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The conquest of vitalism or the eclipse of organicism? The 1930s Cambridge organizer project and the social network of mid-twentieth-century biology


In the 1930s, two concepts excited the European biological community: the organizer phenomenon and organicism. This essay examines the history of and connection between these two phenomena in order to address the conventional ‘rise-and-fall’ narrative that historians have assigned to each. Scholars promoted the ‘rise-and-fall’ narrative in connection with a broader account of the devitalizing of biology through the twentieth century. I argue that while limited evidence exists for the ‘fall of the organizer concept’ by the 1950s, the organicism that often motivated the organizer work had no concomitant fall – even during the mid-century heyday of molecular biology. My argument is based on an examination of shifting social networks of life scientists from the 1920s to the 1970s, many of whom attended or corresponded with members of the Cambridge Theoretical Biology Club (1932–1938). I conclude that the status and cohesion of these social networks at the micro scale was at least as important as macro-scale conceptual factors in determining the relative persuasiveness of organicist philosophy.

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1 Haraway Donna, Crystals, Fabrics, and Fields: Metaphors That Shape Embryos, reprinted with a foreword by S.G. Berkeley: North Atlantic Books, 2004, p. 2.

2 See ‘Correspondence with June [Goodfield] Toulmin of the Nuffield Foundation, London’, August 1962–May 1963, Needham Papers, Cambridge University Library (hereafter JN/CUL), M.89.

3 Joseph Needham to June Goodfield-Toulmin, 26 April 1965, JN/CUL, M.89.

4 C.H. Waddington to June [Goodfield] Toulmin, 9 April 1965, JN/CUL, M.89.

5 Spemann Hans and Mangold Hilde, ‘Induction of embryonic primordia by implantation of organisers from a different species’ (1924) (tr. Viktor Hamburger), International Journal of Developmental Biology (1965) 45, pp. 1338. See also Horder Timothy J., ‘The organizer concept and modern embryology: Anglo-American perspectives’, International Journal of Developmental Biology (2001) 45, pp. 97132.

6 De Robertis E.M., ‘Spemann's organiser and self-regulation in amphibian embryos’, Nature Reviews Molecular Cell Biology (2006) 7, pp. 296302, 297, argues that works by Barth L.G. (‘Neural differentiation without organizer’, Journal of Experimental Zoology (1941) 87, pp. 371383) and Holtfreter Johannes (Concepts on the Mechanism of Embryonic Induction and Its Relation to Parthenogenesis and Malignancy, Growth in Relation to Differentiation and Morphogenesis, New York: Academic Press, 1948) were the final two on the organizer before the 1990s. However, the work of Eakin Richard M. (‘The nature of the organiser’, Science (1949) 109, pp. 195197) and Ficq A. (‘Analyse de l'induction neurale par autoradiographie’, Experientia (1954) 10, pp. 2021) provide clear indications that substantial work on the chemical nature of the organizer continued well after the Second World War, contra Goodfield-Toulmin's suppositions.

7 Hans Driesch was the best-known ‘vitalist’ in the twentieth century. Allen Garland, ‘Mechanism, vitalism and organicism in late nineteenth and twentieth-century biology: the importance of historical context’, Studies in History and Philosophy of Science Part C: Studies in History and Philosophy of Biological and Biomedical Sciences (2005) 36, pp. 261283. However, De Robertis, op. cit. (6), p. 299, recollects the common knowledge from that era that ‘Spemann's organizer set developmental biology back by 50 years’, due to his vitalism. See also Harrington Anne, Reenchanted Science: Holism in German Culture from Wilhelm II to Hitler, Princeton: Princeton University Press, 1996.

8 Non-epistemic values were presumed to motivate the ‘classic’ case of Lysenko's Trofim D. agronomy (for example Julian Huxley, Heredity, East and West: Lysenko and World Science, New York: Schuman, 1949; and Megay E.N., ‘Lysenkoism and the stateless society’, Journal of Politics (1953) 15, pp. 211230).

9 Toulmin Stephen and Goodfield-Toulmin June, The Architecture of Matter, London: Hutchinson, 1962, p. 378.

10 Waddington to June [Goodfield] Toulmin, 9 April 1965 (JN/CUL, M.89).

11 The narrative that mechanistic biology would ensure progress while mystical vitalism would halt it had been made earlier in the century. For example: Elkus A.S., ‘Mechanism and vitalism’, Journal of Philosophy, Psychology and Scientific Methods (1911) 8, pp. 355358; and Lillie Ralph S., ‘The philosophy of biology: vitalism versus mechanism’, Science, new series (1914) 40, pp. 840846.

12 Benson highlighted the ‘organism’ concept still earlier. Benson Keith R., ‘Biology's “phoenix”: historical perspectives on the importance of the organism’, American Zoologist (1989) 29, pp. 10671074.

13 Gilbert Scott F. and Sarkar Sahotra, ‘Embracing complexity: organicism for the twenty-first century’, Developmental Dynamics (2000) 219, pp. 19, 4–5.

14 My focus on network is partly to do with the nature of the relationships between the main actors in this story – Needham, Waddington and Woodger – the core of the Theoretical Biology Club (TBC) in the 1930s. A few studies have been published addressing the work of one or another member of this network, namely Abir-Am Pnina, ‘The discourse of physical power and biological knowledge in the 1930s: a reappraisal of the Rockefeller Foundation's “policy” in molecular biology’, Social Studies of Science (1982) 12, pp. 341382; Abir-Am, ‘The “Biotheoretical Gathering” in England and the origins of molecular biology (1932–38)’, unpublished dissertation, Institut d'histoire et de sociopolitique des sciences, Université de Montréal, Montreal, Canada, 1983; Abir-Am, ‘The Biotheoretical Gathering, trans-disciplinary authority and the incipient legitimation of molecular biology in the 1930s: new perspective on the historical sociology of science’, History of Science (1987) 25, pp. 170; Abir-Am, ‘The philosophical background of Joseph Needham's work in chemical embryology’, in Gilbert Scott F. (ed.), A Conceptual History of Modern Embryology, Baltimore: Johns Hopkins University Press, 1994, pp. 159180; Gilbert Scott F., ‘Epigenetic landscaping: Waddington's use of cell fate bifurcation diagrams’, Biology and Philosophy (1991) 6, pp. 135154; Gilbert, ‘Induction and the origins of developmental genetics’, in Gilbert, A Conceptual History of Modern Embryology, op. cit., pp. 181–206; Gilbert, ‘Diachronic biology meets evo-devo: C.H. Waddington's approach to evolutionary developmental biology’, American Zoologist (2000) 40, pp. 729737; Hall Brian K., ‘Waddington's legacy in development and evolution’, American Zoologist (1992) 32, pp. 113122; Hall, ‘In search of evolutionary developmental mechanisms: the 30-year gap between 1944 and 1974’, Journal of Experimental Zoology: Molecular and Developmental Evolution (2004) 302B, pp. 518; Smocovitis V. Betty, ‘Serious matters: on Woodger, positivism, and the evolutionary synthesis’, Biology and Philosophy (2000) 15, pp. 553558; and Yoxen Edward, ‘Form and strategy in biology: reflections on the career of C.H. Waddington’, in Horder Timothy J., Witkowski J.A. and Wylie C.C. (eds.), A History of Embryology, Cambridge: Cambridge University Press, 1986. My focus on social networks takes cues from Dupré John, The Disorder of Things: Metaphysical Foundations of the Disunity of Science, Cambridge, MA: Harvard University Press, 1993; Laudan Larry, Beyond Positivism and Relativism: Theory, Method, and Evidence, Boulder, CO: Westview Press, 1996; and Latour Bruno, Reassembling the Social: An Introduction to Actor-Network Theory, Clarendon Lectures in Management Studies, Oxford and New York: Oxford University Press, 2005.

15 Huneman Phillipe and Wolfe Charles T., ‘The concept of organism: historical, philosophical, scientific perspectives’, History and Philosophy of the Life Sciences (2010) 32, pp. 147154.

16 Haldane John Scott, Organism and Environment as Illustrated by the Physiology of Breathing, New Haven: Yale University Press, 1917, p. 3 n. 1.

17 Haldane, op. cit. (16), p. 17.

18 As late as 1950, Russell proclaimed, ‘If we look at living things quite simply and objectively we cannot but be struck by one feature of their activities, which seems to mark them off from anything inorganic … [T]here is common to all living things this basic element of directive striving, usually unconscious and blind, only rarely emerging into consciousness to become intelligently purposive’. Russell E.S., ‘The “drive” element in life’, British Journal for the Philosophy of Science (1950) 1, pp. 108116, 108.

19 Russell E.S., Form and Function: A Contribution to the History of Animal Morphology, London: J. Murray, 1916, pp. vvi.

20 Russell E.S., The Interpretation of Development and Heredity: A Study in Biological Method, Oxford: Clarendon Press, 1930, pp. 306307.

21 Ritter William E., The Unity of the Organism; or, the Organismal Conception of Life, 2 vols., Boston: R.G. Badger, 1919, vol. 1, p. xix. Emphasis is his, versus ‘theories’.

22 Smuts J.C., Holism and Evolution, New York: The Macmillan Company, 1926, p. 8.

23 Mayer Jean, ‘L.J. Henderson: a biographical sketch’, Journal of Nutrition (1968) 94, pp. 15; and Cross Steven J. and Albury William R., ‘Walter B. Cannon, L.J. Henderson, and the organic analogy’, Osiris (1987) 3, pp. 165192.

24 E.g. Nordenskiöld Erik, The History of Biology: A Survey (tr. L.B. Eyre), New York and London: A.A. Knopf, 1928, pp. 525527; and Thompson D'Arcy W., On Growth and Form, 2nd edn (ed. Lancelot L. Whyte), Cambridge: Cambridge University Press, 1942, p. 345 : ‘We then deal not with material continuity … but with a continuity of forces, a comprehensive field of force’.

25 C.D. Broad, The Mind and Its Place in Nature, International Library of Psychology, Philosophy, and Scientific Method, London: Routledge & Kegan Paul, 1925, pp. 58–59: Emergentism ‘explain[s] the difference of behavior wholly in terms of difference of structure’.

26 See Meyer-Abich Klaus, ‘Bohr's complementarity and Goldstein's holism in reflective pragmatism’, Mind and Matter (2004) 2, pp. 91103.

27 Whitehead Alfred North, Science and the Modern World, Lowell Lectures, 1925, New York: Free Press, 1953, pp. 138157.

28 American biologist E.E. Just provides another important reference point: ‘[Just's organicism] represent[ed] the standard opinion of most practicing biologists and, as such, refute[s] the dichotomous scheme that sees biology as a war between vitalists and mechanists’. Gould Stephen J., ‘Just in the middle’, in Gould, The Flamingo's Smile: Reflections in Natural History, New York: W.W. Norton & Company, 1987, pp. 377391, 388.

29 Greenwald A.G., Pratkanis A.R., Leippe M.R. and Baumgardner M.H., ‘Under what conditions does theory obstruct research progress?’, Psychological Review (1986) 93, pp. 216229.

30 Spemann Hans, ‘The organiser-effect in embryonic development: Nobel Lecture, December 12, 1935’, in The Nobel Foundation (eds.), Nobel Lectures, Physiology or Medicine 1922–1941, Amsterdam: Elsevier, 1965,, accessed 28 August 2009.

31 Spemann and Mangold, op. cit. (5).

32 Hamburger Viktor, ‘Hans Spemann on vitalism in biology: translation of a portion of Spemann's autobiography’, Journal of the History of Biology (1999) 32, pp. 231243, 233.

33 Horder Timothy J. and Weindling Paul, ‘Hans Spemann and the organizer’, in Horder T.J., Witkowski J.A. and Wylie C.C. (eds.), A History of Embryology, Cambridge: Cambridge University Press, 1986, pp. 183242, 221; and Hamburger, op. cit. (32), p. 234.

34 See, for instance, the relazione published by the Istituto di Istologia-Embriologia (II-E) della R. Università di Padova (1931–1932) sent to Joseph Needham by Tullio Terni, director. Padova's II-E was funded by the Rockefeller Foundation. Spemann and Ross Harrison set up the micro-instrumentation. Terni then travelled to the labs of Spemann and Mangold to learn proper techniques. JN/CUL, E.111.

35 Allen Garland, ‘A pact with the embryo: Viktor Hamburger, holistic and mechanistic philosophy in the development of neuroembryology, 1927–1955’, Journal of the History of Biology (2004) 37, pp. 421475; and Witkowski J.A., ‘Optimistic analysis: chemical embryology in Cambridge 1920–42’, Medical History (1987) 31, pp. 247268.

36 Needham hoped that Chemical Embryology, 3 vols., Cambridge: Cambridge University Press, 1931, would initiate a ‘new branch’ of science, but it nearly led to a nervous breakdown. Henry Holorenshaw (pseud. for Joseph Needham), ‘The making of an honorary taoist’, in Needham Joseph, Teich Mikulás and Young Robert M. (eds.), Changing Perspectives in the History of Science: Essays in Honour of Joseph Needham, London: Heinemann Educational, 1973, pp. 120, 7; and Winchester Simon, The Man Who Loved China, New York: Harper Perennial, 2008, p. 28.

37 Popper Karl R., ‘Obituary: Joseph Henry Woodger’, British Journal for the Philosophy of Science (1981) 32, pp. 328330.

38 Popper, op. cit. (37), p. 328. Popper introduced Alfred Tarski to Woodger, who would translate a number of Tarski's works into English. Popper established a gentle, persistent rivalry with Woodger regarding Tarski and neither Woodger nor the Theoretical Biology Club meeting that Popper attended appear in Popper's autobiography. See Popper Karl R., Unended Quest, New York: Routledge, 2002; and Hacohen Malachi H., Karl Popper, the Formative Years, 1902–1945: Politics and Philosophy in Interwar Vienna, Cambridge: Cambridge University Press, 2000, p. 311 n. 68.

39 Needham Joseph, ‘Review of J.H. Woodger, Biological Principles, a Critical Study’, Mind (1930) 39, pp. 221226.

40 Woodger J.H., Biological Principles: A Critical Study, London: K. Paul, Trench, Trubner & Co., 1929, pp. 482487. While serving in the First World War, Woodger befriended Ian Suttie, a psychotherapist whose holistic concept of the developing self deeply impacted Woodger's philosophy. Floyd W.F. and Harris F.T.C., ‘Joseph Henri [sic] Woodger, Curriculum Vitae’, in Gregg J.R. and Harris F.T.C. (eds.), Form and Strategy in Science: Studies Dedicated to Joseph Henry Woodger on the Occasion of His Seventieth Birthday, Dordrecht: Reidel, 1964, pp. 16; see also the correspondence between Woodger and Suttie, 1919 to 1925, Woodger's Papers, University College London (hereafter JHW/UCL), C1/3/Suttie.

41 Abir-Am, ‘The philosophical background of Joseph Needham's work’, op. cit. (13).

42 Woodger J.H., ‘The “concept of organism” and the relation between embryology and genetics, part III’, Quarterly Review of Biology (1931) 6, pp. 178207, 206.

43 Needham admitted that ‘in all probability [he and von Bertalanffy] could come to agreement’. Needham to Woodger, 12 October 1929; see correspondence from 8 October 1929 to 2 April 1930, JN/CUL, M.97.

44 von Bertalanffy Ludwig, ‘An outline of General Systems Theory’, British Journal for the Philosophy of Science (1950) 1, pp. 134165; and von Bertalanffy, ‘The theory of Open Systems in physics and biology’, Science (1950) 111, pp. 2329.

45 von Bertalanffy Ludwig and Woodger J.H., Modern Theories of Development: An Introduction to Theoretical Biology (tr. J.H. Woodger), London: Oxford University Press, 1933.

46 ‘Mechanism … provides us with no grasp of the specific characteristics of organisms, of the organization of organic processes among one another, of organic “wholeness”, of the problem of the origin of organic “teleology”, or of the historical character of organisms.’ Von Bertalanffy and Woodger, op. cit. (45), p. 46.

47 Von Bertalanffy and Woodger, op. cit. (45), pp. 1, 5.

48 Von Bertalanffy and Woodger, op. cit. (45), pp. 10–11.

49 Harrington, op. cit. (7), pp. xvi–xx; and see, for instance, Heske Frans, Jordan Pascual and Meyer-Abich Adolf (eds.), Organik: Beiträge zur Kulture unserer Zeit, Berlin: Fritz Haller, 1954.

50 Woodger J.H., ‘The “concept of organism” and the relation between embryology and genetics, part I’, Quarterly Review of Biology (1930) 5, pp. 122; Woodger, ‘The “concept of organism” and the relation between embryology and genetics, part II’, Quarterly Review of Biology (1930) 5, pp. 438463; and Woodger, op. cit. (42).

51 Needham Joseph, The Sceptical Biologist, New York: W.W. Norton, 1930, pp. 8586.

52 Woodger, part I, op. cit. (50), p. 7, here quoting G.K. Chesterton.

53 Woodger, part I, op. cit. (50), pp. 6–7; and Woodger, op. cit. (42), p. 207.

54 Goldsmith Maurice, ‘Joseph Needham, honorary taoist’, in Needham J., Mukherjee S. Kumar and Ghosh A. (eds.), The Life and Works of Joseph Needham, Calcutta: The Asiatic Society, 1997, pp. 13.

55 Holorenshaw, op. cit. (36), p. 10; and Needham Joseph, Time: The Refreshing River (Essays and Addresses, 1932–1942), London: G. Allen & Unwin, 1943, pp. 149 and 241–242. Gilbert and Sarkar, op. cit. (13), p. 5, suggested that this more explicit left turn in the political leanings of its supporters ultimately hampered the acceptance of organicism. The cases of both J.B.S. Haldane and H.J. Muller would seem to speak against this reading. Both men were acknowledged leftists and did not lose support for their reductionism.

56 Woodger to Needham, 7 July 1931, JN/CUL, M.97; and Woodger to Needham, 30 April 1932, JN/CUL, M.98.

57 Yoxen, op. cit. (14), pp. 312–313.

58 Woodger to Needham, 23 April 1936, JN/CUL M.99.

59 Abir-Am,‘The philosophical background of Joseph Needham's work’, op. cit. (14), pp. 173–174.

60 Winchester, op. cit. (36), p. 30.

61 Waddington C.H., ‘Pollen germination in stocks and the possibility of applying lethal factor hypothesis to the interpretation of their breeding’, Journal of Genetics (1929) 21, pp. 193206, was written under the direction of Edith Saunders, a long-time collaborator with William Bateson at the John Innes Horticultural Institute. About his collaboration with J.B.S. Haldane (Waddington C.H. and Haldane J.B.S., ‘Inbreeding and linkage’, Genetics (1931) 16, pp. 357374), Waddington could only say that he had ‘tasted the thin gruel of mathematical formalism’. Waddington C.H., The Evolution of an Evolutionist, Edinburgh: Edinburgh University Press, 1975, p. 7.

62 Waddington composed two substantial, though unpublished, essays on the conjunction of philosophy and biology. The first, entitled ‘Philosophy and biology’ (1929), was awarded the Arnold Gerstenberg 1851 Scholarship. Waddington papers, University of Edinburgh Library Special Collections (hereafter CHW/UEL), MS 3024.2. In the second, ‘The process of abstraction and the vitalist – mechanist controversy’ (1931), Waddington extolled the philosophy of C.D. Broad (CHW/UEL, MS 3024.3).

63 Yoxen, op. cit. (14), p. 313.

64 Mukherjee and Ghosh, op. cit. (54), 173–92; and Robertson Alan, ‘Conrad Hal Waddington. 8 November 1905–26 September 1975’, Biographical Memoirs of Fellows of the Royal Society (1977) 23, pp. 575622, 615–621. During the same six-year period, Joseph Needham and Waddington produced a surprising number of papers and books, including Needham Joseph, A History of Embryology, New York: Abelard-Schuman, 1934; Needham, Order and Life: The Terry Lecture, New Haven: Yale University Press, 1936; Needham, Integrative Levels: A Revaluation of the Idea of Progress, Herbert Spencer Lectures, Oxford: Clarendon Press, 1937; and Waddington C.H., How Animals Develop, London: G. Allen & Unwin, Ltd, 1935.

65 Abir-Am, ‘The Biotheoretical Gathering’, op. cit. (14), pp. 34–41, details the multiple marginality of group members due to their political, scientific, philosophical and social views as well as their unstable academic positions.

66 Woodger to Needham, 30 April 1932, JN/CUL M.98. Though initially attendees assembled at Woodger's summer cabin, this spot was not conducive for those to the north and the Woodgers occasionally alternated hosting with the Needhams, who had a holiday home at Ringstead Mill in East Anglia.

67 Needham to Woodger, 19 August 1932, JN/CUL M.98. ‘Theoretical Biology Club’ appeared on the dedication page of Needham's Order and Life (1936). Woodger used ‘biotheoretical gathering’ (uncapitalized) in his 30 April 1932 ‘start-up’ letter to Needham (JN/CUL M.98). Abir-Am adopted ‘Biotheoretical Gathering’ (capitalized) in ‘The Biotheoretical Gathering’, op. cit. (14), p. 55 n. 3. Other participants referred to it as a ‘Conference’ (Dorothy Wrinch to Woodger, 23 May 1935), ‘Geschäftsordnung’ or ‘Woodgery’ (Woodger to Needham, 23 April 1936, JN/CUL M.99). Given that Needham's term made it to print – whereas Woodger's term was informal and only appeared inconsistently and in private letters – I will continue to use ‘Theoretical Biology Club’/‘TBC’.

68 Slack Jonathan M.W., ‘Conrad Hal Waddington: the last Renaissance biologist?’, Nature Reviews, Genetics (2002) 3, pp. 889895.

69 Here Bernal drew from Thompson, op. cit. (24). See Olby Robert C., ‘Structural and dynamical explanations in the world of neglected dimensions’, in Horder T.J., Witkowski J.A. and Wylie C.C. (eds.), A History of Embryology, Cambridge: Cambridge University Press, 1985, pp. 275308, 281.

70 Davis Gregory K., Dietrich Michael R. and Jacobs David K., ‘Homeotic mutants and the assimilation of developmental genetics into the evolutionary synthesis, 1915–1952’, in Cain J. and Ruse M. (eds.), Descended from Darwin: Insights into the History of Evolutionary Studies, 1900–1970, Philadelphia: American Philosophical Society, 2009, pp. 139140. For TBC meeting notes see JN/CUL, E.111 and J.243.

71 Davis, Dietrich and Jacobs, op. cit. (70), p. 139, fault Spemann for categorizing the organizer as ‘an irreducible, holistic phenomenon’. The September 1933 TBC notes demonstrate that the Needham–Waddington critique of Spemann was not his holism but his quasi-vitalism (JN/CUL, E.111).

72 Abir-Am, ‘The Biotheoretical Gathering’, op. cit. (14), p. 2.

73 Popper quoted by Hacohen, op. cit. (38), p. 315. Hunstanton is the seaside town nearest Ringstead, so Popper is referring to the Needham meeting site. Abir-Am, ‘The Biotheoretical Gathering’, op. cit. (14), p. 2, also places the meeting at Ringstead. While Popper recalls Bernal as being in attendance, Abir-Am does not. According to my examination of the 1936 ‘Whitsun[day]’ meetings, Bernal did attend. JN/CUL, J.244.

74 Popper Karl R., ‘Scientific reduction and the essential incompleteness of all science’, in Ayala F.J. and Dobzhansky T. (eds.), Studies in the Philosophy of Biology: Reduction and Related Problems, Berkeley: University of California Press, 1974, pp. 259284.

75 Werskey Gary, The Visible College: The Collective Biography of British Scientific Socialists of the 1930s, New York: Holt, Rinehart and Winston, 1979; Winchester, op. cit. (36); and see Needham's contribution to Waddington's obituary, 1975, JN/CUL, M.95.

76 Woodger to Needham, 2 April 1930, JN/CUL, M.97; and Waddington, The Evolution of an Evolutionist, op. cit. (61), p. 4.

77 Needham, op. cit. (55), p. 178–206; and Slack, op. cit. (68), p. 891.

78 Joseph Needham, ‘An account of the negotiations regarding financial aid for technical assistance, etc., for studies on the borderline between biochemistry and embryology’, unpublished memo, 12 July 1938, JN/CUL, E.114.

79 Abir-Am Pnina G., ‘The Rockefeller Foundation and the rise of molecular biology’, Endeavour (2001) 25, pp. 5560.

80 According to the Rockefeller Foundation's 1938 ‘Annual Report’, quoted in Kay Lily E., The Molecular Vision of Life: Caltech, the Rockefeller Foundation, and the Rise of the New Biology, New York: Oxford University Press, 1993, p. 49. Weaver coined the term ‘molecular biology’ and began explicitly channelling funding in this direction around 1938 (Kay, op. cit., pp. 188–189). See also Kohler Robert E., ‘The management of science: the experience of Warren Weaver and the Rockefeller Foundation programme in molecular biology’, Minerva (1976) 14, pp. 279306.

81 While Abir-Am (The discourse of physical power and biological knowledge in the 1930s, op. cit. (14), and ‘The Biotheoretical Gathering’, op. cit. (14)) has discussed the role of Needham's proposal for an MPC morphology unit in the history of the ‘Biochemical Laboratory Extension’, she has not addressed these other factors that contributed to its demise.

82 It was well known in Oxbridge scientific circles that ‘James Gray has a curious hostility to Experimental Embryology & related subjects.’ That curious hostility translated specifically into shifting grant money away from Waddington even before this Rockefeller Foundation funding incident. Julian Huxley to Needham, 12 June 1934, JN/CUL, M.34.

83 ‘[W]e have reason to believe that the Rockefeller Foundation approves the suggestion that proposals for grants to increase the stipends of University Officers should be referred to the Council and to the General Board’. ‘Memorandum on Applications for Grants’, approved by the Cambridge University Council on 1 March 1937, JN/CUL, B.26; ‘Memorandum on the position of C.H. Waddington’, enclosure with letter from Needham to Tisdale, 20 July 1936, JN/CUL, B.25; and Tisdale to Needham, 9 July 1936, JN/CUL, B.25. Waddington did secure another source of income, which allowed him to stay in Cambridge and work at Strangeways Laboratories at a reduced capacity. ‘Dean, Donaldson, Adrian, and Dale’ of Strangeways to Needham, 29 November 1936, JN/CUL, B.26.

84 Needham to Fredrick G. Hopkins, 26 June 1938, JN/CUL, B.28.

85 Hopkins to Needham, 15 August 1938, JN/CUL, B.28.

86 Hopkins to Needham, 20 September 1938, JN/CUL, B.28.

87 Needham, ‘Memorandum on Cambridge work on Organiser Phenomena’ (undated; I date it contextually to October 1938), JN/CUL, B.27.

88 Needham to Tisdale (marked ‘draft’), 21 November 1936; and Needham to Secretary of Trustees, Strangeways Laboratories, 27 November 1936 (both JN/CUL, B.26). Rockefeller continued this support from 1935 to 1938. Abir-Am, ‘The Biotheoretical Gathering’, op. cit. (14), p. 32.

89 H.M. Miller (Rockefeller Foundation) to Needham, 27 January 1936, JN/CUL, B.25.

90 Witkowski, op. cit. (35), p. 268.

91 Waddington C.H., ‘Studies on the nature of the amphibian organization centre – VII. Evocation by some further chemical compounds’, Proceedings of the Royal Society of London. B, Biological Sciences (1934–1990) (1938) 125, pp. 365372, 370–371.

92 Johannes Holtfreter, ‘Reminiscences on the life and work of Johannes Holtfreter’, in Gilbert, A Conceptual History of Modern Embryology, op. cit. (14), pp. 109–127; and Gerhart John, ‘Johannes Holtfreter: January 9, 1901–November 13, 1992’, Biographical Memoirs National Academy of Sciences (US) (1998) 73, pp. 209228.

93 Joseph Needham, ‘An account of the negotiations …’, 12 July 1938, JN/CUL, E.114.

94 Waddington wanted to return to Cambridge biology after the war, but realized he had ‘no respectable biological job and can't see how to get one. Gray has definitely stated that he will not provide any resources for embryology, even if more funds are available to his department’. Waddington to Needham, 3 October 1944, JN/CUL, M.95.

95 Woodger to Needham, 18 April 1937, JN/CUL, M.99.

96 Mead Margaret, Blackberry Winter: My Earlier Years, New York: Simon & Schuster, 1984, p. 249.

97 See notes from these later conferences in JN/CUL, J.245 and JHW/UCL, C1/M/22.

98 Waddington to Needham, 22 July 1938, JN/CUL, M.94.

99 Robertson, op. cit. (64), p. 592–593. Dobzhansky had just published Genetics and the Origin of Species, 1st edn, New York: Columbia University Press, 1937. Relations between Dobzhansky and Sturtevant were tense at this time and, while sources are silent on the degree to which they worked together in 1938–1939, Dobzhansky may have turned to Waddington for support. They did remain friends for the rest of their lives. Provine William B., ‘Origins of the genetics of natural populations series’, in Lewontin Richard C., Provine William B., Moore John and Wallace Bruce (eds.), Dobzhansky's Genetics of Natural Populations: I–XLIII, New York: Columbia University Press, 1982, p. 30; and see Waddington to Dobzhansky, 1 February 1974, Dobzhansky Papers, American Philosophical Society, Philadelphia (hereafter TD/APS), Mss.B. D65.

100 Needham, Application to ‘Foundation’ containing ‘List of Cambridge Papers on Organiser Phenomena’, 1939, JN/CUL, B.33.

101 Waddington to Needham, 15 November 1963, JN/CUL, M.95.

102 Though Waddington had published few papers on evolution at this time, Gregory Bateson initially piqued his interest in this subject as well as in genetics. See E.L. Peterson, ‘Finding mind, form, organism, and person in a reductionist age: the challenge of Gregory Bateson and C.H. Waddington to biological and anthropological orthodoxy, 1924–1980’, unpublished dissertation, University of Notre Dame, 2010, pp. 183–186.

103 The ‘mechanisms of evolution constitute problems of population genetics’. Dobzhansky, op. cit. (99), p. 11. Needham and Gregory Bateson were Waddington's chief sources of encouragement to undertake the project. Waddington C.H., Organisers and Genes, Cambridge: Cambridge University Press, 1940, p. viii.

104 Many years later, Waddington saw the lack of acceptance of his epigenetics as a corollary to the apathy that greeted T.H. Morgan's attempt at a synthesis between development and genetics. Waddington, The Evolution of an Evolutionist, op. cit. (60), p. 7.

105 Waddington, op. cit. (103), pp. 92–93. Effectively, Waddington anticipated by two decades the landmark paper, François Jacob and Jacques Monod, ‘On the regulation of gene activity’, read at Cold Spring Harbor Symposium on Quantitative Biology, at Cold Spring Harbor, New York, 1961.

106 Waddington later used a ‘crude analogy’ to explain these concepts: ‘you can't easily turn a millionaire's country house into a subway station, but it's not too difficult to turn it into a country club’. Waddington to Errol E. Harris, 10 December 1963, CHW/UEL, MS 3039.4. Raff Rudolph A., ‘Evo-devo: the evolution of a new discipline’, Nature Reviews Genetics (2000) 1, pp. 7479. On these topics see Brigandt Ingo, ‘From developmental constraint to evolvability: how concepts figure in explanation and disciplinary identity’, in Love Alan C. (ed.), Conceptual Change in Biology: Scientific and Philosophical Perspectives on Evolution and Development, Berlin: Springer, forthcoming; Sapp Jan, Beyond the Gene: Cytoplasmic Inheritance and the Struggle for Authority in Genetics, Oxford: Oxford University Press, 1987, p. 17; Burian Richard M., ‘Lillie's paradox – or, some hazards of cellular geography’, in Burian (ed.), The Epistemology of Development, Evolution, and Genetics, Cambridge: Cambridge University Press, 2005, pp. 183209; Horder Timothy J., ‘Syllabus for an embryological synthesis’, in Wake D.B. and Roth G. (eds.), Complex Organizational Functions: Integration and Evolution in Vertebrates, New York: John Wiley & Sons, 1989, pp. 315348, p. 340; Gilbert Scott F., Opitz J.M. and Raff Rudolph A., ‘Resynthesizing evolutionary and developmental biology’, Developmental Biology (1996) 173, pp. 357372, 361–362; and Amundson Ron, The Changing Role of the Embryo in Evolutionary Thought: The Roots of Evo-Devo, Cambridge: Cambridge University Press, 2005, pp. 175180.

107 Waddington C.H., ‘Canalization of development and the inheritance of acquired characters’, Nature (1942) 150, pp. 563565, 563.

108 Here Waddington drew explicitly from the work of Whitehead. See Peterson E.L., ‘The excluded philosophy of evo-devo? Revisiting C.H. Waddington's failed attempt to embed Alfred North Whitehead's “organicism” in evolutionary biology’, History and Philosophy of the Life Sciences (2011) 33, pp. 301320.

109 Waddington, op. cit. (107), p. 564.

110 Waddington, op. cit. (107), p. 565.

111 Waddington to Needham, 23 February (no year; I date it contextually to 1946), JN/CUL, M.95.

112 For a description see Slack, op. cit. (68), p. 891; and Gilbert, ‘Epigenetic landscaping’, op. cit. (13). One of Waddington's clearest arguments for the epigenetic landscape's applicability to modern evolutionary theory appeared in Waddington C.H., ‘Evolutionary adaptation’, in Tax Sol (ed.), Evolution after Darwin – The University of Chicago Centennial, Vol. 1: The Evolution of Life, Chicago: The University of Chicago Press, 1960.

113 Jablonka Eva and Lamb Marion J., Evolution in Four Dimensions: Genetic, Epigenetic, Behavioral, and Symbolic Variation in the History of Life, Cambridge, MA: MIT Press, 2005.

114 Contra Gilbert and Sarkar, op. cit. (13), pp. 4–5; and Harrington, op. cit. (7).

115 Waddington C.H. (ed.), Towards a Theoretical Biology – 1. Prolegomena. I.U.B.S. Symposia, Chicago: Aldine Publishing Company, 1968.

116 See, for instance, Mayr's explicit endorsement of ‘Holism–Organicism’ and his well-known attack on ‘Bean Bag genetics’. Mayr Ernst, The Growth of Biological Thought: Diversity, Evolution, and Inheritance, Cambridge, MA: Belknap Press, 1982, pp. 6667; and Mayr, ‘Where are we?’, Cold Spring Harbor Symposia on Quantitative Biology (1959) 2, 114. See also Dobzhansky Theodosius, ‘Introductory remarks’, in Ayala F.J. and Dobzhansky T. (eds.), Studies in the Philosophy of Biology: Reduction and Related Problems, Berkeley: University of California Press, 1974, pp. 13.

117 Note that only George Simpson included Waddington among the contributors to the contemporary theory of evolution in the 1974 conference, and fellow contributor Viktor Hamburger despaired that Waddington's ‘chapter’ was still ‘missing’ from the Modern Synthesis. Mayr Ernst, ‘G.G. Simpson’, in Mayr Ernst and Provine William B. (eds.), The Evolutionary Synthesis: Perspectives on the Unification of Biology, Cambridge, MA: Harvard University Press, 1980, p. 452; and Viktor Hamburger, ‘Embryology and the Modern Synthesis in evolutionary rheory’, in Mayr and Provine, op. cit., p. 97.

118 Ernst Mayr to Waddington, 6 November 1966, Mayr papers, Harvard University Archives, 74.7 (hereafter EM/HUA), Box 14, Folder 919; and Mayr to Daniel Polikoff, 28 July 1981, EM/HUA 74.7, Box 29, Folder 1339.

119 Dobzhansky to Waddington, 15 August 1959, TD/APS, Mss.B. D65, Series I, ‘Waddington’.

120 For more on this extended organicist network see Haraway, op. cit. (1); and Peterson, op. cit. (102).

121 For example, Varela F.J., Maturana H.R. and Uribe R., ‘Autopoiesis: the organization of living systems, its characterization and a model’, Biosystems (1974) 5, pp. 187196; Wake D.B., Roth G. and Wake M.H., ‘The problem of stasis in organismal evolution’, Journal of Theoretical Biology (1983) 101, pp. 211224; and Lewontin R.C., The Triple Helix: Gene, Organism, and Environment, Cambridge, MA: Harvard University Press, 2001.

122 Four influential symposia were dedicated to the question of organicism and reductionism in the 1960s and 1970s: Arthur Koestler's Alpbach Symposium ( Koestler Arthur and Smythies J.R., Beyond Reductionism: New Perspectives in the Life Sciences: The Alpbach Symposium 1968, New York: Macmillan, 1969); Waddington's theoretical biology group at the IUBS International Biological Program (CHW/UEL and EM/HUA); the September 1972 and June 1974 conferences – Reductionism in Biology and Mind in Nature – both held at the Villa Serballoni in Bellaglo, Italy (Ayala and Dobzhansky, op. cit. (116); and Cobb John B. and Griffin David R., Mind in Nature: Essays on the Interface of Science and Philosophy, Claremont: University Press of America, 1977).

123 Nagel Ernst, ‘Mechanistic explanation and organismic biology’, in Munson Ron (ed.), Man and Nature: Philosophical Issues in Biology, New York: Delta Books, 1971, pp. 19–32. Notably, Nagel focused his critique on the 1920s organicism of E.S. Russell and completely by-passed that of Woodger and von Bertalanffy.

124 Crick Francis, Of Molecules and Men, Seattle: University of Washington Press, 1966.

125 Waddington C.H., ‘No vitalism for Crick; review of Of Molecules and Men by Francis Crick’, Nature (1967) 216, pp. 202203. Other reviewers concurred that Crick was conflating organicism with vitalism to make a point about biological progress (for example Livingston Laura, ‘Review: Of Molecules and Men by Francis Crick’, Yale Journal of Biology and Medicine (1968) 40, p. 328).

126 Crick to Waddington, 9 November 1967, and Waddington to Crick, 27 December 1967, Francis Harry Compton Crick Papers, Wellcome Library for the History and Understanding of Medicine, London, PP/CRI/I/2/6/5.

127 Astbury W.T., ‘Molecular biology or ultrastructural biology?’, Nature (1961) 190, p. 1124.

128 Medawar Peter, ‘A biological retrospect’, Nature (1965) 207, pp. 13271330, 1328.

129 Benson, op. cit. (12), details some of these earlier attempts.

130 Callon Michael, ‘From science as an economic activity to socioeconomics of scientific research: the dynamics of emergent and consolidated techno-economic networks’, in Mirowski Phil and Sent Esther-Miriam (eds.), Science Bought and Sold: Essays in the Economics of Science, Chicago: The University of Chicago Press, 2002, pp. 277317.

131 Waddington C.H., Tools for Thought: How to Understand and Apply the Latest Scientific Techniques of Problem Solving, New York: Basic Books, 1977, p. 23.

132 Waddington C.H., ‘Fifty years on’, Nature (1975) 258, pp. 2021, 21.

133 Waddington, op. cit. (131), pp. 24–25.

Many thanks to Joe Cain and David Depew for an initial nudge in 2010, to Tim Resovsky for patient suggestions along the way, and to the anonymous reviewers for conscientious comments.

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