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Evolutionarily stable strategy

An evolutionarily stable strategy (ESS) is a strategy (or set of strategies) that is impermeable when adopted by a population in adaptation to a specific environment, that is to say it cannot be displaced by an alternative strategy (or set of strategies) which may be novel or initially rare. Introduced by John Maynard Smith and George R. Price in 1972/3, it is an important concept in behavioural ecology, evolutionary psychology, mathematical game theory and economics, with applications in other fields such as anthropology, philosophy and political science.

In game-theoretical terms, an ESS is an equilibrium refinement of the Nash equilibrium, being a Nash equilibrium that is also "evolutionarily stable." Thus, once fixed in a population, natural selection alone is sufficient to prevent alternative (mutant) strategies from replacing it (although this does not preclude the possibility that a better strategy, or set of strategies, will emerge in response to selective pressures resulting from environmental change).

Evolutionarily stable strategies were defined and introduced by John Maynard Smith and George R. Price in a 1973 Nature paper. Such was the time taken in peer-reviewing the paper for Nature that this was preceded by a 1972 essay by Maynard Smith in a book of essays titled On Evolution. The 1972 essay is sometimes cited instead of the 1973 paper, but university libraries are much more likely to have copies of Nature. Papers in Nature are usually short; in 1974, Maynard Smith published a longer paper in the Journal of Theoretical Biology. Maynard Smith explains further in his 1982 book Evolution and the Theory of Games. Sometimes these are cited instead. In fact, the ESS has become so central to game theory that often no citation is given, as the reader is assumed to be familiar with it.

Maynard Smith mathematically formalised a verbal argument made by Price, which he read while peer-reviewing Price's paper. When Maynard Smith realized that the somewhat disorganised Price was not ready to revise his article for publication, he offered to add Price as co-author.

Maynard Smith on p.174 of the Postscript in his, 1982 work Evolution and the Theory of Games argues that the concept of an ESS is itself polyphyletic (i.e. derives from more than one common evolutionary ancestral group or ancestor). In this he not only mentions Price but also list various other sources for the concept. These include the following:

But the concept proper was derived from R. H. MacArthur and W. D. Hamilton's work on sex ratios, derived from Fisher's principle, especially Hamilton's (1967) concept of an unbeatable strategy. Maynard Smith was jointly awarded the 1999 Crafoord Prize for his development of the concept of evolutionarily stable strategies and the application of game theory to the evolution of behaviour.

Uses of ESS:

The Nash equilibrium is the traditional solution concept in game theory. It depends on the cognitive abilities of the players. It is assumed that players are aware of the structure of the game and consciously try to predict the moves of their opponents and to maximize their own payoffs. In addition, it is presumed that all the players know this (see common knowledge). These assumptions are then used to explain why players choose Nash equilibrium strategies.

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