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Evolutionary causes as mechanisms: a critical analysis

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Abstract

In this paper, we address the question whether a mechanistic approach can account for evolutionary causes. The last decade has seen a major attempt to account for natural selection as a mechanism. Nevertheless, we stress the relevance of broadening the debate by including the other evolutionary causes inside the mechanistic approach, in order to be a legitimate conceptual framework on the same footing as other approaches to evolutionary theory. We analyse the current debate on natural selection as a mechanism, and extend it to the rest of the evolutionary causes. We focus on three approaches that we call the stochastic view, the functional view, and the minimalist view. We argue that all of them are unable to account for evolutionary causes as mechanisms. It is concluded that the current mechanistic proposals cannot be accepted as a common framework for evolutionary causes. Finally, we outline some guidelines and requirements that any mechanistic proposal should meet in order to be applied to evolutionary theory.

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Notes

  1. These notions of mechanism, which were analysed by Skipper and Millstein, are ontic. They are based on the idea that mechanisms are part of reality and exist independently of us. However, Daniel J. Nicholson (2012) has proposed an epistemic notion of mechanism for biology and claimed that it can account for natural selection. Nicholson defines mechanisms as “epistemic models that enable the explanation of how phenomena are causally brought about” (Nicholson 2012, 161). Although Nicholson considers that his notion of mechanism allows us to understand natural selection as a mechanism, understanding natural selection (or another evolutionary cause) as a mechanism in this sense is problematic and does not suit biologists’ ideas.

  2. Skipper and Millstein (2005) consider that only MDC’s proposal faces problems related with regularity. Regarding Glennan’s proposal, they argue that it “holds promise for capturing the way in which natural selection is regular” (Skipper and Millstein 2005, 342).

  3. There is disagreement among some philosophers on this causal view, arguing instead for a “statistical view” of evolution (Matthen and Ariew 2002, Walsh et al. 2002). In as much we accept that these phenomena are all causal, then we ask whether there can be a mechanistic account for them.

  4. Joyce C. Havstad (2011) has also raised some other problems of Barros’ proposal. She noted that the model of the mechanism of generalized natural selection proposed by Barros is too general and due to this fact is unable to distinctively characterize natural selection. The model proposed by Barros is not a model of generalized natural selection but a model of selection in general, which fits any selective process.

  5. There are some theoretical works where multiple evolutionary causes, and not only genetic drift, are modelled stochastically. For example, Rice and collaborators (Rice 2008; Rice and Papadopoulos 2009; Rice et al. 2011) have developed a stochastic version of the Price equation that can deal with random variables as stochastic fitness and stochastic migration, and therefore allowing to model selection and migration as stochastic processes. However, we consider that those theoretical works do not accurately represent biologists’ ideas about the nature of evolutionary causes. Biologists often understand evolutionary causes other than drift as deterministic processes. From our point of view, modelling stochastically those evolutionary causes is probably just a result of epistemological limitations. This seems the case of Rice and collaborators, inasmuch they talk repeatedly about our epistemic problems to study evolutionary systems and the necessity to construct stochastic models to supply those shortcomings. For example, they say that “we can not know with certainty how many descendants an individual will leave or what they will look like until after reproduction has taken place” (Rice and Papadopoulos 2009, 2); and that “[b]ecause we can not know with certainty how many descendants each individual in a population will have, we need to treat fitness as a random variable -having a distribution of possible values” (Rice et al. 2011).

  6. It is necessary to point out that we are not claiming that any deviation from fitness expectations should be attributed to drift. That deviation could be the result of other elements (selective pressures and/or other evolutionary causes that were not taken into account, a spurious statistical correlation, etc.).

  7. Another remarkable example of this deterministic conceptualization is provided by Brian Charlesworth: “In the era of multi-species comparisons of genome sequences and genome-wide surveys of DNA sequence variability, there is more need than ever before to understand the evolutionary role of genetic drift, and its interactions with the deterministic forces of mutation, migration, recombination and selection.” (Charlesworth 2009, 195).

  8. We think that, in the case of natural selection, those external sources of irregularity identified by DesAutels –i.e. “non-critical environmental features which are not constitutive of the token mechanism [selection]” (2016, 19)– are very likely to be just instantiations of drift processes.

  9. DesAutels also proposed a third distinction. He distinguishes between abstract and concrete regularity. Abstract regularity is the ability of a type mechanism to subsume different token mechanisms. On the other hand, concrete regularity is the capacity of a type mechanism to give a detailed account of the token mechanisms that it subsumes.

  10. Certainly, there are examples of sterile organisms with positive fitness, like in eusocial insect colonies, where some individuals forego their reproductive capacity in order to support the reproduction of their relatives. These types of organisms successfully pass their genes to the next generation. In our example, there is no such behavior.

  11. As Gillespie says: “To a geneticist, fitness is just another trait with a genetic component. To an evolutionist, it is the ultimate trait because it is the one upon which natural selection acts” (Gillespie 2004, 59).

  12. De Jong endorses a similar claim: “Differences in adaptation will lead to fitness differences, but fitness differences are not necessarily associated with differences in adaptation” (De Jong 1994, 20).

  13. Another example of this is a particular type of selection, the so-called called “survival of the first”, where a best adapted type cannot invade a population because the population growth is superexponential (see Michod 1999, chap. 3; Nowak 2006, chap. 2, for mathematical details). It could be argued that the fact that natural selection does not always produce adaptation, does not imply that adaptation is not the function of natural selection. In order for a mechanism to have a function it does not have to always succeed at implementing that function. For example, heart’s function is to pump blood at a specific rate, but sometimes it does not pump blood at that rate. Nevertheless, it does not seem the case for natural selection and adaptation. When a heart does not pump blood at a proper rate, doctors consider that it is not producing its function in a correct matter. However, when natural selection is acting but it does not produce adaptation, evolutionary biologists do not consider that it is not producing its function in a correct matter, as long as there is non-random differential reproductive success.

  14. There is a related worry. It might also be the case that a mechanistic approach is able to conceptualize evolutionary factors as mechanisms, in an isolated way, but it is not able to express the interactions between different mechanisms.

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Acknowledgements

We would like to thank the editor Staffan Müller-Wille and two anonymous reviewers for fruitful criticism of an earlier version of this paper. Valeriano Iranzo and the members of the Philosophy of Science Reading Group of the University of Valencia have also given helpful comments. Saúl Pérez-González is supported by the Spanish Ministry of Science, Innovation and Universities under Grant FPU16/03274. Victor J. Luque is supported by the Spanish Ministry of Science, Innovation and Universities under Projects FFI2016-76799-P and FFI2017-89639-P.

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Pérez-González, S., Luque, V.J. Evolutionary causes as mechanisms: a critical analysis. HPLS 41, 13 (2019). https://doi.org/10.1007/s40656-019-0250-5

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