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Evolutionary game theory studies how a population's composition changes when the success of a pure strategy depends on which other pure strategies it encounters. The central feedback is a frequency-dependent payoff: becoming common changes the opponents a strategy meets, which can change whether it does well. In a coordination game, common behaviour can reinforce itself; in the coexistence example below, being rare is an advantage. This makes evolutionary game theory useful for asking which behaviours can persist and which initial compositions lead to different outcomes, without assuming that each participant solves an optimization problem.

In the basic model, a large, well-mixed population has finitely many inherited pure strategies; interactions sample opponents according to their population frequencies. Assume that a strategy's expected payoff determines its per capita net reproductive growth rate, with a common baseline allowed. These are modelling assumptions: reproduction must transmit the strategy, encounters must match the proposed sampling rule, and the conversion from payoff to growth must be justified for an application. The replicator equation below describes the resulting continuous-time frequencies. Taylor and Jonker's ^[ref] explicitly derive this growth model.

For learned behaviour, reproduction can instead be replaced by an assumed copying rule: an individual samples another individual uniformly and copies a better-performing pure strategy at a rate proportional to the positive payoff difference. At frequencies $x_i,x_j$, the net flow from $j$ to $i$ is $x_i{x}_j(f_i-f_j)$, because the two directional positive parts subtract to $f_i-f_j$. Summing over $j$ gives $x_i(f_i-\bar{f})$, the replicator equation. This derivation states a particular mechanism; an arbitrary rule for learning or imitation need not give the same dynamical system.

Three questions must be distinguished. A symmetric Nash equilibrium tests whether an individual can improve its expected payoff against the resident composition. An evolutionarily stable strategy tests whether each distinct rare mutant is outperformed after it is introduced. Dynamical stability tests the response of an actual dynamical system to small changes of composition, and asymptotic stability additionally requires nearby trajectories to converge. For a finite symmetric bimatrix game under the continuous-time replicator equation, every evolutionarily stable strategy is a symmetric Nash equilibrium and a locally asymptotically stable rest point; the converse from asymptotic stability to an evolutionarily stable strategy fails in general. See Ross Cressman and Yi Tao, The replicator equation and other game dynamics (2014), Theorems 1–2 and Example 1. These statements refer to stability on the full probability simplex, including perturbations introducing currently absent pure strategies.

A concrete experiment makes the frequency feedback visible. In 2009, Gore, Youk and van Oudenaarden competed two yeast strains: a producer of the enzyme invertase, which enables sucrose hydrolysis, and a mutant lacking the corresponding SUC2 gene. Released sugars can feed neighbours, so avoiding production can be advantageous. They used shaken 5 mL cultures with 5% sucrose at $30{}^{\circ }\mathrm{C}$, diluted daily after 23 hours of growth. Different labelled proteins allowed the two strains to be counted separately by flow cytometry; selective plating periodically checked the counts. The cooperator required externally supplied histidine, so restricting that amino acid imposed an adjustable growth disadvantage. See Snowdrift game dynamics and facultative cheating in yeast, pp. 253 and 255–256, Methods Summary.

Starting with rare producers and, separately, rare nonproducers tested both invasion directions. Under the reported coexistence conditions each rare strain increased; changing histidine or added glucose shifted the eventual composition. Their explanation used nonlinear growth benefits, rather than a fixed pairwise payoff matrix. This motivates testing both rarity regimes before inferring an evolutionarily stable strategy or fitting a replicator equation; it does not identify the illustrative tables below with measured payoffs. See Gore, Youk and van Oudenaarden, Figures 1–3.

An early motivation was understanding why animal contests can involve restraint even when escalation might win an individual encounter. Maynard Smith and Price's ^[ref] introduced the evolutionarily stable strategy in this setting. Taylor and Jonker's ^[ref] then studied explicit continuous and discrete dynamics. Static resistance to invasion and the paths populations follow are related questions, but they require separate definitions.

For a symmetric finite game, pure-action expected payoffs $e(i,p)$ against independent identical opposing mixed strategies are continuous. The displayed map takes the probability simplex continuously into itself. At a Brouwer fixed-point theorem fixed point, put $r_i=[e(i,p)-e(p){]}_{+}$ and $S={\sum }_i{r}_i$. The fixed-point equations give $r_i=p_{i}S$. If $S>0$, every supported action has strictly positive excess payoff, contradicting ${\sum }_i{p}_i(e(i,p)-e(p))=0$. Hence $S=0$ and every supported action is a best response, proving a symmetric Nash equilibrium.

A Nash equilibrium of a multi-person game is a collection of strategies such that each player's strategy is a best response to the others: no unilateral change raises that player's expected payoff. For mixed strategies, every pure action used with positive probability must attain the same maximal payoff. Take $c>0$.

In this costly turnout game, compute the payoff difference between voting and abstaining, holding the others' strategies fixed. The opponent $A$ is pivotal exactly when one of $B,C$ votes. Without $A$ the proposition then passes, while with $A$ the tied ballot fails. The pivotal voting probability is $2\beta (1-\beta )$, soIf $A$ mixes with $0<\alpha <1$, indifference is necessary. Therefore

For supporter $B$, voting is pivotal when $A$ abstains and $C$ abstains, or when $A$ votes and $C$ votes. Thus its voting advantage, and likewise that of $C$, isAt $(\alpha ,\beta )=(1,3/4)$, this equals zero, while ${\Delta }_A=c(16(3/4)(1/4)-3)=0$. Every player is indifferent between voting and abstaining at these opponent strategies; in particular, $A$'s pure vote and both supporters' mixtures are best responses. Hence

It is not unique. At $(\alpha ,\beta )=(0,1/4)$, both supporter advantages are $4c(3/4)-3c=0$, and ${\Delta }_A=0$ again. This gives a second symmetric Nash equilibrium. To see these are the only equilibria with a common supporter probability, note that $\beta =0$ or $1$ makes ${\Delta }_A=-3c$, forcing $\alpha =0$; neither endpoint then satisfies the supporters' best response condition. For $0<\beta <1$, supporter indifference requiresIf $A$ also mixes, its two possible values $\beta =1/4,3/4$ force $\alpha =0,1$ respectively, contradicting strict mixing. Finally $\alpha =0$ gives $\beta =1/4$, and $\alpha =1$ gives $\beta =3/4$. Thus

Allowing $B,C$ to have distinct probabilities $\beta ,\gamma$ gives two additional Nash equilibria. The three voting advantages areIf both supporters mix, subtracting their indifference equations gives $(2\alpha -1)(\gamma -\beta )=0$. At $\alpha =1/2$ a supporter has advantage $-c$, so cannot mix; hence $\beta =\gamma$ and the two symmetric equilibria result. If $B$ always abstains and $C$ mixes, the latter's indifference gives $\alpha =1/4$, and $A$'s indifference gives $\gamma =3/8$. Then ${\Delta }_B=-3c/4<0$, so abstention really is optimal. Interchanging $B,C$ gives the other equilibrium. If $B$ always votes and $C$ mixes, the corresponding equations give $\alpha =3/4$, $\gamma =5/8$, but ${\Delta }_B=-3c/4$, contradicting voting. The four configurations with both supporters pure also fail a best response condition. Consequently the full collection of turnout equilibria with two supporters and one opponent is

A Nash equilibrium is a profile in which each player's strategy is a best response to all the others: no unilateral change increases that player's payoff. In this routing game the payoff can be taken as minus delay. A pure strategy is a single route; a mixed strategy is a probability distribution over routes, with different players randomizing independently. A non-zero-sum game does not require the players' payoffs to add to a fixed total. Nash's theorem guarantees a mixed Nash equilibrium for every finite game with finitely many pure strategies. Pure equilibria need not exist in general; the congestion game here does have them, since a unilateral delay change equals the change in its finite congestion potential.

Put $x=n_1+n_3$ and $y=n_2+n_3$. The three current delays, in the order $ABD,ACD,ABCD$, areFor a player using the first route, switching to the second makes the new second-route load $y+1$, while switching to the third retains that player's use of $AB$ and adds one user to $CD$. The two comparisons are thereforeThey reduce to $x\le y+1$ and $y\ge 74$. Reversing the two outer routes gives $y\le x+1$ and $x\ge 74$ for each second-route user. A third-route user who switches to the first retains their own use of $AB$, and one who switches to the second retains their own use of $CD$. The corresponding comparisons give $y\le 75$ and $x\le 75$. Thus the necessary and sufficient conditions, together with nonnegative integer counts summing to $n$, areThey are sufficient because each used route has been compared with both alternatives, and necessary because each comparison is an available unilateral deviation. The unit changes in loads matter in a finite-player congestion game.

For $n=100$, the counts $(25,25,50)$ give $x=y=75$ and every player's delay is $15/4$. They satisfy all the conditions. To find every pure Nash equilibrium, first note that $n_3=0$ is impossible: if either outer route is empty, its other route users fail the lower-load condition; if both are used, $x,y\ge 74$ contradict $x+y=100$. Thus $n_3>0$, so $x,y\le 75$. Since $x=100-n_2$ and $y=100-n_1$, this forces $n_1,n_2\ge 25$. Both are positive, so the remaining conditions force $x,y\ge 74$ and hence $n_1,n_2\le 26$. Their differences are at most one, so every such pair works. The complete equilibrium count vectors areEach vector also represents many pure profiles obtained by choosing which labeled players use each route. Therefore the displayed equilibrium is not unique, even as an aggregate count vector.

A coordinated allocation with fifty players on each outer route and nobody on $ABCD$ gives every player delay $7/2$, strictly better than $15/4$. It is not a Nash equilibrium: a first-route user can switch to $ABCD$ and obtain delayThis exhibits the conflict between coordinated welfare and unilateral incentives. In fact $7/2$ is below every user's delay in all four pure equilibrium count vectors above.

For a symmetric Nash equilibrium when $n=100$, let each player independently choose the two outer routes with probability $q$ each and the middle route with probability $1-2q$. Conditional on choosing the first route, that player certainly uses $AB$, while each of the other $99$ players uses it with probability $1-q$. The expected delays of the first and third routes are consequentlyThe second has the same expected delay as the first by symmetry. Equalizing these givesAll three probabilities are positive and all three pure routes have expected delay $15/4$, so every route is a best response to the other players' common mixture. This proves a symmetric mixed Nash equilibrium. Using probabilities $1/4,1/4,1/2$ would omit the player's own-load correction and would not make all three expected delays equal.

For completeness, the same calculation works for every positive number of players. If $n\le 75$, every player choosing $ABCD$ is a symmetric pure Nash equilibrium. For $75<n<149$, a symmetric mixed equilibrium hasFor $n\ge 149$, each player choosing either outer route with probability one half is a symmetric mixed equilibrium. In that last regime the unused middle route has expected cost at least that of the outer routes, since their difference is $-3/4+(n+1)/200\ge 0$. These formulas agree at the regime boundaries.

The replicator equation says that a strategy grows in frequency when its expected payoff exceeds the population's current mean payoff. It is an ordinary differential equation for a composition, rather than for total population size. The subtraction of the mean is what makes relative success matter: even a type growing in absolute number can shrink as a fraction if other types grow faster.

Let $x_i(t)$ be the frequency of pure strategy $i$, with $n$ available pure strategies andIn a symmetric bimatrix game, let the payoff matrix $A$ have entry $a_{ij}$ equal to the payoff to strategy $i$ against $j$. The other player's payoff in that encounter is $a_{ji}$: symmetry between player roles does not require $A=A^T$. Uniform random encounters give the expected payoffs and their population meanAfter choosing the time unit so that the growth-rate scale is one, the continuous-time replicator equation isHere a payoff is measured on a scale proportional to net growth per unit time, not merely as an arbitrary preference ranking. More generally one can specify locally Lipschitz frequency-dependent payoffs $f_i(x)$ directly; the same equation with $\bar{f}={\sum }_i{x}_i{f}_i$ then defines a unique frequency trajectory. The linear pairwise model and its reproductive interpretation are given in Taylor and Jonker (1978), sections 2–3; the formulas here also follow by the following direct normalization.

Suppose the number $N_i$ of type $i$ obeys ${\dot{N}}_i=N_i[b(t)+f_i(x)]$, where the baseline $b(t)$ is identical for all types. Set $N={\sum }_i{N}_i>0$ and $x_i=N_i/N$. The quotient rule givesThus the common baseline cancels. Adding the same constant to every entry of $A$ leaves the replicator equation unchanged; multiplying $A$ by a positive constant changes only the speed along trajectories. The same payoff matrix can therefore describe frequency evolution at different growth-rate baselines and time scales.

The probability simplex is invariant in forward time. On ${\sum }_i{x}_i=1$,Normalization alone would not exclude negative frequencies. Nonnegativity follows as well: along a solution,An absent type remains absent, and a positive frequency remains positive at every finite time. For the fixed payoff matrix model, bounded rates on the compact probability simplex give global forward solutions. Frequencies can still tend to zero as $t\to \infty$. Each face is invariant, so this model has no spontaneous introduction of new pure strategies; mutation or outside introductions would require additional terms. For two present types the logarithm of their frequency ratio satisfies $\frac{d}{dt}\log (x_i/x_j)=f_i-f_j$, directly expressing the accumulated payoff advantage.

For two pure strategies, write the composition as $(x,1-x)$, where $x$ is the frequency of strategy 1, and letTheir payoff difference and the resulting scalar ordinary differential equation areIn the interior $0<x<1$, the sign of the payoff difference determines the direction of motion. The endpoints are always equilibrium points, and an additional interior equilibrium point exists precisely when the affine payoff difference has a zero strictly between them. At a simple interior zero $x_{\ast }$, $F^{\prime }(x_{\ast })=x_{\ast }(1-x_{\ast })\Delta f^{\prime }(x_{\ast })$: a negative value gives asymptotic stability, a positive value gives instability. At the endpoints $F^{\prime }(0)=b-d$ and $F^{\prime }(1)=c-a$, with stability understood relative to $[0,1]$. A zero derivative requires examining the actual flow rather than this strict sign test.

For an original coordination game, imagine two conventions that reward matching, with convention 1 giving the larger matching reward. ChooseThe crossing is $x_{\ast }=2/5$. Below it, convention 1 loses frequency; above it, convention 1 gains frequency. The endpoint equilibrium points $0$ and $1$ have one-sided asymptotic stability, while $2/5$ is unstable: the corresponding derivatives are $-2,-3$ and $6/5$, respectively. Every initial $0<x(0)<2/5$ tends to $0$, every initial $2/5<x(0)<1$ tends to $1$, and exactly $x(0)=2/5$ stays there. Even the convention with the larger matching reward can disappear if it starts below the threshold. This is bistability, not selection of a universally best convention.

The compositions giving a symmetric Nash equilibrium in this coordination game are the two pure conventions and the mixed composition $(2/5,3/5)$. Both pure conventions are evolutionarily stable strategies because a mutant convention has strictly lower payoff against the pure resident. The mixed symmetric Nash equilibrium is not an evolutionarily stable strategy: against that resident both conventions tie, but after a pure convention-1 mutant is introduced its payoff advantage is $5(x-2/5)>0$. Equal payoffs at a symmetric Nash equilibrium need not provide resistance to invasion or asymptotic stability.

For an original coexistence model, choose payoffs making each convention advantageous when sufficiently rare:Here $x_{\ast }=3/4$ attracts every initial composition with both types present: $\dot{x}>0$ below it and $\dot{x}<0$ above it. The interior derivative is $F^{\prime }(3/4)=-3/4$, while $F^{\prime }(0)=3$ and $F^{\prime }(1)=1$ make both endpoints unstable to introduction of the other type. The endpoints nevertheless remain stationary if the other type is exactly absent; the attraction statement excludes those boundary initial states.

The sole symmetric Nash equilibrium in this coexistence model is $p=(3/4,1/4)$. It is also an evolutionarily stable strategy. For any different mixed strategy $q=(y,1-y)$, the first comparison in the two-condition criterion for evolutionary stability ties because $Ap=(3/2,3/2)$. Its second comparison is strictly positive:Thus restoring frequency feedback can protect a mixture even though all present types earn equal payoffs at the mixture. Coexistence here means different individuals using different pure strategies; it does not require every individual to randomize with a mixed strategy.

Frequency evolution need not increase the mean payoff. In the coexistence example $\bar{f}=5x-4x^2$, whose maximum occurs at $x=5/8$, whereas the attracting composition is $3/4$. For $5/8<x<3/4$, the frequency $x$ increases while $\frac{d}{dt}\bar{f}=(5-8x)\dot{x}<0$. Individual payoff advantages drive the equation; maximizing collective mean payoff is a different optimization problem.

More generally, a rest point only equalizes payoffs among present types: $x_i>0$ implies $(Ax{)}_i=x^{T}Ax$. A symmetric Nash equilibrium additionally requires $(Ax{)}_i\le x^{T}Ax$ for every absent type. All vertices are rest points even when an absent type could invade. For finite symmetric bimatrix games, a rest point with Lyapunov stability on the full probability simplex is a symmetric Nash equilibrium; an interior trajectory that converges has a symmetric Nash equilibrium as its limit. These are conditional results, not a convergence theorem for every trajectory. See Cressman and Tao (2014), Theorem 1.

The rock paper scissors payoff matrix gives an explicit obstruction to universal convergence:Since $A^T=-A$, $x^{T}Ax=0$, and in the interior $\frac{d}{dt}\log (x_1{x}_2{x}_3)={\sum }_i(Ax{)}_i=0$. Thus $x_1{x}_2{x}_3$ is conserved. Its maximum on the probability simplex occurs only at $(1/3,1/3,1/3)$, the unique symmetric Nash equilibrium, so an interior trajectory starting elsewhere cannot converge to that composition. Every other interior product level is a closed curve without a rest point, giving a periodic orbit. The existence of these cycles is also discussed in Cressman and Tao (2014), Example 1. The useful lesson is to inspect the specified payoff feedback and dynamical system, rather than infer convergence from the existence of an evolutionarily stable strategy or a Nash equilibrium.