We define a general class of models representing natural selection between two alleles. The population size and spatial structure are arbitrary, but fixed. Genetics can be haploid, diploid, or otherwise; reproduction can be asexual or sexual. Biological events (e.g. births, deaths, mating, dispersal) depend in arbitrary fashion on the current population state. Our formalism is based on the idea of genetic sites. Each genetic site resides at a particular locus and houses a single allele. Each individual contains a number of sites equal to its ploidy (one for haploids, two for diploids, etc.). Selection occurs via replacement events, in which alleles in some sites are replaced by copies of others. Replacement events depend stochastically on the population state, leading to a Markov chain representation of natural selection. Within this formalism, we define reproductive value, fitness, neutral drift, and fixation probability, and prove relationships among them. We identify four criteria for evaluating which allele is selected and show that these become equivalent in the limit of low mutation. We then formalize the method of weak selection. The power of our formalism is illustrated with applications to evolutionary games on graphs and to selection in a haplodiploid population.

The recent discovery of zero-determinant strategies for the iterated prisoner's dilemma sparked a surge of interest in the surprising fact that a player can exert unilateral control over iterated interactions. These remarkable strategies, however, are known to exist only in games in which players choose between two alternative actions such as "cooperate" and "defect." Here we introduce a broader class of autocratic strategies by extending zero-determinant strategies to iterated games with more general action spaces. We use the continuous donation game as an example, which represents an instance of the prisoner's dilemma that intuitively extends to a continuous range of cooperation levels. Surprisingly, despite the fact that the opponent has infinitely many donation levels from which to choose, a player can devise an autocratic strategy to enforce a linear relationship between his or her payoff and that of the opponent even when restricting his or her actions to merely two discrete levels of cooperation. In particular, a player can use such a strategy to extort an unfair share of the payoffs from the opponent. Therefore, although the action space of the continuous donation game dwarfs that of the classic prisoner's dilemma, players can still devise relatively simple autocratic and, in particular, extortionate strategies.cooperation | evolutionary game theory | extortion | repeated games G ame theory provides a powerful framework to study interactions between individuals ("players"). Among the most interesting types of interactions are social dilemmas, which result from conflicts of interest between individuals and groups (1, 2). Perhaps the most well-studied model of a social dilemma is the prisoner's dilemma (3). A two-player game with actions, C ("cooperate") and D ("defect"), and payoff matrix,is said to be a prisoner's dilemma if T > R > P > S (4). In a prisoner's dilemma, defection is the dominant action, yet the players can realize higher payoffs from mutual cooperation (R) than they can from mutual defection (P), resulting in a conflict of interest between the individual and the pair, which characterizes social dilemmas. Thus, in a one-shot game (i.e., a single encounter), two opponents have an incentive to defect against one another, but the outcome of mutual defection (the unique Nash equilibrium) is suboptimal for both players. One proposed mechanism for the emergence of cooperation in games such as the prisoner's dilemma is direct reciprocity (5, 6), which entails repeated encounters between players and allows for reciprocation of cooperative behaviors. In an iterated game, a player might forgo the temptation to defect in the present due to the threat of future retaliation-"the shadow of the future"-or the possibility of future rewards for cooperating (4, 7), phenomena for which there is both theoretical and empirical support (8, 9). One example of a strategy for the iterated game is to copy the action of the opponent in the previous round ("tit for tat") (4). Alternatively, a player might choose to re...

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Population structure and spatial heterogeneity are integral components of evolutionary dynamics, in general, and of evolution of cooperation, in particular. Structure can promote the emergence of cooperation in some populations and suppress it in others. Here, we provide results for weak selection to favor cooperation on regular graphs for any configuration, meaning any arrangement of cooperators and defectors. Our results extend previous work on fixation probabilities of rare mutants. We find that for any configuration cooperation is never favored for birth-death (BD) updating. In contrast, for death-birth (DB) updating, we derive a simple, computationally tractable formula for weak selection to favor cooperation when starting from any configuration containing any number of cooperators. This formula elucidates two important features: (i) the takeover of cooperation can be enhanced by the strategic placement of cooperators and (ii) adding more cooperators to a configuration can sometimes suppress the evolution of cooperation. These findings give a formal account for how selection acts on all transient states that appear in evolutionary trajectories. They also inform the strategic design of initial states in social networks to maximally promote cooperation. We also derive general results that characterize the interaction of any two strategies, not only cooperation and defection.

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The environment has a strong influence on a population’s evolutionary dynamics. Driven by both intrinsic and external factors, the environment is subject to continual change in nature. To capture an ever-changing environment, we consider a model of evolutionary dynamics with game transitions, where individuals’ behaviors together with the games that they play in one time step influence the games to be played in the next time step. Within this model, we study the evolution of cooperation in structured populations and find a simple rule: Weak selection favors cooperation over defection if the ratio of the benefit provided by an altruistic behavior, b, to the corresponding cost, c, exceedsk−k′, where k is the average number of neighbors of an individual andk′captures the effects of the game transitions. Even if cooperation cannot be favored in each individual game, allowing for a transition to a relatively valuable game after mutual cooperation and to a less valuable game after defection can result in a favorable outcome for cooperation. In particular, small variations in different games being played can promote cooperation markedly. Our results suggest that simple game transitions can serve as a mechanism for supporting prosocial behaviors in highly connected populations.

Evolutionary game theory is a powerful framework for studying evolution in populations of interacting individuals. A common assumption in evolutionary game theory is that interactions are symmetric, which means that the players are distinguished by only their strategies. In nature, however, the microscopic interactions between players are nearly always asymmetric due to environmental effects, differing baseline characteristics, and other possible sources of heterogeneity. To model these phenomena, we introduce into evolutionary game theory two broad classes of asymmetric interactions: ecological and genotypic. Ecological asymmetry results from variation in the environments of the players, while genotypic asymmetry is a consequence of the players having differing baseline genotypes. We develop a theory of these forms of asymmetry for games in structured populations and use the classical social dilemmas, the Prisoner’s Dilemma and the Snowdrift Game, for illustrations. Interestingly, asymmetric games reveal essential differences between models of genetic evolution based on reproduction and models of cultural evolution based on imitation that are not apparent in symmetric games.

Models in evolutionary game theory traditionally assume symmetric interactions in homogeneous environments. Here, we consider populations evolving in a heterogeneous environment, which consists of patches of different qualities that are occupied by one individual each. The fitness of individuals is not only determined by interactions with others but also by environmental quality. This heterogeneity results in asymmetric interactions where the characteristics of the interaction may depend on an individual's location. Interestingly, in non-varying heterogeneous environments, the long-term dynamics are the same as for symmetric interactions in an average, homogeneous environment. However, introducing environmental feedback between an individual's strategy and the quality of its patch results in rich eco-evolutionary dynamics. Thus, individuals act as ecosystem engineers. The nature of the feedback and the rate of ecological changes can relax or aggravate social dilemmas and promote persistent periodic oscillations of strategy abundance and environmental quality. arXiv:1807.01735v2 [q-bio.PE]