Abstract
This is the second of a two-part essay on the history of debates concerning the creativity of natural selection, from Darwin through the evolutionary synthesis and up to the present. In the first part, I focussed on the mid-late nineteenth century to the early twentieth, with special emphasis on early Darwinism and its critics, the self-styled “mutationists.” The second part focuses on the evolutionary synthesis and some of its critics, especially the “neutralists” and “neo-mutationists.” Like Stephen Gould, I consider the creativity of natural selection to be a key component of what has traditionally counted as “Darwinism.” I argue that the creativity of natural selection is best understood in terms of (1) selection initiating evolutionary change, and (2) selection directing evolutionary change, for example by creating the variation that it subsequently acts upon. I consider the respects in which both of these claims sound non-Darwinian, even though they have long been understood by supporters and critics alike to be virtually constitutive of Darwinism.
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Notes
Note the importance of temporal order here, and the different evolutionary narratives offered by Darwinians vs. mutationists. Another debate taking place during the time frame covered here—the “Fisher-Wright controversy”—also concerned the timing of evolutionary events. Rosales (2017) stresses the narrative differences between Ronald Fisher’s “mass selection” and Sewall Wright’s “shifting balance” views of evolutionary change.
As I mentioned in Part I, there is another (at least one other) important line of discussion concerning creativity that I am not considering in either essay, even though it is especially dear to me. It has to do with a different (though partially overlapping) set of issues—especially indeterminism and vitalism—and a different (though partially overlapping) group of actors. For example, it would add Henri Bergson and Pierre Teilhard de Chardin, and would consider viewpoints of Ronald Fisher and Theodosius Dobzhansky that are different from those discussed here. I hope this will someday be the third part of this project. Fortunately, Gayon (2008) left us with a superb analysis of Bergson’s influence in this regard, to go with Hodge’s (1992) as-always thoughtful and provocative account of Fisher and Bergson.
See also Simpson’s acknowledgment (1953, p. 77).
See also Dobzhansky (1970, pp. 201–209).
Chetverikov does not (here) address the degree of dominance/recessiveness of the genetic variants in question, or how variations that might initially be recessive might subsequently become dominant. If A2 and B2 are both completely recessive, the selection of A2 would have to be preceded by a doubling-up of A2 through sexual crossing. Then as the frequency of A2 increases, it would become more and more likely that A2A2 would occur in combination with a doubled-up B2B2, bringing into existence the combination A2A2/B2B2 that would subsequently be selected over other alternative combinations of genes at those loci.
Interestingly, Muller (1929) offered an account of how otherwise improbable combinations of beneficial variations could be brought about by selection. His account resembled the version that I have associated with the Synthesis. But Muller never presented it as an argument for the creativity of natural selection. Moreover, it is importantly different from the Synthesis version in that it involves new beneficial mutations arising in combination with previously selected alleles. The new combination does not arise until the new mutation occurs; while on the Synthesis version there is no waiting around for new, direction-continuing, or direction-setting mutations. That Muller could be seen as a mutationist is evidenced by the neo-mutationist Masatoshi Nei’s portrayal of him as such, as will be discussed in the next section.
Along these lines, see also Phillips (1996). The title, “Waiting for a Compensatory Mutation: Phase Zero of the Shifting-Balance Process,” says it all. Phillips argues that Wright’s three-phase, shifting-balance theory of evolution ought to be supplemented with an additional “phase zero” at the beginning, corresponding to the mutational production of variation. The shifting-balance theory, as formulated by Wright, had no mutation going on. “A largely untreated aspect of the shifting-balance theory, that of the limiting impact of waiting for the production of new mutations, is analysed here” (Phillips 1996, p. 271).
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Acknowledgements
This paper is dedicated to the memory of Jean Gayon, who contributed so much to the intellectual and social community of historians and philosophers of science; and whose insights and good will, and smile, were especially appreciated by those of us working on evolutionary biology. Thank you Jean. Many thanks also to Philippe Huneman, the editor of this special issue, and to my fellow contributors for your insights into the evolutionary synthesis and your feedback at our “debuts” in Montpellier and Chicago. Special thanks to Cosima Herter, Arlin Stoltzfus, Alirio Rosales, Michael Dietrich, Kele Cable, Philippe Huneman, Richard Burian, David Depew, Brian McLoone, and my two very thoughtful reviewers. Thanks also to my audiences at the 2014 “Philosophy of Biology at Madison” (POBAM) workshop, organized by Elliott Sober; and the 2014 “Life Sciences in the 20th Century” symposium at Washington University, celebrating Garland Allen’s retirement, organized by Jane Maienschein and Mike Dietrich.
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Beatty, J. The Creativity of Natural Selection? Part II: The Synthesis and Since. J Hist Biol 52, 705–731 (2019). https://doi.org/10.1007/s10739-019-09583-4
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DOI: https://doi.org/10.1007/s10739-019-09583-4