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Concept: Evolutionary game theory


We study evolutionary game dynamics on structured populations in which individuals take part in several layers of networks of interactions simultaneously. This multiplex of interdependent networks accounts for the different kind of social ties each individual has. By coupling the evolutionary dynamics of a Prisoner’s Dilemma game in each of the networks, we show that the resilience of cooperative behaviors for extremely large values of the temptation to defect is enhanced by the multiplex structure. Furthermore, this resilience is intrinsically related to a non-trivial organization of cooperation across the network layers, thus providing a new way out for cooperation to survive in structured populations.

Concepts: Structure, Nash equilibrium, Evolutionary game theory, Evolutionarily stable strategy, Game theory, The Evolution of Cooperation, Sociology, Prisoner's dilemma


A tragedy of the commons occurs when individuals take actions to maximize their payoffs even as their combined payoff is less than the global maximum had the players coordinated. The originating example is that of overgrazing of common pasture lands. In game-theoretic treatments of this example, there is rarely consideration of how individual behavior subsequently modifies the commons and associated payoffs. Here, we generalize evolutionary game theory by proposing a class of replicator dynamics with feedback-evolving games in which environment-dependent payoffs and strategies coevolve. We initially apply our formulation to a system in which the payoffs favor unilateral defection and cooperation, given replete and depleted environments, respectively. Using this approach, we identify and characterize a class of dynamics: an oscillatory tragedy of the commons in which the system cycles between deplete and replete environmental states and cooperation and defection behavior states. We generalize the approach to consider outcomes given all possible rational choices of individual behavior in the depleted state when defection is favored in the replete state. In so doing, we find that incentivizing cooperation when others defect in the depleted state is necessary to avert the tragedy of the commons. In closing, we propose directions for the study of control and influence in games in which individual actions exert a substantive effect on the environmental state.

Concepts: Evolutionary dynamics, Evolution and the Theory of Games, Replicator equation, Maxima and minima, Evolutionarily stable strategy, Evolutionary game theory, Game theory


It is often assumed that in public goods games, contributors are either strong or weak players and each individual has an equal probability of exhibiting cooperation. It is difficult to explain why the public good is produced by strong individuals in some cooperation systems, and by weak individuals in others. Viewing the asymmetric volunteer’s dilemma game as an evolutionary game, we find that whether the strong or the weak players produce the public good depends on the initial condition (i.e., phenotype or initial strategy of individuals). These different evolutionarily stable strategies (ESS) associated with different initial conditions, can be interpreted as the production modes of public goods of different cooperation systems. A further analysis revealed that the strong player adopts a pure strategy but mixed strategies for the weak players to produce the public good, and that the probability of volunteering by weak players decreases with increasing group size or decreasing cost-benefit ratio. Our model shows that the defection probability of a “strong” player is greater than the “weak” players in the model of Diekmann (1993). This contradicts Selten’s (1980) model that public goods can only be produced by a strong player, is not an evolutionarily stable strategy, and will therefore disappear over evolutionary time. Our public good model with ESS has thus extended previous interpretations that the public good can only be produced by strong players in an asymmetric game.

Concepts: Evolution and the Theory of Games, Matching pennies, Chicken, Goods, Evolutionary game theory, Evolutionarily stable strategy, Nash equilibrium, Game theory


Abiraterone treats metastatic castrate-resistant prostate cancer by inhibiting CYP17A, an enzyme for testosterone auto-production. With standard dosing, evolution of resistance with treatment failure (radiographic progression) occurs at a median of ~16.5 months. We hypothesize time to progression (TTP) could be increased by integrating evolutionary dynamics into therapy. We developed an evolutionary game theory model using Lotka-Volterra equations with three competing cancer “species”: androgen dependent, androgen producing, and androgen¬†independent. Simulations with standard abiraterone dosing demonstrate strong selection for androgen-independent cells and rapid treatment failure. Adaptive therapy, using patient-specific tumor dynamics to inform on/off treatment cycles, suppresses proliferation of androgen-independent cells and lowers cumulative drug dose. In a pilot clinical trial, 10 of 11 patients maintained stable oscillations of tumor burdens; median TTP is at least 27 months with reduced cumulative drug use of 47% of standard dosing. The outcomes show significant improvement over published studies and a contemporaneous population.

Concepts: Estrogen, Evolutionary game theory, Androgen, Metastasis, Testosterone, Cancer, Game theory, Prostate cancer


Most examples of the application of evolutionary game theory to problems in biology involve highly simplified models. I contend that it is time to move on and include much more richness in models. In particular, more thought needs to be given to the importance of (i) between-individual variation; (ii) the interaction between individuals, and hence the process by which decisions are reached; (iii) the ecological and life-history context of the situation; (iv) the traits that are under selection, and (v) the underlying psychological mechanisms that lead to behaviour. I give examples where including variation between individuals fundamentally changes predicted outcomes of a game. Variation also selects for real-time responses, again resulting in changed outcomes. Variation can select for other traits, such as choosiness and social sensitivity. More generally, many problems involve coevolution of more than one trait. I identify situations where a reductionist approach, in which a game is isolated from is ecological setting, can be misleading. I also highlight the need to consider flexibility of behaviour, mental states and other issues concerned with the evolution of mechanism.

Concepts: Evolution and the Theory of Games, Biology, Evolutionarily stable strategy, Evolutionary game theory, Psychology, Evolution, Natural selection, Game theory


Recent empirical work highlights the heterogeneity of social competitions such as political campaigns: proponents of some ideologies seek debate and conversation, others create echo chambers. While symmetric and static network structure is typically used as a substrate to study such competitor dynamics, network structure can instead be interpreted as a signature of the competitor strategies, yielding competition dynamics on adaptive networks. Here we demonstrate that tradeoffs between aggressiveness and defensiveness (i.e., targeting adversaries vs. targeting like-minded individuals) creates paradoxical behaviour such as non-transitive dynamics. And while there is an optimal strategy in a two competitor system, three competitor systems have no such solution; the introduction of extreme strategies can easily affect the outcome of a competition, even if the extreme strategies have no chance of winning. Not only are these results reminiscent of classic paradoxical results from evolutionary game theory, but the structure of social networks created by our model can be mapped to particular forms of payoff matrices. Consequently, social structure can act as a measurable metric for social games which in turn allows us to provide a game theoretical perspective on online political debates.

Concepts: Strategy, Evolutionarily stable strategy, Scientific method, Evolutionary game theory, Competition, Social network, Sociology, Game theory


Spatial structure is one of the most studied mechanisms in evolutionary game theory. Here, we explore the consequences of spatial structure for a question which has received considerable empirical and theoretical attention in recent years, but has not yet been studied from a network perspective: whether cooperation relies on intuitive predispositions or deliberative self-control. We examine this question using a model which integrates the “dual-process” framework from cognitive science with evolutionary game theory, and considers the evolution of agents who are embedded within a social network and only interact with their neighbors. In line with past work in well-mixed populations, we find that selection favors either the intuitive defector strategy which never deliberates, or the dual-process cooperator strategy which intuitively cooperates but uses deliberation to switch to defection when doing so is payoff-maximizing. We find that sparser networks (i.e., smaller average degree) facilitate the success of dual-process cooperators over intuitive defectors, while also reducing the level of deliberation that dual-process cooperators engage in; and that these results generalize across different kinds of networks. These observations demonstrate the important role that spatial structure can have not just on the evolution of cooperation, but on the co-evolution of cooperation¬†and cognition.

Concepts: Defection, Evolutionarily stable strategy, Evolutionary game theory, Natural selection, Psychology, Science, Scientific method, Game theory


In the animal world, the competition between individuals belonging to different species for a resource often requires the cooperation of several individuals in groups. This paper proposes a generalization of the Hawk-Dove Game for an arbitrary number of agents: the N-person Hawk-Dove Game. In this model, doves exemplify the cooperative behavior without intraspecies conflict, while hawks represent the aggressive behavior. In the absence of hawks, doves share the resource equally and avoid conflict, but having hawks around lead to doves escaping without fighting. Conversely, hawks fight for the resource at the cost of getting injured. Nevertheless, if doves are present in sufficient number to expel the hawks, they can aggregate to protect the resource, and thus avoid being plundered by hawks. We derive and numerically solve an exact equation for the evolution of the system in both finite and infinite well-mixed populations, finding the conditions for stable coexistence between both species. Furthermore, by varying the different parameters, we found a scenario of bifurcations that leads the system from dominating hawks and coexistence to bi-stability, multiple interior equilibria and dominating doves.

Concepts: Evolutionarily stable strategy, Prisoner's dilemma, Chicken, Conflict, Evolutionary game theory, Species, Evolution, Aggression


Game theory provides a quantitative framework for analyzing the behavior of rational agents. The Iterated Prisoner’s Dilemma in particular has become a standard model for studying cooperation and cheating, with cooperation often emerging as a robust outcome in evolving populations. Here we extend evolutionary game theory by allowing players' payoffs as well as their strategies to evolve in response to selection on heritable mutations. In nature, many organisms engage in mutually beneficial interactions and individuals may seek to change the ratio of risk to reward for cooperation by altering the resources they commit to cooperative interactions. To study this, we construct a general framework for the coevolution of strategies and payoffs in arbitrary iterated games. We show that, when there is a tradeoff between the benefits and costs of cooperation, coevolution often leads to a dramatic loss of cooperation in the Iterated Prisoner’s Dilemma. The collapse of cooperation is so extreme that the average payoff in a population can decline even as the potential reward for mutual cooperation increases. Depending upon the form of tradeoffs, evolution may even move away from the Iterated Prisoner’s Dilemma game altogether. Our work offers a new perspective on the Prisoner’s Dilemma and its predictions for cooperation in natural populations; and it provides a general framework to understand the coevolution of strategies and payoffs in iterated interactions.

Concepts: Biology, Nash equilibrium, Evolutionary game theory, Evolution, Natural selection, Evolutionarily stable strategy, Game theory, Prisoner's dilemma


Experiments on the Ultimatum Game (UG) repeatedly show that people’s behaviour is far from rational. In UG experiments, a subject proposes how to divide a pot and the other can accept or reject the proposal, in which case both lose everything. While rational people would offer and accept the minimum possible amount, in experiments low offers are often rejected and offers are typically larger than the minimum, and even fair. Several theoretical works have proposed that these results may arise evolutionarily when subjects act in both roles and there is a fixed interaction structure in the population specifying who plays with whom. We report the first experiments on structured UG with subjects playing simultaneously both roles. We observe that acceptance levels of responders approach rationality and proposers accommodate their offers to their environment. More precisely, subjects keep low acceptance levels all the time, but as proposers they follow a best-response-like approach to choose their offers. We thus find that status equality promotes rational sharing while the influence of structure leads to fairer offers compared to well-mixed populations. Our results are far from what is observed in single-role UG experiments and largely different from available predictions based on evolutionary game theory.

Concepts: Evolutionarily stable strategy, Theory, Evolutionary game theory, Hypothesis, Proposals, Scientific method, Game theory, Proposal