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The Objection Is Sustained: a Defence of the Defense of Beanbag Genetics

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The Objection Is Sustained: a Defence of the Defense of Beanbag Genetics DEFENSE OF BEANBAG GENETICS 451 Conflict of interest: None declared. 6 Kimura M. Evolutionary rate at the molecular level. Nature 1968;217:624–26. 7 Hogben LT. In: Manton MP (ed.). Darwinism and the Study of Society. London: Tavistock Publications, 1961. References 8 Fisher RA. The Genetical Theory of Natural Selection. Oxford: Clarendon Press, 1930. 1 Haldane JBS. A defense of beanbag genetics. Perspect Biol Med 9 Denton M. Evolution: A Theory in Crisis. Bethesda, MD: 1964;7:343–60. Reprinted Int J Epidemiol 2008;37:435–42. Adler and Adler, 1986. 2 Mayr E. Animal Species and Evolution. Cambridge MA: 10 Hagedoorn AL, Hagedoorn AC. The Relative Value of the Belknap Press, 1963. Processes Causing Evolution. The Hague: Martinus Nijhoff, 3 Mayr E. Where are we? Cold Spring Harbor Symp Quant Biol 1921. 1959;24:1–24. 11 Watterson GA. The homozygosity test of neutrality. 4 Haldane JBS. The cost of natural selection. J Genet Genetics 1978;88:405–17. 1957;55:511–24. 12 Tajima F. Statistical methods for testing the neutral Downloaded from 5 Haldane JBS. More precise expressions for the cost of mutations hypothesis by DNA polymorphism. Genetics natural selection. J Genet 1961;57:351–60. 1989;123:585–95. Published by Oxford University Press on behalf of the International Epidemiological Association International Journal of Epidemiology 2008;37:451–454 ß The Author 2008; all rights reserved. doi:10.1093/ije/dyn050 http://ije.oxfordjournals.org Commentary: The objection is sustained: a defence of the defense of beanbag genetics Renee M Borges at Indian Institute of Science on May 5, 2010 Accepted 18 February 2008 Darwin couched his theory of evolution by means of to the theoretical work of Ronald Fisher, Sewall natural selection in verbal arguments. According to Wright and JBS Haldane. In 1959, in a plenary this theory, if the variation observed between individ- address at the Cold Spring Harbor Symposium on the uals was heritable, and if it contributed differentially occasion of the centenary celebration of The Origin of to the survivorship and reproductive success of Species, Mayr2 made disparaging remarks about the individuals, the process of natural selection would type of theoretical population genetics being done by ensure greater representation of individuals bearing Fisher, Wright and Haldane, and called their body of favourable traits chosen from the existing variation in work the equivalent of beanbag genetics. He implied subsequent generations. This concept is powerful, that the type of models that they developed, which whether verbally or mathematically articulated. Ernst often dealt with single genes, was too simplistic to Mayr was Darwin’s champion in the 20th century, reflect the real world that he and other naturalist- defending Darwin and Darwinism vigorously through- systematists and experimentalists such as Theodosius out his long and industrious career that spanned 80 Dobzhansky were investigating. He appeared to throw years.1 However, as outlined in Borges,1 even among down the gauntlet to the trinity of Fisher, Wright and the Darwinists, the 1950s was a time of conflict Haldane, and challenged them to account for the between experimental biologists and naturalist- value of their work; in other words, he appeared to be systematists such as Mayr on one hand and theore- declaring that their recognition as the major architects ticians on the other. The so-called evolutionary of the evolutionary synthesis was unwarranted. This synthesis of the 1930s and 1940s that amalgamated address stimulated Haldane to write his celebrated Darwin’s theory with genetics was largely attributed response: ‘In defense of beanbag genetics’,3 the paper that is being commented upon here. Before examining Haldane’s defence, it is necessary to consider the motivation behind Mayr’s statements. Centre for Ecological Sciences, Indian Institute of Science, Various possibilities have been suggested. First, the Bangalore 560 012, India. E-mail: [email protected] statements may have stemmed from Mayr’s illiteracy 452 INTERNATIONAL JOURNAL OF EPIDEMIOLOGY in genetics;4 most of Mayr’s knowledge of genetics challenge to the legitimacy of his work, for I found had been acquired from reading Dobzhansky, or from these words in Fisher’s Preface to The Genetical Theory discussion with such individuals as Dobzhansky’s of Natural Selection published in 1930:8 student Bruce Wallace;5 thus Mayr’s knowledge of the work of theoretical population geneticists was sketchy ‘The types of mind which result from training in and largely derivative. Furthermore, Dobzhansky mathematics and in biology certainly differ pro- himself, the source of much of Mayr’s genetics, did foundly; but the difference does not seem to lie not understand enough theoretical genetics and took in the intellectual faculty ...What is profoundly Wright’s formulations at face value without truly susceptible of training is the imagination, and comprehending the mathematics (see evidence pre- mathematicians and biologists seem to differ sented in Borges1). Second, Mayr is believed to have enormously in the manner in which their imagi- criticized models that dealt with independently nations are employed ...Perhaps I can best make inherited genes at single loci (the beanbag view) clear that the mathematician’s imagination also Downloaded from because he was more concerned with the ‘unity of the has been trained to some advantage, by quoting genotype’ in which there was epistasis or interactions a remark dropped casually by Eddington in a between genes. Thus Mayr’s beanbag view could have recent book—‘We need scarcely add that the been a symptom of the conflict between reductionist contemplation in natural science of a wider and holistic views of evolution. However, according domain than the actual leads to a far better to De Winter,6 fundamental misconceptions led to understanding of the actual.’ (p. 267, The Nature of Mayr’s caricature of the important work of the the Physical World.) For a mathematician the http://ije.oxfordjournals.org population geneticists. For example, Fisher’s formula- statement is almost a truism. From a biologist, tion of a gene-wise theory of selection was reduc- speaking of his own subject, it would suggest an tionist only in that it assumed that genes were extraordinarily wide outlook. No practical biologist inherited independently, while it did not assume that interested in sexual reproduction would be led to they were either expressed or functioned indepen- work out the detailed consequences experienced dently. This fundamental misconception led to Mayr’s by organisms having three or more sexes; yet what pejorative remarks and disparaging views of beanbag else should he do if he wishes to understand why genetics.6 the sexes are, in fact, always two?’ The third reason for Mayr’s critical remarks may at Indian Institute of Science on May 5, 2010 have been his need to secure for the naturalist- Here is Fisher declaring that mathematical explora- systematists (such as himself) and the experimental tion of the realm of possibilities is what gives better biologists (such as Dobzhansky) an important place in insight to our understanding of the natural world, the history of the evolutionary synthesis. Perhaps and pointing out to the dichotomy between biologists undue emphasis was being given to the work of the and mathematicians, a divide that must surely have theoreticians and not enough to the contributions of contributed to Mayr’s remarks. Haldane also provides others to the synthesis.7 If Darwin could have made several examples of this divide in his defence, which such an impact with a verbal argument, why could in the spirit of Fisher’s words, appears to be more of not others making verbal arguments also acquire a defence of theoretical biology or even of theory in important stature in the history of evolutionary science, rather than a defence of just theoretical biology? Mayr did indeed verbalize valuable ideas population genetics. Lewontin9 believes, however, that such as one that is now recognized as the founder evaluating the contribution of the theoretical work of principle in population biology,5 and therefore may the trio (Fisher–Wright–Haldane) to the evolutionary have assumed that mathematical formulations were synthesis that occurred in the 1930s and 1940s is unnecessary for important insights into evolutionary difficult because of the confusion between the history processes. The fourth reason could be that Mayr and the sociology of science at that time. According to revelled in being polemical and, by making out- Lewontin9 there was a genuine disconnect between rageous statements, he believed that he was stimu- the biologists and the theoreticians, and many lating investigation and advancing the field by important insights into evolutionary processes that provoking responses in clarification (see evidence for were possible from the theory were simply unavail- this reviewed in Borges1). able to the biologists because of their mathematical Whatever the real reason behind Mayr’s challenge, illiteracy; the evolutionary synthesis proceeded with- Haldane’s spirited defence was the result. In this out a true merger of these fields. Leigh10 also feels response, Haldane declares that he would have to pick that the reason why the work of Fisher, for example, up the challenge, because as Fisher was not alive at was little known among evolutionary biologists, the time, he could not defend himself, while Wright especially in the United States, was the difficult was the gentlest of people, and so unlikely to defend mathematics, and that Mayr, who did not understand himself. However, although Fisher had passed away, most of it, exercised a powerful negative influence it appeared that he had already anticipated such a over its assimilation into mainstream evolutionary DEFENSE OF BEANBAG GENETICS 453 biology. Ewens11 questions whether such complex and, echoing Mayr in 1959, raised doubts about the mathematics was actually required for the problems value of theoretical work (cited in Lewontin9): at hand, since Mayr and Dobzhansky seemed to make do with verbal arguments.
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