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Animal agency in the age of the Modern Synthesis: W.H. Thorpe's example



The mechanical and reductive ideals of much of modern science leave it ill-equipped to recognize, let alone account for, the agency of animals. So says a tradition of criticism well represented in the writings of the British behavioural biologist W.H. Thorpe FRS (1902–1986). This paper recovers the range of overlapping debates and developments, philosophical and religious as well as scientific, which led Thorpe to champion animal agency in the period and place much better known now as headquarters for the neo-Darwinian Modern Synthesis. A retracing of Thorpe's path through such forgotten domains as applied-science Lamarckism and the philosophical psychology of Leonard Hobhouse complicates a now-standard picture (which Thorpe himself promoted) of the Modern Synthesis as inimical to animal agency. Largely thanks to Thorpe's work, as publicized by Julian Huxley, the Modern Synthesis revitalized the fortunes of what became one of the mainstays of agential science, the Baldwin effect.

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1 Thorpe, W.H., Science, Man and Morals, London: Scientific Book Club, 1965, pp. 1516 .

2 Thorpe, W.H., Evolution and Christian Belief, London: British Social Biology Council, 1951, p. 11 . Also published as an article in Biology and Human Affairs (1951) 17, pp. 619, 16.

3 Thorpe, W.H., The Origins and Rise of Ethology: The Science of the Natural Behaviour of Animals, London: Heinemann Educational, 1979, pp. 2930 . On the 1927 congress see South-Eastern Union of Scientific Studies: Annual Congress’, Nature (1927) 119, pp. 874875 .

4 Thorpe, op. cit. (3), pp. 26–29, 26; Radick, Gregory, ‘Morgan, Conwy Lloyd (1852–1936)’, in Lightman, Bernard (ed.), Dictionary of Nineteenth-Century British Scientists, vol. 3, Bristol: Thoemmes Continuum, 2004, pp. 14251426 .

5 C. Lloyd Morgan, ‘Territory in bird life’, Transactions of the South-Eastern Union of Scientific Societies (1927), pp. 22–26. Of Howard, Morgan wrote, Few equal him in accuracy of observation, in plain-tale description, and in cautious interpretation with due regard to life-story and to mind-story.’ Morgan, Life, Mind and Spirit, London: Williams and Norgate, 1926, p. 154 . On Morgan and Howard see Burkhardt, Richard W. Jr, Patterns of Behavior: Konrad Lorenz, Niko Tinbergen, and the Founding of Ethology, Chicago: The University of Chicago Press, 2005, pp. 9497 .

6 Morgan, C. Lloyd, Emergent Evolution, London: Williams and Norgate, 1923 ; Morgan, Life, Mind and Spirit, op. cit. (5), esp. pp. 37 ff. for the animal behaviour ‘-story’ distinctions, 61 for the emergentist gloss on the canon, and x for ‘Divine Purpose’. On Morgan's writings in the emergentist tradition see Blitz, David, Emergent Evolution: Qualitative Novelty and the Levels of Reality, Kluwer: Dordrecht, 1992, Chapters 6–9.

7 Thorpe, op. cit. (3), p. 29. For Thorpe's biography see Radick, Gregory, ‘Thorpe, William Homan’, in Noretta Koertge (ed.), New Dictionary of Scientific Biography, vol. 7, Detroit: Charles Scribner's Sons, 2008, pp. 4245 ; and more extensively Hinde, R.A., ‘William Homan Thorpe 1902–1986’, Biographical Memoirs of Fellows of the Royal Society (1987) 33, pp. 620629 .

8 On Kammerer's lecture see Koestler, Arthur, The Case of the Midwife Toad, London: Picador, 1971, Chapter 7; for Thorpe's testimonial see p. 70. On MacBride's Lamarckism see Peter Bowler, J., ‘E.W. MacBride's Lamarckian eugenics and its implications for the social construction of scientific knowledge’, Annals of Science (1984) 41, pp. 245260, esp. 247–250.

9 E.W. MacBride, ‘The nature and origins of mutations’, Transactions of the South-Eastern Union of Scientific Societies (1927), pp. 13–21.

10 W.H. Thorpe, ‘The fauna of brackish pools of the Sussex coast’, Transactions of the South-Eastern Union of Scientific Societies (1927), pp. 27–34.

11 Cf. Lamb, Marion J., ‘Attitudes to soft inheritance in Great Britain, 1930s–1970s’, in Gissis, Snait and Jablonka, Eva (eds.), Transformations of Lamarckism: From Subtle Fluids to Molecular Biology, Cambridge, MA: MIT Press, 2011, pp. 109120, esp. 110; Bowler, Peter J., The Eclipse of Darwinism: Anti-Darwinian Evolution Theories in the Decades around 1900, Baltimore: Johns Hopkins University Press, 1983, pp. 98106 ; and Burkhardt, Richard W. Jr, ‘Lamarckism in Britain and the United States’, in Mayr, Ernst and Provine, William B. (eds.), The Evolutionary Synthesis: Perspectives on the Unification of Biology, Cambridge, MA: Harvard University Press, 1980, pp. 343352 . An excellent starting point for anyone wishing to explore the depth and breadth of scientific interest in Lamarckian inheritance c.1930 is the discussion in Levins, Richard and Lewontin, Richard, ‘The problem of Lysenkoism’, in Levins and Lewontin, The Dialectical Biologist, Cambridge, MA: Harvard University Press, 1985, pp. 163196, 176179 .

12 I am indebted in this section to the pioneering coverage of this period of Thorpe's career in Burkhardt, op. cit. (5), pp. 339–341.

13 Thorpe, W.H., ‘Biological races in Hypomeuta padella L.’, Journal of the Linnean Society of London, Zoology (1929) 36, pp. 621634, 633; Harrison, J.W. Heslop, ‘Experiments on the egg-laying instincts of the sawfly, Potania salicis Christ., and their bearing on the inheritance of acquired characters; with some remarks on a new principle of evolution’, Proceedings of the Royal Society B (1927) 101, pp. 115126 .

14 Thorpe, W.H., ‘Biological races in insects and allied groups’, Biological Reviews of the Cambridge Philosophical Society (1930) 5, pp. 177212 , 208, quoting from Thompson, W.R. and Parker, H.L., ‘Host selection in Pyrausta nubialis, Hübn.’, Bulletin of Entomological Research (1928) 18, pp. 359364, 360.

15 Thorpe, W.H., ‘William Robin Thompson 1887–1972’, Biographical Memoirs of Fellows of the Royal Society (1973) 19, pp. 654678, esp. 661–662. Science and Common Sense was published in 1937.

16 Anna M. Bidder, ‘William Thorpe’, The Friend, 16 May 1986, pp. 621–622, 621. Bidder says (at 621) that Thorpe applied for membership in the Friends in 1945, and Hinde, in his obituary, op. cit. (7), p. 630, that fifteen years before Thorpe had begun attending meetings while working at Farnham Royal.

17 Letters from E.W. MacBride to W.H. Thorpe, 21 April and 1 May 1931, in the Papers of W.H. Thorpe, Manuscripts Reading Room, Cambridge University Library, Add.8784, Box (aka ‘Set’) 6, ‘M’ sheaf. The lice experiments were reported in Nuttall, G.H.F., ‘The systematic position, synonymy and iconography of Pediculus humanus and Phthirus pubis ’, Parasitology (1919) 11, pp. 329346 .

18 MacBride, E.W., ‘Habit: the driving force of evolution’, Supplement to Nature (20 June 1931) 127, pp. 933944 , 942 on Thorpe; 940 on the differences between Lamarck's views and Neo-Lamarckism. Editorial quotations from the first item in the ‘News and views’ section, Nature, 20 June 1931, p. 946.

19 J.B.S. Haldane, ‘The hereditary transmission of acquired characters’, Nature, 4 and 11 June 1932, pp. 817–819, 856–858. For Fisher's critical remarks on Lamarckism around the same time see, for example, R.A. Fisher, ‘Inheritance of acquired characters’, Nature, 15 October 1932, p. 579; Fisher, Indeterminism and natural selection’, Philosophy of Science (1934) 1, pp. 99117, esp. 109–112; and Fisher, Adaptation and mutations’, School Science Review (1934) 1, pp. 294301, esp. 295–297 – all gathered in Collected Papers of R.A. Fisher, vol. 3 (ed. Bennett, J.H.), Adelaide: University of Adelaide, 1973 .

20 Thorpe, W.H. and Jones, F.G.W., ‘Olfactory conditioning in a parasitic insect and its relation to the problem of host selection’, Proceedings of the Royal Society B (1937) 124, pp. 5680, 78. On how J.M. Baldwin rather than Morgan or H.F. Osborn came to be remembered as the discoverer of organic selection see Richards, Robert J., Darwin and the Emergence of Evolutionary Theories of Mind and Behavior, Chicago: The University of Chicago Press, 1987, pp. 480495 .

21 See, for example, Thorpe, W.H., ‘Ecology and the future of systematics’, in Huxley, Julian (ed.), The New Systematics, London: Oxford University Press, 1940, pp. 341364 ; Thorpe, Animal learning and evolution’, Nature (14 July 1945) 156, p. 46 ; Thorpe, The evolutionary significance of habitat selection’, Journal of Animal Ecology (1945) 14, pp. 6770 .

22 In a footnote in his 1930 paper, Thorpe observed that while the term ‘biological races’ was better than ‘physiological races’ for capturing the sense of differences based in preferences rather than structures, he followed his boss Thompson in regarding ‘psychological’ as better still, especially for insects; Thorpe, op. cit. (14), p. 180.

23 Letter from W.H. Thorpe to J.S. Whale, 22 September 1941, Thorpe Papers, Box 8, Bundle i, Folder 1; Whale, J.S., Christian Doctrine, Cambridge: Cambridge University Press, 1941 . I have not identified the source of the quotation from Hobhouse (if indeed it is a quotation), but it is in keeping with the discussion in his Morals in Evolution, 2 vols., London: Chapman & Hall, 1906, vol. 2, pp. 127141 .

24 Thorpe, W.H., Animal Nature and Human Nature, London: Methuen, 1974, pp. 349, 45. See too, for example, Thorpe, Biology and the Nature of Man, London: Oxford University Press, 1963, pp. 54, 109; Thorpe, Biology, Psychology and Belief, Cambridge: Cambridge University Press, 1961, p. 44 ; and as discussed in the next section, Thorpe, op. cit. (2), pp. 8 (pamphlet), 13 (article). On Hobhouse see Radick, Gregory, The Simian Tongue: The Long Debate about Animal Language, Chicago: The University of Chicago Press, 2007, pp. 211214 ; and Renwick, Chris, British Sociology's Lost Biological Roots: A History of Futures Past, London: Palgrave Macmillan, 2012, esp. pp. 98120, 170180 .

25 Hobhouse, L.T., Mind in Evolution, London: Macmillan, 1901 ; Radick, op. cit. (24), pp. 214, 222–223. Morgan later credited Hobhouse and successors such as Köhler with having shown that apes and to a lesser extent monkeys had reached a higher mental level of ‘sub-rational’ foreplanning. Morgan, Life, Mind and Spirit, op. cit. (5), pp. 211–212.

26 Hobhouse, L.T., Development and Purpose: An Essay towards a Philosophy of Evolution, London: Macmillan, 1913, pp. xxvi, 371.

27 On hormism see Morgan, Life, Mind and Spirit, op. cit. (5), pp. 87–93; also Flugel, J.C., A Hundred Years of Psychology 1833–1933, London: Duckworth, 1933, pp. 270278 . As Flugel stressed, hormic psychology was identified most closely with McDougall, who was increasingly well known for Lamarckism-affirming learning experiments with rats (see ibid., p. 278).

28 Letter from W.H. Thorpe to J.S. Whale, 22 September 1941, Thorpe Papers, Box 8, Bundle i, Folder 1, emphases in original. On Thorpe's animal welfare work see Wilson, David A.H., ‘Animal psychology and ethology in Britain and the emergence of professional concern for the concept of ethical cost’, Studies in History and Philosophy of Biological and Biomedical Sciences (2002) 33, 235–261, esp. 249–251, 256.

29 Thorpe, W.H., ‘Types of learning in insects and other arthropods’ (I), British Journal of Psychology (1943) 33, pp. 220235, esp. 222–224, 224; Thorpe, Types of learning in insects and other arthropods’ (II–III), British Journal of Psychology (1944) 34, 20–31, 6676 .

30 Thorpe, W.H., ‘A type of insight learning in birds’, British Birds (1943) 37, pp. 2931, 29; with a follow-up paper the next year: Thorpe, Further notes on a type of insight learning in birds’, British Birds (1944) 38, pp. 4649 .

31 R.A. Fisher, Creative Aspects of Natural Law, Fourth Annual Arthur Stanley Eddington Memorial Lecture, Cambridge: Cambridge University Press, p. 19. On the place of this lecture within Fisher's oeuvre see Hodge, M.J.S., ‘Biology and philosophy (including ideology): a study of Fisher and Wright’, in Hodge, Before and after Darwin: Origins, Species, Cosmogonies, and Ontologies, Aldershot: Ashgate, 2008, XIII, esp. pp. 256262 . From the beginning of the Eddington lecture series in 1946 through to 1975, Thorpe was chairman of the trustees; see Hinde, op. cit. (7), p. 631.

32 Thorpe, op. cit. (2), pp. 8–9, 11 (pamphlet), 13–14, 16 (article); Huxley, T.H. and Huxley, Julian, Evolution and Ethics, 1893–1943, London: Pilot Press, 1947 .

33 Letter from Charles Raven to W.H. Thorpe, 12 January 1951, Thorpe Papers, Box 8, Bundle i, Folder 1, emphasis in original, quoting (inexactly) from Thorpe, op. cit. (2), pp. 5 (pamphlet), 10 (article).

34 Letter from Herbert G. Wood to W.H. Thorpe, 6 November 1951, Thorpe Papers, Box 6, ‘W’ sheaf, quoting from Thorpe, op. cit. (2), pp. 4, 9 (pamphlet), 9, 14 (article). Even if Thorpe's lecture failed to satisfy all comers, it did him no harm professionally, at least to judge by his election in 1951 to the Fellowship of the Royal Society.

35 Thorpe, W.H., Learning and Instinct in Animals, London: Methuen, 1956 .

36 Waddington, C.H., The Nature of Life, London: George Allen & Unwin, 1961, p. 90 . Thorpe took extensive notes (preserved in the archive: Thorpe Papers, Box 8, Bundle ii, Folder 1) on Waddington's book and cited it in Science, Man and Morals. On Waddington's studies in the 1940s and 1950s of what he called ‘genetic assimilation’, but widely interpreted ever since as a variant of the Baldwin effect, see Hall, Brian K., ‘Baldwin and beyond: organic selection and genetic assimilation’, in Weber, Bruce H. and Depew, David J., Evolution and Learning: The Baldwin Effect Reconsidered, Cambridge, MA: MIT Press, 2003, Chapter 8.

37 Polanyi, Michael, Personal Knowledge: Towards a Post-critical Philosophy, Chicago: The University of Chicago Press, 1962 (first published 1958), pp. 382404 ; Thorpe, op. cit. (1), pp. 20 ff. Documents preserved in the archive show that Thorpe and Polanyi spoke in the same session at the 1959 annual conference of the British Society for the Philosophy of Science and corresponded in the summer of 1962 about emergence in biology; Thorpe Papers, Box 8, Bundle ii, Folder 1. On Thorpe on emergence see Gillespie, Neal C., ‘The interface of natural theology and science in the ethology of W.H. Thorpe’, Journal of the History of Biology (1990) 23, pp. 138, 810 .

38 Thorpe, W.H., ‘Molecules and evolution’, Nature (1966) 210, pp. 663664, 664.

39 Gillespie, op. cit. (37), 35.

40 See, in addition to Hodge, op. cit. (31), Moore, James R., ‘R.A. Fisher: a faith fit for eugenics’, Studies in History and Philosophy of Biological and Biomedical Sciences (2007) 38, pp. 110135 .

41 ‘Protest’ was a category invoked at the time, for example by Morgan in Emergent Evolution, op. cit. (6), pp. 7–8: ‘the whole doctrine of emergence is a continued protest against mechanical interpretation, and the very antithesis to one that is mechanistic’.

42 I first considered Lamarckism along these lines in a review essay, Radick, Gregory, ‘Deviance, Darwinian style’, Metascience (2005) 14, pp. 453457 .

43 Koestler, Arthur, Janus: A Summing Up, London: Hutchinson, 1978 , quotation from Thorpe on p. 165. On Koestler's admiring relations with the anti-reductionist scientists of the era – many of them gathered in 1968 in Alpbach for the famous Beyond Reductionism symposium, which Thorpe chaired (the quotation is from his opening remarks) – see Stark, James F., ‘Anti-reductionism at the confluence of philosophy and science: Arthur Koestler and the biological periphery’, Notes and Records of the Royal Society (2016) 70, pp. 269286 .

44 See, for example, Lewontin, Richard C., ‘Gene, organism, and environment’, in Bendall, D.S. (ed.), Evolution from Molecules to Men, Cambridge: Cambridge University Press, 1985, pp. 273285 ; and more expansively Lewontin, The Triple Helix: Gene, Organism, and Environment, Cambridge, MA: Harvard University Press, 2000 . Niche constructionism and developmental systems theory belong to the same family of criticism. Recent indictments of the Modern Synthesis in this tradition include Corning, Peter A., ‘Evolution “on purpose”: how behaviour has shaped the evolutionary process’, Biological Journal of the Linnean Society (2014) 112, pp. 242260 and Walsh, D.M., Organisms, Agency, and Evolution, Cambridge: Cambridge University Press, 2015 .

45 Simpson, G.G., ‘The Baldwin effect’, Evolution (1953) 7, pp. 110117 ; Huxley, Julian, Evolution: The Modern Synthesis, London: Allen & Unwin, 1942, p. 524 .

46 On the Baldwin effect's scientific trajectory see the papers gathered in Weber and Depew, op. cit. (36); and also a review essay on the volume by Sterelny, Kim, ‘Baldwin effects’, Evolution and Development (2004) 6, pp. 295300 .

47 On Thorpe and the Baldwin effect see the brief but excellent discussion in Gillespie, op. cit. (37), pp. 30–31. Oddly, the otherwise comprehensive Weber and Depew volume, op. cit. (36), includes just one minor and misleading mention of Thorpe (p. 143).

48 Huxley recounted the origins of what became Evolution: The Modern Synthesis in Huxley, op. cit. (45), p. 7. Correspondence from March 1938 preserved in the Julian Huxley archive at Rice University, Texas, shows that Thorpe was already at work then on the chapter for Huxley's New Systematics volume, published as Thorpe, ‘Ecology’, op. cit. (21). Many thanks to Emily Herring for sharing copies of these letters with me.

49 Huxley, op. cit. (45), pp. 295–308, 296.

50 Thorpe, ‘Animal learning’, op. cit. (21); Thorpe, ‘Evolutionary significance’, op. cit. (21) (the former is a precis of the latter). For Thorpe's sense of the paper's importance, in explaining how ‘recent developments in genetics and in the study of animal learning make the Baldwin concept at once more probable and easier to understand’, see Thorpe, op. cit. (35), pp. 256–258, 257. Simpson, distinguishing an animal's selecting of its environment from the Baldwin effect strictu sensu, cited Thorpe's paper as linking the two by showing how ‘[s]election of the environment may … be a first step in the Baldwin effect.’ Simpson, op. cit. (45), p. 111.

51 The pluralism of the Modern Synthesis has been a general theme in the historiography of recent decades, beautifully encapsulated in David Depew and Bruce Weber's line about the Synthesis being ‘more like a treaty than a theory’. Depew, David and Weber, Bruce, Darwinism Evolving: Systems Dynamics and the Genealogy of Natural Selection, Cambridge, MA: MIT Press, 1995, p. 300 . On behaviour's lack of full integration into the Synthesis, the prize one-liner is considerably older. In a 1957 review of Thorpe's Learning and Instinct in Animals, the Glasgow zoologist S.A. Barnett wrote, ‘It was once said of certain investigations of population dynamics that the animals concerned did not behave at all: they merely numbered off from the right.’ Barnett, S.A., ‘The new ethology’, New Biology (1957) 24, pp. 118124, 118, emphasis in original.

I am deeply grateful to Mandy Rees for inviting this contribution, for gently but insistently prodding it into being, and for arranging an anonymous review from which I learned a great deal. It is also a pleasure to record my thanks to Jon Hodge, Chip Burkhardt and Emily Herring for much-appreciated discussion and encouragement along the way.


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