Evolutionary game theory
Evolutionary game theory 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.
History
Classical game theory
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.The problem of ritualized behaviour
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.Adapting game theory to evolutionary games
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, 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. 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.
Evolutionary games
Models
Evolutionary game theory encompasses Darwinian evolution, including competition, natural selection, and heredity. Evolutionary game theory has contributed to the understanding of group selection, sexual selection, altruism, parental care, co-evolution, and ecological dynamics. Many counter-intuitive situations in these areas have been put on a firm mathematical footing by the use of these models.The common way to study the evolutionary dynamics in games is through replicator equations. These show the growth rate of the proportion of organisms using a certain strategy and that rate is equal to the difference between the average payoff of that strategy and the average payoff of the population as a whole. Continuous replicator equations assume infinite populations, continuous time, complete mixing and that strategies breed true. Some attractors of the equations are evolutionarily stable states. A strategy which can survive all "mutant" strategies is considered evolutionarily stable. In the context of animal behavior, this usually means such strategies are programmed and heavily influenced by genetics, thus making any player or organism's strategy determined by these biological factors.
Evolutionary games are mathematical objects with different rules, payoffs, and mathematical behaviours. Each "game" represents different problems that organisms have to deal with, and the strategies they might adopt to survive and reproduce. Evolutionary games are often given colourful names and cover stories which describe the general situation of a particular game. Representative games include hawk-dove, war of attrition, stag hunt, producer-scrounger, tragedy of the commons, and prisoner's dilemma. Strategies for these games include hawk, dove, bourgeois, prober, defector, assessor, and retaliator. The various strategies compete under the particular game's rules, and the mathematics are used to determine the results and behaviours.
Hawk dove
The first game that Maynard Smith analysed is the classic hawk dove game. It was conceived to analyse Lorenz and Tinbergen's problem, a contest over a shareable resource. The contestants can be either a hawk or a dove. These are two subtypes or morphs of one species with different strategies. The hawk first displays aggression, then escalates into a fight until it either wins or is injured. The dove first displays aggression, but if faced with major escalation runs for safety. If not faced with such escalation, the dove attempts to share the resource.| meets hawk | meets dove | |
| if hawk | V/2 − C/2 | V |
| if dove | 0 | V/2 |
Given that the resource is given the value V, the damage from losing a fight is given cost C:
- If a hawk meets a dove, the hawk gets the full resource V
- If a hawk meets a hawk, half the time they win, half the time they lose, so the average outcome is then V/2 minus C/2
- If a dove meets a hawk, the dove will back off and get nothing – 0
- If a dove meets a dove, both share the resource and get V/2
The solution of the hawk dove game explains why most animal contests involve only ritual fighting behaviours in contests rather than outright battles. The result does not at all depend on "good of the species" behaviours as suggested by Lorenz, but solely on the implication of actions of so-called selfish genes.
War of attrition
In the hawk dove game the resource is shareable, which gives payoffs to both doves meeting in a pairwise contest. Where the resource is not shareable, but an alternative resource might be available by backing off and trying elsewhere, pure hawk or dove strategies are less effective. If an unshareable resource is combined with a high cost of losing a contest both hawk and dove payoffs are further diminished. A safer strategy of lower cost display, bluffing and waiting to win, is then viable – a bluffer strategy. The game then becomes one of accumulating costs, either the costs of displaying or the costs of prolonged unresolved engagement. It is effectively an auction; the winner is the contestant who will swallow the greater cost while the loser gets the same cost as the winner but no resource. The resulting evolutionary game theory mathematics lead to an optimal strategy of timed bluffing.This is because in the war of attrition any strategy that is unwavering and predictable is unstable, because it will ultimately be displaced by a mutant strategy which relies on the fact that it can best the existing predictable strategy by investing an extra small delta of waiting resource to ensure that it wins. Therefore, only a random unpredictable strategy can maintain itself in a population of bluffers. The contestants in effect choose an acceptable cost to be incurred related to the value of the resource being sought, effectively making a random bid as part of a mixed strategy. This implements a distribution of bids for a resource of specific value V, where the bid for any specific contest is chosen at random from that distribution. The distribution can be computed using the Bishop-Cannings theorem, which holds true for any mixed-strategy ESS. The distribution function in these contests was determined by Parker and Thompson to be:
The result is that the cumulative population of quitters for any particular cost m in this "mixed strategy" solution is:
as shown in the adjacent graph. The intuitive sense that greater values of resource sought leads to greater waiting times is borne out. This is observed in nature, as in male dung flies contesting for mating sites, where the timing of disengagement in contests is as predicted by evolutionary theory mathematics.