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Application of game theory to evolving populations in biology
Evolutionary game theory (EGT) is the application of game theory to evolving populations in biology. It defines a framework of contests, strategies, and analytics into which Darwinian competition can be modelled. It originated in 1973 with John Maynard Smith and George R. Price's formalisation of contests, analysed as strategies, and the mathematical criteria that can be used to predict the results of competing strategies. Evolutionary game theory differs from classical game theory in focusing more on the dynamics of strategy change. This is influenced by the frequency of the competing strategies in the population. Evolutionary game theory has helped to explain the basis of altruistic behaviours in Darwinian evolution. It has in turn become of interest to economists, sociologists, anthropologists, and philosophers.

Classical non cooperative game theory was conceived by John von Neumann to determine optimal strategies in competitions between adversaries. A contest involves players, all of whom have a choice of moves. Games can be a single round or repetitive. The approach a player takes in making their moves constitutes their strategy. Rules govern the outcome for the moves taken by the players, and outcomes produce payoffs for the players; rules and resulting payoffs can be expressed as decision trees or in a payoff matrix. Classical theory requires the players to make rational choices. Each player must consider the strategic analysis that their opponents are making to make their own choice of moves.

Evolutionary game theory started with the problem of how to explain ritualized animal behaviour in a conflict situation; "why are animals so 'gentlemanly or ladylike' in contests for resources?" The leading ethologists Niko Tinbergen and Konrad Lorenz proposed that such behaviour exists for the benefit of the species. John Maynard Smith considered that incompatible with Darwinian thought, where selection occurs at an individual level, so self interest is rewarded while seeking the common good is not. Maynard Smith, a mathematical biologist, turned to game theory as suggested by George Price, though Richard Lewontin's attempts to use the theory had failed.

Maynard Smith realised that an evolutionary version of game theory does not require players to act rationally—only that they have a strategy. The results of a game show how good that strategy was, just as evolution tests alternative strategies for the ability to survive and reproduce. In biology, strategies are genetically inherited traits that control an individual's action, analogous with computer programs. The success of a strategy is determined by how good the strategy is in the presence of competing strategies (including itself), and of the frequency with which those strategies are used. Maynard Smith described his work in his book Evolution and the Theory of Games. Participants aim to produce as many replicas of themselves as they can, and the payoff is in units of fitness (relative worth in being able to reproduce). It is always a multi player game with many competitors. Rules include replicator dynamics, in other words how the fitter players will spawn more replicas of themselves into the population and how the less fit will be culled, in a replicator equation. The replicator dynamics models heredity but not mutation, and assumes asexual reproduction for the sake of simplicity. Games are run repetitively with no terminating conditions. Results include the dynamics of changes in the population, the success of strategies, and any equilibrium states reached. Unlike in classical game theory, players do not choose their strategy and cannot change it: they are born with a strategy and their offspring inherit that same strategy.

In the war of attrition there must be nothing that signals the size of a bid to an opponent, otherwise the opponent can use the cue in an effective counter strategy. There is however a mutant strategy which can better a bluffer in the war of attrition game if a suitable asymmetry exists, the bourgeois strategy. Bourgeois uses an asymmetry of some sort to break the deadlock. In nature one such asymmetry is possession of a resource. The strategy is to play a hawk if in possession of the resource, but to display then retreat if not in possession. This requires greater cognitive capability than hawk, but bourgeois is common in many animal contests, such as in contests among mantis shrimps and among speckled wood butterflies.

Games like hawk dove and war of attrition represent pure competition between individuals and have no attendant social elements. Where social influences apply, competitors have four possible alternatives for strategic interaction. This is shown on the adjacent figure, where a plus sign represents a benefit and a minus sign represents a cost. In a cooperative or mutualistic relationship both "donor" and "recipient" are almost indistinguishable as both gain a benefit in the game by co operating, i. e. the pair are in a game wise situation where both can gain by executing a certain strategy, or alternatively both must act in concert because of some encompassing constraints that effectively puts them "in the same boat". In an altruistic relationship the donor, at a cost to themself provides a benefit to the recipient. In the general case the recipient will have a kin relationship to the donor and the donation is one way. Behaviours where benefits are donated alternatively (in both directions) at a cost, are often called "altruistic", but on analysis such "altruism" can be seen to arise from optimised "selfish" strategies. Spite is essentially a “reversed” form of cooperation where neither party receives a tangible benefit. The general case is that the ally is kin related and the benefit is an easier competitive environment for the ally. Note: George Price, one of the early mathematical modellers of both altruism and spite, found this equivalence particularly disturbing at an emotional level. Selfishness is the base criteria of all strategic choice from a game theory perspective – strategies not aimed at self survival and self replication are not long for any game. Critically however, this situation is impacted by the fact that competition is taking place on multiple levels – i. e. at a genetic, an individual and a group level.

Eusocial insect workers forfeit reproductive rights to their queen. It has been suggested that kin selection, based on the genetic makeup of these workers, may predispose them to altruistic behaviours. Most eusocial insect societies have haplodiploid sexual determination, which means that workers are unusually closely related. This explanation of insect eusociality has, however, been challenged by a few highly noted evolutionary game theorists (Nowak and Wilson) who have published a controversial alternative game theoretic explanation based on a sequential development and group selection effects proposed for these insect species.

Rock paper scissors incorporated into an evolutionary game has been used for modelling natural processes in the study of ecology. Using experimental economics methods, scientists have used RPS games to test human social evolutionary dynamical behaviours in laboratories. The social cyclic behaviours, predicted by evolutionary game theory, have been observed in various laboratory experiments.

Aside from the difficulty of explaining how altruism exists in many evolved organisms, Darwin was also bothered by a second conundrum – why a significant number of species have phenotypical attributes that are patently disadvantageous to them with respect to their survival – and should by the process of natural section be selected against – e. g. the massive inconvenient feather structure found in a peacock's tail. Regarding this issue Darwin wrote to a colleague "The sight of a feather in a peacock's tail, whenever I gaze at it, makes me sick." It is the mathematics of evolutionary game theory, which has not only explained the existence of altruism, but also explains the totally counterintuitive existence of the peacock's tail and other such biological encumbrances. On analysis, problems of biological life are not at all unlike the problems that define economics – eating (akin to resource acquisition and management), survival (competitive strategy) and reproduction (investment, risk and return). Game theory was originally conceived as a mathematical analysis of economic processes and indeed this is why it has proven so useful in explaining so many biological behaviours. One important further refinement of the evolutionary game theory model that has economic overtones rests on the analysis of costs. A simple model of cost assumes that all competitors suffer the same penalty imposed by the game costs, but this is not the case. More successful players will be endowed with or will have accumulated a higher "wealth reserve" or "affordability" than less successful players. This wealth effect in evolutionary game theory is represented mathematically by "resource holding potential (RHP)" and shows that the effective cost to a competitor with a higher RHP are not as great as for a competitor with a lower RHP. As a higher RHP individual is a more desirable mate in producing potentially successful offspring, it is only logical that with sexual selection RHP should have evolved to be signalled in some way by the competing rivals, and for this to work this signalling must be done honestly. Amotz Zahavi has developed this thinking in what is known as the "handicap principle", where superior competitors signal their superiority by a costly display. As higher RHP individuals can properly afford such a costly display this signalling is inherently honest, and can be taken as such by the signal receiver. In nature this is illustrated than in the costly plumage of the peacock. The mathematical proof of the handicap principle was developed by Alan Grafen using evolutionary game theoretic modelling.

A mathematical model analysing the behaviour of a system needs initially to be as simple as possible to aid in developing a base understanding the fundamentals, or “first order effects”, pertaining to what is being studied. With this understanding in place it is then appropriate to see if other, more subtle, parameters (second order effects) further impact the primary behaviours or shape additional behaviours in the system. Following Maynard Smith's seminal work in evolutionary game theory, the subject has had a number of very significant extensions which have shed more light on understanding evolutionary dynamics, particularly in the area of altruistic behaviors. Some of these key extensions to evolutionary game theory are:

Geographic factors in evolution include gene flow and horizontal gene transfer. Spatial game models represent geometry by putting contestants in a lattice of cells: contests take place only with immediate neighbours. Winning strategies take over these immediate neighbourhoods and then interact with adjacent neighbourhoods. This model is useful in showing how pockets of co operators can invade and introduce altruism in the Prisoners Dilemma game, where Tit for Tat (TFT) is a Nash Equilibrium but NOT also an ESS. Spatial structure is sometimes abstracted into a general network of interactions. This is the foundation of evolutionary graph theory.

Many evolutionary games have been modelled in finite populations to see the effect this may have, for example in the success of mixed strategies.