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Fitness “kinematics”: biological function, altruism, and organism–environment development

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Abstract

It’s recently been argued that biological fitness can’t change over the course of an organism’s life as a result of organisms’ behaviors. However, some characterizations of biological function and biological altruism tacitly or explicitly assume that an effect of a trait can change an organism’s fitness. In the first part of the paper, I explain that the core idea of changing fitness can be understood in terms of conditional probabilities defined over sequences of events in an organism’s life. The result is a notion of “conditional fitness” which is static but which captures intuitions about apparent behavioral effects on fitness. The second part of the paper investigates the possibility of providing a systematic foundation for conditional fitness in terms of spaces of sequences of states of an organism and its environment. I argue that the resulting “organism–environment history conception” helps unify diverse biological perspectives, and may provide part of a metaphysics of natural selection.

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Notes

  1. It appears that contributing to fitness can’t be understood in terms of counterfactuals in any simple way; see Wimsatt (1972, pp. 55ff) and Millikan (2002).

  2. Some models (e.g., Ewens 2004) and general accounts of fitness (e.g., Mills and Beatty 1979; Sober 1984) define type fitness as an average of token fitnesses. If this were correct in general, arbitrary fluctuations in token fitnesses should sometimes accumulate and force type fitnesses to diverge from what’s biologically appropriate (Abrams 2007).

  3. Ramsey also described two notions of fitness, ratchet fitness and flux fitness, which do allow fitness changes, but argued that neither is relevant to natural selection. Ramsey didn’t describe relationships between these notions. As I mention below (footnote 8), my approach allows the possibility of seeing static fitness as deriving from flux fitness along with other information.

  4. Ramsey (2006, pp. 492f) does acknowledge that the number of possible lives could be infinite, but only in order to argue that this doesn’t present an epistemological problem for his account of fitness. My point here is that Ramsey’s characterization of what fitness is leaves out an important aspect: the probability distribution over possible lives. Note that this probability distribution may be over an uncountably infinite number of element—lives—but this is mathematically unproblematic; in such cases probabilities are defined by integrals over a probability density function rather than by sums of finite or countably infinite probabilities.

  5. Much of this discussion was inspired by Gillespie (1977), summarizing other work by Gillespie.

  6. It may be that different definitions of fitness are needed in different contexts (Stearns 1989; Brandon 1990; Krimbas 2004; Ariew and Lewontin 2004; Rosenberg and Bouchard 2008); see Abrams (2009a) for a contrary view. Note that biologists often define fitness as a “deterministic” scalar which doesn’t depend on probabilities; this usually seems to be a simplification for modeling convenience. I don’t deal here with arguments that fitness only summarizes actual reproductive success (e.g., Walsh 2007); see Rosenberg and Bouchard (2008) for further references and critiques.

  7. Abrams (2009a) argues that probabilities of numbers of later descendants can be replaced by probabilities of occurrences in a current organism’s life. Ramsey, Neander, and others use inclusive fitness (Hamilton 1964), which incorporates reproductive probabilities for kin; my view is that effects of kin on reproductive probabilities are special cases of more general contextual effects: group selection, frequency- or density-dependent selection, etc. (Michod 1982; Sterelny 1996; Sober and Wilson 1998). Rosenberg and Bouchard’s arguments that fitness might be defined by a token organism “solving the design problems set by [an environment] more fully” than another organism (Rosenberg and Bouchard 2008, and references given there) is not intended to apply to types, but I suspect that an analogous notion of fitness for types would ultimately have to be cashed out in terms of probabilities of reproductive events or other events in possible lives.

  8. Probabilities of branchings are probabilities conditional on the previous state, and they would satisfy a Markov condition, i.e., the probability of an event conditional on a prior event is unaffected by still earlier events. This implies that the probability of a single history would be the product of probabilities of branchings along the way. The probability of an entire set of histories beginning from particular initial conditions would be the sum, or more likely integral over probabilities of individual histories beginning from those conditions. Finally, the probability of a set of histories beginning from a variety of initial conditions would be a sum or integral of products of probabilities of initial conditions and probabilities of histories conditional on the various initial conditions (cf. Abrams 2007). This provides a way of connecting Ramsey’s (2006) notion of flux fitness with his notion of block fitness, defining the latter in terms of the former along with probabilities over initial conditions.

  9. I’m skeptical about this aspect of the PIF (Abrams 2007) despite my appreciation of its intended theoretical role.

  10. Such as a deterministic mechanism which systematically generates outcomes in certain frequencies.

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Acknowledgements

I’m grateful for many individuals’ contributions to the history leading to this paper (without suggesting that they endorse its conclusions): Bill Wimsatt, Joel Velasco, Kim Sterelny, Elliott Sober, Eric Saidel, Alex Rosenberg, Robert Richardson, Grant Ramsey, Jeremy Pober, Jessica Pfeiffer, Ruth Millikan, Dan McShea, Philippe Huneman, Dan Garber, Patrick Forber, Lindley Darden, Frédéric Bouchard, Robert Brandon, Matt Barker, André Ariew, Murat Aydede, anonymous reviewers, and audience members at various presentations.

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  1. Department of Philosophy, University of Alabama at Birmingham, 900 13th South Street, HB 414A, Birmingham, AL, 35294-1260, USA

    Marshall Abrams

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Abrams, M. Fitness “kinematics”: biological function, altruism, and organism–environment development. Biol Philos 24, 487–504 (2009). https://doi.org/10.1007/s10539-009-9153-2

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