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Coadaptation and the Inadequacy of Natural Selection

Published online by Cambridge University Press:  05 January 2009

Mark Ridley
Mark Ridley
Affiliation:
Department of Zoology, South Parks Road, Oxford, OX1 3PS.
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Extract

When Charles Darwin published his theory in 1859 the biological community gave very different receptions to the idea of evolution and to the theory of natural selection. Evolution was accepted as widely and rapidly as natural selection was rejected. Most biologists were ready to accept that evolution had occurred, but not that natural selection was its cause. They preferred other explanations of evolution, such as theories of big directed variation, or admitted that they did not know its cause. Darwin himself never maintained that natural selection was the sole cause of evolution. He thought of it as one among several causes, and did not specify how much evolution had occurred by the natural selection of fortuitous variations, and how much by other factors such as the inherited effects of use and disuse. However, Darwin did maintain that natural selection was in principle capable of explaining all the observed properties of organisms. He did not think that there were some characteristics that were particularly likely to have evolved by natural selection, and other kinds that were not. Against this, many of this critics thought that there were characteristics that natural selection was particularly powerless to explain. Thus it could not account for characteristics that were detrimental, or those that seemed useless (such as species differences), or those that were of too little importance for natural selection to have favoured them. There were also characteristics of such complexity that it was unimaginable that natural selection could have built them up in tiny stages from fortuitous, undirected variants. The present essay will be concerned with just one, the last mentioned, of these kinds of characteristics.

Type
Research Article
Information
The British Journal for the History of Science , Volume 15 , Issue 1 , March 1982 , pp. 45 - 68
Copyright
Copyright © British Society for the History of Science 1982

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References

Based on the essay which was awarded the Society's Singer Prize for 1980.

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22 Russell, , op. cit. (14), pp. 232–41, quote from p. 241.Google Scholar There is truth in Russell's judgement although I would question whether it was a ‘drawback’ (p. 232) in Darwin's thought.

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31 Peckham, , op. cit. (20), pp. 242–3.Google Scholar Also Darwin, to Wallace, , 30.i.1871Google Scholar, in Darwin, , op. cit. (4), vol. iii, pp. 135–6.Google Scholar

32 On the extent of variation, further see: Darwin to Lyell, Darwin, , op. cit. (4), vol. ii, p. 388Google Scholar; Rudwick, , op. cit. (2), p. 203Google Scholar: Schweber, S. S., ‘The young Darwin’, Journal of the history of biology, 1979, 12, pp. 185–6.CrossRefGoogle Scholar

33 Darwin, & Seward, (eds), op. cit. (16), vol. i, p. 145.Google Scholar Darwin here, as elsewhere, is using coadaptation as a synonym of adaptation rather than in the more specific sense of an adaptation in which the parts are necessarily mutually adjusted.

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35 Spencer did not mention Weismann in Factors. According to Spencer's biographer, ‘The earliest notice of Dr. Weismann to be found in the correspondence is in a letter to Mr. Henry Collins (26 February, 1890)’, Duncan, D. (ed. ), Life and latters of Herbert Spencer, London, 1908, p. 343.Google Scholar See also G.J. Romanes to his wife, 21.iii.1893, in Romanes, E. (ed.), Life and letters of George John Romanes, London, 1896, p. 301.Google Scholar

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37 Turner, F. M., Between science and religion, New Haven, 1974, pp. 134–63Google Scholar on Romanes, quotations from p. 142. ‘Physicus’ [Romanes, G. J.], A candid examination of theism, London, 1878; p. 171Google Scholar illustrates splendidly Romanes' beliefs about the relation of morality and evolution.

38 Wallace, A. R., Darwinism, London, 1889, p. 127Google Scholar for covariation, p. 418 for quote; Wallace, A. R., ‘The origin of species controversy’, Nature, 1869, 105–7 and 132–3CrossRefGoogle Scholar, for gradual evolution of eye see p. 106, for artificial selection analogy pp. 106–7. Also see Wallace to Darwin quoted in Hull, , op. cit. (6), p. 302Google Scholar and Darwin, to Wallace, , op. cit. in (31).Google Scholar

39 For Wallace's non-use of the analogy in his original discovery, see Limoges, C., La sélection naturelle, Paris, 1970, pp. 96–9Google Scholar and Bowler, P. J., ‘Alfred Russel Wallace's concepts of variation’, Journal of the history of medicine, 1976, 31, p. 24.Google ScholarPubMed Quote from Wallace, A. R., ‘On the tendency of varieties to depart indefinitely from the original type’, Journal of the proceedings of the Linnean Society, 1859, 3, p. 61.Google Scholar For other examples see Young, R. M., ‘Darwin's metaphor: Does nature select?’, The monist, 1971, 55, pp. 471–3CrossRefGoogle Scholar and Wallace, , Darwinism, op. cit. (38), p. vi.Google Scholar

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41 A point that is further illustrated by the exchange on coadaptation between Raphael Meldola and Romanes in Nature in 1891. Meldola argued that coadaptations evolve in small stages; Romanes accepted this as the only way of reconciling natural selection with coadaptation. The exchange was prolonged by Romanes’ curious conceit (which he cannot have read) that Meldola was repeating Wallace's artificial selection analogy. The letters, mainly entitled ‘Coadaptation’, were in Nature, 1891, 43, 409–12, 489–90, 557–8, 582–3, 44, 7, 28–9, 55.Google Scholar See also Wallace to Meldola in Marchant, , op. cit. (24), vol. ii, pp. 50–1.Google Scholar Meldola, a professor of chemistry and expert on dyes, was also an entomologist, President of the Royal Entomological Society, and translator of Weismann. He held similar views on evolution to Wallace and Weismann. He was passed over by DSB and DNB; the best biographical source is Marchant, J. (ed.), Raphael Meldola, London, 1916.Google Scholar At about this time, Ball, W. P., Are the effects of use and disuse inherited?, London, 1890, pp. 19–20Google Scholar, discussed Spencer on coadaptation and concluded that coadaptations could be built up in small stages by natural selection.

42 Spencer, H., ‘The inadequacy of “natural selection”’, Contemporary review, 1893, 63, 153–66 and 439–56, see pp. 159–60.Google Scholar (This and his subsequent papers in the controversy were republished as separate pamphlets and also in the 2nd edn of Principles of biology, London, 1898, vol. i, pp. 602–91.Google Scholar) Wallace, A. R., ‘Are individually acquired characters inherited?’, Fortnightly review, 1893, 59 (O.S., 53 N.S.), p. 659Google Scholar (original italics), 660, 660–1 (original italics). Wallace introduced this paper as being jointly inspired by Weismann's The germ plasm, Spencer's ‘Inadequacy of “natural selection”’ and a work by the American neo-Lamarckian, C. A. Keeler (who Wallace quotes as approving of Spencer's coadaptation argument).

43 Churchill, F. B., ‘The Weismann–Spencer controversy over the inheritance of acquired characters’, in Forbes, E. G. (ed.), Human implications of scientific advance (Proceedings of the XVth International Congress of the History of Science), Edinburgh, 1978, pp. 451–68Google Scholar; Blacher, L. J., ‘Die Diskussion zwischen Spencer und Weismann über die Bedeutung der natürlichen Zuchtwahl and der drekten Anpassung für die Evolution’, NTM-Schriftenreihe Geschichle, 1973, 2, 50–8.Google Scholar

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48 Ball, W. P., ‘Natural selection and Lamarckism’, Natural science, 1893, 2, p. 347Google Scholar; this paper was another of the replies to Spencer's ‘Inadequacy’.

49 Weismann, , op. cit. (45), p. 328Google Scholar (original italics).

50 Spencer, H., ‘A rejoinder to Professor Weismann’, Contemporary review, 1893, 64, 893–912Google Scholar; Ball, W. P., ‘Neuter insects and Darwinism’, Natural science, 1894, 4, 281–9Google Scholar; Weismann, A., The effect of external influences on development, London, 1894, p. 68Google Scholar; Spencer, , ‘Weismannism’, op. cit. (24), pp. 597–9, 603Google Scholar; Weismann, A., ‘Heredity once more’, Contemporary review, 1895, 68, p. 445.Google Scholar

51 Published as Weismann, , Effect, op. cit. (50)Google Scholar. G. J. Romanes founded the lecture in 1892, the selection of lecturers to be ‘determined with reference either (a) to general eminence in art, literature, or science, or (b) to special claims for discussing any particular subject of high interest at the time.’ (Founder's Preface). The 1893 lecture was Huxley's classic Evolution and ethics. Romanes died soon after Weismann delivered his lecture. The lecture is now given annually by someone from the posher echelons of the artistic establishment.

52 Roux, W., Der Kampf der Theile im Organismus, Leipzig, 1881.Google Scholar Roux sent Darwin a copy, of which Darwin wrote to Romanes: ‘As far as I can imperfectly judge, it is the most important book on Evolution which has appeared for some time. I believe that G. H. Lewes hinted at the same fundamental idea.…’ (Darwin, , op. cit. (4), vol. iii, p. 244Google Scholar, also see Darwin to Huxley in ibid, p. 119). Lewes was one of Spencer's closer friends, and they discussed the inheritance of acquired characters (Duncan, , op. cit. (35), p. 541).Google Scholar

53 Weismann, , Effect, op. cit. (50), pp. 17, 18, 18–19 (italics original).Google Scholar

54 Spencer, , ‘Weismannism’, op. cit. (24), p. 597.Google Scholar

55 Weismann, A., Neue Gedanken zur Vererbungsfrage, Jena, 1894Google Scholar; ‘On germinal selection’, The monist, 1896, 6, 250–93Google Scholar reprinted as a pamphlet (Chicago, 1896). Germinal selection is a precise application of intra-selection to the germ plasm, a possibility vaguely noted but not much explored in the Romanes lecture (Weismann, , Effect, op. cit. (50), pp. 20–1).Google Scholar

56 Weismann, , ‘Heredity’, op. cit. (50), p. 430.Google Scholar

57 Weismann, , ‘on germinal selection’, op. cit. (55), p. 264.Google Scholar The second passage was italicized in the original; it is a remarkable statement, Weismann not having admitted this before even as a possibility.

58 Weismann later accommodated the possibility of non-adaptive orthogenesis in a modified theory of germinal selection, on which see Bowler, P. J., ‘Thomas Eimer and orthogenesis: evolution by ‘definitely directed variations’, Journal of the history of medicine, 1979, 34, 40–73, especially pp. 55–8.Google Scholar Bowler tends to treat germinal selection as if it were solely a response to the debate on orthogenesis (e.g. p. 45), though it was also stimulated by the claim that natural selection could not explain useful structures. The precursor of germinal selection, intra-selection, was first proposed as a direct answer to the problem of coadaptation, in the 1894 Romanes lecture.

59 Spencer, H., Facts and comments, London, 1902, pp. 92–6CrossRefGoogle Scholar; and a short note, ‘Heredity once more’, Contemporary review, 1895, 68, p. 608.Google ScholarWeismann, A., Tne evolution theory, London, 1904, vol. ii, pp. 81–7Google Scholar, and, ‘The selection theory’ in Seward, A. C. (ed.), Darwin and modern science, Cambridge, 1909, pp. 32–42.Google Scholar

60 Robinson, G., ‘Weismann, August Friedrich Leopold’, in Gillespie, C. C. (ed.), Dictionary of scientific biography, New York, 1976, vol. xiv, p. 237.Google Scholar William James, by contrast, praised Spencer: ‘I may say that Spencer's controversy over use-inheritance with Weismann, entered into after he was sixty, seems to me in point of quality better than any other part of his work. It is genuine labor over a puzzle, genuine research.’ (‘Herbert Spencer’, The Atlantic monthly, 1904, 94, p. 107Google Scholar; reprinted in James, W., Memories and studies, London, 1911.)Google Scholar

61 Anonymous, ‘Weismann and Spencer’, Natural science, 1895, 7, p. 233.Google Scholar

62 Simpson, G. G., ‘The Baldwin effect’, Evolution, 1953, 7, 110–17CrossRefGoogle Scholar is an authoritative christening. The idea was first and most clearly stated by Spalding, D. A., ‘Instinet with original observations on young animals’, Macmillan's magazine, 1873, 27, 282–93Google Scholar, which was reprinted in the British journal of animal behaviour, 1954, 2, 2–11Google Scholar, with an introduction by J. B. S. Haldane (p. 1 ) drawing attention to Spalding's priority. See the passage about Robinson Crusoe's parrot on p. 11 of the reprint. Wallace, A. R., ‘Lamarck versus Darwin’, Nature, 1889, 40, p. 619CrossRefGoogle Scholar, also independently published the same idea. Neither Spalding nor Wallace applied the idea to the problem of coadaptation, and none of the authors that did seemed aware of Spalding's or Wallace's papers.

63 Morgan, C. L., Animal life and intelligence, London, 1891, p. 213.CrossRefGoogle Scholar In Morgan, C. L., ‘Factors in the evolution of the Mammalia’, Natural science, 1892, 1, pp. 100–1Google Scholar, he seems to be groping towards (if you'll pardon the anachronism) the Baldwin effect. Morgan, C. L., Habit and Instinet, London, 1896, ch. xiv.CrossRefGoogle ScholarRichards, R. J., ‘Lloyd Morgan's theory of Instinet: from Darwinism to neo-Darwinism’, Journal of the history of the behavioral sciences, 1977, 13, 12–323.0.CO;2-E>CrossRefGoogle ScholarPubMed, documents Lloyd Morgan's close knowledge of Spalding's work (p. 22) and his gradual conversion to Weismannism (pp. 26–30).

64 Wallace, A. R., ‘The problem of Instinct’, Natural science, 1897, 10, p. 168.Google Scholar

65 Baldwin mentioned the idea in various publications in 1895, and he collected the relevant quotations in Baldwin, J. M., ‘A new factor in evolution’, American naturalist, 1896, 30, 441–51 and 536–53CrossRefGoogle Scholar (I have quoted from p. 449). This paper along with all the previous ones was collected in Baldwin, J. M., Development and Evolution, New York, 1902Google Scholar, dedicated to Lloyd Morgan, Osborn and E. B. Poulton. Baldwin's contemporaries found him verbose and obsessively self-important. See Mueller, R. H., ‘A chapter in the history of the relationship between psychology and sociology in America: James Mark Baldwin’, Journal of the history of the behavioral sciences, 1976, 12, p. 2503.0.CO;2-H>CrossRefGoogle ScholarPubMed, where Cooley's advice to Baldwin is quoted: ‘If you wish to produce anything of lasting value, you must see to it that the subject matter, the truth, is the first interest of your mind, not your books, your essay, yourself as discoverer and Communicator of the truth.’ (original italics).

66 Osborn, H. F., ‘[A mode of evolution requiring neither natural selection nor the inheritance of acquired characters]’ Transactions of the New York academy of sciences, 1896, 15, 141–3 and 148Google Scholar; and ‘The limits of organic selection’, American naturalist, 1897, 31, 944–51, especially p. 950.Google Scholar ‘Across the flyleaf he [Osborn] wrote in his large and elegant hand: To George Gaylord Simpson with the appreciation and best wishes of Henry Fairfield Osborn. As this was in heavy, flowing black ink, I reached for a blotter, but he stayed my hand and said, “Never blot the signature of a great man.”’ (Simpson, G. G., Concession to the improbable, New Haven, 1978, p. 40.)Google Scholar

67 Osborn, , ‘Limits’, op. cit. (66), p. 950.Google Scholar

68 Baldwin, , ‘New factor’, op. cit. (65), p. 549.Google Scholar

69 For yet more authors on coadaptation, see Ellegård, A., Darwin and the general reader, Göteborg (Göteborgs Universitets Arskrift, Ixiv, no. 7), 1958, ch. 6 and pp. 247–50.Google Scholar