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Distinguishing Natural Selection from Other Evolutionary Processes in the Evolution of Altruism

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Abstract

Altruism is one of the most studied topics in theoretical evolutionary biology. The debate surrounding the evolution of altruism has generally focused on the conditions under which altruism can evolve and whether it is better explained by kin selection or multilevel selection. This debate has occupied the forefront of the stage and left behind a number of equally important questions. One of them, which is the subject of this article, is whether the word “selection” in “kin selection” and “multilevel selection” necessarily refers to “evolution by natural selection.” I show, using a simple individual-centered model, that once clear conditions for natural selection and altruism are specified, one can distinguish two kinds of evolution of altruism, only one of which corresponds to the evolution of altruism by natural selection, the other resulting from other evolutionary processes.

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Notes

  1. For a recent discussion on the relation between multilevel selection and kin selection see Okasha (2015).

  2. I borrow the term “selective environment” from Brandon (1990). As recognized by Brandon (2014) this concept of natural selection faces several possible objections. One of them is that any case in which the individuals of a population significantly interact with each other (altruism is only one case in which this kind of interaction occurs) will prevent these individuals from being in the same environment. I will regard this problem as non-fatal to this formulation and leave its resolution for further work.

  3. Brandon, in his original definition, refers to the environment as only external factors to the organism. For reasons that cannot be developed here, to be consistent, it should be defined in reference to a phenotype, be it expressed within or beyond the physical boundaries of the organism. See Haig (2012) for a similar notion of the environment in relation to what he calls the “strategic gene.”

  4. I leave for further work to determine which evolutionary force(s) each kind of difference should be attributed to.

  5. This is inspired from Sober and Wilson (1998, pp. 19–21)

  6. This is also a point made in the literature on reciprocal altruism (Trivers 1971; Axelrod 1984).

  7. By “causal factors” I mean “difference makers” following Woodward’s (2003) interventionist account of causation.

  8. Note that β, in the general case, could be inferior to 1 (but superior or equal to 0), in which case altruistic individuals would “avoid” interacting with other altruistic individuals. I consider here only the case in which altruistic individuals “seek” other altruistic individuals, hence why I assume β ≥ 1.

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Acknowledgments

I am thankful to Patrick Forber, Arnaud Pocheville, and two anonymous reviewers for their comments on earlier versions of this paper. I am particularly thankful to Arnaud Pocheville for his extensive help on the most technical parts of the paper and for his thorough proofreading. This research was supported under Australian Research Council’s Discovery Projects funding scheme (Projects DP0878650 and DP150102875).

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  1. Department of Philosophy, University of Sydney, Sydney, NSW, Australia

    Pierrick Bourrat

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Appendix

Appendix

Let us start from (15):

$$\beta (bN_{A} (N_{T} - N_{A} ) - cN_{T} N_{A} ) > bN_{A} (N_{T} - N_{A} ) + cN_{T} \left( {N_{T} - N_{A} } \right)$$

To be meaningful, we assume β ≥ 0. Let us call the term in brackets on the left-hand side of (15) \(P = bN_{A} (N_{T} - N_{A} ) - cN_{T} N_{A}\). Let’s call Q the right hand side.

If P > 0 we have:

$$\beta > Q / P$$

We know that\(cN_{T} (N_{T} - N_{A} ) > 0 > - cN_{T} N_{A}\), thus Q > P which means that β > 1.

If P = 0 then 0 > Q, which is impossible.

If P < 0 then either 0 > Q, which is impossible, or β < 0, which is impossible.

Thus, if (15) holds, β > 1.

Then P > 0 implies N A  > 0 and \(b > c\frac{{N_{T} }}{{N_{T} - N_{A} }}\), in the case where \(N_{T} \ne N_{A}\). This is an interesting constraint bearing on b which is consistent with the hypotheses classically made in models on the evolution of altruism, namely that the benefit received by the focal altruistic individual is larger than the cost it pays.

Thus if (15) holds, β > 1 (which satisfies the intuition), N A  > 0 (which is expected), and \(b > c\frac{{N_{T} }}{{N_{T} - N_{A} }}\), if \(N_{T} \ne N_{A}\).

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Bourrat, P. Distinguishing Natural Selection from Other Evolutionary Processes in the Evolution of Altruism. Biol Theory 10, 311–321 (2015). https://doi.org/10.1007/s13752-015-0210-6

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