Evolutionarily stable strategy  

A solution concept in game theory  
Relationship  
Subset of  Nash equilibrium 
Superset of  Stochastically stable equilibrium, Stable Strong Nash equilibrium 
Intersects with  Subgame perfect equilibrium, Trembling hand perfect equilibrium, Perfect Bayesian equilibrium 
Significance  
Proposed by  John Maynard Smith and George R. Price 
Used for  Biological modeling and Evolutionary game theory 
Example  Hawkdove 
An evolutionarily stable strategy (ESS) is a strategy (or set of strategies) which, if adopted by a population in a given environment, is impenetrable, meaning that it cannot be invaded by any alternative strategy (or strategies) that are initially rare. It is relevant in game theory, behavioural ecology, and evolutionary psychology. An ESS is an equilibrium refinement of the Nash equilibrium. It is a Nash equilibrium that is "evolutionarily" stable: once it is fixed in a population, natural selection alone is sufficient to prevent alternative (mutant) strategies from invading successfully. The theory is not intended to deal with the possibility of gross external changes to the environment that bring new selective forces to bear.
First published as a specific term in the 1972 book by John Maynard Smith,^{ [1] } the ESS is widely used in behavioural ecology and economics, and has been used in anthropology, evolutionary psychology, philosophy, and political science.
Evolutionarily stable strategies were defined and introduced by John Maynard Smith and George R. Price in a 1973 Nature paper.^{ [2] } Such was the time taken in peerreviewing the paper for Nature that this was preceded by a 1972 essay by Maynard Smith in a book of essays titled On Evolution.^{ [1] } 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 .^{ [3] } Maynard Smith explains further in his 1982 book Evolution and the Theory of Games .^{ [4] } 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 peerreviewing 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 coauthor.
The concept was derived from R. H. MacArthur ^{ [5] } and W. D. Hamilton's^{ [6] } 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.^{ [7] }
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.
Evolutionarily stable strategies are motivated entirely differently. Here, it is presumed that the players' strategies are biologically encoded and heritable. Individuals have no control over their strategy and need not be aware of the game. They reproduce and are subject to the forces of natural selection, with the payoffs of the game representing reproductive success (biological fitness). It is imagined that alternative strategies of the game occasionally occur, via a process like mutation. To be an ESS, a strategy must be resistant to these alternatives.
Given the radically different motivating assumptions, it may come as a surprise that ESSes and Nash equilibria often coincide. In fact, every ESS corresponds to a Nash equilibrium, but some Nash equilibria are not ESSes.
An ESS is a refined or modified form of a Nash equilibrium. (See the next section for examples which contrast the two.) In a Nash equilibrium, if all players adopt their respective parts, no player can benefit by switching to any alternative strategy. In a two player game, it is a strategy pair. Let E(S,T) represent the payoff for playing strategy S against strategy T. The strategy pair (S, S) is a Nash equilibrium in a two player game if and only if this is true for both players and for all T≠S:
In this definition, strategy T can be a neutral alternative to S (scoring equally well, but not better). A Nash equilibrium is presumed to be stable even if T scores equally, on the assumption that there is no longterm incentive for players to adopt T instead of S. This fact represents the point of departure of the ESS.
Maynard Smith and Price ^{ [2] } specify two conditions for a strategy S to be an ESS. For all T≠S, either
The first condition is sometimes called a strict Nash equilibrium.^{ [9] } The second is sometimes called "Maynard Smith's second condition". The second condition means that although strategy T is neutral with respect to the payoff against strategy S, the population of players who continue to play strategy S has an advantage when playing against T.
There is also an alternative, stronger definition of ESS, due to Thomas.^{ [10] } This places a different emphasis on the role of the Nash equilibrium concept in the ESS concept. Following the terminology given in the first definition above, this definition requires that for all T≠S
In this formulation, the first condition specifies that the strategy is a Nash equilibrium, and the second specifies that Maynard Smith's second condition is met. Note that the two definitions are not precisely equivalent: for example, each pure strategy in the coordination game below is an ESS by the first definition but not the second.
In words, this definition looks like this: The payoff of the first player when both players play strategy S is higher than (or equal to) the payoff of the first player when he changes to another strategy T and the second player keeps his strategy S and the payoff of the first player when only his opponent changes his strategy to T is higher than his payoff in case that both of players change their strategies to T.
This formulation more clearly highlights the role of the Nash equilibrium condition in the ESS. It also allows for a natural definition of related concepts such as a weak ESS or an evolutionarily stable set.^{ [10] }


In most simple games, the ESSes and Nash equilibria coincide perfectly. For instance, in the prisoner's dilemma there is only one Nash equilibrium, and its strategy (Defect) is also an ESS.
Some games may have Nash equilibria that are not ESSes. For example, in harm thy neighbor (whose payoff matrix is shown here) both (A, A) and (B, B) are Nash equilibria, since players cannot do better by switching away from either. However, only B is an ESS (and a strong Nash). A is not an ESS, so B can neutrally invade a population of A strategists and predominate, because B scores higher against B than A does against B. This dynamic is captured by Maynard Smith's second condition, since E(A, A) = E(B, A), but it is not the case that E(A,B) > E(B,B).


Nash equilibria with equally scoring alternatives can be ESSes. For example, in the game Harm everyone, C is an ESS because it satisfies Maynard Smith's second condition. D strategists may temporarily invade a population of C strategists by scoring equally well against C, but they pay a price when they begin to play against each other; C scores better against D than does D. So here although E(C, C) = E(D, C), it is also the case that E(C,D) > E(D,D). As a result, C is an ESS.
Even if a game has pure strategy Nash equilibria, it might be that none of those pure strategies are ESS. Consider the Game of chicken. There are two pure strategy Nash equilibria in this game (Swerve, Stay) and (Stay, Swerve). However, in the absence of an uncorrelated asymmetry, neither Swerve nor Stay are ESSes. There is a third Nash equilibrium, a mixed strategy which is an ESS for this game (see Hawkdove game and Best response for explanation).
This last example points to an important difference between Nash equilibria and ESS. Nash equilibria are defined on strategy sets (a specification of a strategy for each player), while ESS are defined in terms of strategies themselves. The equilibria defined by ESS must always be symmetric, and thus have fewer equilibrium points.
In population biology, the two concepts of an evolutionarily stable strategy (ESS) and an evolutionarily stable state are closely linked but describe different situations.
In an evolutionarily stable strategy, if all the members of a population adopt it, no mutant strategy can invade.^{ [4] } Once virtually all members of the population use this strategy, there is no 'rational' alternative. ESS is part of classical game theory.
In an evolutionarily stable state, a population's genetic composition is restored by selection after a disturbance, if the disturbance is not too large. An evolutionarily stable state is a dynamic property of a population that returns to using a strategy, or mix of strategies, if it is perturbed from that initial state. It is part of population genetics, dynamical system, or evolutionary game theory. This is now called convergent stability.^{ [11] }
B. Thomas (1984) applies the term ESS to an individual strategy which may be mixed, and evolutionarily stable population state to a population mixture of pure strategies which may be formally equivalent to the mixed ESS.^{ [12] }
Whether a population is evolutionarily stable does not relate to its genetic diversity: it can be genetically monomorphic or polymorphic.^{ [4] }
In the classic definition of an ESS, no mutant strategy can invade. In finite populations, any mutant could in principle invade, albeit at low probability, implying that no ESS can exist. In an infinite population, an ESS can instead be defined as a strategy which, should it become invaded by a new mutant strategy with probability p, would be able to counterinvade from a single starting individual with probability >p, as illustrated by the evolution of bethedging.^{ [13] }
Cooperate  Defect  
Cooperate  3, 3  1, 4 
Defect  4, 1  2, 2 
Prisoner's Dilemma 
A common model of altruism and social cooperation is the Prisoner's dilemma. Here a group of players would collectively be better off if they could play Cooperate, but since Defect fares better each individual player has an incentive to play Defect. One solution to this problem is to introduce the possibility of retaliation by having individuals play the game repeatedly against the same player. In the socalled iterated Prisoner's dilemma, the same two individuals play the prisoner's dilemma over and over. While the Prisoner's dilemma has only two strategies (Cooperate and Defect), the iterated Prisoner's dilemma has a huge number of possible strategies. Since an individual can have different contingency plan for each history and the game may be repeated an indefinite number of times, there may in fact be an infinite number of such contingency plans.
Three simple contingency plans which have received substantial attention are Always Defect, Always Cooperate, and Tit for Tat . The first two strategies do the same thing regardless of the other player's actions, while the latter responds on the next round by doing what was done to it on the previous round—it responds to Cooperate with Cooperate and Defect with Defect.
If the entire population plays TitforTat and a mutant arises who plays Always Defect, TitforTat will outperform Always Defect. If the population of the mutant becomes too large — the percentage of the mutant will be kept small. Tit for Tat is therefore an ESS, with respect to only these two strategies. On the other hand, an island of Always Defect players will be stable against the invasion of a few TitforTat players, but not against a large number of them.^{ [14] } If we introduce Always Cooperate, a population of TitforTat is no longer an ESS. Since a population of TitforTat players always cooperates, the strategy Always Cooperate behaves identically in this population. As a result, a mutant who plays Always Cooperate will not be eliminated. However, even though a population of Always Cooperate and TitforTat can coexist, if there is a small percentage of the population that is Always Defect, the selective pressure is against Always Cooperate, and in favour of TitforTat. This is due to the lower payoffs of cooperating than those of defecting in case the opponent defects.
This demonstrates the difficulties in applying the formal definition of an ESS to games with large strategy spaces, and has motivated some to consider alternatives.
The fields of sociobiology and evolutionary psychology attempt to explain animal and human behavior and social structures, largely in terms of evolutionarily stable strategies. Sociopathy (chronic antisocial or criminal behavior) may be a result of a combination of two such strategies.^{ [15] }
Evolutionarily stable strategies were originally considered for biological evolution, but they can apply to other contexts. In fact, there are stable states for a large class of adaptive dynamics. As a result, they can be used to explain human behaviours that lack any genetic influences.
The prisoner's dilemma is a standard example of a game analyzed in game theory that shows why two completely rational individuals might not cooperate, even if it appears that it is in their best interests to do so. It was originally framed by Merrill Flood and Melvin Dresher while working at RAND in 1950. Albert W. Tucker formalized the game with prison sentence rewards and named it "prisoner's dilemma", presenting it as follows:
Two members of a criminal gang are arrested and imprisoned. Each prisoner is in solitary confinement with no means of communicating with the other. The prosecutors lack sufficient evidence to convict the pair on the principal charge, but they have enough to convict both on a lesser charge. Simultaneously, the prosecutors offer each prisoner a bargain. Each prisoner is given the opportunity either to betray the other by testifying that the other committed the crime, or to cooperate with the other by remaining silent. The possible outcomes are:
In game theory, the Nash equilibrium, named after the mathematician John Forbes Nash Jr., is a proposed solution of a noncooperative game involving two or more players in which each player is assumed to know the equilibrium strategies of the other players, and no player has anything to gain by changing only his own strategy.
Tit for tat is an English saying meaning "equivalent retaliation". It developed from "tip for tap", first used in 1558.
The game of chicken, also known as the hawk–dove game or snowdrift game, is a model of conflict for two players in game theory. The principle of the game is that while the outcome is ideal for one player to yield, but the individuals try to avoid it out of pride for not wanting to look like a 'chicken'. So each player taunts the other to increase the risk of shame in yielding. However, when one player yields, the conflict is avoided, and the game is for the most part over.
In game theory, the best response is the strategy which produces the most favorable outcome for a player, taking other players' strategies as given. The concept of a best response is central to John Nash's bestknown contribution, the Nash equilibrium, the point at which each player in a game has selected the best response to the other players' strategies.
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.
In game theory, coordination games are a class of games with multiple pure strategy Nash equilibria in which players choose the same or corresponding strategies.
In game theory, the centipede game, first introduced by Robert Rosenthal in 1981, is an extensive form game in which two players take turns choosing either to take a slightly larger share of an increasing pot, or to pass the pot to the other player. The payoffs are arranged so that if one passes the pot to one's opponent and the opponent takes the pot on the next round, one receives slightly less than if one had taken the pot on this round. Although the traditional centipede game had a limit of 100 rounds, any game with this structure but a different number of rounds is called a centipede game.
In game theory, grim trigger is a trigger strategy for a repeated game.
In game theory, a solution concept is a formal rule for predicting how a game will be played. These predictions are called "solutions", and describe which strategies will be adopted by players and, therefore, the result of the game. The most commonly used solution concepts are equilibrium concepts, most famously Nash equilibrium.
"A population is said to be in an evolutionarily stable state if its genetic composition is restored by selection after a disturbance, provided the disturbance is not too large. Such a population can be genetically monomorphic or polymorphic." —Maynard Smith (1982).
In game theory an uncorrelated asymmetry is an arbitrary asymmetry in a game which is otherwise symmetrical. The name 'uncorrelated asymmetry' is due to John Maynard Smith who called payoff relevant asymmetries in games with similar roles for each player 'correlated asymmetries'.
In game theory, folk theorems are a class of theorems about possible Nash equilibrium payoff profiles in repeated games. The original Folk Theorem concerned the payoffs of all the Nash equilibria of an infinitely repeated game. This result was called the Folk Theorem because it was widely known among game theorists in the 1950s, even though no one had published it. Friedman's (1971) Theorem concerns the payoffs of certain subgameperfect Nash equilibria (SPE) of an infinitely repeated game, and so strengthens the original Folk Theorem by using a stronger equilibrium concept: subgameperfect Nash equilibria rather than Nash equilibria.
The Bishop–Cannings theorem is a theorem in evolutionary game theory. It states that (i) all members of a mixed evolutionarily stable strategy (ESS) have the same payoff, and (ii) that none of these can also be a pure ESS. The usefulness of the results comes from the fact that they can be used to directly find ESSes algebraically, rather than simulating the game and solving it by iteration.
In game theory, the war of attrition is a dynamic timing game in which players choose a time to stop, and fundamentally trade off the strategic gains from outlasting other players and the real costs expended with the passage of time. Its precise opposite is the preemption game, in which players elect a time to stop, and fundamentally trade off the strategic costs from outlasting other players and the real gains occasioned by the passage of time. The model was originally formulated by John Maynard Smith; a mixed evolutionarily stable strategy (ESS) was determined by Bishop & Cannings. An example is an allpay auction, in which the prize goes to the player with the highest bid and each player pays the loser's low bid.
In game theory, the purification theorem was contributed by Nobel laureate John Harsanyi in 1973. The theorem aims to justify a puzzling aspect of mixed strategy Nash equilibria: that each player is wholly indifferent amongst each of the actions he puts nonzero weight on, yet he mixes them so as to make every other player also indifferent.
Risk dominance and payoff dominance are two related refinements of the Nash equilibrium (NE) solution concept in game theory, defined by John Harsanyi and Reinhard Selten. A Nash equilibrium is considered payoff dominant if it is Pareto superior to all other Nash equilibria in the game. When faced with a choice among equilibria, all players would agree on the payoff dominant equilibrium since it offers to each player at least as much payoff as the other Nash equilibria. Conversely, a Nash equilibrium is considered risk dominant if it has the largest basin of attraction. This implies that the more uncertainty players have about the actions of the other player(s), the more likely they will choose the strategy corresponding to it.
In game theory, an epsilonequilibrium, or nearNash equilibrium, is a strategy profile that approximately satisfies the condition of Nash equilibrium. In a Nash equilibrium, no player has an incentive to change his behavior. In an approximate Nash equilibrium, this requirement is weakened to allow the possibility that a player may have a small incentive to do something different. This may still be considered an adequate solution concept, assuming for example status quo bias. This solution concept may be preferred to Nash equilibrium due to being easier to compute, or alternatively due to the possibility that in games of more than 2 players, the probabilities involved in an exact Nash equilibrium need not be rational numbers.
Subjective expected relative similarity (SERS) is a normative and descriptive theory that predicts and explains cooperation levels in a family of games termed Similarity Sensitive Games (SSG), among them the wellknown Prisoner's Dilemma game (PD). SERS was originally developed in order to (i) provide a new rational solution to the PD game and (ii) to predict human behavior in singlestep PD games. It was further developed to account for: (i) repeated PD games, (ii) evolutionary perspectives and, as mentioned above, (iii) the SSG subgroup of 2x2 games. SERS predicts that individuals cooperate whenever their subjectively perceived similarity with their opponent exceeds a situational index derived from the game’s payoffs, termed the similarity threshold of the game. SERS proposes a solution to the rational paradox associated with the single step PD and provides accurate behavioral predictions. The theory was developed by Prof. Ilan Fischer at the University of Haifa.
Reciprocal altruism in humans refers to an individual behavior that gives benefit conditionally upon receiving a returned benefit, which draws on the economic concept – ″gains in trade″. Human reciprocal altruism would include the following behaviors : helping patients, the wounded, and the others when they are in crisis; sharing food, implement, knowledge.