In evolutionary games the fitness of individuals is not constant but depends on the relative abundance of the various strategies in the population. Here we study general games among n strategies in populations of large but finite size. We explore stochastic evolutionary dynamics under weak selection, but for any mutation rate. We analyze the frequency dependent Moran process in well-mixed populations, but almost identical results are found for the Wright-Fisher and Pairwise Comparison processes. Surprisingly simple conditions specify whether a strategy is more abundant on average than 1/n, or than another strategy, in the mutation-selection equilibrium. We find one condition that holds for low mutation rate and another condition that holds for high mutation rate. A linear combination of these two conditions holds for any mutation rate. Our results allow a complete characterization of n*n games in the limit of weak selection.
The multi-population replicator dynamics (RD) can be considered a dynamic approach to the study of multi-player games, where it was shown to be related to Cross' learning, as well as of systems of coevolving populations. However, not all of its equilibria are Nash equilibria (NE) of the underlying game, and neither convergence to an NE nor convergence in general are guaranteed. Although interior equilibria are guaranteed to be NE, no interior equilibrium can be asymptotically stable in the multi-population RD, resulting, e.g., in cyclic orbits around a single interior NE. We introduce a new no…
We model evolution according to an asymmetric game as occurring in multiple finite populations, one for each role in the game, and study the effect of subjecting individuals to stochastic strategy mutations. We show that, when these mutations occur sufficiently infrequently, the dynamics over all population states simplify to an ergodic Markov chain over just the pure population states (where each population is monomorphic). This makes calculation of the stationary distribution computationally feasible. The transition probabilities of this embedded Markov chain involve fixation probabilities o…
We propose a stochastic model in evolutionary game theory where individuals (or subpopulations) can mutate changing their strategies randomly (but rarely) and explore the external environment. This environment affects the selective pressure by modifying the payoff arising from the interactions between strategies. We derive a Fokker-Plank integro-differential equation and provide Monte-Carlo simulations for the Hawks vs Doves game. In particular we show that, in some cases, taking into account the external environment favors the persistence of the low-fitness strategy.
In this paper, we study analytically the statistics of the number of equilibria in pairwise social dilemma evolutionary games with mutation where a game's payoff entries are random variables. Using the replicator-mutator equations, we provide explicit formulas for the probability distributions of the number of equilibria as well as other statistical quantities. This analysis is highly relevant assuming that one might know the nature of a social dilemma game at hand (e.g., cooperation vs coordination vs anti-coordination), but measuring the exact values of its payoff entries is difficult. Our d…
Cooperation is ubiquitous in every level of living organisms. It is known that spatial (network) structure is a viable mechanism for cooperation to evolve. Until recently, it has been difficult to predict whether cooperation can evolve at a network (population) level. To address this problem, Pinheiro et al. proposed a numerical metric, called Average Gradient of Selection (AGoS) in 2012. AGoS can characterize and forecast the evolutionary fate of cooperation at a population level. However, stochastic mutation of strategies was not considered in the analysis of AGoS. Here we analyzed the evolu…
The emergence and maintenance of cooperation has attracted intensive scholarly interest and has been analysed within the framework of evolutionary game theory. The role of innovation, which introduces novel strategies into the population, is a relatively understudied aspect of evolutionary game theory. Here, we investigate the effects of two sources of innovation---mutation and heterogeneous updating rules. These mechanisms allow agents to adopt strategies that do not rely on the imitation of other individuals. The model introduces---in addition to canonical imitation-based strategy updating--…
In this paper, we study the number of equilibria of the replicator-mutator dynamics for both deterministic and random multi-player two-strategy evolutionary games. For deterministic games, using Decartes' rule of signs, we provide a formula to compute the number of equilibria in multi-player games via the number of change of signs in the coefficients of a polynomial. For two-player social dilemmas (namely, the Prisoner's Dilemma, Snowdrift, Stag Hunt, and Harmony), we characterize (stable) equilibrium points and analytically calculate the probability of having a certain number of equilibria wh…
In order to accommodate the empirical fact that population structures are rarely simple, modern studies of evolutionary dynamics allow for complicated and highly-heterogeneous spatial structures. As a result, one of the most difficult obstacles lies in making analytical deductions, either qualitative or quantitative, about the long-term outcomes of evolution. The "structure-coefficient" theorem is a well-known approach to this problem for mutation-selection processes under weak selection, but a general method of evaluating the terms it comprises is lacking. Here, we provide such a method for p…
Cancer cells obtain mutations which rely on the production of diffusible growth factors to confer a fitness benefit. These mutations can be considered cooperative, and studied as public goods games within the framework of evolutionary game theory. The population structure, benefit function and update rule all influence the evolutionary success of cooperators. We model the evolution of cooperation in epithelial cells using the Voronoi tessellation model. Unlike traditional evolutionary graph theory, this allows us to implement global updating, for which birth and death events are spatially deco…
A new approach to understanding evolution [Val09], namely viewing it through the lens of computation, has already started yielding new insights, e.g., natural selection under sexual reproduction can be interpreted as the Multiplicative Weight Update (MWU) Algorithm in coordination games played among genes [CLPV14]. Using this machinery, we study the role of mutation in changing environments in the presence of sexual reproduction. Following [WVA05], we model changing environments via a Markov chain, with the states representing environments, each with its own fitness matrix. In this setting, we…
Evolutionarily stable strategy (ESS) is an important solution concept in game theory which has been applied frequently to biological models. Informally an ESS is a strategy that if followed by the population cannot be taken over by a mutation strategy that is initially rare. Finding such a strategy has been shown to be difficult from a theoretical complexity perspective. We present an algorithm for the case where mutations are restricted to pure strategies, and present experiments on several game classes including random and a recently-proposed cancer model. Our algorithm is based on a mixed-i…
Repeated games have a long tradition in the behavioral sciences and evolutionary biology. Recently, strategies were discovered that permit an unprecedented level of control over repeated interactions by enabling a player to unilaterally enforce linear constraints on payoffs. Here, we extend this theory of "zero-determinant" (or, more generally, "autocratic") strategies to alternating games, which are often biologically more relevant than traditional synchronous games. Alternating games naturally result in asymmetries between players because the first move matters or because players might not m…
Direct reciprocity is a mechanism for sustaining mutual cooperation in repeated social dilemma games, where a player would keep cooperation to avoid being retaliated by a co-player in the future. So-called zero-determinant (ZD) strategies enable a player to unilaterally set a linear relationship between the player's own payoff and the co-player's payoff regardless of the strategy of the co-player. In the present study, we analytically study zero-determinant strategies in finitely repeated (two-person) prisoner's dilemma games with a general payoff matrix. Our results are as follows. First, we …
Game theory deals with strategic interactions among players and evolutionary game dynamics tracks the fate of the players' populations under selection. In this paper, we consider the replicator equation for two-player-two-strategy games involving cooperators and defectors. We modify the equation to include the effect of mutation and also delay that corresponds either to the delayed information about the population state or in realizing the effect of interaction among players. By focusing on the four exhaustive classes of symmetrical games -- the Stag Hunt game, the Snowdrift game, the Prisoner…
Cooperation is ubiquitous at every level of living organisms. It is known that spatial (network) structure is a viable mechanism for cooperation to evolve. A recently proposed numerical metric, average gradient of selection (AGoS), a useful tool for interpreting and visualizing evolutionary dynamics on networks, allows simulation results to be visualized on a one-dimensional phase space. However, stochastic mutation of strategies was not considered in the analysis of AGoS. Here we extend AGoS so that it can analyze the evolution of cooperation where mutation may alter strategies of individuals…
In this paper, I model and study the process of natural selection between all possible mixed strategies in classical two-player two-strategy games. I derive and solve an equation that is a natural generalization of the Taylor-Jonker replicator equation that describes dynamics of pure strategy frequencies. I then investigate the evolution of not only frequencies of pure strategies but also of total distribution of mixed strategies. I show that the process of natural selection of strategies for all games obeys the dynamical Principle of minimum of information gain.
The theory of repeated games analyzes the long-term relationship of interacting players and mathematically reveals the condition of how cooperation is achieved, which is not achieved in a one-shot game. In the repeated prisoner's dilemma (RPD) game with no errors, zero-determinant (ZD) strategies allow a player to unilaterally set a linear relationship between the player's own payoff and the opponent's payoff regardless of the strategy that the opponent implements. In contrast, unconditional strategies such as ALLD and ALLC also unilaterally set a linear payoff relationship. Errors often happe…
It is known that learning of players who interact in a repeated game can be interpreted as an evolutionary process in a population of ideas. These analogies have so far mostly been established in deterministic models, and memory loss in learning has been seen to act similarly to mutation in evolution. We here propose a representation of reinforcement learning as a stochastic process in finite "populations of ideas". The resulting birth-death dynamics has absorbing states and allows for the extinction or fixation of ideas, marking a key difference to mutation-selection processes in finite popul…
Evolutionary game theory is a successful mathematical framework geared towards understanding the selective pressures that affect the evolution of the strategies of agents engaged in interactions with potential conflicts. While a mathematical treatment of the costs and benefits of decisions can predict the optimal strategy in simple settings, more realistic settings such as finite populations, non-vanishing mutations rates, stochastic decisions, communication between agents, and spatial interactions, require agent-based methods where each agent is modeled as an individual, carries its own genes…
In a single-state repeated game, zero-determinant strategies can unilaterally force functions of the payoffs to take values in particular closed intervals. When the explicit use of a determinant is absent from the analysis, they are instead called autocratic. We extend their study to the setting of finite state games with deterministic transitions. For a given game we find that the endpoints of the intervals of enforceable values must satisfy fixed point equations. From these extreme enforceable values we show it is always possible to construct finite memory strategies to enforce a particular …
我们发现Jan，Stauffer和Moseley [Theory in Biosc.，119，166（2000）]对于性的进化，即设计出逃避由于太多有害突变而导致的灭绝的策略，对于在有限种群内成功进化有性个体的稳态种群是足够的，但不是必需的。只要允许每一代中有一个有限的转化为有性生殖的概率，就可以产生一个稳定的有性生殖群体，因为每个个体的有害突变数量有一个上限。对于这个概率的大值，我们发现一个相变到一个间歇性的，多稳定的制度。另一方面，在极慢驱动的限制下，另一个过渡发生到不同的稳态分布，在无性群体中具有较少的有害突变。
We consider the Moran process with two populations competing under an iterated Prisoners' Dilemma in the presence of mutation, and concentrate on the case where there are multiple Evolutionarily Stable Strategies. We perform a complete bifurcation analysis of the deterministic system which arises in the infinite population size. We also study the Master equation and obtain asymptotics for the invariant distribution and metastable switching times for the stochastic process in the case of large but finite population. We also show that the stochastic system has asymmetries in the form of a skew f…
We consider a two-dimensional model of three species in rock-paper-scissors competition and study the self-organisation of the population into fascinating spiraling patterns. Within our individual-based metapopulation formulation, the population composition changes due to cyclic dominance (dominance-removal and dominance-replacement), mutations, and pair-exchange of neighboring individuals. Here, we study the influence of mobility on the emerging patterns and investigate when the pair-exchange rate is responsible for spiral waves to become elusive in stochastic lattice simulations. In particul…
The ultimatum game has been a prominent paradigm in studying the evolution of fairness. It predicts that responders should accept any nonzero offer and proposers should offer the smallest possible amount according to orthodox game theory. However, the prediction strongly contradicts with experimental findings where responders usually reject low offers below $20\%$ and proposers usually make higher offers than expected. To explain the evolution of such fair behaviors, we here introduce empathy in group-structured populations by allowing a proportion $α$ of the population to play empathetic stra…
