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The conquest of vitalism or the eclipse of organicism? The1930s Cambridge organizer project and the social networkof mid-twentieth-century biology

ERIK PETERSON

The British Journal for the History of Science / FirstView Article / November 2013, pp 1 - 24DOI: 10.1017/S0007087413000435, Published online: 29 October 2013

Link to this article: http://journals.cambridge.org/abstract_S0007087413000435

How to cite this article:ERIK PETERSON The conquest of vitalism or the eclipse of organicism? The 1930s Cambridgeorganizer project and the social network of mid-twentieth-century biology. The British Journal forthe History of Science, Available on CJO 2013 doi:10.1017/S0007087413000435

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The conquest of vitalism or the eclipseof organicism? The 1930s Cambridgeorganizer project and the social networkof mid-twentieth-century biology

ERIK PETERSON*

Abstract. In the 1930s, two concepts excited the European biological community: theorganizer phenomenon and organicism. This essay examines the history of and connectionbetween these two phenomena in order to address the conventional ‘rise-and-fall’ narrative thathistorians have assigned to each. Scholars promoted the ‘rise-and-fall’ narrative in connectionwith a broader account of the devitalizing of biology through the twentieth century. I arguethat while limited evidence exists for the ‘fall of the organizer concept’ by the 1950s, theorganicism that often motivated the organizer work had no concomitant fall – even duringthe mid-century heyday of molecular biology. My argument is based on an examination ofshifting social networks of life scientists from the 1920s to the 1970s, many of whom attendedor 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 asimportant as macro-scale conceptual factors in determining the relative persuasiveness oforganicist philosophy.

For it is a fact that the history of science does raise passions, and the basic perspective adoptedtoward the status of scientific worlds does condition our poetry and our politics.

Donna Haraway, Crystals, Fabrics, and Fields1

Old wounds, reopened

In March of 1965, June Goodfield-Toulmin wrote to Joseph Needham at CambridgeUniversity and C.H. Waddington at the University of Edinburgh inquiring about the‘growth and downfall’ of the ‘organiser concept’ in the 1930s. She was initiating researchfor a ‘history of experimental embryology from about 1920 up to 1950’, and this rise-and-fall narrative would feature as its central theme. Though their interactions up to thispoint had been cordial,2 Joseph Needham responded to her critically. ‘[You] will

* History Department at the University of Alabama, 208 Ten Hoor Hall, Tuscaloosa, AL 35487-0212,USA. Email: elpeterson@as.ua.edu. 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 conscientiouscomments.1 Donna Haraway, 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.

BJHS, Page 1 of 24. © British Society for the History of Science 2013doi:10.1017/S0007087413000435

undoubtedly fail’, he insisted, ‘unless you can achieve a more open mind than is to beinferred from the phrase [growth and downfall]’. He saw the ‘drama’ she hoped tonarrate in quite a different light: ‘the perennial struggle of conceptual opposites; herestimulus vs. reactivity, as perennial as the continuity vs. discontinuity antithesis inphysics’.3

Goodfield-Toulmin’s other letter recipient, Waddington, reacted more strongly:

There never was any downfall of the concept of the organiser . . . The only thing that ever felldown was a straw man erected by some American biologists for the express purpose of pushingit over. They performed this simple task so much to their own satisfaction that they had somesuccess in persuading American biologists in general that the problem of embryonic inductionis not important . . .4

Rather than simply being a stylistic division between genetics in Europe and genetics inthe United States, Waddington viewed this division between genetics and embryology asa fundamental blind spot in the whole of American molecular biology:

It is because the essential connection between [genetics and embryology] . . . has beenobscured for the last quarter of a century, that molecular biologists are now saying thatthey will have to move in and solve the problems of differentiation from scratch, as thoughnothing of value had yet been discovered. And of course when they do turn their attentionto the subject, they light immediately on . . . the phenomena of embryonic induction, i.e., ‘theorganiser’.

What was at stake in this private disagreement? At first glance, Goodfield-Toulminsought merely first-hand insight into the ‘organizer project’: a series of embryologicaland biochemical experiments conducted at Cambridge by Needham and Waddington,among others, from 1933 to 1938. They derived their work on the phenomenon ofembryonic induction from the Nobel Prize-winning experiments of Hans Spemann andHilde Mangold in 1935.5 Though a vigorous science in its day, few scientists continuedinvestigating the organizer phenomenon in the decades immediately after the SecondWorld War.6 By the time Goodfield-Toulmin wrote to Needham and Waddington,

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 Hans Spemann and Hilde Mangold, ‘Induction of embryonic primordia by implantation of

organisers from a different species’ (1924) (tr. Viktor Hamburger), International Journal of DevelopmentalBiology (1965) 45, pp. 13–38. See also Timothy J. Horder, ‘The organizer concept and modernembryology: Anglo-American perspectives’, International Journal of Developmental Biology (2001) 45,pp. 97–132.6 E.M. De Robertis, ‘Spemann’s organiser and self-regulation in amphibian embryos’, Nature Reviews

Molecular Cell Biology (2006) 7, pp. 296–302, 297, argues that works by L.G. Barth (‘Neural differentiationwithout organizer’, Journal of Experimental Zoology (1941) 87, pp. 371–383) and Johannes Holtfreter(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 finaltwo on the organizer before the 1990s. However, the work of Richard M. Eakin (‘The nature of the organiser’,Science (1949) 109, pp. 195–197) and A. Ficq (‘Analyse de l’induction neurale par autoradiographie’,Experientia (1954) 10, pp. 20–21) provide clear indications that substantial work on the chemical nature of theorganizer continued well after the Second World War, contra Goodfield-Toulmin’s suppositions.

2 Erik Peterson

many historians and philosophers of science regarded Spemann’s organizer experimentsas vitalism.7 Goodfield-Toulmin intended her history of the organizer to reveal the nega-tive impact that non-epistemic values such as vitalism have upon scientific research.8 ForWaddington and Needham, Goodfield-Toulmin was perpetuating a divide betweencontemporary molecular biology, which she and others believed had unified the lifesciences, and vitalism, which she viewed as faulty metaphysics.9 But the ‘rise-and-fall’narrative troubled Waddington. The misreading of this event was symptomatic of alarger problem: ‘the extremely unfortunate divorce that has grown up between geneticsand embryology in the States’.10

How should the organizer phenomenon be interpreted? Was it the last bastionof vitalism finally toppled by molecular biology? Or were Needham and Waddingtoncorrect: the organizer experiments were being wrongly, if not deliberately, misinter-preted to the ultimate detriment of biology? Both sides of this argument have merit. Theorganizer experiments did cease to be carried out in the UK after the late 1930s. ButGoodfield-Toulmin and others did inappropriately recast the organizer experimentsin order to fit a longer historical narrative. This narrative, as Waddington hinted, wasnot merely a recounting of experiments. Instead the story was about elevating the tropeof progress in biology.11 Needham and Waddington suspected that authors of thenarrative of the ‘rise and fall of the organiser’ conflated organicism with discreditedvitalism, then asserted that molecular biology progressed biology by vanquishing thisfalse ‘organicism’.

A decade ago, Gilbert and Sarkar reclaimed organicism and persuasively argued thatorganicism was the right approach for twenty-first-century ‘evo-devo’.12 Organicismhad been discarded, they asserted, because it kept ‘bad company’ with fascism,communism and marginal New Age groups.13 This article tacitly supported the idea that

7 Hans Driesch was the best-known ‘vitalist’ in the twentieth century. Garland Allen, ‘Mechanism, vitalismand organicism in late nineteenth and twentieth-century biology: the importance of historical context’, Studiesin History and Philosophy of Science Part C: Studies in History and Philosophy of Biological and BiomedicalSciences (2005) 36, pp. 261–283. However, De Robertis, op. cit. (6), p. 299, recollects the common knowledgefrom that era that ‘Spemann’s organizer set developmental biology back by 50 years’, due to his vitalism. Seealso Anne Harrington, Reenchanted Science: Holism in German Culture fromWilhelm II to Hitler, Princeton:Princeton University Press, 1996.8 Non-epistemic values were presumed to motivate the ‘classic’ case of Trofim D. Lysenko’s agronomy (for

example Julian Huxley,Heredity, East andWest: Lysenko andWorld Science, NewYork: Schuman, 1949; andE.N. Megay, ‘Lysenkoism and the stateless society’, Journal of Politics (1953) 15, pp. 211–230).9 Stephen Toulmin and June Goodfield-Toulmin, 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: A.S. Elkus, ‘Mechanism and vitalism’, Journal of Philosophy,Psychology and Scientific Methods (1911) 8, pp. 355–358; and Ralph S. Lillie, ‘The philosophy of biology:vitalism versus mechanism’, Science, new series (1914) 40, pp. 840–846.12 Benson highlighted the ‘organism’ concept still earlier. Keith R. Benson, ‘Biology’s “phoenix”: historical

perspectives on the importance of the organism’, American Zoologist (1989) 29, pp. 1067–1074.13 Scott F. Gilbert and Sahotra Sarkar, ‘Embracing complexity: organicism for the twenty-first century’,

Developmental Dynamics (2000) 219, pp. 1–9, 4–5.

The conquest of vitalism or the eclipse of organicism? 3

organicism was an obscure view held by a few minor scholars decades ago. I argue,instead, that historians have underestimated the pervasiveness of the philosophy and thescope of the organicist social network. My research suggests that, in fact, a substantialnumber of influential scholars in multiple biological subfields held organicist beliefsthrough the majority of the twentieth century. From this perspective, we should vieworganicism as a robust philosophy of biology in triadic tension with two others –vitalism and reductionistic mechanism.In this effort to revise the conventional narrative that organicism ‘fell’ to be

rehabilitated in the twenty-first century, I focus on the formation, dissolution andattempted reinstantiation of organicist social networks – including most prominently theCambridge Theoretical Biology Club. Perhaps the organizer project ‘fell’. Nevertheless,the reasons for its fall and its connection to the philosophy of organicism are neitherstraightforward nor strictly conceptual. Departmental politics, budgets and personalpreferences may have sealed the fate of Cambridge’s organizer project, but theorganicism espoused by Needham and Waddington did not follow the same path asthat of the organizer project. Based on previously unexplored archival materials, thisessay brings to the fore the importance of collections of interacting individuals who heldsimilar views on the philosophy of the organism and a commitment to apply thoseholistic views to the life sciences and beyond.14

14 My focus on network is partly to do with the nature of the relationships between the main actors in thisstory –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, namelyPnina Abir-Am, ‘The discourse of physical power and biological knowledge in the 1930s: a reappraisal of theRockefeller Foundation’s “policy” in molecular biology’, Social Studies of Science (1982) 12, pp. 341–382;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 deMontréal, Montreal,Canada, 1983; Abir-Am, ‘The Biotheoretical Gathering, trans-disciplinary authority and the incipientlegitimation of molecular biology in the 1930s: new perspective on the historical sociology of science’,History of Science (1987) 25, pp. 1–70; Abir-Am, ‘The philosophical background of Joseph Needham’s workin chemical embryology’, in Scott F. Gilbert (ed.), A Conceptual History of Modern Embryology, Baltimore:Johns Hopkins University Press, 1994, pp. 159–180; Scott F. Gilbert, ‘Epigenetic landscaping: Waddington’suse of cell fate bifurcation diagrams’, Biology and Philosophy (1991) 6, pp. 135–154; Gilbert, ‘Induction andthe 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 evolutionarydevelopmental biology’, American Zoologist (2000) 40, pp. 729–737; Brian K. Hall, ‘Waddington’s legacy indevelopment and evolution’, American Zoologist (1992) 32, pp. 113–122; Hall, ‘In search of evolutionarydevelopmental mechanisms: the 30-year gap between 1944 and 1974’, Journal of Experimental Zoology:Molecular and Developmental Evolution (2004) 302B, pp. 5–18; V. Betty Smocovitis, ‘Serious matters: onWoodger, positivism, and the evolutionary synthesis’, Biology and Philosophy (2000) 15, pp. 553–558; andEdward Yoxen, ‘Form and strategy in biology: reflections on the career of C.H. Waddington’, in TimothyJ. Horder, J.A. Witkowski and C.C. Wylie (eds.), A History of Embryology, Cambridge: Cambridge UniversityPress, 1986. My focus on social networks takes cues from John Dupré, The Disorder of Things: MetaphysicalFoundations of the Disunity of Science, Cambridge, MA: Harvard University Press, 1993; Larry Laudan,Beyond Positivism and Relativism: Theory, Method, and Evidence, Boulder, CO: Westview Press, 1996; andBruno Latour, Reassembling the Social: An Introduction to Actor-Network Theory, Clarendon Lectures inManagement Studies, Oxford and New York: Oxford University Press, 2005.

4 Erik Peterson

What was/is twentieth-century organicism?

As mentioned above, Needham andWaddington held to a philosophy called organicism,a concept that pre-dated any work on the organizer problem.15 A decade or more beforeNeedham and Waddington took up organicism, John Scott Haldane and EdwardStewart Russell pronounced against the reductionistic mechanism that they saw in thelife sciences of their day. Haldane hoped to retain the common-sense appeal of vitalismwhile simultaneously insisting upon testable entities and forces attributed to mechanisticbiology.16 He thought this ‘organicism’ provided a way to avoid the dualistic vitalism ofHans Driesch, but without conceding that the organic world was nothing but physico-chemical mechanism.17 According to his version of organicism, traits of live organisms –their ability to self-regulate within certain constraints, for instance – emerged from prop-erties shared by non-living materials to create something entirely new; this did not mean,however, that there were two different kinds of substance, living and non-living.Haldane’s organicism was both monistic (that is to say, there is but one quality of ma-terial substance in the world) and emergent. By emergent, Haldane referred to the pecu-liar ‘popping up’ of structures or behaviours or powers unique to organized systems, notexclusively reducible to the physico-chemical states from which they are constructed. Hiscontemporary, Glaswegian ichthyologist Russell, found it more difficult to downplay theapparent distinction between living and non-living.18 Like Haldane, Russell adopted the‘simple everyday conception’ that organisms ‘are active, purposeful agents’ while avoid-ing the ‘vitalism’ of Lamarck or Samuel Butler.19 For Russell, organicism was possiblebecause organisms possessed an ‘essential unity’, the individuated parts of which ‘cannever fully explain the activity of the whole; they are to be regarded rather as conditions,both limiting and implementing that activity’.20 The structure of organisms as wholes, inother words, seemed to constrain, and therefore precede, the function of parts.

Aside from Haldane and Russell, organicism drew adherents from a plurality ofbackgrounds. American marine biologist William Emerson Ritter composed a two-volume tome defending ‘the facts . . . of organic unity and integration’ before going on tolead the United States Science Service.21 Prime minister of South Africa Jan C. Smuts

15 Phillipe Huneman and Charles T. Wolfe, ‘The concept of organism: historical, philosophical, scientificperspectives’, History and Philosophy of the Life Sciences (2010) 32, pp. 147–154.16 John Scott Haldane, 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’. E.S. Russell, ‘The “drive” elementin life’, British Journal for the Philosophy of Science (1950) 1, pp. 108–116, 108.19 E.S. Russell, Form and Function: A Contribution to the History of Animal Morphology, London:

J. Murray, 1916, pp. v–vi.20 E.S. Russell, The Interpretation of Development and Heredity: A Study in Biological Method, Oxford:

Clarendon Press, 1930, pp. 306–307.21 William E. Ritter, 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’.

The conquest of vitalism or the eclipse of organicism? 5

criticized both the notion that life and mind were substances distinct from the materialand the reduction of ‘life and mind to a subsidiary and subordinate position as mereepiphenomena’.22 Harvard University biochemist Lawrence J. Henderson spent fiveyears sketching his alternative to both mechanism and vitalism.23 Still others insistedthat reductionistic mechanism no longer sufficed to explain the overwhelming complex-ity of the vast majority of phenomena in the living world.24 Just as J.S. Haldane claimedthat organisms exhibited properties belonging to a seemingly different class than thematerials from which they emerged, Cambridge philosopher of mind C.D. Broad spelledout a version of mental emergence that he contrasted to ‘Substantial Vitalism’.25

Alfred North Whitehead drew these organicist threads together into a morecomprehensive world view. In his Science and the Modern World, Whitehead criticizedthe tendency of scientists, as well as the general public, to view modernity through thelenses of materialism, mechanism and reductionism. Paradoxically, modern physics didnot depict that sort of universe.26 Whitehead suggested that other modern sciencesfollow the trend in physics, elevating the prominence of networks – he often used theterm nexus – over entities. Process, form, development, exchange, reciprocity andinteraction across integrated systems should displace descriptions of the behaviour ofindividuals by the functions and compositions of parts. In other words, Whiteheadcalled for a science focused on multi-perspectival networks of relationships subjectivelyanalysable – though not objectively definable – as concrete ‘events’ or ‘chreods’ ratherthan as the behaviour of aggregated atomic units.27 This was the core of organicism.By the mid-1920s, organicism could not be regarded as an idiosyncratic idea confined

to the fringes, but neither was it unified into a movement. Even at Cambridge, so deeplyimpacted by the work of Whitehead, D’Arcy Thompson and others, no ‘school’ oforganicism existed. Nevertheless, as we will see below, it was at Cambridge that a cadreof philosophically sophisticated scientists inspired by these intellectual ancestors beganto gather into a tighter network during the 1930s to clarify this organicist ‘third way’between dualistic vitalism and reductionistic mechanism.28 Spemann’s organizer becameanother catalyst in the formation of that social network.

22 J.C. Smuts, Holism and Evolution, New York: The Macmillan Company, 1926, p. 8.23 JeanMayer, ‘L.J. Henderson: a biographical sketch’, Journal of Nutrition (1968) 94, pp. 1–5; and Steven

J. Cross andWilliam R. Albury, ‘Walter B. Cannon, L.J. Henderson, and the organic analogy’,Osiris (1987) 3,pp. 165–192.24 E.g. Erik Nordenskiöld, The History of Biology: A Survey (tr. L.B. Eyre), New York and London:

A.A. Knopf, 1928, pp. 525–527; and D’Arcy W. Thompson, On Growth and Form, 2nd edn (ed. LancelotL. 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 differenceof behavior wholly in terms of difference of structure’.26 See Klaus Meyer-Abich, ‘Bohr’s complementarity and Goldstein’s holism in reflective pragmatism’,Mind

and Matter (2004) 2, pp. 91–103.27 Alfred North Whitehead, Science and the Modern World, Lowell Lectures, 1925, New York: Free Press,

1953, pp. 138–157.28 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

6 Erik Peterson

Metaphysics and the ‘organizer problem’ (1924)

How do the organizer experiments fit into the history of organicism? A main problemwith concepts like organicism has less to do with their content than with their use. Howdoes a metaphysical commitment – vitalism, organicism or reductionist mechanism, forthat matter – actually impact scientific work? Can it only inhibit research progress?29

Hans Spemann believed that his metaphysics had led him to investigate and interpret theorganizer phenomenon that led to a Nobel Prize.

In 1924, Spemann and his assistant, Hilde Mangold, transplanted a small group ofcells from the upper blastopore lip of one very young salamander embryo, Tritoncristatus, into another, Triton taeniatus. Usually, transplants of this sort and with em-bryos of this young age resulted in the conformity of the new tissue to the new surround-ing area. But in this case, transplanted blastopore lip would not conform. Instead, thetiny lip transplant transformed the host tissue. The area around the transplant foldedinto a neural plate – a developing nervous system. In the later experiments, Spemannreported that the region ‘bulge[d] out into optical vesicles and add[ed] lenses andauditory vesicles’.30 These experiments convinced Spemann that the blastopore lipcontained a cell area, potentially a hormone-producing complex, that ‘organized’ theembryonic tissue of any region into a new brain stem.31 Spemann refused to interpretthe phenomenon by referencing merely the physico-chemical constituents. In his mind,the organizer was a whole-organism problem.32 Unfortunately, biologists and historianstended to interpret Spemann’s holism as evidence of vitalism, despite the fact thatSpemann rejected Driesch’s philosophy.33

Questions over Spemann’s philosophical proclivities notwithstanding, the organizergrew to be a fashionable international pursuit bridging biochemistry and embryology inthe pre-Second World War period.34 Excited researchers expected that the biochemicalidentification of the organizer would be the key to unlocking the most basic problem of

scheme that sees biology as a war between vitalists and mechanists’. Stephen J. Gould, ‘Just in the middle’,inGould, The Flamingo’s Smile: Reflections in Natural History, New York: W.W. Norton & Company, 1987,pp. 377–391, 388.29 A.G. Greenwald, A.R. Pratkanis, M.R. Leippe and M.H. Baumgardner, ‘Under what conditions does

theory obstruct research progress?’, Psychological Review (1986) 93, pp. 216–229.30 Hans Spemann, ‘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, http://nobelprize.org/nobel_prizes/medicine/laureates/1935/spemann-lecture.html, accessed 28 August2009.31 Spemann and Mangold, op. cit. (5).32 Viktor Hamburger, ‘Hans Spemann on vitalism in biology: translation of a portion of Spemann’s

autobiography’, Journal of the History of Biology (1999) 32, pp. 231–243, 233.33 Timothy J. Horder and Paul Weindling, ‘Hans Spemann and the organizer’, in T.J. Horder,

J.A. Witkowski and C.C. Wylie (eds.), A History of Embryology, Cambridge: Cambridge University Press,1986, pp. 183–242, 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 theRockefeller Foundation. Spemann and Ross Harrison set up the micro-instrumentation. Terni then travelled tothe labs of Spemann and Mangold to learn proper techniques. JN/CUL, E.111.

The conquest of vitalism or the eclipse of organicism? 7

development, morphological patternedness.35 A small group of multidisciplinaryscholars at Cambridge were numbered among the pursuers of the organizer. Unliketheir contemporaries, however, the Cambridge group insisted that a search for a singlebiochemical entity would not suffice.

Building the organicist base of the Cambridge organizer project (1929–1931):Needham, Woodger and Von Bertalanffy

The Cambridge organizer project had its gestation outside any laboratory. Con-versations in the late 1920s between Joseph Needham and J.H. Woodger in England andWoodger and Ludwig von Bertalanffy in Austria helped to refine organicism and thesocial network that would promote it.From 1920 to 1942 Needham used his influential position in the laboratory of

Frederick Gowland ‘Hoppy’ Hopkins to transgress the boundaries between biochem-istry and embryology. Initially, he fixated on seemingly insoluble problems relating tothe development of chicks in their eggs, eventually writing the interdisciplinary magnumopus of his young career, the three-volume Chemical Embryology. Spemann’s organizerphenomenon appeared to be the next challenge.36 Needham’s interactions with JosephH. Woodger were among the factors contributing to his interest in the organizer project.At Woodger’s death, even Karl Popper called him ‘Socrates’.37 Over a career nearly

seven decades long, Woodger had cast a quiet shadow over the philosophy of biology inthe Anglo-American world. Members of the Vienna circle met him in 1926; philosophersall over the world, including Alfred Tarski, Rudolf Carnap and W.V.O. Quine, thoughthim a friend and ally. However, later scholars tended to overlookWoodger’s importanceas an isolated example of overly analytic times, not offering much for either the benchscientist or the philosopher.38

Though they had corresponded before, the first major public exchange betweenWoodger and Needham centered on Needham’s review of Woodger’s Biological

35 Garland Allen, ‘A pact with the embryo: Viktor Hamburger, holistic and mechanistic philosophy in thedevelopment of neuroembryology, 1927–1955’, Journal of the History of Biology (2004) 37, pp. 421–475; andJ.A. Witkowski, ‘Optimistic analysis: chemical embryology in Cambridge 1920–42’, Medical History (1987)31, pp. 247–268.36 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 Joseph Needham, Mikulás Teich and RobertM. Young (eds.), Changing Perspectives in the History of Science: Essays in Honour of Joseph Needham,London: Heinemann Educational, 1973, pp. 1–20, 7; and Simon Winchester, The Man Who Loved China,New York: Harper Perennial, 2008, p. 28.37 Karl R. Popper, ‘Obituary: Joseph HenryWoodger’, British Journal for the Philosophy of Science (1981)

32, pp. 328–330.38 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 regardingTarski and neither Woodger nor the Theoretical Biology Club meeting that Popper attended appear in Popper’sautobiography. See Karl R. Popper, Unended Quest, New York: Routledge, 2002; and Malachi H. Hacohen,Karl Popper, the Formative Years, 1902–1945: Politics and Philosophy in Interwar Vienna, Cambridge:Cambridge University Press, 2000, p. 311 n. 68.

8 Erik Peterson

Principles in Mind. Needham reviewed Biological Principles positively. Still Woodgerbelieved he had completely missed the point. Needham, like J.S. Haldane, stressed the in-principle reconcilability – he used ‘harmony’ – of mechanism and vitalism, as long aseach remained consigned to its right place.39 Woodger, by contrast, bluntly denied thatany via media could be constructed between these two camps as they were defined in the1920s. Too many of the obvious problems of biology were ‘absolutely insoluble’, hethought, so long as mechanists and vitalists each argued over a concept of entities. ForWoodger, levels of organization, internal and multiple relations, and the counterentropicchange most easily observed during the process of development were the kinds oforganic factor that rendered biology epistemologically, if not qualitatively, parallel(rather than reducible) to the physico-chemical.40 Woodger (following Whitehead)believed that the quantum revolution in physics had emancipated biology. Like physi-cists, biologists no longer had to speak in terms of determined entities using merelyphysico-chemical language. Biology could be autonomous.

Despite their brief misunderstanding, Woodger and Needham developed a solidfriendship, continuing to meet regularly through the early 1930s.41 Needham did not‘grudge the effort needed for a precise analysis of our logical procedure, and for theaccurate expression of our results’ that Woodger offered; Woodger recognized that heneeded the backing of an experimentalist to win him a broader hearing in philosophicaland scientific circles.42 Needham was one such source and Woodger gladly included himin the discussions defining organicism that he had already initiated with Austriansystems biologist Ludwig von Bertalanffy.43

Woodger connected with von Bertalanffy through their mutual acquaintances withthe Viennese circle of logical empiricists. Von Bertalanffy would not introduce hisgeneral systems theory, for which he became widely known, until after the SecondWorldWar.44 But Woodger had already convinced him to join in on the ‘organicist’ project bythe late 1920s. He translated and elaborated von Bertalanffy’s Kritische Theorie derFormbildung; the resultingModern Theories of Development became a cornerstone text

39 Joseph Needham, ‘Review of J.H. Woodger, Biological Principles, a Critical Study’, Mind (1930) 39,pp. 221–226.40 J.H. Woodger, Biological Principles: A Critical Study, London: K. Paul, Trench, Trubner & Co., 1929,

pp. 482–487. While serving in the First World War, Woodger befriended Ian Suttie, a psychotherapist whoseholistic concept of the developing self deeply impacted Woodger’s philosophy. W.F. Floyd and F.T.C. Harris,‘Joseph Henri [sic] Woodger, Curriculum Vitae’, in J.R. Gregg and F.T.C. Harris (eds.), Form and Strategy inScience: Studies Dedicated to Joseph Henry Woodger on the Occasion of His Seventieth Birthday, Dordrecht:Reidel, 1964, pp. 1–6; see also the correspondence between Woodger and Suttie, 1919 to 1925, Woodger’sPapers, University College London (hereafter JHW/UCL), C1/3/Suttie.41 Abir-Am, ‘The philosophical background of Joseph Needham’s work’, op. cit. (13).42 J.H. Woodger, ‘The “concept of organism” and the relation between embryology and genetics, part III’,

Quarterly Review of Biology (1931) 6, pp. 178–207, 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 Ludwig von Bertalanffy, ‘An outline of General Systems Theory’, British Journal for the Philosophy of

Science (1950) 1, pp. 134–165; and von Bertalanffy, ‘The theory of Open Systems in physics and biology’,Science (1950) 111, pp. 23–29.

The conquest of vitalism or the eclipse of organicism? 9

in theoretical embryology.45 As with other organicists, von Bertalanffy was convincedthat neither reductionist mechanism nor dualistic vitalism could serve as the truefoundation of biology.46 Significantly, von Bertalanffy believed that as a bridge betweencommon-sense vitalism and reductionisic mechanism, organicism served the purpose oftheoretical unification that the other philosophies had also claimed. Biology, likechemistry and physics, required a ‘methodological clarification’ as ‘protection againstthe fallacies of hurried hypotheses’.47 This new theoretical biology should beginembryology because theoretical rigour in embryology would align future progress in therest of experimental biology and remedy the inefficient infighting of reductionisticmechanists and dualistic vitalists.48

From the point of view of von Bertalanffy, Austro-German gestalt theorists, and evenSpemann, a metaphysical monism could in principle connect levels from subatomicphysics to psychology.49 Here Woodger differed from the Germans. He believed that‘flattening out’ would not work. Instead an important middle-range theory needed to beinserted and explicated: a structural connection between physico-chemical non-livingobjects and biological ones. He sketched out his approach in a series of dense articlespublished in the Quarterly Review of Biology over 1930–1931.50 Here, he criticizedthe epistemological dead end of Driesch’s vitalism, but he also directed gentle jabsat ‘neo-mechanism’ – the theory endorsed by Needham at that time51 – and, notably, J.S.Haldane’s ‘concept of organism’. Because of the vagueness of definitions such asHaldane’s, organicism had not necessarily ‘been tried and found wanting’, but had ‘beenfound difficult and not tried’.52 Woodger’s goals, then, were dual: (1) to clarify the messylinguistic conventions of biology, and (2) to convince working biologists that ‘new waysof thinking’more related to hypothetico-deductive algorithms than to ‘intuitions’ shouldbe pursued if biology was going to be put on firmer (that is to say, non-Cartesian ordualist) footing.53 Capturing the ethos of organicism, Woodger wanted to render or-ganismic biology philosophically respectable while acknowledging the intuitions ofvitalists and mechanists alike.Two events occurred in 1931 that decisively changed the history of this Needham–

Woodger–von Bertalanffy social network and, consequently, organicism. The first is that

45 Ludwig von Bertalanffy and J.H. Woodger, Modern Theories of Development: An Introduction toTheoretical 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, Frans Heske, Pascual Jordan and Adolf Meyer-

Abich (eds.), Organik: Beiträge zur Kulture unserer Zeit, Berlin: Fritz Haller, 1954.50 J.H. Woodger, ‘The “concept of organism” and the relation between embryology and genetics, part I’,

Quarterly Review of Biology (1930) 5, pp. 1–22; Woodger, ‘The “concept of organism” and the relationbetween embryology and genetics, part II’,Quarterly Review of Biology (1930) 5, pp. 438–463; andWoodger,op. cit. (42).51 Joseph Needham, The Sceptical Biologist, New York: W.W. Norton, 1930, pp. 85–86.52 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.

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Woodger and Needham attended the Second International Congress of the History ofScience and Technology held in London that year. There, popular Bolshevik revo-lutionary and philosopher of science Nikolai I. Bukharin delivered his address, ‘Theoryand practice from the standpoint of dialectical materialism’. Bukharin’s perspectivegreatly impressed Needham.54 By the end of the conference Needham had adopted amore radical posture in line with Bukharin’s exposition: he was ready to discardEurocentrism, the collusion of science with oppressive capitalism, and the dyadic op-position of vitalism and mechanism at the heart of Western biology.55 In the seconddecisive event of 1931, Needham connected Woodger with another Cambridge scientistglad to explore theoretical biology –Conrad Hal ‘Wad’Waddington.56 Waddington hadrecently transformed himself from palaeontologist to population geneticist to embryolo-gist.57 Soon C.H. Waddington would become the main promulgator of organicism (or‘Woodgery’, as he liked to call it).58

The organicist network evolves (1931–1938): the influence of the original TheoreticalBiology Club on the Cambridge organizer project

Waddington shared many interests with Needham: he had been profoundly impacted bythe philosophy of Alfred North Whitehead, leaned left in his politics, and straddleddisciplinary lines throughout his career.59 Like Needham, Waddington was a dedicatedmorris dancer and shared non-traditional attitudes regarding marital relationships.60

With so much in common, Needham and Waddington jumped at the chance to worktogether at Strangeways Laboratory in Cambridge. Their friendship resulted in a deep-ening of the organicist social network, established Cambridge as a leading centre forexperimental embryology until 1938, and would strongly influence the remainder ofWaddington’s career as he explored the intersection of development and evolution.

Waddington came to Strangeways as a novice in embryology in 1929 following abrief period of unfulfilling work in population genetics with Edith Saunders andJ.B.S. Haldane.61 At the same time, from 1929 to 1931, Waddington had obtained a

54 Maurice Goldsmith, ‘Joseph Needham, honorary taoist’, in J. Needham, S. Kumar Mukherjee andA. Ghosh (eds.), The Life and Works of Joseph Needham, Calcutta: The Asiatic Society, 1997, pp. 1–3.55 Holorenshaw, op. cit. (36), p. 10; and Joseph Needham, 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 ultimatelyhampered the acceptance of organicism. The cases of both J.B.S. Haldane and H.J. Muller would seem to speakagainst 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; andWoodger 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 C.H. Waddington, ‘Pollen germination in stocks and the possibility of applying lethal factor hypothesis

to the interpretation of their breeding’, Journal of Genetics (1929) 21, pp. 193–206, was written under thedirection of Edith Saunders, a long-time collaborator with William Bateson at the John Innes HorticulturalInstitute. About his collaboration with J.B.S. Haldane (C.H. Waddington and J.B.S. Haldane, ‘Inbreeding and

The conquest of vitalism or the eclipse of organicism? 11

scholarship in philosophy and had made his own excursion into the vitalism–mechanismdebate.62 He read Spemann, and from that reading decided that he would attempt thetissue transfer techniques on much more difficult avian embryos. Surprisingly, he wassuccessful enough to receive an invitation from Richard Goldschmidt to continue workin Germany. After receiving a Rockefeller Medical Fellowship in 1931, Waddingtonjoined Otto Mangold’s Berlin-Dahlem lab.63 Later, Joseph Needham and his wife,Dorothy, joined Waddington to investigate the physico-chemical process of amphibiandevelopment. This partnership produced ten co-authored papers on amphibiandevelopment from 1931 to 1937.64

In retrospect, it seems almost inevitable that this small cadre would develop a moretightly knit relationship. They soon brought leftist crystallographer J.D. Bernal andhis associate Dorothy Crowfoot Hodgkin into their social network.65 In April 1932,Needham suggested to Woodger that they have a small excursion to discuss theoreticalissues. Woodger spoke with Waddington, who enthusiastically agreed to attend.Woodger volunteered his family’s summer cabin in Epsom Down and suggested thattheir informal ‘biotheoretical gathering’ expand to include E.S. Russell, among others(Russell did not attend).66 Needham would later give the ‘Conference’ a more formid-able moniker from the Cambridge lexicon: the ‘Theoretical Biology Club’ (hereafterTBC).67 Together, this collection of young scientists and philosophers inspired

linkage’, Genetics (1931) 16, pp. 357–374), Waddington could only say that he had ‘tasted the thin gruel ofmathematical formalism’. C.H. Waddington, The Evolution of an Evolutionist, Edinburgh: EdinburghUniversity 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 1851Scholarship. 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 Alan Robertson, ‘Conrad Hal Waddington.

8 November 1905–26 September 1975’, Biographical Memoirs of Fellows of the Royal Society (1977) 23,pp. 575–622, 615–621. During the same six-year period, Joseph Needham and Waddington produced asurprising number of papers and books, including Joseph Needham, 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 C.H.Waddington,HowAnimals 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 unstableacademic 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 occasionallyalternated 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’sOrder and Life (1936). Woodger used ‘biotheoretical gathering’ (uncapitalized)in his 30 April 1932 ‘start-up’ letter to Needham (JN/CULM.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 toNeedham, 23 April 1936, JN/CUL M.99). Given that Needham’s term made it to print –whereas Woodger’sterm was informal and only appeared inconsistently and in private letters – I will continue to use ‘TheoreticalBiology Club’/‘TBC’.

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interdisciplinary work in the Oxbridge–London academic triangle and refined morecarefully both the concept and the application of organicism.68

Their meetings covered a gamut of issues in theoretical biology. At the first meetingin August 1932, Bernal drew from his training in crystallography to suggest that amorphological scale of bodily form could be drawn that included everything fromthe level of quantum structures to metazoa.69 Mathematician Dorothy Wrinch spokeof ‘geometrical botany’ and the mathematical complexity of morphological forms.Waddington presented short papers combining Drosophila genetics and embryology;Needham’s meeting notes record that even in 1932 Waddington was thinking abouthomeotic mutations in Drosophila – the basis for his important aristapedia experimentsyears later.70 A preoccupation with topology and geometrical patterning as a non-atomistic explanatory framework set the tone for the organicist ethos of the early TBC.But discussions ranged well beyond the confines of embryology, mathematics or crys-tallography.

This feature, their extra-academic common interests, distinguished this small groupfrom other contemporaries interested in theoretical biology. The TBC not only stageddetailed discussions of biological philosophy and practice, it established bonds offriendship extending beyond their careers or institutional affiliations. Fuelled by talk ofWhitehead, D’Arcy Thompson, Friedrich Engels and others, they discussed the scien-tific ordering of the state, Russian communism versus British Christian socialism,and – drawing from Waddington’s background in palaeontology – evolution.

TBC members were interested in the organizer phenomenon in particular. And inthe summer of 1933 the Needhams travelled with Waddington to Otto Mangold’sBerlin-Dahlem laboratory, where they would collaborate on the organizer. There theydemonstrated for the first time that the phenomenon –Waddington by then had labelledit ‘evocation’ – could be produced by non-living tissue. This finding convinced Needhamand Waddington that Spemann’s explanations of the phenomenon were not quiteright.71 When they returned to Britain, they agreed to invite participants to the TBCfrom outside the initial group to help address what had become a far more difficultscientific and philosophical problem: how could non-living tissue instigate embryonicdevelopment?

One of those invited was Viennese philosopher Karl Popper, who attended the May1935 meeting held at the Woodgers’ cabin.72 He returned a year later to meet with the

68 Jonathan M.W. Slack, ‘Conrad Hal Waddington: the last Renaissance biologist?’, Nature Reviews,Genetics (2002) 3, pp. 889–895.69 Here Bernal drew from Thompson, op. cit. (24). See Robert C. Olby, ‘Structural and dynamical

explanations in the world of neglected dimensions’, in T.J. Horder, J.A. Witkowski and C.C. Wylie (eds.),A History of Embryology, Cambridge: Cambridge University Press, 1985, pp. 275–308, 281.70 Gregory K. Davis, Michael R. Dietrich and David K. Jacobs, ‘Homeotic mutants and the assimilation of

developmental genetics into the evolutionary synthesis, 1915–1952’, in J. Cain and M. Ruse (eds.), Descendedfrom Darwin: Insights into the History of Evolutionary Studies, 1900–1970, Philadelphia: AmericanPhilosophical Society, 2009, pp. 139–140. 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 theNeedham–Waddington critique of Spemann was not his holism but his quasi-vitalism (JN/CUL, E.111).

The conquest of vitalism or the eclipse of organicism? 13

TBC at the ‘old windmill at Hunstanton’.73 The June 1936 meeting was the apex of theTBC – Popper, J.B.S. Haldane, Hyman Levy and others joined Waddington, Needham,Woodger and Bernal. Conversation during meals at the 1936 meeting circulated aroundsocialism, science and state planning. Though Popper disagreed with the politicalleanings of the group, he found organicism appealing. Even many decades later, herefused to support reductive mechanism.74

TBC meetings unquestioningly impacted the organicist science of its core members.75

As a result, they would stand by two claims in particular: (1) ‘the properties of acomponent depend upon the component of which it is a component (relational prop-erties) as well as on the parts into which it is analysable’, and (2) scientists experiment on‘occasions of experience’ (that is to say, temporal, unified, holistic systems that areprocesses rather than entities) by artificially dividing them up to render them potentiallyanalysable.76 The first point is an ontological claim about the integration of organismicstructures; the second an epistemological one about the provisionality of scientists’knowledge. Inspired by discussions at the TBC, Needham and Waddington expectedsystemic rather than atomic explanations for the evocation process, to appreciate aplurality of inputs at different levels inside the organism (for example, genetic, epi-genetic, cytological and so on), and to regard any isolable set of developmental causesas incomplete explanatory factors.77 Not only did this mean that the organizer workof Needham, Waddington and their occasional co-authors did not resemble thecontributions of other international organizer groups, it would also eventually makethe acquisition of funding more difficult: funding agencies believed the organizer to be adiscrete biochemical compound.

The gradual ‘fall’ of the Cambridge organizer project and the TBC (1934–1942)

Early in 1934, however, the Cambridge organizer work seemed exceptionallypromising. Inspired by their cooperative, multidisciplinary attack on a ‘borderline sub-ject’, W.E. Tisdale of the Rockefeller Foundation approached Needham in April andasked him to suggest ways in which the foundation could advance ‘chemical

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 Karl R. Popper, ‘Scientific reduction and the essential incompleteness of all science’, in F.J. Ayala and

T. Dobzhansky (eds.), Studies in the Philosophy of Biology: Reduction and Related Problems, Berkeley:University of California Press, 1974, pp. 259–284.75 Gary Werskey, 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 toWaddington’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.

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embryology’.78 By June, Needham had submitted an extremely ambitious plan for anInstitute of Embryology. When that plan was rejected, Needham sent a hardly moremodest proposal to establish a formal institute devoted to ‘Mathematico-physico-chemical [MPC] Morphology’.79 We do not know the true motivations for supportingNeedham’s plan, nor do we know why it was eventually rejected. There are hints thatthe explication of morphology with reference to physics and chemistry fit theRockefeller’s preference for funding projects interested in the ‘ultimate littleness ofthings’.80 In the end, however, disciplinary interstices represented precariously rockysoil for Rockefeller seed money and dreams of an MPC morphology unit at Cambridgenever materialized.

The survival of the Biochemical Laboratory Extension was further complicatedby problems involving Waddington’s salary and the fracturing of the Strangeways–Cambridge collaboration caused by miscommunications between F.G. Hopkins,Needham and Tisdale. The combination of all three of these quotidian issues did indeedlead to a ‘fall’ of the organizer project at Cambridge and contributed to the break-up ofthe Cambridge TBC.81

Conflict over Waddington’s salary ended up severely, if temporarily, hampering hisscientific career. From 1933, Waddington held a fellowship at Christ’s, a part-timelectureship in zoology, and, on account of his philosophical prowess, the RoyalCommissioners of the Exhibition of 1851 Senior Studentship. In 1936, the studentshipwas cut by two-thirds; after paying his alimony, this left him with a meager £250 perannum. When Needham turned to the Rockefeller Foundation on his behalf seekingemergency salary assistance, the Cambridge Council – spurred by Waddington’scolleague James Gray – took offence.82 After some directed consultation, the Counciland General Board changed their policy to allow for direct oversight, which blockedfuture efforts to secure salary funding for Waddington.83

78 Joseph Needham, ‘An account of the negotiations regarding financial aid for technical assistance, etc., forstudies on the borderline between biochemistry and embryology’, unpublished memo, 12 July 1938, JN/CUL,E.114.79 Pnina G. Abir-Am, ‘The Rockefeller Foundation and the rise of molecular biology’, Endeavour (2001)

25, pp. 55–60.80 According to the Rockefeller Foundation’s 1938 ‘Annual Report’, quoted in Lily E. Kay, The Molecular

Vision of Life: Caltech, the Rockefeller Foundation, and the Rise of the New Biology, New York: OxfordUniversity Press, 1993, p. 49. Weaver coined the term ‘molecular biology’ and began explicitly channellingfunding in this direction around 1938 (Kay, op. cit., pp. 188–189). See also Robert E. Kohler, ‘Themanagement of science: the experience of Warren Weaver and the Rockefeller Foundation programme inmolecular biology’, Minerva (1976) 14, pp. 279–306.81 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 MPCmorphology unit in the history of the ‘Biochemical Laboratory Extension’, she has not addressed these otherfactors 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 awayfrom 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

The conquest of vitalism or the eclipse of organicism? 15

Understandably, haggling over Waddington’s salary led to bad feelings betweenNeedham and Waddington on the one side and Cambridge University officials on theother. By 1938, Tisdale revealed to Needham that even Hopkins’s support for his andWaddington’s organizer work was waning and, with Hopkins’s retirement nearing,Tisdale warned Needham that Hopkins’s successor would likely cut off Needhamentirely.84 Hopkins rejected Tisdale’s suggestion as ‘absurd’ and vowed to support ‘yourwork on the Organiser’.85 Little improved, however. By September 1938, Needhamapplied to the Royal Society for a grant; Hopkins wrote in support.86 In the grant ap-plication, Needham attempted to convey the precariousness of the situation:

If the grant is not obtained: Nothing further on the nature of the primary organiser will be donein Cambridge, though Cambridge has at present the lead in this subject. If it is not done inCambridge, there is no chance of its being done anywhere else in England . . . On the whole, itmay be said that the subject would be held up for 10 to 20 years . . . it seems truly shortsighted,especially in a country where belief in pure science has always been boasted, not to recognisethe value of research into the deepest processes of early embryonic development.87

The Royal Society did fund Needham, but permanent damage had been done.Waddington transitioned most of his research to Strangeways. Cambridge failed tosupport experimental embryology consistently after this point.Of course the relationship with the Rockefeller Foundation also suffered. Needham

and Waddington had been so successful in their pursuit of the organizer in 1933–1934that Weaver felt compelled to invest in the work straddling Hopkins’s biochemistry laband the Strangeways Laboratory through the mid-1930s.88 At one point the Rockefellereven funded laboratory technicians to work with Needham and Waddington atCambridge.89 Yet the succession of failed attempts at identifying the specific biochemicalessence of the organizer was not the solution the Rockefeller Foundation had hoped for,even if it sustained Waddington and Needham’s organicist suppositions.90 By 1938,Waddington reported that even some toxic substances induced the embryo to sym-metrically develop just as any other ‘natural’ compound. To Waddington, such a resultindicated that the ‘evocator . . . appears impossible to discover’ given some of the rulingassumptions that ‘the artificial [i.e. experimental] process gives a true picture of what

Board’. ‘Memorandum on Applications for Grants’, approved by the Cambridge University Council on1March 1937, JN/CUL, B.26; ‘Memorandum on the position of C.H. Waddington’, enclosure with letter fromNeedham 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 atStrangeways Laboratories at a reduced capacity. ‘Dean, Donaldson, Adrian, and Dale’ of Strangeways toNeedham, 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 from1935 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.

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happens in nature’.91 He introduced the speculation he and Needham shared that, ratherthan a biochemical substance that induced change, the evocation principle was distri-buted throughout a field or gradient in the developing embryo but suppressed. Once thesuppressor was released by adding the synthetic chemical compound, the embryo beganthe developmental process. Johannes Holtfreter –who escaped Germany to work withWaddington and Needham in 1939–1940 – later confirmed this experimental result.92

Nevertheless, while it was still an active field of research, the Rockefeller would nolonger support embryology at Cambridge without requisite university support.93

Lacking prospects of permanent financial support doing experimental embryology atCambridge, Needham and Waddington began pursuing other interests.94 Consideringthe troubles the others were having at Cambridge, even Woodger began devoting moretime to meetings with Tarski and ‘a large number of new friends’.95 In 1937, he travelledto Poland; he was at Yale on a Rockefeller grant in 1938, meeting Waddington’s closefriends Gregory Bateson and Margaret Mead, among others.96 After his return, he spentincreasing amounts of time with scholars at the London School of Economics. When hereconvened the TBC again after the war, most of the original members had moved on.97

Waddington travelled to America on a Rockefeller grant to study population genetics.He began the summer of 1938 at the Marine Biological Laboratory at Woods Hole,Massachusetts, visited Ross Harrison at Yale and L.C. Dunn at Columbia, and ended atT.H. Morgan’s Drosophila lab at the California Institute of Technology.98 Over the1938–1939 academic year, he worked on Drosophila development alongside Morgan’scomrades Alfred Sturtevant and Theodosius Dobzhansky.99

91 C.H. Waddington, ‘Studies on the nature of the amphibian organization centre –VII. Evocation by somefurther chemical compounds’, Proceedings of the Royal Society of London. B, Biological Sciences (1934–1990)(1938) 125, pp. 365–372, 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 John Gerhart, ‘JohannesHoltfreter: January 9, 1901–November 13, 1992’, Biographical Memoirs National Academy of Sciences (US)(1998) 73, pp. 209–228.93 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 forembryology, 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 Margaret Mead, 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 publishedGenetics and the Origin of Species,

1st edn, New York: Columbia University Press, 1937. Relations between Dobzhansky and Sturtevant weretense 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.William B. Provine, ‘Origins of the genetics of natural populations series’, in Richard C. Lewontin,William B. Provine, John Moore and Bruce Wallace (eds.), Dobzhansky’s Genetics of Natural Populations:I–XLIII, New York: Columbia University Press, 1982, p. 30; and see Waddington to Dobzhansky, 1 February1974, Dobzhansky Papers, American Philosophical Society, Philadelphia (hereafter TD/APS), Mss.B. D65.

The conquest of vitalism or the eclipse of organicism? 17

By then the Cambridge organizer project was most certainly on its deathbed.Needham remained in biochemistry, but ceased writing grants to support work on theorganizer by 1939.100 But at Cambridge the ‘fall of the organizer’ had little to do with aconcomitant failure of organicism or the death of vitalism. It was a contingent victim oflack of funding by the Rockefeller and disinterest, even active opposition, by othermembers of the faculty – ironically due to Needham’s aggressive pursuit of that funding.While work specifically on the constitution of the organizer stopped at Cambridge, theorganicism motivating the ‘rise of the organizer’ was not dead at all and in fact hadbegun to infiltrate genetics.

Organizers, genes and an evolutionary synthesis (1938–1942)

Waddington’s year of work in the United States reinforced his belief that Americangenetics did not really know what to do with the problems of development dealt with byexperimental embryology or the organicist interests he entertained.101 Before he hadeven completed his first genetics text, Introduction to Modern Genetics, he began tocontemplate a far more complex project – a recontextualization of evolutionary theoryas a change in organismic morphology.102

Stimulated by T.H. Morgan’s Embryology and Genetics (1934), and challenged byDobzhansky’s Genetics and the Origin of Species (1937) –which seemed to deny anyrole for embryology in evolutionary theory –Waddington chiselled away at his ownmodern synthesis, Organisers and Genes (1940).103 Despite his ambitions to bring em-bryology into the genetic fold, his unique experimental expertise, and the tradition ofthis genetic–embryological synthesis inspired by Morgan, Waddington perceived resist-ance from the American genetics network.104 He believed that his contemporariesrejected his contextualization of genetics as a limited means to a greater end: understand-ing the development of complex organismal form. Genes themselves were best con-sidered heuristically, as transitive points in a continuum rather than all-important atomicunits. And in Organisers and Genes, Waddington stressed both individual genes actingto create phenotypic (largely morphological) traits and genetic complexes working inconcert with environmental inputs to produce processual gradients and fields, evolving

100 Needham, Application to ‘Foundation’ containing ‘List of Cambridge Papers on OrganiserPhenomena’, 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, andperson in a reductionist age: the challenge of Gregory Bateson and C.H. Waddington to biological andanthropological 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 theproject. C.H. Waddington, 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, TheEvolution of an Evolutionist, op. cit. (60), p. 7.

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‘equilibria’ rather than randomly combining entities.105 Waddington also elucidatedthree additional concepts in line with his organicism that his genetic contemporarieswould have reacted against: developmental constraint, the ‘paradox of nuclearequivalence during cellular differentiation’, and the Causal Completeness Principle.106

Each of these notions arose and was discussed, albeit utilizing different terminology, atTBC meetings in the 1930s. And, most significantly, each concept challenged the geneticunderpinnings of neo-Darwinism.

Waddington’s indirect attack on neo-Darwinism ultimately made the formation of apost-war social network of organicists more difficult and less cohesive. While he adopteda conciliatory rhetorical style toward orthodox evolution in Organisers and Genes, hissubsequentNature article, ‘Canalization of development and the inheritance of acquiredcharacters’, was more blunt. He leapt right into the ‘battle, which raged for so longbetween the theories of evolution supported by geneticists on one hand and bynaturalists on the other’. Waddington proclaimed scepticism that an ‘exclusive relianceon the natural selection of merely chance mutations’ could simultaneously account for(1) the profound diversity of the organic world; (2) the common-sense discontinuitybetween species; (3) functional fitness of trait to environment; (4) sharp differentiation ofcellular tissues into organs during development; and (5) the tendency –whether at thelevel of tissues, organs, phenotypes, or species – for the developing system to produce‘the standard end-product’.107 Rather than evolution proceeding by the ‘sorting out ofrandom mutations by the natural selective filter’ – that is to say, the neo-Darwinism ofR.A. Fisher and J.B.S. Haldane – he proposed developmental ‘canalization’.108

105 Waddington, op. cit. (103), pp. 92–93. Effectively, Waddington anticipated by two decades thelandmark paper, François Jacob and Jacques Monod, ‘On the regulation of gene activity’, read at Cold SpringHarbor 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 toErrol E. Harris, 10 December 1963, CHW/UEL, MS 3039.4. Rudolph A. Raff, ‘Evo-devo: the evolution of anew discipline’, Nature Reviews Genetics (2000) 1, pp. 74–79. On these topics see Ingo Brigandt, ‘Fromdevelopmental constraint to evolvability: how concepts figure in explanation and disciplinary identity’, in AlanC. Love (ed.), Conceptual Change in Biology: Scientific and Philosophical Perspectives on Evolution andDevelopment, Berlin: Springer, forthcoming; Jan Sapp, Beyond the Gene: Cytoplasmic Inheritance and theStruggle for Authority in Genetics, Oxford: Oxford University Press, 1987, p. 17; Richard M. Burian, ‘Lillie’sparadox – or, some hazards of cellular geography’, in Burian (ed.), The Epistemology of Development,Evolution, and Genetics, Cambridge: Cambridge University Press, 2005, pp. 183–209; Timothy J. Horder,‘Syllabus for an embryological synthesis’, in D.B. Wake and G. Roth (eds.), Complex OrganizationalFunctions: Integration and Evolution in Vertebrates, New York: John Wiley & Sons, 1989, pp. 315–348,p. 340; Scott F. Gilbert, J.M. Opitz and Rudolph A. Raff, ‘Resynthesizing evolutionary and developmentalbiology’, Developmental Biology (1996) 173, pp. 357–372, 361–362; and Ron Amundson, The ChangingRole of the Embryo in Evolutionary Thought: The Roots of Evo-Devo, Cambridge: Cambridge UniversityPress, 2005, pp. 175–180.107 C.H. Waddington, ‘Canalization of development and the inheritance of acquired characters’, Nature

(1942) 150, pp. 563–565, 563.108 Here Waddington drew explicitly from the work of Whitehead. See E.L. Peterson, ‘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. 301–320.

The conquest of vitalism or the eclipse of organicism? 19

Canalization merely takes the appearance of the stability of phenotypes acrossgeological time seriously, rather than assuming it to be illusory. As Waddington saw it,‘constancy’ in phenotypes

must be taken as evidence of the buffering of the genotype against minor variations not only inthe environment in which the animals developed but also in its genetic make-up. That is to say,the genotype can, as it were, absorb a certain amount of its own variation without exhibitingany alteration in development.109

For Waddington, canalization also accounted for the immediate, or intra-generational,fitness that so lured naturalists to neo-Lamarckism. He suggested that canalized develop-ment could account for an example familiar to naturalists of that persuasion, theprenatal origins of sternal and alar callosities in ostriches. Long ago, these callositiesarose only with the appropriate environmental abrasions. But as time went on, thethreshold of skin reactivity became much more sensitive. Ostriches who developedcallouses faster under ever-slighter stimulus would be selected until some sort ofdevelopmental ‘switch’ occurred. After this point, the ostriches would develop callositiesin the appropriate places without environmental stimulus.110

To depict the process of canalization, Waddington adopted the illustration of an‘epigenetic landscape’. As in many of his endeavours, Waddington hoped that byinvoking an unfamiliar term he would highlight the intricate processual relationshipof genetics and development.111 Epigeneticists would investigate the causes of develop-ment without either privileging genes-only explanations or ignoring genetics altogether.But he knew that this truly robust overarching Entwicklungsmechanik was notnecessarily intuitive to Anglo-American biologists. Geneticists would be reluctantto pry open the black box of development; embryologists, on the other hand, would bereluctant to cede any additional territory to genetics. Waddington hoped that hisepigenetic landscape would serve as neutral ground allowing geneticists and embryolo-gists to work together on the mechanics of development. The different levels oforganization at the genetic and embryological/developmental levels would, as Woodgerproposed in TBC meetings, retain their distinctiveness while still allowing for inter-levelanalysis.Waddington’s epigenetic landscape has drawn a great deal of attention, positive and

negative, and it is outside the scope of this paper to address it in detail.112 The significantpoint here is that Waddington’s genetic and evolutionary work from 1942 on wasproperly organicist. Epigenetic changes are indeterminate: many-gene to many-trait.In his model, alterations occur to the entire developmental geography, or, as he called it,

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 theoryappeared in C.H. Waddington, ‘Evolutionary adaptation’, in Sol Tax (ed.), Evolution after Darwin –TheUniversity of Chicago Centennial, Vol. 1: The Evolution of Life, Chicago: The University of Chicago Press,1960.

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‘potentiality’. Should these alterations in the epigenetic landscape occur far enoughupstream in the developmental process, substantial and seemingly discontinuous changewould result in the eventual adult organism. Hence what would get passed on – the unitof heredity and, therefore, evolution –would not be a code-script detailing a particularphenotype but a ranged system of relatively discrete propensities – tendencies rather thaninevitabilities, constraints rather than ‘random walks’. Phyletically, what evolve are deepstructural propensities or combinations of relationships (‘chreods’). Any evolutionaryconcept that could hope to accommodate epigenetic processes would be organicistalmost necessarily.113

The significance of the falling organizers and failing networks (1940s–1970s)

Neither Waddington’s critique of neo-Darwinism nor the ‘fall of the organizer’ caused aconcomitant fall of organicism. Quite simply, organicism did not fall, disappear orbecome discredited due to its association with National Socialism.114 How, then, couldhistorians persuasively argue that it did fall? By the time of Goodfield-Toulmin’sinvestigation in the 1960s, two factors had indeed changed, contributing to the ‘fall’narrative.

First, the social networks of scholars had changed drastically. Needham andWoodgerinvested in other pursuits. As the head of a genetics department at Edinburgh moredevoted to practical animal breeding than the unsolved problems of development,Waddington found few local colleagues with whom he could discuss organicism.115

Between the 1940s and the 1960s, Waddington befriended evolutionary biologistsinterested in theory – Ernst Mayr and Theodosius Dobzhansky most prominently.116

While potentially influential allies, Mayr, Dobzhansky and their network never offeredWaddington the kind of support that the TBC members once had.117 Nor was there amutual experimental focus, as Waddington and Needham shared during their organizerwork. Waddington’s challenges to conventional neo-Darwinism had something to do

113 Eva Jablonka and Marion J. Lamb, 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 C.H. Waddington (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’. Ernst Mayr, The Growth of Biological Thought: Diversity, Evolution, and Inheritance,Cambridge, MA: Belknap Press, 1982, pp. 66–67; and Mayr, ‘Where are we?’, Cold Spring Harbor Symposiaon Quantitative Biology (1959) 2, 1–14. See also Theodosius Dobzhansky, ‘Introductory remarks’, inF.J. Ayala and T. Dobzhansky (eds.), Studies in the Philosophy of Biology: Reduction and Related Problems,Berkeley: University of California Press, 1974, pp. 1–3.117 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 thatWaddington’s ‘chapter’ was still ‘missing’ from the Modern Synthesis. Ernst Mayr, ‘G.G. Simpson’, inErnst Mayr and William B. Provine (eds.), The Evolutionary Synthesis: Perspectives on the Unification ofBiology, Cambridge, MA: Harvard University Press, 1980, p. 452; and Viktor Hamburger, ‘Embryology andthe Modern Synthesis in evolutionary rheory’, in Mayr and Provine, op. cit., p. 97.

The conquest of vitalism or the eclipse of organicism? 21

with this. Mayr never agreed with Waddington on the basic features or the implicationsof their biological theories.118 And Dobzhansky regularly neglected to mentionWaddington’s role in the larger context of twentieth-century evolutionary biology andgenetics.119 In the end, this transatlantic network interested in philosophy, genetics andevolution never solidified around extra-academic pursuits the way the TBC membershad. Furthermore, over time, travel became more difficult as Needham, Waddington,Dobzhansky, Mayr, von Bertalanffy and others – including Gregory Bateson, ArthurKoestler, Michael Polanyi and Paul Weiss – all aged.120 Nevertheless, it would beinappropriate to claim that organicism or the supportive social network entirely dis-appeared. In fact, their followers continued to promote vigorously these conceptsthrough to the end of the century.121

However, even while the network of individuals interested in organicism persisted insubfields of developmental and evolutionary biology, a growing network of analyticphilosophers and molecular biologists declared the concept unhelpful or worse.122 In the1950s, Ernest Nagel pointed out that claims by biologists including E.S. Russell thatorganisms could not be reduced strictly to their constitutive physico-chemical mech-anisms were short-sighted. Our present inability to reduce organisms to the behaviour ofphysico-chemical units should not be regarded as support for irreducibility in principle;for all we know, future biologists might be able to resolve the apparent irreducibility. YetNagel sympathized with the ‘protest of organismic biology against the dogmatismfrequently associated with mechanistic approaches to biology’.123 Francis Crick, whospoke for molecular biologists in the mid-1960s, did not share even these slightsympathies. In Of Molecules and Men, Crick declared that progress in the twentieth-century life sciences depended upon the removal of vitalism. Echoing critiques fromdecades earlier, Crick pinned the vitalist label on organicism and then argued that

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, F.J. Varela, H.R. Maturana and R. Uribe, ‘Autopoiesis: the organization of living systems,

its characterization and a model’, Biosystems (1974) 5, pp. 187–196; D.B. Wake, G. Roth and M.H. Wake,‘The problem of stasis in organismal evolution’, Journal of Theoretical Biology (1983) 101, pp. 211–224; andR.C. Lewontin, The Triple Helix: Gene, Organism, and Environment, Cambridge, MA: Harvard UniversityPress, 2001.122 Four influential symposia were dedicated to the question of organicism and reductionism in the 1960s

and 1970s: Arthur Koestler’s Alpbach Symposium ( Arthur Koestler and J.R. Smythies, 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 – bothheld at the Villa Serballoni in Bellaglo, Italy (Ayala and Dobzhansky, op. cit. (116); and John B. Cobb andDavid R. Griffin, Mind in Nature: Essays on the Interface of Science and Philosophy, Claremont: UniversityPress of America, 1977).123 Ernst Nagel, ‘Mechanistic explanation and organismic biology’, in RonMunson (ed.),Man and Nature:

Philosophical Issues in Biology, New York: Delta Books, 1971, pp. 19–32. Notably, Nagel focused his critiqueon the 1920s organicism of E.S. Russell and completely by-passed that of Woodger and von Bertalanffy.

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vestiges of vitalism were a drag on the advance of biology.124 Waddington pointedout in his review of Of Molecules and Men that Crick debated straw men.125

While privately Crick admitted that his critique lacked precision, he believed it did notmatter: the reductionistic ‘genes-for’ language Crick deployed was the way moleculargeneticists described the organic world.126 If organicism resisted the perspective ofmolecular biologists that all explanations in biology were ultimately reducible tomicroscopic physico-chemical entities, then ultimately organicists would be discredited.The molecular approach, as W.T. Astbury reminded Waddington in an exchangepublished in Nature, ‘is the coming biology’.127 In addition, Crick and Astbury hadthe support of former post-war TBC member Sir Peter Medawar, who had debatedwith Woodger, Needham and Waddington about these very issues in the 1940s and1950s.128 It was conversations with Medawar around the time of this address thatinspired Goodfield-Toulmin to write her proposed narrative of the ‘rise and fall of theorganizer’.

Revolutionary claims of this sort about the reducibility of whole organisms tomicroscopic entities had been made earlier in the twentieth century – they motivatedWhitehead to write Science and the Modern World.129 But by the 1960s and 1970s,there was a second reason the situation had changed: after the Second World War, theprofessional ethos of the life sciences slowly began to shift in accordance with economicprinciples favouring short-term results.130 At its core, organicism is a philosophy thatasks questions about the nature of the organism and only indirectly promises practicalbiological control, as the organizer project itself demonstrated. Waddington sensed theincongruity of this kind of philosophy with the state of biology in the second half of thecentury: ‘reductionism is a recipe for action’.131 When what is called for is making a‘quick (scientific) buck by discovering some useful practical information’, the reduc-tionistic mechanism embraced by Crick and Astbury trumped organicism. By the latterhalf of the twentieth century, the metaphysics of Crick and Astbury had become an‘epidemic intellectual disease, which causes people to argue that the reality of anything is

124 Francis Crick, Of Molecules and Men, Seattle: University of Washington Press, 1966.125 C.H. Waddington, ‘No vitalism for Crick; review of Of Molecules and Men by Francis Crick’, Nature

(1967) 216, pp. 202–203. Other reviewers concurred that Crick was conflating organicism with vitalism tomake a point about biological progress (for example Laura Livingston, ‘Review: Of Molecules and Men byFrancis Crick’, Yale Journal of Biology and Medicine (1968) 40, p. 328).126 Crick to Waddington, 9 November 1967, andWaddington 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 W.T. Astbury, ‘Molecular biology or ultrastructural biology?’, Nature (1961) 190, p. 1124.128 Peter Medawar, ‘A biological retrospect’, Nature (1965) 207, pp. 1327–1330, 1328.129 Benson, op. cit. (12), details some of these earlier attempts.130 Michael Callon, ‘From science as an economic activity to socioeconomics of scientific research: the

dynamics of emergent and consolidated techno-economic networks’, in Phil Mirowski and Esther-Miriam Sent(eds.), Science Bought and Sold: Essays in the Economics of Science, Chicago: The University of Chicago Press,2002, pp. 277–317.131 C.H. Waddington, Tools for Thought: How to Understand and Apply the Latest Scientific Techniques

of Problem Solving, New York: Basic Books, 1977, p. 23.

The conquest of vitalism or the eclipse of organicism? 23

proportional to the precision with which it can be defined in molecular or atomicterms’.132 Organicism, he protested, was the ‘method for making major advances inhuman comprehension’, like those of ‘Darwin, Freud, Einstein [and] the quantumphysicists’. On the other hand, reductionistic mechanism promised predictabilityand control. By the end of his life, this approach appeared to take its place as the‘Conventional Wisdom of the Dominant Group’; Waddington shortened the phrase to amemorable acronym, ‘COWDUNG’.133

In this essay, I have attempted to ground the history of organicism in the twentiethcentury in a shifting social network. In some ways, the resulting picture reflects themetaphor that Needham, Waddington and, indeed, Whitehead suggested applied to theuniverse as a whole: that of a river. A concept runs its particular course because of socialnetworks of adherents: scientists gathered in pools, sometimes mere eddies, discussingand applying their philosophy to their biology. Once viewed through the lens of thesesocial networks, the fate of any particular set of experiments in the larger researchstream has less to do with macroscopic social or conceptual factors than with micro-scopic ones. For instance, the apparent failure of the organizer project was not a stage inthe conquest of vitalism. Rather, political intrigue at Cambridge, funding priorities at theRockefeller Foundation, Needham’s political interests, and the financial problems ofWaddington had more to do with its demise. If the concept of organicism seemed tosuffer the same fate as the organizer project, that also had less to do with its misappliedassociation with fascism, for instance, than with the character of the social groupsupporting it. Though organicism did not disappear to be rediscovered concomitantlywith evo-devo, a plurality of factors weakened the social network of adherents.Molecular biology did not ‘rise’ to displace organicism, but the simplicity and rhetoricalpower of the progress-through-devitalizing-biology narrative meant that organicists hadto carefully qualify their dissent.

132 C.H. Waddington, ‘Fifty years on’, Nature (1975) 258, pp. 20–21, 21.133 Waddington, op. cit. (131), pp. 24–25.

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