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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

Published online by Cambridge University Press:  29 October 2013

ERIK PETERSON
ERIK PETERSON*
Affiliation:
History Department at the University of Alabama, 208 Ten Hoor Hall, Tuscaloosa, AL 35487-0212, USA. Email: elpeterson@as.ua.edu.
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Abstract

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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Type
Research Article
Information
The British Journal for the History of Science , Volume 47 , Issue 2 , June 2014, pp. 281 - 304
Copyright
Copyright © British Society for the History of Science 2013 

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

Donna Haraway, Crystals, Fabrics, and Fields Footnote 1

Old wounds, reopened

In March of 1965, June Goodfield-Toulmin wrote to Joseph Needham at Cambridge University 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 research for 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 this point had been cordial,Footnote 2 Joseph Needham responded to her critically. ‘[You] will undoubtedly fail’, he insisted, ‘unless you can achieve a more open mind than is to be inferred from the phrase [growth and downfall]’. He saw the ‘drama’ she hoped to narrate in quite a different light: ‘the perennial struggle of conceptual opposites; here stimulus vs. reactivity, as perennial as the continuity vs. discontinuity antithesis in physics’.Footnote 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 fell down was a straw man erected by some American biologists for the express purpose of pushing it over. They performed this simple task so much to their own satisfaction that they had some success in persuading American biologists in general that the problem of embryonic induction is not important …Footnote 4

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

It is because the essential connection between [genetics and embryology] … has been obscured for the last quarter of a century, that molecular biologists are now saying that they will have to move in and solve the problems of differentiation from scratch, as though nothing of value had yet been discovered. And of course when they do turn their attention to the subject, they light immediately on … the phenomena of embryonic induction, i.e., ‘the organiser’.

What was at stake in this private disagreement? At first glance, Goodfield-Toulmin sought merely first-hand insight into the ‘organizer project’: a series of embryological and biochemical experiments conducted at Cambridge by Needham and Waddington, among others, from 1933 to 1938. They derived their work on the phenomenon of embryonic induction from the Nobel Prize-winning experiments of Hans Spemann and Hilde Mangold in 1935.Footnote 5 Though a vigorous science in its day, few scientists continued investigating the organizer phenomenon in the decades immediately after the Second World War.Footnote 6 By the time Goodfield-Toulmin wrote to Needham and Waddington, many historians and philosophers of science regarded Spemann's organizer experiments as vitalism.Footnote 7 Goodfield-Toulmin intended her history of the organizer to reveal the negative impact that non-epistemic values such as vitalism have upon scientific research.Footnote 8 For Waddington and Needham, Goodfield-Toulmin was perpetuating a divide between contemporary molecular biology, which she and others believed had unified the life sciences, and vitalism, which she viewed as faulty metaphysics.Footnote 9 But the ‘rise-and-fall’ narrative troubled Waddington. The misreading of this event was symptomatic of a larger problem: ‘the extremely unfortunate divorce that has grown up between genetics and embryology in the States’.Footnote 10

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

A decade ago, Gilbert and Sarkar reclaimed organicism and persuasively argued that organicism was the right approach for twenty-first-century ‘evo-devo’.Footnote 12 Organicism had been discarded, they asserted, because it kept ‘bad company’ with fascism, communism and marginal New Age groups.Footnote 13 This article tacitly supported the idea that 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 the scope of the organicist social network. My research suggests that, in fact, a substantial number of influential scholars in multiple biological subfields held organicist beliefs through the majority of the twentieth century. From this perspective, we should view organicism 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 and attempted reinstantiation of organicist social networks – including most prominently the Cambridge Theoretical Biology Club. Perhaps the organizer project ‘fell’. Nevertheless, the reasons for its fall and its connection to the philosophy of organicism are neither straightforward nor strictly conceptual. Departmental politics, budgets and personal preferences may have sealed the fate of Cambridge's organizer project, but the organicism espoused by Needham and Waddington did not follow the same path as that of the organizer project. Based on previously unexplored archival materials, this essay brings to the fore the importance of collections of interacting individuals who held similar views on the philosophy of the organism and a commitment to apply those holistic views to the life sciences and beyond.Footnote 14

What was/is twentieth-century organicism?

As mentioned above, Needham and Waddington held to a philosophy called organicism, a concept that pre-dated any work on the organizer problem.Footnote 15 A decade or more before Needham and Waddington took up organicism, John Scott Haldane and Edward Stewart Russell pronounced against the reductionistic mechanism that they saw in the life sciences of their day. Haldane hoped to retain the common-sense appeal of vitalism while simultaneously insisting upon testable entities and forces attributed to mechanistic biology.Footnote 16 He thought this ‘organicism’ provided a way to avoid the dualistic vitalism of Hans Driesch, but without conceding that the organic world was nothing but physico-chemical mechanism.Footnote 17 According to his version of organicism, traits of live organisms – their ability to self-regulate within certain constraints, for instance – emerged from properties 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 material substance in the world) and emergent. By emergent, Haldane referred to the peculiar ‘popping up’ of structures or behaviours or powers unique to organized systems, not exclusively reducible to the physico-chemical states from which they are constructed. His contemporary, Glaswegian ichthyologist Russell, found it more difficult to downplay the apparent distinction between living and non-living.Footnote 18 Like Haldane, Russell adopted the ‘simple everyday conception’ that organisms ‘are active, purposeful agents’ while avoiding the ‘vitalism’ of Lamarck or Samuel Butler.Footnote 19 For Russell, organicism was possible because organisms possessed an ‘essential unity’, the individuated parts of which ‘can never fully explain the activity of the whole; they are to be regarded rather as conditions, both limiting and implementing that activity’.Footnote 20 The structure of organisms as wholes, in other words, seemed to constrain, and therefore precede, the function of parts.

Aside from Haldane and Russell, organicism drew adherents from a plurality of backgrounds. American marine biologist William Emerson Ritter composed a two-volume tome defending ‘the facts … of organic unity and integration’ before going on to lead the United States Science Service.Footnote 21 Prime minister of South Africa Jan C. Smuts criticized both the notion that life and mind were substances distinct from the material and the reduction of ‘life and mind to a subsidiary and subordinate position as mere epiphenomena’.Footnote 22 Harvard University biochemist Lawrence J. Henderson spent five years sketching his alternative to both mechanism and vitalism.Footnote 23 Still others insisted that reductionistic mechanism no longer sufficed to explain the overwhelming complexity of the vast majority of phenomena in the living world.Footnote 24 Just as J.S. Haldane claimed that organisms exhibited properties belonging to a seemingly different class than the materials from which they emerged, Cambridge philosopher of mind C.D. Broad spelled out a version of mental emergence that he contrasted to ‘Substantial Vitalism’.Footnote 25

Alfred North Whitehead drew these organicist threads together into a more comprehensive world view. In his Science and the Modern World, Whitehead criticized the tendency of scientists, as well as the general public, to view modernity through the lenses of materialism, mechanism and reductionism. Paradoxically, modern physics did not depict that sort of universe.Footnote 26 Whitehead suggested that other modern sciences follow the trend in physics, elevating the prominence of networks – he often used the term nexus – over entities. Process, form, development, exchange, reciprocity and interaction across integrated systems should displace descriptions of the behaviour of individuals by the functions and compositions of parts. In other words, Whitehead called for a science focused on multi-perspectival networks of relationships subjectively analysable – though not objectively definable – as concrete ‘events’ or ‘chreods’ rather than as the behaviour of aggregated atomic units.Footnote 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 deeply impacted by the work of Whitehead, D'Arcy Thompson and others, no ‘school’ of organicism existed. Nevertheless, as we will see below, it was at Cambridge that a cadre of philosophically sophisticated scientists inspired by these intellectual ancestors began to gather into a tighter network during the 1930s to clarify this organicist ‘third way’ between dualistic vitalism and reductionistic mechanism.Footnote 28 Spemann's organizer became another catalyst in the formation of that social network.

Metaphysics and the ‘organizer problem’ (1924)

How do the organizer experiments fit into the history of organicism? A main problem with concepts like organicism has less to do with their content than with their use. How does a metaphysical commitment – vitalism, organicism or reductionist mechanism, for that matter – actually impact scientific work? Can it only inhibit research progress?Footnote 29 Hans Spemann believed that his metaphysics had led him to investigate and interpret the organizer phenomenon that led to a Nobel Prize.

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

Questions over Spemann's philosophical proclivities notwithstanding, the organizer grew to be a fashionable international pursuit bridging biochemistry and embryology in the pre-Second World War period.Footnote 34 Excited researchers expected that the biochemical identification of the organizer would be the key to unlocking the most basic problem of development, morphological patternedness.Footnote 35 A small group of multidisciplinary scholars at Cambridge were numbered among the pursuers of the organizer. Unlike their contemporaries, however, the Cambridge group insisted that a search for a single biochemical 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. Conversations in the late 1920s between Joseph Needham and J.H. Woodger in England and Woodger and Ludwig von Bertalanffy in Austria helped to refine organicism and the social 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 biochemistry and embryology. Initially, he fixated on seemingly insoluble problems relating to the development of chicks in their eggs, eventually writing the interdisciplinary magnum opus of his young career, the three-volume Chemical Embryology. Spemann's organizer phenomenon appeared to be the next challenge.Footnote 36 Needham's interactions with Joseph H. Woodger were among the factors contributing to his interest in the organizer project.

At Woodger's death, even Karl Popper called him ‘Socrates’.Footnote 37 Over a career nearly seven decades long, Woodger had cast a quiet shadow over the philosophy of biology in the Anglo-American world. Members of the Vienna circle met him in 1926; philosophers all over the world, including Alfred Tarski, Rudolf Carnap and W.V.O. Quine, thought him a friend and ally. However, later scholars tended to overlook Woodger's importance as an isolated example of overly analytic times, not offering much for either the bench scientist or the philosopher.Footnote 38

Though they had corresponded before, the first major public exchange between Woodger and Needham centered on Needham's review of Woodger's Biological Principles in Mind. Needham reviewed Biological Principles positively. Still Woodger believed 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 as each remained consigned to its right place.Footnote 39 Woodger, by contrast, bluntly denied that any via media could be constructed between these two camps as they were defined in the 1920s. Too many of the obvious problems of biology were ‘absolutely insoluble’, he thought, so long as mechanists and vitalists each argued over a concept of entities. For Woodger, levels of organization, internal and multiple relations, and the counterentropic change most easily observed during the process of development were the kinds of organic factor that rendered biology epistemologically, if not qualitatively, parallel (rather than reducible) to the physico-chemical.Footnote 40 Woodger (following Whitehead) believed that the quantum revolution in physics had emancipated biology. Like physicists, biologists no longer had to speak in terms of determined entities using merely physico-chemical language. Biology could be autonomous.

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

Woodger connected with von Bertalanffy through their mutual acquaintances with the Viennese circle of logical empiricists. Von Bertalanffy would not introduce his general systems theory, for which he became widely known, until after the Second World War.Footnote 44 But Woodger had already convinced him to join in on the ‘organicist’ project by the late 1920s. He translated and elaborated von Bertalanffy's Kritische Theorie der Formbildung; the resulting Modern Theories of Development became a cornerstone text in theoretical embryology.Footnote 45 As with other organicists, von Bertalanffy was convinced that neither reductionist mechanism nor dualistic vitalism could serve as the true foundation of biology.Footnote 46 Significantly, von Bertalanffy believed that as a bridge between common-sense vitalism and reductionisic mechanism, organicism served the purpose of theoretical unification that the other philosophies had also claimed. Biology, like chemistry and physics, required a ‘methodological clarification’ as ‘protection against the fallacies of hurried hypotheses’.Footnote 47 This new theoretical biology should begin embryology because theoretical rigour in embryology would align future progress in the rest of experimental biology and remedy the inefficient infighting of reductionistic mechanists and dualistic vitalists.Footnote 48

From the point of view of von Bertalanffy, Austro-German gestalt theorists, and even Spemann, a metaphysical monism could in principle connect levels from subatomic physics to psychology.Footnote 49 Here Woodger differed from the Germans. He believed that ‘flattening out’ would not work. Instead an important middle-range theory needed to be inserted and explicated: a structural connection between physico-chemical non-living objects and biological ones. He sketched out his approach in a series of dense articles published in the Quarterly Review of Biology over 1930–1931.Footnote 50 Here, he criticized the epistemological dead end of Driesch's vitalism, but he also directed gentle jabs at ‘neo-mechanism’ – the theory endorsed by Needham at that timeFootnote 51 – and, notably, J.S. Haldane's ‘concept of organism’. Because of the vagueness of definitions such as Haldane's, organicism had not necessarily ‘been tried and found wanting’, but had ‘been found difficult and not tried’.Footnote 52 Woodger's goals, then, were dual: (1) to clarify the messy linguistic conventions of biology, and (2) to convince working biologists that ‘new ways of thinking’ more related to hypothetico-deductive algorithms than to ‘intuitions’ should be pursued if biology was going to be put on firmer (that is to say, non-Cartesian or dualist) footing.Footnote 53 Capturing the ethos of organicism, Woodger wanted to render organismic biology philosophically respectable while acknowledging the intuitions of vitalists 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 Woodger and Needham attended the Second International Congress of the History of Science and Technology held in London that year. There, popular Bolshevik revolutionary and philosopher of science Nikolai I. Bukharin delivered his address, ‘Theory and practice from the standpoint of dialectical materialism’. Bukharin's perspective greatly impressed Needham.Footnote 54 By the end of the conference Needham had adopted a more radical posture in line with Bukharin's exposition: he was ready to discard Eurocentrism, the collusion of science with oppressive capitalism, and the dyadic opposition of vitalism and mechanism at the heart of Western biology.Footnote 55 In the second decisive event of 1931, Needham connected Woodger with another Cambridge scientist glad to explore theoretical biology – Conrad Hal ‘Wad’ Waddington.Footnote 56 Waddington had recently transformed himself from palaeontologist to population geneticist to embryologist.Footnote 57 Soon C.H. Waddington would become the main promulgator of organicism (or ‘Woodgery’, as he liked to call it).Footnote 58

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

Waddington shared many interests with Needham: he had been profoundly impacted by the philosophy of Alfred North Whitehead, leaned left in his politics, and straddled disciplinary lines throughout his career.Footnote 59 Like Needham, Waddington was a dedicated morris dancer and shared non-traditional attitudes regarding marital relationships.Footnote 60 With so much in common, Needham and Waddington jumped at the chance to work together at Strangeways Laboratory in Cambridge. Their friendship resulted in a deepening of the organicist social network, established Cambridge as a leading centre for experimental embryology until 1938, and would strongly influence the remainder of Waddington's career as he explored the intersection of development and evolution.

Waddington came to Strangeways as a novice in embryology in 1929 following a brief period of unfulfilling work in population genetics with Edith Saunders and J.B.S. Haldane.Footnote 61 At the same time, from 1929 to 1931, Waddington had obtained a scholarship in philosophy and had made his own excursion into the vitalism–mechanism debate.Footnote 62 He read Spemann, and from that reading decided that he would attempt the tissue transfer techniques on much more difficult avian embryos. Surprisingly, he was successful enough to receive an invitation from Richard Goldschmidt to continue work in Germany. After receiving a Rockefeller Medical Fellowship in 1931, Waddington joined Otto Mangold's Berlin-Dahlem lab.Footnote 63 Later, Joseph Needham and his wife, Dorothy, joined Waddington to investigate the physico-chemical process of amphibian development. This partnership produced ten co-authored papers on amphibian development from 1931 to 1937.Footnote 64

In retrospect, it seems almost inevitable that this small cadre would develop a more tightly knit relationship. They soon brought leftist crystallographer J.D. Bernal and his associate Dorothy Crowfoot Hodgkin into their social network.Footnote 65 In April 1932, Needham suggested to Woodger that they have a small excursion to discuss theoretical issues. Woodger spoke with Waddington, who enthusiastically agreed to attend. Woodger volunteered his family's summer cabin in Epsom Down and suggested that their informal ‘biotheoretical gathering’ expand to include E.S. Russell, among others (Russell did not attend).Footnote 66 Needham would later give the ‘Conference’ a more formidable moniker from the Cambridge lexicon: the ‘Theoretical Biology Club’ (hereafter TBC).Footnote 67 Together, this collection of young scientists and philosophers inspired interdisciplinary work in the Oxbridge–London academic triangle and refined more carefully both the concept and the application of organicism.Footnote 68

Their meetings covered a gamut of issues in theoretical biology. At the first meeting in August 1932, Bernal drew from his training in crystallography to suggest that a morphological scale of bodily form could be drawn that included everything from the level of quantum structures to metazoa.Footnote 69 Mathematician Dorothy Wrinch spoke of ‘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 about homeotic mutations in Drosophila – the basis for his important aristapedia experiments years later.Footnote 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 crystallography.

This feature, their extra-academic common interests, distinguished this small group from other contemporaries interested in theoretical biology. The TBC not only staged detailed discussions of biological philosophy and practice, it established bonds of friendship extending beyond their careers or institutional affiliations. Fuelled by talk of Whitehead, D'Arcy Thompson, Friedrich Engels and others, they discussed the scientific 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 in the summer of 1933 the Needhams travelled with Waddington to Otto Mangold's Berlin-Dahlem laboratory, where they would collaborate on the organizer. There they demonstrated for the first time that the phenomenon – Waddington by then had labelled it ‘evocation’ – could be produced by non-living tissue. This finding convinced Needham and Waddington that Spemann's explanations of the phenomenon were not quite right.Footnote 71 When they returned to Britain, they agreed to invite participants to the TBC from outside the initial group to help address what had become a far more difficult scientific and philosophical problem: how could non-living tissue instigate embryonic development?

One of those invited was Viennese philosopher Karl Popper, who attended the May 1935 meeting held at the Woodgers’ cabin.Footnote 72 He returned a year later to meet with the TBC at the ‘old windmill at Hunstanton’.Footnote 73 The June 1936 meeting was the apex of the TBC – Popper, J.B.S. Haldane, Hyman Levy and others joined Waddington, Needham, Woodger and Bernal. Conversation during meals at the 1936 meeting circulated around socialism, science and state planning. Though Popper disagreed with the political leanings of the group, he found organicism appealing. Even many decades later, he refused to support reductive mechanism.Footnote 74

TBC meetings unquestioningly impacted the organicist science of its core members.Footnote 75 As a result, they would stand by two claims in particular: (1) ‘the properties of a component depend upon the component of which it is a component (relational properties) 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 are processes rather than entities) by artificially dividing them up to render them potentially analysable.Footnote 76 The first point is an ontological claim about the integration of organismic structures; the second an epistemological one about the provisionality of scientists’ knowledge. Inspired by discussions at the TBC, Needham and Waddington expected systemic rather than atomic explanations for the evocation process, to appreciate a plurality of inputs at different levels inside the organism (for example, genetic, epigenetic, cytological and so on), and to regard any isolable set of developmental causes as incomplete explanatory factors.Footnote 77 Not only did this mean that the organizer work of Needham, Waddington and their occasional co-authors did not resemble the contributions of other international organizer groups, it would also eventually make the acquisition of funding more difficult: funding agencies believed the organizer to be a discrete biochemical compound.

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

Early in 1934, however, the Cambridge organizer work seemed exceptionally promising. Inspired by their cooperative, multidisciplinary attack on a ‘borderline subject’, W.E. Tisdale of the Rockefeller Foundation approached Needham in April and asked him to suggest ways in which the foundation could advance ‘chemical embryology’.Footnote 78 By June, Needham had submitted an extremely ambitious plan for an Institute of Embryology. When that plan was rejected, Needham sent a hardly more modest proposal to establish a formal institute devoted to ‘Mathematico-physico-chemical [MPC] Morphology’.Footnote 79 We do not know the true motivations for supporting Needham's plan, nor do we know why it was eventually rejected. There are hints that the explication of morphology with reference to physics and chemistry fit the Rockefeller's preference for funding projects interested in the ‘ultimate littleness of things’.Footnote 80 In the end, however, disciplinary interstices represented precariously rocky soil for Rockefeller seed money and dreams of an MPC morphology unit at Cambridge never materialized.

The survival of the Biochemical Laboratory Extension was further complicated by 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 indeed lead to a ‘fall’ of the organizer project at Cambridge and contributed to the break-up of the Cambridge TBC.Footnote 81

Conflict over Waddington's salary ended up severely, if temporarily, hampering his scientific career. From 1933, Waddington held a fellowship at Christ's, a part-time lectureship in zoology, and, on account of his philosophical prowess, the Royal Commissioners of the Exhibition of 1851 Senior Studentship. In 1936, the studentship was cut by two-thirds; after paying his alimony, this left him with a meager £250 per annum. When Needham turned to the Rockefeller Foundation on his behalf seeking emergency salary assistance, the Cambridge Council – spurred by Waddington's colleague James Gray – took offence.Footnote 82 After some directed consultation, the Council and General Board changed their policy to allow for direct oversight, which blocked future efforts to secure salary funding for Waddington.Footnote 83

Understandably, haggling over Waddington's salary led to bad feelings between Needham and Waddington on the one side and Cambridge University officials on the other. By 1938, Tisdale revealed to Needham that even Hopkins's support for his and Waddington's organizer work was waning and, with Hopkins's retirement nearing, Tisdale warned Needham that Hopkins's successor would likely cut off Needham entirely.Footnote 84 Hopkins rejected Tisdale's suggestion as ‘absurd’ and vowed to support ‘your work on the Organiser’.Footnote 85 Little improved, however. By September 1938, Needham applied to the Royal Society for a grant; Hopkins wrote in support.Footnote 86 In the grant application, 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 done in Cambridge, though Cambridge has at present the lead in this subject. If it is not done in Cambridge, there is no chance of its being done anywhere else in England … On the whole, it may 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 recognise the value of research into the deepest processes of early embryonic development.Footnote 87

The Royal Society did fund Needham, but permanent damage had been done. Waddington transitioned most of his research to Strangeways. Cambridge failed to support 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 1933–1934 that Weaver felt compelled to invest in the work straddling Hopkins's biochemistry lab and the Strangeways Laboratory through the mid-1930s.Footnote 88 At one point the Rockefeller even funded laboratory technicians to work with Needham and Waddington at Cambridge.Footnote 89 Yet the succession of failed attempts at identifying the specific biochemical essence of the organizer was not the solution the Rockefeller Foundation had hoped for, even if it sustained Waddington and Needham's organicist suppositions.Footnote 90 By 1938, Waddington reported that even some toxic substances induced the embryo to symmetrically develop just as any other ‘natural’ compound. To Waddington, such a result indicated that the ‘evocator … appears impossible to discover’ given some of the ruling assumptions that ‘the artificial [i.e. experimental] process gives a true picture of what happens in nature’.Footnote 91 He introduced the speculation he and Needham shared that, rather than a biochemical substance that induced change, the evocation principle was distributed throughout a field or gradient in the developing embryo but suppressed. Once the suppressor was released by adding the synthetic chemical compound, the embryo began the developmental process. Johannes Holtfreter – who escaped Germany to work with Waddington and Needham in 1939–1940 – later confirmed this experimental result.Footnote 92 Nevertheless, while it was still an active field of research, the Rockefeller would no longer support embryology at Cambridge without requisite university support.Footnote 93

Lacking prospects of permanent financial support doing experimental embryology at Cambridge, Needham and Waddington began pursuing other interests.Footnote 94 Considering the troubles the others were having at Cambridge, even Woodger began devoting more time to meetings with Tarski and ‘a large number of new friends’.Footnote 95 In 1937, he travelled to Poland; he was at Yale on a Rockefeller grant in 1938, meeting Waddington's close friends Gregory Bateson and Margaret Mead, among others.Footnote 96 After his return, he spent increasing amounts of time with scholars at the London School of Economics. When he reconvened the TBC again after the war, most of the original members had moved on.Footnote 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 at T.H. Morgan's Drosophila lab at the California Institute of Technology.Footnote 98 Over the 1938–1939 academic year, he worked on Drosophila development alongside Morgan's comrades Alfred Sturtevant and Theodosius Dobzhansky.Footnote 99

By then the Cambridge organizer project was most certainly on its deathbed. Needham remained in biochemistry, but ceased writing grants to support work on the organizer by 1939.Footnote 100 But at Cambridge the ‘fall of the organizer’ had little to do with a concomitant failure of organicism or the death of vitalism. It was a contingent victim of lack of funding by the Rockefeller and disinterest, even active opposition, by other members 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, the organicism motivating the ‘rise of the organizer’ was not dead at all and in fact had begun to infiltrate genetics.

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

Waddington's year of work in the United States reinforced his belief that American genetics did not really know what to do with the problems of development dealt with by experimental embryology or the organicist interests he entertained.Footnote 101 Before he had even completed his first genetics text, Introduction to Modern Genetics, he began to contemplate a far more complex project – a recontextualization of evolutionary theory as a change in organismic morphology.Footnote 102

Stimulated by T.H. Morgan's Embryology and Genetics (1934), and challenged by Dobzhansky's Genetics and the Origin of Species (1937) – which seemed to deny any role for embryology in evolutionary theory – Waddington chiselled away at his own modern synthesis, Organisers and Genes (1940).Footnote 103 Despite his ambitions to bring embryology into the genetic fold, his unique experimental expertise, and the tradition of this genetic–embryological synthesis inspired by Morgan, Waddington perceived resistance from the American genetics network.Footnote 104 He believed that his contemporaries rejected his contextualization of genetics as a limited means to a greater end: understanding the development of complex organismal form. Genes themselves were best considered heuristically, as transitive points in a continuum rather than all-important atomic units. And in Organisers and Genes, Waddington stressed both individual genes acting to create phenotypic (largely morphological) traits and genetic complexes working in concert with environmental inputs to produce processual gradients and fields, evolving ‘equilibria’ rather than randomly combining entities.Footnote 105 Waddington also elucidated three additional concepts in line with his organicism that his genetic contemporaries would have reacted against: developmental constraint, the ‘paradox of nuclear equivalence during cellular differentiation’, and the Causal Completeness Principle.Footnote 106 Each of these notions arose and was discussed, albeit utilizing different terminology, at TBC meetings in the 1930s. And, most significantly, each concept challenged the genetic underpinnings of neo-Darwinism.

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

Canalization merely takes the appearance of the stability of phenotypes across geological 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 in the 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 exhibiting any alteration in development.Footnote 109

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

To depict the process of canalization, Waddington adopted the illustration of an ‘epigenetic landscape’. As in many of his endeavours, Waddington hoped that by invoking an unfamiliar term he would highlight the intricate processual relationship of genetics and development.Footnote 111 Epigeneticists would investigate the causes of development without either privileging genes-only explanations or ignoring genetics altogether. But he knew that this truly robust overarching Entwicklungsmechanik was not necessarily intuitive to Anglo-American biologists. Geneticists would be reluctant to pry open the black box of development; embryologists, on the other hand, would be reluctant to cede any additional territory to genetics. Waddington hoped that his epigenetic landscape would serve as neutral ground allowing geneticists and embryologists to work together on the mechanics of development. The different levels of organization at the genetic and embryological/developmental levels would, as Woodger proposed in TBC meetings, retain their distinctiveness while still allowing for inter-level analysis.

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.Footnote 112 The significant point here is that Waddington's genetic and evolutionary work from 1942 on was properly 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, ‘potentiality’. Should these alterations in the epigenetic landscape occur far enough upstream in the developmental process, substantial and seemingly discontinuous change would result in the eventual adult organism. Hence what would get passed on – the unit of heredity and, therefore, evolution – would not be a code-script detailing a particular phenotype but a ranged system of relatively discrete propensities – tendencies rather than inevitabilities, constraints rather than ‘random walks’. Phyletically, what evolve are deep structural propensities or combinations of relationships (‘chreods’). Any evolutionary concept that could hope to accommodate epigenetic processes would be organicist almost necessarily.Footnote 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 a concomitant fall of organicism. Quite simply, organicism did not fall, disappear or become discredited due to its association with National Socialism.Footnote 114 How, then, could historians persuasively argue that it did fall? By the time of Goodfield-Toulmin's investigation in the 1960s, two factors had indeed changed, contributing to the ‘fall’ narrative.

First, the social networks of scholars had changed drastically. Needham and Woodger invested in other pursuits. As the head of a genetics department at Edinburgh more devoted to practical animal breeding than the unsolved problems of development, Waddington found few local colleagues with whom he could discuss organicism.Footnote 115 Between the 1940s and the 1960s, Waddington befriended evolutionary biologists interested in theory – Ernst Mayr and Theodosius Dobzhansky most prominently.Footnote 116 While potentially influential allies, Mayr, Dobzhansky and their network never offered Waddington the kind of support that the TBC members once had.Footnote 117 Nor was there a mutual experimental focus, as Waddington and Needham shared during their organizer work. Waddington's challenges to conventional neo-Darwinism had something to do with this. Mayr never agreed with Waddington on the basic features or the implications of their biological theories.Footnote 118 And Dobzhansky regularly neglected to mention Waddington's role in the larger context of twentieth-century evolutionary biology and genetics.Footnote 119 In the end, this transatlantic network interested in philosophy, genetics and evolution never solidified around extra-academic pursuits the way the TBC members had. Furthermore, over time, travel became more difficult as Needham, Waddington, Dobzhansky, Mayr, von Bertalanffy and others – including Gregory Bateson, Arthur Koestler, Michael Polanyi and Paul Weiss – all aged.Footnote 120 Nevertheless, it would be inappropriate to claim that organicism or the supportive social network entirely disappeared. In fact, their followers continued to promote vigorously these concepts through to the end of the century.Footnote 121

However, even while the network of individuals interested in organicism persisted in subfields of developmental and evolutionary biology, a growing network of analytic philosophers and molecular biologists declared the concept unhelpful or worse.Footnote 122 In the 1950s, Ernest Nagel pointed out that claims by biologists including E.S. Russell that organisms could not be reduced strictly to their constitutive physico-chemical mechanisms were short-sighted. Our present inability to reduce organisms to the behaviour of physico-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. Yet Nagel sympathized with the ‘protest of organismic biology against the dogmatism frequently associated with mechanistic approaches to biology’.Footnote 123 Francis Crick, who spoke for molecular biologists in the mid-1960s, did not share even these slight sympathies. In Of Molecules and Men, Crick declared that progress in the twentieth-century life sciences depended upon the removal of vitalism. Echoing critiques from decades earlier, Crick pinned the vitalist label on organicism and then argued that vestiges of vitalism were a drag on the advance of biology.Footnote 124 Waddington pointed out in his review of Of Molecules and Men that Crick debated straw men.Footnote 125 While privately Crick admitted that his critique lacked precision, he believed it did not matter: the reductionistic ‘genes-for’ language Crick deployed was the way molecular geneticists described the organic world.Footnote 126 If organicism resisted the perspective of molecular biologists that all explanations in biology were ultimately reducible to microscopic physico-chemical entities, then ultimately organicists would be discredited. The molecular approach, as W.T. Astbury reminded Waddington in an exchange published in Nature, ‘is the coming biology’.Footnote 127 In addition, Crick and Astbury had the support of former post-war TBC member Sir Peter Medawar, who had debated with Woodger, Needham and Waddington about these very issues in the 1940s and 1950s.Footnote 128 It was conversations with Medawar around the time of this address that inspired Goodfield-Toulmin to write her proposed narrative of the ‘rise and fall of the organizer’.

Revolutionary claims of this sort about the reducibility of whole organisms to microscopic entities had been made earlier in the twentieth century – they motivated Whitehead to write Science and the Modern World.Footnote 129 But by the 1960s and 1970s, there was a second reason the situation had changed: after the Second World War, the professional ethos of the life sciences slowly began to shift in accordance with economic principles favouring short-term results.Footnote 130 At its core, organicism is a philosophy that asks questions about the nature of the organism and only indirectly promises practical biological control, as the organizer project itself demonstrated. Waddington sensed the incongruity of this kind of philosophy with the state of biology in the second half of the century: ‘reductionism is a recipe for action’.Footnote 131 When what is called for is making a ‘quick (scientific) buck by discovering some useful practical information’, the reductionistic mechanism embraced by Crick and Astbury trumped organicism. By the latter half 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 proportional to the precision with which it can be defined in molecular or atomic terms’.Footnote 132 Organicism, he protested, was the ‘method for making major advances in human comprehension’, like those of ‘Darwin, Freud, Einstein [and] the quantum physicists’. On the other hand, reductionistic mechanism promised predictability and 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 a memorable acronym, ‘COWDUNG’.Footnote 133

In this essay, I have attempted to ground the history of organicism in the twentieth century in a shifting social network. In some ways, the resulting picture reflects the metaphor that Needham, Waddington and, indeed, Whitehead suggested applied to the universe as a whole: that of a river. A concept runs its particular course because of social networks of adherents: scientists gathered in pools, sometimes mere eddies, discussing and applying their philosophy to their biology. Once viewed through the lens of these social networks, the fate of any particular set of experiments in the larger research stream has less to do with macroscopic social or conceptual factors than with microscopic ones. For instance, the apparent failure of the organizer project was not a stage in the conquest of vitalism. Rather, political intrigue at Cambridge, funding priorities at the Rockefeller Foundation, Needham's political interests, and the financial problems of Waddington had more to do with its demise. If the concept of organicism seemed to suffer the same fate as the organizer project, that also had less to do with its misapplied association with fascism, for instance, than with the character of the social group supporting it. Though organicism did not disappear to be rediscovered concomitantly with evo-devo, a plurality of factors weakened the social network of adherents. Molecular biology did not ‘rise’ to displace organicism, but the simplicity and rhetorical power of the progress-through-devitalizing-biology narrative meant that organicists had to carefully qualify their dissent.

Footnotes

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. 13–38. See also Horder, Timothy J., ‘The organizer concept and modern embryology: Anglo-American perspectives’, International Journal of Developmental Biology (2001) 45, pp. 97–132.

6 De Robertis, E.M., ‘Spemann's organiser and self-regulation in amphibian embryos’, Nature Reviews Molecular Cell Biology (2006) 7, pp. 296–302, 297, argues that works by Barth, L.G. (‘Neural differentiation without organizer’, Journal of Experimental Zoology (1941) 87, pp. 371–383) 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. 195–197) and Ficq, A. (‘Analyse de l'induction neurale par autoradiographie’, Experientia (1954) 10, pp. 20–21) 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. 261–283. 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. 211–230).

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. 355–358; and Lillie, Ralph S., ‘The philosophy of biology: vitalism versus mechanism’, Science, new series (1914) 40, pp. 840–846.

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. 1067–1074.

13 Gilbert, Scott F. and Sarkar, Sahotra, ‘Embracing complexity: organicism for the twenty-first century’, Developmental Dynamics (2000) 219, pp. 1–9, 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. 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 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. 1–70; 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. 159–180; Gilbert, Scott F., ‘Epigenetic landscaping: Waddington's use of cell fate bifurcation diagrams’, Biology and Philosophy (1991) 6, pp. 135–154; 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. 729–737; Hall, Brian K., ‘Waddington's legacy in development and evolution’, American Zoologist (1992) 32, pp. 113–122; 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. 5–18; Smocovitis, V. Betty, ‘Serious matters: on Woodger, positivism, and the evolutionary synthesis’, Biology and Philosophy (2000) 15, pp. 553–558; 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. 147–154.

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. 108–116, 108.

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

20 Russell, E.S., The Interpretation of Development and Heredity: A Study in Biological Method, Oxford: Clarendon Press, 1930, pp. 306–307.

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. 1–5; and Cross, Steven J. and Albury, William R., ‘Walter B. Cannon, L.J. Henderson, and the organic analogy’, Osiris (1987) 3, pp. 165–192.

24 E.g. Nordenskiöld, Erik, The History of Biology: A Survey (tr. L.B. Eyre), New York and London: A.A. Knopf, 1928, pp. 525–527; 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. 91–103.

27 Whitehead, Alfred North, 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 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. 377–391, 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. 216–229.

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, http://nobelprize.org/nobel_prizes/medicine/laureates/1935/spemann-lecture.html, 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. 231–243, 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. 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 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. 421–475; and Witkowski, J.A., ‘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 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. 1–20, 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. 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 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. 221–226.

40 Woodger, J.H., 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 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. 1–6; 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. 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 von Bertalanffy, Ludwig, ‘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.

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. 1–22; Woodger, , ‘The “concept of organism” and the relation between embryology and genetics, part II’, Quarterly Review of Biology (1930) 5, pp. 438–463; and Woodger, op. cit. (42).

51 Needham, Joseph, 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.

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. 1–3.

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. 193–206, 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. 357–374), 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. 575–622, 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. 889–895.

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. 275–308, 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. 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 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. 259–284.

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. 55–60.

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. 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 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. 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 Gerhart, John, ‘Johannes Holtfreter: 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 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. 74–79. 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. 183–209; 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. 315–348, p. 340; Gilbert, Scott F., Opitz, J.M. and Raff, Rudolph A., ‘Resynthesizing evolutionary and developmental biology’, Developmental Biology (1996) 173, pp. 357–372, 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. 175–180.

107 Waddington, C.H., ‘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 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. 301–320.

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. 66–67; and Mayr, , ‘Where are we?’, Cold Spring Harbor Symposia on Quantitative Biology (1959) 2, 1–14. 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. 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 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. 187–196; Wake, D.B., Roth, G. and Wake, M.H., ‘The problem of stasis in organismal evolution’, Journal of Theoretical Biology (1983) 101, pp. 211–224; 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. 202–203. 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. 1327–1330, 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. 277–317.

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. 20–21, 21.

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