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Ernst Mayr’s interactions with J. B. S. Haldane

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Abstract

Ernst Mayr and J. B. S. Haldane, major contributors to the ‘modern synthesis’ in evolutionary theory, set an example of how scientific disagreements need not come in the way of friendship. After getting acquainted, they kept discussing issues related to evolution until just before Haldane’s death in 1964. Their dissimilar backgrounds meant that they adopted different approaches. A major disagreement emerged regarding the right way to look at the role of genes in evolution. Mayr felt that the elementary models of population genetics were oversimplifications and therefore inadequate for representing evolutionary processes, though he was not consistent in his attitude. Haldane, on the other hand, maintained that the mathematical treatment of simple models had an important role to play. The Mayr-Haldane interactions illustrate divergent viewpoints concerning the utility of mathematics in biology.

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Notes

  1. See for example the standard texts by Li (1955) or Futuyma (2013).

  2. Trivial in principle, that is. In practice, the combinatorial complexity increases rapidly with the number of loci; and, once recombination is considered, it becomes increasingly difficult to arrive at analytical solutions.

  3. The classical books by Fisher (1930/1958/1999) and Haldane (1932/1990) are full of examples. Among them, one might cite the results that in the absence of natural selection, variation would be lost very slowly in large populations and relatively rapidly in small populations, that recurrent mutation can allow genes with deleterious effects to persist in stable equilibria and that “a selective advantage of the heterozygote over both homozygotes will preserve both of a pair of allelomorphs indefinitely” (Haldane 1953). Haldane (1953) goes on to say: “Such ideas as these pass rapidly from being mathematical theorems to being common sense. Other ideas with a superficial appeal to common sense, for example that dominants must oust recessives, appear to require mathematical disproof to prevent their spread”, the last being a comment on what led to Hardy’s formulation of the celebrated Hardy–Weinberg principle (see Crow 1999).

  4. Nor—eventually—did W. H. Provine, historian of population genetics. In contrast to the favourable stand he had taken previously, his opinion as expressed in the 2001 re-issue of The Origins of Theoretical Population Genetics was “Now I see these theoretical models of the early 1930s, still widely used today, as an impediment to understanding evolutionary biology…” (Provine 1971/2001). Regarding “Evolutionary—or “Modern”—Synthesis”, see Provine (1980). The appropriateness of the term, and whether a new synthesis is necessary, or even under way today, has been debated.

  5. Mayr (1963, p. 263). The implied negative assessment was leavened with occasional positive statements. Earlier he had granted that beanbag genetics had made useful contributions, had been a ‘‘necessary step in the development of our thinking” and had “restored the prestige of natural selection’’ (Mayr 1959). But on the whole he seems to have thought that its contribution to evolutionary theory was questionable.

  6. Haldane (1964).

  7. A list is provided in Rao and Nanjundiah (2010/2011), p. 236; also see Sarkar (2005, 2007, 2013).

  8. According to SS “There is no explicit indication of why or when it was written”.

  9. For that matter, we have not tried to distinguish between them and other sources of genetic load either, e.g. the load due to recombination or segregation.

  10. Or, equivalently, “cost”. Mayr (1963, p. 253) says that it can be thought of as the price to be paid for “prevent[ing] the depletion of genetic variability” and “thwart[ing] the homogenizing tendencies of natural selection”. Slightly later he refers to “some six or seven kinds of genetic load”.

  11. Incidentally this illustrates Mayr’s essential reservation regarding ‘beanbag genetics’, namely that it glossed over the importance of interactions between alleles at different loci (also see later). VR/VN lists others who raised the same objection.

  12. In VR/VN the year of R. A. Fisher’s death is stated incorrectly as 1960; it should have been 1962. The (wrong) date was used to infer that because that Fisher was no longer alive when Mayr’s 1963 book appeared, he could not have responded to it. Fortunately the inference stands. Had he chosen to do so, Fisher could have responded to the ‘beanbag’ criticisms that Mayr had made before 1962. It would be interesting to learn whether he knew of Mayr’s talk at the 1959 Cold Spring Harbor Symposium.

  13. Ernst Mayr to Veena Rao, Museum of Comparative Zoology letterheads; M1 is typed and dated 18 March 2004 and M2 is handwritten with the date given as 28. Sept. 04.

  14. According to the biographical memoir of Mayr by Walter J. Bock, he was born on 5 July 1904 and died on 3 February 2005 (see http://www.nasonline.org/publications/biographical-memoirs/memoir-pdfs). VR/VN (page 240) errs in giving the dates as 1905–2005.

  15. Jepsen et al. (1949).

  16. See Cain and Mayr (2004).

  17. Except possibly in the case of Wright, Mayr’s acquaintance with the work of the population geneticists appears to have begun after they had begun to make substantial contributions. Prior to mentioning Haldane, he says (on the same page) “People have often asked me what impact Fisher, Haldane and Wright had on my thinking. My answer is quite embarrassing. I knew nothing of Fisher until I read Dobzhansky (1955), and even then I did not read Fisher’s book”; and further, “at the time [the impression one gets is that he means in 1931 or 1932] I had not yet heard of Wright and genetic drift”.

  18. Letter from Mayr to Haldane dated 16 March 1951 (No. HUGFP 74.7, Box 8, F384; Pusey Library, Harvard University Archives, Cambridge, MA, USA), see p. 242, footnote 25 in Rao and Nanjundiah (2010/2011).

  19. Ibid.

  20. Undated letter (inferred to be sent in summer 1951) from Haldane to Mayr (No. HUGFP 74.7, Box 9, F 406; Pusey Library, Harvard University Archives, Cambridge, MA, USA), see footnote 26 on p. 243 of Rao and Nanjundiah (2010/2011).

  21. Letter from Mayr to Haldane dated 26 September 1951 (No. HUGFP, 74.7, Box 9, F 406; Pusey Library, Harvard University Archives, Cambridge, Mass, USA), see footnote 27 on p. 243 of Rao and Nanjundiah (2010/2011).

  22. Letter from Haldane to Mayr dated October 3, 1951 (no. HUGFP 74.7, Box 9, F 406; Pusey Library, Harvard University Archives, Cambridge, Mass, USA).

  23. Mayr had told Haldane that he was going to “lecture to Buzzati-Traverso’s department” in the University of Pavia (footnote 25 on p. 242 of Rao and Nanjundiah 2010/2011).

  24. The proceedings were published in Buzzati-Traverso 1954. Nevertheless the 1953 Pavia Symposium is significant as a forerunner of the Mayr-Haldane dispute on the usefulness of population genetics calculations based on Mendelian genes. See discussion on p. 245 of Rao and Nanjundiah 2010/2011.

  25. Letter from Mayr to Haldane dated 13 April 1959 (No. HUGFP 74.7, Box 6, F 731; Pusey Library, Harvard University Archives, Cambridge, Mass., USA), footnote 29 on p. 244 of Rao and Nanjundiah (2010/2011).

  26. Letter from Haldane to Mayr dated 30 December 1959 (no. HUGFP 74.7, Box 6, F 731; Pusey Library, Harvard University Archives, Cambridge, Mass., USA), footnote 41 on p. 249 of Rao and Nanjundiah (2010/2011).

  27. Director, Anthropological Survey of India, January 1959–January 1964. See http://ignca.nic.in/Lectures_PDFs/NKBose/NKBose007a.pdf.

  28. It was the published version of Mayr’s 1959 Cold Spring Harbor presentation too that elicited a strong objection to the ‘beanbag’ characterisation from Wright (Wright 1960), providing further justification for highlighting the significance of 1959. The immediate provocation for Haldane’s counter-attack appears to have been Mayr’s 1963 book “Animal Species and Evolution”. Haldane cites it first in his paper, after which he moves on to the 1959 talk as “another place” in which “Mayr made a more specific challenge” (Haldane 1964).

  29. Basically, that no “noteworthy quantitative statements about evolution” had emerged from it. As well, “qualitatively new ideas” had been “very few”. See “The Dispute Anticipated” on p.244 of Rao and Nanjundiah, 2010/2011.

  30. Mayr (1959) cites with approval Waddington’s misgivings regarding classical population genetics, and it may be this that SS may be referring to in saying “the elaboration of Waddington’s views later in The Strategy of the Genes (1957) had a direct role in emboldening Mayr”. The information that Mayr had read Waddington’s critical remarks after they were repeated in 1957, and agreed with them, appears to have been communicated personally by Mayr to Sarkar (2007, p. 74, footnote 34).

  31. Buzzati-Traverso (ed.) (1954). Dobzhansky and Lerner spoke in the 1953 Pavia symposium; both questioned the utility of population genetics theory based on Mendelian genes. As we have mentioned, Mayr shared their negative opinion and mounted his own attack later. Besides Mayr and Haldane, Fisher too was at the Pavia symposium. Like them, he did not contribute to the published proceedings either. Even if he did not make a formal presentation, he had ample opportunity to intervene and rebut the contentions of Dobzhansky and Lerner. It would be worth investigating whether he did so. See footnote 12 here and pp. 245–246 and footnote 34 on p. 246 in Rao and Nanjundiah 2010/2011.

  32. See Appendix.

  33. Also stressed by VR/VN and SS.

  34. Three words immediately preceding “like Mahatma Gandhi”are scratched out in the original of M1. They appear to read “not a sati”. Perhaps initially Mayr meant to write “not a saint”.

  35. Evidently, by “all my life” Mayr means the time since he first got to know of Haldane’s work and also met him, which was in 1947 as mentioned. See note 17 and related text.

  36. See footnote 31.

  37. Mayr himself thought their approaches were harbingers of a new framework for evolutionary genetics that he called “the genetic ‘theory of relativity’” (Mayr 1959, alluding to Mayr 1955). Provine (2004) refers to the terms in the course of discussing what he calls Mayr’s “love/hate relationship with genetics”.

  38. Haldane (1964) and Wright (1960). Wright’s comments on Mayr’s 1959 address at the Cold Spring Harbor Symposium came the very next year in a review of the Symposium volume. They were worded strongly, beginning with “..he [i.e., Mayr] has seriously misinterpreted the roles of these various lines [of research in population genetics], as well as the contributions of the one with which I am most familiar”, and going on to detail the misrepresentations. Haldane’s stance overlapped with that of Wright but was not identical (Rao and Nanjundiah 2010/2011). Provine states that in a 1986 interview Mayr “regret[ted] classifying Wright as a beanbagger, except in his one-locus-two-allele models. His physiological genetics, emphasis upon gene interaction, and shifting balance theory all were more congenial to Mayr’s views” (Provine 2004). Haldane (1964, Defense) seems to have been unaware of Wright’s riposte. In the Defense Haldane cites and lauds Wright’s contributions extensively. Along the way he remarks “Wright is one of the gentlest men I have ever met, and if he defends himself, will not counterattack” (which, as the quotation from Wright (1960) shows, was not true) and “If I have not defended Sewall Wright, this is largely because I should like to read his defense”.

  39. The tensions are all the more sharp when they involve, as they do in this case, the traditionally non-overlapping fields of biology and mathematics (discussed in Nanjundiah 2005 and Rao and Nanjundiah 2010/2011, p. 272 onwards).

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Acknowledgments

VR thanks Ernst Mayr for graciously agreeing to provide his recollections of Haldane, and for interceding with the Harvard University Archives, Pusey Library, for permission to publish archival material. This contribution forms part of the research project titled ‘J. B. S. Haldane’s Indian Period’, supported by the Indian National Science Academy, New Delhi.

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Authors and Affiliations

  1. National Institute of Advanced Studies, Bangalore, 560012, India

    Veena Rao

  2. Stellenbosch Institute for Advanced Study (STIAS), Wallenberg Research Centre at Stellenbosch University, Marais Street, Stellenbosch, 7600, South Africa

    Vidyanand Nanjundiah

  3. Centre for Human Genetics, Electronic City (Phase I), Bangalore, 560100, India

    Vidyanand Nanjundiah

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  1. Veena Rao
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  2. Vidyanand Nanjundiah
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Correspondence to Veena Rao.

Appendix

Appendix

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    M1

  2. (ii)

    M2

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Rao, V., Nanjundiah, V. Ernst Mayr’s interactions with J. B. S. Haldane. HPLS 38, 138–150 (2016). https://doi.org/10.1007/s40656-016-0098-x

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