IV.5.1.1 Evolution ultimately fixes those traits, characteristics or patterns of behaviour that represent evolutionarily stable strategies, not those that provide their bearer with the greatest biological fitness.

The fact that the advantageous or disadvantageousness of a certain strategy (in general a certain trait) depends on the frequency of alternative strategies (traits) in the population indicates that it is necessary to quite fundamentally reassess the original Neodarwinist concepts of the mechanisms of biological evolution.While the simple Neodarwinist model quite naturally assumes that the criterion of evolutionary success of a particular trait (characteristic or pattern of behaviour) consists in the average biological fitness of its bearers, contemporary theory indicates that, in the long-term perspective, the evolutionary stability of the given strategy is a more important criterion (given by patterns of behaviour or the presence of a particular trait).At the very least since 1973, when John Maynard Smith and George Price published the concept of evolutionarily stable strategy, this rendered completely irrelevant the popular dispute between the proponents and opponents of Darwinism as to whether the principle of natural selection and the principle of biological fitness are or are not circular definitions and whether Darwin’s explanation of the formation of adaptive traits is or is not a tautology from the standpoint of formal logic (see I.10.1)The development of the theory of evolutionarily stable strategy demonstrated that this explanation is primarily a scientific error – mutants whose mutated trait gives them greater fitness at the present time or in the future do not predominate in evolution, but rather mutants whose mutated traits represent evolutionarily stable strategy in the sense of game theory.We could, of course, begin to consider evolutionary stability to be a criterion of fitness.However, this would constitute a very fundamental redefinition of the Neodarwinist concept of fitness (as a technical term encompassing a set of quite specific and, under various circumstances, different characteristics affecting the chance of an organism to leave progeny, see I.10.1) and the term fitness would then really lose meaning to a substantial degree.

If we return to the original model of the dove and the hawk, we can see that, from the viewpoint of classical Neodarwinist theory, the dove strategy should win out in a structured metapopulation, as local subpopulations consisting of only doves would contain individuals with the greatest average biological fitness and migrants originating from this population would contribute to the greatest degree to the subpopulation of migrants, so that they would “infect”, with their genes, the greatest number of surrounding populations and would establish the greatest percentage of new populations.“Infection” of already existing populations would, however, be unsuccessful in a great many cases or would be only temporarily successful, as the dove strategy is not ESS.The theory of evolutionarily stable strategies shows that, in the long-term perspective, the substantially less advantageous mixed strategy from the standpoint of average biological fitness “behave as a hawk with probability p and like a dove with probability 1-p” is more successful.While this strategy is suboptimal from the standpoint of the average reward that the two individuals carry away from the encounter (compared to the average reward in a population consisting of only doves), it is, however, evolutionarily stable and, because the relevant population cannot be infected with any other strategy, it will, in time, predominate in all the local populations, and thus also in the subpopulation of migrants – potential founders of new subpopulations.

            The concept of evolutionarily stable strategy thus apparently represents one of the deepest and most important blows to the very foundations of Darwin’s theory of evolution since the emergence of Neodarwinism.It is interesting and apparently quite significant that this aspect of the theory of evolutionarily stable strategies is currently not assigned its true worth and is even not much taken into account amongst evolutionary biologists.

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The classical Darwinian theory of evolution can explain the evolution of adaptive traits only in asexual organisms. The frozen plasticity theory is much more general: It can also explain the origin and evolution of adaptive traits in both asexual and sexual organisms Read more
Draft translation from: Evoluční biologie, 2. vydání (Evolutionary biology, 2nd edition), J. Flegr, Academia Prague 2009. The translation was not done by biologist, therefore any suggestion concerning proper scientific terminology and language usage are highly welcomed. You can send your comments to flegratcesnet [dot] cz. Thank you.