact that populations become completely homozygotic. “As more successive self-fertilizations are carried out, progeny is each time more uniform. The increase of uniformity is fast in the first generations, but it becomes slower as the number of self-fertilizations escalates, if this number is large enough (about five or ten successive generations) populations become completely uniform.” 37 In this way, a self-fertilized line of maize by itself cannot be used as an agronomic seed. “When studying the decrease of productivity and the decrease of heterozygotic genotype, it is observed that the parallelism between both processes is considerably narrow. It is confirmed that the productivity of successive populations is strongly linked to the amount of heterozygotes present in those populations.”38

32

33

34

During Cárdenas’ administration 18 million hectares were distributed among communities and common lands (ejidos). In this manner, the amount of hectares in the social sector increased to 25 million (land outside private property). The object of the agronomic distribution during Cárdenas’ administration sought not only the satisfaction of a popular demand stated in the Constitution of 1917, but also the formation of small productive units, with self-feeding capacity. The basic unity of the Reform model was the conformation of common lands (ejidos). This refers to an endowment of lands that were given to a population nucleus so they could make use of it in the way they saw the fittest. Besides the endowment of lands and financing, the Agronomic Reform of the Cardenato included the establishment of an education system that allowed the formation of technical professionals to help in the development of common lands. Therefore, in association with common land nucleus, schools were created where children and young people should acquire knowledge of agriculture, cattle and other specific activities that the ecologic medium allowed. Between 1938-1940 from 20 to 30 people were working on sesame, sorghum, maize and beans. Among Taboada’s most oustanding collaborators were Eduardo Limón in El Bajío and Clemente Juárez in Torreón, Coahuila. Limón’s knowledge and training in experimental genetics was superior to Taboada’s. Limón worked in obtaining inbred lines of maize in order to cross them and obtain hybrid seed. See, Olea Franco, 2002. See Barahona and Gaona, 2001, and Gaona and Barahona, 2001.

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Taboada dedicated himself to produce what he called stabilized maize varieties starting from the open pollination varieties created during earlier years. “There are several types of high yield corn seeds. The highest yields are obtained with the so called “hybrid” corn, but their exceptional productivity only lasts the first cycle. In subsequent cycles, the productivity decreases so rapidly that sometimes its yield is inferior to that obtained with ordinary seeds, forcing the farmer to acquire new seeds each year […] Improved stabilized varieties with open pollination are another type of high yield maize […] Thanks to their characteristics, the open pollination varieties are better for our poorest farmers and are nearly as productive as the hybrid types.” 39 The seeds were collected from the peasants in different regions of the country, and tested in experimental stations, private fields, and even church gardens. Taboada and collaborators knew nothing about the characteristics of the seeds, thus they began planting them, describing the traits of plants and their growth, establishing times for sowing and harvesting, registering their reaction under different growth conditions and so on.40 In this way, research was seen as a learning process to improve agricultural practices. To obtain the stabilized corn varieties, Taboada first obtained lines with the fewest agronomic deficiencies and exhibiting good crossing results. Taboada would first cross any two given lines and select those particular combinations that exhibited high yield, obtaining eventually several combinations of lines that would be genetically stable, i.e., with productivity that remained high from one planting season to the next. These were distributed in the 1950s among Mexican farmers, especially in areas of small traditional farms (some of these varieties are still sown nowadays).41 In this way Taboada was able to develop the “stabilized” maize that has been widely used by farmers since, in spite of not having the highest yield, as it could be used in successive generations with no additional cost. Taboada claimed that farmers needed to get it only once and then could go on cultivating it and selecting seed from their own harvest. Taboada and his group accepted some varieties of hybrid maize then sown in the United States, from American colleagues in the 1940s. They did cultivation tests, but they concluded that the hybrid maize seed could not simply be planted elsewhere and, of course, the objetive of the IIA was to produce improve stabilized varietes of maize, not hybrid ones, that could be planted in all

35

36

37 38 39

Another institution created during Cárdenas’ administration was the Biotechnic Institute (Instituto Biotécnico, IB, 1934) within the Ministry of Agriculture and Development. The IB had a section on plant genetics and a botanical laboratory. It only lived 6 years, during which researches explored the evolutionary history of corn and used this information to purify indigenous maize crops. This is not surpising since its leader, Enrique Beltrán, was the most distinguished student of Alfonso L. Herrera (whose points of view towards biology and genetics has been discussed earlier in this paper). Researches at the IB motivated the notion that science ought to be used to purify farmer’s varieties by inbreeding and hybridization, to solve scientific, economic, social as well as cultural problems. When President Avila Camacho came to power in 1940, the IB was reorganized and closed doors, and the researchers were dispersed. See, Matchett, 2006. For Taboada, scientific research was necessary to improve the Mexican field, and the establishment of an agronomic experimental method, based on the laws of inheritance, was necessary as it could improve the scientific character of this discipline. The use of genetics was mainly about: 1) obtaining pure lines of native varieties; 2) the formation of new varieties through hybridization; and 3) the improvement, through hybridization, of created varieties, other already existing native varieties or imported ones. Taboada, E. 1981. Ibid. Secretaría de Agricultura y Ganadería, 1952.

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regions in the country inasmuch as they facilitated the acceptance by peasants and farmers, who did not require buying seed for every new crop. In1960 the IIA was merged with the Office of Special Studies, created in 1944 as part of a program of cooperation between the Mexican government and the Rockefeller Foundation.

3.2 THE OFFICE OF SPECIAL STUDIES AND THE ROCKEFELLER FOUNDATION The first activity of the Rockefeller Foundation (RF) in México was the 1923 campaign against yellow fever. After a major reorganization in 1928, the RF continued its emphasis on public health and medicine, but began to pay more attention to scientific education. Between 1940 and 1949 the RF launched a major agricultural program in Mexico, with two main goals, to improve food-crop production (corn and wheat) and to train Mexicans in agricultural techniques. 42 In the beginning of the 1940’s with the change of administration from Cárdenas to Manuel Ávila Camacho (1940-1946), the project of a capitalist orientation to agriculture reappeared in the government; the tendency was to increase production in the proper private sector of Mexican agriculture so that it could provide a surplus to feed the ever growing cities and could supply the new industries. This was due to rearrangements in the Mexican political class and the problems of supplying the great metropolitan areas. So, another group of Mexican researchers was formed, integrated in the Mexican Agricultural Program (Programa Agrícola Mexicano, PAM), specifically within the Office of Special Studies (Oficina de Estudios Especiales, OEE), a product of the involved cooperation between the Mexican government and the Rockefeller Foundation (FR) of the United States, in the introduction of the “technological package” characteristic of the “green revolution” that began in Mexico as a pilot project and was later transferred into other Third World countries. Since 1936, there was talk in the RF about beginning conversations with the Mexican government with the intention of cooperating in agronomic politics. However, the oil expropriation in 1938, that had an effect on North American and British enterprises, complicated the bilateral relations and set back the possibility of establishing a joint program. When General Manuel Ávila Camacho stepped into the presidency he began negotiation with the Rockefeller Foundation in 1941 and established a program of agronomic cooperation with the intent of working on the increase of agronomic productivity in Mexico. The committee sent by the Rockefeller foundation was formed by E.C. Stakman, head of the Phytopathology Division of the University of Minnesota, Paul Mangelsdorf, director of the Botanic Museum of Harvard University and Richard Bradfield, head of Cornell University’s Agronomic Department. The implementation of the PAM began in 1943 with Jacob G. Harrar as its first director. This program

40

41 42

Although it was easy to multiply the seed in the experimental stations without further expense, the peasants had no funds for doing that, because after harvesting the crop they would need money for packaging, treatment, and transportation. Then, “Taboada proposed to the State Government to get a loan and buy all Celaya improved seed, then store it and distribute it with the collaboration of the heads of the main maize-growing municipalities. The later should be asked to promote the seed for the following growing season and to set the price of seed by the ton…. Agricultural researchers would keep an amount of the income to continue seed multiplication.” Olea Franco, 2002. Taboada, 1960. See Cueto, 1994. For the role played by the RF in the rise of biology, see Kay, 1993.

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had as main objective the basic research of useful methods and materials to increase basic crops and enhancement of the formation and training of professionals. After the program officially began, Dr. Stakman returned to Mexico with the director of the program for the purpose of consulting Mexican scientists and to establish the basis and specific actions that should follow. Both proceeded according to the consulting committee’s preliminary report that recommended an initial focus on the following branches of agronomic science: 1) soils; 2) genetics; 3) disease and pest control; and 4) cattle. American specialists and Mexican researchers agreed, after an exhaustive study, on a two-objective plan: the central activity would be the fundamental research of useful methods and materials for the increase of basic alimentary sustenance crops; but since this program had to be, in time, totally Mexican, there was an agreement to enhance, as a second task, a training program for training Mexican researchers. 43 In that manner the OEE was born mainly dedicated to the breeding of maize and wheat, 44 and the introduction of a technological package of incomes and practices, improved seeds, chemical fertilizers, insecticides and herbicides, and irrigation, necessary elements for the exploitation of new, genetically improved, varieties. The OEE’s program reflected a scientific, technological and methodological content that was traditional of the agricultural colleges of the United States, whose work was intrumental in the commercialization of agriculture. The PAM headed in Mexico by the RF depended also on the connections with the financial and industrial institutions associated with the introduction of new agricultural technologies. Mexican government sponsored, through the OEE, international loans to Mexico for petrochemical inputs, agricultural machinery and equipment, genetically improved varieties of cattle, and pharmaceutical drugs. According to Olea Franco, these research activities were a case of tutelage of Mexican agricultural researchers by American scientists who were chosen by the RF to constitute the different research teams who worked in Mexico for about seventeen years. The social orientation carried out by the OEE was only one of the many expressions of the abandonment by the Mexican state of the agrarian reform program launched by Cardenas few years ago. 45 The “green revolution” had no success in Mexico regarding maize, 46 since it brought the polarization of different sectors of agriculture, due mostly to the fact that they were orientated to major producers that could buy machinery and incomes, and not to all farmers, and because wheat being a self-fertilizing plant that is very sensitive to latitude and altitude changes, it was assumed that farmers already knew their plants and had their techonologies and seed. 47 This revolution was later exported to India, Philippines and Pakistan, and from there to Afghanistan, Ceylon, Indonesia, Iran, Kenya, Morocco, Thailand, Tunes, and Turkey. 48 However, in terms of scientific research and technological application, the activities of the OEE were unprecedent in Mexico. By the end of 1945, seven American scientists employed and 43 44 45 46 47 48

Harrar, 1950, p. 14. In that time, 72% of the national surface was dedicated to the farming of these two grains. Olea Franco, 2002. The program of wheat breeding, carried out by Norman Borlaug was successful in Mexico. Borlaug was awarded the Nobel Peace Price for his work with wheat in 1970. The IIA tried to improve what the farmer knew about wheat, but did not substitute it for somenting else. Barahona and Gaona, 2001, and Gaona and Barahona, 2001. For the Green Revolution discussion in Mexico, see Fitzgerald, 1994; Frankel, 1963; Griffin, 1971, and Reyes, 1981. See also, Jennings, 1988.

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paid by the RF, with Ph.D. degrees, were heading the research programs. The number of American scientists with the same degree doubled. By the end of 1950, twenty-five Mexican graudates were part of the OEE, and their number increased in the following years. The RF had a scholarship program for graduate studies in the United States to young Mexican agrononomists who where part of the OEE. Mexican agronomists with graduate studies abroad became the directors of agricutural schools, experimental stations, research laboratories, and government head offices. It must be said, that during the presidency of Avila Camacho, the cooperation with the United States was crucial. In 1942, the Second Inter-American Conference on Agriculture took place. Seventy-six official delegates accompanied by 43 other members of official delegations, and 77 collaborating delegates took part of it. The first conference was held at Washington in 1930, but the second was a great success, bringing a large number of agricultural and plant scientists from all the Americas attending. A summary of the subjects discused at this conference included: soil conservation, soil surveys, and the standardization of techniques and terminology; control of agricultural pests and diseases; agricultural education, especially the provision of more scholarships for Latin Americans; and livestock improvement and the provisions of unified systems of registration for thoroughbred stock.49 Although the OEE was referred as part of the Ministry of Agriculture, the RF headed, staffed, and directed all its activities. It grew to 21 American scientists and 100 Mexican associates. By the 1950s more than two thousand maize varieties had been collected mostly in Mexico but also in North and South America, These varieties were kept in seed banks and treated as private property. The OEE contributed importantly to other research programs, such as the Institute of Research on Rice in the Philippines, the International Institute of Tropical Agriculture in Nigeria, The International Center of Tropical Agriculture in Colombia, and the International Center for the Improvement of Maize and Wheat (Centro Internacional del Mejoramiento del Maíz y Trigo) in Mexico. Towards the end of the decade of the 1980s, it became apparent that there was no reason to keep two institutions dedicated to plant improvement. The OEE was being increasingly directed by Mexican specialists who had been trained with the aid of the RF, while the latter’s interests were focused on the exportation of the Green Revolution’s new technology to other South American countries, especially Colombia, so that it increasingly left the running of the Office in Mexican hands. In 1961, the IIA merged with the OEE, forming the National Institute of Agricultural Research (Instituto Nacional de Investigaciones Agrícolas). This institution took control of all the experimental fields, equipment, and personnel.

Conclusions The introduction of genetics in Mexico had a parallel with the development in the United States and Europe only in some of its currents. Before 1900, where there was not a discourse on genetics, 49

Olea Franco, 2002. Although one of the main subjects of the Conference was the creation of a unified system of registration, intellectual property in México only began in the 1990s. In previous times, the only way to guarantee the intellectual property was to hide all the procedures to obtain the varieties.

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but rather of a broader term of inheritance, it was handled and conceptualized in Mexico within medicine. After 1900, when genetics surface as a science, its more practical or technological side, phytotechnics or plant breeding was developed in Mexico. Interest in plant breeding was developed by agronomists with the support of the Mexican governments as long as this represented higher yields and therefore an increase in the economical value of their crops. Nevertheless, research in classic genetics with the intent to discover general principles, the construction of genetic maps, or the explanation of evolutionary patterns, was not practiced in Mexico until the 1960s, in the way that it was practiced in other countries like the United States. However, Mexican researchers that were instructed in phytotechnics, possessed the full body of basic genetic knowledge necessary to understand the hereditary mechanisms that took place during experimentation (as proven by Apuntes de Genética written in 1938 by E. Taboada). They also had a considerable understanding of population genetics, which was being developed independently from maize genetics. The programs developed by the IIA and the PAM were specific and comprised almost exclusively the improvement of varieties with commercial and economic value. Independently from the political tendencies and the social and economic level of agronomic producers towards whom the positive results of genetic improvement carried out by Edmundo Taboada and the PAM were intended, methodologically, both lines followed a common research pattern. It always began by collecting genetic material conformed by the seeds of the plants subject to experimentation (maize, wheat, beans etc.) coming from different parts of the Republic or imported, its planting in experimental fields and the observation of characteristics of phytotechnic interest that each possessed, followed by the selection of those varieties that presented the most adequate characteristics for the intended purpose (greater yield, greater resistance to disease, early maturation, etc.). Once these varieties were obtained, experimentation could proceed by means of crossbreeding with the intention of producing hybrid varieties with even better characteristics than the parental varieties. However, the conception and use of breeding techniques lead to the instrumentation of different agronomic practices according to political positions. One of the main objectives of IIA was the implementation of experimental stations in different parts of the country in order to increase the production of wheat, corn, cocoa, rice, sesame seed, and beans. For Taboada, the goal of Mexican geneticists was to genetically improve varieties and successfully adapt them for planting in the different agricultural regions of the country. One of the biggest successes of the Institute was to get a variety of corn with high productivity similar to that of hybrid corn, but which would retain its high productivity from one harvest to the next, without the need of producing new hybrid seed for each planting season. Taboada’s stabilized corn varieties were the most important achievement of Mexican agriculture. This work was focused on solving farmers’ problems and improving maize genetics research in Mexico. The stabilized corn varieties benefited farmers but didn’t contribute significantly to economic change in large-scale agriculture. Economically, the introduction of hybrid seed was more important. It led to the capitalization of the farms and the creation of a flourishing business, namely, the production and sale of seed. Taboada argued there were some substantial differences between research and extension programs at IIA and OEE. Taboada claimed that an agricultural researcher must know the

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characteristics not only of land farmers crops, but also their needs and problems, and local practices for sowing and harvesting, otherwise he would be isolated without knowing the reality, nor frame of reference or idea, even if he knows what books say. 50 In regard to maize improvement, the approach of the OEE and the RF was contrary to Taboada’s. The OEE dedicated most resources to the production of high yield hybrid seed that could only be purchased by farmers who had substantial financial resources. This seed performed best with fertilizers and its efficiency depended upon being planted in irrigated areas. The OEE’s approach had prevailed during the 1950s in the Department of Agriculture. In 1948, 80% of corn cultivars had been planted with open pollination varieties, but by 1956 the production program of the Department dedicated 96% of its capacity to hybrid seed production, which benefited the commercial production of corn and irrigated agriculture.51 These two tendencies shared the same objectives, to achieve an increase in basic food production in Mexico, and also the same methodologies of Mendelian hybridization; however, they focused agronomic research in different manners. Besides, there was an important difference in the economic and administrative support provided by the federal government during the 40’s and 50’s. The political discrepancies generated a rather distant relationship between both institutions during those decades. Ana Barahona Departamento de Biología Evolutiva Universidad Nacional Autónoma de México [email protected]

50 51

Taboada, 1985. For discussion about the influence of the PAM on Mexican Agriculture, see Fitzgerald, 1994, and Cotter, 1994.

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References Barahona, Ana and Ana L. Gaona. 2001. “The History of Science and the Introduction of Plant Genetics in México.” History and Philosophy of the Life Sciences 23: 157-168. Barahona, Ana and Ismael Ledesma. 2002. “El Positivismo y los Orígenes de la Biología Mexicana.” Archives Internationales d’Histoire des Sciences 149(52): 277-305. Barahona, Ana, Susana Pinar and Francisco J. Ayala. 2003. La Genética en México. Institucionalización de una Disciplina. México: Universidad Nacional Autónoma de México. Basalla, George. 1967. “The Spread of Western Science.” Science 156: 611-622. Beltrán, Enrique. 1951. “Panorama de la Biología Mexicana.” Revista de la Sociedad Mexicana de Historia Natural 12: 69-99. ————. 1968. “Alfonso L. Herrera, Primera Figura de la Biología Mexicana.” Revista de la Sociedad Mexicana de Historia Natural 29: 37-91. Beltrán, Enrique. 1982. Contribución de México a la Biología. México: Ed. Cecsa. Chambers, David Wade. 1993. “Locality and Science. Myths of Centre and Periphery.” In Antonio Lafuente, Alberto Elena and María Luisa Ortega (eds.) Mundialización de la Ciencia y Cultura Nacional. Madrid: Doce Calles/Universidad Autónoma de Madrid. 605-617. Cueto, Marcos. 1994. “Visions of Science and Development. The Rockefeller Foundation’s Latin America Surveys of the 1920s.” In Marcos Cueto (ed.) Missionaries of Science. The Rockefeller Foundation & Latin America. Bloomington: Indiana University Press. 1-22. East, Edward Murray. 1936. “Heterosis.” Genetics 21: 375. Fitzgerald, Deborah. 1994. “Exporting American Agriculture. The Rockefeller Foundation in Mexico, 19431953.” In Marcos Cueto (ed.) Missionaries of Science. The Rockefeller Foundation & Latin America. Bloomington: Indiana University Press. 72-96. Frankel, Sir Otto H. 1963. “Some Conceptual Aspects of Technical Change.” In Jean Maynaud (ed.) Social Change and Economic Development. Leiden. Gaona, Ana L. and Ana Barahona. 2001. “La Introducción de la Genética en México. La Genética Aplicada al Mejoramiento Vegetal. ” Asclepio 53(2): 23-40. Griffin, Keith. 1971. The Green Revolution and Some Issues of Economic Policy. Geneva: United Nations Research Institute for Social Development. Guevara Fefer, Rafael. 2002. Los últimos años de la historia natural y los primeros días de la biología mexicana. La práctica científica de Alfonso Herrera, Manuel Ma. Villada y Mariano Bárcenas. México: Instituto de Biología, Universidad Nacional Autónoma de México. Harrar, Jacob George. 1950. The Mexican Agricultural Program. A Review of the First Six Years of Activity Under the Joint Auspices of the Mexican Government and the Rockefeller Foundation. Rockefeller Foundation, New York: Leo Hart Co. Herrera, Alfonso. L. 1897. Recueil des Lois de Biologie Générale. México: Oficina de Tipografía de la Secretaría de Fomento. ————. 1904. Nociones de Biología. México: Imprenta de la Secretaría de Fomento (Facsimilar Edition, México: Universidad Autónoma de Puebla, 1992). ————. 1924. Biología y Plasmogenia. México: Ed. Herrero Hermanos Sucesores. Hewitt de Alcántara, Cynthia. 1985. La Modernización de la Agricultura Mexicana, 1940-1970. México: Siglo XXI Editores. Home, Roderick Weir and Sally Gregory Kohlstedt (eds.). 1991. International Science and National Scientific Identity. Australia between Britain and America. Dordrecht: Kluwer Academic Publishers. Instituto Nacional de Investigaciones Agrícolas (INIA). 1985. Edmundo L. Taboada Ramírez. Una semblanza, 1906-1983. México: Secretaría de Agricultura y Recursos Hidráulicos. Jennings, Bruce. H. 1988. Foundations of the International Agricultural Research. Science and Politics in Mexican Agriculture. Boulder: Westview Press. Kay, Lily E. 1993. The Molecular Vision of Life. CalTech, the Rockefeller Foundation, and the Rise of the New Biology. New York: Oxford University Press. Latour, Bruno. 1987. Science in Action. Milton Keynes: Open University Press. Ledesma, Ismael and Ana Barahona. 1999. “Alfonso Luis Herrera e Isaac Ochoterena. La institucionalización de la Biología en México.” Historia Mexicana 48-3(191): 635-674. Ledesma, Ismael and Ana Barahona. 2003. “The Institutionalization of Biology in Mexico in the early 20th Century. The Conflict Between Alfonso Luis Herrera (1868-1942) and Isaac Ochoterena (1885-1950).” Journal of the History of Biology 36(2): 285-307. Lewontin, Stephen. 1983. The Green Revolution and the Politics of Agricultural Development in Mexico since

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1940. Thesis (Ph. D.) University of Chicago. Matchett, Karin. 2006. “At Odds over Inbreeding. An Abandoned Attempt at Mexico/United States Collaboration to “Improve” Mexican Corn, 1940-1940.” Journal of the History of Biology 39(2): 345-372. Mayr, Ernst. 1982. The Growth of the Biological Thougth, Cambridge: Harvard University Press. Meyer, Lorenzo. 1977. Mexico an the United States in the Oil Controversy, 1917-1942. Austin: University of Texas Press. McClellan, James Edward. 1992. Colonialism and Science. Saint Dominguez in the Old Regime. Baltimore: Johns Hopkins University Press. Olea Franco, Adolfo. 2002. One Century of Higher Education and Research in Mexico (1850s-1960s) with a Preliminary Survey on the Same Subjetc in the United States. Thesis (Ph. D.) Harvard University. Palladino, Paolo. 1993. “Between Craft and Science. Plant Breeding, Mendelian Genetics, and British Universities, 1900-1920.” Technology and Culture 34(2): 300-323. Palladino, Paolo and Michael Worboys. 1993. “Science and Imperialism.” Isis 84:91-102. Paul, Diane and Barbara Kimmelman. 1988. “Mendel in America. Theory and Practice, 1900-1919.” In Ronald, Rainger, Keith Rodney Benson and Jane Maienschein (eds.) The American Development of Biology. Philadelphia: University of Pennsylvania Press. 281-312. Petitjan, Patrick. 1992. “Sciences et Empires. Un Thème Promotteur, des Enjeux Cruciaux.” In Petitjean, Patrick, Catherine Jami, and Anne Marie Moulin (eds.) Science and Empires. Historical Studies about Scientific Development and European Expansion. Dordrecht: Kluwer Academic Publishers. 3-12. Portes Gil, Emilio. 1929. Informe del C. Presidente de la República al H. Congreso de la Unión en la parte relativa al ramo de Agricultura y Fomento. México: Ed. Cultura. Reyes, Pedro. 1981. Historia de la Agricultura. Información y síntesis. México: A.G.T. Editor S. A. Secretaría de Agricultura y Ganadería, 1952. Informe de Labores. 1951-1952. México. Shull, George Harrison. 1946. “Hybrid Seed Corn.” Science 103: 549. Taboada, Edmundo. 1938. Apuntes de Genética. México: Escuela Nacional de Agricultura, Chapingo. ————. 1960. Líneas Homocigotas I. México: Instituto de Investigaciones Agrícolas, Secretaría de Agricultura y Ganadería. ————. 1981. “Mejoramiento Genético.” Conference dictated in CAEZAC, in Calera, Zac. México. ————. 1985. “Ingeniero Edmundo Taboada Ramírez.” In Edmundo L. Taboada Ramírez. Una Semblanza 1906-1983. México: Secretaría de Agricultura y Recursos Hidráulicos. Vessuri, Hebbe. 1994. “The Institutionalization Process.” In Jean-Jacques Salomon, Francisco R. Sagasti, and Céline Sachs-Jeantet (eds.) The Uncertain Quest. Science, Technology and Development. New York: United Nations University Press. 168-200. Webster, G, von. 1992. “La Agricultura en el Porfiriato. 1876-1911.” In Primer Simposio de ciencias agronómicas. México: Colegio de Postgraduados en Ciencias Agrícolas de Chapingo. 135-137.

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Herbert Spencer Jennings, Heredity, and Protozoa as Model Organisms, 1908-1918 Judy Johns Schloegel

In early 1908, Herbert Spencer Jennings published his first research exploring hereditary phenomena in protozoa. The article, which explored the fate of “new or acquired” structural characteristics in protozoa, was the first in a series of articles continuing through 1917 that appeared under the heading of “Heredity, Variation and Evolution in Protozoa.” Jennings conceived of the research in this series as a logical progression from his earlier research on the behavior of lower organisms, which he carried out in the ten years following earning his Ph.D. at Harvard in 1896. As he explained at the outset of this article, although this new research appeared to be a “complete departure” from his earlier experimental program, it was “in reality a logical continuation” of that earlier work, which foregrounded the question of how behavior “happens to become so largely adaptive.” 1 Jennings explained that many behaviors, identified as processes such as learning or habit formation, were found to arise in the lifetime of the individual organism. Some adaptive behavioral features, however—referred to with such terms as reflex, tropism, or instinct—did not arise in the lifetime of the individual, but were said to be inherited from one generation to the next. Uncovering the processes by which adaptive characters were inherited, Jennings explained, was the problem that he subsequently aimed to tackle. 2 The two research programs were unified by a further feature, however—namely, the use of protozoa as experimental research organisms. Jennings’ decision to use the unicellular protozoa in his earlier behavioral studies was informed by two critical lines of thought: (1) the nascent conceptualization of protozoa as models of biological phenomena and of other living entities—in this case, of cells in multicellular organisms; and (2) adherence to an evolutionary framework that emphasized the biological importance of the protozoa due to their apparent simplicity and primordial nature. Both concepts emerge in Jennings’ thought in 1896, when he began to plan and carry out a semester of post-doctoral research funded by Harvard University in the laboratory of the physiologist, Max Verworn, in Jena. Following the completion of his doctoral dissertation, which was largely critical of numerous accounts of cell cleavage grounded in developmental mechanics (Entwicklungsmechanik), Jennings aimed in his postdoctoral research to produce what he believed would be a more satisfactory account of early development. In Verworn’s laboratory, he embarked on experimental study of the behavior of unicellular organisms with the objective of producing models of the actions of groups of embryological cells. Jennings largely abandoned this original project, however, as he became interested in the behavioral responses of unicellular organisms as scientific objects in their own right. He nonetheless continued to develop a conception of protozoa as models of general biological phenomena in the years that followed. Jennings’ conceptualization of protozoa as models was circumscribed by his adherence to progressive evolution, which he adopted during his residence in Jena, well-known as the 1 2

Jennings (1908a), p. 578. Ibid., pp. 578-9.

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intellectual center of German evolutionary thinking. His adoption of a progressive evolutionary framework was informed particularly by the teachings and writings of Verworn, who was himself a devoted student at Jena of Ernst Haeckel and the evolutionist and physiologist, Wilhelm Preyer. Verworn championed the utility of protozoa for physiologists due to their standing “nearest to the first and simplest forms of life.”3 Jennings, like Verworn, believed that such organisms in turn exhibited the simplest and most evolutionarily fundamental living phenomena that generally were obscured in higher organisms by the evolution and overlayering of increasingly complex phenomena in higher organisms.4 In this paper I consider the rationale for Jennings’ use of protozoa as model organisms of hereditary phenomena and the central role these organisms played in his articulation and defense of a broad conception of heredity in the early decades of the twentieth century. Further, I wish to consider Jennings’ relationship to a genetics/heredity enterprise, or epistemic space, as it was being defined between 1907, when Jennings embarked on hereditary research, until the end of the Great War.5 From the outset, Jennings was motivated in his hereditary studies—as was the case with other leading American geneticists such as William Castle and T. H. Morgan—to demonstrate evolution experimentally.6 Unlike many early geneticists, however, Jennings adhered to the principle of progressive evolution, leading him to prioritize the study of hereditary phenomena in the simplest organisms possible. In each of the major hereditary problems that he and others debated during the period—including the efficacy of the inheritance of acquired characteristics and of selection, and the significance of pure lines and Mendelian inheritance, Jennings turned to the asexually-reproducing protozoa for experimental insights. While the effort to uphold a generalized conception of heredity on the basis of research with specific organisms may appear counter-intuitive, for Jennings, the unique evolutionary status of the protozoa as the simplest of the cellular organisms made them perfect for illuminating hereditary mechanisms at their most fundamental level.

*** By the time of the publication of his first article on heredity on the eve of his fortieth birthday, in 1908, Jennings had established himself as one of the United States’ foremost zoologists. His 1906 monograph, The Behavior of Lower Organisms, while widely criticized and debated for its progressive evolutionary assumptions, was hailed as methodologically incontrovertible; in the same year Jennings earned the coveted position as successor to William Keith Brooks at the Johns Hopkins University, and served as director of the Zoological Laboratory until he retired in 1938; he served as president of both the American Society of Zoologists in 1909 and the American Society of Naturalists in 1910; and, during the same period, he became recognized as a rare and formidable philosophical thinker in the American biological institutional landscape. It was with a

3 4 5 6

Verworn (1899), p. 51. See Schloegel (2006), especially pp. 49-69. On the epistemic space of heredity at the turn of the century, see Müller-Wille and Rheinberger (2005). See also Rheinberger (1997). See for example, Allen (1978) on Morgan and Rader (1998) on Castle.

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certain level of confidence that Jennings set out to tackle fundamental problems of heredity at a moment of great ferment in hereditary thought.7 In his first article on heredity, published in 1908, Jennings maintained that the primary interest that guided his study was that of the evolution of unicellular organisms. 8 Later that year, in the second article in the series, he refined the “central problem” that concerned him as one of heredity.9 Such an evolution in thinking was certainly not unusual among zoologists, many of whom came to the problems of heredity through their preoccupation with the unresolved problems of evolution. More specifically, however, Jennings, like his contemporary, T. H. Morgan, was guided by a primary concern about the mechanisms underlying the evolution of adaptive features.10 Jennings is notable throughout his early publications on heredity for his agnosticism about the possible mechanisms of heredity and evolution. In his first article on heredity in 1908, for example, he indicated his agnosticism as he defined his use of the term: “I use the word ‘heredity’ merely as a brief and convenient term for the ‘the resemblance between parents and progeny,’ without implying any underlying entity, and without prejudice about the grounds of this resemblance.” While rejecting the likelihood that natural selection among individual organisms could be the sole mechanism of evolution, Jennings aimed to turn attention to the mechanisms underlying the “internal adaptations” within organisms. 11 In his plan for elucidating “how organisms have arisen,” Jennings aimed first to clarify what he considered to be the “normal” processes of heredity and variation, i.e., to uncover the similarities and differences that are normally found to arise in the passing from one generation to the next. With such baseline information in hand, he envisioned, it would then be possible to intervene experimentally in these normal—or “racial”—processes to investigate the primary question of how inherited modifications arise.12 Critical to this undertaking was the use of the “simplest organisms.” The protozoa were valuable, Jennings explained, because of their rapid rate of reproduction (at least one generation a day) and most especially because reproduction occurred in the “simplest forms.” 13 In particular, the protozoa were of interest due to the widespread assumption among zoologists that, since they don’t separate into somatic and germ cells, they possess fundamentally different hereditary processes than those of the metazoa. Since reproduction in the protozoa occurs by simple division, i.e., they reproduce asexually, many had concluded that the protozoan progeny are the same as the parents—as Jennings himself wrote in 1906, that “the offspring are the parents, merely subdivided.”14 This assumption was generally accompanied by the reasoning that, since there is no distinction between the soma and the germ in protozoa, characteristics attained by the parents would be perpetuated in the offspring. Consequently, as Jennings explained, “if the difference really exists, the Protozoa are much more plastic in evolution than are the Metazoa.” 15 The first 7 8 9 10 11 12 13 14 15

On Jennings, see Schloegel (2006); Kingsland (1987); Sonneborn (1975); Ritter (1912). Jennings (1908a), pp. 577-583. Jennings (1908b), pp. 393-4. Morgan (1903). Jennings, 1908a, pp. 584 (footnote), 582. Ibid., p. 583. Ibid. Ibid., pp. 584-5; Jennings (1976 [1906]), p. 320. Jennings (1908a), p. 584.

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experimental task that Jennings set for himself in his hereditary studies then was to determine whether characteristics acquired by the individual during its lifetime are in fact perpetuated in their progeny. Specifically, Jennings turned his attention to the inheritance of localized, structural characters in the ciliate Paramecium, as compared to the unlocalized characters brought about by such processes as acclimatization, which affect the organism as a whole. The inheritance of localized characters was viewed by some, particularly August Weismann, to achieve a higher standard of proof of inheritance. Through isolation and culturing, Jennings pursued a series of experiments that followed the transmission of structural abnormalities that appeared naturally in the population.

Figure 1. Transmission of structural abnormalities in Paramecium. From H. S. Jennings, “Heredity, Variation, and Evolution in Protozoa. I. The Fate of New Structural Characters in Paramecium, in Connection with the Problem of the Inheritance of Acquired Characters in Unicellular Organisms,” The Journal of Experimental Zoology 5 (1908): 577-632, 594.

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In several series cultured from an unusually bent individual paramecium over many generations, the abnormality was transmitted to only one of the two individual progeny, thus failing to produce a new race. In one case, however, Jennings was able to observe the formation of what he considered a new race, when he followed a line in which the individuals resulting from fission remained united in chains. Chains of individuals were inherited in all subsequent generations of both the anterior and posterior fission products, demonstrating the basic insight that, if a new character is to be inherited, the modification to the parent cell causes it to somehow behave differently at reproduction, thus causing it to produce the characteristic anew in each progeny . 16

Figure 2. Inheritance of Chains in Paramecium. Ibid., p. 600.

Jennings, of course, realized that this demonstration of the inheritance of chain formation did not involve some kind of germinal material—and this was exactly his point. The case of chain formation enabled him to demonstrate the fundamental similarity between protozoa and metazoa—a similarity that he had come to recognize only recently—and, at the same time, to demonstrate the value of protozoa as model organisms for the study of heredity. As Jennings noted,

16

Ibid., pp. 618-25.

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It is of course possible that the origin of new permanently inherited characters is not normally through mere modifications of the external parts of the cell, such as we see in our illustrative cases [of chain formation.] Possibly there must be originally some modification of more recondite parts—nucleus, chromosomes, or the like—and that these then secondarily act upon and change the outer parts. This would add farther complication, but would not change the essential point, which is that in order that a characteristic may be inherited, it must be due to some modification that causes a change in the processes of reproduction.17

In the first case, Jennings maintained that his observations demonstrated that the protozoa were not, in fact, more plastic than the metazoa. Furthermore, his investigations demonstrated that the barrier to the inheritance of acquired characters was not the separation of the germ and soma, but rather the process of cell division, which meant that “the problem of how new inherited characters arise is the same in Protozoa as in Metazoa.”18 This consequently supported his second point that protozoa were ideal organisms for the study of how new inherited characters arise, since: (1) the basic hereditary process was fundamentally the same in both protozoa and metazoa; (2) at a practical level, the protozoa multiply rapidly, for expedient results; and (3) finally, the single cellularity of protozoa made their exposure to environmental influences more feasible and, at the same time, hereditary effects in them more readily observable.19 Over the course of the next few years, Jennings became increasingly committed to the value of protozoa as model organisms in the study of heredity despite the largely negative results that he continued to receive. In his second heredity paper published in 1908, Jennings turned his attention from the inheritance of acquired characteristics to selection, as another possible means for demonstrating evolution experimentally. In these investigations, focused on the size of Paramecium, he found that the large amount of variability in the progeny descended from a single individual was largely attributable to growth of the individuals in the course of the life cycle and to different environmental conditions. When growth and environment were controlled to the extent possible, the remaining variability exhibited by the line of descendents could not be affected by selection and thus was not heritable: despite persistent efforts to select for the largest and smallest offspring, the individuals in the resulting line did not deviate from the mean size of the line.20 These results gave way to Jennings’ adoption of Johannsen’s notion of a “pure line” as a label that more tellingly communicated the imperviousness of races, or the series of individuals descended from a single individual, to selective pressures. 21 Jennings’ thinking about pure lines appears to have cemented his commitment to the utility of protozoa as model organisms. In three different articles appearing in The American Naturalist between 1909 and 1911, Jennings again championed the various virtues of protozoa for use in the study of heredity and variation. In particular, he emphasized the value of protozoa for shedding light on debates about the effectiveness of selection and in making concrete the controversial notions of “pure line” and “genotype,” which were held by many to be hypothetical or theoretical 17 18 19 20

21

Ibid, p. 625 (footnote). Ibid., p. 627. Ibid. Jennings maintained that if growth and environmental conditions could be completely controlled, “all the evidence indicates that the standard deviation and coefficient of variation would be zero.” Jennings (1909), p. 333. Jennings (1908b), pp. 521-4.

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terms. In the well-known symposium on pure lines and genotypes at the American Society of Naturalists meeting in December 1910—the year that he served as president of the society— Jennings declared of pure lines and genotypes that These things, whatever we call them, are concrete realities—realities as solid as the diverse existence of dogs, cats and horses. I find in many biologists not working in genetics an incorrigible bent for seeking under such a term as genotype something deeply hypothetical or metaphysical, and for characterizing it therefore boldly as purely imaginative. This is merely because such workers have not the things themselves before them.22

The material nature of these entities, Jennings explained in another article appearing during the same period, was readily apparent when protozoa were employed for hereditary study, since “unicellular organisms are essentially free germ cells,”23 which when perpetuated in isolation series, are in fact a pure line, or the material embodiment of a genotype. These entities, however, “become a little elusive, a little abstract,” in higher organisms that interbreed and therefore are not “pure.”24 In cases where selection had been shown to take place, he maintained it was because the investigator had not first isolated their material into pure races. In addition to the value that he attributed then to the protozoa as model organisms for the study of heredity and variation, Jennings increasingly championed the value of the pure line itself as a tool for hereditary analysis. The “absolutely permanent” nature of the pure line made it a “dissecting knife” that cut away obscurity and confusion, transcended the acrimonious debate, and focused the investigator of heredity on the still unanswered question: what are the sources of the minute hereditary differences upon which evolution operates? 25 The rhetorical positioning in this statement and in the paper as a whole is, I argue, critically important. This moment in 1910 is one of the few times prior to 1918 that Jennings presents himself as a “geneticist” engaged in an enterprise called “genetics.” Far more typically, as has been seen here, he referred to his research during this period as concerned with problems of “heredity.” In general, Jennings aimed to uncover the most fundamental mechanisms of heredity and advance a generalized conception of heredity—both of which he believed would surpass Mendelian inheritance or even the theory of the gene, in significance. In this particular context, however, before a large group of biologists and naturalists many of whom indeed were skeptical of the new genetics/heredity enterprise as a whole, Jennings presented himself as a “geneticist.” He did this not because of an investment in any particular theory or account of inheritance or any particular mechanism—because he largely did not adhere to one. Rather it was because of his considerable and growing investment in protozoa as model organisms for the study of heredity and the legitimacy that the scientism and precision of the pure line or genotype concept conferred on the protozoa. Jennings aimed as much, perhaps even more, for pure lines to serve protozoa as he did for protozoa to serve pure lines. Outside of rare moments like these, however, Jennings couched his experimental research program with protozoa during this period in terms of studies of heredity. He moved from his 22 23 24 25

Jennings (1911), p. 80. Jennings (1909), p. 322. Jennings (1910), p. 139. Ibid., pp. 137, 141.

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studies in animal behavior to heredity in 1907 not because of any specific interest in or commitment to Mendelism, but because of the many unanswered questions about heredity that were highlighted by the rediscovery of Mendel’s laws. Generally speaking, then, Jennings aimed to position his studies of protozoa within the epistemic space of heredity so as to emphasize the fundamentality and generality that he pursued and to distance himself from the more limited conceptions that were connoted by the term “genetics.”

Postscript Jennings reclaimed the mantle of “genetics” and engaged in much of the same type of rhetorical positioning of himself as a geneticist like that seen in the 1910 discussion of pure lines and genotypes when he engaged in his public critiques of eugenics throughout the 1920s. He clearly found it advantageous for the success of his critique to maximize his identity as an “insider” to the genetics enterprise.26 By the second half of the 1920s, Jennings was teaching a course entitled “Non-Mendelian Genetics,” which he viewed to be a term synonymous with “uniparental genetics,” and was the basis of a monograph that he published by the name of “the genetics of protozoa” in 1929. As I have suggested here, these terms would have seemed rather nonsensical to Jennings fifteen years earlier. At the outset of his course in 1927, he defined genetics as “the analysis of the production of the differential characteristics of organisms” and non-Mendelian genetics as the “lesser known portion” of genetics “which is not subject to the laws grouped under the name of Mendelism.” 27 Non-Mendelian genetics was essentially a replacement for Jennings’ earlier use of the term “heredity,” and he continued to maintain that the insights generated by the program would possess a generality not offered by Mendelian genetics. That Jennings adopted the term “genetics” in the late 1920s suggests, at least in the American context, the large extent to which Mendelism, the theory of the gene, and also eugenics had largely defined the terms of hereditary discourse. 28 Judy Johns Schloegel Independent Scholar, Illinois [email protected]

26 27 28

See Schloegel (2006), especially pp. 72-111. Sonneborn (1927-28), pp. 1-3. Schloegel (2006), especially pp. 127-39.

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References Allen, Garland E. 1978. Thomas Hunt Morgan. The Man and His Science. Princeton: Princeton University Press. Jennings, Herbert Spencer. 1908a. “Heredity, Variation, and Evolution in Protozoa. I. The Fate of New Structural Characters in Paramecium, in Connection with the Problem of the Inheritance of Acquired Characters in Unicellular Organisms.” The Journal of Experimental Zoology 5: 577-632. _____. 1908b. “Heredity, Variation and Evolution in Protozoa. II. Heredity and Variation of Size and Form in Paramecium, with Studies of Growth, Environmental Action and Selection.” Proceedings of the American Philosophical Society 47: 393-546. _____. 1909. “Heredity and Variation in the Simplest Organisms.” The American Naturalist 43: 321-37. _____. 1910. “Experimental Evidence on the Effective of Selection.” The American Naturalist 44: 136-45. _____. 1911. “Pure Lines in the Study of Genetics of Lower Organisms.” The American Naturalist 45: 79-89. _____. 1976 [1906]. The Behavior of Lower Organisms. Bloomington, Indiana: Indiana University Press. Kingsland, Sharon. 1987. “A Man Out of Place: Herbert Spencer Jennings at Johns Hopkins, 1906-1938.” American Zoologist 27: 807-817. Morgan, Thomas Hunt. 1903. Evolution and Adaptation. New York: Macmillan Co. Müller-Wille, Staffan and Hans-Jörg Rheinberger. 2005. “Introduction: A Cultural History of Heredity III: 19th and Early 20th Centuries.” Preprint 294. Max Planck Institute for the History of Science, Berlin. Rader, Karen A. 1998. “‘The Mouse People’: Murine Genetics Work at the Bussey Institution, 1909-1936.” Journal of the History of Biology 31: 327-354. Rheinberger, Hans-Jörg. 1997. Toward a History of Epistemic Things. Stanford: Stanford University Press. Ritter, William E. “Professor Jennings as a Biological Philosopher.” Science 35 (February 16, 1912): 267-269. Schloegel, Judith Johns. 2006. Intimate Biology: Herbert Spencer Jennings, Tracy Sonneborn, and the Career of American Protozoan Genetics. Ph.D. dissertation, Indiana University. Sonneborn, T. M. 1927-28. “Lecture Notes of ‘Genetics: Chiefly Non-Mendelian; A Course of Lectures, 1927-1928.” Bloomington, Indiana: Tracy M. Sonneborn Manuscripts, Lilly Library. Sonneborn, T. M. 1975. “Herbert Spencer Jennings, April 8, 1868-April 14, 1947.” Biographical Memoirs of the National Academy of Sciences 47: 143-223. Verworn, Max. 1899. General Physiology: An Outline of the Science of Life. Translated from the German edition and edited by Frederic S. Lee. New York: Macmillan and Co.

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Introduction On December 29th, 1910, scientists of the American Society of Naturalists met at a symposium with the title: “The Study of Pure Lines or Genotypes.” Just a year before, the Danish botanist Wilhelm Johannsen had published his book Elemente der exakten Erblichkeitslehre, where he had introduced the neologism “gene” and the differentiation between the concepts of a “genotype” and a “phenotype.”1 At this symposium in December 1910, Herbert Spencer Jennings, a zoologist at John Hopkins University, was determined to convince the audience that Johannsen’s genotypes were not merely hypothetical things or something “purely imaginative,” 2 as some critics had argued. Instead, Jennings wanted to persuade his colleagues of the reverse: that genotypes had a real existence and were “facts that strike you in the face.”3 He claimed that they were “concrete realities—realities as solid as the diverse existence of dogs, cats and horses.” 4 Jennings realism was highly influenced by the research object he had been working with for a couple of years: the unicellular organism Paramecium. At the same symposium, Jennings tried to illustrate what “pure lines”—or real genotypes—looked like in the case of this research object. In translating the world of Paramecium into the world of higher organisms, Jennings drew the following metaphorical picture: To get a clear grasp of the matter I believe that those not working with lower organisms will find it worth while to try to realize the condition which the investigator in this field (that is research on Paramecium, C.B.) has before him. A comparison may help. In lower organisms the genotype is actually isolated, each in a multitude of examples, which live along without admixture, visibly different from all others, for many generations, before again plunging into the melting pot of crossbreeding. In higher organisms we should have the same thing if every rabbit, every dog, every human being, multiplied by repeated division into two like itself, till there were whole counties inhabited by persons that were replicas of our absent president; cities made up of copies of our secretary, and states composed of duplications of the janitor I saw outside. Every human being, as it now stands, represent a different genotype (save perhaps in the case of identical twins), and these genotypes become inextricably interwoven at every generation. It is therefore easy to see how the genotype idea might appeal to workers among higher organisms as a mere hypothesis.5 What might have sounded slightly fanciful to Jennings audience is a common image in today’s debates on cloning. Representations of mass duplication of human beings are frequently used in 1 2 3 4 5

Johannsen 1913 [1909]. Jennings 1911. p. 80. Ibid, p. 80/81. Ibid, p. 81. Ibid, p. 81.

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the realm of popular media and science fiction to illustrate the horror of future cloning possibilities. In such images, cloning is understood as a kind of a serial mass production, as the production of an endless series of identical human beings out of one prototype. One could easily be tempted to think that these images of clones as the results of serial reproduction are very recent phenomena. However, what I want to argue today is that the concept of the clone as referring to serial (re)production or serial replication is something that was developed very early in the 20 th century life sciences. Later, I will return to Jenning’s quotation, with which I have introduced my discussion because it comes from precisely the historical period in which the terminology of the clone originated. In my paper, I will be dealing with one chapter of the very early history of the “clone” (as a scientific concept) and cloning (as a scientific practice). I will address the transition of the clone concept from its origins in horticultural and plant breeding into a laboratory research tool in the first two decades of the 20th century. With this transition, a specific aspect in the history of the concept emerges, namely the use of cell clones as technical objects, that is: as specific research tools for experimentation. My argument in this paper is the following: although the term “clone” was introduced within the context of horticultural breeding (where it designates a group of plants that were asexually propagated from a single ancestor plant) it soon became a concept used in the space of the laboratory. During the 1910s and 1920s, the term “clone” was introduced to refer to research objects which had standardized qualities because they were serially reproduced from one original or ancestral cellular unit. I will focus on early genetical work on protozoa, because it was in this field of protozoology that “pure lines”—then redefined as “clones”—of unicellular organisms began to be used as standardized research tools. Scientists such as Herbert S. Jennings or the German zoologist Victor Jollos started to apply the method of “pure line breeding” (developed by the botanist Wilhelm Johannsen in his well known work on beans) to the research on unicellular organism, especially on Paramecium. Their research on clones of Paramecium aimed at the general investigation of genetic constitution and heredity as well as questions about the inheritance of acquired characteristics. Thus, in the beginning of the 20th century, the scientific career of the “clone” concept started with the rise of the genetics of unicellular organisms, its demand for uniformity, and the quest for standardized research objects—clones that could be used as a kind of model organism. But before I go into detail, let me start with a few sentences on my general perspective: This paper is part of a broader project on the history of cloning, in which I regard the concept of the clone as a kind of “boundary concept” that went through very different scientific and non-scientific fields over the course of the 20th century. The main questions of the project are: what kinds of semantic shifts, what kinds of exchange and feedback between different spheres are related to these processes of circulation? The project addresses the relationship between material practices and concept formation within bioscientific research. Furthermore, I am interested in the impact of popular representations and cultural images that arose around the figure of “the clone." During the 20th century, the concept of the clone circulated among very different research fields (such as plant breeding, botany, cytology, tissue culture, genetic engineering and

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developmental biology as well as reproductive medicine.) As this occured, the epistemic status of the clone changed over time: Thus, in my perspective, there are three interrelated fields that are important for understanding the history of cloning: 1) the emergence of the clone as an epistemic and as a technical object within different experimental systems during the 20 th century 2) the clone as an interdisciplinary and inter-discursive element, and 3) the clone as a cultural phenomenon. In this paper, I will focus on questions concerning the first aspect: My main question today is, how the notion of the clone, which was first used to refer to some plant products, such as some special fruits, was introduced into the laboratory. Thus, the question is: How did the “clone” become an experimental object and concept? The paper has two parts: First, I will briefly follow the history of the clone concept and its introduction to the study of unicellular organisms. I will outline the discussion about the scope and limits of the concepts of “clones” and “pure lines” that began around 1910. The passages I am about to quote will show that the definition of what a “clone” was oscillated between two opposite poles: a presumed genetic identity, on the one hand, and on the other: the idea of genealogical origins. In the second part I will discuss one specific experimental approach, working with clones in more detail, namely that of the German protozoologist Victor Jollos. His research, which was very much stimulated by Jennings, led to the concept of “Dauermodifikationen” (persisting modifications). The existence of Dauermodifikationen was highly debated among German geneticists in the 1920s. Here, I will turn to questions of today’s workshop: the changing practices of early genetics and their impacts on the notion of heredity. In the last part I will discuss how Jollos concept of Dauermodifikationen stimulated the discussions of cytoplasmic inheritance in Germany during the 1920s.

“Clones” and “pure lines”: struggle for exact meanings Like the gene concept, the notion of the clone was first introduced to biology at the beginning of the 20th century. Whereas the “gene” referred to an abstract or even ideal unit (according to Wilhelm Johannsen’s use of the term), the notion of the clone referred from the very beginning to a concrete material object. It was Herbert J. Webber, a botanist from the Plant Breeding Laboratory of the US Department of Agriculture, who coined the term “clone“ in 1903—having searched for more than two years for a “suitable term to apply to those plants that are propagated vegetatively by buds, grafts, cuttings, suckers, runners, slips, bulbs, tubers, etc.” 6 Webber argued that “the plants grown from such vegetative parts are not individuals in the ordinary sense, but are simply transplanted parts of the same individual, and in heredity and in all biological and physiological sense, such plants are the same individual.“7 Webber then defined “clons” (sic) as “groups of plants that are propagated by the use of any form of vegetative parts (....) and which are simply parts of the same individual seedling.“8 6 7 8

Webber 1903, p. 502. Ibid, p. 502. Ibid. p. 502.

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This was the original definition of a “clone” in plant breeding at the beginning of the 20 th century. If you trace the history of the concept, you will soon find the following short articles, published in the journal Science in the years 1911 and 1912 (that is only one year after the above mentioned symposium of the American Society of Naturalists) with the titles: “Genotypes, Biotypes, Pure Lines and Clones” respectively: “Genotype and Pure Line.”9 The authors are Herbert Jennings and the botanist and plant breeder Geo H. Shull. Responding to each other, both scientists tried to clarify the semantics of the newly arising concepts in the emerging discoursive framework of genetics. Many historians have already emphasized the importance of “pure lines” for the emerging field of genetics. In his well known work, Johannsen had designated a group of beans, which were all descendants of the same orginal self-fertilizing plant, a “pure line.” He had observed that the variations in weights of all beans descending from beans that belonged to the same “pure line,” could always be described with the same variation curve. Besides playing an important role in the field of plant breeding (and its economics), “pure lines” or “pure cultures” were also central for the field of applied microbiology and the brewery industry from the last decades of the 19 th century onward.10 Pointing to the diverse uses of the term “pure line,” Jennings argued that an expansion of that definition, originally introduced by Johannsen, was inevitable. Moreover, he argued that a new term was needed, a term that included—in addition to the feature of a “genealogical series”—the issue of “genetic identity.” According to Jennings, “pure lines” designate a “genealogical series in which there arises no diversity in hereditary characteristics”11 and he gave a list of such cases of pure lines: Pure lines in this sense might be expected, from what we thus far have learned, (1) in cases of vegetative reproduction, (2) in at least some cases of parthenogenesis (where no reduction division occurs), (3) in case of self-fertilization of homozygotic organisms, (4) in case of inbreeding of a group of genotypically identical homozygotic organisms.12

Only the third group referred to Johannsen’s definition of a “pure line.” Therefore, Jennings concluded that “it appears that we badly need a term that will include ‘genotypically identical’ series of forms”13 arising in other cases than in Johannsen’s definition of the term. 9 10

11 12 13

Jennings 1911b; Shull 1912; Shull 1912b. For the analysis of Johannsen’s work see Roll-Hansen 1978; Roll-Hansen 2005; for the central importance of “pure lines,” and the notion of “purity” in early genetics and breeding research, see Bonneuil (this volume), and Müller-Wille (2007). Jennings 1911b, p. 841. Ibid. p. 841. Ibid. p. 842.

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Jennings’ redefinition and his search for a new term that included the aspect of “genotypically identical series of forms” did not remain unanswered: Only three weeks later, in January of 1912, the US botanist George H. Shull responded to Jennings with a short article on “Genotypes, Biotypes, Pure lines, and Clones.”14 Discussing the cases Jennings had mentioned, Shull recommended the use of Webber’s notion of “clone” or “clonal varieties” to refer to the “vegetatively reproduced potato and paramecium (...), in contradistinction to the self-fertilizing ’pure lines’ of beans.” 15 Furthermore, he suggested “the general adoption of the word ‘clone’ for all groups of individuals having identical genotypic character, and arising by asexual reproduction of any sort, including apogamy (i.e. so-called ‘parthenogenesis’ unaccompanied by a reduction division).” 16 Whereas the individuals of the “pure line” were necessarily homozygous, the individuals of a clone were not. This difference was one reason to argue for a new terminology. But the main goal was, similar to Jennings’, to find a term that would denote the “identical genotypic character” of a series of descendants arising by asexual reproduction (instead of self-fertilization). In his first note from January 1912, Shull was very enthusiastic about the important fact that clones could be regarded as having “identical genotypic characters.” Nevertheless, something—or somebody—must have unsettled the author’s view. Only three weeks later, Shull published a revision of his proposed definition. On February 2nd, 1912, we find another notice in Science, in which Shull explained: “Further consideration convinces me that this restriction (that is: the restriction of the term clone to ’groups of genotypically identical individuals,’ C.B.) is highly undesirable because it is impracticable.”17 The botanist now argued that, “it would be quite impossible to know for a certainty that two twigs used as cuttings or cions from the same tree had the same genotypic constitution.”18 Thus, he revoked his previous emphasis on genotypic identity and redefined the clone as “a group of individuals traceable through asexual reproductions (including parthenogenesis (...)) to a single ancestral zygote, or else perpetually asexual.” 19 “This definition,” he concluded, “puts the word ‘clone’ on exactly the same footing as the expression ‘pure line’, making it a purely genealogical term and involving no implication whatever as to the genotypic equality of the individuals included in the single clone.”20 From these quotations we can see two things: With the application of the method of “pure line breeding” to protozoa research, there arose a need for a new terminology. This led to the introduction of the concept of the “clone” into this field. But secondly, at the same time, there was a discussion about whether clones could be regarded as genetically stable objects. At this time, the “clone” was mainly a genealogical term. Statements about the genetic stability of clones could not be made with certainty; on the contrary the stability of the genotypic identity of the material was itself the subject of the research. Or as Jennings had emphasized in 1910, the questions were: Can 14 15 16 17 18 19 20

Shull 1912. Ibid. p. 27. Ibid. p. 28. Shull 1912b, p. 183. Ibid. p. 183. Ibid. p. 183. Ibid. p. 183.

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selection change genotypes? “Can environmental action permanently modify them?” 21 These questions resulted from the controversial debates about the evolutionary role of selection and variation. Once Johannsen had shown that selection had no effect on pure lines, he—like many other geneticists—concluded that mutations (or discontinous variations) were more important than selection in bringing about evolutionary changes. Darwinists, on the other hand, emphasized the primary power of selection and regarded evolution as a process based on continous hereditary variations. With this I now come to my second issue: the work of Victor Jollos, whose research on “variability and heredity in microorganisms”22 also started within this realm of evolutionary questions.

Victor Jollos’ work on Paramecium: the clone as a technical object Let me start with some biographical notes:23 Victor Jollos was one of the early coworkers of Max Hartmann, whose department at the Kaiser Wilhelm Institute for Biology emerged as one of the leading centers for genetic work on unicellular organisms in Germany in the 1920s. Born in Odessa (Ukraine) in 1887, Jollos spent his entire youth in Germany. After completing his Abitur he studied with Richard Hertwig in Munich and received his PhD in zoology in 1910. In 1912, Jollos became research assistant at the Institute for Infectious Diseases in Berlin (the “Robert Koch Institut”), where he worked in Max Hartmann’s Department for Protozoological Research. When Hartmann was appointed to the newly founded Kaiser Wilhelm Institute (KWI) for Biology in 1915, Jollos followed him there. From 1925 to 1929, he spent a few years as professor of zoology at the University in Cairo/Egypt. After that, he continued working in Hartmann’s group at the KWI in Berlin. With the rise of the Nazi Regime, Jollos was forced to leave Germany. In 1934, he emigrated with his family to the USA. Despite the support offered by colleagues such as Jennings and Tracy Sonneborn, Jollos never got an adequate position in the USA, possibly because of the differences in research styles between the German and the US scientific landscapes. 24 In 1941, Jollos died in Madison/Wisconsin. Jollos started his research on Paramecium in 1910 shortly after his PhD, when he was still in Munich, and he continued this research when he went to Berlin. Except for the years during World War I, when Jollos was engaged in medical work (he got an additional degree in medicine in 1918),25 the research on Paramecium kept him occupied until the early 1920s. Stimulated by Jennings’ approach, Jollos regarded research on Paramecium from the very beginning as a promising tool for experimentation around general questions concerning heredity. Even in his first papers on this subject Jollos emphasized that there was in principle no border

21 22 23 24 25

Jennings 1911, p. 81. See title Jollos 1914. For Jollos’ biography see Brink 1941. For this aspect see the discussion in: Dietrich 1996. See Brink 1941.

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between protozoa (or unicellular organisms) and higher organisms. Protozoa research, he argued, could offer valuable insights into general questions of heredity.26 The original material for Jollos’ work—the Paramecium populations—came from different lakes around Munich and Berlin (see Fig. 1). He isolated 9 different strains of Paramecium caudatum and 3 different strains of Paramecium aurelia from these populations. These strains showed differences in three categories: 1) their length, 2) their resistance to extreme temperatures, and 3) their resistance to arsenic acid. This material (as it is listed on the table) provided the basis for a complex system to produce "pure cultures" or "clones" of Paramecium. Until the early 1920s, Jollos developed a complex system consisting of a huge number of clonal lines derived from his original 12 strains of Paramecium.

Figure 1. Jollos research material: Different clones of Paramecium and their origins. Source: Hämmerling 1929, p. 5.

When Jollos started this work, he was still speaking of “individual lines” (“Individuallinien”), 27 explicitly in reference to Johannsen’s concept of a “pure line.” Yet, we also find the debate about the diverse semantics of a “pure line” mentioned in Jollos’ writings. The need for a term to better fit the conditions of asexual propagating objects also influenced Jollos’ work. Soon he began to speak of “clones” to designate his experimental material. In principle, each Paramecium that started to propagate by cell division could be the origin of a new clone, that is: of standardized objects for research. The crucial point of this system was the calibration of the environmental conditions. Jollos had to make sure that no conjugation or parthenogenesis occurred, and that the Paramecium propagated only by cell divisions. 26 27

See for example Jollos 1914, p. 33-34. Jollos 1913, p. 225, p. 227, p. 229

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The questions with which Jollos began his research were about the influence of environmental conditions on the genotype. He varied parameters such as temperature (or treatment with poison). He tried to change the clones in such a way that their reaction norm (“Reaktionsnorm”) was altered and that this changes became stable and inherited. To induce inherited changes, Jollos did the following: He exposed these different clones of Paramecium over a specific time period to ever increasing levels of arsenic acid; clones from those cultures of Paramecium that survived within the first level of arsenic were transferred into another, slightly higher level of arsenic medium and so on. Each step involved the production of new clonal lines for further experimentations. By this procedure, Jollos was able to generate clones that showed an increased resistance to arsenic. When clones of these lines were placed into an arsenic-free medium (and the procedure was started again) this acquired resistance persisted over periods of hundreds or more cell divisions, sometimes periods of longer than half a year. However, there was always a point in the end when all clones declined to the original level of arsenic sensibility. Especially after a conjugation (and an exchange of nuclear material) had occurred, there was a return to the original characteristics. The most important outcome of this research, was the concept of Dauermodifikationen (persisting modifications)—a notion which was introduced by Jollos to explain these surprising findings. As early as 1914, Jollos spoke of Dauermodifikationen as a third kind of variation that could neither be regarded as a mutation nor as mere modification. 28 In the following years, he devoted his whole energy to a clarification of this phenomenon. Jollos’s research on Dauermodifikationen was widely discussed. As Jan Sapp has shown, a lot of neo-Lamarckian scientists “viewed it as providing evidence for the inheritance of acquired characteristics.”29 However, Jollos himself did not share this opinion; he was convinced that the observed persisting modifications did not challenge the stability of the genotype. Nevertheless, Jollos’ research led to another important distinction that was part of another discussion in the 1920s, namely the issue of cytoplasmic inheritance: In 1921, as a result of his Paramecium work, Jollos distinguished two different kinds of transmission phenomena (“Übertragungserscheinungen”) and variations, which he related to two different cellular substances: 1) The transmission of hereditary factors (genes) and their variations, which are related to the structures of cell nuclei (chromosoms), and 2) the transmission of changes (“Übertragung von Veränderungen”) that are based on modifications (“Umstimmungen”) of the cytoplasm or other structures. Only variations belonging to the first group can be regarded as genotypic variations, or mutations, and only this kind of variation is a hereditary variation in the stricter sense. All variations of the second group are variations that belong to the category of modifications and Dauermodifikationen.30

During the 1920s, Jollos research was regarded as a major experimental contribution to the view that the cytoplasm played a significant role in the hereditary processes. In particular, Joachim Hämmerling, another coworker of Max Hartmann, interpreted Jollos’ results in the context of the 28 29

Jollos 1914, p. 20. Sapp 1987, p. 61.

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so-called “Kernplasmaproblem”—the discussion of nuclear-cytoplasmic relation in genetics. Hämmerling, who also popularized Jollos’ concept of Dauermodifikationen in a small book (with the same title) in 1929, argued along the lines of scientists like Carl Correns and Fritz von Wettstein. 31 For him, the existence of Dauermodifikationen showed that the cytoplasm had a strong impact on the gene action (“Genentfaltung”). Whereas Jollos himself spoke more cautiously of an active role of the cytoplasm in hereditary processes, Hämmerling speculated whether the cytoplasm could be regarded as an autonomous genetical constitutive element just as important—or maybe even more important—as the nucleus. I mention this last because we find here another interesting link to the history of cloning. Around that time, Hämmerling started his nuclear transplantation experiments with Acetabularia—which can be seen as some of the first successful nuclear transfer experiments. Let me conclude: One could certainly discuss the debate about Dauermodifikationen and cytoplasmic inheritance in more detail. However, my interest here has been to outline the transition of the notion of a “clone” (that was developed in the field of horticultural breeding) into a concept that was introduced to the space of laboratory practices. My argument has been that the scientific career of the clone concept (as referring to a standardized research material/research tool with all of its semantics of identiy and purity) started with the rise of the genetics of unicellular organisms. How far we can understand these developments in the broader context of the rise of model organisms at the beginning of the 20th century is an issue of further discussions. Christina Brandt Max-Planck-Institut für Wissenschaftsgeschichte, Berlin [email protected]

30

31

“An Stelle der alten Begriffsbestimmung der Vererbung als Übertragung von Anlagen auf die Nachkommen, einer Begriffsbestimmung, die die Mannigfaltigkeit der Übertragungserscheinungen nicht berücksichtigt, setzen wir also die Einteilung: 1. Übertragung von Erbanlagen (Genen) und deren Veränderung, die mit Kernstrukturen (Chromosomen) in Zusammenhang stehen und 2. Übertragung von Veränderungen, die auf Umstimmung des Plasmas oder bestimmter gesonderter Strukturen beruhen. Nur Abänderungen, die zur ersten Gruppe gehören, sind als genotypische, als Mutationen oder nach der in dieser Arbeit verwandten Ausdrucksweise als “im strengen Sinne” erbliche Abänderungen zu bezeichnen. Alle Umstimmungen der zweiten Art gehören zur Kategorie der Modifikation und Dauermodifikation.” Jollos 1921, p. 207. Hämmerling 1929, p. 59-65.

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References Brink, R.A. 1941. “Victor Jollos 1887-1941.” Science 94: 270-272. Dietrich, Michael R. 1996. “On the Mutability of Genes and Geneticists: The ‘Americanization’ of Richard Goldschmidt and Victor Jollos.” Perspectives on Science 4: 321-345. Hämmerling, Joachim. 1929. Dauermodifikationen. (Handbuch der Vererbungswissenschaft, Bd. I, hrsg. v. E. Baur u. M. Hartmann), Berlin. Jennings, Herbert Spencer. 1911. “Pure Lines in the Study of Genetics in Lower Organisms.” The American Naturalist 45: 79-89. Jennings, Herbert Spencer. 1911b. “‘Genotype’ and ‘Pure Line’.” Science 34 (No. 885): 841-842. Johannsen, Wilhelm. 1913 [1909]. Elemente der exakten Erblichkeitslehre. Mit Grundzügen der biologischen Variationsstatistik. Jena, 2. Aufl. Jollos, Victor. 1913. “Experimentelle Untersuchungen an Infusorien.” Biologisches Centralblatt 13: 222-236. ————. 1914. “Variabilität und Vererbung bei Mikroorganismen.” Zeitschrift für induktive Abstammungs- und Vererbungslehre 12: 14-35. ————. 1921. “Experimentelle Protistenstudien. I. Untersuchung über Variabilität und Vererbung bei Infusorien.” Archiv für Protistenkunde 43:1-223. ————. 1924. “Variabilität und Vererbung bei Protisten.” Zentralblatt für Bakteriologie u. Parasitenkunde, Heft 13: 22-37. Müller-Wille, Staffan. 2007. “Hybrids, pure cultures, and pure lines: From nineteenth-century biology to twentieth century genetics.” Studies in History and Philosophy of the Biological and Biomedical Sciences 38: 796-806. Roll-Hansen, Nils. 1978. “The Genotype Theory of Wilhelm Johannsen and its Relation to Plant Breeding and the Study of Evolution.” Centaurus 22: 201-235. ————. 2005. “Sources of Johannsen’s Genotype Theory.” In A cultural history of heredity III. 19th and early 20th century. Preprint Nr. 294. Berlin: Max Planck Institute for the History of Science. 43-52. Sapp, Jan. 1987. Beyond the Gene. Cytoplasmic Inheritance and the Struggle for Authority in Genetics. New York/ Oxford. Shull, George H. 1912. “‘Genotypes’, ‘Biotypes’, ‘Pure Lines’ and ‘Clones’.” Science 35 (No. 888): 27-29. ————. 1912b. “‘Phenotype’ and ‘Clone’.” Science 35 (No. 892): 182-183. Webber, Herbert J. 1903. “New Horticultural and Agricultural Terms.” Science 18 (No. 459): 501-503.

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Pedigree vs. Mendelism. Concepts of Heredity in Psychiatry before and after 1900 Bernd Gausemeier

Psychiatry is one of the most important contexts for the emergence of a science of heredity. In the late 19th and early 20th century, the inheritance of mental diseases was not only a major concern to the medical community, but also a highly disputed matter of public discourses. Nevertheless, our knowledge about psychiatric (and, in general, medical) concepts of “hereditary diseases” in this era is still quite fragmentary. This may be due to the fact that the pre-Mendelian use of the term “heredity” might appear vague and incoherent from a modern perspective. But if we want to understand the meaning of “heredity” in 19th century psychiatry, we have to apply other criteria than theoretical congruity: since we are dealing with a medical discourse, we have to consider the aetiological and nosological concepts in which the notion of “hereditary diseases” was embedded. Moreover, we have to look at the practices that were used to record and to analyze “hereditary” phenomena. In this paper, I will primarily focus on the latter aspect: the impact of statistical and genealogical techniques on the perception of hereditary diseases. On this basis, I will discuss the precarious state of Mendelism in early 20th century medicine. Mendelian theory, as historians of medicine are beginning to realize, was by no means enthusiastically accepted by physicians and psychiatrists.1 I want to argue that this resistance was not due to a lack of scientific understanding among medical researchers, but to the specific place of the concept “heredity” in medical thinking and practice.

Statistics Most 19th century psychiatrists were convinced that hereditary disposition was the major cause of mental diseases. Accordingly, it was a part of their professional routine to inquire about indications of madness or other abnormalities in a patient’s family. Ever since the emergence of modern psychiatric institutions in the early 19th century, mental asylums—especially in France —collected such information and converted them into statistical records stating the number of “hereditary burdened” cases among their inmates.2 These figures—the earliest form of quantitative data concerning human heredity—provided the major basis for 19th century discussions about the pathogenetic role of heredity. Their value, however, appeared questionable since the numbers published by different asylums diverged strikingly. The influential German alienist Maximilian Jacobi criticized in 1844 that most mental hospitals contented themselves with a cursory inquiry of the patient’s relatives.3 While Jacobi held that a more conscientious style of investigation would provide more reliable data, Wilhelm Griesinger, the founding father of German academic psychiatry, raised more basic objections concerning the uses of asylum statistics. Besides the ratio of cases allegedly based on a “hereditary disposition,” alienists 1 2 3

Rushton 1994, p. 144. Cartron 2007. Jacobi 1844, p. 598 ff.

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meticulously listed the number of patients who had suffered from alcoholism, physical diseases or professional setbacks, and often used these data to assess the aetiological importance of the respective phenomena. For Griesinger, this approach was both inadequate and deceptive, because it created an artificial separation between aetiological factors whose true meaning could only be understood in the context of a particular life history. It was the foremost task of the scientific psychiatrist to examine the whole genesis (Bildungsgeschichte) of the mainfest disease, by grasping “all those subtle threads at their beginnings which are at their ends interwoven into a web of delusion (Wahngespinst).”4 Griesinger had no doubt that heredity was usually the most important aspect in this complex of pathogenic factors. Family history, thus, was a crucial part of psychiatric anamnesis, but it had to be treated in conjunction with the patient’s individual biography, as its pre-history. This view was not unusual among 19th century psychiatrists. Even if the hereditary disposition was commonly regarded as the prime aetiological factor, it was not seen as a force that necessarily “produced” a certain disease, but rather as a potential that could be evoked and altered by certain environmental influences. Griesinger’s objections reflect the fundamental conflict between the clinical and the statistical meaning of “heredity”: while for the practitioner, it was one aspect in a complex psychopathological process, it became an isolated category in asylum records. The administrative practices of large clinical institutions, thus, created a space in which “heredity” became a visible entity.5 From an aetiological perspective, the presence or absence of “hereditary influences” was primarily a hint regarding the curability of the case. Though older convictions that “hereditary” diseases were necessarily incurable6 were quite out of fashion by the mid-19th century, many psychiatrists held that “hereditary” cases tended to develop into specific, severe forms of mental diseases. Bénédict Morel, the renowned degeneration theorist, regarded the “hereditary” diseases as one nosological class. Morel’s approach was developed further into the concept of hereditary diathesis, which was widely accepted in late 19th-century psychopathology, especially in France. 7 According to this idea, all mental and nervous diseases emanated from the same kind of hereditary disposition. Its manifestation was believed to aggravate in the course of generations—severe forms were regarded as the result of continued neuropathic degeneration in a family, while all kinds of excentric or aberrant behaviour could be interpreted as its primary forms. This perception accorded with the most simple forms of asylum statistics, in which any information about insane, criminal or just “suspicious” relatives similarly generated an entry into the column for “hereditary” cases. In the second half of the 19th century, however, some statistical surveys issued by psychiatric institutions went beyond giving the mere percentage of “predisposed” patients. Since asylums collected personal data like age, religion or profession from their new entries, it was possible to establish the most diverse correlations between these items and “hereditary” disposition, or to calculate if “hereditary burdened” patients were more likely to recover or to relapse.8 More sophisticated reports differentiated the notion of heredity according 4 5 6 7

Griesinger 1867, p. 132 f. This development may partly explain why—as López-Beltrán (1992, 36 f) states—the noun “heredity” began to be used as a general concept in the French medical community during the 1830s. Waller 2007. Dowbiggin 1991, p. 119 ff.

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to degree of kinship—“direct” heredity (i.e. observed diseases in the parental generation), “indirect” respectively “atavistic” heredity (grandparents) or “collateral” heredity (siblings).9 This specific style of categorization is characteristic for 19th century medical concepts of heredity: the term “heredity” primarily referred to a pathogenic force appearing in different “grades,” not to the constant transmission of a certain trait. Psychiatrists, thus, had no lack of statistical data referring to the factor “heredity.” Those inclined to laborious statistical work rather ended up producing hundreds of tables which yielded no results of scientific value. Asylum directors who realized this unsatisfactory situation nevertheless blamed it on the insufficient state of psychiatric record-keeping. A standardization of statistical methods in all major psychiatric institutions, they hoped, would generate a basis for truly scientific studies.10 In the 1860s and 70s, the reform of asylum statistics became a vividly discussed topic among German alienists. Following the 1867 International Congress for Psychiatry in Paris, there had been international efforts to develop standard schemes for asylum statistics.11 After these plans had been thwarted by the 1870/71 war, the Association of German Alienists authorized a commission to conceive census forms and statistical tables for the common use in all German asylums. Similar discussions took place in Switzerland, where a national standard scheme was adopted in 1872.12 Psychiatrists engaged in these discussions were aware that a kind of statistics generating more precise data about aetiological factors like heredity required more specific categories, more detailed information and, consequently, more work. Characteristically, the German Alienist’s commission declared that it was preferable to restrict the new schemes to data whose “scientific value was recognized from all sides” and to simplify statistical procedures by “leaving out the administrative and aetiological considerations (with the exception of heredity).”13 Apparently, it went without saying that “heredity” was a most crucial aetiological aspect that could easily be determined, while the “scientific value” of almost any other kind of information turned out to be controversial—especially the question of nosological categories. In the 1870s, there was at best a rudimentary system for the classification of mental diseases. Different psychiatric schools followed different nosological concepts. As the asylum director Friedrich Wilhelm Hagen put it, psychiatrists were no “typesetting machines” who were able to sort aetiological facts into predefined categories, but human beings dealing with individual life histories. 14 He feared that statistical standardization would only result in time-consuming work that produced nothing but an ever-increasing accumulation of numbers and tables.15 Once again, Hagen’s position exemplifies the tension between the administrative practice of record-production and the customs of anamnesis: while statistical surveys demanded well-defined and fixed categories, the individual cases often ruled out such clear-cut evaluations.

8 9 10 11 12 13 14 15

Tigges 1867. Hagen 1876, p. 208 f. Tigges 1867, p. 119. Anon. 1873a, p. 459-460. Anon., 1873a; Anon., 1873b. Nasse 1873/74, p. 241. Hagen 1871, p. 278. Ibid., p. 269.

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Yet nosological specifications were necessary if alienists wanted to understand the distribution of certain forms of mental diseases and the respective role of heredity in their aetiology. In 1874, the Association of Alienist’s commission issued standard forms which distinguished seven forms of mental disease.16 Further, they asked if cases of mental disease, nervous disease, alcoholism, suicide, “remarkable character“ and criminal acts were noted in parents, siblings, grandparents, uncles and aunts. In contrast to prior forms of asylum records, the Association of Alienist’s scheme allowed to determine the percentage of “hereditary” cases for each nosological category and to specify the nature of “familial disposition.” Yet its application only aggravated the inherent problems of asylum statistics: due to the large array of categories, the absolute figures often became too low to be of statistical significance. The first survey using the scheme abstained from specifying the ancestors’ diseases because this practice would have fragmented the material even more.17 Characteristically, the same survey noticed only one regularity of heredity—the “law” that psychopathological phenomena were most likely to be transmitted from mother to son and from father to daughter. The structure of asylum records favored speculations about specific maternal and paternal “influences” on the offspring, since they always listed male and female patients separately. Apart from such traditional beliefs, even the more sophisticated statistical practices produced nothing that could be regarded as new, commonly accepted knowledge about heredity. Despite the fact that it increasingly moved to the center of psychiatric studies, heredity was not yet a scientific object in its own right. It was not because the prevailing methods of asylum statistics were essentially about establishing correlations. They allowed to relate “heredity” to other phenomena (e.g. sex, age, nosological categories) but not to analyze the ways of hereditary transmission—a problem that only began to bother psychiatrists by the time when a science of heredity emerged.

Genealogy By the 1890s, more and more voices in the German psyciatric community rejected the statistical approach altogether and called for a turn to genealogical methods. The Jena psychiatrist Otto Binswanger, for example, categorically stated that “the questions raised by the recent works about heredity will, in clinical research, only be solved through the accurate study of individual family trees but not through mass statistics.”18 The phrase “recent works about heredity” referred, of course, to August Weismann’s theory of the continuity of germ plasm and the debates it had sparked off. This new biological discourse about heredity did not only exert a strong influence on the medical community because medical scientists had to take sides in the struggle over the possibility or impossibility of an inheritance of acquired characters. It evoked a new awareness that the notion of heredity, as it was traditionally used in medicine, was rather a means of clinical classification than a biological concept. As the psychiatrist Robert Sommer clarified, one had to distinguish between the mere observation of certain pathological traits within a family history 16 17 18

Anon. 1874. The scheme distinguished melancholia, mania, secondary psychic disorder (“Sekundäre Seelenstörung”), paralytic psychic disorder, imbecility, “delirium potatorum”. Hagen 1876, p. 207 f. O. Binswanger, preface to Rohde 1895, p. IX.

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(“Heredität”), and the positive proof that such a case was in fact based on biological inheritance (“Vererbung”).19 Psychiatrists became aware that their old use of the term “heredity” referred to an analogy between phenomena in different generations, while the scientific problem was to establish a causal relation between these phenomena. In other words, it was only at the end of the 19th century that heredity became a scientific object in medicine, i.e. that medical scientists began to ask how a certain disease was transmitted and what about it was transmitted. The means to investigate this was the pedigree. Doubtlessly, psychiatrists had studied family histories long before the 1890s. Ideas about the “hereditary transformation” of diseases or the progressive degeneration of families clearly rested upon observations on several succesive generations. Richard Krafft-Ebing’s statement that his knowledge about the heritability of nervous diseases was based “on the exhaustive study of the pedigrees of many hundreds of sick persons.”20 indicates that genealogy provided a form of tacit knowledge about heredity. The common form to represent and to analyze heredity in 19th century psychiatric journals and books, however, were statistical tables. Large-scale genealogical studies on the inheritance of mental diseases were first initiated by the the influential Swiss alienist August Forel. In 1895, his assistant Jenny Koller published a study that was not only based on family histories of patients from Forel’s Burghölzli asylum, but also on genealogical material collected among other social groups. 21 The comparison of the occurence of mental diseases in the pedigrees of “healthy” and “mentally ill” people showed that also the former were often “hereditary burdened,” though to a lesser extent than asylum patients. This approach was, effectively, an extended version of old-style asylum statistics. It responded to critics who objected that inquiries only based on patient records necessarily exaggerated the role of the “hereditary burden,” because they comprised no comparative material.22 The inclusion of “normal” genealogies indicated that traces of “hereditary” madness were not confined to certain degenerated families, but omnipresent. This aspect, however, was not the main reason why genealogical methods became an increasingly popular matter of discussion in German medicine after 1900. Many physicians and psychiatrists were influenced by Ottokar Lorenz’s 1898 handbook of genealogy. 23 Lorenz, an historian, did not only define genealogy as a borderline method linking the historical and the biological sciences, he also forcefully argued that scientists would only be able to understand the problem of heredity by applying proper genealogical methods. As stated before, psychiatrists were open to this kind of advice, because it appeared as a counterdraft to the practices of asylum statistics. While asylums and clinics collected masses of family data based on oral information that was, in addition, mostly restricted to two generations, individual family studies opened up a “vertical,” more specific perspective. Wilhelm Strohmayer, Lorenz’ most ardent follower in psychiatry, stressed that medical genealogy was about going “more into the deep than into the breadth” since “a few, but thorough investigations about heredity are more useful than countless

19 20 21 22 23

Sommer 1901, p. 67. Krafft-Ebing 1869, p. 443 f. Koller 1895. Grassmann 1896, p. 1018. Lorenz 1898.

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inaccurate ones.”24 Extensive pedigrees including reliable psychopathological data would allow to understand the laws of transmission. Most adherents of medical genealogy, however, believed the scientific study of human heredity could only be advanced through larger samples of adequate genealogical material. Robert Sommer, the most influential advocate of medical genealogy, and other psychiatrists called for the establishment of regional or national centers compiling pedigrees assembled in psychiatric asylums and clinics.25 The idea of such large-scale projects raised the question of methodical standardization. Medical genealogists devotedly argued about the best way to compile data, the number of generations to be included, and the most concise form to depict family relations. A crucial question was in how far family histories provided data satisfying the needs of a medical investigation. The idea of following a certain trait over a longer succession of generations was adapted to the history of aristocratic dynasties—most notably the much-discussed example of the facial features running in the Habsburg family.26 Beyond that, it was only applicable to cases of rare and specific diseases in families with an unusually good genealogical tradition. The most remarkable study of this kind was published in 1913, after more than a decade of fieldwork, by the Swedish neurologist and eugenicist Herman Lundborg, who traced a particular epileptic illness over 6-7 generations.27 His claim that the disease could be attributed to a Mendelian recessive gene rested on a comparatively reasonable line of argument. However, Lundborg also listed other features of the ramified kinship he had studied: alcoholism, madness, violence, aberrant behaviour, crime—in a word, the expressions of proceeding degeneration. Lundborgs work represents two basic tendencies of medical genealogy: on the one hand, he claimed to provide an “exact” analysis of a hereditary disease, on the other, he compiled diverse biographical particulars which were unreservedly taken to be related by means of heredity. Further, its paper-wasting reproduction of 50 large pedigrees strikingly documented the practical intricacies of the genealogical approach. Most practitioners conducting medico-genealogical studies were not faced with this problem, but rather with families hardly providing reliable biographical information beyond the grandparental generation. They were aware that the genealogical structure of most families could at best be reconstructed up to the great-grandparental generation. 28 A controversial topic among medical genealogists was the question if studies in human heredity had to consider only the direct ancestry of a proband or also the collateral lineages, i.e. aunts/uncles and grandaunts/granduncles. Following the latter option, the ophthalmologist Arthur Crzellitzer conceived a “kinship chart” that arranged the ancestors around the living proband in the center. 29 Printed on a squarish sheet of paper, Crzellitzer’s form provided a solution for an urgent practical problem of medical genealogy: it allowed to collect a large number of family studies concerning a certain trait in a reasonably concise way—e.g. in a folder or a flip box.

24 25 26 27 28 29

Strohmayer 1908, p. 483. Sommer 1913, p. 394 f. E.g. Lorenz 1898, p. 402-408; Strohmayer 1911; Haecker 1911. Lundborg 1913. Crzellitzer 1908, p. 575; Jolly 1913, p. 382. Crzellitzer 1908.

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Crzellitzer’s charts achieved some attention, but not a wide distribution. In the years between 1900 and World War I, large parts of the German medical community were concerned with practical questions of genealogical research, but there never emerged a large, concerted project like the Eugenics Record Office in the USA. The idea of such a central institution was propagated by a group centered around the Zentralstelle für deutsche Personen- und Familiengeschichte in Leipzig, an association of genealogists closely in touch with psychiatrists and eugenicists. 30 It was based on the belief that a treasury of well-reconstructed pedigrees, compiled with the help of hobby genealogists and physicians, would generate valuable material both for historical and for medical purposes. This comprehensive outlook explains much of the popularity of genealogy in the medical community, but it also contributed to its eventual failure. Historical and medical family research, after all, did not only have different objectives, but also required different techniques of data-arrangement.

Mendelian Statistics The flaws of the medico-genealogical discourse were obstinately pinpointed by the practising physician and medical statistician Wilhelm Weinberg. The difficulties to obtain reliable medical data about past generation, as he first stated in 1903, did not only make family studies problematic, but utterly useless for the study of disease inheritance. 31 While most medical genealogists supposed that good genealogical material would inevitably generate insights into the inheritance of certain traits, Weinberg held that studies in human heredity were only possible if there was a sharply defined phenomenon to be investigated and a sufficient supply of reliable data about it. Weinberg was by no means a dispraiser of genealogy as such: he was experienced in the use of family records for studying diseases like cancer and tuberculosis. But he argued that if medical scientists wanted to understand the regularities of heredity, they had to disengage from singular case studies and to process genealogical material according to rigid statistical methods. This was also necessary, according to Weinberg, „to eliminate the influence of social factors.” Since families usually formed a constant „environment,” genealogical case studies tended to blur heredity and tradition.32 After he became acquainted with the Mendelian theory by the mid-1900s, Weinberg began to develop a method that stood out both against pedigree studies and older forms of medical statistics. Many adherents of medical genealogy accepted Mendelism, most notably the group of American eugenicists associated with the Eugenics Record Office (ERO). The ERO approach was based on the collection of pedigrees (usually comprising three generations) containing a certain trait. From its average distribution in the respective generations, it was deduced whether the trait was transmitted according to the recessive or dominant Mendelian mode. 33 Weinberg’s approach was clearly different: he understood that doing Mendelian genetics was not about investigating pedigrees, but about constructing generations. Between 1908 and 1914, Weinberg published a large 30 31 32 33

Breymann 1912. Weinberg 1903. Weinberg 1908, p. 378. Cannon/Rosanoff 1911; Rosanoff/Orr 1911.

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number of contributions to a statistical theory of Mendelian human genetics which place him, in retrospect, among the founders of population genetics.34 His ideas were by no means ignored, but their practical implications were barely realized by medical scientists. A significant exception was the psychiatrist and eugenicist Ernst Rüdin, who adopted Weinberg’s views on medical genealogy and statistics in the early 1910s.35 In the following years, he developed a project aiming at the proof that the main forms of nervous diseases were transmitted as Mendelian traits, drawing constantly on Weinberg’s statistical concepts and his personal advice.36 In 1916, Rüdin published his study on the inheritance of Dementia praecox (Schizophrenia). For this purpose, he had compiled a large number of schizophrenia patients in clinics and asylums. The next step was to detect all the siblings (who potentially comprised additional Dementia praecox-cases) and the parents of the patients. All these relatives were, as far as possible, examined by a medical expert. In this way, Rüdin was able to construct a Mendelian ‘F1’- and a ‘P’generation, with secure medical data on each individual. The Rüdin/Weinberg method was based on the hypothesis that the disease in question emanated from a recessive factor. If this was assumed, all the parents had to be regarded as heterozygote bearers of the factor, except for those families where one parent also displayed the disease. These cases were separately treated as heterozygote/homozygote crossings. If the “recessive” hypothesis was correct, the ratio of ill people among the sibling series born from two ‘normal’ parents had to be 25%, respectively 50% in the group born from one sick parent. The statistical proof was, of course, a little more complex. In 1912, Weinberg had pointed out that it was impossible to achieve correct Mendelian ratios with a sample only comprising manifest bearers of an assumedly recessive trait. Since the records necessarily missed all the double-heterozygote couples that produced no manifestly ill offspring at all, both the parental and the filial generation were incomplete and the ratios were distorted. Weinberg developed mathematical tools to eliminate this source of error. 37 It is especially this methodical contribution that shows what distinguished Weinberg from the practices of medical genealogy—not only his statistical skills, but primarily his awareness that Mendelism was about calculating with the unseen. In theory, the Weinberg-Rüdin method was doubtlessly the first genuinely Mendelian approach to human heredity. But apart from accurate statistics, a Mendelian approach requires a clearly defined trait. If such traits exist at all in humans—striking and rare features like polydactily or haemophilia had been the first viable objects of Mendelian studies—this was surely not the case in the realm of psychiatry. Rüdin’s object of study, Dementia praecox, was not only a complex disease, but a highly contested nosological concept. As stated above, the classification of mental diseases had been a major problem in all attempts to standardize psychiatric statistics in the late 19th century. In the 1890s, a large part of psychiatrists began to accept Emil Kraepelin’s sophisticated nosological system as the new gold standard for clinical classification. Dementia praecox was a central category in Kraepelin’s system, a nosological “unit” that was characterized by a specific aetiology and psychopathology. Nevertheless, influential colleagues disagreed over this point: other concepts of schizophrenia included a much wider scope of symptoms and morbid 34 35 36 37

On Weinberg’s biography cf. Früh 1996. Rüdin 1911. Weber 1993, 109 f. Weinberg 1912, 166 ff.

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phenomena than Kraepelin’s definition.38 The question at stake here was if nervous and mental diseases could be divided into clearly distinct pathological “units“ or if they formed a “family” of interrelated clinical phenomena. When Rüdin intended to prove that Dementia praecox could be attributed to a Mendelian factor, thus, he also ventured to demonstrate that Kraepelin’s definition and his entire systematic approach were correct. Yet Rüdin’s description of his work shows how difficult it was to treat Dementia praecox as a constant “trait.” According to Kraepelin, the disease became manifest up to the age of 40, so siblings who died before that age could not clearly be counted as “ill” or “normal” cases. Further, the manifestation was considerably affected by environmental influences. Finally, the whole psychophysical personality of the patient affected the manifestation of the trait. 39 It largely depended on triggering factors—traumata, alcoholism or extreme physiological situations like childbirth—if and when it appeared. In short, Rüdin had to include the whole genesis of the disease into his calculations. But he mainly considered the other exogenous and endogenous factors because they “obscured” or distorted the clear picture of Mendelian inheritance. Heredity was not the only pathogenic cause, but the only one that counted. Nevertheless, his reductionist efforts were not crowned with success. After all calculations and corrections, his sibling series only showed less than 5% instead of the expected 25% of Dementia patients. He resorted to the alternative explanation that the disease was caused by two recessive factors, but measured by his own methodical claims he had failed to establish a clear Mendelian scheme. The same was the case for his subsequent studies on the transmission of other Kraepelinean “disease units.“ When he was about to become Nazi Germany’s most influential eugenicist in 1933, he frankly admitted that for none of the major mental and nervous diseases a Mendelian mode of inheritance had been unquestionably established. 40 The most sophisticated approach of Mendelian human genetics in the first half of the 20th century clearly demonstrated that it was not possible to “mendelize” complex mental diseases. This failure notwithstanding, Rüdin’s reductionism offered a new perspective for the psychiatric concept of heredity. The focus on a nosological “unit” like Dementia praecox was a turn against traditional concepts of “hereditary transformation” or “neuropathic disposition,” that is against the practice of counting diverse pathological features as appearances of the same hereditary tendency. Rüdin did not categorically rule out that there might be hereditary dispositions manifesting themselves in various clinical phenomena rather than specific Mendelian factors causing certain pathological “traits.” But he argued that his approach was the only way to test both options.41 There is no doubt, however, that he was convinced of the fundamental correctness of his Mendelian hypothesis. Being an ardent eugenicist, he held that a psychopathology based on Mendelian principles—in place of the old cloudy notion of “hereditary disposition”—would allow to identify a risk of hereditary ill progeny with scientific certainty. 42 Further, Rüdin’s approach profoundly changed the scope of genealogical methods. Rüdin’s co-workers directly interrogated patients and their relatives, but they also collected material from 38 39 40 41 42

Roelcke 1996. Rüdin 1911, p. 547. Rüdin 1934, p. 134. Rüdin 1916, p. 139 ff. Rüdin 1911, p. 495.

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registry offices, church registers, hospital files, records of police, military and legal authorities, and finally family histories. The fruit of this genealogical field work was processed in the form of index cards.43 This method was geared to compute data relevant for a certain problem (like the inheritance of Dementia), but it also allowed to recombine the material for different projects. Pedigrees were only internally used to visualize family relations. Incited by the structuralist idea of Mendelism, Rüdin had transformed medical genealogy into a database technique.

Non-Mendelian genealogy Rüdin’s Mendelian research program was exceptional in the medical realm. Many other medical researchers in Germany accepted Mendelism as an interesting theory which provided, for example, a nice way to explain why hereditary diseases „skipped” generations.44 But they were generally unable to integrate Mendelism into their aetiological and nosological thinking, most notably the concept of the unit-character. Psychiatrist and occasional genealogist Robert Sommer offers a characteristic example for this tendency. Though he did not explicitly reject Mendelism, it was incompatible with his medicogenealogical approach. When he argued for area-wide family research projects, it was because he hoped for insights into the “familial relations of the mental diseases and their distribution in the whole country.”45 In this view, the question was not how certain diseases were transmitted but how they were related. Pathological characters were not at all transmitted in a constant form, they were transformed. In the same spirit, the asylum director Hans Roemer stated that the crucial question about pathological heredity was “according to which rules the slight and the severe alterations of psychic health are interlinked.”46 Understanding heredity was a way to understand the nosological relations between diseases, not their respective modes of transmission. In this perspective, the enthusiasm for pedigrees so vigorously attacked by Weinberg made perfect sense: “familial relations” between diseases and anomalies were best studied by using extensive genealogies of “interesting” families. Sommer’s perspective was even broader: for him, the essential question was how mental diseases and “normal” mental qualities were related. The „normal personality” already alluded to its psychopathological potential—not only in individuals, but also in whole families. While Rüdins approach was based on the clear definition of a disease as a distinct feature, Sommer’s idea of family research was about “finding the family type in its various manifestations.”47 Rüdin collected and arranged specific data about populations, Sommer envisaged psycho-pathological family histories. Mendelism, thus, was incompatible with a widespread medical concept of heredity, but the problematic relation between Mendelism and medicine was scarcely discussed in an explicit way. A remarkable exception in this respect was the Rostock pathologist Friedrich Martius. Like Sommer, Martius accepted Mendelism in principle, but he disputed its applicability to the human 43 44 45 46 47

Rüdin 1916, p. 25 ff. E.g. Kraepelin 1909, p. 177. Sommer 1913, p. 394. Römer 1912, p. 293. Sommer 1907, p. 108.

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domain. His attack on the poor quality of pedigree studies claiming the “Mendelian” transmission of ill-defined characters like “musicality” would have met approval from Weinberg, but his critique went beyond methodical flaws: The now fashionable attempts to adjust pedigree material—which is so abundantly at hand in the medical literature—to the Mendelian numerical proportions suggest a congruence which does not exist so far and is little likely to exist. For the human material is by its nature contrary to the application of the experimental method working with pure lines.48

Weinberg or Rüdin would have objected that even if “pure lines” in the sense of Wilhelm Johannsen did not exist in human populations, it was possible to construct statistical purity with respect to a certain trait. But for Martius, diseases were not traits at all,49 and statistical purity was not an indisputable value in clinical medicine. Argueing from the viewpoint of the practitioner, he consequently asked for the benefits of Mendelism for clinical practice. Martius pinpointed the implicit motivation for many adherents of Mendelism in medicine, namely the claim that if it was once established how a disease was transmitted, it would be possible to predict it with mathematical precision and—if the state once adopted eugenic principles—to eliminate it. Even if it was proven that certain diseases behaved like dominant or recessive factors, Martius asked, what was won? It was still impossible to predict the offspring’s state of health with complete certainty, and accordingly the exact knowledge of the Mendelian scientist did not lead any further than the experience of the pre-Mendelian physician. 50 For the practicing physician, he claimed, it was sufficient and much more valuable to know the principles of “scientific genealogy.” While the Mendelian view only created an illusion of certainty, accurate family research exhibited the facts of heredity “as they really exist in human beings.”51 This was not a naive statement of an unimaginative traditionalist. Martius realized that Mendelism introduced the reckoning with virtual realities into medicine. He sensed that with the interest for the invisible logic of the genotype, the focus shifted away from the actual object of the medical study, the patient and the disease. In so far, Martius formulated a sharp-sighted criticism of eugenic aspirations, though he was himself an old-school hereditarianist and eugenicist. While Martius’ vision of eugenics was a “reasonable” form of premarital “counselling“ based on accurate family studies, Rüdin’s approach implied the control of populations.

Conclusion The two concepts of genealogical methods discussed here—represented by Sommer and Martius on the one hand and by Weinberg and Rüdin on the other—do not only correspond to diverging ideas about heredity, but also to essentially different concepts of disease. While the “traditional” medical concept of heredity attributed plasticity and variability to hereditary diseases, the Mendelian view implied their stability and specificity. And exactly this view was hardly compatible 48 49 50 51

Martius 1913, p. 222. Ibid., p. 190. Ibid., p. 186. Ibid., p. 188.

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with the experiences of the clinical practitioner. A physician or psychiatrist investigating a patient and his/her family history was not confronted with “traits,” but with dynamic and variable phenomena. Further, he had little need to know how exactly a certain disease was transmitted. Nevertheless, Mendelism entered medicine mainly due to the efforts of eugenicists. But it was much more successful as an idea than as a formative method. The Weinberg/Rüdin approach was clearly exceptional in this regard. While it strikingly demonstrated that it was inadequate to conceive complex diseases as simple Mendelian traits, it still marked a break with the medical fascination for genealogy. Wilhelm Weinberg’s attack against medical genealogy did not only rest on the insight that pedigrees were an inappropriate tool for a Mendelian approach, but also on the awareness that genealogical methods imprinted their own logic on the concept of heredity. In a way, his arguments preceded Wilhelm Johannsen’s much more explicit 1911 diatribe against the idea of “ancestral influence.”52 Johannsen’s insistent statement that the „modern view of heredity” was not at all about the transmission of ancestor’s “qualities,” but about something strictly nonpersonal and non-physical—the Mendelian factors—was most likely inspired by his knowledge of contemporary medical genealogy. Both for Weinberg and Johannsen, the concept of heredity had to be cleared from all remnants of genealogical thinking before it could become scientific. This claim, however, made little impact on the medicine of their time. But even after a “century of the gene” and the rise of molecular techniques, genealogical practices are still a significant part of contemporary human genetics and genomics. Does this mean that we still adhere to genealogical rather than to a “modern” view of heredity? At least it shows that the science of human genetics, after all, deals with humans and human relations, not only with genes. If we talk about heredity, we still touch questions about descent, ancestry, and personal identity. Bernd Gausemeier Max-Planck-Institut für Wissenschaftsgeschichte, Berlin [email protected]

52

Johannsen 1911, p. 130.

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References Anon. 1873a. “Bericht über die Sitzung des Vereins der dt. Irrenärzte zu Leipzig 13.8.1872.” Allgemeine Zeitschrift für Psychiatrie 29: 459-466. Anon., 1873b. “Versammlung der schweizerischen Irrenärzte am 25. und 26.9.1872 in Burghölzli.” Allgemeine Zeitschrift für Psychiatrie 29: 579-584. Anon. 1874. “Zählkarten und Tabellen für die Statistik der Irrenanstalten. Aufgestellt von dem Verein der deutschen Irrenärzte.” Suppl. Nr. 6 to Allgemeine Zeitschrift für Psychiatrie 30. Breymann, Hans. 1912. “Über die Notwendigkeit eines Zusammengehens von Genealogen und Medizinern in der Familienforschung.” Archiv für Rassen- und Gesellschaftsbiologie 9: 18-29. Cannon, Gertrude L., and A. J. Rosanoff. 1911. “Preliminary Report of a study of heredity in insanity in the light of the mendelian laws.” Eugenics Record Office Bulletin Nr. 3. Cartron, Laure. 2007. “Pathological Heredity as a bid for greater recognition of medical authority in France, 1800-1830.” In Müller-Wille and Rheinberger. 2007. Crzellitzer, Arthur. 1908. “Sippschaftstafeln, ein neues Hilfsmittel zur Erblichkeitsforschung.” Medizinische Reform 16: 573-578, 604-605, 624-629. Dowbiggin, Ian R. 1991. Inheriting Madness. Professionalization and Psychiatric Knowledge in NineteenthCentury France. Berkeley. Früh, Dorothee. 1996. “Wilhelm Weinberg (1862-1937), Armenarzt und Populationsgenetiker. Anmerkungen zu Leben und Werk.” Biologisches Zentralblatt 115: 112-119. Grassmann, Karl. 1895/96. “Kritischer Ueberblick über die gegenwärtige Lehre von der Erblichkeit der Psychosen.” Allgemeine Zeitschrift für Psychiatrie 52: 960-1022. Griesinger, Wilhelm. 1867. Die Pathologie und Therapie der psychischen Krankheiten für Aerzte und Studirende. Stuttgart. Haecker, Valentin. 1911. “Der Familientypus der Habsburger.” Zeitschrift für induktive Abstammungs- und Vererbungslehre 6: 61-89. Hagen, Friedrich Wilhelm. 1871. “Ueber Statistik der Irrenanstalten.” Allgemeine Zeitschrift für Psychiatrie 27: 266-294. ————. 1876. Statistische Untersuchungen über Geisteskrankheiten. Nach den Ergebnissen der ersten 25 Jahre der Kreis Irrenanstalt zu Erlangen unter Mitwirkung von deren Hülfsärzten. Erlangen. Jacobi, Maximilian. 1844. Die Hauptformen der Seelenstörungen in ihren Beziehungen zur Heilkunde. Leipzig. Johannsen, Wilhelm. 1911. “The Geneotype Conception of Heredity.” In The American Naturalist 45:129159. Jolly, Philipp. 1913. “Die Heredität der Psychosen.” Archiv für Psychiatrie und Nervenkrankheiten 52: 377436, 492-715. Koller, Jenny. 1895. “Beitrag zur Erblichkeitsstatistik der Geisteskranken im Kanton Zürich.” Archiv für Psychiatrie und Nervenkrankheiten 27: 268-294. Kraepelin, Emil. 1909 (-1915). Psychiatrie. Ein Lehrbuch für Studierende und Ärzte. Leipzig. Krafft-Ebing, Richard von. 1869. “Ueber die prognostische Bedeutung der erblichen Anlage zum Irresein.” Allgemeine Zeitschrift für Psychiatrie 26: 438-456. López-Beltrán, Carlos. 1992. Human Heredity 1750-1870. The Construction of a Domain. Ph. Diss, London. Lorenz, Ottokar. 1898. Lehrbuch der gesammten wissenschaftlichen Genealogie. Berlin. Lundborg, Herman. 1913. Medizinisch-biologische Familienforschungen innerhalb eines 2232köpfigen Bauerngeschlechtes in Schweden (Provinz Blekinge). Jena. Martius, Friedrich. 1913. Konstitution und Vererbung in ihren Beziehungen zur Pathologie. Berlin. Morel, Bénédicte Auguste. 1860. Traité des Maladies Mentales. Paris. Müller-Wille, Staffan, and Hans-Jörg Rheinberger (eds.) 2007. Heredity Produced. At the Crossroads of Biology, Politics, and Culture, 1500-1870. Cambridge/Mass. Nasse, C. F. 1873/74. “Vorlage für eine deutsche Irren-Anstalt-Statistik.” Allgemeine Zeitschrift für Psychiatrie 30: 240-248. Roelcke, Volker. 2000. “Naturgegebene Realität oder Konstrukt? Die Debatte über die “Natur” der Schizophrenie, 1906 bis 1932.” Fundamenta Psychiatrica 14: 44-53. Römer, Hans. 1912. “Über psychiatrische Erblichkeitsforschung.” Archiv für Rassen- und Gesellschaftsbiologie 9: 292-329. Rohde, Friedrich. 1895. Über den gegenwärtigen Stand der Frage nach der Entstehung und Vererbung individueller Eigenschaften und Krankheiten. Jena. Rosanoff, A. J., and Florence I. Orr. 1911. “A Study of heredity of insanity in the light of the mendelian theory.” Eugenics Record Office Bulletin Nr. 5.

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Rüdin, Ernst. 1911. “Einige Wege und Ziele der Familienforschung mit Rücksicht auf die Psychiatrie.” Zeitschrift für die gesamte Neurologie und Psychiatrie 7: 487-585. ————. 1916. Studien über Vererbung und Entstehung geistiger Störungen. Vol. 1: Zur Vererbung und Neuentstehung der Dementia Praecox. Berlin. ————(ed.). 1934. “Empirische Erbprognose.” Erblehre und Rassenhygiene im völkischen Staat. München. Rushton, Alan R. 1994. Genetics and Medicine in the United States 1800 to 1922. Baltimore/London. Sommer, Robert. 1901. Diagnostik der Geisteskrankheiten für praktische Ärzte und Studierende. Berlin/Wien. ————. 1907. Familienforschung und Vererbungslehre. Leipzig. ————. 1913. “Familiengeschichtliche Quellenkunde im Gebiete der Psychiatrie und Anthropologie.” In Heydenreich, E. (ed.). 1913. Handbuch der praktischen Genealogie. Leipzig. Strohmayer, Wilhelm. 1908. “Zur Kritik der Feststellung und der Bewertung psychoneurotischer erblicher Belastung.” Archiv für Rassen- und Gesellschaftsbiologie 5: 479-497. ————. 1911. “Die Vererbung des Habsburger Familientypus.” Archiv für Rassen- und Gesellschaftsbiologie 8:150-164. ————. 1912. “Die Vererbung des Habsburger Familientypus.” Archiv für Rassen- und Gesellschaftsbiologie 9: 775-785. Tigges, Wilhelm. 1867. “Statistik, betreffend 3115 Aufnahmen in Marsberg, nebst vergleichender Statistik der der eigenen Untersuchung unterworfenen Verhältnisse.” Allgemeine Zeitschrift für Psychiatrie 24 (Suppl.): 117-475. Waller, John C. 2007. “Poor Old Ancestors. The Popularity of Medical Hereditarianism, 1770-1870.” In Conference: A Cultural History of Heredity II: 18th and 19th Centuries. Max Planck Institute for the History of Science, Preprint 247. Berlin: 131-144. Weber Matthias M. 1993. Ernst Rüdin. Eine kritische Biographie. Berlin. Weinberg, Wilhelm. 1903. “Pathologische Vererbung und genealogische Statistik.” Deutsches Archiv für Klinische Medizin 78: 521-540. ————. 1908/09. “Über Vererbungsgesetze beim Menschen.” Zeitschrift für induktive Abstammungsund Vererbungslehre 1: 377-392, 440-460. ————. 1912. “Weitere Beiträge zur Theorie der Vererbung.” Archiv für Rassen- und Gesellschaftsbiologie 9: 165-174.

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Pedigree Charts as Tools to Visualize Inherited Disease in Progressive Era America Philip Wilson

Within the burgeoning field of eugenics history, pedigree charts have received minimal attention. This remains somewhat puzzling if one accepts the claim that “almost all studies of human heredity” in the early 1900s “tended to involve the collection of pedigrees.” It was, after all, these studies in particular that provided the “facts of human inheritance necessary for the construction of eugenic breeding programs.”1 This lack of attention may, in part, be due to the longstanding marginalization of the study of images in favor of text, at least within the history of science and medicine.2 The relatively minor study of the interpretation of images overall suggests, as art historian Barbara Maria Stafford argued, that images have long been “shunted to the edge of what really matters.”3 Pedigree charts were hardly a new concept of representing information during the Progressive Era, at least for the genealogically minded. Indeed, they had been used for centuries in attempt to trace human lineages back to the Biblical Adam. The term “Pedigree,” or etymologically, pied de grue (a crane’s foot), derives from the symbol used in medieval genealogical tables or trees that, looking like the multi-pronged avian’s foot, denoted a succession of generations. 4 Charting pedigrees was primarily performed “from historical or legal motives.” Biology, per se, had “no place” in the early study of these charts.5 The medical use of pedigree charts in the U.S. was pioneered in 1845 by Philadelphia physician Pliny Earle as he visually documented five generations of one family’s history of color blindness. 6 Yet, this representation of heredity from a medical viewpoint was little copied throughout the nineteenth century. Rather, these charts were predominantly used by animal breeders to record and to predict favorable matings. But as Progressive Era America clamored over the U.S. Government devoting considerably more resources to the proliferation of its agriculture and farm animals than it did its own human population, the pedigree chart reemerged in the study of humans during the “classical era” of genetics.7 (See Figure 1) Slowly throughout this period, the pedigree chart became a standardized scientific tool to medical audiences, using simple, readily recognizable symbols to denote particular meaning regarding heredity and disease. In due course, this tool eased communication about the developing understanding of hereditary patterns of human disease, bridging classical genetics from the theoretical, to the experiential, to the clinical.

1 2

3 4 5 6

Ludmerer (1972), p. 55. Sander Gilman (1988) and Barbara Maria Stafford (1991) have long noted this point. For an excellent overview of the importance of carefully chosen images, see Tufte (2001). One notable exception is Mark Jackson (1995) who has focused upon the visual representation of feeblemindeness in early 20th century eugenic literature. Stafford (1991), p. 6. For an interesting etymological ramble through pedigrees and the nomenclature of nature, see Potter and Sargent (1974). Popenoe and Johnson (1922), p. 39. Rushton (1994), pp. 12-14.

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Figure 1. American Eugenics Society Pedigree Cartoon. Quips on Eugenics - Archive Problems. Source: Folder, Box C - 2 - 6 : 15, Harry H. Laughlin Papers, Pickler Memorial Library, Truman State University, Kirksville, Missouri, USA.

In the U.S., the greatest popularization of the pedigree chart as a tool to visualize inherited human characteristics or traits emanated from the work of the Eugenics Record Office (ERO), in Cold Spring Harbor, Long Island, New York. ERO-trained fieldworkers organized data gathered from throughout the U.S. on charts—what they termed “Mendelian Blanks,” a phrase that attested the bias of their outlook—to represent the incidence and prevalence of particular traits or characteristics that were thought to be hereditarily passed along familial lines. 8 These traits included physically visible conditions such as eye and hair color, multiple births and birthsdefects including hare-lip and cleft palate as well as diseases including tuberculosis, syphilis, and alcoholism. The leading U.S. publication of popular science, Scientific American, claimed that these pedigree charts represented a true “inventory of the blood” of the nation. 9 This paper explores ways in which Harry H. Laughlin, superintendent of the ERO from 1910 to 1939, used this tool to maneuver the flow of information gathered within this repository of human inheritance data to scientific and medical communities as well as to the public. It focuses in particular upon the relationship between inheritance and disease as represented in the pedigree charts that the ERO prepared and distributed during Laughlin’s superintendency. Laughlin’s published writings and correspondence relating to inherited human disease that appeared during 7

8 9

As an example of the apparent lack of attention on humans, Downing (1918), p. 149, argued that the “expert dairyman carefully inquires into the purity of strain and ancestral performance of the animal he mates with his choice cows. The farmer insists on a hog with certified ancestors. We have sense enough to apply such knowledge of heredity as we possess to our farm stock. It seems little enough to ask that we should exercise as much good sense in producing children as we do in the production of hogs and corn.” Such claims were still pouring forth a decade later. M.R. Ferris (1929), secretary to the Council of the Institute of American Genealogy, The National Clearinghouse for Genealogical Information, wrote to Laughlin with the sentiment, “Certainly you will agree that the systematic preservation of the lineages of human beings in the interest of better citizenship is infinitely more important than the registration of livestock pedigrees in the interest of better beef.” Kimmelman (1983) analyzed the agricultural context within which human eugenics arose. The Mendelian leaning of the ERO has been widely noted. See, for example, Rushton (1994) and Turney & Balmer (2000). Collins (1913).

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this period will also be examined in order to better contextualize his use of ERO pedigree charts. The popularity of these particular ERO tools of visualization throughout the United States and Europe is also briefly explored. Their explicit and implicit intended uses in various venues will be examined, including their appearance at the International Eugenic Congresses of 1912, 1921, and 1932, at the Chicago World’s Fair of 1933-34, in many country and state fairs across the U.S., in routine ERO mailings across the country and to Europe, in the correspondence between physicians who sought to update classifications of inherited human disease, and in popular biology textbooks and marriage manuals. In conclusion, special attention is focused upon how the pedigree chart as a spatial arrangement of hereditary patterns of disease prompted discussions about the need for a new healing space—the hereditary clinic.

Human Heredity, Disease and the ERO in Progressive Era Americ The concept of a Progressive Era in U.S. history (approximately 1890–1920) invokes myriad views. One such view represents the time when the U.S. strengthened its position in relation to other leading nations worldwide. Doing so required a double-faced, Janus-type look into both its past and its future. As a nation just over a century old, the U.S. had expended considerable effort, first in fighting to maintain its independence, and more recently, to hold itself intact as a nation. Over that century, the U.S. had also accumulated an expanding genealogical record. Within some circles, it was thought that the nation’s strength and endurance was closely correlated with the physical constitution of its people. The New England physician, Edwin M. Fuller argued that the relatively young U.S. still had a chance to fend off becoming laden with hereditary disease. The older a nation grows, the larger the percentage of hereditary diseases are manifest, and ... after a century’s growth, our nation appeals in silent language to our profession for remedies and intelligent barriers which may be stationed at the portals of society, that the ignorant and easily captivated masses may be warned of the approaching dangers to society and individuals.10

Later during the Progressive Era, and particularly following the Great War (World War I), the U.S. had become globally recognized as a supreme world power. A concomitant need arose in the minds of many to maintain the healthy stock of the American peoples. Should the U.S. population become less pure and “infected” with socially undesirable traits, they argued that the country’s political and economic stronghold would begin to crumble. Looking toward the future, many progressive-minded thinkers argued that in order to prosper even further—and more rapidly—a need existed to better understand the genealogy, breeding potential, and healthiness of the nation’s human reproductive stock. A major shift in thoughts about heredity and disease followed the rediscovery of Gregor Mendel’s work during this era. While working at the newly opened University of Chicago, Harvard-trained zoologist, Charles B. Davenport summarized Mendel’s findings for an English 10

Fuller (1887), p. 206.

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reading audience.11 Within a few years, Mendel’s principles of genetics were being applied not only to plants and animals, but to humans as well. The recessive nature of the trait for albinism in humans was first reported in 1903, followed by similar findings about hereditary deafness in 1905. Two years later, Davenport reported that eye color and hair form (i.e., straight versus curly) pedigree charts of family lineages were interpretable in terms of Mendelian inheritance. No longer did heredity pursue “vague” questions, argued geneticist E.B. Wilson. Rather, it had become approached in terms of a quest to answer very “clear, concise mathematical problems.” 12 Another contemporary explained that heredity was “not the outcome of constitutional transmitted qualities and the condition but is the transmitted quality itself. . . . Of course, heredity, being an abstract noun, cannot be measured except as it is made manifest in concrete things such as stature and other measurable qualities.”13 The rediscovery of Mendel’s principles ushered in a new phase of investigating heredity; a phase shifting from descriptive, morphological explanations of hereditary tendencies to an experimental and statistical based science of genetics. By 1910, a number of diseases had become labeled as hereditary. 14 Geneticists and physicians claimed that certain disorders, including Huntington’s chorea, presenile cataract, and chronic familial jaundice were derived from specific hereditary “determiners” in the “germ plasm.” The presence of the determiner in these disorders was manifest in the visible signs of the disease. If individuals with this type of determiner reproduced, Davenport claimed that “at least half” of the offspring would be “similarly affected.”15 Another major group of disorders was viewed as having originated from a “normal” (i.e., disease-free or at least symptom-free) individual carrying a specific defect in his or her germ cells that did not induce any physically apparent signs but which, when transmitted and “unite[d] with a similarly defective germ cell from the other parent,” inflicted the offspring with disease. 16 His view was consistent with that of the “Constitutionalists” who envisioned human bodies as “carriers of the pathological histories of their race or type” and who argued that it was by passing along “defects” of these “histories” from one generation to the next tended towards “racial degeneration.”17 Preventing the hereditary transmission of such disorders as epilepsy, manic depressive insanity, alcoholism, and cleft palate presented the additional challenge of identifying apparently healthy carriers of particular disease traits. Davenport became Director of the Carnegie-funded Station for Experimental Evolution in Cold Spring Harbor, Long Island, New York, in 1904. Six years later, he established a division of this station, the Eugenic Record Office (ERO), that focused solely upon eugenics. For the next thirty years, America’s most significant advances in promoting eugenics stemmed from this 11 12 13

14 15 16 17

Davenport (1901). For a biographical overview of Davenport and his contributions to hereditary thinking, see MacDowell (1946) and Kevles (1985). Wilson (1908), pp. 200-222. Laughlin, “What is Heredity?”, p. 2. Henceforth, all references to the Harry H. Laughlin Collection at Truman State University’s Pickler Memorial Library are described as “Harry Laughlin Papers.” A useful finding aid entitled “Guide to the Harry H. Laughlin Papers” is available both in print at Pickler Library and online at . For historical explorations into the hereditary thinking underlying venereal disease and tuberculosis— two major chronic diseases of the period—see Wilson (2003, 2006) Davenport (1912). Davenport (1912), p. 7. Cantor (2000), p. 356.

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office.18 The ERO promoted a widespread understanding of the hereditary propensity of disease together with solutions for preventing such diseases in future generations. In 1910, Davenport hired Harry Laughlin to supervise work at the ERO. Laughlin began his career teaching agriculture, natural science and a course in early civilizations at his alma mater, Kirksville State Normal School in Missouri (now Truman State University). 19 While teaching science, he became interested in the new field of genetics. He attracted considerable attention from cattle breeders for his research into the heredity of coat color in shorthorn cattle. In his agricultural lab, Laughlin exposed his students to Mendelian concepts of heredity through breeding experiments involving some uncommon varieties of poultry. Desiring information to classify his newly bred products, he had initially contacted Davenport, who later invited him to the Brooklyn Institute of Arts and Sciences to take his genetics course during the summer of 1907. The two remained in contact, and, in January 1909, Davenport visited Laughlin while traveling to the annual Animal Breeder’s Association (ABA) meeting in Columbia, Missouri. At the ABA meeting, Davenport convinced Laughlin to turn his interests toward the hereditary study of another animal: humans. The study of human heredity required different approaches than that of other animals. For humans were “slowly reproducing” animals who were “not subject to laboratory experimentation for genetic research like drosophila or the white mouse,” thus, Laughlin argued, “it is necessary” in man to consider “as experiments” the history already made in migration, mating, and size of family, and to secure firsthand description of individual persons, their “personal case histories,” and “records of their blood-kinship.”20 The analysis of such pursuits, however, required more patience than in laboratory animals. For to secure the “final correction of the measure of hereditary traits” in humans, one had to wait for a time when specific “generations shall have passed.” Only then would “all of the descendants and collateral kin . . . have developed and exhibited their inborn traits of character.” Only then will these “facts . . . be available for throwing light upon the innate qualities of the propositus.” It is this passage of time, he continued, that “secures unbiased judgments” and “treats defects and talents with equal impartiality” when “arriving at personal and family estimates.” Time again was critical for transforming the hereditary “gossip of one generation . . . [into] cold historical data in the next.” 21 Laughlin was already experienced with human genetic studies having previously engaged his college students in gathering family data about traits that were presumed to be hereditarily influenced. As an advocate of the pedagogical power of visual displays, he had guided them in preparing their own families pedigree charts in an attempt to discern hereditary patterns in the repetition and variance of eye color in successive generations.22 It was regrettable, Laughlin argued, that “the study of humanity is not an exact science like chemistry.” For by establishing 18 19

20 21 22

For extensive historical accounts of the ERO, see Allen (1986) and Watson (1991). Substantial biographical material on Laughlin appears in Hassencahl (1970), Reilly (1991), King (2000), and Bruinius (2006). Barkan (1991) provides a significant treatment of the immigration concerns of eugenisists during Laughlin’s period. See also Laughlin (1922). Harry Laughlin Papers (1939), p. 13. Laughlin (1921a), p. 23. Laughlin (1910, 1914). Laughlin published his eye color heredity work with ten “advanced students” in 1919. As a college student, Laughlin (1899) had advocated his belief in the importance of using expositions as a venue for publicly displaying knowledge.

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such a science, he envisioned its practitioners taking young individuals, analyzing their character, and “improving” them by “supplying” qualities that were lacking and “modifying” those perceived as “abnormalities.”23

Laughlin and the ERO Fieldworkers Laughlin and Davenport organized the ERO around several missions of operation. Accordingly, the ERO was designed to: 1. Serve eugenical interests in the capacity of repository and clearing house. 2. Build up an analytical index of the traits of American families. 3. Train field workers to gather data of eugenical import. 4. Maintain a field force actually engaged in gathering such data. 5. Cooperate with other institutions and with persons concerned with eugenical study. 6. Investigate the manner of the inheritance of specific human traits. 7. Advise concerning the eugenical fitness of proposed marriages, and 8. Publish results of researches. Several of these goals specifically involved the production, storage, and analysis of pedigree charts. Above all else, Laughlin repeatedly distinguished the need for ERO pedigree charts to delve beyond those typically used by genealogists (see figure 2). The genealogist, he argued, “strives to work out the family net-work, giving the names, dates, and connections.” What was missing, however, was “a description of the natural, physical, mental, and temperamental qualities of each member listed . . . .” Once this information was provided, Laughlin concluded, we would have a “record of practical pedigree-value, one which can be used in tracing the descent and recombination of natural qualities within the family-tree.”24 Laughlin summarized, the “usual outline of the genealogist . . . is merely the skeleton” upon which ERO efforts must “clothe it with the sinews and organs of Natural Traits” if pedigree charts are to “have any scientific value.” 25 Even then, he noted, the mere charting of biological information was only the beginning. “Individual Analysis Cards,” listing all of each pedigree members’ constitutional traits, tendencies, and disorders were also required to complete the “critical biological biography” for each family. For when displayed in this manner, the “bare facts concerning the natural capacities and shortcomings of various members of a family . . . constitute an instructive guide for the family.” 26 23 24 25

26

Harry Laughlin Papers, “Ideal Young Man.” “Eugenics and Other Sciences.” (1920), p. 77. Harry Laughlin Papers, “A Few Points to Observe in Writing up Notes.” Elsewhere (Harry Laughlin Papers, 1939a, p. 15), Laughlin acknowledges that the genealogists” biographical accounts were of some help to eugenics research as “records of human functioning which check[ed] constitutional traits diagnosed or collected from other sources.” See also “Eugenics and other Sciences.” (1920). Harry Laughlin Papers, (1915) Sections II and III. Banker (1923), p. 306, suggested the word “ecography” to account for the complete biological and historical component of family histories. The ERO was not alone in providing instructions of the construction of human pedigree charts. J.F. Munson (1910), a physician working at the Craig Colony for Epileptics in Sonyea, New York, published easy-to-follow guidelines in the New York Medical Journal.

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Figure 2. Sample Pedigree Chart, Davenport and Laughlin (1915) How to Make a Eugenical Family Study.

Laughlin’s ERO efforts were aligned with those that eugenicists Paul Popenoe and Roswell Hill Johnson articulated in Applied Eugenics (1922). There they claimed that genealogy has its application to science by “furnish[ing] means for getting knowledge of the laws of heredity.” Such an application made it possible for individuals to “better understand” their “place in the world” and “to marry better.”27 With this collaborative approach in mind, it was argued that the “Golden Age of genealogy is yet to come.” For in such an Age, genealogy would “become the study of heredity, rather than the study of lineage.” Or rather, insofar as humans were concerned, “heredity” would become functionally defined as “the interpretation of genealogy.” 28 When constructed with critical care, pedigree charts and the accompanying analysis cards— collectively referred to by the ERO as “scientific genealogies”—would be able to serve multiple 27 28

Popenoe and Johnson (1922), p. 330. Ibid., pp. 335–337.

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purposes. On one hand, they provided essential information for every individual to “inquire into the natural endowment of its . . . members and by pedigree study to find out how the traits of each would be transmitted in given matings—to be calculating and forehanded in mate selection, so that the offspring will present fortunate combinations of desirable family traits.” To this end, Laughlin argued, “every family should establish a permanent Family Pedigree Archive, for only through the information conveyed by such may the facts of fortune be worked out,—or, to put it in the old way, may one see where the finger of destiny points.” Indeed, it would serve a greater value still if “several branches of one’s own family” could have their investigations “coordinated by a Family Association” whereby a “most excellent and useful scientific pedigree record” of the whole family would be produced, a task requiring “but little effort on the part of each branch.” 29 Such a study, he noted, becomes “almost priceless” to a given family, particularly after the “oldest person consulted in preparing it has passed away.” For, “as a rule, an individual is personally acquainted with but three generations of his or her kin and connections, and without personal knowledge and care [,] character analysis is very difficult.”30 Indeed, it “should be considered a filial duty as well as a duty to society to secure at the earliest opportunity from the oldest living members of one’s family detailed facts concerning those who still live in the memory of their contemporaries.”31 It will be “a happy day for our national welfare,” indeed, “when the keeping of . . . [a family pedigree] archive becomes a national family habit.” Each family merely “needs but an organizer” to accomplish this goal. 32 Ever the organizer himself, Laughlin envisioned his own pedigree archiving task on a much grander scale. Similar to what he urged each family to acquire, Laughlin sought for the ERO to become the national pedigree archive. By acquiring “all authentic family history studies,” the ERO “seeks ultimately to have an index of the network of the family kin, and of the natural heritable traits of all of our better American families.” As this “ideal[ized goal] becomes realized, it will become less difficult,” he concluded, for “representative families by using the [ERO’s] files . . . to work out their pedigrees in practical pedigree—i.e., trait prediction—fashion.” 33 To achieve this national aim, Laughlin coordinated the collecting and recording of family data through an extensive outreach program. From 1910 through 1924, he and Davenport trained teams of “field workers” in the principles of human genetics and provided them with skills necessary to gather extensive family histories.34 The field workers were mostly young collegeeducated women. As political historian Diane B. Paul has argued, women were “especially well suited” for eugenic fieldwork. They had the ability to form the sympathetic relationships with families in order to persuade them to divulge familial information. Additionally, the women’s “intuition and sharp eye for detail” allowed them to “swiftly and accurately” assess an individual’s physical, mental, and tempermental traits. And, alas, reproductive matters by convention fell into

29 30 31 32 33 34

Harry Laughlin Papers, “The Permanent Family Pedigree Archive”. Harry Laughlin Papers, (1915). Davenport and Laughlin (1915), p. 3. Harry Laughlin Papers, “The Permanent Family Pedigree Archive”. Harry Laughlin Papers, “Eugenics”, p. 6. For a telling account, see Bix (1997). Laughlin (1929) claimed to have overseen the training of 258 field workers between 1910 and 1924.

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a woman’s provenance.35 Not surprisingly, one of the desired qualities of a field worker was that she professed to “like pedigree study.”36 Relying upon his pedagogical prowess, Laughlin exposed field workers to a series of lectures and lab activities on eugenics. The range of topics he addressed included chromosomal structure, anthropological measurement, elementary statistics and discussions of the medical conditions deemed to be, at least in part, hereditary such as skin pigmentation, insanity, cataracts, and epilepsy. Readings centered around Brown University biologist Herbert E. Walter’s Genetics and were also drawn from intelligence measurement authors Alfred Binet, Lewis M. Terman, and Robert M. Yerkes. Additionally, Laughlin led these students through an experimental study of cross fertilized and pure bred corn in order to allow them to personally uncover the Mendelian laws regarding the segregation and recombination of hereditary traits. In subsequent discussions, students used visible evidence obtained from their corn experiments as analogies for the transfer of “defective” traits and “unfit” matings in the human population. Students were also provided with ERO-established guidelines instructing them how to make a eugenic study of a family. Laughlin posed questions including the following to acquaint his students with charting pedigrees. 1) Peter’s wife’s mother was feebleminded. Peter was normal and so was his wife, but their son was affected. Where else must the taint have existed? 2) I married a widow that had a grown-up daughter. My father visited us and married my stepdaughter. I had a son born and my father had a son. Tabulate the relationships. 3) Mr. Harold Leek married Ida Smith, daughter of Egbert Smith. The bride was the daughter by first marriage. His (Egbert’s) marriage was with the daughter of Joseph Leek, Harold’s father. Chart and indicate in words the curious relationship existing.37 Field worker students also gained experience in analyzing pedigrees of “social defectives.” In one comparative pedigree exercise, students first examined Family A, which showed a “great susceptibility to manic depressive insanity.” It was known that among some of the members of this family, a “very light exciting cause was sufficient to call for an attack.” Family B’s pedigree showed only one of the near kin to have become insane, and that only “through the most formidable array of exciting causes.” However, other members of Family B “had traits shown on the pedigree chart which were sufficient indicators of insanity, which might attack a given member.” Students then compared the analysis of these families with “that of so-called normal families” in which no exciting cause “would have been sufficient to break down an individual to the point of insanity.” 38 To gain experience in charting family pedigrees of actual “social defectives,” students were sent on supervised educational visits to study the patient populations in nearby clinics at King’s Ridge, Amityville, Letchworth Village, and Central Islip. They also visited immigration control facilities on Ellis Island.39

35 36 37 38

Paul (1995), pp. 54-7. Harry Laughlin Papers, “Qualities Desired in a Eugenical Field Worker”. Harry Laughlin Papers, “Pedigree to be Charted by Class”, pp. 1-2. Harry Laughlin Papers, “Outline of Notes for Condensed Statement and Examples of the Principals of Eugenics,” pp. 1-2.

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In contrast to Laughlin’s encouragement of America’s best families to submit their own family pedigree studies to the ERO, he focused ERO field workers’ efforts towards documenting the pedigrees of those he deemed as “socially defective” or “socially inadequate.” Fear was already looming over the increasing numbers of “degenerates” in the U.S. before the Great War. State legislators deemed such individuals as the “greatest problem that confronts our nation,” and they claimed the “degenerates” were present in “a greater multitude” than anyone could count. 40 Supportive of their concern, Laughlin and his field workers provided the essential ingredient that legislators had been missing: specific quantification of the “social deviants” who, it was argued, by their “inferior blood” were viewed as a great and costly “menace to society.” Sociologists had long engaged in elaborate discourse over the “3Ds” of society (the defective, the dependent, and the delinquent classes). By the mid 1910s, many found this classification scheme to be too inclusive to guide specific actions toward improving societal discord. More precise definitions were needed to identify those special classes of society who “need special care, restraint or direction, who as a group do not contribute in net to the general welfare . . . but who on the contrary . . . entail a drag upon those members of the community who have sufficient insight, initiative, competency, physical strength, and social instincts to enable them to live effective lives without particular social custody.”41 Typical of his immodest proposals, Laughlin sought to rectify this nosological nuisance, and he campaigned for the official adoption of the term “socially inadequate” as a more precise designation of the “3Ds” within society. According to many ERO publications, Laughlin subdivided the “socially inadequate” to include 1) the feeble-minded, 2) the insane, 3) the criminalistic, 4) the epileptic, 5) the inebriate, 6) the diseased—including those with tuberculosis, leprosy, and venereal disease, 7) the blind, 8) the deaf, 9) the deformed, and 10) the dependent— including orphans, old folks, soldiers and sailors in homes, chronic charity aid recipients, paupers and ne’er-do-wells.42 Every state institution soon became eager to host or hire an ERO-trained field worker who collected information about the ancestry of the insane, the feeble-minded, the criminals, the diseased, and the paupers housed therein. Field workers became veritable “human research machines.”43 Data was organized on what were called “Mendelian Blanks”—family pedigree charts that contained particular information about the incidence of specific traits or characteristics thought to be hereditarily linked. These traits included physically visible traits such as eye and hair color, multiple births and birth defects including hare-lip and cleft palate together with diseases including tuberculosis, syphilis, and alcoholism. Although special talents in music, math, sports, or invention were also recorded, particular focus was given to the subjective 39

40 41 42 43

Harry Laughlin Papers, “A Corn Breeding Experiment.” Henry H. Goddard (1910), noted eugenicist and superintendent of the care of the institutionalized feeble-minded in Vineland, New Jersey, also supplied specific instructions for fieldworkers in the preparation of pedigree charts. Report of the Commission on the Segregation, Care and Treatment of Feeble-Minded and Epileptic Persons in the Commonwealth of Pennsylvania, (1911). Laughlin (1921), p. 68. For further discussion of these categories, see Wilson (2002). Bix (1997), p. 640. Zenderland (1998), p. 159, commented upon the similar training of field workers and social workers, yet noted that the former worked under the rubric of nature whereas the later worked under the rubric of nurture.

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assessments of mental ability and physical defects. It was hoped that analyzing traits in the form of pedigree charts would enhance the understanding of inheritance patterns of particular diseases. As an example, the ERO’s “Schedule for Recording First-hand Pedigree-data in Hereditary Eye Defect and Blindness” stated that it was the immediate aim to “secure for study [the] authentic pedigrees of families with hereditary eye defect to the end that the rules of inheritance of definite eye defects may be more clearly determined.”44 Field workers carried their own copies of the ERO’s special bulletins including No. 2 Study of Human Heredity, No. 6 The Trait Book, No. 7 The Family-history Book, and No. 13 How to Make a Eugenical Family Study. These publications, which attempted to standardize pedigree reporting methods, were also readily available for purchase by the public. In order to better appreciate the ERO fieldworkers’ use of pedigree charts in the field, let’s turn briefly to the instruction that they received from Laughlin.

Pedigree Charts and the ERO Although the ERO acknowledged that information about family histories had “for many years” been obtained through the application material, medical examinations, and letters from relatives regarding “defectives” in “the better organized Hospitals and Institutions,” such information was “far from satisfactory.” The ERO claimed that “experience had shown that there is only one way to get a satisfactory family history of a stranger and that is to go, or to secure a trained assistant to go, to the various members of the family and with tact and patience and time secure the necessary facts.”45 Using field workers to “go to the homes” and to “interview persons that can and will give the desired information” would, it was claimed, enhance the precision and accuracy of the data obtained. Such workers were to first learn all they could about a patient from the office files at the institution, even obtaining addresses of patient’s relatives and friends. Although they were encouraged to focus upon the specific trait being studied (i.e., the primary trait), field workers were also urged to embrace further opportunities to “learn of other traits that may be significantly or incidentally associated with the primary trait.”46 “Just before starting out to visit the relatives and friends,” the field worker is to visit the patent “in his ward or cottage.” Then, “armed with recent personal knowledge of the patient, which assures her cordial welcome” the fieldworker proceeds to visit the patient’s home and “interviews the relatives, friends, and family physician.” The field worker must endeavor to “see as many relatives as possible,” as “facts omitted or overlooked by one [relative] are often recalled and told in full detail by another.” And, “by this means information already obtained is confirmed.” Once this data is collected and recorded, a pedigree chart is to be constructed.47 Field workers were sent out with the assurance that “the parents or other relatives of the patient” would be “pleased to think that the hospital or school takes such an interest in the patient as to send a visitor to the home.” 48 44 45 46 47

Harry Laughlin Papers, "Schedule for Recording First-hand Pedigree-data on Hereditary Eye Defect and Blindness", p. 1. Davenport (1915), p. 18. Davenport, Laughlin, Weeks, Johnstone, and Goddard (1911), p. 7. Davenport, Laughlin, Weeks, Johnstone, and Goddard (1911), pp. 1-2.

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As many members of the “restricted” and “extended” families as possible were to be recorded on the pedigree chart.49 Fieldworkers were urged to “lay great stress upon the reliability of the sources” of the information that they obtained, checking the “testimony of one informant against another.” The traits and personalities of those individuals in the collateral lines (i.e., any line other than your direct ancestors) of the pedigree were to be strongly considered since a better understanding of their genotype would “throw light upon the germ-plasm of the propositus.” Field workers were warned “Don’t diagnose!”—and to use terms including ‘insane,’ ‘feebleminded,’ ‘criminal,’ ‘neurotic,’ and ‘normal’ with great caution. Rather, they were instructed to provide sufficient details to “enable an expert to draw some conclusions from the data.”50 Standard symbols were to be used to represent afflicted and unafflicted individuals, specific lines of generational lineage, and specific traits and afflictions (see figure 3). The ERO produced a Trait Book to ensure that standard symbolic representations were known. 51 Some disease or “defective” conditions were so frequently studied that they acquired specific color representations on pedigree charts. For example, red was used to encode for epilepsy, green for insanity, violet for criminalistic tendencies, and black for feeble-mindedness. 52 Finally, fieldworkers were alerted to provide the names and addresses of “defectives who need Institutional care.” As such, the data that they collected became particularly “useful information . . . when application is made for admission” to respective institutions.53 The ERO relied upon the pedigree chart as their most common tool of assimilating and promulgating information about the nation’s reproductive stock, both the lineages of favorable stock as well as those of the “socially inadequate.” Such charts served practical measures for the ERO by “determin[ing] . . . the eugenical fitness” of a contemplated marriage, “gauging the specific educability or the hereditary potentialities of a given individual,” and “evaluating the intrinsic value of . . . [a] family, whenever such knowledge may aid . . . [that] family in directing along profitable lines the education of its youth and in encouraging biologically fortunate matings of its marriageable members.”54 Originally a tool for genealogists and biographers, this chart was modified by fieldworkers and others at the ERO so that it could just as easily be used to express biological aspects of all the individuals within a given family. By incorporating all of the known

48 49

50 51

52 53 54

Davenport (1915), p. 18. The “restricted” family consisted of the propositus, his siblings, and the consorts and children of these siblings; the father of the propositus and the father’s siblings and consorts and their children; the father’s father and the father’s mother as well as the corresponding relations on the mother’s side of the family. The “extended” family included, in addition to the restricted family, a history of the uncles and aunts by marriage, the consorts and children of the cousins, the siblings of the grandparents and their consorts and children, as well as their children’s children, and of the eight great-grand-parents. Davenport and Laughlin (1915), p. 6. Harry Laughlin Papers, “A Few Points to Observe in Writing Up Notes”. Among the disease traits or characteristics listed were: alcoholic, blindness, Bright’s disease, cancer, chorea, cripple, criminalistic, deafness, dementia, dropsy, eccentricity, encephalitis, epileptic, goiter, general paralysis of the insane, gonorrheal, hysteria, ill defined organic disease, insane, kidney disease, locomotor ataxia, manic depressive insanity, migrainous, neuropathic condition, obesity, paralytic, paranoia, pneumonia, senile, sexually immoral, shiftlessness, softening of the brain, syphilitic, traumatic insanity, tubercular, vagrant, varicose veins, and vertigo. Davenport, Laughlin, Weeks, Johnstone, and Goddard (1911), p. 4. Davenport, Laughlin, Weeks, Johnstone, and Goddard (1911), p. 2. Harry Laughlin Papers, (1915).

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and gathered data about a particular family on one sheet of paper, these charts maintained a visual simplicity.

Figure 3. Key to Symbols used in ERO Pedigree Charts, ERO Bulletin No. 2 (1911) The Study of Human Heredity (1911).

Overall, pedigree charts objectified, quantified and visualized many previously invisible aspects of disease. They penetrated into the germ layer giving new insight into the genotypic level regardless of whether any aspect of the disease was phenotypically expressed. 55 In that way, they allowed for better discrimination of hereditary difference between individuals. But as ERO efforts demonstrated, they also provided a new way of imaging or re-presenting disease. 56 As such, they 55

For a contemporary discussion of genotype, see Johannsen (1911). Sapp (1983) further contextualizes the genotype-phenotype distinction as iterated during this period. The word “idiotype” was used somewhat synonymously with “genotype” in literature of the period, particularly in that of the Constitutionalists’ writings on the body.

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became a conceptual tool for more fully appreciating patterns of inheritance for particular diseases. They also revealed a new structural knowledge that gave a better glimpse of the movement of disease via the germ plasm throughout a given family. In and of themselves, these tools exhibited connections and offered some cautions as to what to look for in existing and future generations. On their own, however, they did not offer infallible explanations of particular patterns of inheritance. Many different humans had gathered information for pedigree charts, and thus, this process left considerable sources of error. Perhaps an even greater source of error arose from the potential of missing information in one or more generations. Quite often, field workers and others relied solely upon the subjective views of one family member to account for various states of disease in all of that individual’s known relatives. Even if that individual divulged all that he or she knew, much of this view may have stemmed from hearsay. Others, it was noted, may have been cajoled by fieldworkers into giving information that they thought the fieldworkers wanted to hear. Finally, no system was in place to verify either the information that was collected or its recording. Although pedigree charts did exemplify one attempt of the scientification of society that marked much activity in the U.S. during the Progressive Era, the reality of data collection, at times, failed to uphold objectives of valid and verifiable information gathering according to the expectations of the scientific method. Or, in other words, as critics claimed, they were “insufficiently critical to establish what actually is true.”57 Within the world of medicine, pedigree charts became shorthand representations of the presence and potential patterns of disease. As with any shorthand system of symbolization, minimalist abstractions are rendered. In this case, humans were disembodied into some type of representational simulacrum in which they appeared as only bits or bytes of select information. This idea advanced reductionistic representations of humanity by offering a tool that diminished the concept of the human. The disembodiment of humans to mere boxes and circles encoded with information was consistent with reductionist thinking common of this era that encouraged medical thinkers to look at the body more as distinct components rather than as a whole patient. Within a short timeframe during the “classical era of genetics,” pedigree charts gained an iconic status.58 Though mere lines, circles, and squares, they held a power to persuade viewers to think about heredity within their own family. Following art historian Barbara Maria Stafford, it is precisely these kind of forms—the simplest forms of artistic expression—that represent “ideal forms,” the very forms that “should be imposed on disorderly biota” in order to clarify the desired image to be represented.59 Thanks to the efforts of all of those who have contributed to our better understanding of the “mapping cultures of twentieth-century genetics,” we have learned to see and to read new meaning in the design of linkage and genomic maps.60 As a form of visualizing meaning, pedigree 56 57 58 59

For an overview of the social construction of genetic disease, see Yoxen (1984) in contrast to Child’s(1999) history of ideas approach. Ludmerer (1972), p. 59. For coverage of other icons related to heredity, see Nelkin and Lindee (1995) and Rheinberger and Gaudillière (2004). Stafford (1991), p. 3. Further exploration into the semiotics and symbolic significance of the components of pedigree charts would be most helpful, though it lies beyond the scope of this paper.

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charts are a kind of mapping as well. Like maps, they artfully produce a linear image that both spatially and temporally delineate patterns of connectedness between generations. They provide a means of orientation and direction; they concisely depict phenomena in relationship to each other; they convey meaning through the power of synopsis; and they are perspectival in that what features are included depended upon the aims, needs, interests, and mindset of the pedigree chart designers. Curiously, these little mini-exhibitions of knowledge served both individual and societal needs. On one family’s pedigree chart, each individual was highlighted as was his or her interconnectedness with everyone in an entire family, at least regarding a particular trait or disease. These charts seemed to introduce labels of either normality or deviance upon potentially all members of the family represented therein. But the ERO also used vast collections of pedigree charts as a form of collective data, expanding their apparent range of observation, in a manner that supported their overarching efforts of societal reform. Such efforts were aimed, in part, to convince American society that eugenics was working well within mainstream science of the era. Turning specifically to Laughlin, we find that he relied upon both the scientific and the simplistic ways that pedigree charts conveyed information as part of his rhetorical strategy to persuade various audiences about the potential that eugenics held for individual families, for the nation, and for the world. As an example, Laughlin found the pedigree chart to be useful in his persuasive proposals to gain support for what he viewed as the best means of eliminating the social burden created by the “socially inadequate.” The “conscious striving for race betterment on the part of the socially inadequate,” he argued, “is impossible . . . . Therefore society must control their reproduction.” It ought to be a “eugenic crime,” he claimed, to “turn a possible parent of defectives loose upon the population.”61 As secretary to the Committee to Study and Report on the Best Practical Means of Cutting off the Defective Germ Plasm in the American Population, Laughlin issued the committee’s report detailing ten possible “cures” of the problem. 62 Ranging from segregation to euthanasia, the committee strongly favored reproductive sterilization as the “least objectionable” and the “most cost-effective” solution.63 Pedigree charts were, so Laughlin argued, an “obvious” choice to unambiguously document and visualize the “practical application” of eugenics schemes.64 With considerable rhetorical skill and sheaves of pedigree charts, Laughlin convinced many states to adopt a model law that he had drafted to serve as the official legislative organ to involuntarily control the reproduction of their institutionalized populations. By 1921, the year before the publication of Laughlin’s Eugenical Sterilization in the United States, 3200 individuals 60 61 62

63 64

See the companion volumes of Rheinberger and Gaudillière (2004), and Gaudillière and Rheinberger, (2004). As cited in Kevles (1985), p. 108. See Laughlin (1920) for an example of his rhetorical prowess in promoting sterilization. Other committee members and consultants included prominent New York lawyer, Bleeker Van Wagenen, Johns Hopkins physician, Lewellys F. Barker, Henry Goddard, the “psychometrician” at the Vineland Training School in New Jersey who introduced IQ testing into the US, Johns Hopkins geneticist Raymond Pearl, and Louis Marshall, leader of the American Jewish Congress. Reilly (1991), p. 60. Laughlin (1912), p. 121.

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across the nation were reported to have been sterilized. That number tripled by 1928, and by 1938, nearly 30,000 met this fate. More than half of the states in the US adopted Laughlin’s law, with California, Virginia, and Michigan boasting of their lead.65 Laughlin also used pedigree charts to secure the staunch support of the U.S. judiciary. In a precedent-setting case, that of Buck v. Bell in 1927, Supreme Court Justice Oliver Wendell Holmes, Jr. upheld the Virginia Statute and claimed, “It is better for all the world if, instead of waiting to execute degenerate offspring for crime, or to let them starve for their imbecility, society can prevent those who are manifestly unfit from continuing their kind.” In specific reference to the reputed feeble-mindedness of Carrie Buck and her ancestors as attested on pedigree charts, Justice Holmes deemed in words that have continued to ring loudly, “Three generations of imbeciles is enough.” Following this pronouncement, Buck was reproductively sterilized against her will but in accordance to the highest law in the land.66

Popularizing Eugenics Laughlin spearheaded several efforts to popularize eugenics beyond the confines of the ERO. After witnessing the success of the first international congress on eugenics in London in 1912, Laughlin facilitated two additional international congresses; one held at New York City’s American Museum of Natural History in 1921 and another at the same venue in 1932. Although aimed primarily at professionals, these conferences drew international attention to U.S. efforts to curb the reproduction of “degenerates” and promote the proliferation of the genetically well endowed. Eugenicists devoted considerable effort to enhance public awareness about eugenics. Princeton geneticist and cytologist Edwin Grant Conklin noted that the “widespread ignorance” regarding heredity was profound. “Any general reform,” he argued, “must rest upon enlightened public opinion . . . the schools, the churches and the press can do no more important work for mankind than to educate the people, after they educate themselves, on this important matter.” 67 Campaigns centered around educating the public in order to foster a general “eugenic conscience.”68 Effort must be expended, another eugenicist argued, such that the public gains a sensitivity in favor of eugenic fitness similar to what they have against incest and miscegenation. 69 As the ERO was actively involved in educating the public, Laughlin worked diligently to keep the message of eugenics paraded before the populace. As a public servant, he oversaw the design of a multitude of easy-to-understand handouts which, using simplistic diagrams and brief accompanying text, were used to relay particulars about the genetic principles underlying human eugenics for the lay public. He distributed these handouts freely to thousands of individuals who 65 66

67 68 69

Reilly (1991), p. 97. Court records were used as the basis of Smith and Nelson (1989). Stephen Jay Gould (1984) has briefly addressed Carrie Buck’s plight , and much of the sentiment of this case, though not all factual, was portrayed in the 1994 made-for-TV movie, “Against Her Will: The Carrie Buck Story.” The 1994 Worldview Pictures Production documentary, “The Lynchburg Story: Eugenic Sterilization in America” is considerably more accurate in its presentation of this case. Conklin (1922), p. 308. Walter (1914), p. 251. Ibid., p. 252.

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contacted the ERO. As part of his routine, Laughlin would ask people to complete two family pedigree charts which he included in his mailings. He urged the recipient to be as accurate and complete as possible in identifying all the hereditary traits in each family member according to the list he enclosed. If desired, the ERO was “glad to supply . . . small rubber stamps” of squares and circles, free of charge, to ease the completion of the charts. He urged people to “recast” the chart “two or three times” before drawing up a final copy, to incorporate all “new kinsmen . . . discovered” in the process.70 After completing both forms identically, the recipients were to return one of them to the ERO for “secure filing” where it would remain “permanently available for reference by persons with legitimate concern” for such records. 71 The other, he suggested, should be kept for their own family records. His actions were aimed at providing families with a tool that expanded the genealogical tree recorded in family bibles, helping them to better visualize the genetic traits present in their family’s recent past. This task also fulfilled Laughlin’s self-serving interest of supplying data to the ERO beyond that generated by the field workers. Similar letters were sent to community clubs and organizations as well as to libraries. Here again Laughlin distinguished the typical genealogical family tree from a pedigree record of familial traits. By completing ERO charts, he noted, one can “trace the descent and recombination of natural qualities in the family tree in true pedigree fashion.” He closed his form letter to libraries acknowledging their help in “aiding pedigree study of the human family” by “securing valuable permanent records which otherwise would not be prepared, or if prepared, would be lost to the family and the state.”72 College courses at the time, as well as their accompanying textbooks, devoted increasing coverage to human eugenics. By the late 1920s, nearly 400 U.S. college courses were taught on eugenics.73 Laughlin directed a series of letters to professors of biology, sociology, and psychology urging them to adopt his pedigree charting methods. Professors were asked to supervise student’s completion of the ERO’s standardized pedigree forms, and Laughlin left it up to the professors to collect the forms to return to the ERO or to “eliminate” any of the pedigree charts that were, according to the professor, “inaccurate or scantily prepared.”74 Professor U.G. Weatherly of Indiana University claimed that this project “furnished the very best possible kind of laboratory material.” There “could be no more effective method of getting young people in contact with the serious problems of family inheritance,” he added. Students are “led not only to take a vital interest in the family history,” but this pedigree analysis gave them “a sound and impelling interest in the future fate of their own groups and of the race.”75 In another effort, Laughlin oversaw the ERO’s sale of sets of lantern slides that cold be easily used to deliver pre-organized lectures. One section of these sets, some 21 slides, illustrated family pedigree study, whereas another set of 23 slides showed “Pedigrees of Defectives.” This non-profit ERO venture was available for anyone “interested in eugenics studies.” 76 70 71 72 73 74 75 76

Davenport and Laughlin (1915), p. 9. Harry Laughlin Papers, “Family-Tree Folder,” p. 1. For a discussion of the confusion over various attempts in analyzing these pedigrees, see Harry Laughlin Papers, (1937). Harry Laughlin Papers, “Letter to Libraries”, p. 2. Allen (1983), p. 116. Harry Laughlin Papers, “Memorandum of Suggestions to Instructors”. Harry Laughlin Papers, “Family Pedigree Study as College Laboratory Work”. Harry Laughlin Papers, (1938), title page.

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In addition to his public outreach through mailings and lectures, Laughlin supported public exhibitions as a means to further educate people regarding the benefits of eugenics. In a 1921 exhibition, Laughlin provided an overview of the usefulness of human pedigree study in a case study devoted to the naturalist, John Burroughs. In selections of the text that accompanied this exhibit, we find a rationale for pursuing pedigree study. With the materials called inborn traits—physical and mental—nature casts new human personalities by the process of segregation and recombination in heredity. Here also the laws of chance hold good, but the whole problem of human inheritance is so complex that at present only an ancestral trait here and a quality there may be safely wagered, by a definite chance to enter a given offspring. The task is further complicated by the fact that in heredity there is occasionally a new value—a mutation the geneticists call it, not previously present in any form. In such cases the same does not accord with the usual known rules. Nevertheless the Science of human pedigree-analysis is making headway.77

In what was undoubtedly his single greatest success in educating the masses, Laughlin organized a eugenics exhibit around the theme “Pedigree-study in Man” as part of the Chicago World’s Fair held in 1933 and 1934. Consistent with the Fair’s “Century of Progress” theme, Laughlin incorporated many recent eugenic advances within his exhibition. 78 He created a series of panels which, when viewed according to a specific order, presented the principles of human heredity as a puzzle which exhibit goers could solve based upon their own personal and family experience. Since “no one was stationed permanently at the eugenics exhibit,” it was “necessary that the charts be self-explanatory and well adapted for conversation among mutually interested visitors.” 79 To ensure that his exhibit caught the attention of every age and social class, he employed a variety of practical laboratory set-ups. Some stations were set up with the Midwestern farmer in mind, invoking parallels between human stock and live stock breeding and crop production. The socially elite were catered to with a “Test for Instinctive Appreciation of Quality and Elegance.” In this test, ten animal fur samples of varying quality were placed on a table. Using score cards, fair goers were asked to “consider quality and elegance in relation to the appeal [that the furs made] to you personally,” and then to rank the samples from best-liked to least-liked. Their findings were then applied to corresponding pedigree charts that outlined how certain favorable traits in a human population could best be propagated.80 Part of the effort to improve general eugenic knowledge was aimed at approaching marriage in a more discriminating manner. If young people, “before picking out their life partners . . . are taught to realize the fact that one marries not an individual but a family, then “better matings will be made.”81 Others advocated that stricter marriage laws were essential to reduce the future threat of heritable bad habits. As the popular sexual hygiene manual, Safe Counsel or Practical Eugenics, advocated, one of the “simplest and most effective methods of improving the human race” was by 77 78

79 80 81

Harry Laughlin Papers, (1921), p. 1. The “Century of Progress” theme was selected in attempt to “demonstrate to an international audience the nature and significance of scientific discoveries and the methods of achieving them.” History Files— A Century of Progress, 1998. Laughlin (1935), p. 161. Harry Laughlin Papers, (1932). Popenoe and Johnson (1922), p. 164

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requiring a “certificate of freedom from transmissible disease before a marriage license could be issued.”82 The authors noted that such laws existed in a few states, but that they “have never been, and are not now systematically enforced.” Nothing prevented persons forbidden to marry in one jurisdiction from doing so in another. Others noted that the marriage laws as they existed were somewhat contradictory to eugenics aims. For instance, “sexual offenders” were often “forced” to marry in order to “legalize the offense” and “save the woman’s honor.” 83 Implementing a new “health certificate plan” for a “eugenic marriage license” would require a “clean bill of health, both mental and physical, from every applicant for a marriage license, both male and female—that certificate to be signed by a reputable physician who would not dare risk his professional reputation without a rigid, thorough and final examination. And let us make it a felony to go outside the jurisdiction of the state to evade the letter of the law.”84 ERO efforts also warned that unfit marriages would bring about distasteful and unproductive offspring. Laughlin’s exhibit at the Chicago World’s Fair incorporated pedigree charts showing how both desirable and undesirable traits could be passed along family lines. By placing two pedigrees side by side, he drew particular contrasts between the presidential Roosevelt family and the “degenerate” Ishmael family. Similar to the Jukes and the Kallikaks, the Ishmaels from Indiana were used as a representative family of over 1,750 individuals in which eugenicists traced the linear passage of “defective germ-plasm.”85 By studying the passage of ancestral lineage, viewers were urged to drop any lingering views that marriage was purely a human choice and adopt the more socially desirable belief, at least according to the eugenicists, that responsible Americans pursued marriage mindful of eugenics.86 Laughlin wanted to convey through pedigree charts that bearing children should be envisioned more as a social privilege than merely as an individual right.

Pedigrees and Hereditary Disease Clinics New York University Professor Rudolph Binder argued in 1923, that eugenics would be “more readily adopted by a community which knows the value of good health.” Moreover, communities that held a “high ideal of health” and in which physicians worked towards preventing disease via improvements in sanitation and hygiene, and towards “producing a finer type of man,” would eventually come to view disease itself as “something abnormal.” 87 Shifting the focus of medical practice from cure to prevention and to the elimination of disease became a rallying point for a number of health care reformers in the 1930s. Indeed, many came to view achieving these goals through eugenical means as the panacea for disease control. 82 83 84 85 86

87

Jefferis and Nichols (1922), p. 16 Walter (1914), p. 251. Jefferis and Nichols (1922), p.17. Claims that pedigrees “ought to be as obligatory as a birth certificate or a marriage license” were still being made through the end of the 1950s. See Montagu (1959), pp. 297-98. Rev. Oscar C. McCulloch (1888) traced the lineage of this family’s “degenerates” in “The Tribe of Ishmael: A Study in Social Degeneration.” As cited in East (1929), p. 233. For Laughlin’s own account of the success of this exhibit, see Laughlin (1935). For a comparable assessment of the eugenics exhibition at the Second International Congress of Eugemics in 1921, see Laughlin (1923). Binder (1923).

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Laughlin’s interest in the eugenic control of disease continued to flourish. His correspondence indicates a growing interest in various methods to apply eugenical research to a wider range of health care reform measures. By doing so, Laughlin created the need for even more family pedigree assemblage and analysis. No longer were the institutionalized viewed as his only target population; rather, he now expanded his purview to cover the entire U.S. population. The realm over which eugenics ruled was, in the eyes of many, expanding. The role that ERO field workers and institutional physicians had once played in gathering pedigree data would now fall upon the shoulders of every family physician. Laughlin faced one major problem—few family physicians, in his opinion, had sufficient practical knowledge to apply eugenics to their own patient populations. The 1910s campaign to restructure American medical education by providing all medical students a sound working knowledge of human genetics and eugenics was not realized by the 1930s. 88 Having personally witnessed Davenport’s crusade for this reform, Laughlin knew well the previous strategies that had been employed. Therefore, instead of trying to reintroduce previously ineffective stratagems, Laughlin looked for a new approach. After considerable exploration, he concluded that a human genetics clinic would best fit the public’s need and serve his own agenda as well. A clinic would serve as a training center for physicians, a gathering point for geneticists and public health workers to share information about best approaches for preventing disease, as an outlet to provide direct patient care, and as a counseling center for patients who were concerned about marriageability issues. Laughlin devoted considerable efforts towards forming such a clinic which he hoped would flourish and, like his successful eugenics sterilization law, become the model adopted throughout the States. Eugenicists’ broad support of a national genetics clinic signified their vision to become more directly serviceable and in control of the campaign against disease. Eugenics-trained clinicians could expand and maintain a national database of familial pedigrees as well as provide hereditary counseling services to an increasing number of inquiries from individuals regarding marriagability. Eugenicists viewed the growing demand for such services as the lay public’s acknowledgement of their expert ability to accurately predict the likelihood of disease on future matings. Publications from such a clinic, Laughlin argued, would alert physicians nationwide to the potential hereditary background of many diseases. Through intimate professional knowledge about their patients, physicians who knew the clinic’s latest findings would be even better able to judge the reproductive fitness between potential marriage partners. In this way physicians would become increasingly powerful figures at the forefront of promoting eugenics. A human genetics clinic would also, it was argued, provide an organized interdisciplinary team or network of professionals collectively working towards the betterment of humanity. Moreover, an interdisciplinary approach was viewed as the ideal method to obtain a greater understanding of the hereditary predisposition thought to underlie a multitude of chronic diseases. By leading the charge to reduce the incidence of an increasing number of diseases found to be caused, in part, by hereditary predisposition, the clinic was viewed as central to improving civilization. 88

For examples of the drive to enhance eugenics education within the medical school curriculum, see Jordan (1912) and Davenport (1912).

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With an abundance of pedigree charts at his disposal, Laughlin directed his initial attention regarding a clinic towards adding a genetics component to physician George Draper’s highly successful Constitutional Clinic at Presbyterian Hospital in New York City. Pedigree charts of Draper’s patients were studied with the aim of identifying particular “types” of constitutional makeup. For example, individuals with peptic ulcer were “subjected to study as one group; all with gall bladder disease as another; and those with pernicious anemia, diabetes, acute rheumatic fever, etc., into others.” It was by focusing upon differences in the constitutional makeup between different “types” of patient classes that Draper believed physicians could best build a more accurate perception of the pathogenesis of particular diseases as well as more readily acquaint themselves with the best measures to prevent disease in different constitutional “types.” 89 Appearing not to have forged the connections he desired with Draper’s Clinic, Laughlin gathered more pedigree charts and attempted to gain support for his clinic on another front. As a member of the National Research Council’s Committee on Human Heredity, Laughlin worked to secure Rockefeller Foundation funds to support, among other ventures, the establishment of a genetics clinic. Again, his efforts were rebuffed. At this point, Laughlin divested further efforts to achieve his goal through two different private ventures. With the primary support of investor Wicliffe Preston Draper, Boston Lawyer Malcolm Donald, U.S. Supreme Court Justice, John Marshall Harlan, and president of the American Eugenics Society, Frederick Osborn, Laughlin helped co-found a group in 1937 whose initial plans included the establishment of “The Institute of American Eugenics.” One component of this Institute provided for the foundation and maintenance of a “marriage clinic” to which “persons seriously interested in the inheritance of human racial and family-stock qualities could present their specific problems for advice and information in accordance with the known facts on human heredity.”90 The work of this newly-founded group, who initially considered naming themselves The Eugenics Fund, Inc., or The Genetics Fund, veered in slightly different directions. This group, eventually named the Pioneer Fund, attained many of its initial eugenical goals under Laughlin’s directorship. It failed, however, to garner the support needed to found a eugenics-based clinic. 91 Exemplifying his characteristic fortitude, Laughlin sought yet another pathway to gain support for his clinic. James E. Eddy, founder of the Institute of Forest Genetics in Placerville, California, shared Laughlin’s vision of enabling eugenics to be better used in the fight against chronic disease. What was needed, Eddy argued, was a Clinic of Human Heredity. He discussed his plan with California Institute of Technology geneticist, T.H. Morgan who regarded the idea as “excellent” and suggested that it should be “started and carried out in connection with some established laboratory such as that at Cold Spring Harbor.” He cautioned Eddy to ensure “permanent [financial] support,” arguing that it was one thing to start a clinic of this kind but quite another to “ensure its future support.”92 89 90 91

92

George Draper outlined his overall views of basic human types in a number of publications, among the earliest being “Man as a Complete Organism—In Health and Disease” (1934). Pioneer Foundation, “Notes on Getting the Work Underway”. The Pioneer Fund’s current director, Harry F. Weyher, outlined this organization’s somewhat controversial history in “Contributions to the History of Psychology: CXII. Intelligence, Behavioral Genetics, and the Pioneer Fund” (1998). T. H. Morgan (1938).

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Eddy had also shared Morgan’s views of a Cold Spring Harbor base and approached Laughlin about the possibility of establishing the clinic in close proximity to the ERO. Laughlin quickly joined Eddy’s campaign for a clinic and drafted an “Outline of the Organization, Staff and Service, Proposed for ‘The Clinic of Human Heredity’” which he presented to John C. Merriam, President of the Carnegie Institute in July 1937. The team proposed a 100,000 cubic foot clinic to be situated near the ERO on Carnegie property. The building would consist of the clinic’s headquarters, an office, a laboratory as well as archives and library space for pedigree analysis. Staffing would include “one eugenicist-in-charge,” one investigator “skilled in the diagnosis and measurement of human traits,” one geneticist “skilled in the rules of inheritance of human traits,” one field worker, one secretary-stenographer who would also act as archivist/librarian, and one janitorial caretaker. After an initial $70,000 investment for the building and equipment, the team proposed the clinic could be run on $20,000 per annum. One well organized clinic would, it was argued, become the “model for similar clinics, many of which would be required to serve the whole field effectively, and which . . . could be established in universities, medical schools, . . . social centers, and possibly [stand] independently” as well. Team members anticipated patients (or clients) “who are faced with or who are especially concerned with a particular problem in human heredity” to first “supply the evidence from several near-kin with a description of the presence or absence [of a trait] or the degree of development of the same . . . [or “allied”] trait in each named near-kin.” After analysis of the evidence, the “findings would be reported to the inquirer.” The findings, it was noted, do “not necessarily [need to] include advice,” but they should state “as accurately as possible the behavior of Nature in reference to the inheritance of the particular subject-trait” as well as the “probabilities of the particular trait being transmitted along certain branches of a specified family tree.” The proximity of the clinic to the ERO was suggested because “the world’s stock of knowledge of rules of inheritance of a given trait are on hand and available for critical application to the specific [hereditary] problem.” The team admitted that “hundreds” of such inquiries had already been answered by the ERO, but that with time and labor constraints, the practice was generally discouraged. A “competent clinic” organized to meet such demands from the public and the medical profession was, they concluded, sorely needed. The team considered whether the staff should offer their services to the public for a small fee or in exchange for completed pedigree information. “First-hand pedigrees of human traits” were, it was noted, of “great use to the archives of human heredity.” Such records, when procured by field workers, were claimed to “cost much more” than by acquiring them in exchange for free clinical services.93 Merriam’s support would be critical for the birth of this clinic. His response, however, was not all that Eddy, Goethe, and Laughlin hoped for. Merriam argued that since the clinic would have “as its normal function a broad relation to health and medical problems,” it might best be “connected with a great university hospital or with some independent institution of the hospitalresearch laboratory type.” He acknowledged that the pedigree data assembled by the ERO was indeed “indispensable” for such a clinic, but he envisioned that rather than adding a clinic on their 93

Harry Laughlin Papers, “Outline of the Organization, Staff, and Service, Proposed for ‘The Clinic of Human Heredity’” n.d.

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own grounds, the ERO could better establish links with an existing clinic whereby the latter could “obtain use of [the ERO] materials without serious interference with the progress of the [existing ERO] research program.” He saw such a relationship to be of advantage to both participants but argued it would be wiser to keep the two participants “sharply separate as to responsibilities and administration.”94 Laughlin and his co-organizers discussed establishing ties with The Johns Hopkins Hospital in Baltimore. Despite their persistent efforts along these lines, the Human Hereditary Clinic they envisioned never materialized. On the surface, Carnegie President Merriam’s doubts were never successfully overcome. Furthermore, Merriam retired from his post and was succeeded by Vannevar Bush in January 1939. Laughlin immediately apprised him of the plans for the clinic, but the new president put him off until he became more familiar with overall Carnegie operations. Like an increasing number of medical science the Carnegie Foundation began to distance itself from anything labeled eugenics. The pedigree chart proved to be a valuable tool for the developing field of human genetics in several important ways. It offered a concise and clear way of demonstrating a perceived hereditary linkage regarding a particular disorder or disease. Laughlin’s coordinated gathering and distribution of family pedigree information was designed, in part, for the eugenic attempt to maintain a healthy reproductive stock within the U.S. population. As such, his use of these charts further substantiated the “hardening” that had occurred in beliefs about the nature of heredity during the late nineteenth century. In particular, the regular appearance of these tools strengthened “hard hereditarian” claims that inherited defects and disease were solely dependent upon a non-malleable nature.95 As such, alteration of the reproductive stock of the American people during the Progressive Era became intensely focused upon nature rather than nurture. More investigations remain to be undertaken to more fully appreciate the roles whereby these valuable tools have secured a place of permanence in the field of human genetics.

Philip K. Wilson, Ph.D. Penn State University College of Medicine Hershey, PA [email protected]

94 95

Merriam, John C. (1938). Carlos López-Beltrán (1994) described that this malleable view existed in the “soft hereditarianism” beliefs of the early nineteenth century in contrast to the more objective qualifications of a nature-based, “hard hereditarianism” later in the century.

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References 1920. “Eugenics and Other Sciences.” Journal of Heredity 11: 77-78. 1927. “Family Pedigree Study as College Laboratory Work.” Eugenical News (July) 84. Allen, Garland E. 1983. “The Misuse of Biological Hierarchies. The American Eugenics Movement, 19001940.” History and Philosophy of the Life Sciences 5(2): 105-28. ————. 1986. “The Eugenics Record Office at Cold Spring Harbor, 1910-1940. An Essay in Institutional History.” Osiris, 2nd series 2: 225-64. Asbell, Bernard. 1995. The Pill. A Biography of the Drug that Changed the World. New York: Random House. Banker, Howard J. 1923. “The Ideal Family History.” In Eugenics, Genetics and the Family. Scientific Papers of the Second International Congress of Eugenics. Vol 1. Baltimore: Williams and Wilkins. 306-311. Barkan, Elazar. 1991. “Reevaluating Progressive Eugenics. Herbert Spencer Jennings and the 1924 Immigration Legislation.”Journal of the History of Biology 24(1): 91-112. Binder, Rudolph. 1923. “Health and Eugenics.” Eugenics in Race and State, Scientific Papers of the Second International Congress of Eugenics. Vol.2. 292-297. Bix, Amy Sue. 1997. “Experiences and Voices of Eugenics Fieldworkers. Women’s Work’ in Biology.” Social Studies of Science 27: 625-68. Bruinius, Harry. 2006. Better for All the World. The Secret History of Forced Sterilization and America”s Quest for Racial Purity. New York: Alfred A. Knopf. Bureau of Vital Statistics, State Board of Health. 1924. Eugenics in Relation to the New Family and the Law on Racial Integrity. Richmond, VA: Public Printing. Cantor, David. 2000. “The Diseased Body.” In Roger Cooter and John Pickstone (eds.). Medicine in the Twentieth Century. Amsterdam: Harwood. 347-366. Child, Baron. 1999. Genetic Medicine. A Logic of Diseases. Baltimore, MD: Johns Hopkins University Press. Collins, P. 1913. “The Progress of Eugenics.” Scientific American 109: 459. Conklin, Edwin Grant. 1922. Heredity and Environment in the Development of Men. Princeton: Princeton University Press. Davenport, Charles B. 1901. “Mendel’s Law of Dichotomy in Hybrids.” Biological Bulletin 2: 307-310. ————. 1912. “Eugenics and the Physician.” New York Medical Journal. reprint (8 June). ————. 1915. “Field Work an Indispensable Aid to State Care of the Socially Inadequate.” In Basis for the Joint Employment of Field Workers by the Eugenics Record Office and Institutions for the Socially Inadequate. Cold Spring Harbor, N.Y.: Carnegie Institute of Washington Eugenic Record Office. ———— and Harry H. Laughlin. 1915. “How to Make a Eugenical Family Study.” Eugenic Record Office Bulletin No. 13, Cold Spring Harbor, NY. ————, Harry H. Laughlin, David F. Weeks, E.R. Johnstone, and Henry H. Goddard. 1911. The Study of Human Heredity. Methods of Collecting, Charting and Analyzing Data. Eugenic Record Office Bulletin No. 2, Cold Spring harbor, NY. Downing, Elliot Rowland. 1918. The Third and Fourth Generation. An Introduction to Heredity. Chicago: University of Chicago Press. Draper, George. 1934. “Man as a Complete Organism—In Health and Disease.” New York State Medical Journal 34. Reprint. 1-12. East, Edward M. 1929. Heredity and Human Affairs. New York: Charles Scribner’s Sons. Ferris, M.R. 1929. “Ferris MR. to Harry H. Laughlin.” 8 November. In Institute of American Genealogy Magazine. Folder, box D-4-2:9, Harry Laughlin Papers. Fuller, Edwin M. 1887. “Pedigree in Health and Disease.” Transactions of the Maine Medical Association (Portland). 89-210. Gaudillière, Jean-Paul and Hans-Jörg Rheinberger. 2004. From Molecular Genetics to Genomics. The Mapping Cultures of Twentieth-Century Genetics. London: Routledge. Gilman, Sander L. 1988. Disease and Representation. Images of Illness from Madness to AIDS. New York: Cornell University Press. Goddard, Henry H. 1910. “Heredity of Feeble-Mindedness.” American Breeders Magazine 1(3): 165-178. Gould, Stephen Jay. 1984. “Carrie Buck’s Daughter.” Natural History 93(7): 14-8. Harry Laughlin Papers. 1915. “Brief Instructions on How to Make a Eugenical Study of a Family.” In Eugenics in National Reconstruction; Eugenics Study Instructions. Folder, box D-2-5:2. ————. 1921. “A Pedigree Study of John Burroughs, Naturalist and Man of Letters.” Manuscript. In Pedigree of John Burroughs. Folder, box D-5-1:8. ————. 1932. “Harry H. Laughlin to Norman C. Meier.” 23 March. Chicago Fair-Arrangements Correspondence. Folder, box D-2-4.

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————. 1936. Honorary degree-University of Heidelberg, Germany. Folder, box E-1-3, Harry Laughlin Papers. ————. 1937. “Report of the Archivist of the Eugenics Record Office for the Year September 1, 1936 to September 1, 1937.” Typescript. Classifying Eugenics Material. Folder, box C-2-3:2. ————. 1938. “Classification of Lantern Slides.” In Classification of Lantern Slides. Folder, box C-2-4:6. ————. 1939. “Report on Researches in Eugenics and Heredity.” Manuscript. In Report on Researches in Eugenics and Heredity. Folder, box c-2-6:1. ————. n.d. “A Corn Breeding Experiment. To provide laboratory material for students of human heredity.” In Corn Breeding Experiment. Folder, box D-4-3. ————. n.d. “A Few Points to Observe in Writing up Notes.” In Eugenics Field Worker-Material Used in Classes-1921. Folder, box C-2-4:9. ————. n.d. “Eugenics typescript.” In Eugenics Office Established. Folder, box C-2-3:6. ————. n.d. “Family-Tree Folder.” Forms used by Eugenic Record Office. Folder, box C-4-3:3. ————. n.d. “Ideal Young Man. Professor Laughlin Fixed Per Centage [sic] of Elements of Ideal Character. Unidentified newspaper clipping discussing Laughlin’s address when High School Principal (1900-1905).” Miscellaneous. Folder, box E-1-3. ————. n.d. “Letter to Libraries.” In Ideas for Disseminating Research Materials. Folder, box D-5-2:10. ————. n.d. “Memorandum of Suggestions to Instructors who are Using the Records of Family Traits and the Family Tree Folders of the Eugenics Record Office as Guides in Required Laboratory Work in Human Pedigree Study in Course in Biology, Sociology, and Psychology.” In Eugenics Work for College Women. Folder, box D-5-3:12. ————. n.d. “Outline of Notes for Condensed Statement and Examples of the Principals of Eugenics.” In Eugenics Field Worker Materials used in Classes. 1921. Folder, box C-2-4:9. ————. n.d. “Outline of the Organization, Staff, and Service, Proposed for ‘The Clinic of Human Heredity’.” In Outline of the Organization, Staff, and Service, Proposed for ‘The Clinic of Human Heredity’. Folder, box E-2-2. ————. n.d. “Pedigree to be Charted by Class.” In Summer School Class. Folder, box E-2-1:20. ————. n.d. “Personal notes.” In Corn Breeding Experiment. Folder, box D-4-3. ————. n.d. “Qualities Desired in a Eugenical Field Worker.” In Eugenics Field Worker-Material Used in Classes-1921. Folder, box C-2-4:9. ————. n.d. “Schedule for Recording First-hand Pedigree-data on Hereditary Eye Defect and Blindness.” In Forms Used by Eugenics Office. Folder, box C-4-3:3. ————. n.d. “The Permanent Family Pedigree Archive.” In Eugenics Office Estyablished. Folder, box C-23:6. ————. n.d. “The Pioneer Foundation. n.d. Notes on Getting the Work Underway.” In Draper-OsbornPioneer Fund. Folder, box D-2-3. ————. n.d. “What is Heredity?” In A Problem in Probability . . . What is Heredity? Folder, box D-5-2:1. Hassencahl, Frances J. 1970. Harry H. Laughlin, ‘Expert Eugenics Agent’ for the House Committee on Immigration and Naturalization, 1921 to 1931. Ph.D. diss., Case Western Reserve University. Herrnstein Richard J. and Charles Murray. 1994. The Bell Curve. The Reshaping of American Life By Difference in Intelligence. New York: Free Press. History Files — A Century of Progress. 1998. (28 November 2006. Jackson, Mark. 1995. “Images of Deviance. Visual Representations of Mental Defectives in Early TwentiethCentury Medical Texts.” British Journal of the History of Science 28: 319-337. Jefferis, B.G. and J.L. Nichols. 1922. Safe Counsel or Practical Eugenics. Naperville, Illinois: J.L. Nichols. Johannsen, W. 1911. “The Genotype Conception of Heredity.” The American Naturalist 45: 129-159. Jordan, H.E. 1912. “The Place of Eugenics in the Medical Curriculum.” In Problems in Eugenics, Papers Communicated to the First International Eugenics Congress. London: The Eugenics Education Society. 396-399 Kevles, Daniel J. 1985. In the Name of Eugenics. Genetics and the Uses of Human Heredity. Berkeley: University of California Press. Kimmelman, Barbara A. 1983. “The American Breeder’s Association. Genetics and Eugenics in an Agricultural Context, 1903-13.” Social Studies of Science 13: 163-204. King, Desmond. 2000. Making Americans. Immigration, Race, and the Origins of the Diverse Democracy. Cambridge: Harvard University Press. Laughlin, Harry H. 1899. “School Expositions.” The School Journal: 488-489. ————. 1910. “Thremmatology in the Kirksville Normal School, and the Nature of Eye Color in Man.”

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Missouri School Journal 27: 160 a-d. ————. 1912. “An Account of the Work of the Eugenics Record Office.” American Breeders Magazine 3: 119-123. ————. 1914. “Honors for the Normal School and for Kirksville.” Kirksville Journal, 1 January, Character References, etc. Folder, box E-1-3, Harry Laughlin Papers. ————. 1920. “Eugenical Sterilization in the United States.” Reprinted from Social Hygiene 6(4): 499532. ————. 1921. “The Socially Inadequate. How Shall We Designate and Sort Them?” American Journal of Sociology 27(1): 54-70. ————. 1922. “U.S. House Committee on Immigration and Naturalization.” Analysis of the Metal and Dross in America’s Modern Melting Pot. 67th Congress, 3rd session, series 7-C, 725-831. ————. 1923. The Second International Exhibition of Eugenics. Baltimore: Williams and Wilkins. ————. 1929. “The Progress of American Eugenics.” Eugenics 2(2): 3-16. ————. 1935. “The Eugenics Exhibit at Chicago.” Journal of Heredity 26(4): 155-62. ————. 1936. “The Coil-Spring Properties of Chromosomes.” Genetica 28(5/6): 126-45. ————. 1939. “Special Committee on Immigration and Naturalization Report.” In Immigration and Conquest. New York: New York State Chamber of Commerce. López-Beltrán, Carlos. 1994. “Forging Heredity. From Metaphor to Cause, a Reification Story.” Studies in the History and Philosophy of Science 25: 211-235. Ludmerer, Kenneth M. 1972. Genetics and American Society. A Historical Appraisal. Baltimore: Johns Hopkins University Press. MacDowell, E. Carleton. 1946. “Charles Benedict Davenport, 1866-1944. A Study of Conflicting Influences.” Bios 17: 3-50. McCulloch, Oscar C. 1888. “The Tribe of Ishmael. A Study in Social Degeneration.” Proceedings of the Fifteenth National Conference on Charities and Correction. Buffalo: New York. Merriam, John C. 1938. “Merriam, John C. to James G. Eddy, January 13.” In Correspondence-Eddy Clinic of Human Heredity. Folder, Box E-2-1, Harry Laughlin Papers. Montagu, Ashley. 1959. Human Heredity. Cleveland: World Publising Co. Morgan, T.H. 1938. “Morgan, T. H. to James G. Eddy, January 29.” In Correspondence-Eddy Clinic of Human Heredity. Folder, box E-2-1, Harry Laughlin Papers. Munson, J.F. 1910. “An Heredity Chart.” New York Medical Journal 91: 437-438. Nelkin, Dorothy and M. Susan Lindee. 1995. The DNA Mystique. The Gene as a Cultural Icon. NewYork: W.H. Freeman Paul, Diane B. 1995. Controlling Human Heredity. 1865 to the Present. Atlantic Highlands: Humanities Press. Popenoe, Paul and Roswell Hill Johnson. 1922. Applied Eugenics. New York: Macmillan. Potter, Stephen and Laurens Sargent. 1974. Pedigree. The Origins of Words from Nature. New York: Taplinger. Reilly, Philip R. 1991. The Surgical Solution. A History of Involuntary Sterilization in the United States. Baltimore: Johns Hopkins University Press. Report of the Commission on the Segregation, Care and Treatment of Feeble-Minded and Epileptic. 1911. Persons in the Commonwealth of Pennsylvania. Legislation Pursuant to Joint Resolution, 14 June. Rheinberger, Hans-Jörg and Jean-Paul Gaudillière. 2004. Classical Genetic Research and its Legacy. The Mapping Cultures of Twentieth-Century Genetics. London: Routledge. Rushton, Alan R. 1994. Genetics and Medicine in the United States, 1800 to 1922. Baltimore: Johns Hopkins University Press. Sapp, Jan. 1983. “The Struggle for Authority in the Field of Heredity, 1900-1932. New Perspectives on the Rise of Genetics.” Journal of the History of Biology 16: 311-342. Smith, J. David and K. Ray Nelson. 1989. The Sterilization of Carrie Buck. Far Hills, New Jersey: New Horizon Press. Stafford, Barbara Maria. 1991. Body Criticism. Imaging the Unseen in Enlightenment Art and Medicine. Cambridge: MIT Press. Tufte, Edward R. 2001. The Visual Display of Quantitative Information. Cheshire, Connecticut: Graphics Press. Turney, Jon and Brian Balmer. 2000. “The Genetic Body.” In Roger Cooter and John Pickstone (eds.) Medicine in the Twentieth Century. Amsterdam: Harwood. 399-415. Walter, Herbert E. 1914. Genetics. An Introduction to the Study of Heredity. New York: Macmillan. Watson, Elizabeth L. 1991. Houses for Science. A Pictorial History of Cold Spring Harbor Laboratory. Cold Spring Harbor: Cold Spring Harbor Press.

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Weyher, Harry F. 1998. “Contributions to the History of Psychology. CXII. Intelligence, Behavioral Genetics, and the Pioneer Fund.” Psychological Reports 82: 1347-1374. Wilson, E.B. 1908. “Some Recent Studies In Heredity.” The Harvey Lectures, 1906-1907 (200-222) as cited in A. R. Rushton, 1994. Genetics and Medicine in the United States 1800-1922. Baltimore, MD: Johns Hopkins University Press. 66. Wilson, Philip K. 2002. “Eugenicist Harry Laughlin’s Crusade to Classify and Control the Socially Inadequate in Progressive Era America.” Patterns of Prejudice 36: 49-67. ————. 2003. “Bad Habits from Bad Genes. Early 20th-Century Eugenic Attempts to Eliminate Syphilis and Associated ‘Defects’ from the United States.” Canadian Bulletin of Medical History 20: 11-41. ————. 2006. “Confronting ‘Hereditary’ Disease. Eugenic Attempts to Eliminate Tuberculosis in Progressive Era America.” Journal of Medical Humanities 27: 19-37. Yoxen, E.J. 1984. “Constructing Genetic Diseases.” In T. Duster & K. Garrett (eds.). Cultural Perspectives on Biological Knowledge. Norwood, NJ: Ablex Publishing Company. 41-62. Zenderland, Leila. 1998. Measuring Minds. Henry Herbert Goddard and the Origins of American Intelligence Testing. Cambridge: Cambridge University Press.

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Biohistorical Narratives of Jewish History. Contextualizing the Studies of Wilhelm Nussbaum (1896-1985) Veronika Lipphardt

Towards the end of the 19th century, anthropologists in Germany, Great Britain, France, Russia and elsewhere regarded “the Jews”—persons who were considered to be Jews—as a fascinating object of research. A complex assemblage of social, political and cultural factors led to this focussing of attention. The aim of this paper is to highlight scientific factors: It seeks to demonstrate how scientists began to tell Jewish history in biological terms, in accordance with tremendous theoretical and practical changes in the life sciences around 1900. For physical anthropologists, one difficult problem was to find a well defined group to study. To investigate the essential features of a biologically coherent group, one had to make sure that all persons under study belonged to the group “by nature.” In the light of Darwin’s theories, this meant “reproductive isolation,” a genealogical cohesion for many generations. Anthropologist Bernhard Blechmann from Dorpat was sure that, with the Jews, he had found a group which had doubtlessly been homogenous and isolated for a very long time: It is quite a mysterious fact—and acknowledged by most experts—that the Jewish tribe, from its emergence 4000 years ago up to today, has scarcely undergone any changes, and that no other racial type can be traced back through the millenias with as much precision as the Jewish.1

Commenting on the persistence of the alledgedly “pristine and pure” Jewish race, Blechmann’s remark was—inside and outside the scientific community—hardly unique. In the late 19 th century, Jews were generally seen as hardy, indestructible, obstinate, tenacious, brash, ruthless and resolutely self-serving.2 Jews were considered an isolated group, clinging to conservative traditions, refusing to accommodate and change their way of life, and resisting all kinds of environmental influences. Although pejorative, such stereotypes also resonated ambivalent appreciation of admirable and outstanding characteristics, as, for example, assertiveness—thereby rendering the Jews even more harmful. Although historians have shown in detail how these notions easily spread as a “cultural code of anti-Semitism”3 and were taken up by National Socialists later, their persistenc in scientific practices and discourses has not been traced yet.4 While important studies on Zionism and racial theory have appeared recently5, it is less well known that from the beginning of the 20th century onward the “Jewish race,” or the “biology of the Jews,” was a topic of serious study. Many scientists 1 2 3 4

5

Blechmann (1882) p. 1-2. Driesmans (1912/13) p. 158. Volkov (2002). Touching upon this aspect, see: Efron (1994); Kiefer (1991); Hödl (1997); Gilman (1991); Gilman (1984); Doron (1980); Falk (1998); Hart (1999); Hart (2000), especially Chapt. 4 “The Pathological Cycle;” Essner (1995); Lilienthal (1993); Bacharach (1980). Especially Falk (2006).

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up to the 1930es claimed that the Jews were an isolated, unmixed, and persistent type and thus the “ideal object” for investigating human heredity and race.6 In German-speaking countries, the “biology of the Jews” was discussed by Non-Jewish and Jewish scientists after 1900.7 Among the latter, secular “integrationists,” as I call them, clearly outweighed Zionists and religious Jews, especially after World War I. In addition to theoretical speculations, most scientific publications drew on empirical data and employed all kinds of contemporary techniques, such as anthropometry, statistics, genealogy, psychiatry, pathology, and serology. Contemporary theories of heredity informed many of those investigations. The transferrability of Lamarckian, Darwinian, Weismannian and Mendelian concepts onto the “biology of the Jews” was hotly debated. Indeed, this scientific debate on the “biology of the Jews” became a forum for discussions on human heredity and races. The identity politics and harsh polemics which pervaded this debate are, though certainly a very interesting and important aspect, not the main focus of this paper. Instead, I want to show how concepts of heredity were used in empirical studies on Jews and how methods of genealogy became important tools in this context. Neolamarckism and Neodarwinism are certainly the most obvious concepts to be considered, and indeed they dominated many arguments of the debate. However, I want to concentrate on concepts that were associated with Mendelism, mainly because Mendelism offered empirical approaches as well as theoretical inspiration. After presenting some general thoughts on what I call biohistorical narratives and their representation in the debate, I will discuss the example of Wilhelm Nussbaum who pursued research in the “biology of the Jews” between 1933 and 1935.

Biohistorical narratives and the projection of experiments onto history Investigations and explanations of biological diversity are inevitably accompanied by stories about the historical emergence of the diversity studied. Plants and animals do not take much interest in the stories biologists tell about their ancestry, and they do not leave behind historical records and documents. In the case of human diversity, these stories describe historical events using biological terms. Such stories may be called “biohistorical narratives” because they integrate many historical “facts” with a few biological mechanisms, such as selection, evolution, adaptation, cross-breeding, and environmental influences. Biohistorical narratives are not confined to the domain of science—quite the contrary: They constitute integral elements of the identity building of many nations, families, ethnic groups or other social entities—in addition to or intertwined with other narrative identity constructions. 8 However, since genetics and evolutionary biology have become the predominant source of knowledge on diversity and heredity, most of those rather mystic narratives need to be aligned

6 7 8

Among others: Andree (1881); Blechmann (1882), p. 2; a rather crititical account: Fishberg (1913), S. 9. See also: Hart (1999), p. 270. Lipphardt (2006). The distinction between “Jewish” and “Non-Jewish“ scientists in this paper is in accordance with their self-ascriptions. Recently, many studies on narrative identity constructions from all fields of humanities have been published. Crucial for this trend: Hobsbawm and Ranger (1983); Anderson (1992).

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with modern genetics in order to be consistent with contemporary understanding of “how life works.” To explain why children resemble their parents, families draw on their understanding of modern genetics. To explain, for example, how the early ancestors of modern Europeans became European, geneticists tell stories about historical events that shaped what today we know as ethnic diversity or human populations. While narratives of inheritance—for example in families— concentrate on a few generations and identifiable individuals, narratives of diversity can stretch over centuries and large masses of living beings. Let us return to the so called “biology of the Jews.”9 One reason why the Jews were considered an “ideal object” for research in heredity was their well documented history. Many religious and historical texts, of both Jewish and non-Jewish origin, supplied a great variety of biohistorical narratives about Jews. At the beginning of the 20th century, the following narrative of Jewish history prevailed in scientific publications: Allegedly, the Jewish race had resulted from an ancient cross between three oriental racial types (Amorites, Semites and Hethites) and could thus not be considered a pure race, but a race mixture.10 After the destruction of the Temple, Jews were dispersed throughout Europe—a story of migration into various geographic environments. Because alledgedly they did not intermarry with non-Jewish societies, the ancient race mixture became a “pure stock” that reproduced only within its own community—it was called an “inbred” or even “incestuous” race. Variations between the dispersed Jewish groups were explained as local adaptations, either induced by climate or other environmental factors. In the Middle Ages, so the narration continues, ghetto life had tremendous selective effects upon the biological make-up of this race, and emancipation in modern times obviously was supposed to lead to race mixture and adaptation. 11 According to this narrative, Jews were ideal for testing key concepts of biology: variation, geographic and reproductive isolation, selection and so on. Both Jewish and non-Jewish scientists drew on these narratives, yet by combining single narratives very differently and with different motives and outcomes. The dominant narration given above was subject to minor alterations: For example, in terms of “compatibility,” it made a difference whether the oriental race mixture was regarded as closely related or alien to the “European race mixture.” While Non-Jewish authors tended to blame Jews for voluntary social isolation, Jewish authors explained isolation with the discrimination by the Christian surrounding. Whether the selection effects of the ghetto time had been harmful or enhancing for the genetic make-up was a contentious topic, as well as the question whether the negative genetic effects could be altered or not by exercise, education, and positive environmental influences, such as equal opportunities and social acceptance—or climate and hygiene. 9

10

11

I borrowed this term from Franz Weidenreich, who tried to found a “Wissenschaftliches Institut zur Erforschung der Biologie der Juden” in 1934. Weidenreich Papers, American Museum of Natural History (AMNH). It was Prof. Felix von Luschan who first published on Jews as a “Rassengemisch;” Luschan (1892). The “Rassengemisch”-hypothesis came to be the standard doctrine in the “biology of the Jews” in the beginning 20th century. For an interesting version of the narration, see: Auerbach (1907a); Auerbach (1907b). As PhD-student, Auerbach published an article against the “Rassengemisch”-thesis of his academic teacher Felix von Luschan; later, in the 1920es, he integrated the notion of a “Rassengemisch” into his narration of Jewish history.

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It is barely surprising that Neodarwinism helped to support notions of inalterability and was supposed mostly—but not only—by Non-Jewish authors, while Neolamarckism, as a narrative of personal adaptability and hereditary flexibility, prevailed in the narratives of some Jewish biologists. But there were many exceptions to this rule, and although there might be a tendency towards Neodarwinism among Zionists, as suggested by other authors, the dividing line cannot be drawn easily. Biographical factors played an important role in this context. For a biologist, a biological theory was more than just a working hypothesis: It had to explain all kinds of striking phenomena in his social world as well, especially at a time when biological theory did not provide the coherence of the later Great Synthesis. Certainly, biologists agreed upon some “prominent examples,” serving as “test cases” or “touchstones” for biological theories: the mule as a proof of hybrid infertility, or the Axolotl for Neolamarckian inheritance. Vice versa, such “prominent examples” could be the target of publically performed deconstruction of scientific antagonists. 12 Often, however, “well known” and “obvious” phenomena of human heredity or social life were at the center of those strategies of exemplification. And hence, according to their own social experiences, some authors—from Jewish or Non-Jewish or “mixed” families—considered the example of the Jews a very important test case for any theory of inheritance. It is this specific interaction between personal narratives and scientific theories which was at work in their scientific inquiries.13 Mendelian genetics provided not only new terms, metaphors and theoretical frameworks with which to interprete the history of the Jews, but also an applied empirical methodology. Anthropologists constantly complained about an inherent problem of human genetics: No experiments were allowed, no pure line inbreeding technology applicable. But exactly for this reason, the Jews seemed to allow for an alternative approach. For to investigate the “biology of the Jews” along Mendelian lines, the research object had to meet certain requirements: It had to be a pure line—that is, an inbred group—which had not undergone mixture with other groups. Projects could be designed to demonstrate the inheritance of certain characteristics within this isolated homogenous group or to analyse the results of so-called “bastardisations” between Jews and non-Jews. Either way, it was necessary to take extensive notes on families, pedigrees, inherited characteristics, and any striking feature. Variation within the isolated group was admitted, but considered to be insignificant compared with the enormous differences between Jews and nonJews. For some anthropologists, purity was not necessarily a feature of the group itself: it was rather visible in contrast to other groups.14

12

13

14

For example, the Neo-Lamarckist Kammerer was famous for his amphibian experiments. When a colleague claimed in 1926 that Kammerer had manipulated his specimens, not only Kammerer, but NeoLamarckism in general was discredited. Others considered this example a rather marginal or irrelevant one, not worthy of any serious scientific discussion. At the time, however, “being objective about ‘the Jewish question’” implied an active, scientific approach rather than the so called “ignoring” or “denying” attitude which was suspected to be driven by interest. This runs contrary to today’s intuition to render the latter as the only “serious scientists” and those who were interested in studying the Jews as “pseudoscientists” (which expresses a moral judgement I agree to; but it omits the alledgedly “serious scientists” and their motives from historical investigation, which I find asymmetric). My claim is that in both cases individual factors on the socio-cultural level help to explain the respective attitude. Lenz (1914/15); Auerbach (1920/21); Auerbach (1919); Auerbach (1930); Gutmann (1925).

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The notion of the Jews as a pure and unalterable race turned into an indispensable presupposition for empirical research. Conversely, the operational sequence of Mendelian experiments had been projected onto Jewish history: The supposed reproductive isolation of the Jews led researchers to discriminate the “Jewish line” from the “Non-Jewish line” and render the children of intermarriages as F1-generation. Although both lines showed considerable variation, the distinction seemed much clearer than any distinction between so-called European groups. And that seemed to make sense because European history was, at least if contrasted with intercontinental constellations, narrated as a story of exchange, mutual relations, and kinship between Christians. To give an example of such a projection, I would briefly like to draw on the botanist Redcliffe Salaman’s study of the Jewish facial expression.15 The facial expression of the Jews was generally said to display a melancholic and permanently suffering condition and to be “durchschlagend,” persistent, inheritable, and distinguishable—in Mendelian terms: dominant. To scrutinize these assumptions, Salaman took photos of children from Jewish-Christian couples in Great Britain and showed them to assistants (who were not familiar with his research design), asking whether the photo showed a Jew or a non-Jew. The assistants he had recruited were all Jewish—and Salaman regarded this as methodological advantage: Most of my observers were quite ignorant of the purpose of my examination and of the results I expected, whilst none were conversant with Mendelian or other theories of heredity. All who have assisted me have been themselves Jews and I have noted a distinct tendency on their part to claim, wherever possible, a Jewish type or face for the children they have examined, and also, as I shall show, the results are entirely in the opposite direction, yet what error there is, is distinctly towards increasing the number of supposed Jewish faces in the offspring of mixed marriage.16

In spite of their “tendency,” the assistants identified a large majority of the faces as non-Jewish. Even though they had been looking for Jewish faces, they were unable to detect the “Jewishness” of the “mixed offspring.” Salaman drew the conclusion that “the Jewish facial type […] is a character which is subject to the Mendelian law of Heredity,” and that “the Jewish features have been shown to be recessive to the Northern European.”17 This, in reverse, seemed to prove that the Jews had indeed been an inbreeding group for the longest time, because otherwise the facial expression—according to Salaman—would simply have disappeared over the centuries. […] complex as the origin of the Jew may be, close inbreeding for at least two thousand years has resulted in certain stable or homozygous combinations of factors which react in accordance with the laws of Mendel.18

German reactions to Salaman’s study were very ambivalent.19 On the one hand, rejecting old stereotypes of Jewish persistency, ineradicability and “dominance,” and then claiming “recessiveness” instead, evoked notions of the Jews as being less aggressive and more submissive, 15 16 17 18

Salaman (1911). On Salaman’s study, see Falk (1998). Salaman (1991) p. 280. Salaman (1991), pp. 285, 288. Salaman (1991) p. 290.

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adaptable and integratable. On the other hand, “recessiveness” connoted the subtle danger of hidden and invisible enemies and maneuvers. For Zionists, it may have been difficult to accept such a proof for the “defencelessness” of Jews against “dissolving” into the so-called European population.20 This, again, shows how strongly influenced by personal believes the multiple interpretations of such scientific findings could be. The next subchapter will examine an exceptional and drastic case of those interactions between personal and scientific agenda.

Wilhelm Nussbaum and the “Arbeitsgemeinschaft für Jüdische Erbforschung und Erbpflege” After the Nazis assumed power in 1933, discussions and investigations on the “biology of the Jews” came to an end in Germany.21 However, Wilhelm Nussbaum, a young Jewish gynaecologist who had trained as an anthropologist with Eugen Fischer and Otmar von Verschuer until 1933, 22 set up an institution in Nazi Germany that over the course of eight months investigated more than 1100 Jews. The “Arbeitsgemeinschaft für Jüdische Erbforschung und Erbpflege” was founded in the summer of 1933 and existed until March of 1935 (fig. 1).23 Nussbaum received considerable support from Jewish institutions and was able to conduct his work with the permission of state authorities. The very interesting political implications of this story will be discussed elsewhere; this paper shall concentrate on the way Nussbaum organized his research.

Figure 1. Letterhead of the "Arbeitsgemeinschaft für Jüdische Erbforschung und Erbpflege."

19

20 21

22 23

For the discussion of Salaman’s study of the “Jewish facial expression” and the alledgedly dominant inheritance of the “Jewish type,” see: Fishberg (1913) pp. 176-194; Till (1913); Feist (1925) pp. 187-188; Kaznelson (1913) p. 489; Marcuse (1921) p. 327; Wagenseil (1923/1925) p. 88; Auerbach (1930) column 1178; Michelson (1929) pp. 65-70; Iltis (1930) p. 67. Theilhaber (1911). With exceptions: Walter Dornfeldt, a student of Eugen Fischer, published an anthropological study in 1940 (Dornfeldt 1940). In Austria, anthropologists examined Jews who had been imprisoned before their deportation to concentration camps. In Auschwitz, skeletons of prisoners were collected for research purposes (Rupnow 2006). On Eugen Fischer and Otmar von Verschuer: Lösch (1997); Schmuhl (2005). All references in this subchapter relate to material in the William Nussbaum Collection, Archive of the Leo Baeck Institute, New York. I have worked with the collection in 2004. The collection has since been completely reorganized and not been accessible during that time. It is being microfilmed at present (Feb 2007) and will be accessible in summer 2007. Until then, the location of the material cited in this paper within the collection cannot be provided. The new finding aid is already available online: Guide to the Papers of William Nussbaum (1896-1985), 1773-1975 (bulk 1932-1935), AR 10750, processed by Michael Simonson, October 2006, http://findingaids.cjh.org/?fnm=WilliamNussbaum&pnm=LBI (last accessed 7.3.2007).

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Nussbaum was well aware that managing a scientific undertaking of this sort under such political conditions was quite a challenge. The most important thing for him was an elaborate inscription regime, a “Bezifferungssystem,” as he termed it: “The Bezifferungssystem for such investigations must be simple, manageable and consistent. I have set up such a system. ”24 And indeed, Nussbaum’s system contained preprinted questionnaires and examination forms, pedigrees and report sheets that were filled out for each of the hundreds of probands. With the help of assistants, photos were taken, collected and registered; hundreds of lists, charts, calculations, graphs, tables and diagrams were derived from them; manuscripts and papers summarized findings and implications; and thousands of letters to and from Jews in Germany were sent to gather more information.

Figure 2. Examination form from the Nussbaum Collection.

24

Nussbaum, Wilhelm, manuscript, untitled (“Die erbbiologische Betrachtungsweise…”), undated, 18pp., here: p. 3 (author’s translation).

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The system’s backbone was a central register where each proband was listed by name or by place of origin and supplied with a number. Other registers were derived from the central register, such as a family register or a twin register. Nussbaum’s interest in twins dated back to his PhD-time with Fischer: He had planned to conduct a twin study according to Verschuer’s methodology before 1933. Each proband underwent medical examination and was measured according to anthropometrical standards. The data were recorded on an examination form; of course one was filled out for Nussbaum himself (fig. 2). The blank form, published by Verschuer, sought more than just detailed anthropometric information. It also reported on nutritional and health conditions, illnesses, social status, religion, the form of genitals and so on. On the back, a wide space was left open for “special observations.”25

Figure 3. Handwritten examination form devised by Nussbaum.

25

In Nussbaum’s form, it contains a general description of the bodily condition of the proband; for other probands, psychic pecularities or abnormalities were noted.

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Obviously, the form did not suit Nussbaum’s needs: he devised a hand-written one (fig. 3). On this form, Nussbaum included a special entry for hereditary diseases—in this case it says “neurasthenic”—and three entries for the geographic origin of the proband, one for East European Jews, one for Germans and an empty one for any other. On each form, the register number from the main register was noted. The parents’ register numbers were printed in their name fields (unless the proband was an orphan), and the family and pedigree register number were displayed there too. A large white field was used for manifold purposes in other cases, such as indicating whether the proband was a “Mischling,” a “Langschädel,” or had mental problems. Numbers referring to other documents were also printed in this large field, as, for example, the “group number” which will be explained below. For gynaecological information—menstruation, births, menopause etc.—as well as the proband’s peculiarities and talents Nussbaum included extra fields.

Figure 4. Pedigree form from Nussbaum Collection.

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Certain probands underwent more thorough investigation than others, and additional forms needed to be filled out. There were three different psychogramm forms, one for children and two for adults. There were two very detailed forms for twins. Pedigree forms and “family forms” were filled out for each family (fig. 4). As can be seen from the legend and additional notes, Nussbaum noted any peculiarity which ran in the family—talents, habits, likings and quirks. From pedigrees such as these, he derived trait pedigrees, such as the one for homosexuality (fig. 5) or the one for “recessive enervation” (fig. 6).

Figure 5. Pedigree for homosexuality from Nussbaum Collection.

All the forms were linked with one another by references, numbers and symbols. Each proband could easily be traced through the system by just following the references. Proband groups could be sorted according to many aspects, as for example origin: Berlin Jews and South German Jews, together representing German Jews; and German Jews together with East European Jews made up the European group, in contrast to the Sephardim.

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Figure 6. Pedigree for "recessive ennervation" from Nussbaum Collection.

To analyse this mass of information, Nussbaum devised hundreds of documents which represent a stepwise refining process of information with several intermediary analytical steps, such as sorting, counting, calculating, comparing etc. The table titled “Central European physical characteristics in Jews” was, for example, the result of many calculations for those values Nussbaum had gathered from the probands’ forms; it was meant to demonstrate that Jews were not a foreign race, but had become a central European “Bevölkerungsgruppe” (fig. 7).

Figure 7. Table on "Central European physical characteristics in Jews" from Nussbaum Collection.

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Figure 8. "Gruppenformel" from Nussbaum Collection.

Very curious documents are those that refer to what Nussbaum called the “Gruppenformel” or “Rassenformel.” Nussbaum had selected four characteristics for a special analysis: eye color, hair color, head shape, and face shape. Each character was supposed to occur in two or three variations, and Nussbaum assumed that theoretically there were 36 combinations (fig. 8). Each proband’s examination form was supplied with one of those 36 “Gruppenformel” that reported on his or her specific combination of characteristics. Then Nussbaum analysed which combinations appeared in the various “geographical” groups he had examined, in East European Jews, Sephardim, German Jews etc. He found that certain combinations appeared in none of the groups, others only in one or two groups, and some in all groups. He did what every student of human diversity does: He sought clear lines of demarcation between ethnic groups. Whereas Salaman had sought the historical dividing line between the “pure inbred Jewish race” and the Europeans, Nussbaum sought diversity within the Jewish “Bevölkerungsgruppe.” However, apart from that, he also analysed his data on twins and compared “Jewish twins” and “Aryan twins” (fig. 9).

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Figure 9. Comparison of "Jewish" and "Ayran" twins from Nussbaum Collection.

He also, but not only, tried to determine Mendelian patterns in the inheritance of those characteristics. Other than Salaman, he was not interested in “pure groups,” but rather in the inheritance of single characteristics in families. As many scrawled notes and drafts show, he tried to apply Mendelian methodology whenever he suspected Mendelian ratios to appear (fig. 10). It seems that he sought for an ultimate scientific legitimation for what he was doing, a finding that would have rendered his work valuable in the eyes of famous geneticists, but without success.

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Figure 10. Calculation of Mendelian ratios from Nussbaum Collection.

Beyond these efforts, Nussbaum supported a rather liberal kind of eugenics and examined patients in Jewish charity institutions for blind, deaf and “feeble-minded” persons. One table compared the percentage of those diseases given in the Reichsgebrechlichenzählung in 1925 with an estimation for 1934, allegedly yielding alarming results: The percentage of heterozygotic carriers of those diseases among Jews seemed to increase (fig. 11/12). Nussbaum saw the Nazi segregation laws as the most dangerous threat to the Jewish community: According to him, the new political situation obviously caused more inbreeding than ever before.26 Besides, Nussbaum saw healthy Jewish citizens leaving the country in much higher proportions than those with hereditary defects. Both events, he concluded, increased the occurrence of heritable diseases among Jews in Germany. Nussbaum turned the dominant biohistorical narrative about the Jews upside down: They were now being forced to be the inbreeding group that they had never before been. In order to avert such dangers of degeneration, Nussbaum set up a marriage counselling service for Jews (fig. 13/14).

26

Nussbaum, Wilhelm, manuscript, untitled (“Die erbbiologische Betrachtungsweise…”), u.d., 18 pp., here: p. 16 (author’s translation).

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Figure 11. Table "Erbkranke und Erbbelastete in Deutschland" from Nussbaum Collection.

Nussbaum was also busy publishing articles, giving lectures at Jewish institutions, writing letters to potential supporters, and launching new campaigns in order to get more information, such as the addresses of twins. When he published a call for pedigrees dating back to the early 18 th century, hundreds submitted pedigrees and provided additional information on their families. 27 Nussbaum’s intention was to find out about the geographical distribution of Jews in Central Europe in Early Modern times, but he also offered a certain kind of orientation: Many hoped Nussbaum could help them overcome the shock of the “Ariernachweis”-policy of the Nazis. Their growing interest in genealogy, and their need for positive identification, run parallel to a new interest in Jewish culture among German Jews after 1933.

27

Arthur Czellitzer, a doctor and geneticist, worked along similar lines like Nussbaum and tried to convince German Jews to be proud of their origin, ancestry and heritage; Czellitzer (1934).

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Figure 12. Graph "Erbkranke u. Belastete Juden in Dtsch." from Nussbaum Collection.

Nussbaum wanted to convince Jews in Germany that his work was of outstanding importance and could help the Jewish community. He gained much support by numerous Jewish institutions, such as the Jüdische Frauenbund, the Jüdische Kulturbund and the Reichsvertretung der Deutschen Juden.28 His manuscripts and lectures contain biohistorical narratives taken from scientific discourse, and they powerfully resonated with biohistorical narratives that were embedded in what Nussbaum saw as Jewish traditions, including those of emancipation and integration. Backed with genealogical and anthropological data, he offered a remarkable alternative biohistorical narration: According to Nussbaum, the Jews—an ancient oriental race mixture of three types—had fully adapted to the European surrounding by environmental influences, education and intermarriages since the Middle Ages.29

28 29

Leo Baeck helped Nussbaum with his emigration with a very positive reference letter. For a similar account, see Fishberg (1913).

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Figure 13. Advertisment for eugenic counselling from Nussbaum Collection.

Although some of Nussbaum’s aims and perspectives come out clear from the archival material, it leaves many questions unanswered. The most confusing aspect is probably the rhetoric style of many of his texts: Some passages echo the biopolitical language of the Nazis, and it is hard to imagine that Nussbaum could succeed with this imitation when seeking support from Jewish institutions. On the other hand, he was dependent on the acceptance and support by German authorities as well. It remains unclear whether he tried to avoid censorship, or whether he really embraced that rhetorical mix of Darwinism, genetics and racial theory. He was obviously fascinated by biology, to an extent that might be called religious, but at the same time he seemed to have been aware of the dangerous political situation. It is thus difficult to say what Nussbaum’s intention really was. Certainly, he was convinced that his endeavours were the only salvation available to German Jews. He clearly considered race biology and eugenics to be the best weapons for defending Jews from Nazi persecution. Whether he deemed it possible that his institution could win a sovereign position within Nazi Germany, remains unclear.

Figure 14. Advertisment for eugenic counselling from Nussbaum Collection.

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But apart from that, Nussbaum was also pursuing a number of scientific objectives, some of which were demonstrated above. My suggestion is that Nussbaum did not devise all of these objectives from the outset, but that they changed over time. He surmised how difficult it would be to acquire such a sample in the future, because he saw people leaving the country. Long-term investigations were out of the question. What Nussbaum decided to do was to collect as much information as ever possible in the shortest time-span possible, even if the usefulness of much data remained unclear to him at the time. He recognized how important it was to link all the various inscriptions, in order to acquire a very dense system of information. He assumed that later on, after the examinations were completed, the data would reveal many regularities and “striking effects” of great interest. Nussbaum was obviously aware that the unfinished nature of his project was also the best strategy for coping with his awkward situation: Any challenge to his work, be it from Jewish, state, or any other institutional or private source, could be mitigated because he could provide a convincing biohistorical narrative derived from his data. That data could be interpreted in countless different ways; any of the various steps of analysing data offered new directions for interpretation. For example, Nussbaum gave a lecture on the inheritance of gynaecological traits in twins in London 1934.30 In 1948 he lectured in Brussels on his “Gruppenformel.”31 His inscription system was extensive, flexible, and all-encompassing, but its comprehensiveness would only be appreciated in the future, once methods to interpret the data correctly would become available. I think this “deferred science” is one of the most remarkable aspects about genealogical and many other sorts of inscriptions.32 His project also helped Nussbaum to emigrate and work with Franz Boas, who had been looking for a young German-Jewish anthropologist, trained under his opponent Fischer, to disprove race theories. Boas and the Warburg family supported Nussbaum with a stipend and a research position in New York.33 The data Nussbaum had collected in Germany were integrated into a large-scale research project on bodily conditions of children of various ethnic groups that Boas and Shapiro had initiated in New York in 1935.34 It remains unclear just how serious Nussbaum was about his own rather moderate eugenic views and whether he may simply have used them to avoid conflicts with German state authorities. It was not unusual for German-Jewish doctors to promote liberal eugenics before 1933. However, in a Boasian context, these ideas had no place. After emigration, Nussbaum never wrote about eugenics again. Although Nussbaum’s inscription system resembled the research strategies he had been taught to use at Fischer’s laboratory, Nussbaum modified all the forms to suit his own unique needs. Thus he gathered more, and at the same time more specific information than the published forms of his 30 31 32 33 34

Unpublizierter Bericht über den internationalen Kongreß für Ethnologie und Anthropologie in London. Anthropological Studies on German Jews (1933/34) 16pp., manuscript, undated; printed abstracts of Congress papers. The rhetoric of incomplete data/methodology and hence the deferral of prospective benefits is also among the most consistent characteristics of positivist science. See Franz Boas Papers, American Philosophical Society Archive, Philadelphia, Correspondence BoasNussbaum. See Harry Shapiro Collection, American Natural History Museum, Box 68, folder: Research data, Negroe infants, White infants, Hebrew Orphan Asylum.

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former supervisors allowed. The specific information he sought was designed to account for the unique situation of the probands: Jews in Germany between 1933-1935, a time when biohistorical narratives on Jews were not only part of a cultural code, but also deployed as arguments in the discrimination and persecution of Jews.

Concluding remarks Looking back from the perspective of today, it might seem irritating that Jewish anthropologists joined in the biological debate on Jews, a debate that also circulated narratives compatible with Nazi ideology. John Efron, who has studied Zionist doctors mainly, has argued that participating in the debate was a form of active resistance and self-assertion, and that racial discourse was so universal at the time that, for a trained scientist, no other concepts and terms came into question than those of biology.35 This is a convincing explanation, however, there are more differentiations to be made. Racial discourse was not homogenous; biology provided more concepts than race; and Efron does not explain why Jewish scientists agreed to pejorative accounts of the Jewish people. Furthermore, his argument captures only the socio-political agenda of the scientists he studied, but not their scientific agenda. And it certainly fails to capture the complex and manifold agendas of a non-Zionist like Wilhelm Nussbaum. Michael Bernstein’s concept of side shadowing is helpful in this context.36 In his analysis of literary narrativions of the life stories of Holocaust victims and survivors, Bernstein criticizes what he calls back shadowing: a kind of retroactive foreshadowing in which the shared knowledge of the outcome of a series of events by narrator and listener is used to judge the participants in those events as though they too should have known what was to come.37

With such a narrative strategy, Bernstein criticizes the repression of the “value of the quotidian, the counter-authenticity of the texture and rhythm of our daily routines and decisions, the myriad of minute and careful adjustments that we are ready to offer in the interest of a habitable social world.”38 He suggests to practice a strategy he calls “sideshadowing: a gesturing to the side, to a present dense with multiple, and mutually exclusive, possibilities for what is to come.” 39 Sideshadowing’s attention to the unfulfilled or unrealized possibilities of the past is a way of disrupting the affirmations of a triumphalistic, unidirectional view of history in which whatever has perished is condemned because it has been found wanting by some irresistible historico-logical dynamic. […] Instead of the global regularities that so many intellectual and spiritual movements claim to reveal, sideshadowing stresses the significance of random, haphazard, and unassimilable contingencies [...].40

35 36 37 38 39

Efron (1994). Bernstein (1994). Bernstein (1994), p. 16. Bernstein (1994), p. 121. Bernstein (1994), p. 1.

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Bernstein’s critique suggests that one should not project today’s knowledge about the Holocaust back into pre-1933 Germany. Post-war Historians have, with much success, traced the ideological roots of Nazism back to Enlightenment times. But that does not mean that before 1933, it should have been predictable how those traditions would lead to genocide. What is even more important in this context is the wide frame of possible futures that was open for the imagination of the contemporaries: None of the Jewish scientists who participated in the debate knew what racial biology would be used for under the Nazis, or that biologists would support genocide. Instead, they imagined all kinds of futures for racial biology. For some of them it seemed a promising professional career; or they aimed to prove that, by pure scientific evidence, racism had no future at all. Science itself was full of unassimilable contingencies, had various possible futures, and it was not foreseeable which of those would come to pass. For Nussbaum, many different hopes and aims were connected to his scientific efforts. Even the research design of his study, including Mendelian methods, was set up flexible enough to serve various, contradicting aims he pursued all at the same time. He lived in a “present dense with multiple, and mutually exclusive, possibilities for what is to come,” even if he knew that most of those possible futures were dangerous and unhappy for him and his family. Seen against the backdrop of his situation, it was not opportunistic, but a desperate attempt to make the best of an unpredictable future. Veronika Lipphardt Humboldt-Universität zu Berlin [email protected]

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Bernstein (1994), p. 3-4. “To concentrate on the sideshadowed ideas and events, on what did not happen, does not cast doubt on the historicity of what occured but views it as one among a range of possibilities, a number of which might, with equal plausibility, have taken place instead. The one that actually was realized, though, exists from then on with all the weight afforded by the singularity of what we might call its event-ness.” p. 7.

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Photo Credits All pictures: William Nußbaum Collection, AR 10750, Leo Baeck Institute New York.

References Anderson, Benedict. 1992. Imagined Communities. Reflections on the Origin and Spread of Nationalism. New York. Andree, Richard.1881. Zur Volkskunde der Juden. Bielefeld/Leipzig. ————. 1907a. “Die jüdische Rassenfrage.” Archiv für Rassen- und Gesellschaftsbiologie 4: 332-361. Auerbach, Elias. 1907b. “Bemerkungen zu Fishberg’s Theorie über die Herkunft der blonden Juden.” Zeitschrift für Demographie und Statistik der Juden 3, 6: 92-93. ————. 1920/21. “Rassenkunde (Rubrik: Forschung und Erkenntnis).” Der Jude 5, 1: 49-57. ————. 1919. “Rasse und Kultur.” Neue jüdische Monatshefte 4, 1:11-17. ————. 1930. “Vererbung.” In Jüdisches Lexikon, Vol. 4.2, column 1178-1181. Berlin. Bacharach, Walter Zwi. 1977. “Ignaz Zollschans ’Rassentheorie’.” Jahrbuch des Instituts für deutsche Geschichte 6: 179-197. ————. 1980. “Jews in Confrontation with Racist Antisemitism, 1879–1933.” Year Book of the Leo Baeck Institute 25: 197–219. Bernstein, Michael A. 1994. Foregone Conclusions. Against Apocalyptic History. Berkeley. Blechmann, Bernhard. 1882. Ein Beitrag zur Anthropologie der Juden. Dorpat. Czellitzer, Arthur. 1934. Mein Stammbaum. Eine genealogische Anleitung für deutsche Juden. Berlin. Dornfeldt, Walter. 1940. Studien über Schädelform und Schädelveränderung von Berliner Ostjuden und ihren Kindern. Berlin. Doron, Joachim. 1980. “Rassenbewusstsein und naturwissenschaftliches Denken im deutschen Zionismus während der wilhelminischen Ära.” Jahrbuch des Instituts für deutsche Geschichte 9, Beiheft 10: 388–427. Driesmans, Heinrich. 1912/13. “Zur Biologie der jüdischen Rasse.” Politisch-Anthropologische Revue 11: 149159. Efron, John. 1994. Defenders of the race. Jewish doctors and race science in fin-de-siècle Europe. New Haven. Essner, Cornelia. 1995. “Antisemitismus, Rassenforschung und Judentum im Deutschen Kaiserreich.” In Nora Goldenbogen et. al. (ed.). Hygiene und Judentum. Dresden. Falk, Raphael. 1998. “Zionism and the Biology of the Jews.”Science in Context 11, 3–4: 587–607. ————. 2006. “Zionism, Race and Eugenics.” In Geoffry Cantor,and Marc Swelitz (eds.). Jewish Tradition and the Challenge of Darwinism. Chicago. 137-165. Feist, Sigmund. 1925. Stammeskunde der Juden. Leipzig. Fishberg, Maurice. 1913. Die Rassenmerkmale der Juden. München. Franz Boas Papers. PP B:B61. American Philosophical Society. Correspondence with William Nussbaum. Franz Boas Papers. PP B:B61. American Philosophical Society. Correspondence with Franz Weidenreich, letter Weidenreich to Boas, 12.5.1934. Gilman, Sander. 1984. “Jews and Mental Illness. Medical metaphors, Antisemitism and the Jewish Response.” Journal of the History of Behavioral Sciences 20: 150–159. ————. 1991. The Jew’s Body. New York/London. Gutmann, Moses Julius. 1925. “Die Krankheiten der Juden.” Zeitschrift für Demographie und Statistik der Juden N.F. 2: 51-54. Harry L. Shapiro Collection. American Museum of Natural History (AMNH). Research Library, Special Collections, Box 69, Folders “Michelson & Nussbaum’s material”. Hart, Mitchell B. 1999. “Racial Science, Social Science, and the politics of Jewish Assimilation.” Isis 90: 268– 297. ————. 2000. Social Science and the Politics of Modern Jewish Identity. Stanford Hobsbawm, Eric and Terence Ranger. 1983. The Invention of Tradition. Cambridge: Cambridge University Press. Hödl, Klaus. 1997. Die Pathologisierung des jüdischen Körpers. Antisemitismus, Geschlecht und Medizin im Fin-de-siècle. Wien. Iltis, Hugo. 1930. Volkstümliche Rassenkunde. Jena. Kaznelson, Paul. 1913. “Über einige ‚Rassenmerkmale’ des jüdischen Volkes.” Archiv für Rassen- und

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Gesellschaftsbiologie 10: 484-499. Kiefer, Annegret. 1991. Das Problem einer “Jüdischen Rasse“. Eine Diskussion zwischen Wissenschaft und Ideologie (1870–1930). Frankfurt a.M. Lenz, Fritz. 1914/15. Review of Semi-Kürschner (Berlin 1913). Archiv für Rassen- und Gesellschaftsbiologie 11: 546. Lilienthal, Georg. 1993. “Die jüdischen ’Rassenmerkmale’. Zur Geschichte der Anthropologie der Juden.” Medizinhistorisches Journal 28: 173–198. Lipphardt, Veronika. 2006. Biowissenschaftler mit jüdischem Hintergrund und die ‚Biologie der Juden’. Debatten, Identitäten, Institutionen. PhD thesis, Humboldt University: Berlin. Lösch, Niels. 1997. Rasse als Konstrukt. Leben und Werk Eugen Fischers. Frankfurt a.M. Luschan, Felix von. 1892. “Die anthropologische Stellung der Juden.” Correspondenzblatt der Deutschen Gesellschaft für Anthropologie 23: 94-100. Marcuse, Max. 1921. “Rassen- und Krankheitsfrage bei den Juden. Bemerkungen zu der Schrift von M.J. Gutmann”. Zeitschrift für Sexualwissenschaft 7: 326-329. Michelson, J. 1921. “Die jüdische Rasse in der modernen Anthropologie.” Monistische Monatshefte 1929: 6570. Rupnow, Dirk. 2006. “Antijüdische Wissenschaft im “Dritten Reich”. Wege, Probleme und Perspektiven der Forschung.” In Simon Dubnow-Institut Jahrbuch/Yearbook 5. Salaman, Redcliffe N. 1911. “Heredity and the Jew.” Journal of Genetics 1: 273-292. Schmuhl, Hans-Walter. 2005. Grenzüberschreitungen. Das Kaiser-Wilhelm-Institut für Anthropologie, menschliche Erblehre und Eugenik 1927-1945. Göttingen. Theilhaber, Felix. 1911. Der Untergang der deutschen Juden. München. Till, Eberhard. 1913/14. “Review of Fishberg, Maurice. Die Rassenmerkmale der Juden, München 1913.” Politisch-Anthropologische Revue 12: 516-527. Volkov, Shulamit. 2002. Antisemitismus als kultureller Code. München. Wagenseil, Fritz. 1923/25. “Beiträge zur physischen Anthropologie der spaniolischen Juden und zur jüdischen Rassenfrage.” Zeitschrift für Morphologie und Anthropologie 23: 33-150. Weidenreich Papers. American Museum of Natural History (AMNH), Research Library, Special Collections, Box I, Research proposal. William Nussbaum Papers. Archive of the Leo Baeck Institute, New York. Guide to the Papers of William Nussbaum (1896-1985), 1773-1975 (bulk 1932-1935), AR 10750, processed by Michael Simonson, October 2006, http://findingaids.cjh.org/?fnm=WilliamNussbaum&pnm=LBI (7.3.2007).

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William Bateson’s Pre- and Post-Mendelian Research Program in ‘Heredity and Development’ Marsha L. Richmond

The rediscovery of Mendel’s work in the spring of 1900 sparked a flurry of new activity in biology. Within a short time, many researchers all over the world began breeding experiments to see whether this new approach might be the long-sought breakthrough in understanding the basis of heredity. William Bateson (1862-1926) was in the vanguard of these investigators. Having studied variation for over a decade as a means of studying evolutionary change, Bateson remarked, soon after reading Mendel’s 1865 paper, that “we are in the presence of a new principle of the highest importance,” only a year later to proclaim that because of this work “the whole problem of heredity has undergone a complete revolution” (Bateson, 1900, 60; Bateson and Saunders, 1902a, 4). Although certainly not everyone shared this view, with hindsight Bateson’s prediction seems prescient indeed. Mendel did in fact revolutionize modern biology. Historians of genetics have long agreed that the year 1900 marked a significant watershed in understanding the problem of heredity. Indeed, the literature on this period predominantly focuses on the period after Mendel’s rediscovery and particularly the decade after 1910, which witnessed the rise of “Morganian genetics,” or the Mendelian chromosome theory of heredity. Yet, in order to appreciate the full extent of the impact Mendel’s work exerted on early twentieth century biology, it is helpful to have a better understanding of the kind of work on heredity that was being pursued immediately prior to the rediscovery. That is, we need to compare and contrast the work on heredity in the pre-Mendel years with that in the immediate post-Mendel period. Bateson is a particularly good subject for such a comparison, given that he was arguably the one who did more to promote Mendelism in the English-speaking world than any other biologist. Contrasting his 1890s publications with those after May 1900 thus provides a particularly good indication of the importance of Mendel’s rediscovery: it easily allows us to see the ways in which Mendel’s work transformed Bateson’s previous inchoate research program in “variation” into a targeted study of the “physiology of heredity,” or what in 1906 he christened “genetics” (Bateson, 1907, 91). A brief (and albeit somewhat schematized) sketch of Bateson’s situation circa 1900 helps clarify this claim. Although not one of the “re-discoverers” of Mendel, Bateson was perhaps better positioned to appreciate the significance of Mendel’s laws of heredity than were either Hugo de Vries, Carl Correns, or Erik von Tschermak (Kottler, 1979; Lenay, 2000; Saha, 1984; Rheinberger, 2000; Stamhuis, Meijer, and Zevenhuizen, 1999). For more than a decade he had been focusing on discontinuity in nature, cataloguing cases of variation that represented alternative rather than gradualistic change. In 1894 he published an encyclopedic compendium of notable cases of variation in his Materials for the Study of Variation (Bateson, 1992). The next year he began a new phase of this project, undertaking an experimental investigation of variation through carrying out hybrid crosses. For five years he collected data without discovering any patterns or underlying process that could explain his findings. When directed to Mendel’s paper, Bateson at once realized that the laws of heredity Mendel formulated based on the inheritance of discrete character-pairs

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in hybrid crosses could explain his own findings of the persistence of alternate characters rather than their blending. Bateson thus became an immediate and ardent convert to Mendelism. He changed his experimental design to reflect Mendel’s method of crossing and back-crossing, and began reinterpreting his previous results in the light of “Mendelian” analysis. He also began to proselytize, trying to entice many more workers to become committed disciples. In this he was successful: by 1906, Bateson was recognized as a leading figure in the new field, serving as the head of an active band of followers, not simply in Cambridge but in England and abroad (Olby, 1985, 2004; Falk, 1995). There is, however, a certain irony in this turn of events. Prior to 1900 Bateson operated on the fringe of British biology. He was somewhat of a pariah in Cambridge, the seat of Balfourian postDarwinian evolutionary morphology, not so much because he abandoned the search for phylogenetic progenitors but because he so brazenly regarded this approach as worthless (see Ridley, 1985; Geison, 1978; Blackman, 2003; and Hall, 2004). He also was regarded as a maverick among the Darwinians owing to his well publicized views about the discontinuous basis of evolutionary change and vocal challenge to the all-sufficiency of natural selection. Championing Mendel’s laws served to propel Bateson from the periphery of British biology to a place within its inner circle, gaining not just national but also international prominence (Lock, 1906, viii). Just how did such a remarkable transformation—in both theory change as well as scientific stature— come about? A full-scale biography of Bateson remains a great desideratum in the history of genetics. Nonetheless, there is an abundant corpus of literature that leaves few aspects of Bateson’s career untouched. One area that remains hazy, however, is the work he did during the interregnum between the publication of Materials for the Study of Variation and the rediscovery of Mendel. This is precisely the focus of the present paper, which aims to examine the work Bateson carried out in the 1890s and compare it with that conducted during the first two years after finding Mendel. It aims to gauge critical changes that occurred in Bateson’s technical procedures, standards of analysis, and problem orientation in order to assess more fully the changing architecture of knowledge that marked his dramatic shift from a study of variation to the new Mendelian program in “genetics.” In so doing it highlights the importance of his long-time scientific collaboration with women biologists, which has not to date been sufficiently appreciated. Contrasting Bateson’s study of variation before and after 1900, I argue, allows us better to recognize the continuities with previous practice as well as new modes of conceptualization wrought by Mendel, and to highlight the main features that shaped the new epistemic space Bateson carved out for the new field of genetics.

Bateson’s Study of Variation THE INFLUENCE OF WILLIAM KEITH BROOKS Bateson, by his own admission, noted that his interest in studying the origin of variation was sparked by his association with the American zoologist William Keith Brooks during the summers they worked together in 1883 and 1884. In his contribution to Brooks’s memorial volume, Bateson

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provided an insightful profile of how Brooks’s rather novel understanding of variation influenced his own thinking and, by implication, his subsequent career in biology. It is worth quoting this passage at length: For myself I know that it was through Brooks that I first came to realize the problem which for years has been my chief interest and concern. At Cambridge in the eighties morphology held us like a spell. That part of biology was concrete. The discovery of definite, incontrovertible fact is the best kind of scientific work, and morphological research was still bringing up new facts in quantity. It scarcely occurred to us that the supply of that particular class of fact was exhaustible, still less that facts of other classes might have a wider significance. In 1883 Brooks was just finishing his book “Heredity,” and naturally his talk used to turn largely on this subject. He used especially to recur to his ideas on the nature and causes of variation, and to the conception which he developed in “Heredity,” that the functions of the male and female germ cells are distinct. The leading thought was that which he expresses in his book (p. 312) that “the obscurity and complexity of the phenomena of heredity afford no ground for the belief that the subject is outside the legitimate province of scientific enquiry.” He deplored the fact that he had no opportunity for the requisite experiments in breeding, but he saw plainly that such experiments were the first necessity for progress in biology. To me the whole province was new. Variation and heredity with us had stood as axioms. For Brooks they were problems. As he talked of them the insistence of these problems became imminent and oppressive. It all sounded rather inchoate and vaporous at first, intangible as compared with the facts of development which we knew well how to pursue, but with the lapse of time the impression became strong that Brooks was on the right line. That autumn I went home feeling that though in technique we were a long way ahead of Johns Hopkins—I had the pleasure of showing off the Jung microtome, then the latest thing in progress, to the admiring Baltimore men—yet somehow Brooks had access to novelties of a more serious description. (Bateson, 1910, 6-7; see also Bateson, 1922, 55-56)

Although a professor of morphology, Brooks looked to variation for clues about the workings of evolution. Written only two years after Darwin’s death, Brooks’s 1884 book, The Law of Heredity. A Study of the Cause of Variation, and the Origin of Living Organisms was not only dedicated to Darwin but paid homage to The Variation of Animals and Plants under Domestication (1868). Indeed, Brooks offered a revision to Darwin’s theory of heredity, to counter objections to pangenesis and reflect recent research in cytology (Brooks, 1883; Benson, 1979). Brooks noted that Darwin’s theory of pangenesis, which posited that the ovum contains “not the perfect animal in miniature, but a distinct germ for each distinct cell or structural element of the adult,” had been undermined empirically by Francis Galton and theoretically by Lamarckians, and hence required modification (Brooks, 1883, 78). Galton had tested pangenesis experimentally by infusing rabbits with the blood of different varieties (and hence, presumably, containing different pangenes) and found they were not transformed as was to be expected on the theory of pangenesis (Pearson, 1914-30, 2: 160-73; Gillham, 2001). For their part, Lamarckians favored more rapid response to environmental change than natural selection operating to alter pangenes seemed to allow. Brooks characterized his theory as lying “midway between that accepted by Darwin and that advocated by Semper and other Lamarckians,” and thus offering a good compromise. “If the hypothesis of pangenesis could be so remodelled,” he wrote, “as to demand

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the transmission of only a few gemmules from the various parts of the body to the reproductive elements, instead of the countless numbers which are demanded by the hypothesis in its original form, we should escape many of the objections which have been urged against it” (Brooks, 1883, 80). These few gemmules, he assumed, would be collected by the ovum, which thus served a conservative role in heredity. Variation and adaptation arose from the male element’s “peculiar power to gather and store up germs,” allowing for faster response to changes in the environment (Ibid., 82, 84-85). Hence Bateson’s statement that for Brooks “the functions of the male and female germ cells are distinct.” Brooks obviously supported a “particulate” view of heredity suggested by pangenesis, pointing to his view of “the egg as containing material particles of some kind to represent each of the hereditary congenital peculiarities of the race.” Yet in understanding how variation could come about, he referred to the “physicalistic” view of forces operating on matter proposed by St. George Jackson Mivart (1827-1900), quoting at length from Mivart’s On the Genesis of Species (1871): It is quite conceivable that the material organic world may be so constituted that the simultaneous action upon it of all known forces, mechanical, physical, chemical, magnetic, terrestrial and cosmical, together with other as yet unknown forces which probably exist, may result in changes which are harmonious and symmetrical, just as the internal nature of vibrating plates causes particles of sand scattered over them to assume definite and symmetrical figures when made to oscillate in different ways by the bow of a violin being drawn along their edges. The results of these combined internal powers and external influences might be represented under the symbol of complex series of vibrations (analogous to those of sound and light) forming a most complex harmony or a display of most varied colors. . . . Also as the atoms of a resonant body may be made to give out sound by the juxtaposition of a vibrating tuning-fork, so it is conceivable that the physiological units of a living organism may upset the previous rhythm of such units, producing modifications in them—a fresh chord in the harmony of Nature—a new species. It seems probably, therefore, that new species may arise from some constitutional affection of parental forms—an affection mainly if not exclusively of their generative system. (Ibid., 87)

Qualitative change in the gemmules thus ultimately derived from forces operating on matter to cause some kind of physical rearrangement. Brooks indicated that his view of variation well accorded with Mivart’s: “a new variation is caused in essentially the manner which Mivart suggests as probable. The accumulated influence of surrounding conditions, organic and inorganic, does upset the previous rhythm of the physiological units of the living organism, and causes them to give rise to gemmules, and the tendency of the corresponding units of the offspring to vary, is directly due to this constitutional affection of the parental forms” (Ibid.). This way of conceptualizing variation had major implications for the understanding of species change. If variation arose from environmental changes affecting the physiological activity of the cells of an organism and causing them to throw off altered gemmules, then the possibility of sudden, abrupt changes of form was likely. This presented a challenge to the assumption of gradualistic evolution. As Brooks stated: There are many reasons for believing that variations under nature may not be so minute as Darwin supposes, but that evolution may take place by jumps or saltations. According to our

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view a change in one part will disturb the harmony of related parts, and will cause their cells to throw off gemmules. A slight change in one generation may thus become in following generations a very considerable modification, and there is no reason why natural selection should not be occasionally presented with great and important saltations. (Ibid., 328)

In his book, Brooks, then, despite paying homage to Darwin, offered significant revisions to the tenets of Darwin’s theory of evolution and hereditary theory of pangenesis (see Endersby, 2003). Ten years later, Bateson joined suit, also by means of publishing a work dedicated to studying “the nature and causes of variation.” FROM MORPHOLOGY TO VARIATION Historians have mainly focused on Bateson’s morphological training as having promoted his deviation from neo-Darwinian gradualistic evolution. As Peter Bowler stated, “the particular research problems that had engaged Bateson as a morphologist must have predisposed him to see discontinuous variation as more important” (Bowler, 1992, xxi). Yet it was shortly after working with Brooks at the Chesapeake Zoological Laboratory that Bateson decided to change his line of morphological investigation, abandoning the search for ancestral forms he had previously pursued. In taking up Brooks’s suggestion that studying variation was a better means of understanding the process of evolution than tracing phylogenetic relationships, he explicitly rejected the morphological research program that Francis Balfour had championed at Cambridge (Olby, 2004; Hall, 2005). This had serious implications for his career, both in terms of altering his scientific methodology as well as hindering his ability to garner institutional and financial support for his work. In 1886 and 1887, Bateson undertook a research trip to the Aral Sea and Egypt, with the aim of investigating “the relation between the variations of animals and the conditions under which they live” (Bateson, 1889). He was disappointed to find no clear-cut evidence to support this claim. Rather, the shells of the mollusk Cardium edule, living in the brackish water of the Aral Sea, presented only slight differences from the forms inhabiting the much saltier water of the Mediterranean. He regarded the results of this study as negative evidence, showing “that, while such variations do occur in certain species, in the majority they do not.” (B. Bateson, 1928, 35). According to his wife Beatrice Bateson, “[h]e always regarded these expeditions as failures,” and yet ultimately as putting him on the right road after all: they “proved very stimulating: he had to make good: if he had followed a false clue, the greater the need to find the right one” (Ibid., 27). Bateson’s sense of failure reinforced a growing conviction that variation was not produced by the affect of external stimuli acting on organisms, but rather by the operation of some unknown mechanism internal to the organism, an idea, as we have seen, that was not dissimilar to Brooks’s views. As Bowler points out, the germ of this conception was already apparent in Bateson’s oftcited 1886 work on the ancestry of the chordata. There he identified a tendency in nature toward the repetition of parts, by which means he explained the origin of (segmented) chordates from an (unsegmented) invertebrate, Balanoglossus. “The duplication of an existing structure was selfevidently a discontinuous process,” Bowler noted, “and he suspected that it would occur whether or not the results were useful. In effect, evolution would be driven by a process arising from within the organism, forcing the species to evolve in a certain direction whatever the environment to

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which it was exposed” (Bowler, 1992, xix; Hall, 2005). Although Bateson made no explicit mention of Brooks’s and Mivart’s ideas about the basis of variation, he was obviously moving toward looking internally rather than externally for causal factors prompting organic change. By his own admission, upon concluding this study Bateson experienced an epiphany, a kind of personal “revolt from morphology.” As he described it: “On finishing these investigations I became dissatisfied with this mode of attacking biological problems and resolved to seek a new field of inquiry.” As a result, some time in 1885 he embarked on “work of an entirely different kind” (B. Bateson, 1928, 31). The main problem he set for himself was to identify “the nature of the forces by which the forms have been produced and fixed. . . . Hence, if we seek to know the steps in the sequence of animal forms, we must seek by studying the variations which are now occurring in them, and by getting a knowledge of the modes of occurrence of those variations and, if possible of the laws which limit them” (Ibid., 33-34). He increasingly came to believe in “the Discontinuity of Variation” as a “new point of attack on the problem of Species,” and conceived of variation from a physicalistic standpoint not unlike that expressed by Mivart and Brooks. The underlying elements of his thinking moved him in certain directions, “from observation of Discontinuity to Meristics, Symmetry and the Repetition of Parts, and to tentative suggestions of Rhythm, which he never put aside from his considerations of the forms of life” (Ibid., 56). These were topics present in all of Bateson’s publications in the 1890s. One of the first fruits of Bateson’s new line of work on variation appeared in 1890, in a short paper entitled “On Some Cases of Abnormal Repetition of Parts in Animals.” This brief description of cases of “abnormal repetitions of normal structures” reveals his acute interest in meristic phenomena, or repetitive patterns in nature. While apologizing for the descriptive nature of this work, he nonetheless pointed out its relevance, stating that “the key to some of the problems of variation is to be sought by an analysis of this class of facts, yet such an analysis can only be attempted after a wide survey of the whole ground” (see Bateson, 1928, 1: 113-23). Another paper presented the same year to the Cambridge Philosophical Society continued with this theme. He exhibited a number of insect specimens he had collected and reported on “about 220 recorded cases of extra legs, antennae, palpi or wings, and particulars” in various species of insects. He concluded by offering his views about the “mode of occurrence of these structures,” but unfortunately these remarks were omitted from the published abstract of his talk (Ibid., 1: 125). Until this point Bateson had kept pretty close to the vest about the new conceptual reorientation prompted by his new direction of research. By 1891, however, it was clear that he was beginning to deviate both from mainstream morphology and also from core neo-Darwinian tenets. The first public admission of his new “epistemic space” came in a seemingly descriptive paper discussing cases of floral symmetry. Coauthored with his sister Anna Bateson (1863-1928), a graduate in botany at Newnham College, Cambridge, this paper discussed the irregular forms of corollas (the petals of flowers) in four species, Linaria spuria, Veronica buxbaumii, Gladiolus hybrids, and Streptocarpus. Despite its seeming descriptive nature, this paper in fact harbored well considered general reflections on the nature of variation and its role in evolutionary change. The authors began by focusing on variation without discussing its cause, noting that such an attempt should wait “until a much fuller knowledge of the modes of Variation shall have been

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attained.” They did, however, address the evolutionary significance of irregular corollas, stating that such cases appeared to be connected “with their adaptation to the purposes of crossfertilisation, and that their perfection and persistence have consequently been achieved by the agency of Natural Selection” (Bateson and Bateson, 1891, 386). While granting a significant role to natural selection, they did not believe selection operated on a series of small variations, resulting in gradual evolution. Rather, they envisioned the kind of variation important in evolutionary change as discrete and discontinuous in nature, as opposed to the continuous variation the neoDarwinians assumed. They indeed speculated about the kind of processes that might result in discontinuous as opposed to gradual variation: The success of any attempt to comprehend the nature of the forces which are at work in the production of Variation will depend very largely on the precision with which we shall be able to answer these questions [about whether the series of ancestors of new forms is continuous or discontinuous], and to determine the degree of continuity which is present in the process of Evolution. For if, on the one hand, the transition from form to form shall be found to occur by insensible and minimal changes which are so small that no integral change can ever be perceived, we should recognise an analogy with the continuous action of mechanical forces; but if it should appear that the series is a discontinuous one, and that there are in it lacunae which are filled by no intermediate form, the analogy would rather hold with the phenomena of chemical action, which is known to us as a discontinuous process, leading to the formation of a discontinuous series of bodies, and depending essentially on the discontinuity of the properties of the elementary bodies themselves. (Bateson and Bateson, 1891, 387-88)

Evolution, in other words, may result from the selection of discontinuous rather than continuous variation. In proposing a kind of causal juxtaposition of continuous variation as based on the operation of mechanical forces versus discontinuous variation derived from chemical combination, the authors seemingly deviated from both Brooks’s and Mivart’s more morphological emphasis on physical forces as the source of organic variation. They found the idea of elements combining in different proportions to produce discrete chemical compounds a better analogy for understanding sudden qualitative change in living forms. In this they were unusual, for few biologists in the late nineteenth century speculated about the chemical basis of heredity; rather, following Darwin, most thought in terms of some kind of particulate inheritance (Robinson, 1979). Throughout his career, however, Bateson often invoked chemical analogies to illustrate his views about hereditary change.1 Continuing along these lines, if it were found, the authors reasoned, that many variations were indeed discontinuous, then “the necessity for supposing each structure to have been gradually modelled under the influence of Natural Selection is lessened, and a way is suggested by which it may be found possible to escape from one cardinal difficulty in the comprehension of Evolution 1

See, for example, Bateson and Saunders, 1902b, 147: “Remembering that we have no warrant for regarding any hereditary character as depending on a material substance for its transmission, we may, with this proviso, compare a compound character with a double salt, such as an alum, from which one or other of the metals of the base can be dissociated by suitable means, while the compound acid-radicle may be separated in its entirety, or again be decomposed into its several constituents. Though a crude metaphor, such an illustration may serve to explain the great simplification of the physiology of heredity to which the facts now point.”

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by Natural Selection” (Bateson and Bateson, 1891, 388). The “difficulty” to which they refer was the origin of complex structures like alternative forms of flowers, and specifically the problem of explaining the selective value of incipient stages of useful structures. The question of what adaptive advantage a new character would represent at its earliest stage of formation that could trigger natural selection was first posed by Mivart in Genesis of Species (1871) and answered by Darwin in the 6th edition of Origin of Species (1872; Gayon, 2003, 24344). The Batesons returned to this issue in the context of flower morphology, noting, “it cannot be supposed that the mechanism was at all periods of its evolution so beneficial as to be selected . . . in short, that the evolution of a special contrivance for adaptation is not compatible with constant and perpetual usefulness.” Rather, in all the cases of irregular corollas they analyzed the new form appeared to be an instance of a “sudden variation” rather than a gradual change from one form to another. Moreover, unlike meristic phenomena, in species exhibiting flowers with irregular corollas, the “change of symmetry [was] attained not by an alteration in the number of parts, but by the selection of a different morphological plane about which the symmetry is developed.” Such a change could only have come about by a sudden alteration in the basic morphology of a plant’s petal structure. “It is easy,” they claimed, “to conceive the steps between forms differing in the degree of expression of some character, such as size or intensity of coulour, but in trying to pass from a form with one kind of symmetry to a form with another we often cannot even conceive the transitional steps.” (Bateson and Bateson, 1891, 388-89; 417). What, they implied, could be the possible selection value to cause a 2-petaled flower symmetrical form to generate a 3-petaled asymmetrical flower? (See Figure 1.) In this way the Batesons thus declared their allegiance to discontinuous variation as the predominant force in evolution. Darwin, they noted, was well aware of forms that had characters that did not blend. “Our object now is to show,” they stated, “that this principle is widely true of variations which are of the nature of specific changes, and to point out that it may help us to measure the size of the integral steps of Variation” (Bateson and Bateson, 1891, 419-20). Such a position was not only at odds with contemporary neo-Darwinian tenets, but also challenged the core ancestrian view of the biometricians. Galton’s law of ancestral heredity posited that individuals inherited characters proportionally from their remote ancestors as well as their parents. Thus the well known phenomenon of “reversion” could be explained as the reappearance of ancestral traits (Gayon, 1998, chap. 4). Both Galton and W. F. R. Weldon, the authors noted, had recently analyzed forms exhibiting continuous variations in size, Galton studying human stature and Weldon the proportional sizes of the limbs of shrimp. Both attempted to test the applicability of Galton’s “Law of Error,” according to which “the greater the departure from the normal form, the rarer will be the Variation” (Bateson and Bateson, 1891, 420-21; Galton, 1889; Weldon, 1890). However, such an explanation could not well apply to the cases of floral variation they were considering, which were not “instances of reversion to an ancestral type.” Indeed, under their view, they noted, “[i]t is likely that the study of Variation will hereafter lead to and necessitate a revision of the whole question of the nature of Reversion, but this is no part of our purpose at present” (Bateson and Bateson, 1891, 415). Reversion, considered from a discontinuous point of view, rather suggested the persistence of latent traits, not ancestral reminiscence.

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Figure 1. Variations in Floral Symmetry. Table from Bateson and Bateson 1891, Plate LI.

This line of thought led them to another “fact of great importance,” namely, that “there are at least two classes of Variation,” those representing a “Variation in kind,” such as the cases of floral symmetry they studied, and other “Variations in degree,” like the phenomena investigated by Galton and Weldon. Variations in kind were discrete and qualitative, exhibiting no transitional forms. There must, therefore, be some underlying mechanism accounting for this property. “It seems, in fact, in cases where changes of symmetry are concerned, that the intermediate forms are, as it were, points of unstable equilibrium, and that the body therefore assumes these forms rarely, as in some instances, or never, as in others” (Bateson and Bateson, 1891, 421). The notion of

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“unstable equilibrium” built on Galton’s notion of “organic stability” as the basis of discontinuous change introduced in Natural Inheritance (1889) (Galton, 1889, 27; Gayon, 1998, 170-74; Gillham, 2001). Variations in degree, on the other hand, were by nature quantitative, and, they argued, “no one as yet has ever indicated the way by which such Variations could lead to the constitution of new forms, at all events under the sole guidance of Natural Selection” (Bateson and Bateson, 1891, 421). Evolution, they thus implicitly argued, did not result from selection operating on continuous characters (as assumed by the neo-Darwinians and biometricians alike) but was rather fueled by the sudden appearance of new “variations in kind,” upon which selection could operate. Evolution, in other words, was saltatory, not gradualistic, in nature. Having made a series of bold theoretical claims about the nature of evolutionary change, the two authors next turned to the question of the best methodology to study evolution. In this context they challenged the efficacy of morphology to deal with this problem. As a “deduction from facts,” they reasoned that only variations arising through a continuous process of change will leave traces of their history that can be investigated by means of a “comparative study of form and development,” that is, by employing the conventional morphological toolkit. If, as they believed, the kind of variation important for evolution is variation in kind rather than variation in degree, and hence produced through a sudden event, then “comparative morphology [will] cease to be an effectual guide to the history of Descent.” As they explained: We are therefore disposed to think that the first teaching of the facts of Variation is this: that comparison of forms is not likely to be a good guide to the history of those forms; and that there is no evidence that degrees of apparent relationship of form are an indication of degrees of actual relationship by descent; and that nothing short of an actual knowledge of the processes of Variation and a discernment of the changes which are possible to living things from those which are impossible to them, can be of any use in the solution of the problem of Descent. (Bateson and Bateson, 1891, 4)

In this way, the Batesons called into question the value of morphology for investigating evolutionary change. This conclusion, they well realized, “touches the nature and soundness of the received principles by which morphological facts are interpreted” (Ibid., 416). If morphology can only study variation in degree but not variation in kind, its importance as a means of exploring evolutionary change was thus significantly diminished. Hence, the 1891 paper by the Bateson siblings was not at all a merely empirical contribution to botany. Rather it should be read as a daring theoretical manifesto, offering a radical revision, not just of the approach to variation and heredity, but of foundational concepts guiding modern biological investigation. They challenged such fundamental Darwinian tenets as gradualism and the role played by natural selection. They also applied boundary conditions that restricted the applicability, and hence efficacy, of morphology and biometry to evolutionary problems. In short, the 1891 paper laid out a completely new direction of research as well as a new mode of conceptualizing evolutionary change. William Bateson soon followed up this declaration of principles with an even more extensive avowal of his new, revisionist views of variation and speciation in a monograph devoted to exploring “variation in kind.”

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MATERIALS FOR THE STUDY OF VARIATION (1894) Two years after the paper outlining his maverick views of variation, Bateson’s book, Materials for the Study of Variation, Treated with Especial Regard to Discontinuity in the Origin of Species (1894), was published. In this book, he extensively catalogued notable cases of variation, paying particular attention to discontinuous variations that appeared to have arisen suddenly and without any apparent adaptive value. His view of the discontinuous nature of evolutionary change shared similarities with the ideas Brooks expressed in his 1884 book. Yet despite pointing fifteen years later to the formative influence Brooks had on his own thinking, it is curious to note that Bateson did not refer to Brooks or his proposed “law of heredity” in his own book on variation. In Materials for the Study of Variation, Bateson emphasized the importance of this topic, proclaiming that “Variation, whatever may be its cause, and however it may be limited, is the essential phenomenon of Evolution. Variation, in fact, is Evolution” (Bateson, 1992, 6). In the few pages he devoted to speculating about the cause of variation, he presented a physicalistic description that shared essential features with the views expressed earlier by Mivart and Brooks. Meristic or repetitive variation was, he believed, the result of mechanical changes in the pattern of a developing organism. He explained his view as follows: “Patterns into which the tissues of animals are divided represent positions in which the forces that effect the division are in equilibrium. On this view the lines or planes of division would be regarded as lines or planes at right angles to the directions of the dividing forces; and in the lines of Meristic Division we are perhaps actually presented with a map of the lines of those forces of attraction and repulsion which determine the number and positions of the repeated parts, and from which Symmetry results” (Ibid., 70). As suggested in the 1891 paper, he thus assumes that such variations in degree are the result of mechanical forces operating on living matter to effect change (see Coleman, 1970). Bowler has rightly noted that the “evidence presented in Materials is not, of course, experimental in character. Bateson gathered his evidence for the existence of discontinuity from the study of natural varieties and occasional monstrosities” (Bowler, 1994, xxii). Yet we should not overlook Bateson’s statements that emphasize the pressing need for an experimental approach to the problem. “So long as systematic experiments in breeding are wanting,” he wrote, “and so long as the attention of naturalists is limited to the study of normal forms, in this part of biology which is perhaps of greater theoretical and even practical importance than any other, there can be no progress” (Bateson, 1992, 76). In so doing, he echoed Brooks’s earlier statement that the nature of variation could best be approached by undertaking extensive hybridization experiments (Brooks, 1883, vii). In the conclusion to his book, Bateson reiterated this point, stating: “The only way in which we may hope to get at the truth is by the organization of systematic experiments in breeding, a class of research that calls perhaps for more patience and more resources than any other form of biological inquiry” (Bateson, 1992, 574). In the event, with considerable patience but few resources, he himself took up the call. BATESON’S HYBRIDIZATION EXPERIMENTS, 1895-1900 Immediately after publishing Materials for the Study of Variation, Bateson initiated a new line of research to study variation experimentally. In part, this appears to have resulted from his doubts

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about the persuasive value of simply amassing facts on discontinuous variation. As he admitted to Anna shortly before the work appeared, “Every day almost makes me misgive more and more about this book. Several of the sections of evidence seem very weak now they are actually ‘floated and rendered’ as the builders say, and half of the best things seem never to have been got in at all” (B. Bateson, 1928, 54). These misgivings intensified after the negative reception given to his book. Adam Sedgwick, Balfour’s successor as the head of Animal Morphology at Cambridge, called Bateson’s approach “stupid & narrow” and thereafter provided him little institutional support. 2 His earlier mentor and friend, Frank Weldon, viciously attacked Bateson’s new research program in a scathing review of Materials (Weldon, 1894). Despite his criticism of biometry in the 1891 paper, Bateson found this lack of support and harsh criticism difficult to accept. In 1895, he began a series of hybridization experiments in the hope of providing evidence to support his view that discontinuous or saltatory variations, rather than the small, adaptive variations envisioned by Darwin and the neo-Darwinians, was the basis for evolutionary change. He thus aimed to prove that alternative characters did not, in fact, generally blend but rather were discrete and somehow self-perpetuating. In pursuing this line of work, Bateson well realized the need for collaborators, but his marginal status at Cambridge worked against this. Even Anna, although sharing his views about variation and evolution, was no longer able to assist with such a study. In 1890, when her mother left Cambridge and thus broke up the family home, she decided to abandon academic biology to support herself as a tradeswoman, using her inheritance to purchase a nursery in a small village in Hampshire, in the south of England.3 Although William disapproved of this decision, believing it lowered her social status, Anna was realistically confronting her limited options. As a student of botany at Newnham College, Cambridge, Anna had only obtained a second class in both parts 1 and 2 of the Natural Sciences Tripos (1884, 1886). Although this qualified her for a position as a botanical instructor (demonstrator) at the Balfour Biological Laboratory for Women as well as earned her a Bathurst Studentship to pursue postgraduate study, a future career in academic science was not well assured. The reality was that a woman with a degree in science in 1890 had precious few options for employment. Although she had gained recognition among botanists through her publications, her tripos rankings made a secure position at Cambridge unlikely (Richmond, 1997).4 While many women science graduates looked for teaching positions at a 2

3

4

Sedgwick to Bateson, 9 October 1890: “I also think that yr. views on Zoology—on the morphology side— are stupid & narrow, but that is a very different thing from thinking that yr. work is stupid & unprofitable.” Bateson Correspondence, ADD 8634, Manuscripts Room, Cambridge University Library. Anna Bateson obtained second-class honors in the Natural Sciences Tripos, 1884 and 1886. She taught advanced botany at the Balfour in 1886 and was appointed assistant demonstrator in botany in 1887. She held a Bathurst studentship, 1887-1889, which supported post-graduate research. She assisted Francis Darwin in the University Botanical Laboratory, 1886-1890. In 1890, she left Cambridge to establish a nursery in Bashley, Hampshire, that became quite profitable. See Cock, 1979. That Anna Bateson’s work had gained attention, particularly among German botanists, is shown by Carl Correns’s letter to Bateson, 21 October 1900, who asked him: “Sind Sie mit Miss A. Bateson verwandt, die die hübschen Untersuchungen über die Wirkung von Kreuz- und Selbstbefruchtung bei kleinblüthigen Pflanzen ausgeführt hat? Ich habe diese Arbeit mit grossem Interesse gelesen!” [“Are you related to Miss A. Bateson, who published the wonderful investigation of the effect of cross- and selffertilization in small-flowered plants? I read this work with great interest!”] Correns refers to A. Bateson, 1888. This same paper was cited by Munich professor of botany Karl Goebel in his essay on the biology of flowers in the 1909 volume commemorating Darwin (Goebel, 1909, 421).

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secondary school that offered science instruction, in the 1890s few such positions were open (Perrone, 1993). In the event, while Anna’s decision well suited her needs and inclination, it left William without a suitable intellectual confident and experimental ally. With Anna gone, Bateson turned to another Newnham botanist, Dorothea Frances Matilda (Dora) Pertz (1859-1939) to help carry out an experimental study of variation. A niece of geologist Charles Lyell, Dora Pertz was an accomplished botanist with solid experimental skills (Browne 2004). Bateson designed crosses for Pertz to carry out in Veronica, aiming “to test whether there is any difference between offspring raised from abnormal flowers, and the offspring of normal flowers borne by the same plant.” Pertz pursued this work over four growing seasons, from 1892 through 1895, self-crossing abnormal flowers and normal flowers and then comparing the proportion of normal flowers to abnormal ones in the progeny. In the joint paper they published in 1898, they noted that normal flowers appeared about 80-90 percent of the time, no matter whether the parents were normal or abnormal (Bateson and Pertz, 1898). These results were disappointing, and thus the approach was abandoned. In the mid-1890s Bateson entered into what became a highly profitable mutual collaboration with another young Newnham botanist, Edith Rebecca (Becky) Saunders (1865-1945). In many respects, Becky Saunders’s background was similar to Anna Bateson’s. Saunders took second class honors in Part I of the Natural Sciences Tripos in 1887, but the following year she gained distinction by gaining a first in Part II (botany). She was awarded a Bathurst research studentship, which allowed her to pursue postgraduate research, and served as botanical demonstrator at the Balfour Biological Laboratory for Women. But in 1899, upon the resignation of her friend Marion Greenwood, Saunders was appointed director of the laboratory, a position she held until its closure in 1914, also holding various college positions. Saunders was thus more fortunate than either Anna Bateson or Dora Pertz in being able to pursue an academic career in science. This was fortunate for Bateson. With an extensive knowledge of botany and a strong background in research, Saunders proved to be an excellent colleague. Her independent research conducted on problems of variation ultimately provided critical evidence that supported his views of discontinuous variation (Richmond, 1997, 2001; Creese, 1998). Using seeds that Bateson brought back with him from a trip to Italy, in the summer of 1895 Saunders initiated a series of breeding experiments on an allotment they rented from the Cambridge Botanic Garden. In the Italian Alps, Bateson had observed two distinct forms of the perennial herb Biscutella laevigata growing side-by-side, one with hairy (hoary) leaves and the other with smooth, or glabrous, leaves. He found this especially curious given that the two forms “intercrossed readily” and yet apparently bred true to form: offspring generally exhibited either hoary or glabrous leaves, with relatively few intermediates. Saunders set out to determine how this distinctness was maintained. As she noted in her paper of 1897, “on the supposition that hairiness and smoothness are characters capable of blending freely, it might be expected that offspring derived from a cross between hairy and smooth parents would tend constantly to regress to a mean condition of texture.” Since this biometrical prediction was not fulfilled, it appeared this case “might lead to interesting results bearing upon the views which have recently been brought forward with regard to discontinuous variation and its value as a factor in the origin of species”

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(Saunders, 1897). The case, in other words, appeared to support Bateson’s hypothesis of discontinuous variation. Presenting her findings qualitatively, Saunders noted that the alternative characters generally persisted in the offspring. Out of 208 plants raised, 127 were hairy, 36 appeared to be intermediates, and 45 were smooth leaved. When she crossed two hairy plants, out of the 76 offspring that matured, 61 were hairy, 13 intermediate, and 2 smooth. These results were thus promising. Hairy and smooth-leaved Biscutella appeared to represent two stable discontinuous characters, with only a handful of “blended” offspring appearing in cross-bred plants. Moreover, the hairy character appeared to be “prepotent” over smooth leaves, a fact that Saunders wished to follow up (Ibid., 17, 18). In the summer of 1896, Saunders set out “to ascertain the nature and amount of the variations occurring among the offspring of unlike parents,” intercrossing hairy-leaved plants with smoothleaved ones. The seeds she procured from the offspring of the cross were planted the same year. These results she presented in tabulated form:

Classification of 120 Cross-Bred Seedlings

Surface hairy

Surface intermediate

Surface smooth

Totals

Number of seedlings derived from five hairy plants x smooth plants

4

7

26

37

Number of seedlings derived from five smooth plants x hairy plants

5

32

28

65

Number of seedlings derived from one plant, surface, marginal hairs numerous x hairy plant

12

6

0

18

Totals

21

45

54

120

These data were not as supportive of the discontinuity hypothesis. Saunders attempted to downplay the negative evidence, reporting that the results “show that a blending of parental characters as regards hairiness and smoothness occurs to a certain extent in the offspring of plants of dissimilar types, giving rise to intermediate forms.” She could only add that plants exhibiting an intermediate condition early in life often grow distinctly more glabrous with maturity (Ibid., 23). Saunders’s investigation of Biscutella illustrates the evolving approach she and Bateson took toward the study of variation. In setting up their hybrid crosses, they looked for data that indicated the persistence of “variations in kind” in heredity. They did not carry out a quantitative and statistical analysis of “variation in degree,” as biometricians did in recording the proportion of individuals showing ancestral rather than parental inheritance through the regression to the population mean. In concentrating on the retention of parental traits among the progeny, they rather aimed, as Robert Olby has noted, to predict “the probability that a given transmissible

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character of the parents will be possessed by the offspring” (Olby, 1987, 400-401). Olby also tellingly points out that Bateson “was thus concerned not with variation in general, but with those forms of variation which he saw as significant in the origin of species” (Olby, 1989, 308). As Bateson described the approach he and Saunders employed: “Cross-breeding, then, is a method of investigating particular cases of evolution one by one, and determining which variations are discontinuous and which are not, which characters are capable of blending to produce a mean form and which are not” (Bateson, 1899, 64). It was thus not blending characters but rather alternative ones that held their attention. Encouraged by her findings in Biscutella, Saunders embarked on a new series of crosses using other species with forms exhibiting alternative variations. To carry out this scheme, however, she needed a larger plot of land. Bateson thus began to solicit outside funding, approaching the Royal Society as a member of its newly reconstituted Evolution Committee. Formed in 1894 as the Committee for the Measurement of Plants and Animals, with Galton as chair and Weldon as secretary, the committee soon encountered differences of opinion over its orientation, whether evolution was to be approached mathematically (as did the biometricians) or biologically. One consequence was the invitation of Karl Pearson (a mathematician) and Bateson (a biologist) to join the committee in 1896 (Froggatt and Nevin, 1971; Provine, 1971; Kim, 1994). An early initiative of the committee was inviting animal breeders and horticulturists to submit proposals for investigations on heredity “such as relate to the means whereby new races of plants and animals come into existence, and old ones are modified,” for which small sums were available to support. Although now a nurserywoman, Anna Bateson was among those submitting a proposal to study heredity in the Lady Slipper orchid, Cypripedium. Weldon mentioned Anna’s application in a letter to Bateson, saying: “The scheme of your sister’s is worth twenty programmes.” Although funding was apparently granted, it is not clear whether Anna ever carried out the project (Cock, 1979, 62). By January 1900, the growing animosity between the biometricians and Bateson came to a head. As a result, Galton, Pearson, and Weldon all resigned from the committee, and Bateson took over as secretary. He took this opportunity to redirect the committee’s mission to reflect his own approach. Funding from the Royal Society allowed Bateson and Saunders to expand their experimental work, even if they continued primarily to rely on domestic resources (Richmond 2006). The Society also eventually supported the publication of their findings, although not without considerable debate (B. Bateson, 1928, 60-61). Using the small grant from the Royal Society to procure a larger plot of land and some assistance, in the growing season of 1897-1898, Saunders thus began a study of inheritance in four new species, all of which exhibited alternate characters. This included: (1) Matthiola, the common garden stock, whose hairy and glabrous forms offered “excellent material for statistical experiments upon cross-breeding;” (2) Lychnis, which exhibited hairy and glabrous varieties as well as differently colored flowers; (3) varieties of Atropa with contrasting flower and fruit colors; and (4) Datura varieties in which the fruits borne were either prickly or smooth. By the beginning of the third growing season (1899-1900), Saunders had accumulated sufficient data to conclude that this study supported Bateson’s views concerning discontinuous inheritance, but she did not yet feel enough advanced to publish a full account, which indeed was not forthcoming until 1902.

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During these years Bateson also carried out breeding experiments in plants and animals. In 1895 he began crossing butterflies and in 1898 initiated extensive cross-breeding experiments in poultry (Bateson, 1897, 1898a, and 1898b; see also Olby, 1985, 125; Cock, 1971, 29-34). 5 However, he encountered technical problems in crossing butterflies and was generally not as successful as Saunders in finding support for his views of discontinuity. He found it hard to interpret his results, which appeared “irregular and puzzling” (Cock, 1971, 6). In the end, he did not publish any of the results of this work until after he learned about Mendel’s experiments in hybridization (Bateson, 1902b). Some understanding of the conceptual basis of Bateson’s and Saunders’s early research program comes from general remarks he made in several papers published before May 1900. In a paper of 1897, for example, Bateson briefly noted Saunders’s preliminary findings, stating that her experiments indicated that “the two characters of smoothness and hairiness do not completely blend, and the offspring do not regress to one mean form, but to two distinct forms. The variety, in short, is not ‘swamped by intercrossing’” (Bateson, 1897, in Bateson 1928, 1: 354). In July 1899, in a paper presented at the first international conference on plant hybridization organized by the Royal Horticultural Society, he again referred to Saunders’s work. In this oft-cited paper, “Hybridisation and Cross-Breeding as a Method of Scientific Investigation,” Bateson described in detail the nature of the investigations he and Saunders were pursuing. He presented a brief synopsis of his views about variation, admitting that “we are far from knowing which kinds of variations may thus be definite and palpable, and which are not.” To gain such answers, more extensive data was needed, and he implored practical breeders, and “especially the cross-breeder of plants or of animals,” to help provide “first-hand evidence as the magnitude of variations.” The question of the day, in his view, was “Why do not nascent varieties become obliterated by crossing with the type form?” This required crossing a variety “with its nearest allies” and recording “how many of the offspring resembled each parent and how many shewed characters intermediate between those of the parents” (Bateson, 1899, 60-61, 63). This, in short, was precisely the approach he and Saunders were using. This paper is frequently cited because it presents a vivid statement of Bateson and Saunders’s research program just prior to his reading Mendel’s 1865 paper. Some earlier historians have interpreted Bateson’s statements as anticipating Mendel’s experimental approach (see Cock, 1971). Yet, it is clear, judging by the work he had pursued since 1890, that this is mistaken. Both in terms of methodology and conceptualization, Bateson and Saunders were far removed from the approach Mendel brought to his hybridization studies. To be sure, like Mendel Bateson and Saunders crossed forms exhibiting alternative characters and recorded how many of the offspring 5

See Bateson, 1898, which includes a description of his experimental set-up for butterfly breeding: “We had the pleasure of seeing Mr Bateson’s garden in which these experiments are carried out, and admired the simplicity and completeness of the arrangement for these experiments. He finds no difficulty in getting the butterflies to pair and oviposit. The apparatus consists of a box placed in the garden, open to the weather, and covered with gauze. The box is about 30” x 18”, and contains a supply of flowers in a glass and of the food-plant growing in a pot. Some shade is provided by a partial covering of canvas thrown loosely over. Mr Bateson has long ranges of these boxes and of pots of the food-plants to which the insects may be removed and on which they are sleeved after oviposition has been completed.” A notebook containing records of Bateson’s crosses of moths and butterflies is in the Bateson Collection, John Innes Centre, Norwich.

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resembled one or the other parent and how many showed characters intermediate between the two. However, they generally did not know the ancestry of the parental generation, and they seldom bred an F2 generation. Saunders, for example, only did so in Lychnis, repeating experiments made by Hugo de Vries. In an F1 cross between hairy and smooth leaved plants that produced all hairy progeny, the offspring were “backcrossed” with the smooth parent, producing an F2, “of which some were hairy and others smooth” (Bateson, 1899, 64-65). Neither she nor Bateson generally carried out reciprocal crosses. This largely was because Bateson was operating with a working hypothesis that “in-breeding may have a specific effect in modifying the power of transmitting parental character to offspring” (Bateson and Saunders, 1902a, 3; see also Olby, 1987, 412-13; Cock, 1971, 6). The differences between their work and Mendel’s systematic approach could not be starker. With regard to data collection, Bateson and Saunders recorded the results of their hybrid crosses, but only to note whether the parental character persisted in the offspring or was blended. This reflected the ultimate goal of their research, described by Bateson in 1899: How did the one form separate from the other? By crossing the two forms together and studying the phenomena of inheritance, as manifested by the cross-bred offspring, we may hope to obtain an important light on the origin of the distinctness of the parents, and the causes which operate to maintain that distinctness. (Bateson, 1899, 63)

Thus, they solely focused on tracing the inheritance of parental discontinuous characters in the offspring. This explains the small number of crosses made and paucity of data collected in comparison with Mendel. Indeed, Bateson downplayed the amount of data required, telling breeders that only “rough statistics” were needed: “All that is really necessary is that some approximate numerical statement of the result should be kept,” he noted, and just “a few words” might suffice to describe the outcome of a cross (Ibid.). The generally qualitative character of their work is particularly apparent in Bateson’s discussion of Saunders’s results in Biscutella. In mentioning the “well-marked discontinuity between the two varieties,” Bateson referred to the difference between “the nature of the relationship of the two forms to each other,” explaining in a footnote his use of the term relationship: “It is used to denote not simply the blood-relationship of the forms to each other, but those physiological relations subsisting between them which are manifested by experimental crossing. The word is thus used in a sense similar to that which it bears when we speak of the chemical relations of one substance to another” (Ibid., 64n.). He spoke of the “transmitting powers” of diverse varieties being unequal, noting that “in each the mechanism of inheritance works differently.” Yet he recognized that if “tested by the method of breeding and by study of the transmitting powers, the relation of varieties and species would be shewn in an entirely new light” (Bateson, 1900, 55). Unlike Mendel, Bateson could discern no pattern in his results, let alone any indication of the physical basis of heredity. In short, Bateson and Saunders continued to work within the same hybridizing tradition that Darwin pursued (Olby, 1985; Bartley, 1992). Hence it is clear that despite some tantalizing resemblance, both Bateson’s and Saunders’s experimental design and interpretation of results were a far cry from the focused experimental analysis developed by Mendel (Balen, 1986, 181). After five years of work, then, neither Bateson nor

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Saunders could discern any general patterns to describe the results of their crosses. A. G. Cock described the situation facing Bateson in 1900 as follows: “he had to some extent reached an impasse as far as any explanation of his experimental results was concerned, an impasse which was resolved when he came to learn of Mendel” (Cock, 1971, 7). Robert Olby has painstakingly and convincingly reconstructed a chronology of the critical events in the spring of 1900. Bateson apparently first read Hugo de Vries’s April 1900 paper in French on the “law of the disjunction of hybrids” on 8 May 1900, while on the train to London to read a paper to the Royal Horticultural Society. But he only learned about Gregor Mendel’s 1866 paper on the laws of heredity after reading de Vries’s subsequent paper published a month later that was directed at German botanists (de Vries, 1900a, 1900b; Olby, 1987). He presumably read the papers of Carl Correns and Erik von Tschermak shortly afterwards (Correns, 1900; Tschermak, 1900). Certainly, it is easy, given Bateson’s and Saunders’s previous inability to recognize any law-like patterns in their own crosses, to understand the incredible impact these works must have had on them. Having long attempted to gather evidence in support of the hypothesis of discontinuous variation, de Vries’s proposal of a “law of disjunction,” based on a modification of Darwin’s hypothesis of pangenesis, must have been very interesting indeed. Describing crosses between parents distinguished by a single character, de Vries noted that there was no possibility of the progeny evidencing blending heredity because “l’hybride ne saurait tenir le milieu entre eux; car le caractère simple doit être considéré comme une unité non divisible” (the hybrid would not be able to blend, because the simple character must be considered as an indivisible unit) (de Vries, 1900a). Mendel’s paper, however, as Olby has noted, provided “the key he needed. It gave a causal explanation for the production of variation which was independent of the environment; it showed how hereditary differences were separated from hybrid mixture because of the purity of the germ cells. The theory offered an algorithm with which to predict the outcome of experiments in crossing” (Olby, 2004). It provided, in short, both the critical conceptual principles and method of analysis that Bateson and Saunders had long been seeking. There was also another aspect of this flurry of papers that engaged Bateson and Saunders. The data de Vries and Correns presented were based on many of the same species they were studying, namely, Veronica, Lychnis, Datura, Stramonium, and Matthiola. Not only did these papers provide critical conceptual keys to aid in the interpretation of their findings, but they also may have triggered a desire to publish their own data as soon as possible, given that neither Bateson nor Saunders had as yet published full accounts of their work of the past five years. THE IMPACT OF MENDEL The extent to which the “rediscovery” of Mendel’s work prompted a radical reorientation in Bateson and Saunders’s research program can readily be gauged from noting the change in tenor and substance between his previous pronouncements and the published version of Bateson’s May 1900 paper to the Royal Horticultural Society. He began with the bold statement: “An exact determination of the laws of heredity will probably work more change in man’s outlook on the world, and in his power over nature, than any other advance in natural knowledge that can be foreseen,” followed by the confident assertion that “these laws can be determined.” Repeating a call to horticulturists to conduct controlled breeding experiments, Bateson spoke of the effort to

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discover a law of heredity as of great practical importance in helping to predict “the degree with which the purity of a strain may be increased by selection in each successive generation.” He mentioned Galton’s well-known work on this problem, but noted that Galton’s law of ancestral heredity did not apply to cases like those he and Saunders considered, in which “on crossing two varieties the character of one variety is almost always transmitted to the first generation,” or those in which “the characters of one variety very largely, though not exclusively, predominate in the offspring,” and thus could not be regarded as a general law of heredity. This was little changed from earlier presentations. It was in this context, however, that Bateson mentioned de Vries’s recent “brief account” of his study of variation (citing, in the footnote, both the French and German texts) and pointed out the essential aspects of this work: “The cases are all examples of discontinuous variation: that is to say, cases in which actual intermediates between the parent forms are not usually produced on crossing. It is shown that the subsequent posterity obtained by self-fertilising these cross-breds or hybrids break up into the original parent according to fixed numerical rule” (Bateson, 1900, 57; my emphasis). Bateson then referred to Mendel’s work, calling his account “excellent and complete” and stating that “the principles which he is able to deduce from them will certainly play a conspicuous part in all future discussions of evolutionary problems.” He drew attention to Mendel’s use of dominant to refer to the prevailing character, and recessive to the other, which avoided “the complications involved by use of the expression ‘prepotent.’” There could be no doubt, Bateson claimed, that “Mendel’s law is a substantial reality”: already his findings had been confirmed by de Vries, Correns, and Tschermak. It is clear, he concluded, that “we are in the presence of a new principle of the highest importance.” Moreover, the “facts of crossing prove that each ovule and each pollen grain is pure in respect of each character to which the law applies.” The direction of future work was thus set: Mendel’s “hypothesis of perfect purity in the reproductive cells” required validation and “the subjects of experiment should be chosen in such a way as to bring the laws of heredity to a real test.” (Bateson, 1900, 57, 59, 60). Bateson and Saunders were ideally situated to do just that: they immediately embarked on revising their pre-1900 experimental design to reflect the new orientation of the Mendelian laws and methodology (Cock, 1971, 4-7). By October 1900, they were well under way, having identified new problems as well as reinterpreted their previous data in order to test the general validity and extent of Mendelian heredity.6 The era of Mendelism had begun. The earliest publications of Bateson and Saunders after the rediscovery of Mendel well reveal the contrast between their pre- and their post-Mendelian research. These include the co-authored first part of the five-part series (1902-1909) entitled Reports to the Evolution Committee of the Royal Society (submitted in December 1901, with additions dated March 1902), and Bateson’s Mendel’s Principles of Heredity: A Defense (preface dated March 1902), which was written in consultation with Saunders. In the introductory essay to the Report entitled “Experimental Studies in the Physiology of Heredity,” Bateson and Saunders described their research program investigating discontinuous 6

That this is true comes from the statement: “After the re-discovery of Mendel’s work it seemed desirable to use varieties differing in a pollen-character. Searching for such forms, it was found in October, 1900, that the Sweet Pea, Emily Henderson (referred to as E. H.), a pure white var., usually has pollen distinct from that of normal vars.” See Bateson, Saunders, and Punnett, 1905, 80.

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characters and the effect of inbreeding “in modifying the power of transmitting parental character to offspring.” Since beginning this study in 1895, they noted, “the whole problem of heredity has undergone a complete revolution,” and the evidence they had collected on prepotency, the longheld belief that in hybrid crosses one race (generally the “older”) would stamp its character on the progeny, “is now capable of different interpretations, and it is clear that to obtain a definite result on this point, a new set of precautions must be used.” In the wake of Mendel, Bateson now recognized that prepotency could be interpreted as a consequence of dominance, and reversion— the reappearance of ancestral traits—as the recurrence of latent characters. Although admitting that their investigations were not as yet sufficiently advanced, the authors stated that “we feel that with the re-discovery and confirmation of the principle which will henceforth be known as Mendel’s Law, the study of heredity and the cognate problems of evolution must enter a new phase” (Bateson and Saunders, 1902a, 3, 4-5, 5). They could wait no longer to publicize their findings. After laying out Mendel’s basic principles and noting the confirmation provided by de Vries, Correns, and Tschermak, they concluded that “the truth of the law enunciated by Mendel is now established for a large number of cases of most dissimilar characters, beyond question.” The key element of Mendel’s law in their view was the following: “The essential part of the discovery is the evidence that the germ-cells or gametes produced by cross-bred organisms may in respect of given characters be of the pure parental types and consequently incapable of transmitting the opposite character,” such that “there may be, in short, perfect or almost perfect discontinuity between these germs in respect of one of each pair of opposite characters.” (Ibid., 11, 12; italics in original). This statement is remarkable given that previously neither had ever discussed the cellular basis of heredity, not even, as Brooks had done, referencing the hypothesis of pangenesis. But conceptualizing discontinuous variation as a “variation in kind” easily permitted them to associate the “physiological relations subsisting between them [two discontinuous varieties] which are manifested by experimental crossing” that Bateson spoke of in 1899 with qualitatively different characters in “the germ-cells or gametes” that Mendel proposed. As Lindley Darden has noted, “That Bateson immediately saw the conceptions as important indicates that they had connections with approaches and problems he had already formulated. That Bateson called them new indicates they differed from some of the conceptions he had held previously. Thus both continuity with old problems and change from old conceptions occurred with Bateson’s adoption of the Mendelian approach” (Darden, 1977, 89). Several statements in the separate pieces that make up the first Reports to the Evolution Committee support this, indicating the ways in which Bateson and Saunders’s previous work differed from the approach taken by Mendel. In the general section discussing “The Facts of Heredity in the Light of Mendel’s Discovery,” they noted how many earlier breeding experiments now “must be re-stated in terms of Mendel’s hypothesis,” noting that it “would be a useful task to go similarly through the literature of breeding and translate the results into Mendelian terms. Such an exercise would show that the change which must now come over the conceptions of biology can only be compared with that which in the study of physical science followed the revelations of modern chemistry” (Bateson and Saunders, 1902b, 125). This was particularly true of their own work.

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In terms of change to their experimental design, after describing Mendel’s use of back-crosses (“crossing the first crosses with pure D and pure R forms respectively”), they noted that in their crosses, “almost all our breedings of cross-breds with pure types have been in the form cross-bred 6 x pure 7, but reciprocal experiments are in progress” (Ibid., 10n.) In discussing her botanical experiments, especially comparing her 1897 paper on Biscutella with the 1902 account of subsequent crosses, Saunders’s noted she had not controlled for the purity of parent plants, and those they originally believed showed complete dominance may have included cross-breeds carrying both a dominant and recessive trait (that is, a heterozygote) (Saunders, 1902, 22). Also, she had not always been careful to prevent unwanted cross-fertilization through the agency of insect pollination. Yet despite these lapses, she believed her results did “follow Mendel’s law with considerable accuracy, and no exceptions that do not appear to be merely fortuitous were discovered” (Ibid., 44). In hybrid crosses between Lychnis vespertina (white flowers; hairy) and L. diurna (red flowers; glabrous), for example, Saunders recalculated the proportion of hairy (dominant) to glabrous (recessive) in the F2 generation as 3.2:1 (rather than the expected 3:1). The same was true for Atropa (color of fruit) and Datura (two pairs of characters: smooth or prickly fruit; white or violet flowers). She included a series of tables to present the data, revising the earlier tabulation appearing in the 1897 paper into categories that reflected Mendelian modes of analysis. (See, for example, Figures 2 and 3, providing data for Lychnis crosses.)

Figure 2. Table from Saunders, 1902, p. 16.

In Matthiola, however, the situation was not as clear-cut as in the other genera, for “the phenomena are much more complex.” She initially set out to follow only one character—leaf surface (hoary or glabrous)—using five different types or races. However, she “found that the results differed widely according to the variety, and occasionally according to the individual, with

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which the crosses were made,” so she expanded her analysis to include other characters as well, including seed color, flower color (white, cream, red, pinkish white, bluish pink), and time of flowering. While several simple cases did appear to “follow Mendelian principles,” others did not. For example, she found that hoariness was not always dominant, as expected. In fact, even after considerable effort to come up with a system to present her results, they remained “complicated and difficult to follow” (Ibid., 15, 32). These seemingly aberrant cases required further analysis and indeed continued to occupy Saunders for the next few years.

Figure 3. Table from Saunders, 1902, p. 17.

With respect to Bateson, it is interesting to contrast the account he provided of the importance of Mendel’s work in 1900 with the revision of this paper included in Mendel’s Principles of Heredity. This well indicates the rapid conceptual and methodological transformation that he was experiencing by virtue of translating his previous views on variation into a new Mendelian mode of analysis. This contrast was sharpened by the need for him to clarify and “defend” the tenets of Mendelism in the wake of the attack launched by Weldon in the second number of the new journal Biometrika, published in January 1902. Weldon downplayed the various accounts of the Mendelian 3:1 ratio of dominants to recessives by pointing to deviations from the expected ratio and claiming these were not simply accidental but indicated the additional operation of ancestral inheritance, not just parental. As he put it, “the degree to which a parental character affects

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offspring depends not only upon its development in the individual parent, but on its degree of development in the ancestors of that parent” (Weldon, 1902a, 244, 251). This paper prompted Bateson to respond, drafting Mendel’s Principles of Heredity—A Defense in only a month. The most significant modification Bateson made to the text of his earlier paper of 1900, announcing Mendel’s laws to the English-speaking public, was an illustration of the union of distinct pairs of characters in a cross between varieties differing in two characters—that is, a dihybrid cross. This material was not included in Reports to the Evolution Committee, and hence must have been added in February or March 1902. For this reason, his description is presented in its entirety below: Mendel made further experiments with Pisum sativum, crossing pairs of varieties which differed from each other in two characters, and the results, though necessarily much more complex, showed that the law exhibited in the simpler case of pairs differing in respect of one character operated here also. In the case of the union of varieties AB and ab differing in two distinct pairs of characters, A and a, B and b, of which A and B are dominant, a and b recessive, Mendel found that in the first cross-bred generation there was only one class of offspring, really AaBb. But by reason of the dominance of one character of each pair these first crosses were hardly if at all distinguishable from AB. By letting these AaBb’s fertilise themselves, only four classes of offspring seemed to be produced, namely, AB

showing

both dominant characters.

Ab

showing

dominant A and recessive b.

aB

showing

recessive a and dominant B.

ab

showing

both recessive characters a and b.

The numerical ratio in which these classes appeared were also regular and approached the ratio: 9 AB : 3Ab : 3aB : 1ab. But on cultivating these plants and allowing them to fertilise themselves it was found that the members of the Ratios ab class produce only ab’s.

1

3

{1

aB class may produce either all aB’s,

{2

or both aB’s and ab’s

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Ratios

3

9

{1

Ab class may produce either all Ab’s,

{2

or both Ab’s and ab’s

{1

AB class may produce either all AB’s,

{2

or both AB’s and Ab’s

{2

or both AB’s and aB’s

{4

or all four possible classes again, namely, AB’s, Ab’s, aB’s, and ab’s,

and the average number of members of each class will approach the ratio 1 : 3 : 3 : 9 as indicated above. (Bateson, 1902a, 11-12.)

Although this material looks elemental to anyone familiar with Mendelian genetics, it was a novel presentation for those just being introduced to Mendel and different from that in Mendel’s 1866 paper. Bateson’s formulation provided a clear presentation of the essence of Mendel’s hypothesis of the “purity of the germ cells” and the law of segregation—the key features that provided solid theoretical grounding for Bateson’s conception of discontinuous variation. Together with the subsidiary assumption of dominant and recessive characters, an explanation was provided for the regular, proportional appearance of parental characters in hybrid offspring, the “blending” of characters in the “heterozygote,” and the reappearance of apparently long lost characters, not because of a “reversion” to ancestral traits but because of the reappearance of latent recessive factors. Mendel, in short, provided an explanation for many of the phenomena Bateson had been grappling with for over a decade. Bateson’s development of a schematized representation of Mendel’s principles may also have come in response to a table Weldon included in his criticism of Mendel that presented all the possible combinations of characters in hybrid crosses along with their frequency (Weldon, 1902a, 235). From this analysis, Weldon concluded that Mendel’s 9 : 3 : 3 : 1 ratio was an approximation that held only in certain crosses. This was because, he maintained, the “degree to which a parental character affects offspring depends not only upon its development in the individual parent, but on its degree of development in the ancestors of that parent.” Hence, since the “law of segregation, like the law of dominance, appears therefore to hold only for races of particular ancestry,” Mendel’s laws could not be considered to be general laws of heredity (Weldon, 1902a, 244, 251). It is possible that Bateson’s presentation of a “mathematized” schema to illustrate Mendel’s analysis of hybrid crosses was calculated to neutralize this challenge by Weldon. Scholars have pointed to the positive as well as negative aspects of the ensuing biometricalMendelian controversy that resulted in a series of vociferous public exchanges after 1902 (Nordman, 1992, 68). It seems clear that Weldon’s consistent, targeted, and clever criticism of Mendelian terminology, experimental design, and data analysis forced Bateson and Saunders to

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think more critically about, and thus sharpen their elaboration of, Mendelian phenomena. For example, Weldon’s second major challenge, published in Biometrika in November 1902 and entitled “On the Ambiguities of Mendel’s Categories,” successfully identified Bateson and Saunders’ “Achilles heel”; he charged that the criteria both used to characterize “intermediates” in their first Report to the Evolution Committee were highly subjective. As he noted: “The confusion between resemblance to a race and resemblance to an individual involved in Mr Bateson’s treatment of Mendel’s work is one of the many unfortunate results which follow when Mendel’s system of dividing a set of variable characters into two categories, and of using these categories as statistical units, is carried too far” (Weldon, 1902b, 44-5, 46). After Bateson and Saunders began to attract more followers to the Mendelian fold in subsequent years, the biometricians’ continuing criticism of their research program was difficult to deal with, but it nonetheless forced them to think carefully about their experimental design and interpretation.

Conclusion William Bateson holds an ambivalent place in the history of early genetics. On the one hand, he is lauded as the champion of Mendel in the English-speaking world, the founder of one of the most important research programs in the first decade of Mendelism, and the founding father of the science of “genetics.” After 1910, with the rise of the Morgan school’s chromosome theory of heredity, he again became somewhat marginalized owing to his long-time refusal to accept the chromosomes as the seat of the Mendelian factors or “genes.” However, both of these stages in his career become more understandable when one juxtaposes them against the earlier period of the 1890s. As we have seen, Bateson came to Mendelism with well developed conceptions about the physical basis of variation (both discontinuous and gradualistic), the limited role of natural selection, and the saltatory nature of evolutionary change. He was not prone to think in terms of particulate heredity, but rather envisioned physical forces and different chemical combinations as effecting changes in organic matter. His ideas seem to have been greatly influenced by Brooks’s view of heredity and development, and hence to be intimately linked to late nineteenth-century concepts. The rediscovery of Mendel crystallized previously inchoate conceptions he held about the “physiology of heredity,” but his understanding of Mendel was superimposed on existing conceptual categorizations. Such a perspective helps us better understand both Bateson’s early Mendelism and later seeming conservatism with regard to genes and chromosomes. Contrasting Bateson’s approach to variation in the 1890s with that after 1900 reveals the tremendous transformation in thought wrought by Mendel’s work. While Bateson’s earlier study led him along paths similar to the ones Mendel followed, it seems clear that he would have continued to flounder had he not been introduced to Mendel’s mode of analysis. Identifying the working hypotheses that fueled his early problematic also highlight differences in his understanding of Mendelism and that of other early Mendelians, especially de Vries and Correns. Ultimately, their conceptualization of variation and heredity prior to learning of Mendel’s work shaped their subsequent Mendelian interpretations (Stamhuis, 1996, 2005; Saha, 1984; Rheinberger, 1995).

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Bateson’s research program was rightly considered unorthodox and even heretical to neoDarwinians and evolutionary morphologists alike. Even before Mendel’s rediscovery, he forced British biologists to confront two opposing approaches to heredity, the biometricians’ ancestrian view of heredity based on Galton’s law of heredity and his own rival understanding grounded in a qualitative, physiological approach to variation. But there was more to their dispute than simply intellectual property rights. At a time of scarce resources to support the increasingly experimental work in biology, these men were also competing for authority and patronage within the scientific establishment of late Victorian Britain (Farrall, 1975; Sapp, 1987; Marie, 2004). Bateson was fortunate in being able to tap a major new resource within the scientific workforce: the talented pool of women among the first generation to gain university training in the life sciences (Richmond, 1997; 2001; 2006). This gave him a significant edge. On a personal level, contrasting Bateson’s pre- and post-Mendelian work better illuminates radical revisions in his architecture of knowledge, which, in turn, provoked major changes in technical procedures, with the new “Mendelian” regime provoking the adoption of different standards of experimental design and control. This resulted in a new means of presenting data, of analyzing crosses mathematically rather than qualitatively, and application of the new “laws” of heredity to understanding variation. Thus Bateson’s former notion of discontinuous variation produced by physical forces or chemical combinations became refashioned into a new Mendelian knowledge regime. Old concepts were translated into new ones: alternative characters became allelomorphs, parental traits were either dominant or recessive, reversion resulted from the reappearance of latent recessive factors, and swamping balanced by the purity of the gametes. In short, Bateson’s previous study of the physiology of heredity was refashioned into the new field of genetics. Contrasting Bateson’s pre- and post-Mendelian work thus explains how a previously marginal line of work could be transformed into a vigorous research program that emerged as the leading approach to heredity in the decade before the introduction of the chromosome theory of heredity. Marsha L. Richmond Wayne State University, Detroit, Michigan, USA [email protected]

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————. 2004. “Bateson, William (1861–1926).” In Oxford Dictionary of National Biography. Oxford: Oxford University Press, 2004. Pearson, Karl. 1914-30. The Life, Letters and Labours of Francis Galton. Cambridge: Cambridge University Press. Perrone, Fernanda. 1993. “Women Academics in England, 1870–1930.” History of Universities 12: 339–67. Provine, William B. 1971. The Origins of Theoretical Population Genetics. Chicago: University of Chicago Press. Rheinberger, Hans-Jörg. 1995. “When Did Carl Correns Read Gregor Mendel’s Paper?” Isis 86: 612-16. ————. 2000. “Mendelian Inheritance in Germany Between 1900 and 1910. The Case of Carl Correns (1864–1933).” Comptes Rendu de l’Académie des Sciences, ser. 3, Sciences de la vie 323: 1089–96. Richmond, Marsha L. 1997. “‘A Lab of One’s Own’. The Balfour Biological Laboratory for Women at Cambridge University, 1884–1914.” Isis 88, no. 3: 422–55. ————. 2001. “Women in the Early History of Genetics. William Bateson and the Newnham College Mendelians, 1900–1910.” Isis 92: 55–90. ————. 2006. “The ‘Domestication’ of Heredity. The Familial Organization of Geneticists at Cambridge University, 1895–1910.” Journal of the History of Biology 33 (2006): 565–605. Ridley, Mark.1985. “Embryology and Classical Zoology in Great Britain.” In J. A. Witkowski T. J. Horder, and C. C. Wylie (eds.). A History of Embryology. Cambridge: Cambridge University Press. 35–67. Robinson, Gloria. 1979. A Prelude to Genetics. Theories of a Material Substance of Heredity, Darwin to Weismann. Lawrence, KS: Coronada Press. Saha, Margaret Samosi. 1984. “Carl Correns and an Alternative Approach to Genetics. The Study of Heredity in Germany Between 1880 and 1930.” Ph.D. diss., Michigan State University. Sapp, Jan. 1987. Beyond the Gene. Cytoplasmatic Inheritance and the struggle for Authority in Genetics. Oxfort: Oxfort University Press. Saunders, Edith Rebecca. 1897. “On a Discontinuous Variation Occurring in Biscutella Laevigata.” Proceedings of the Royal Society 62: 11–26. ————. 1902. “Part I.—Experiments with Plants.” In William Bateson and Edith Rebecca Saunders (eds.) Reports to the Evolution Committee of the Royal Society 1: 13–87. London: Harrison. Stamhuis, Ida H. 1996. “The Rediscovery of Mendel’s Laws Was not Important to Hugo de Vries (1848– 1935). Evidence from His Letters to Jan Willem Moll (1851–1933).” Folia Mendeliana. ————. 2005. “Hugo de Vries’s Transitions in Research Interest and Method.” In Staffan Müller-Wille and Hans-Jörg Rheinberger (eds.). A Cultural History of Heredity III: 19th and Early 20th Centuries. 115– 36. Berlin: Max Planck Institute for the History of Science, preprint series. Stamhuis, Ida H., Onno G. Meijer, and Erik J.A. Zevenhuizen. 1999. “Hugo de Vries on Heredity, 1889– 1903. Statistics, Mendelian Laws, Pangenes, Mutations.” Isis 90: 238–67. Stern, Curt, and Eva R. Sherwood. (eds.). 1966. The Origin of Genetics. A Mendel Source Book. San Francisco: W. H. Freeman and Co. Tschermak, Erik. 1900. “Über künstliche Kreuzung bei Pisum sativum.” Berichte der deutschen botanischen Gesellshaft 18: 232–39. Weldon, W. F. R. 1890. “The Variations Occurring in Certain Decapod Crustacea. 1. Crangon Vulgaris.” Proceedings of the Royal Society 47: 445–53. ————. 1894. “The Study of Animal Variation.” Nature 50: 25–26. ————. 1902a. “Mendel’s Laws of Alternative Inheritance in Peas.” Biometrika 1: 228–54. ————. 1902b. “On the Ambiguity of Mendel’s Categories.” Biometrika 2: 44–55. Zevenhuizen, Erik. 1998. “Hugo de Vries 1848–1998.” Acta Botanica Neerlandica 47: 405–528.

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“Now that Shull has left us to go to Princeton I fear the oenothera work will suffer,” Charles Davenport, director of the Cold Spring Harbor Station for Experimental Evolution, wrote to Hugo de Vries in 1916. “Still I think that Dr. Blakeslee will keep up with some of it and I hope the opportunity will arise for us to have a man who shall devote a good share of his time to the oenothera. I have not forgotten that in your opening address at this Station in 1904 you recommended this plant especially to our care.”1 Various cytological investigators had struggled for years and would continue to struggle to come to grips with Oenothera’s “normal” karyokinetic idiosyncrasies and the sheer complexity of the phenomena it presented (including, at various points, Davis, Renner, Gates, and Cleland). It wasn’t until Albert F. Blakeslee’s early work at the Station, however, and his even more focused studies on his own chosen model organism, the jimsonweed Datura—that these chromosomal shenanigans began to shed light beyond cytogenetics onto the nature of evolution itself. Blakeslee’s studies of the jimsonweed and his attention to its chromosomal dynamics thus highlight an otherwise largely forgotten early genetical study of the production of non-genic hereditary differences—a “genetics without genes,” as it were. Long before the “genetical systems” and other pathbreaking work of C. D. Darlington and later investigators, Blakeslee’s work in the 1920s relating mutations to differences in chromosomal number and arrangement provides a compelling example within the history of genetics of how the unstable meaning of mutation (the discovery of viable non-genic alterations in the hereditary material), and the choice of a particular model organism kept “heredity” and “genes”—at least for a time—from any easy and instant equation with each other.

Albert F. Blakeslee Having begun his botanical career at Harvard under the mycologist Thaxter in 1904, Albert Francis Blakeslee (“Bert”) first encountered de Vries’ mutation theory while teaching at the Connecticut Agricultural College in Storrs. As he recalled in an autobiographical account, it was in 1909 that he first had “the thrill” of reading de Vries’ theory: “[I] thought that if I scoured the country I too might be able to find a species in the process of mutation.” 2 The mutation-theory was at the core of Blakeslee’s interest in genetics, and both its promise and its unanswered questions sparked his imagination on more than one occasion: “I have always felt that the Mutation Theory was a strong factor in turning my interests and research toward genetics,” Blakeslee later remembered. His interest in de Vries’ theory remained strong for the rest of his life.3 Even as late as 1949, Blakeslee continued to say that de Vries was “perhaps the greatest biologist of all time” and that “[t]he mutation theory is one of the corner stones of genetic research.”4 1 2

Davenport to de Vries, March 2, 1916. Blakeslee to de Vries, January 16, 1933.

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Blakeslee began to search for possible organisms on which to conduct his research. At first he thought he had found a suitable choice with the yellow daisy known as the Black-eyed Susan (Rudbeckia hirta), but he was soon forced to move on to another choice when the daisy proved to be self-sterile and too “reduced in vigor” after two or three crosses to withstand generations of inbreeding—not to mention the generations of inbreeding required for proper detailed research. 5 It was around 1909, while at Storrs, that Blakeslee received from the United States Department of Agriculture “a batch of seeds of Datura stramonium as an example of an economic weed.” The seed “happened to give both purple- and white-flowered seedling,” Blakeslee recalled, “and for several years this species was used to demonstrate Mendel’s laws of inheritance,” in his teaching. 6 (According to Edmund Sinnott, Blakeslee offered what was “probably the first organized course in genetics in the United States” in 1914-1915.)7 On leave from Storrs during the 1912-13 year, Blakeslee went to work at Cold Spring Harbor before finally joining the staff there as a resident investigator in genetics in 1915, replacing the departing Shull. Having devoted considerable attention to genetics in his botany work, it was only natural that he chose to bring his work on the “coarse, weedy plant with its beautiful flowers” with him when he moved.8 Blakeslee was on the hunt not only for the “the best possible ‘Versuchstier,’” as he put it, but for the best possible means to do research with it. Now able to work full-time on genetical problems with better facilities at his command, Blakeslee would over the course of the next 27 years make extensive use of six greenhouses and various agricultural test fields, and ran experiments on a grand scale. Blakeslee had been drawn to the jimson weed for a variety of reasons including its hardy toughness, the ease with which it could be grown, and the fact that four generations could be grown per year in greenhouse environments. “At first,” Blakeslee recalled, echoing newly emerging concerns with Oenothera, his own choice Datura “seemed to have too many chromosomes, but we kept at it as a side problem since it was so easy to work with.” 9 The decision paid off. Blakeslee’s assistant, B. T. Avery, found the first novel type in Datura—the so-called “Globe” mutant—in the summer of 1915.10 As he later reported: “The Globe mutant differs from 3

4

5 6 7 8 9

On the occasion of de Vries’ 85th birthday, Blakeslee wrote to de Vries saying “It is a pleasure to have known such a founder of modern genetics who has been an inspiration to my own work” (Blakeslee to de Vries, May 24, 1933). And in a letter to de Vries’ wife, Blakeslee recalled: “at the quarter centennial of the founding of the Brooklyn Botanic Garden I pointed out an instance of his wonderful prevision in suggesting in 1904, in an address at the dedication of our Department here, that attempts be made to induce mutations by the use of X-rays and radium. My own researches owe much to him. In a measure, I feel that I have been carrying on the torch which he has laid down.” Blakeslee to Mrs. de Vries, May 23, 1935. Blakeslee, “Seventy-Five Years of Progress in Genetics,” This lecture, delivered on November 10, 1949, was “one of a series on Development in Some Fields of Science; organized in connection with the 75th anniversary of the founding of Smith College by Sophia Smith.” Ibid., p. 4; cf. Blakeslee to de Vries, April 7, 1933. Blakeslee, “Lebenslauf,” p. 5; cf. obituary by Edmund W. Sinnott, 1955. Sinnott, 1955. Ibid. Blakeslee, “Lebenslauf,” p. 6. Though not in all respects—as Blakeslee later wrote to a colleague, “some of these species [of Datura] give very poor germination—sometimes not over a tenth of one percent.” Blakeslee to O. L. Inman, December 11, 1934. Blakeslee was also later interested in animal polyploidy, but found this a considerably more difficult task as animals were, as he characteristically put it, “functionally dioecious.” Blakeslee to Emmeline Moore, November 15, 1937.

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normals apparently in all parts of the plant. It forms a complex of characters readily recognized whether the plants in question have purple or white flowers, many or few nodes, or spiny or smooth capsules.”11 This was no ordinary genetic mutation like those found in Drosophila. True to de Vries’ theory, much more than one factor had been affected—the entire plant was different from its ancestor, in a whole suite of traits. Blakeslee became convinced that he had found a new species, and labeled the original new plant specimen as such (“N.S.”), including a photograph of the plant in the 1919 paper reporting the discovery. Although the plant proved sterile with other “normal” plants, it could be selfpollinated successfully and with the appearance of progeny that bred true, producing further generations with “depressed globose capsules,” Blakeslee concluded that it “seems to have established itself as a distinct new race.”12 He continued: This physiological incompatibility between a mutation and the parent species from which it arose suggests that we have actually been witnessing in our controlled pedigrees the birth of a new species which may be capable of maintaining itself in a mixed population uncontaminated by crossing with its ancestral line. The race is relatively vigorous.13

In the caption to the photograph included in the paper, Blakeslee put the point more plainly: “Tests have shown that this mutant differs from all others investigated in that it breeds true as a distinct new race. Here we appear to be witnessing the birth of a new species.” 14 As Blakeslee, Avery, and his other assistants bred the “globe” mutants, they rapidly discovered that still “other types appeared as mutants in our cultures, and Datura soon became practically our sole object of investigation.”15 As one observer at the Station recalled: One new form after another began to appear in his cultures. Some were gene mutations but many were evidently different. These produced some offspring like themselves but threw many normal plants. For an outsider to recognize these forms was difficult, since most of their differences were subtle ones. It was the despair of his colleagues to see Blakeslee go down a row of plants and pick out these mutants unerringly. This he could do partly because of his acute powers of observation and partly because he was personally familiar with his material and did not leave the observing and recording to his assistants alone… The size of the Datura cultures increased and in the summer as many as 70,000 plants were grown. Work was actively carried on in the winter, as well, in the six greenhouses and laboratories.16

Blakeslee was even able to identify types that while “indistinguishable in gross appearance from each other,” nevertheless, “in respect to a whole series of characteristics [are] strikingly different 10 11 12 13 14

15 16

Blakeslee, “Lebenslauf,” p. 5. According to Sinnott, Blakeslee had encountered “one or two Jimson weeds which were different from the typical ones and had begun to study them,” while at Storrs (Sinnott, 1954). Blakeslee 1921. Blakeslee and Avery, 1919. Blakeslee and Avery, 1919. Blakeslee and Avery, 1919. As Blakeslee later recounted in 1921, “It may be mentioned that the tetraploid datura was called ‘New Species’ before its tetraploid nature was suspected. It satisfied the requirements of an independent species. The pollen was relatively good, and the mutant formed a distinct race, selffertile and fertile inter se, while practically sterile with the parent stock.” Blakeslee, “Types,” 1921. Blakeslee, “Lebenslauf,” p. 6. Sinnott, 1955, pp. 9, 8.

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from the normal Jimson Weed from which they have been made up to order, as it were, with definite plan and purpose.” Blakeslee eventually found three in particular that he thought “perhaps merit the term of synthesized new ‘species,’ since they satisfy the criterion of breeding true and are more different from the normal type than some of the species which already have been described in the genus Datura.”17 He took these newly encountered mutants to be indicative that they had encountered a situation in Datura similar to that which de Vries had encountered in Oenothera. A year after his initial discoveries, Blakeslee made further explicit reference to the “increasing rôle in experimental evolution” of the de Vriesian “theory of mutations” that had first been laid forth two decades earlier.

Chromosomes Regnant Unlike the drosophilists who fairly readily shared their stocks and data across the fly room and with other centers of fly research, Blakeslee “kept strict control” of his Datura results.18 But Blakeslee was nothing if not collaborative: having collected the seeds of ten different species of Datura from around the world,19 he engaged in a series of ongoing collaborative ventures over the years, working with (among others) the geneticist Edmund W. Sinnott, an expert in the internal anatomy of the Daturas who could recognize most mutants from tissue sample alone (and who also happened to come from Blakeslee’s old stamping grounds in Storrs), and J. T. Buchholz, an expert on “the growth of pollen-tubes and the abortion of ovules as problems in developmental selection.”20 One of Blakeslee’s earliest and ongoing collaborations was with the cytologist John Belling, who had joined Blakeslee’s group in 1920 and helped him in his “study of the nuclear condition of our mutants.”21 Blakeslee, Belling, and the greenhouse manager M. E. Farnham published a “preliminary report” of their findings in Science in 1920.22 And it was Belling’s cytological work— studying the appearance and behavior of chromosomes—that was later held to have given “the greatest possible assistance in the interpretation of the originally baffling phenomenon of mutation in Datura.”23 Indeed, it was largely as a result of this “fruitful association” with Belling— as well as the “development of the aceto-carmine staining method” that permitted chromosomes to be “counted directly in smear preparations”—that Blakeslee was rapidly able to establish that “each mutant was the result not of a gene difference but of a third chromosome added to a particular pair of the twelve in this plant.” Such mutants were termed “trisomics” or “2n+1” types. More generally, this discovery enabled Blakeslee at last to interpret his results on a chromosomal rather than a genic basis that would presumably have required the joint mutation of a number of different genes at the same time.24 Blakeslee’s mutant plants differed by a whole “complex of 17 18 19 20 21 22 23 24

Blakeslee 1932. Demerec, 1959. Sophie Satina, undated. “Department of Genetics,” 1922, p. 95. Blakeslee, 1922, p. 18. Blakeslee, Albert F., J. Belling and M. E. Farnham, 1920. “Department of Genetics,” 1921, p. 108. Demerec, 1959.

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characters,” that were “transmitted collectively,” and that segregated “in a very unusual fashion”—and chromosomal observation and analysis was soon to explain exactly how these phenomena came to pass. While acknowledging that it was “sudden germinal changes, large or small in amount” that were the basis of “perhaps the most fundamental work in modern genetics,” Blakeslee noted that “mutations could not be confined to cells associated with sexual reproduction.” In an apparent reference to the remarkably productive and groundbreaking work of the drosophilists and other more gene-oriented investigators, Blakeslee emphasized that botany had already applied the mutation concept in ways that extended far beyond the genes that many animal geneticists were most concerned with. Somatic mutations, for instance, were those mutations that took place in cells in which sexual processes were not involved. While fairly “less common phenomena in animals,” such somatic mutations—or “bud sports” as they were also frequently called—were common in plants and many were even quite well-known. Such instances of mutation were real, and yet they were clearly beyond the ken and the techniques of the drosophilists—no matter how powerful and innovative these investigators were in identifying and mapping mutant genes. Blakeslee argued that these characteristics, including those whose “inheritance could not be established by breeding experiments,” had been and should continue to be called “mutations.” 25 Blakeslee also held, therefore, that his new discoveries of polyploidy, trisomy, and the other phenotypic effects of chromosomal alterations were thus novel additional worthy instances of mutation: To us, one of the most interesting features of the Datura work is the possibility afforded of analyzing the influence of individual chromosomes upon both the morphology and physiology of the plant without waiting for gene mutations… Our work so far we believe adds evidence to the conclusion that the mature organism—plant or animal—is not a structure like a child’s house of blocks, made up of separate unit characters, nor is it determined by separate and unrelated unit factors. It is rather the resultant of a whole series of interacting and more or less conflicting forces contained in the individual chromosomes.26

Blakeslee fully acknowledged that classical Mendelian research up until this time had “dealt almost exclusively with disomic inheritance.” 27 But, he noted, “[d]istinct variations, provisionally termed mutations… [have] regularly recurred whenever a sufficiently large number of plants have been subjected to observation,” and that these, “[s]o far as investigated… have been found to be connected with a duplication of one or more of the normal chromosomes.” 28 Blakeslee’s mutant plants thus revealed that phenotypically distinct mutations could result from genically identical types, simply with different arrangements or numbers of chromosomes. 29 Mutation could thus 25

26 27 28

Intriguingly, Blakeslee held that the “failure” of a particular mutation in the adzuki bean “to appear more than once in so large a number of individuals indicates that it is a variation genotypic in nature, since it could scarcely be attributed to the reappearance of a character through normal segregation nor be considered a mere modification induced by environmental factors.” The sheer rarity of the mutation was an argument for its genotypic, rather than its chromosomal, basis. Blakeslee, 1919. Blakeslee, 1922, p. 31. Emphasis added. Blakeslee, 1922, p. 27. Blakeslee, 1921, p. 255. Blakeslee later realized, of course, that duplication was not the only means. Following Calvin Bridges’ work on nondisjunction, he acknowledged that there was room for a “rather novel study of trisomic, tetrasomic and pentasomic inheritance.” Blakeslee, 1922, p. 27.

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take place at a level that was neither organismic nor genic. A mutation, therefore, did not need to be genic in order to be genetic. In short order, Blakeslee and his collaborators, colleagues, and competitors identified many other varieties of “chromosomal mutants” over the years—including reciprocal translocation among trisomics, the existence of haploids in higher plants (theretofore unknown), and even mutants with chromosomes arranged in sets and rings (precisely that phenomenon determined to be responsible for the seemingly endless bedeviling of an earlier generation of investigators of Oenothera). While the drosophilists had of course acknowledged the phenomenon of nondisjunction at the microscopic level, it was Blakeslee who connected the dots to the effects at the phenotypic level and brought the effects of trisomy, nondisjunction and other chromosomal phenomena into the realm of possible mechanisms for mutation. As Davenport noted: “it has remained for Datura to reveal in the hands of Blakeslee and his associates, Belling, Farnham, and others, an extensive system of inter-chromosomal mutation and corresponding somatic change the like of which had been entirely unknown.”30

The Meaning of Mutation The wider community of geneticists and other students of heredity were already well aware that it appeared possible to make a definite distinction between the two kinds of mutation thus far readily observed, and whether chromosomal abnormalities were “mutations” became a matter of debate in the field. Although Shull initially seemed to agree with the designation of chromosomal aberrations as mutations—“You go so far in the solution of the change which brings about the occurrence of the Globe mutants that it seemed to me you were justified in applying the more fundamental term ‘mutation’ as a title of your contribution”—a week later in 1921 he coined a new word for such chromosomal mutations and tried to get Blakeslee to use it. (The word was “anomozeuxis.” As Shull noted, “I feel fairly certain that your first reaction to these words will be unfavorable, but they are words which grow easier to say and pleasanter to look at as you become more familiar with them.”)31 By traditional observable botanical and morphological criteria, and by the simple fact that they bred true, Blakeslee’s plants were clearly mutants and any botanist (as de Vries himself had often remarked) would have classified such new organisms discovered in the field as belonging to a new species. By the standards of the drosophilists and some other geneticists, however, these were clearly not new mutants but merely chromosomal aberrants. And yet, as Blakeslee himself reported, “The pure breeding types are more distinct from the original form from which they arose than some of the species of Datura which have been founded on single factor differences. Our types we have ventured to call artificial or synthesized ‘new species.’” In the early 1920s, Blakeslee was fully aware of the polyvalent meaning of “mutation” and of the declining influence of De Vries’ theory among biologists of all stripes. Having laid out the 29 30 31

Relating the existence of these chromosomal types to geographic distribution patterns also did much to help illuminate the evolutionary history of Datura (Sinnott, 1954, pp. 394-8). “Department of Genetics,” 1922, p. 93. Shull to Blakeslee, April 26, 1921.

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relevant details—from the drosophilist H. J. Muller’s work on balanced lethals in the teens to the importance of the study of the behavior, association, and mechanism of chromosomes and chromosomal duplication and polyploidy—Blakeslee asked in 1921: What then is a mutation? I do not feel we need to be bound by its application to the evening primrose for reasons of priority, since Waagen… had previously used the term in paleontology in an entirely different sense. I believe, with the idea that mutations must involve a qualitative change, that we shall ultimately confine the term to mutations of genes, although such mutations may later be shown to be as different from our present conceptions of them as are mutations in the Oenotheras from the conceptions in de Vries’s classical publication, ‘The Mutation Theory.’ It may still be desirable to employ the word mutation as a collective term to designate the sudden appearance of any apparent genetic novelty—whatever its real cause—until we know better.32

Despite claiming that the fundamental meaning of mutation might ultimately be genic, Blakeslee therefore recommended agnosticism on the matter. His own research program, however, was structured around the idea that chromosomal aberrations were not only an important source of variation, but were perhaps even the fundamental mechanism for the production and maintenance of many “new species” of mutants plants. In fact, Blakeslee spent almost no time whatsoever discussing genic mutations in his writings, confining his attention to the significance of chromosomal mutations alone. In a 1921 article purporting to address the various “Types of Mutation,” Blakeslee discussed some of the many varieties of chromosomal mutation, and concluded by saying: There is not time at my disposal to discuss mutations of genes... It has not been possible in this brief presentation to give an extended classification of mutations, nor to discuss in detail their possible significance in evolution. It will be sufficient if I have made clear the distinction which must be kept in mind, in any discussion of the subject, between mutations in individual genes and those brought about by chromosomal aberrations. 33

In all, Blakeslee’s approach represented a distinct modification and reworking of de Vries’ theory.34 Although Blakeslee acknowledged that “[s]trictly speaking I should not call chromosomal aberrations mutations when the changes are purely quantitative”—such as in the case of polyploidy—the accompanying table in his article on “Types of Mutations and Their Possible Significance in Evolution” labeled just these forms precisely in that way. 35 The meaning of mutation for geneticists was unstable, and it was unstable even for Blakeslee himself. He was 32 33 34

35

Blakeslee, 1921, p. 261. Blakeslee, 1921, pp. 262, 265-6. For a case of tetraploidy, for example, not to be considered a mutation was as significant an alteration of de Vries’ theory as is conceivable, as de Vries himself considered the origin of the tetraploid Oenothera gigas to be “the one absolutely typical case of species-formation in all my cultures.” He prefaced his remark by saying: “Please tell Miss Lutz that I enjoyed her discovery of the double number of chromosomes in Oenothera gigas immensely” (de Vries to Davenport, December 31, 1907). Blakeslee went on to insist, however, that “[t]he occurrence of tetraploidy would therefore be no more a mutation than the doubling of chromosomes at the origin of the sporophyte from the gametophyte ferns.” Blakeslee, 1921, pp. 262-3. Blakeslee, 1921, pp. 262-3.

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never opposed to gene mutation: he was fully aware of the drosophilists’ genocentric focus, and gave their understanding of mutation a certain priority (as he once said: “We have seen that chromosomal duplications and related phenomena may simulate gene mutations in their effects upon the individual.”) And yet, in his reworking of de Vries’ theory, Blakeslee’s focus always resolutely remained on understanding the significance of what he called alternately “chromosomal mutations” and “chromosomal aberrations”: “What is their possible significance in evolution?”

Blakeslee and Gager Blakeslee laid out the problem: if plant mutants were due to alterations in chromosomes and not just in genes, then “it should be possible by breeding tests to connect up mutants with as many chromosome sets as there are known Mendelian factors, or factor groups.” This, however, was not readily the case, as there were unusual situations (such as various forms of chromosome duplication) where these varied effects also needed to be taken into account. The discovery of what were termed “balanced” and “unbalanced” types—that is, mutative variants with all paired chromosomes, and types where an additional chromosome was left unpaired—provided for a new means of exploring the influence of general mutation. In effect, Blakeslee argued, it meant there was now a means to avoid having to depend on the random appearance of mutations in a population: The unbalanced condition gives us an opportunity, never before realized, of analyzing the influence of individual chromosomes without waiting for the appearance of gene mutations. Heretofore, the number of factors determined in the chromosomes has been dependent upon the number of mutated genes available for crossing with the normal type. In the jimsons, however, we may study the sum total of all the factors in individual chromosomes by the unbalancing effect upon the structure and physiology of the plant when a single specific chromosomal set has 1 or 2 extra chromosomes.36

Already by 1921, Blakeslee claimed to have discovered three so-called “factor” mutations and twelve “chromosome” mutations in the jimson weed, all of which were “identified by various external characters.” But these numbers were to continue to steadily increase throughout the 1920s and 1930s. “Knowing the mechanism to be affected,” Blakeslee noted in 1921—that is, the behavior, mechanism, and association of the chromosomes—“we may be able ultimately to induce chromosomal mutations by the application of appropriate stimuli.” 37 Radium was one of the first of those stimuli to which Blakeslee turned. Charles Stuart Gager, the director of the Brooklyn Botanic Garden, was the first to investigate the effects of the rays of radium on plants in 1908. In 1921, Blakeslee began a fruitful collaboration

36 37

“Department of Genetics,” 1921, p. 104. Blakeslee even cited Muller’s work on balanced lethals, which he said “strongly suggests that such of the Œnothera mutants as are not caused by chromosomal duplication are due to cross-overs from a balanced lethal condition.” Blakeslee, 1921, pp. 257, 260, 262.

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with Gager, and described the goals of the collaboration in a presentation to the Botanical Society of America on December 28 of that year: to study and compare the structure of these mutant forms, both as to gross external morphology and as to internal anatomy; and thus to determine the structural effects produced by a single factor and those produced by a single entire chromosome. In this way it may be possible to begin an analysis of the factorial constitution of each of the chromosomes.38

Blakeslee’s approach to mutation studies was thus intended to complement other studies in the field, and to better highlight the different ways in which mutations could be produced—both chromosomally and genically, and both by the addition of single triplicate chromosomes to the mix (his current focus), and by other changes that had already been identified (such as rings of linked chromosomes). Phenotypic effects—mutant plants—could result from any of these mechanisms. Although Blakeslee and Gager couched their approach in terms of gene-based genetics, their discoveries were soon to push them ever further toward acknowledging the primacy of chromosomal variation in evolution. Already by 1921, Blakeslee and Gager encountered a peculiar mutant, “Nubbin,” which they noted clearly arose from a “radium-treated parent,” and which was likely the particular result of ray-induced “breaking up and the reattachment of parts of non-homologous chromosomes.” 39 (As Blakeslee later reported in the Year Book, some of the “three chromosomes were fragments, and the fragments of one were attached each to a fragment of the other two.”) 40 With its interchanged chromosomes, Blakeslee thought Nubbin was thus “probably the first induced chromosomal mutation.” 41 He also held that an albino character might also have been due to radium treatment.42 In short, Blakeslee believed that the radium treatment certainly increased the proportion of mutants, but he remained open-minded as to whether it could cause new gene mutations—such as the albino mutant—waiting for evidence that such traits acted as mendelizing characters.43 By the following year, the two men had begun to compose a draft paper, eventually to be published in the Proceedings of the National Academy of Sciences. Production of their paper became bogged down for a period of years, as both the inherent difficulties of the project and Gager’s other commitments kept him away from the radium work. By the dawn of 1927, Gager wrote to Blakeslee saying, “I have just glanced the paper through. Apparently, it will need very considerable revision, if not re-writing. Among other things, it might be desirable to mention the results of Mavor on the production of non-disjunction and crossing over… by X-rays, though reference to those papers should, I think, be very brief.”44 James Mavor’s results, published in Science in 1922 38 39 40

41 42 43 44

Ibid. Gager and Blakeslee, 1923, pp. 75-6. “Nubbin, a type obtained following radium treatment by Dr. Gager in 1921 in which 3 chromosomes were fragments and the fragments of one were attached each to a fragment of the other two, has been of considerable service in the analysis of the cryptic types in nature.” “Department of Genetics,” 1929, p. 45. Blakeslee, “Lectures, Papers, Etc.,” “Control of Evolution and Life Processes in Plants.” “Department of Genetics,” 1922, p. 98. Blakeslee also acknowledged, however, that some mutations were not expected to be mendelizing. Blakeslee to Gager, January 14, 1923. Gager to Blakeslee, January 3, 1927.

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under the title “The Production of Non-Disjunction by X-Rays,” had indicated that the phenomena of nondisjunction first identified in Drosophila by Bridges (the cause of various heritable traits though not specifically a genic mutation) could also be produced artificially.45 Fully aware that some of D. T. MacDougal’s earlier successes with induced mutation had come into question, Blakeslee and Gager were concerned that their own work not fall prey to the same criticisms. Though certain that they had discovered two radium-induced mutations, Blakeslee nonetheless advocated caution: “It seems to me that in view of the trouble which McDougall [sic] got into with his induction of mtations [sic] it behooves us to be extremely cautious, perhaps unnecessarily so, in claiming much for our preliminary experiment.” 46 All in all, he concluded, “I am wondering if we ought not to do a little more work with the radium and get more than an isolated capsule effected [sic] before we get out a formal paper.”47 After years of delay, their joint paper “Chromosome and Gene Mutations in Datura Following Exposure to Radium Rays” finally appeared in the Proceedings of the National Academy of Sciences in February 1927.48 While they acknowledged that when they first presented their results in 1922 they had not yet “a sufficient body of data in regard to the mutability of untreated parents to permit us properly to evaluate the significance of the results,” they now claimed to have accumulated “considerable” data regarding both “gene and chromosomal mutations in closely comparable normal material which can be handled as control to the treated material.” 49 Finding great surprise in their success, they reported that they had discovered a variety of what they called “chromosomal mutants” mostly of the 2n+1 form—having a complete diploid set of chromosomes with an additional chromosome. Although these types of chromosomal mutants had first been mentioned in the Anatomical Record as early as 1923, what was significant in Blakeslee and Gager’s new publication was the sheer rate of production of these mutants.50 While overall they had discovered some 73 “2n+1” forms from 15,417 progeny in the controls (a rate of 0.47%), in one case they found “[a] percentage of 17.7 chromosomal mutants in over 100 offspring from a single capsule”—a rate they described as “enormously greater than [that] we have ever obtained before or since.” They concluded: “In view of the above figures, we believe the radium treatment was responsible for the increased proportion of chromosomal mutations, as also for the appearance of the compound chromosomal type Nubbin.”51 (Recall that drosophilists had, at this time, discovered about 400 visible mutants from their study of some 20 million flies—Blakeslee and Gager’s results were thus by all standards remarkable.) The end result of their collaboration was clear. There was no longer any doubt that the radium could transmute species, and that it did so in at least two different ways: gene mutants (as the drosophilists had found) and chromosomal mutants. 52 45 46 47 48 49 50 51 52

Mavor, 1922. Blakeslee to Gager, January 14, 1923. Blakeslee to Gager, January 14, 1923. Gager and Blakeslee, 1927, pp. 75-79. Ibid. p. 75 Gager and Blakeslee, 1923, p. 424; Blakeslee, 1923, p. 389. Gager and Blakeslee, 1927, p. 78. “[I]t is our belief that most, if not for all, of these three types of results”—the compound chromosomal type Nubbin, the chromosomal mutants, and the gene mutants—“the radium treatment may be held largely responsible,” they concluded (ibid., p. 79).

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Much of Blakeslee’s work in the 1920s thus centered around identifying the various kinds of “apostles” and “acolytes,” as he termed these different varieties of “chromosomal types.” He mapped out the theoretical possibilities of combinations, and charted which ones he observed and with what frequency. And he found these chromosomal types to be related to phenotypically distinct and self-perpetuating “new species” of Datura. He invented a whole terminology for these new chromosomal types, to categorize the cytogenetic differences: “primaries” were 2n+1 trisomics with an additional but unmodified chromosome; “secondaries” were such trisomics with two like ends, the result of further chromosomal interchange; and “tertiaries” were trisomics with ends from two different chromosomes. He invented diagrammatic karyotypes, explaining these processes of chromosomal interchange and the creation of mutant chromosomes, which in turn were responsible for chromosomally mutant plants, and he related these diagrammatic karyotypes to the phenotype. Blakeslee and Gager established that “synthesized pure breeding types, which correspond to synthesized new ‘species’” resulted from radiation treatment. Blakeslee was firmly convinced that these synthesized pure breeding types—the result simply of chromosomal and not gene mutation—were indeed new species in an evolutionary sense: they bred true, generation after generation, and presented themselves as new types to the botanist. 53 Although one of the first to strongly advocate polyploidy, Blakeslee was also aware of other effects that were clearly the result of gene mutations—although the direct relevance of these for evolutionary processes (the emergence and maintenance of new species) was not as readily apparent. (These visible effects included altered pollen tube growth, the non-germination of pollen, and the early or late abortion of pollen grains.) Outside the world of drosophilists, it was not at all clear that gene mutations were in any way more fundamental to the nature of evolution and the origin of species than the chromosomal mutations Gager and Blakeslee were uncovering. Blakeslee’s emphasis on the significance of chromosomal mutation was long-standing. He had written to MacDougal as early as 1923 saying, “I feel very strongly that a study of the chromosomal distribution is likely to explain irregularities in behavior in other plants than the Datura and that chromosomal changes in number have been responsible for evolution.” 54 In the wake of Gager’s work and the widespread realization of the complexity of Oenothera’s chromosomal system, Blakeslee was also aware however, and most especially at the Boston meeting in 1922, “that I have been obliged to caution people with whom I have talked about the Datura work from being overenthusiastic and thinking the chromosome irregularities would explain phenomena which appeared to be explainable on ordinary factorial basis.”55 Enthusiasm for chromosomal mutations as the basis for evolution apparently outstripped enthusiasm for gene mutations in some quarters. Both chromosomal and gene mutations were important for Blakeslee. 56 They were not equally important for everyone else at the time. Gager and Blakeslee had published their paper on “Chromosome and Gene Mutations in Datura Following Exposure to Radium Rays” in the February 1927 edition of the Proceedings of the National Academy of Sciences. By July 22, Science 53 54 55

Blakeslee to Gager, September 5, 1933. Blakeslee to MacDougal, February 15, 1923. Ibid.

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published results on the induction of mutations in Drosophila under the provocative title “Artificial Transmutation of the Gene.”57 The author was none other than one of the archetypal figures in the history of genetics, the ever-priority-conscious Hermann J. Muller. History was about to be rewritten—and Blakeslee and Gager’s successes with radium were about to be written out of the picture in favor of Muller’s experiments with X-rays, and his focus on the gene as “the basis of life.”58 Blakeslee’s work was rapidly overshadowed as Muller’s own remarkable successes—a 150% increase in genic mutation—hit the headlines, contributing to the establishment of mutation as a fundamentally genic phenomenon. Chromosomes were important, certainly, but they had been dethroned.

Conclusion Recognition of the importance of chromosomes—and not just genes—in the phenomena and study of heredity was widespread, especially in botanical circles, in the early twentieth century. Blakeslee’s work linking questions of mutation to questions of chromosome structure, directly influenced by de Vries’ own focus on plants, thus provides a counternarrative to the dominant tale of gene-centered Drosophila genetics, and recovers a history otherwise lost in the afterglow of Muller’s 1927 experiment. Mutation meant many things to many people, even among “geneticists.” While Muller regularly racked his brain trying to make further distinctions between “true” gene mutations and the smallest conceivable changes in chromosomes, in order to discover the “basis of life” as it kept on slipping through his fingers, Blakeslee was a more ecumenical mutationist concerned to study the effects of both gene and chromosomal mutations, recognizing polyploidy, trisomy, and various forms of multiple linkages and translocations all as distinct and proper forms of chromosomal mutation with definite, observable, inducible, and manipulable phenotypic effects. Moreover, while some plants deal well with polyploidy and trisomy and other vagaries of chromosomal interchange—and these include Oenothera and Datura, for whom these mechanisms are chromosomal normalities, not abnormalities—fruit flies simply do not. The choice of experimental organism mattered for the meaning of mutation in this period. Muller’s meteoric rise to fame after 1927 conspired with his focus on the fruit fly, the X-ray, and the gene to eclipse chromosomes from their rightful place in the story of evolution until the later “genetic systems” of Darlington. Although largely forgotten today, Blakeslee’s work on the jimsonweed led him to a vision of a more pluralistic genetics. Not just producing karyotypes and mechanistic explanations for de Vries’ oddities, Blakeslee had in fact, sometimes even against the grain of his 56

57 58

This was in distinct contrast to some other earlier investigators. In an article for the American Naturalist in 1910 entitled “Mendelian Phenomena Without De Vriesian Theory,” William Spillman had proposed four distinct types of “variation”: the Mendelian recombination of characters; fluctuation due to the environment; the discontinuous hereditary “irregularities in the distribution of chromosomes… amenable to the action of natural selection” (or as he also labeled it, in light of new understandings of what was going on cytologically with Oenothera, “de Vriesian mutation”); and what he called “fundamental change in… the germ plasm,” this last of which he believed to be “by far the most important type of evolutionary change.” Muller, 1927. I have addressed Muller’s rewriting of history elsewhere; see Campos, “Mutatis Mutandis: H. J. Muller and the Meaning of Mutation,” forthcoming.

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own inclinations and earliest pronouncements, initiated productive new ways of doing “genetics without genes.” Luis Campos Drew University [email protected]

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Archival Correspondence Cited correspondence is archived in the Blakeslee papers and the Davenport papers at the American Philosophical Society in Philadelphia, Pennsylvania (“APS”); and in the Blakeslee papers at the Cold Spring Harbor Archives (“CSH”). Blakeslee, Lebenslauf of A. F. B. In APS Blakeslee, biographical materials, folder 2. Blakeslee, Lectures, Papers, Etc., Control of Evolution and Life Processes in Plants, APS Blakeslee. Blakeslee to de Vries, January 16, 1933. APS Blakeslee, “de Vries.” Blakeslee to de Vries, April 7, 1933. APS Blakeslee, “de Vries.” Blakeslee to Mrs. de Vries, May 23, 1935. APS Blakeslee, “de Vries.” Blakeslee to Gager, January 14, 1923. APS Blakeslee, “Gager.” Blakeslee to Gager, September 5, 1933. APS Blakeslee, “Gager.” Blakeslee to O. L. Inman, December 11, 1934. CSH. Blakeslee to MacDougal, February 15, 1923. APS Blakeslee, “MacDougal.” Blakeslee to Emmeline Moore, November 15, 1937. CSH. Blakeslee, Seventy-Five Years of Progress in Genetics. In APS Blakeslee, “Blakeslee, Lectures, Papers, Etc.” Davenport to de Vries, March 2, 1916. APS Davenport, “Vries, Hugo de.” de Vries to Davenport, December 31, 1907. APS Davenport, “Vries, Hugo de.” Gager to Blakeslee, January 3, 1927. APS Blakeslee, “Gager.” Sophie Satina, biography of Blakeslee. APS Blakeslee. Shull to Blakeslee, April 26, 1921. APS Blakeslee, “Shull,” folder 3.

References Blakeslee, Albert F. 1919. “A Unifoliolate Mutation in the Adzuki Bean.” Journal of Heredity 10: 153-55. Blakeslee, Albert F. 1921. “The Globe Mutant in the Jimson Weed (Datura stramonium).” Genetics 6: 24164. Blakeslee, Albert F. 1921.“Types of Mutations and their possible significance in evolution.” American Naturalist 55: 254-67. Blakeslee, Albert. F. 1922. “Variations in Datura due to changes in chromosome number.” American Naturalist 56: 16-31. Blakeslee, Albert F. 1923. “Distinction Between Primary and Secondary Mutants in Datura.” Anatomical Record 26: 389. Blakeslee, Albert F. 1932. “Methods of Synthesizing Pure-Breeding Types with Predicted Characters in the Jimson Weed.” Science 76: 571. Blakeslee, Albert F. and B. T. Avery, Jr. 1919. “Mutations in the Jimson Weed.” Journal of Heredity 10: 111120. Blakeslee, Albert F., J. Belling and M. E. Farnham. 1920. “Chromosomal duplication and Mendelian Phenomena in Datura Mutants.” Science 52: 388-90. Campos, Luis. Forthcoming. Mutatis Mutandis: H. J. Muller and the Meaning of Mutation. Demerec, M. 1959. “Albert Francis Blakeslee.” Genetics 44 (1959): 1-4. Department of Genetics. 1921. Carnegie Institution of Washington (CIW) Year Book 20: 101-156. Department of Genetics. 1922. Carnegie Institution of Washington (CIW) Year Book 21: 93-125. Department of Genetics. 1929. Carnegie Institution of Washington (CIW) Year Book 28: 33-66. Gager, C. Stuart and A. F. Blakeslee. 1923. “Induction of gene and chromosome mutations in Datura by exposure to radium rays.” Anatomical Record 24: 424. Gager, C. Stuart, and A. F. Blakeslee. 1927. “Chromosome and Gene Mutations in Datura Following Exposure to Radium Rays.” Proceedings of the National Academy of Sciences 13: 75-79. Mavor, James W. 1922. “The Production of Non-Disjunction by X-Rays.” Science 55: 295-7. Muller, H. J. 1927. “The Artificial Transmutation of the Gene.” Science 66: 84-7. Sinnott, Edmund W. 1954. “Albert Francis Blakeslee (1874-1954).” American Philosophical Society Year Book: 394-8. Sinnott, Edmund W. 1955. “Albert Francis Blakeslee, November 9, 1874 – November 16, 1954.” National

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Academy of Sciences Biographical Memoirs: 1-38. Spillman, W. J. 1910. “Mendelian Phenomena Without De Vriesian Theory.” American Naturalist 44: 216.

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Seeing, Breeding and the Organisation of Variation: Erwin Baur and the Culture of Mutations in the 1920s Alexander von Schwerin

This paper seeks to introduce the research on mutations of the German physician, botanist and geneticist Erwin Baur (1875-1933). It traces Baur’s experimental studies on the heredity of plants from 1903 onwards, particularly focusing on the snap dragon (Antirrhinum majus). A sophisticated approach to the detection of mutations evolved out of these studies around 1920. In the mid-1920s, Baur managed to create a sudden blast of mutations in his experimental object. The crucial invention was a special system of breeding. In this respect, Baur’s experiments were special since in the 1920s mutation research had become dominated by manipulative trials. The artificial induction of mutations turned out to be the more spectacular result in the perspective of the contemporaries. Highlighting the practical and conceptual trajectory of Erwin Baur’s research on mutations, this paper suggests that this view is limited on the conceptual impact of mutations. However, the impact of mutations and the techniques directly related to them were not limited to the very field of mutation research. They increasingly became a technical boundary between variant interests. Baur and other geneticists held not only a fundamental interest in the causes of the variation of organisms, but mutants bound together agricultural interests, evolutionary commitments and eugenic sentiments. Last but not least, mutants were tools and became part of the growing technical spectrum of biological and biomedical research. 1 Instead of reducing variation, geneticists became involved in broadening variations for multiple purposes.2

Starting the story: A new type of mutation as a tool for innovative research In 1934 two leading German geneticists, Alfred Kühn and Nicolai Timoféeff-Ressovsky, utilized a simple distinction when speaking to a mixed auditorium of biologists and physicians at the “scientific week” in Frankfurt (Main). “Good mutations” were those mutations that produced quite visible and striking changes in an organism. Correspondingly, “bad mutations” resulted in nearly invisible changes.3 By using this distinction, Kühn and Timoféeff-Ressovsky resumed the common view of experimentalists in hereditary research. “Good mutations” were “good” in a pragmatic sense because they were useful for ordinary genetic research, such as the mapping of genes or the analysis of traits. Of course when scanning hundreds of flies under the microscope, it was easier to detect a mutant of the Drosophila fly with striking morphological deformities such as

1 2

3

For the history of the institutionalisation of experimental animal breeding see Rader, Mice 2004. This approach towards a history of the formation of a mutational dispositive supplements, see Christophe Bonneuil’s history of the generation of genetic pureness as an “experimental/combinatory/ industrial time-space”. Same preprint. Timoféeff-Ressovsky, Verknüpfung 1935, pp. 101 f.

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missing wings rather than “small changes” such as slight variations in the structure of the wings or of the mean life time.4 However, small changes were “bad” only from the perspective of common genetic practice. The message of Kühn and Timoféeff was actually that small mutants had a special value for research. These mutants were not only the proper material of evolution, but were also useful for innovative genetic research. “Small or weak and variably manifesting” mutations were good as a tool for innovative research such as that on the physiology of genes. 5 Kühn argued that “today the complicated cases are more important for us because they pave the way into the effective gearing (“Wirkungsgetriebe”) of the genes.”6 Speaking of “small mutations” (“Kleinmutationen”), Kühn and Timoféeff-Ressovsky suggested that there was a special class of mutations. However, the term “Kleinmutationen” was not coined by them. In using it they referred to another German geneticist: Erwin Baur. Baur (b. 1875) belonged to the first generation of Mendelian geneticists in Germany and had died in late 1933. Until then, he had managed the large Kaiser Wilhelm Institute for Breeding Research at Müncheberg near Berlin. Baur was known for his style leading the institute like a business. 7 He and his assistants held close relations to farmers and the agricultural industry—seed breeders, breeders of poultry and rabbits and the coat manufacturing industry. 8 Baur was also actively engaged in research politics and had been a notorious promoter of human genetics and eugenics. 9 Less well known are his studies in mutations, though these were quite well esteemed by geneticists at that time. To be sure, “Kleinmutationen” were by definition somewhat unusual in genetics. They were not new in terms of their mode of inheritance and were actually alterations of a hereditary factor, or a so-called factor mutation. Small mutations were noteworthy because of their effects. Geneticists usually referred to those mutations that were not initially noticeable as “recessive” mutations. In contrast to dominant alleles, they became visible only when they became homozygous. Still, in the case of small mutations the manifested effects were only slight, thus, small mutations were somewhat of a subclass of recessive mutations. But natural kinds are always on the move. The distinction between “recessive” and “dominant” genes was fading in the twenties. Timoféeff-Ressovsky classified mutations only phenomenologically since there was a “complete scale” of mutations “beginning with quite visible morphological changes to all kinds of small physiological changes,” that is “from ‘big’ to very ‘small’ mutations.” 10 Small mutations opened a new space that fell between changes induced by mutations and the realm of non-hereditary modification. However, a special situation was necessary that made this

4 5

6 7 8 9 10

Timoféeff-Ressovsky’s models in the late 1920s were flies with slight changes of the wings and with reduced viability. Timoféeff-Ressovsky, Verknüpfung 1935, pp. 95-99. Timoféeff-Ressovsky, Verknüpfung 1935, p. 102; Schwerin, Experimentalisierung 2004, p. 172. “Physiological developmental genetics” was coined by Kühn who tried to study the physiological and chemical steps that were induced by a hereditary factor. Rheinberger, Ephestia 2001, pp. 542-544. Comment by Kühn in Timoféeff-Ressovsky, Verknüpfung 1935, pp. 117 f.; for Kühn on the significance of mutation research see also Kühn, Genwirkung 1934, p. 218. Harwood, Styles 1993, pp. 214-218; Harwood, Ökonomie 2002. Schwerin, Experimentalisierung 2004, pp. 56-83. Lösch, Rasse 1997, pp. 168-175; Schiemann, Baur 1935. Timoféeff-Ressovsky, Verknüpfung 1935, p. 102.

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space possible. Since small mutations were not easy to handle and required special experimental efforts, the question arose as to how these mutations could become something like a natural kind.

Erwin Baur: settling as a botanist within the space of heredity Baur was psychiatrist by education.11 He was however interested in biology. When he was studying in Kiel, a city on Germany’s eastern coast, he received private lessons from the botanist Johannes Reinke and worked at the marine zoological institute. He finally succeeded in getting an assistantship at the botanical institute of Berlin`s university in 1903. Baur said that he was deeply impressed by the rising experimental trend in biology at that time. 12 One of those experimental idols was the botanist Hugo de Vries, whose “Mutation Theory” had been widely discussed at the time. From the perspective of de Vries’ mutations, changes of an organism’s form occurred in a single large and sudden step. Baur admired the experimental courage of de Vries, but he was not quite convinced by his concept of mutations. When Baur started work he was not especially interested in mutations. However, the first problem he had to tackle at the institute in Berlin led him straight into the ongoing biological debates about the reasons for the variation of organisms. The well-known phenomenon that had puzzled the Berliner botanists for some time was the mixed-colour of leaves that were typical for some domestic plants. De Vries had presented these plants as an example for his concept of mutations calling them the “ever sporting species.”13 In the institute’s garden was a variety of the snapdragon, Antirrhinum majus, that showed variegated leaves. When Baur began experiments with one variety of Antirrhinum called Aurea, he was determined to show ordinary botanists the potential of the new experimental methods. He criticised the studies of de Vries and others following him as weak because they confused nonhereditary modifications and real hereditary variants.14 It became a mission for Baur and his assistants to show the methodological pitfalls of contemporary studies in heredity and Mendelian genetics using statistics without distinguishing the genetic status of variants. 15 Thus, the early Mendelian geneticists seemed to intervene as methodologists in the first respect. Baur was not interested in mutations in and of themselves, but his ideas on mutations were influenced by his methodological efforts and work on Aurea. Baur was convinced that the effects of mutations were not as big as de Vries suggested, but evident in the range of the effects of a Mendelian factors. This idea was in accordance with other geneticists. However, Baur’s view on mutations was special in a certain respect; he was convinced that there were even smaller mutations than common Mendelian traits like the colour of blossoms. This conviction derived 11 12 13 14

15

For a detailed biographical sketch of Baur’s early life see Schiemann, Baur 1935. Baur, Einführung 1911, p. 1. Baur, Untersuchungen 1907, p. 443. Baur, Untersuchungen 1907, pp. 443, 448 and 450. Baur showed that the new forms of Aurea were the result of the segregation of mixed characters and the influence of the environment, respectively. In the example of Baur the “ever sporting varieties” of de Vries were only special cases of modification. Baur, Untersuchungen 1907, p. 447. This critique showed Baur to be an early follower of the Danish geneticist Wilhelm Johannsen who strengthened the distinction between non-hereditary variation and heredity. Baur, Untersuchungen 1907, p. 449.

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from Baur’s critique of the confusion with modifications and hereditary factors. He suggested that the reason for the difficulties of distinguishing these types of influences was that the effects of modifications and factors largely overlapped. This assumption rested on a certain idea of how hereditary factors functioned. As a by-product, this idea informed Baur’s view of mutations. When Baur tried to deduce the cause for the variegation of the leaves of Aurea, he turned to the physiological studies of the botanist Georg Klebs. Klebs was successful in systematically showing the influence of the environment on the development of plants. 16 Baur concluded that the appearance of an organism is the result of both the influence of a Mendelian trait and a reaction to the environment. Referring to a formulation of Klebs, Baur specified that the Mendelian factor determined only the range of reactions of an organism to the environment. 17 In 1908, he explained this view when he presented physicians with the basics of Mendelian genetics: “When we see the colour of a blossom or some other outward appearance of a plant, we always see only a result of the reaction of this special individual to the outer conditions.” 18 On this basis he then continued to explain the difference between changes due to the environment (modification) and the heritage (mutations). Sometimes variations are not due to environmental influence, but rather to “what has been so often recalled mutation in the past years […]. However, the case of the newly-appeared, deviating characteristic rests on a change in the mode of reaction—even if it is only a very small one—that means commonly that the change is hereditary; we might then speak of a mutation.”19 The effects of mutations might only be a smart shift in the norm of a plant’s reaction. This view was nothing other than the translation of Johannsen’s statistic scale of continuous modification in a functional and reactive relationship of Mendelian factors and the environment. However, this environmental perspective of hereditary effects drove Baur to emphasise that mutants were not generally recognisable phenomenologically because they could be just as small as the slightest modification.20 Baur performed most of these early experiments on Antirrhinum near his home where he had leased some property.21 However, Baur’s occupation at the university’s institute was also relevant. The experience that Baur had gained working with microorganisms at the zoological institute in Kiel met the interests of the botanists at the Berlin institute working with bacteria and fungi. 22 Since the relationship of “Erbeinheiten und Außeneigenschaft”—the environmental influence 16

17

18 19 20 21

22

Baur, Untersuchungen 1907, pp. 448 f. Klebs was not a Mendelian geneticists but he also criticised de Vries. Klebs, Studien 1907, p. 99. In his view the variations of plants were not due to chance; instead he tried to show experimentally that the plants reacted to the different conditions of the environment. Klebs, Studien 1907, pp. 31 u. 102. Baur, Untersuchungen 1907, p. 449. The formulation used by Baur was close to the concept of the “norm of reaction” formulated one year later by Woltereck that was rather influential in German genetics. Harwood, Culture 1996. Baur, Ergebnisse 1908, p. 286. Emphasis by AS. Baur, Ergebnisse 1908, p. 286. Emphasis by AS. Baur, Ergebnisse 1908, p. 286. Thus, Baur turned de Vries’ theory around: large changes of a trait could simply be modifications, and, vice versa: small changes could be mutations. Baur also held lectures and university courses at that site in Friedrichshagen near Berlin. Schiemann emphasises that this large venue was of quite a bigger scale than the facilities of the university and was a definitive resource for Baur’s move from “pure botany to experimental genetics.” Schiemann, Baur 1935, pp. 65 f. and 70 f. Baur got his “Habilitation” for a work about myxobacteria in 1904. Schiemann, Baur 1935, p. 63.

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and the distinction of modifications and hereditary characters—had become Baur’s main scientific problems, he asked two doctoral students to test claims that extreme conditions or the treatment with poisons, temperature and other agents would induce mutations in bacteria and fungi—Claims which appeared to be quite popular at that time. 23 Actually, they became a junction between hereditary research and bacteriology since bacteriologists were under the proponents of that view.24

40 Mendelian traits and 642 races of the snap dragon: an experimental system on the edge of Mendelian genetics, agriculture and evolutionary theory This early research led Baur into the virulent discourse on the scope of Mendelian genetics. Baur had already attended collected variants of Antirrhinum majus when he was working on variegated leaves, but he stepped fully into that field after he had finished that work in 1908. He did not hesitate to outline what was at stake: he sought to show that the appearance of Antirrhinum was completely influenced by Mendelian factors.25 The attitude of Baur was the same he would adopt in the 1920s: to defend Mendelian genetics against those who tried to limit Mendelian validity. He remarked that experimental geneticists were usually too compliant critics when they regularly admitted that a trait was non-Mendelian; but more than ever one shouldn’t give up the claims since geneticists knew about the complicated relation between Mendelian factors and the environment.26 Obviously, the idea that heredity was disguised by the influences of the environment encouraged far-reaching claims. In his university lecture on hereditary research, Baur suggested that not only common racial traits of plants are in accordance with the Mendelian laws:27 “It looks like the slightest differences between races and species are in accordance with Mendelian laws.”28 This was the context when Baur started his the analysis of Antirrhinum majus published as “Vererbungs- und Bastardisierungsversuche mit Antirrhinum.”29 He was not the only one at that time who used Antirrhinum as a genetic model. The American botanist Ms. Muriel Wheldale had published extensive breeding experiments in 1907, and de Vries had done some work with Antirrhinum as well.30 However, Baur’s aim was to go as far as possible and to include increasingly more traits into the analysis. 23

24 25 26

27 28 29 30

Baur, Vererbungsversuche I 1910, p. 34; Schiemann, Baur 1935, p. 63. Baur was convinced that environmental conditions could induce mutations, referring to earlier experiments of the American zoologist Tower on the Colorado buck. The experiments that were performed by Elisabeth Schiemann and Franz Wolf produced no clear results. However, Baur was satisfied because the experiments seemed to contradict the claim of some bacteriologists that extreme environmental influences result in adaptive and not arbitrary mutations. Baur, Einführung 1911, pp. 203 f. For a short review of the literature see Wolf, Modifikationen 1909; Baur, Einführung 1911, p. 204 Baur, Vererbungsversuche 1910, p. 53. Baur, Vererbungsversuche 1910, p. 93. Baur made the criticism that many people would think that the Mendelian laws are only valid for bastards because they were called “Spaltungsgesetz der Bastarde.” Baur, Ergebnisse 1908, p. 288. Baur, Ergebnisse 1908, p. 288. Baur, Einführung 1911, p. 182. Baur, Vererbungsversuche 1910; Baur, Vererbungsversuche II 1912; Baur, Untersuchungen 1924. Baur, Vererbungsversuche 1910, pp. 88 f. See also Richmond, Birth 2007

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Baur took the first variants of Antirrhinum from the university’s garden. He then searched the nurseries near his house and later, systematically, the seed companies around Berlin; he also collected wild Antirrhinum variants on his hikes through the countryside.31 The variants Baur had bought or collected became “Stammpflanzen,” the units he intended to analyse. 32 He started with the most prominent racial traits such as the colour and form of the blossoms, but he widened successively his scope looking for the colour and form of leaves. 33 In 1908, Baur was confident that the 250 races of Antirrhinum majus he was able to distinguish were the product of the combination of Mendelian differences.34 In 1910, Baur specified 13 “Erbeinheiten” and estimated that there were all together about 40-50 of them.35 This number, he added, would be enough to predict the appearance of a race by looking at the genetic formula. And Baur was sure that the countless races of the species A. majus were “just combinations of these few differences that behave like Mendelian units.”36 The involved number of plants was successively increased. From 1909 on, Baur was allowed to use a part of the university’s garden where there was space for 25,000 individuals. 37 In 1911, Baur changed from the university to the agricultural university of Berlin (“Landwirtschaftliche Hochschule”) and became the head of the newly founded Institute of Heredity Research. First based at Berlin’s city centre with no extensive possibilities to grow plants, Baur managed to get a field and barracks in the nearby city of Potsdam.38 Now Baur and his workers could double their breeds and grew about 50.000 single plants each year.39 Until 1919, the number of analysed genetic units had increased up to 40 causing the characteristic differences of 642 races. Baur’s project could also be linked to the explanation of the evolution of species—a question that had also been addressed by de Vries. Baur began making this connection in 1911. 40 He was quite convinced that Mendelian heredity was a key to understanding the process of evolution, but he was also ambivalent. In 1911, Baur claimed: “The faith of selection theory depends on whether it can be shown that mutations are frequent enough in order to enable an effective process of selection or not.”41 Thus, Baur was quite aware that the analysis of mutations would be a big task for future genetic research.42 However, Baur admitted that at the basis of his environmentalist view there was a major practical obstacle. “It is possible and probable that small mutations, which have curves of modification that overlap with those of the wild type, will be overseen in most of the cases.”43 31 32 33 34 35 36 37 38 39 40 41

42 43

Baur, Untersuchungen 1924, pp. 2-4. In 1912, he cultivated 642 individuals as “Stammpflanzen.” Baur, Vererbungsversuche II 1912, p. 202. Baur, Vererbungsversuche 1910, p. 34. Baur, Ergebnisse 1908, p. 288; Baur, Vererbungsversuche 1910, p. 39. Baur, Vererbungsversuche 1910, pp. 50 f. and 91. Baur, Vererbungsversuche II 1912, p. 202. Baur, Wesen 1908, p. 333. Schiemann, Baur 1935, pp. 79 f. Baur, Untersuchungen 1924, p. 1. Baur, Vererbungsversuche 1910, pp. 34 and 53. Baur, Einführung 1911, p. 265. Early in the lecture, Baur was sceptical as to whether mutations were frequent enough to cause the “‘individual variability’ of a species.” He presumed that the witnessed variations of plants were due mostly to modifications, while variations of animals were due primarily to the new combination of factors. Baur, Einführung 1911, p. 190. Baur, Einführung 1911, p. 202; see also ibid., pp. 186 and 188. Ibid., p. 266.

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Figure 1: The chart shows painted variants of the blossoms of the snap dragon, Antirrhinum majus, which Baur analysed genetically. Source: Baur, Einführung 1911, table 1.

While evolution and mutations were more a topic for the future, the application of Mendelian genetics to agriculture was already then acute and would become even more so when World War I began. Baur foresaw the upsurge of rationalised animal and plant breeding and compared himself and his colleagues to the chemists who deliberately combined atoms and molecules. 44 The result of the genetic analysis was that “once identified genes become more available, I can work with the hereditary formulas exactly as the chemist works with his atoms, molecules and his formulas.”45 With respect to the Mendelian methodology it made no difference to speculate about hereditary changes as material for evolution or to envision them as material for the breeder. 46

Mutations coming through the backdoor Baur’s general strategy to analyse the genetic composition of the snap dragon was a mixture of inbreeding and cross-breeding. The basis of his experiments formed his “Stammpflanzen”system. “Stammpflanzen” became those plants Baur wanted to analyse. 47 The plants selected to

44 45 46 47

First he did so in 1910. Baur, Vererbungsversuche 1910, p. 90. Baur, Vererbungsversuche II 1912, p. 202. This equation should become problematic only later when Baur began to distinguish pathologic and valuable mutations. Baur, Vererbungsversuche 1910, p. 35 f.

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become a “Stammpflanze” were grown in pots for a number of years and served as source for seeds and reference for comparison in cross-breeding experiments. “Stammpflanzen” were the living standards in the ongoing experiments. The “Stammpflanzen” turned out to be homozygous in most of their genetic factors because most of the wild or commercial breeds had been inbred for some time.48 In the perspective of Baur, the “Stammpflanzen” represented the varieties of Antirrhinum that is the tableau of races. Entering the experimental phase, Baur seeded about 1,000 seeds, growing a blooming bed of the cross-generation (F1) that should reveal a Mendelian ratio. However, the genealogic chart presented by Baur does not show this progeny, but only schematically details the steps of his breeding strategy (figure 2). A single plant or few plants were selected for another experiment or to establish new “Stammpflanzen” by self pollination. The breeding strategy was to create not a genealogic tree but an ever increasing net. Step by step, the system of mutual references expanded and each new experiment would reveal a new detail of the genetic constitution of even a distant ancestor or “Stammpflanze.”

Figure 2: This genealogic chart describes schematically the “Stammpflanzen” breeding system used by Baur to analyse the genetic composition of the races of Antirrhinum majus. “A” signifies “Stammpflanzen.” Source: Baur, Mutationen 1918, p. 187.

While Baur repeated this alternate procedure of cross-breeding and inbreeding several hundred times,49 mutations came through the backdoor. In 1910, Baur noticed the first mutant plant in his sowings. Already the circumstance of this observation should become decisive for the 48 49

Ibid., p. 91. In January 1912, Baur reported 300 crossings that were analysed until the second and third generation (F2 and F3)—“a very time-consuming work.” Baur, Vererbungsversuche II 1912, p. 202.

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experimental domestication of later mutations. In 1908, Baur had crossed two plants that were special in the colour of their blossoms because he was interested in the heredity of those colours. The following summer, the progeny split up in a quite normal Mendelian ratio. Afterwards, Baur selected two plants out of the “normal” looking bed which would become “Stammpflanzen.” After self fertilisation, he seeded the progeny. However, when the second generation (F2) grew up, Baur noticed plants with an unusual form of the blossom in the progeny of one of the new hopeful “Stammpflanzen.” He was certain he had found a mutation and named the variety A. majus globosa.50 This kind of incident occured a number of times in the following years. 51 Baur’s interpretation was that a heterozygous and recessive mutation had occurred in the gametes of the parental plants. However, the mutation only became visible in a homozygous condition after he had selected and inbred one offspring carrying that mutation. There was another incident that dramatised this game of visibility. One day Baur stood in front of his experimental field in Potsdam and admired the fresh green of the growing, not yet blooming plantations ordered into chessboard-like units. He was surprised when he noticed that one section differed in the overall colour with the leaves being somewhat lighter than normal.52 The inscription revealed that the lighter plants were the inbred progeny of a cross-breeding in the third generation. Why had Baur not noticed the mutants already in the second generation like in the case of the other mutations? Baur thought he had been inattentive and, in any case, the effect of the mutation was as small as not hereditary changes that often occurred. 53 There was no chance to notice the mutation as long as there was only a small number of mutants. The mutation became only visible when one homozygous mutant was inbred and all plants of a generation were homozygous for the mutation.54 The trained view of the experimenter was betrayed by the ever modifiable plants such that the original event, the mutation, only showed up by difference (e.g. as the pattern of the experimental field). In Baur`s view this game of visibility fit well into his struggle against the pitfalls of “seeing” since he started with debunking the myth of variegated plants appearing as de Vriesian mutations. What was the true difference between mutants and modifications? There was neither a visible nor a clearly distinguishable borderline between heredity and nurture. Since the hereditary effects were often within the range of environmental influences, there was no reason to trust the eye. The recent experiences warned him that he himself had been the victim of his eyes. Nevertheless, Baur’s first experimental mutation system was based on his visual skills. Mutations arose while Baur bred thousands of snap dragons. Looking back Baur stated, “In the first years, I have not seen and investigated most of the mutants because my glance was not skilled [“geschärft”].”55 Often there was a deficit of time to look more closely since Baur went for military service during World War I. He was reluctant when there was time to distinguish just the prominent newly bred variations of Antirrhinum.56 Over the years, Baur’s visual acuity became more and more attuned, so he would see (or “suspect”) an anomalous individual earlier. 57 By 50 51 52 53 54 55 56

Baur, Einführung 1911, p. 201. Baur, Einführung 1914, p. 292. Baur, Einführung 1911, p. 189. Baur, Bedeutung 1925, p. 112. Baur, Untersuchungen 1924, p. 144. Ibid., p. 142. Baur, Mutationen 1918, p. 181; Baur, Bedeutung 1925, p. 111.

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1918, Baur had grown 200,000 individuals during his original experiments and found some 20 mutants.58 However, this was only the number Baur knew for certain. He suggested that there were many more mutations but he had not been able to analyse them because they behaved with too much variation.59 However, this vague assumption was obviously encouraged by his conviction that many hereditary units produced only slight changes in the normal range of variability. In general, Baur`s regime of mutation detection was simply based upon the ‘naked glance.’ Mutants showed up as a disturbance of the homogeneity Baur intended to maintain by inbreeding his cross-breeding products.

Mutations as objects of research: dividing chance/technique and substance/nature Mutations arising as a by-product of ongoing breeding experiments may remind one of the case of the fruit fly Drosophila melanogaster, transformed into a model organism for genetic research around the same time Baur found his first mutants. New Drosophila mutants emerged one after another among the new inhabitants of the Columbia University in New York during the experiments on the coupling of genes. Kohler has described this system as “a breeder reactor.” 60 Thus in 1919, Thomas H. Morgan and his crew had found some 100 mutants in Drosophila so far. 1919 was not a big year in the history of genetics. Nevertheless, there were three important— albeit different—publications that entailed a summary of the knowledge about mutations. All three believed in the decisive role of mutations for the evolutionary process. 61 However, Baur put ‘water into the wine’: “What we have is enough to describe the formation of new races but we have not enough variations to describe the formation of new species.” 62 Baur’s primary problem was that most of the mutations found so far were recessive in genetic terms. According to a common interpretation a recessive mutation was just the loss of the normal function of a gene. This became now a problem because from an evolutionary perspective some of the mutations might have been good for agricultural purposes but, in general, the pool of valuable mutations was limited. 63 Baur realized this in 1919 and actually became depressed. However, Baur was a go-getter by conviction and blamed the theory for finding no more mutants. His target became the presence-absence theory that was originally about the difference between recessive and dominant genes. With respect to this common idea, a recessive mutation meant a loss in the function of a hereditary unit. As early as 1911, Baur had been sceptical about 57 58 59 60 61

62 63

Baur, Bedeutung 1925, p. 111. Baur, Mutationen 1918, pp. 177 and 188. Baur, Untersuchungen 1924, pp. 100 f. Baur estimated to have found over 40 (recessive, heterozygous) mutations. Baur, Einführung 1919, p. 287. Kohler, Lords 1994, p. 47; for a recent approach see Dotan, Interrogation 2006. I refer here to Morgan’s “The Physical Basis of Heredity” (p. 269), a paper of Hermann Muller and Edgar Altenburg on “The Rate of Change of Hereditary Factors in Drosophila” (Muller, Studies 1962, pp. 217220), and the third edition of Baur’s handbook “Introduction into Heredity Research” (p. 346). Baur, Einführung 1919, p. 345. In 1911, Baur still agreed with the “presence-absence theory.” He pointed out that his experimental results were in accordance to those of his friend Hermann Nilsson-Ehle at the Svalöf agricultural station. Baur, Einführung 1911, p. 197. However, Nilsson-Ehle was more consequent in concluding that the combination of genes provided the material for evolution not mutations. Baur, Untersuchungen 1924, 146 f.

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that concept. It is not “imaginable that the complete variety of forms is due to the progressive loss of genetic units.”64 Some years later, he complained that the idea of a loss of function had slowly changed into a statement about the material essence of mutations. Most geneticists would now think that a mutation meant the material loss of the substance of a gene.65 Baur annotated in a sudden philosophical inspiration that this story should be a cautioning example of how language controls our conceptions.66 Of course, the main message of Baur was that the distinction between genetic material and the object of experiment, the genetic factor, was more important than ever. Thus following the isolation of World War I, Baur angrily reappeared on the scientific scene, directly doubting the value of the presence-absence theory.67 Its “fatal role” had been to block the idea that mutations are rather diverse.68 To direct the attention from the quantity to the diversity of mutations was a crucial move in the transformation that Baur’s experimental system would experience in the next years. The year of 1919 became the actual turning point of Baur’s efforts, for now mutations became the key object of his research. Two years later, Baur presented a plan how to check on the mutations on the first congress of the German Society for Genetics. His confidence to domesticate the tramped and invisible mutations was formed by two choices: 1. He now interpreted his former findings of mutants not so much as a product of a lucky glance, but as the product of the conditions of the experiment; in other words by artificial chance. Most mutations were overseen by researchers, “myself included,” and are only found “by chance” through ongoing experiments “representing material that was selected by their striking appearance.”69 The findings had now changed into technical shortcomings of the experimental system that meant, practically speaking, that Baur had to eliminate chance. 2. Using the new term diversity (“Mannigfaltigkeit”), Baur introduced a new distinction into the realm of the so far known mutants. By that phenomenological term he redefined the problem of mutation research. Of course, there was the question of the frequency of mutations. “A completely different question is whether the diversity of mutations is sufficient to provide material that is rich enough for the selection process.”70 This new emphasis on the quality of mutations redirected the problem of the frequency of mutations that was even more virulent because of the shortcomings of the presence-absence theory. The mutations Baur had found became his ‘white knight.’ Baur’s chief witness was the case of the mutant that had shown up only in the third generation as the pattern of the bet. Nobody had thought that these mutations could be different because they had not been noticed until then.71 This idea became more plausible in the light of the “norm of reaction” that implicated that there were mutations that overlapped with the slightest non-hereditary modifications of the individual.

64 65 66 67 68 69 70 71

Baur, Einführung 1911, p. 198. Baur, Einführung 1914, p. 149. Ibid., p. 150. Baur, Mutationen 1918, p. 178. Baur, Einführung 1919, p. 344. Baur, Anzahl 1921, p. 241. Emphasis by AS. Baur, Untersuchungen 1924, p. 142. Baur, Einführung 1919, p. 343. Ibid., p. 344.

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It is possible, no, it is probable that small mutations, whose curves of modification overlap with those of the wild race, are usually overlooked.72

Baur had already considered that possibility in 1911. In 1921, it became an apodictic proposition that guided the upcoming practical transmutation of the experimental system. Eliminating chance meant to enter into a competition to find more of the “little mutants” and above all the “‘barely noticeable’ [“eben noch gerade”], slightest mutants.”73 It is not possible to notice most mutations as such, neither most of the mutations in the controlled breeding cultures of our best studied experimental animals and plants! In order to find all mutations that have been arisen one has to introduce special experimental condition never realised until now.74

The mutations had been lucky moments in Baur’s experimental system that could now be described as statistical events. Now they became a matter of technical chance. The production depended on the correct arrangement of the experiment. The path Baur would choose was already predetermined by his conceptual idiosyncrasy that originated in experiments 15 years prior; namely his obsession with the evolutionary process and the recent experimental experience made on the soil of Potsdam.75 Thus, the key target of Baur’s experimental calculus became the recessive mutations that usually first appeared in a heterozygous form and in almost unchanged plants. Also, these mutations began to naturalize—but not in the sense that Baur now introduced a material correlate. Instead he introduced a classification into the former continuous realm of small to big mutations. The big divide became the distinction of normal mutations and pathological mutations suggesting that these both classes correlated with the scale of effects. 76 Actually, only around that time Baur associated eugenic sentiments with the epistemic problem of mutations. The impact of eugenic normalisation legitimized Baur to introduce a dual classification equating big mutations with pathologic mutants and small mutations with the useful diversity representing the material of evolution.77 Baur’s message when speaking to the geneticist society was that “people in general would think today that most mutants are deformities. Therefore they are not candidates for evolution […] However, the many small mutations, which are quite good for evolution are normally overlooked.”78 It was in this social-technical constellation that Baur developed a naturalising speech about small and large mutations, suggesting that they represented a sort of natural type of their own right, although they still belonged to the same recessive type of Mendelian inheritance.

72 73 74 75 76 77

78

Baur, Einführung 1911, p. 266. Baur, Bedeutung 1925, p. 114. Baur, Einführung 1922, p. 32. Not mentioned here are further reasons why just recessive and heterozygous mutations became central in Baur’s experiments, see footnote 99. Canguilhem has described this pattern of modern normalisation when he showed how distinctions were reintroduced in Broussais’ quantitative order of continuous scales. Canguilhem, Normal 1991, p. 56. The role of mutations for the eugenic mobilisation of geneticists has not until now been well investigated. It seems that the mobilisation came together with the rise of the mutational dispositive in the 1920s. The examples of Muller and Baur at the very least confirm this guess. Baur, Anzahl 1921.

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Baur’s inbreeding system: a new breeding system for the domestication of mutations Beginning in 1922, in order to overcome all the hindrances that made the detection of small mutations a matter of chance, Baur introduced a new breeding system. It was however not completely new, but a transformation of his “Stammpflanzen” system.

Figure 3: This genealogic chart depicts Baur’s strategy to detect recessive, inconspicuous mutations. It was based purely on inbreeding and showed hierarchically the accumulated progeny of one individual (A.7526). Source: Baur, Untersuchungen II 1926, p. 253.

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The genealogic chart in figure 3 shows the descent of the plant A.7526. This kind of pedigree differed at some decisive points from the schema that represented Baur’s “Stammpflanzen” breeding system. First of all, the experiment did not start with the crossing of two individuals, but rather with the self-fertilisation of plant A.7526. Baur now profited from the resources he had established by then with over 4.000 “Stammpflanthat were supposed to be pure lines. 79 But the reference system of the “Stammpflanzen” was now outstripped. The experiment system focussed on the very progeny of one plant. Although now the experiment was limited to one ancestral lineage, it was still open-ended. Each breeding extended the statistic basis of the calculation. And the numbers grew rapidly because Baur did not just select one or two individuals from the offspring (F1). He now claimed that this limitation had been the critical chance moment as it was unlikely that he caught the single plant carrying a recessive mutation out of 1.000 offspring.80 Baur now chose several individuals in order to inbreed them.81 He used his experienced glance to select just those individuals who seemed to be promising candidates for mutants.82 Afterwards, he inspected the second generation to see whether a mutation arose. In the example (see figure 3) Baur selected seven plants (A.7744 to A.7750). The progeny of these plants is shown in lines b to h. However, at that point the experiment was still not complete. Baur repeated the same procedure now with 53 plants of the second generation (A.8194 to A.8255). The principle of his inbreeding scheme now becomes clear: the chart shows only the plants of one generation that were selected for further inbreeding. The family tree expanded rapidly. The small numbers in brackets showed the individuals grown in the third generation (F3) which grew to 3,366 plants. 83 Each plant functioned not only as a test of the mutational status of the grand-mother generation, but also as the starting point for the investigation in the next generations. The calculus of Baur was that he could accumulate the number of inbred plants and of mutants deliberately over generations and calculate the rate of mutation. The transformation of the old system narrowed the scope of the experiment in terms of a clear-cut hierarchical descent schema forming a eugenicallybased model inbreeding family tree. However, at the same time the experiment was widened with respect to its statistical basis. Baur was rather content. Mutants proliferated and in 1924, Baur published a comprehensive overview of his experiments with Antirrhinum and all mutations found. He reported that he had found 5% mutation rate in one inbred family.84 This was a true success since, in 1918, he had calculated a rate of recessive mutations of only 0.2% and the scale now seemed to be open. The experiment illustrated in figure 3 was introduced by Baur one year later. He reported five mutants and calculated a rate of mutation of 6-7%.85 When Baur presented these results to the surprised 79 80 81

82 83 84

See the examples in Baur, Mutationen 1918. Baur, Untersuchungen 1924, p. 143. Baur, Untersuchungen II 1926, p. 253. Baur announced that one should select at least 30 or 100 siblings. Baur, Einführung 1930, p. 316; Baur, Einführung 1922, p. 372. However, the number was not essential because the experiment did not depend on the number selected in one generation but the accumulated number across all generations. Baur, Mutationen 1918, p. 184. Altogether, Baur checked 4.000 individuals including the third generation. Baur, Untersuchungen II 1926, pp. 255 f. Baur, Untersuchungen 1924, p. 144.

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German geneticists in 1925, he pointed out that he had expected a mutation rate of 10%—an immense proportion.86 There was an immediate resonance amongst the German geneticists who were already speaking of the revival of the selection theory. Asked by a newspaper, Max Hartmann pointed out that the experimental “Vererbungsforschung” was on its way to forming the basis for the return of the “doctrine of the natural selection”: “Small mutations that are very frequent can have an especially significant value for selection—supported by environment and bastardisation.”87 Referring to “small mutations,” Hartmann recalled Baur’s results.

The conditions of a new type of mutation: the constellation of experiment, evolution and eugenics The transformation of the frequency problem of mutations into the frequent variety of mutations paid off. Baur had successfully merged the methodological pitfalls of genetic research, the needs of the evolutionary process and the evaluation of mutations. The breeders—and Baur included himself—had not realised that this realm of small mutants existed because they were both only able to use their eyes and had been only interested in striking mutants. 88 Although Baur had learned to consider this situation as an artefact of the normal breeding dispositiv, now—when the mutations proliferated—there was no hindrance to re-naturalise the whole setting of the experiment. This was now an easy step after Baur had started to introduce the evaluation of the mutants in evolutionary and eugenic terms. Thus, by linking the agricultural dispositive, evolutionary mechanisms and eugenic normalization, it became obvious that the striking mutants were an artefact of the selection of the breeder as they normally would have been extinguished by natural selection.89 Nature and evolution left no doubt about the abnormal character of the commonly found mutants and their artificial existence. Most would actually admit that most of the “conspicuous” mutants are “almost without exception deformations” or “distinct pelories.” 90 By contrast, the small mutations were in reality not only more frequent, but also they “do not belong to the field of the pathological, but are absolutely viable types.”91 The small mutations […] are of very different kind, they determine small differences in the colour of leaves, the colour of blossoms, the relative length of anthera, the manner of hair patterns, the size of seeds, etc. In short they form an enormous variety [“Mannigfaltigkeit”]!92

Thus, small mutations affect 85 86 87 88 89 90 91 92

There were 70 plants (in the chart: A.7526 to A.8255) whose progeny had been screened for mutations. Baur, Untersuchungen II 1926, pp. 255 f. Baur, Bedeutung 1925, p. 112. Max Hartmann: Die Lehre von der natürlichen Zuchtwahl überholt?, in: Berliner Tageblatt, Nr. 496, 1. Beiblatt, 1926. Baur, Einführung 1930, p. 398. Baur, Untersuchungen 1924, p. 147. Baur, Anzahl 1921, p. 241; Baur, Bedeutung 1925, p. 111. Baur, 1925 Bedeutung, p. 111. Baur, Bedeutung 1925, p. 113.

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all possible morphological and physiological attributes of an organism. They do not determine changes that are monstrosities or pathologies but changes that are within the norm, that do not decrease viability, but can increase it.93

The effects of small mutations were small in scale but potent. As such, they were similar to those traits that constituted the species. Small mutations are normally within the frame of physiology. In the case that factor mutations will in general likely provide the material for natural selection, and thus for evolution, small mutations will be the candidates.94

Baur’s mutual mobilisation of techniques, concepts and beliefs finally resulted in the deification of both small mutations and large ones. Against the common thinking, Baur pointed out that striking mutations were not usual but rather the “extreme cases.” 95 Instead, the small and “smallest mutations” (Baur) turned out to be the normal, that is, natural cases. To make this difference clear, Baur introduced the proper name “Kleinmutationen.” 96 Most of all mutations are—as far as I can see—mutations of this mode.97

Recalling the statements of Kühn and Timoféeff-Ressovsky in the beginning of the 1930s, small mutations and large ones became types with an epistemic status somewhere between their use as tool and true natural kinds. The new system that bore Baur’s wave of mutations was full of traces that reflected the material work of Baur as a biologist and his convictions as an evolutionist. He always believed in selection theory and was deeply impressed by de Vries’ project of an experimental approach towards the “synthesis of species.” Beginning in 1907, the concept of the norm of reaction gave rise to the idea of “small changes.” Around 1910, Baur mentioned small hereditary changes as a subject of evolutionary change and the material for agricultural domestication. Additionally, Baur’s first breeding system pre-formed the development of his mutation research. Practically speaking, the “Stammpflanzen” system became the starting point of the inbreeding system. The pure lines stored so far provided the material for the experiments started since 1922. Additionally, the choice of his experimental object mattered because the mutations detection system was only realisable with an autogamous plant such as Antirrhinum and not with model organisms like Drosophila.98 Another condition and line of research should be mentioned here. The experimental system for the detection of mutations entailed a crucial decision about the moment when mutations occur in the development of an organism. Baur performed extended experiments on that problem that finally emphasised the relevance of recessive mutations and of artificial chance. 99 It is noteworthy that this constellation was strong enough to rule out conceptual claims such as the “presence-absence theory,” the combinatory concept of selection theory and, last but not 93 94 95 96 97 98

Baur, Untersuchungen 1924, p. 143. Baur, Einführung 1930, p. 398. Baur, Bedeutung 1925, p. 111. Baur, Untersuchungen 1924, p. 146. Baur, Bedeutung 1925, p. 112. Emphasis by AS. Baur, Bedeutung 1925, pp. 113 and 115; Baur, Einführung 1930, p. 316.

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least, the concept of pure lines. In 1924, Baur explained that the path breaking concept of pure lines had been revealed as a hindrance for the new view of mutations because it favoured perceiving modifications where actually small mutations were at work. The comment of Baur is worth citing at length: We have—impressed by the basic studies of Johannsen on the constancy of pure lines—overestimated constancy. I do not want to criticize Johansson, neither his method nor his conclusions. […] But also in this case the successors of Johannsen were rather orthodox [“päpstlicher als der Papst”]. Johannsen himself did not speculate on the occurrence of mutations. But in general, the idea became widely appreciated that mutations and factor mutations were something uncommon and rare and that there are only monstrosities coming out of them. In effect, we have a completely false estimation of the frequency and the variability of factor mutations, probably because of a total “ignorabimus” in the question of evolution or because of trials to explain evolution by natural selection of combinations (see Lotsy and Heribert Nilsson).100

Outlook: Baur’s story in the context of biomedical research from the 1920s on The history presented shows how a specialised experimental system for the domestication of mutations evolved out of the practical and situated heritage of the botanical and agricultural research of Erwin Baur. Considering the conceptual and practical constellation, it becomes clear why no other geneticists chose the path that Baur did. Baur was not the only one who was puzzled about small mutants, which appear to be part of the longer plan of Neo-Darwinian geneticists. For example, Thomas Hunt Morgan made up his mind about small mutations, but those mutations fell behind because he was too busy counting the larger ones.101 Also, Hermann Muller prominently mentioned Baur’s “elusive class of ‘invisible’ mutations” in his article on the artificial inducement of mutations in 1927.102 Muller himself reported cases of mutants that decreased viability or produced other “invisible” and “inconspicuous” effects. Baur’s experiments on Antirrhinum in particular had proven the existence of mutants “that approached or overlapped the normal type to such an extent that ordinarily they would have escaped observation.” 103 However, those mutations that were especially important for the question of natural selection “were not subjected to study” as Muller put it.104 In other words, the trajectory of Muller’s

99

Baur recognised this as a key question of mutation research since roughly 1918. Baur was finally convinced that most mutations happened just at the moment when a sexual cell formed in the plant. Thus, normally only one egg was mutated and, consequently, only one plant of the offspring carried a mutation. Baur, Untersuchung II 1926, p. 255. This model of the generation of mutations supported Baur’s view of chance because he was unlikely to select just the single plant carrying a heterozygous mutation. 100 Baur, Untersuchungen 1924. 101 Kohler, Lords 1994, pp. 39 ff. For Morgan’s mutation experiments in particular see Dotan, Interrogation 2006. 102 Muller, Studies 1962 [Artificial Transmutation of the Gene, in: Science 1927], p. 246. 103 Ibid., p. 247. 104 Ibid., p. 246.

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experimental work established another specialized experimental system for detecting mutations. His system had its own special protagonists: lethal mutations.105 In general, Baur shared a growing interest in mutations among geneticists. His inbreeding system was just one approach that evolved in order to proliferate mutations. Other approaches would be part of a more comprehensive history. This history would show that different interests were connected to variant trials to domesticate mutations. It would embrace both the artificial induction of mutations and the detection of “natural” ones. Not all of these activities were tied to evolution or used the same model organism, but there was a field of activities grouped around mutants forming. Thus, the German geneticists Kühn and Timoféeff-Ressovsky considered “Kleinmutationen” as a research tool and envisioned special breeding stations for experimental animals that should serve biomedical research by screening animals for hidden mutants. 106 Mutants became a type of “leading technology” in a variety of fields including medicine, agriculture, eugenics and biomedical research. The domestication of mutations became an aim that served different interests and resulted in a growing experimental culture of mutation that embraced different experimental objects, manipulative agents, breeding techniques and interests. Alexander von Schwerin TU Braunschweig [email protected]

105 106

Muller, Studies 1962 [The Problem of Genic Modification, lecture in Berlin 1928], p. 253. Schwerin, Experimentalisierung 2004, pp. 175 f.

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References Baur, Erwin. 1907. “Untersuchungen über die Erblichkeitsverhältnisse einer nur in Bastardform lebensfähigen Sippe von Antirrhinum majus.” Berichte der deutschen botanischen Gesellschaft 25: 442454. ————. 1908. “Einige Ergebnisse der experimentellen Vererbungslehre.” Medizinische Klinik 4: 265-292. ————. 1908. “Das Wesen und die Erblichkeitsverhältnisse der “Varietates albomarginatae hort.” von Pelargonium zonale.” Zeitschrift für induktive Abstammungs- und Vererbungslehre 1: 330-351. ————. 1910. “Vererbungs- und Bastardisierungsversuche mit Antirrhinum.” Zeitschrift für induktive Abstammungs- und Vererbungslehre 3: 34-98. ————. 1911. Einführung in die experimentelle Vererbungslehre. Berlin. ————. 1912. “Vererbungs- und Bastardisierungsversuche mit Antirrhinum. II. Faktorenkoppelung.” Zeitschrift für induktive Abstammungs- und Vererbungslehre 6: 201-216. ————. 1914. Einführung in die experimentelle Vererbungslehre, 2. neubearbeitete Aufl., Berlin. ————. 1918. “Mutationen von Antirrhinum majus.” Zeitschrift für induktive Abstammungs- und Vererbungslehre 19: 177-198. ————. 1919. Einführung in die experimentelle Vererbungslehre, 3. u. 4. neubearbeitete Aufl., Berlin. ————. 1922. Einführung in die experimentelle Vererbungslehre, 5. u. 6. neubearbeitete Aufl., Berlin. ————. 1921. “Herr E. Baur-Dahmsdorf führt eine Anzahl Mutanten von Antirrhinum vor.” Zeitschrift für induktive Abstammungs- und Vererbungslehre 27: 241. ————. 1924. Untersuchungen über das Wesen, die Entstehung und die Vererbung von Rassenunterschieden bei Antirrhinum Majus. Leipzig. ————. 1925. “Die Bedeutung der Mutation für das Evolutionsproblem.” Zeitschrift für induktive Abstammungs- und Vererbungslehre 37: 107-115. ————. 1926a. “Untersuchungen über Faktormutationen I. Antirrhinum majus mut. phantastica.” Zeitschrift für induktive Abstammungs- und Vererbungslehre 41: 47-53. ————. 1926b. “Untersuchungen über Faktormutationen II. Die Häufigkeit von Faktorenmutationen in verschiedenen Sippen von Antirrhinum majus.” Zeitschrift für induktive Abstammungs- und Vererbungslehre 41: 251-258. ————. 1930. Einführung in die Vererbungslehre, 7.-11. völlig neubearbeitete Aufl., Berlin. Canguilhem, Georges. 1991. The Normal and the Pathological. New York . Dotan, Igal. 2006. Interrogation of a Fly. Paper presented at the MPI colloquium, Berlin. Harwood, Jonathan. 1993. Styles of Scientific Thought. The German Genetics Community 1900-1933. Chicago. ————. 1996. “Weimar Culture and Biological Theory. A Study of Richard Woltereck (1877-1944).” History of Science 34: 347-377. ————. 2002. “Politische Ökonomie der Pflanzenzucht in Deutschland, ca. 1870-1933.” In S. Heim (ed.), Autarkie und Ostexpansion. Pflanzenzucht und Agrarforschung im Nationalsozialismus. Göttingen. 1433. Klebs, Georg. 1907. “Studien über Variation.” Archiv für Entwicklungsmechanik der Organismen 24: 29-113. Kohler, Robert. 1994. Lords of the Fly. Drosophila Genetics and the Experimental Life. Chicago. Kühn, Alfred. 1934. “Genwirkung und Artveränderung.” Der Biologe, 3: 219-227. Lösch, Niels. 1997. Rasse als Konstrukt, Leben und Werk Eugen Fischers. Frankfurt. Morgan, Thomas Hunt. 1919. The Physical Basis of Heredity. Philadelphia. Muller, H. J. and Joshua Lederberg (eds.). 1962. Studies in Genetics. The Selected Papers of H. J. Muller. Bloomington. Rader, Karen A. 2004. Making Mice. Standardizing Animals for American Biomedical Research, 1900-1955. Princeton. Rheinberger, Hans-Jörg. 2001. “Ephestia. The Experimental Design of Alfred Kühn’s Physiological Developmental Genetics.” Journal of the History of Biology 33: 535-576. Richmond, Marsha L. 2007. “Muriel Wheldale Onslow and Early Biochemical Genetics.” Journal of the History of Biology 40: 389-426. Schiemann, Elisabeth. 1935. “Erwin Baur.” Berichte der Deutschen Botanischen Gesellschaft 52: 51-114. Schwerin, Alexander von. 2004. Experimentalisierung des Menschen. Der Genetiker Hans Nachtsheim und die vergleichende Erbpathologie, 1920-1945. Göttingen. Timoféeff-Ressovsky, N. W. 1935. “Verknüpfung von Gen und Außenmerkmal (Phänomenologie der Genmanifestierung).” In W. Kolle (ed.), Wissenschaftliche Woche zu Frankfurt a.M., 2.-9. September

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1934. Band 1: Erbbiologie. Leipzig. 92-115. Wolf, Franz. 1909. “Über Modifikationen und experimentell ausgelöste Mutationen von Bacillus prodigiosus und anderen Schizophyten.” Zeitschrift für induktive Abstammungs- und Vererbungslehre 2: 90-132.

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Heredity and the Century of the Gene Raphael Falk

Concluding remarks This has been the 4th of the series of workshops on A Cultural History of Heredity, and it was entitled “heredity and the century of the gene.” Let me start by challenging the headline of the workshop and claim that it gives a twisted view of the cultural history of heredity. As has been pointed out repeatedly by different speakers during the last days, to the extent that the gene took a central position in the history of genetics, this changed dramatically in the 1960s. It is true that, as noted by Hans-Jörg Rheinberger, “a gene is a gene, is a gene.” This gene, however, became increasingly a bewildering concept and it has served more and more just as a generic term, like ‘table’ or ‘chair’, rather than as a specific structural entity. In spite of an attempt like that of Lenny Moss to rescue the gene concept by introducing two distinctive modes, of a gene-P and a gene-D, and Evelyn Fox Keller’s suggestion of a shift from a “discourse of gene action” to that of “gene activation” —although, as we learned from her talk last night, she has been accused to lead a jihad against the gene—I suggest that the shift was elsewhere in the cultural level of the history of heredity, namely in the role of reductionist conceptions in genetic research. In two workshops on “Representing genes,” organized a couple of years ago by Karola Stotz and Paul Griffiths in Pittsburgh, we ended up with more than a dozen different phenomena in our attempts to define “a gene.” In the recent book of Bob Weinberg, The Biology of Cancer, the term “gene” as such does not appear at all in the glossary; there are only items like “gene amplification,” “gene family,” “gene pool” etc. It is not in vain that the previous workshops of this series on the Cultural History of Heredity started with the 17th and 18th centuries. Indeed, if we insist on parsing the centuries of the cultural history of research on heredity, I suggest that we start with Linnaeus. The century from 1750 to1860 being the Century of Rationality (for the lack of a better name), with Darwin at its peak; the century from 1860 to1960—starting with Mendel and ending with Crick—as the Century of Reductionism; and starting in the 1960s, the so-far half Century of Integration, of the genome, the proteome, and the return of evo-devo. The publication of Linnaeus’ Systema naturae in 1735 and the fixation of the systems of nature on the one hand, and the publication of the first three volumes of Buffon’s Histoire naturelle in 1749, on the flexibility of nature, introduced the century of claims for rational research of biological synthesis of change. It was characterized by two major research modes: The one, leading from Linnaeus through Koelreuter and Gärtner to Mendel, put the emphasis on hybridization as a mode of research. The other, leading from Buffon, through Lamarck and Geoffroy Saint Hilaire to Darwin, puts the emphasis on morphogenesis, comparative anatomy and embryology as modes of research. Whereas Lamarck’s publication of Philosophie zoologique in 1802 may be noticed as the significant mid-century event, Darwin’s Origin of 1859 and Mendel’s Versuche of 1864 introduce a new century.

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Figure 1. Which one is the gene ? Forms proposed by Paul Griffiths and Karola Stotz at a conference in Pittsburgh, 2003.

The second century, which started with the 1860s soon confronted the dialectical conflicts of methodological reduction and conceptual reduction of biological diversity. The Century of Reductionism of hereditary research reached its peak with Crick’s paper “On protein synthesis” at the Society of Experimental Biology in which he formulated the Central Dogma of Inheritance, and with the statement, attributed to Jacques Monod: “Anything found to be true of E. coli must also be true of elephants.” Morgan’s “Chromosomes and heredity” may be considered as the significant mid-century event. The controversies initiated with the publication of Richard Dawkins’ The Selfish Gene and Edward O. Wilson’s Sociobiology were the more spectacular indicators of the demise of the Century of Reductionism. Britten and Kohne’s discovery that what is true of prokaryotes is not true of eukaryotes was one of the earliest experimental findings that heralded the end of the Century of Reductionism. Roberts and Sharp’s discovery that genes in eukaryotes are not contiguous strings but contain introns, and that the splicing of messenger RNA to delete introns can occur in different ways, yielding different proteins from the same DNA sequence, opened a new era. Gould and Lewontin’s paper on “The spandrels of San Marco and the Panglossian paradigm,” challenging the New Synthesis’s reduction of evolution to changes in gene frequencies, may be considered as one of the first signals on the conceptual level of the emergence of the third century, the Century of Integration.

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Figure 2. Heredity over the centuries. Source: Raphael Falk.

It was no accident that of the eighteen odd talks in the workshop’s program, all but one dealt with the period preceding 1960, that is, with the century of reductionism. Only one, the programmatic talk of Jean Gayon “Widening heredity: From soft to hard inheritance and back” promised to deal with abridging the centuries of reductionism and integration (unfortunately Jean could not present his talk); also Christina Brandt attempted to reach out to integration with the modern notion of clones. Notwithstanding Jon Hodges’ comment on the origin of the term “soft inheritance,” as Gayon pointed out in his abstract, “The emergence of the concept of ‘hard inheritance’ was crucial to the constitution of an experimental science of heredity” in the 1910s: These were the years of bitter but constructive dialectic confrontation between “experimental” and “conceptual” reductionism in hereditary research. It is too bad that Jean could not elaborate more on this subject. As Jean Gayon wrote in his abstract, the late “relaxing of conceptual and empirical constraints imposed by ‘hard’ inheritance,” was another consequence of the change of the culture of hereditary research, from that of reductionism to integrationism. For Johannsen pure lines were significant as instruments to discern phenotypic from genotypic phenomena. As Judy Schloegel has shown, for Jennings the pure line, and even more so the clone, provided empirical means for the establishment of the conceptual reduction of the phenotype to an ultimate genotype. Victor Jollos’ Dauermodifikationen of “soft” or epigenetic inheritance posed, however, a threat to deterministic reductionist genetics, which could not be tolerated in the

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century of reductionism, and was denoted as neo-Lamarckism. I wonder to what extent Jollos’s difficulties in emigrating from Germany and his short, unhappy period in the US were not another aspect of Veronika Lipphardt’s claim that many Jewish-German scientists, who had been immanently neo-Lamarckians because of the aggressive Darwinian claims of German racial scientists, had difficulties when forced to immigrate to the Anglo-American world in which neoDarwinism prevailed. It needed conceptual integration (without abandoning reductionism as a heuristic) to reintroduce the top-down perspective as a respected aspect of genetic research, and to investigate group selection, developmental constraints, or the role of epigenetic mechanisms. When I congratulated Marion Lamb on her and Eva Jablonka’s first book Epigenetic Inheritance and Evolution, suggesting how fortunate they were to publish the right book at the right moment, Marion corrected me, pointing out that it was the other way round: It was their book that helped create the right moment. By the way, the inventor of the term “gene,” Johannsen himself, never liked the term. He consistently rejected the reductionist “unit character,” and to the end of his career he continued to talk of genotypical (and phenotypical) variation, and remained reserved towards the meaning of the concept of the gene. As suggested by Staffan Müller-Wille, a scientific basis meant for him a thoroughly instrumental basis. Conceptually Johannsen was an organicist, though instrumentally he was a reductionist, or as Staffan put it in his talk: “The assumption that the organism is an ensemble of individually reproducing parts drove Johannsen mad.” I am sorry that I cannot dwell on many of the interesting papers of the Workshop in these brief comments. Greg Radick redirected our attention from the Bateson-Pearson polemic to its more profound foundations, which is actually the Bateson-Weldon polemic. Whereas Bateson—as well as de Vries—abandoned the traditional morphogenic approach to the study of variation of species, to join the competing hybridists’ tradition, Weldon adhered to the old tradition of the morphogenicists. Radick highlighted Weldon’s criticism of the Mendelians’ inbuilt methodological bias of discontinuous classification into a finite number of categories. This may indeed be a serious problem, as shown years later by Raymond Pearl who asked fifteen trained geneticists to classify 532 F2 corn kernels from a cross of yellow starchy and white sweet varieties. However, one has to keep in mind that Mendel was very much aware of the possible biases of his empirical methodology and devoted two years selecting the proper strains and traits before testing his hypothesis. Indeed, large scale repeats of Mendel’s experiments over the years proved upholding his classification. It may also be kept in mind that already in1902 Udny Yule showed that much of the Pearson/ Weldon—Bateson polemics concerned the formulation of the problems: Mathematically the Law of Ancestral Heredity could be reduced to that of Mendelian inheritance. Mendelism, Yule pointed out, was focusing on hybridization—the study of specific difference-characteristics between individuals; The Law of Ancestral Heredity, on the other hand, was concerned with heredity. Heredity represents the population-aspect of inheritance; it regards the correlation of variance in one generation of the population with that of another. Accordingly, Yule opened his

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1903 paper: “The statistical theory of heredity, as developed in the work of Galton and Pearson, concerns itself with aggregates or groups of the population and not with single individuals.”

Figure 3. In 1911 Raymond Pearl crossed yellow-starchy with white-sweet homozygous varieties of maize. Fifteen trained scientists (abscissa) counted the same set of 532 F2 kernels (ordinate). The horizontal dotted line gives the Mendelian expectation, and the horizontal dashed line the average of the counts of the fifteen observers. Source: R. S. Root-Bernstein (1983). "Mendel and methodology." History of Science 21: 275-295.

Figure 4. Summary of F2 results of seed color in pea crosses, counts of seven studies. Source: E. W. Sinnott & L. C. Dunn (1932). Principles of Genetics. (2nd ed.) New York: McGraw-Hill. p. 48.

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Finally, allow me to relate briefly to the two papers to the “left” and to the “right” of Muller’s research program of the nature of the gene: Luis Campos’s paper on Blakeslee’s “chromosomal mutations” in Datura, and Alexander Schwerin’s discussion of Erwin Baur’s “Kleinmutationen” in Antirrhinum. Muller was, of course, aware of both chromosomal aberrations and minute mutations and on their role in evolution. However, in his programmatic paper of 1922, Muller emphasized that “it is not inheritance and variation which bring about evolution, but the inheritance of variation, and this in turn is due to the general principle of gene construction which causes persistence of autocatalysis despite the alteration in the structure of the gene itself.” His research program was not to review the kinds of mutations and their relative contribution to population structure and evolution, but to disclose the nature of the gene. Muller was explicitly after the property that makes genes unique, namely mutations, which maintain autocatalysis in spite of change of function, and to do this he needed a quantitative analysis of mutagenesis. Blakeslee appropriately opposed the Drosophilists’ relentless reductionism, treating organisms “like a child’s house of blocks,” but it is necessary to discern the difference between Muller’s conceptual reductionism and his reductionist heuristics. The collective concept of “mutation” employed by de Vries and his followers only confused matters. Many devices and materials were tried out and proved to be mutagenic in those years. However, it was the establishment of the ClBmethod that provided the heuristics for the quantitative determination of the efficiency of a mutation inducing agent, which made Muller’s 1927 contribution to genetics so unique and justifiably entitled him to the priority he was seeking. And I think it still is a central contribution to the century of hereditary reductionism, in which the gene concept played a central role. Let me finish my comments by thanking—I hope in the name of all of us—the organizers of this Workshop both at the Centre for Genomics in Exeter, and at the Max-Planck Institute in Berlin, for a pleasant and fruitful conference. Thank you! Raphael Falk The Hebrew University of Jerusalem [email protected]

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Workshop Heredity in the Century of the Gene (A Cultural History of Heredity IV) December 11-14, 2006 ESRC Research Centre for Genomics in Society, University of Exeter, UK in collaboration with the Max-Planck-Institute for the History of Science, Berlin, Germany Organizers: Staffan Müller-Wille (Exeter), Hans-Jörg Rheinberger (Berlin), John Dupré (Exeter) Venue: Reed Hall

Monday, Dec 11 14:00

Registration

14:30

Welcome John Dupré, Staffan Müller-Wille, Hans-Jörg Rheinberger

15:30

Coffee break

16:00

Session I: The Mendelian Break Staffan Müller-Wille, University of Exeter Leaving inheritance behind: Wilhelm Johannsen and the politics of Mendelism Greg Radick, University of Leeds The Professor and the Pea: Weldon’s Critique of Mendelism Barry Barnes, University of Exeter comments

17:50

Drinks and standing buffet

Tuesday, Dec 12 09:00

Session II: Contexts of Heredity Ilana Löwy and Jean Paul Gaudillière, Centre de recherche médicine, sciences, santé et societé, Villjuif Transmission of human pathologies 1900-1940: the elusive “mendelization” of the clinic Bert Theunissen, University of Utrecht Breeding Dutch dairy cows (1900-1950): Heredity without Mendelism Soraya de Chadarevian, University of California, Los Angeles comments

10:50

Coffee break

11:20

Daniel Kevles, Yale University, New Haven, Ct. Innovation and Ownership in New Fruits: The Horticultural Industry and Intellectual Property in the United States, 1880-1930

Maria Kronfeldner, Max-Planck-Institute for the History of Science, Berlin Coalition and Opposition: Heredity, Culture, and the Boundaries of Anthropology in the Work of Alfred L. Kroeber Edna Suarez, Max-Planck-Institute for the History of Science, Berlin comments 13:10

Lunch

14:30

Social event and dinner

Wednesday, Dec 13 09:00

Session III: Breeding and Inheritance Christophe Bonneuil, Centre Alexandre Koyré, Paris What genes could not do: French plant breeders’ reception of Mendelism (1900-1930) Ana Barahona, Universidad Nacional Autonoma México, Mexico City Mendelism and agriculture in the first decades of the 20th century in Mexico Jonathan Harwood, University of Manchester comments

10:50

Coffee Break

11:20

Session V: Heredity and the Creation of Model Organisms Judy Jones Schloegel, independent scholar, Clarendon Hills/Ill. Herbert Spencer Jennings, Heredity, and Protozoa as Model Organisms, 1908-1918 Christina Brandt, Max-Planck-Institute for the History of Science, Berlin Clones, pure lines and heredity. The work of Victor Jollos Hans-Jörg Rheinberger, Max-Planck-Institute for the History of Science, Berlin comments

13:10

Lunch break

14:30

Session IV: Genealogy and its Uses Bernd Gausemeier, Max-Planck-Institute for the History of Science, Berlin Human Heredity and Mendelism: the Case of Psychiatry Philip Wilson, Penn State University College of Medicine, Hershey (Pennsylvania) Pedigree charts as tools to visualize inherited disease in progressive era America

15:50

Coffee break

16:20

Veronika Lipphardt, Humboldt University, Berlin Jews as an object of Mendelian research (1900-1935) Carlos López Beltrán, Universidad Nacional Autonoma México, Mexico City comments

17:30

Break

19:30

Public evening lecture (Venue: Queens Lecture Theatre 1) Evelyn Fox Keller, Massachusetts Institute of Technology, Cambridge, MA What's in a Word? Genes, Heredity, and Heritability

Thursday, Dec 14: 09:00

Session VI: Managing Variation Marsha Richmond, Wayne State University, Detroit William Bateson’s Pre- and Post-Mendelian Research Program in `Heredity and Development’ Luis Campos, Harvard University, Cambridge/Mass. Genetics Without Genes: Blakeslee, Datura, and ‘Chromosomal Mutations’

10:20

Coffee break

10:50

Alexander von Schwerin, Technical University Braunschweig Seeing, breeding and the organisation of variation – model organisms in the genetics of the twenties Jonathan Hodge, University of Leeds comments

12:00

Lunch break

12:45

Lenny Moss, University of Exeter comments Raphael Falk, Hebrew University, Jerusalem comments Final discussion

14:15

End of workshop