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Is genetic drift a force?

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Abstract

One hotly debated philosophical question in the analysis of evolutionary theory concerns whether or not evolution and the various factors which constitute it (selection, drift, mutation, and so on) may profitably be considered as analogous to “forces” in the traditional, Newtonian sense. Several compelling arguments assert that the force picture is incoherent, due to the peculiar nature of genetic drift. I consider two of those arguments here—that drift lacks a predictable direction, and that drift is constitutive of evolutionary systems—and show that they both fail to demonstrate that a view of genetic drift as a force is untenable. I go on to diagnose the reasons for the stubborn persistence of this problem, considering two open philosophical issues and offering some preliminary arguments in support of the force metaphor.

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Notes

  1. Note that, contra the claim of Filler (and many others) that the magnitude of drift is represented by population size (Filler 2009, p. 777), population size only determines the distribution of drift outcomes, and hence the expected magnitude of drift. I thank an anonymous reviewer for pointing out this fact.

  2. I have described the statisticalist position in a univocal way here, though I suspect that there is as great a diversity of positions among those in the “statisticalist” camp as that in the “causalist” camp which I will describe later. Compare, for example, the positions reflected in Lewens (2010), Walsh (2010), and Matthen and Ariew (2009).

  3. I should also pause to set aside another facet of this debate: the distinction between the processes and products of evolution (the classic source here is Millstein 2002). In the following, I mean to refer only to the processes of evolution, as it is clear that this is the feature to which the force interpretation is directed.

  4. I will continue using “the force interpretation” and “the force metaphor” interchangeably in the following, recognizing throughout that the question up for debate is the extent to which force language can be said to truly describe evolving systems.

  5. A very similar explanation of the general structure of ‘force’-theories is offered by Ellis (1963, 1976), who divides states of a system between “natural” states of affairs that do not require “a continuing causal explanation” and “unnatural” states which do require such explanation—and this separation is a matter of a domain-specific demarcation of the relevant “natural” states.

  6. This point, as with all issues in the causalist/statisticalist debate, is also a matter of some debate. Importantly, if this analogy is to hold, a suitable analogue of “vector addition” must be found for the evolutionary case. Matthen and Ariew (2002, pp. 66–68) push this point forcefully, framing it in terms of the inability to compare different values of “vernacular fitness.” Considering the debate over fitness would take us too far afield here; the causalist can, however, respond by providing a new model of fitness which can be compared across different biological contexts (Pence and Ramsey 2013), or elucidating a non-additive model of force composition (Stephens 2010).

  7. Early in the debate between causalists and statisticalists, this point was often missed—Matthen and Ariew (2002), for example, take it to be a point against the causal interpretation itself that genetic drift cannot be described as a force. This entails, at best, that the force metaphor should be discarded, not that the causal interpretation is untenable, a point stressed by Stephens (2004) and Millstein (2006).

  8. This conception of genetic drift as indiscriminate, lethal natural disaster is pervasive in the philosophy of biology (see, e.g., Beatty 1984; Millstein 2002; Sterelny 2003; Gildenhuys 2009), and unfortunate insofar as biologists are often much more concerned with many of the other notions of drift mentioned here, as these are more frequently found in natural populations.

  9. This view comes, of course, from one side of the causalist/statisticalist debate, and hence is a matter of significant controversy. The characterization of drift it provides, however, is well known, and serves as an example of the general trend I identify below.

  10. Beatty (1992) also mentions neutral mutations, the founder effect, and even (though such a view is now outmoded) the causes of any non-adaptive characters as factors which have, at various historical moments, been considered to be varieties of drift.

  11. This means that, at a minimum, there is a subjective sense of “chance” and “randomness” at work here (i.e., we are unable to predict the outcome of the process of drift). Whether or not there exists a stronger type of “chance” underlying genetic drift, and what exactly this sense might amount to, seems to hinge in large part on the result of the debate over drift’s causal potency (see Rosenberg 2001).

  12. Though see Pigliucci and Kaplan (2006, Chap. 8) for some of the difficulties with the adaptive landscape metaphor.

  13. I thank an anonymous reviewer for pointing out this objection.

  14. Another way to see this is to note that in a population with only selection acting (with a non-zero selection coefficient), in the absence of mutation and migration, we could also equally well predict that the population will arrive at an absorbing barrier and stay there. The existence of the barriers has nothing to do with the process driving the population change. I thank an anonymous reviewer for pointing me toward this analogy.

  15. The claim that forces must have specifiable directions appears, at least, in Matthen and Ariew (2002); Stephens 2004); Brandon 2005, 2006); Wilson 2007); and Massin (2009).

  16. Though it is certainly the case that the argument in favor of mathematical unification is relatively straightforward, given that “drift” explanations unify a wide variety of empirical/causal phenomena. Thanks to an anonymous reviewer for pointing this out.

  17. Notably, if evolutionary forces may be stochastic, then it is likely that selection is best considered as a stochastic force as well. While I lack the space here to pursue all the consequences of this claim, it sharpens the debate between Millstein and Brandon over the distinction between natural selection and genetic drift (Brandon and Carson 1996; Millstein 2002, 2005; Brandon 2005), as it no longer becomes possible to simply sort evolution into its “deterministic” and “indeterministic” components. I thank an anonymous reviewer for noting this implication.

  18. Such conventionalism about what is to count as a force is echoed in several places in the literature (e.g., Ellis 1976; Stephens 2010); see Forster (1988) for opposition.

  19. Or, to be precise, almost any—McShea and Brandon define drift as a certain kind of population-level outcome, and it is logically possible (though practically impossible) that drift could produce precisely the outcomes expected of pure natural selection, over and over again. Any real-world evolving population, however, will drift in almost all circumstances.

  20. I should note that my argument against Brandon and McShea’s objection in this section does not extend to the coherence or utility of their own, positive ZFEL view. As part of their defense of ZFEL, however, McShea and Brandon (Brandon 2006; McShea and Brandon 2010) argue that Sober’s traditional view, on which genetic drift is considered as a second-law force, is incoherent. It is that argument alone which I claim fails. It has also been briefly evaluated (and rejected) by Stephens (2010, p. 721); the approach I offer here goes farther, I believe, toward telling us why this “default-cause” argument fails.

  21. With a small, but predictable, fraction of newly-arisen mutants. Strictly speaking, this discussion concerns behavior in the limit as population size approaches infinity, as an actually infinite population cannot be divided into proportions in this way. This example was benefited by the discussion of drift in Ramsey (2013).

  22. For one, there exist further objections to the force metaphor in the literature—Stephens (2010), for example, considers an objection due to Walsh (2007).

  23. Notably, this is a different enterprise than attempting to search for forces as a project lying within the metaphysics of science. I believe considerations like those raised by McShea and Brandon’s conventionalism and Maudlin’s characterization of quasi-Newtonian theories are fairly decisive that the appropriate question concerns not the existence or ontology of forces, but the explanatory utility of different types of force explanations in different circumstances.

  24. Wilson (2007, pp. 179–184) raises the interesting possibility that forces in Newtonian mechanics are not a fundamental depiction of the world, but rather are elements posited by Newtonian mechanics insofar as it is a special science, just as biology is. This would be yet more evidence closing the analogical gap between forces in Newtonian theory and forces in biology.

  25. Sober (1984, p. 126) invokes a very similar reference to generality in his defense of the separation of natural selection from genetic drift.

  26. Of course, I have likely left off—or unduly lumped together—further ways in which one could adopt the causalist position. The relevant, and I believe underappreciated, claim remains, however: that there is a massive variety of “causalist interpretations” on offer in the literature.

  27. To cite a few particular instances, Millstein (2006) spends a significant amount of time defending the claim that the causal processes of selection and drift ought to be located at the population level, Huneman (2012) takes it as assumed that both selection and drift are population-level phenomena, while Ramsey (2013) situates both at the individual level. Of course, there are manifold problems inherent in ‘ontological-levels’ talk (Batterman 1995; Kim 2002; Heil 1999, 2003), but we lack any better way to make reference to the issues I describe here.

  28. The fact that there are no real populations which are actually in Hardy-Weinberg equilibrium parallels the well-known fact that there are no inertial frames in real-world Newtonian systems, only approximations thereto. Also, while I lack the space to pursue the matter here, I believe this dovetails nicely with biological practice on the units/levels of selection problem. See, for illuminating discussion and analysis, Pigliucci (2010).

  29. Alternatively, one might modify the account of mechanism, making room for stochastic mechanism, as recently and persuasively advocated by DesAutels (2015).

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Acknowledgments

Special thanks to an audience at the APA Eastern Division Meeting, 2012, and particularly my commentators at that meeting, Lindley Darden and Lindsay Craig, without whom several of the best ideas here would be missing. Helpful comments were also provided at the APA by Tyler Curtain, Marc Lange, Massimo Pigliucci, and Beth Preston. Thanks as well to an audience at the 2012 PSA, especially Joshua Filler and Michael Goldsby, and an audience at the Notre Dame History and Philosophy of Science Colloquium, especially Anjan Chakravartty, Melinda Gormley, Christopher Hamlin, Pablo Ruiz de Olano, and Tom Stapleford. Finally, thanks to Edward Jurkowitz, Roberta Millstein, Grant Ramsey, and six anonymous referees for comments on various drafts of this paper. As usual, commentary should not be taken to imply endorsement, and all flaws are undoubtedly mine.

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Pence, C.H. Is genetic drift a force?. Synthese 194, 1967–1988 (2017). https://doi.org/10.1007/s11229-016-1031-2

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