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Evolution and Directionality: Lessons from Fisher’s Fundamental Theorem

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Abstract

As is well-known, the second law of thermodynamics has the property of being ‘time asymmetric’, unlike for example the laws of Newtonian mechanics, which are time reversal-invariant. The second law tells us that in a closed thermal system, the entropy can never decrease, and attains its maximal value at equilibrium. This confers a kind of directionality on thermodynamical processes. If you were given a sequence of state-descriptions of a closed thermal system, i.e., a complete specification of the system’s physical state (including its entropy) at consecutive points in time, you would be able to deduce which direction time is running in. For the second law tells you that that ‘lower entropy’ corresponds to ‘earlier than’.

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Notes

  1. 1.

    This is one version of Wright’s adaptive landscape; the other version depicts the fitness of an organism as a function of its genotype. See Provine (1971) or Edwards (1994) for discussion of the difference.

  2. 2.

    For recent assessments of the adaptive landscape concept, see Plutynski (2008), Kaplan (2008) and Pigliucci (2008).

  3. 3.

    In talking about the change in mean fitness from one generation to another, rather than the instantaneous rate of change, I have switched to a ‘discrete time’ formulation of the FTNS. This makes no difference, for as Ewens (1989) showed, the ‘partial change’ FTNS holds in both discrete time and continuous time models.

  4. 4.

    The only situation in which average effects are not functions of allele frequency is if there is perfect additivity, i.e. no dominance and no epistasis. See Okasha (2008) for further explanation.

  5. 5.

    Thus Price wrote: “what Fisher’s theorem tells us is that natural selection at all times acts to increase the fitness of a species to live under the conditions that existed an instant earlier. But since the standard of ‘fitness’ changes from instant to instant, this constant improving tendency of natural selection does not necessarily get anywhere in terms of increasing ‘fitness’ as measured by any fixed standard (1972, p. 131).

  6. 6.

    The qualification ‘directly’ is necessary in order to exclude the indirect effect of selection on mean fitness that occurs via changes to the environment.

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Acknowledgements

This work was supported by the Arts and Humanities Research Council, Grant No. AH/F017502/1, which I gratefully acknowledge.

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Correspondence to Samir Okasha .

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Okasha, S. (2010). Evolution and Directionality: Lessons from Fisher’s Fundamental Theorem. In: Suárez, M., Dorato, M., Rédei, M. (eds) EPSA Philosophical Issues in the Sciences. Springer, Dordrecht. https://doi.org/10.1007/978-90-481-3252-2_18

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