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Heredity, development and evolution: the unmodern synthesis of E.S. Russell

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Abstract

In 1930, while R.A. Fisher, J.B.S. Haldane, E.B. Ford and S.G. Wright were laying the foundations of what a decade later J.S. Huxley dubbed “Modern Synthesis”, E.S. Russell published a groundbreaking work, The Interpretation of Development and Heredity. In this book Russell not only condemned Mendelian genetics and neo-Darwinism, but also proposed an alternative synthesis unifying heredity, development, and evolution. The book did not represent the work of a mind operating in isolation. Rather, it was a synthetic work connecting ideas and doctrines of many influential scientists working in Europe and the USA. Through the analysis of archival documents and rarely or never mentioned sources, this article provides an unconventional picture of Russell’s theoretical biology. It will be shown that Russell was an international celebrity; he was at the centre of a large network of scholars who shared his ideas and insights. He was one of several biologists arguing for a different synthesis; a synthesis perhaps less visible, less institutionalised, and less ‘modern’, nevertheless with its influential advocates and international support. Finally, this study shows that Russell’s synthesis was not rooted in the classic pantheon of towering figures in the history of biology, i.e. Darwin, Wallace, and Mendel, but was based on the teachings of Kant, Goethe, Cuvier, von Baer, and Müller.

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Notes

  1. See Levit et al. (2006), see also Reif et al. (2000).

  2. Smocovitis (1992, 1996).

  3. See also McRae (1961) and Hjort (1921).

  4. See Holton (2003).

  5. See Holton (2003).

  6. The English edition was revised by E.S. Russell.

  7. Source: http://icesjms.oxfordjournals.org/content/60/6/1169.full.

  8. See John Graham Kerr." Encyclopædia Britannica. Encyclopædia Britannica Online Academic Edition. Encyclopædia Britannica Inc., 2013. Web. 09 Jan. 2013.

  9. See Graham (1954).

  10. On Thomson’s background see Heron-Allen (1933).

  11. See the archive of PhD theses at the University of Glasgow. See the following link: http://eleanor.lib.gla.ac.uk/record=b1633964 (accessed January 7, 2013).

  12. See Haines (2004).

  13. See Roll-Hansen (1984).

  14. The translation of the letter is my own from German.

  15. Letter from von Bertalanffy to Ritter, October 22nd, 1931, Ritter Papers, 71/3 c, Box 6, Folder, Bertalanffy 1901, Bancroft Library Archive, University of Berkeley, California.

  16. Quoted in von Bertalanffy (1933, p. 4).

  17. For a general history of the journal, see Linguerri (2005).

  18. Source: Sarton (1931, p. 2).

  19. See Stonequist (1930), and Sarton (1931).

  20. Enriques (1930).

  21. Rignano (1926a).

  22. Rignano (1931). The German edition of Biological Memory (Rignano 1926b), with von Bertalanffy’s introduction, was published after Rignano’s death in 1930. The British edition was published in 1926 and introduced by the embryologist E.W. MacBride (1866–1940).

  23. Not surprisingly, the English edition was translated in Chicago; indeed, Rignano was quite well known by the biologists based at the Zoological Department of the University of Chicago. As an example of how mnemonic theories of heredity and development were regarded as relevant by the Chicago group, there is an interesting letter from the director, Whitman, in 1909, to the celebrated psychologist and primatologist R. Yerkes (1876–1956). Whitman said: “Mnemonic phenomena are in my opinion at the basis of vital phenomena of every order. Psychology and physiology seem blend here, and I find no point at which they become separate. Yet it may be useful to classify as psychological, actions in which sense organs and brain are conspicuously concerned. On this point I have no right to express an opinion, as you will readily grant.” (Whitman’s letter to Yerkes, October, 24th, 1909 in Lillie Papers, Box IIA, Folder: 99, MBL Archive, Woods Hole). Relations between psychology and physiology, as well as ontogeny and phylogeny, were all central topics in Rignano’s book translated the following year in Chicago. .

  24. Thomson, in his Wonder of Life (1914) defined well mnemonic theories of heredity: “The term ‘mnemic’… is applied to the theories…according to which the germ-cells are supposed to treasure up some of the results of the organism’s experience, as it were, by unconscious memory, so that when they come to develop they reproduce in some measure the traits which their parents or their ancestors acquired as the result of experience. The idea is that the germ-cells become stored with the latent ‘memories’ of past generations, or less metaphorically that the germ-cells are changed or impressed in a definite and specific way by the organism’s experience. Development is in part the ‘recollection’ of these germinally treasured ‘memories” (Thomson 1914, p. 753).

  25. My translation from the French. Yet, in 1912 Russell (1912) reviewed Rignano’s English edition and concluded that: “We are convinced that the solution for the problem of heredity and development must be sought in the particular phenomena that Rignano, with a rare acquaintance, has chosen as specifically important, and we deeply recommend his work, which constitutes to our eyes the best attempt so far to produce a simpler and more rational solution in the framework of a positive philosophy.”(Russell 1912, p. 439) My translation from French.

  26. Russell (1911, pp. 329–345).

  27. See Russell (1932).

  28. Owen first introduced the distinction between analogical and homological organs which represented, Russell argued, a distinction between structure and function: indeed, function, as a teleological property, characterised analogical organs.

  29. Russell here is referring to the law according to which the structures visible during the earlier stages of ontogeny are more common among all organisms than structures appearing in later ontogenic stages. This subject is treated in Richards (1992).

  30. In Russell’s own words: “we must regard the organism as an historical being and interpret its present structure and activities in the light of its past history.” (Russell 1911, p. 343) .

  31. See Russell (1924, p. VII).

  32. Russell: “…it is mainly through perception that life becomes individualized and separates itself out from the environing flux. Through perception the organism clears, as it were, a space around it in which to live, and disposes of time in which to protect itself against the surrounding influences which would imminently destroy it.” (Russell (1932, p. 59).

  33. Russell reported several examples of creative behaviour among Protista and unicellular organisms, see Russell (1924, pp. 65–81).

  34. Russell, in referring to the work of Kepner and Edwards on Amoeba and Pelomyxa feeding habits (Kepner and Whitlock 1917), stated: “From this review of the feeding responses of Amoeba and Pelomyxa, as observed with care and without prejudice by skilled and competent workers, it is apparent how much simpler and adequate is the psychological interpretation than a materialistic. Any attempted restatement in terms of surface tension or rigidly determined tropisms would be lame and hypothetical in comparison.” (Russell 1924, p. 76).

  35. On the way to study animal behaviour in general, Russell proposed a list of rules that the functional biologist should follow: (1) actions are responses of the organism as a whole; (2) actions must be studied in relation to the reciprocal influence of the hormic impulse and the perceived situation; (3) the living organism must be studied in his natural environment first; (4) in an experimental setting, the responses of an organism must always be interpreted as if the organism was responding to stimuli happening in its natural setting; (5) the analysis of physical stimuli must be centred on the analysis of the ‘meaning’ of the overall situation in the environment, i.e., what is important to the organism or what is not, etc.; (6) any response of an organism has a goal or aim toward an object; it is significant, (7) the aim of an action can be found in observing the results of specific responses, or from the observation of the all chain of actions bringing to satisfaction of cessation of the activity; (8) responses can be classified as instances of aims without any respect to the physical nature of the stimulus. See Russell (1924, pp. 79–80).

  36. Thompson to Russell, 29/12/1916, letter no. 14329, DTP-SAA.

  37. Russell to Thompson, 6/01/1917, letter no. 14333, DTP-SAA.

  38. On Ludwig Plate see Levit and Hossfeld (2006).

  39. My translation from the French.

  40. My translation from the French.

  41. About the multiple historical relations between developmental biology and evolutionary studies, see Olsson et al. (2010).

  42. The similarity between von Baer’s and Child’s conception of reproduction is striking.

  43. Von Baer quoted in Russell (1930, p. 35).

  44. See also Dembowski (1926).

  45. On the Mendelian tradition see Bowler (1989); Simunek et al. (2010, 2011).

  46. See Johannsen (1909).

  47. As Russell described it: “In a short but fundamental paper he lays [Johannsen] his finger on the limitations of modern genetical theory, and assesses its general significance in masterly fashion.” (1930, p. 63). As F. Churchill reports, Johannsen was also harshly critical of the British school of biometricians, as exemplified by Weldon and Pearson. He used to say indeed: “We must pursue the science of heredity but not as mathematics”, Johannsen, in Churchill (1974, p. 8).

  48. Then Johannsen concluded: “…the phenotype being the reaction of the genotype (nature) with the ambient conditions (nurture).” (Johannsen 1923, p. 141). See also Johannsen 1911.

  49. In 1922, so before that The Theory of the Gene was published, E. Bauer, a Czechoslovak zoologist, gave a very clear and succinct list of ideas defining Mendelism in his own time. As he saw it, Mendelism is based on the ideas that “…gametes contain certain kind of particles we can consider as physical entities, and these cause an organism developing by to ontogenesis to have certain characters. This process should be conceived of as follows: each of the above mentioned particles has the ability to cause in a given organism one particular character. Thus if for example in a given gamete ten such particles are present, and we number label them one to ten, then these ten particles can cause only ten characters in the developing organism. The first particle cause one character only that corresponds to number one, particle two only the character corresponding to number two, etc. Conversely, to each character exhibited by an organism corresponds a particle, that is, as one would say in mathematics, particles in gametes and properties of an organism developed from such gamete are unequivocally, bi-directionally coordinated.

    Another idea is that these particles that determine the properties of a developing organism are completely independent from each other, i.e., they do not blend during fertilisation but only mix. Therefore, any combination of these particles may occur in descendants of a given fertilisation. These are the basic tenets of Mendelian theory. The particles we have discussed are called genes, and the theory developed from the above-mentioned basic ideas is called Mendelism.” (Bauer, in Simunek et al. 2010, pp. 234–235). This was precisely the doctrine that Russell rejected; both on theoretical and empirical grounds.

  50. See Brachet (1917), see also Mulnard (1992).

  51. As Russell claimed: “It is these special resemblances and differences that have been the subject of the modern study of heredity, whether by biometrical or by genetical methods. The broad general resemblances of type give no hold for experimental or statistical treatment, and have accordingly on the whole been ignored. But it is this general hereditary resemblance which constitutes the main problem. We saw in discussing the gene theory that it deals only with differences between closely allied forms, and with the modes of inheritance of these differences; it leaves the main problem quite untouched as to why, for example, from a pair of Drosophila only Drosophila arise. It takes for granted the inheritance of Johannsen’s ‘great central something’—the general hereditary equipment of the species” (Russell (1930, p. 270).

  52. In particular Lillie’s “The Gene and the Ontogenetic Process”, an article published in Science in 1927 (Lillie 1927). In this paper, Lillie had argued that gene theory could not help to embryology and developmental theories.

  53. Russell also included in his list Whitehead, who, as a philosopher, had provided the theoretical scaffolding for organismal theory.

  54. Russell referred to Bernard’s La Science Experimentale, a book first published in 1878 where Bernard effectively admitted: “In saying that life is a directive idea or an evolutive force of being, we simply convey the idea that there is an unity in all morphological and chemical changes that are produced, at beginning, by the germ until the end of life”, my translation from French, see Bernard (1878, p. 430).

  55. As Russell wrote of the book: “He exhibits throughout a clarity of thought, fidelity to the facts of observations, and a sobriety of hypothesis that make his book one of the finest contributions to general biology ever written”(Russell 1930, p. 84).

  56. As Russell specified: “All parts and organs are, as Kant would say, reciprocally means and ends, and all cooperate in the life of the organism as a whole. For, as D’Arcy Thompson, ‘the life of the body is more than the sum of the properties of the cells of which it is composed.” (Russell 1930, p. 148).

  57. See Russell (1930, pp. 92–93). As Russell emphasised: “This principle of unity, or action of the organism as a whole, corresponds to Whitman’s concept of organization…the same view, that development is essentially an activity of the organism as a whole, has also been upheld by others—by Conklin and Child for instance, and by Ritter…” (Russell 1930, p. 244).

  58. See Russell (1930, p. 189).

  59. As Russell explained: “If the activity of the organism as a whole is not completely reducible to the modes of action of its parts, then it follows that the modes of action of the whole, whether actual or potential, can be transmitted only by a whole, i.e. by the egg in its entirety, which at very beginning of development is the new individual. Subordinate parts of the egg-organism can transmit only their own particular modes of action, and not the modes of action of the whole; they cannot transmit even their own modes save as integral parts of the whole.” (Russell 1930, p. 283).

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Acknowledgments

I would like to thank the Institute of Philosophical Research at UNAM for supporting this work. I am also grateful for the comments and advices I received at the Philosophy of Biology seminars held at the Metropolitan Autonomous University of Mexico (Cuajimalpa), where a first draft of this paper was presented.

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    Maurizio Esposito

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Esposito, M. Heredity, development and evolution: the unmodern synthesis of E.S. Russell. Theory Biosci. 132, 165–180 (2013). https://doi.org/10.1007/s12064-013-0177-4

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