The collapse of cooperation in evolving games

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' strategies as well as their payoffs 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 co-evolution of strategies and payoffs in arbitrary iterated games. We show that, as payoffs evolve, a trade-off between the benefits and costs of cooperation precipitates a dramatic loss of cooperation under the Iterated Prisoner's Dilemma; and eventually to evolution away from the Prisoner's Dilemma altogether. The collapse of cooperation is so extreme that the average payoff in a population may decline, even as the potential payoff for mutual cooperation increases. 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 co-evolution of strategies and payoffs in iterated interactions.Comment: 33 pages, 13 figure

Coevolutionary games - a mini review

Prevalence of cooperation within groups of selfish individuals is puzzling in that it contradicts with the basic premise of natural selection. Favoring players with higher fitness, the latter is key for understanding the challenges faced by cooperators when competing with defectors. Evolutionary game theory provides a competent theoretical framework for addressing the subtleties of cooperation in such situations, which are known as social dilemmas. Recent advances point towards the fact that the evolution of strategies alone may be insufficient to fully exploit the benefits offered by cooperative behavior. Indeed, while spatial structure and heterogeneity, for example, have been recognized as potent promoters of cooperation, coevolutionary rules can extend the potentials of such entities further, and even more importantly, lead to the understanding of their emergence. The introduction of coevolutionary rules to evolutionary games implies, that besides the evolution of strategies, another property may simultaneously be subject to evolution as well. Coevolutionary rules may affect the interaction network, the reproduction capability of players, their reputation, mobility or age. Here we review recent works on evolutionary games incorporating coevolutionary rules, as well as give a didactic description of potential pitfalls and misconceptions associated with the subject. In addition, we briefly outline directions for future research that we feel are promising, thereby particularly focusing on dynamical effects of coevolutionary rules on the evolution of cooperation, which are still widely open to research and thus hold promise of exciting new discoveries.Comment: 24 two-column pages, 10 figures; accepted for publication in BioSystem

Comparing reactive and memory-one strategies of direct reciprocity

Direct reciprocity is a mechanism for the evolution of cooperation based on repeated interactions. When individuals meet repeatedly, they can use conditional strategies to enforce cooperative outcomes that would not be feasible in one-shot social dilemmas. Direct reciprocity requires that individuals keep track of their past interactions and find the right response. However, there are natural bounds on strategic complexity: Humans find it difficult to remember past interactions accurately, especially over long timespans. Given these limitations, it is natural to ask how complex strategies need to be for cooperation to evolve. Here, we study stochastic evolutionary game dynamics in finite populations to systematically compare the evolutionary performance of reactive strategies, which only respond to the co-player's previous move, and memory-one strategies, which take into account the own and the co-player's previous move. In both cases, we compare deterministic strategy and stochastic strategy spaces. For reactive strategies and small costs, we find that stochasticity benefits cooperation, because it allows for generous-tit-for-tat. For memory one strategies and small costs, we find that stochasticity does not increase the propensity for cooperation, because the deterministic rule of win-stay, lose-shift works best. For memory one strategies and large costs, however, stochasticity can augment cooperation.Comment: 18 pages, 7 figure

The Evolution of Extortion in Iterated Prisoner's Dilemma Games

Iterated games are a fundamental component of economic and evolutionary game theory. They describe situations where two players interact repeatedly and have the possibility to use conditional strategies that depend on the outcome of previous interactions. In the context of evolution of cooperation, repeated games represent the mechanism of reciprocation. Recently a new class of strategies has been proposed, so called 'zero determinant strategies'. These strategies enforce a fixed linear relationship between one's own payoff and that of the other player. A subset of those strategies are 'extortioners' which ensure that any increase in the own payoff exceeds that of the other player by a fixed percentage. Here we analyze the evolutionary performance of this new class of strategies. We show that in reasonably large populations they can act as catalysts for the evolution of cooperation, similar to tit-for-tat, but they are not the stable outcome of natural selection. In very small populations, however, relative payoff differences between two players in a contest matter, and extortioners hold their ground. Extortion strategies do particularly well in co-evolutionary arms races between two distinct populations: significantly, they benefit the population which evolves at the slower rate - an instance of the so-called Red King effect. This may affect the evolution of interactions between host species and their endosymbionts.Comment: contains 4 figure

Evolution of cooperation in spatial traveler's dilemma game

Traveler's dilemma (TD) is one of social dilemmas which has been well studied in the economics community, but it is attracted little attention in the physics community. The TD game is a two-person game. Each player can select an integer value between $R$ and $M$ ($R < M$) as a pure strategy. If both of them select the same value, the payoff to them will be that value. If the players select different values, say $i$ and $j$ ($R \le i < j \le M$), then the payoff to the player who chooses the small value will be $i+R$ and the payoff to the other player will be $i-R$. We term the player who selects a large value as the cooperator, and the one who chooses a small value as the defector. The reason is that if both of them select large values, it will result in a large total payoff. The Nash equilibrium of the TD game is to choose the smallest value $R$. However, in previous behavioral studies, players in TD game typically select values that are much larger than $R$, and the average selected value exhibits an inverse relationship with $R$. To explain such anomalous behavior, in this paper, we study the evolution of cooperation in spatial traveler's dilemma game where the players are located on a square lattice and each player plays TD games with his neighbors. Players in our model can adopt their neighbors' strategies following two standard models of spatial game dynamics. Monte-Carlo simulation is applied to our model, and the results show that the cooperation level of the system, which is proportional to the average value of the strategies, decreases with increasing $R$ until $R$ is greater than the threshold where cooperation vanishes. Our findings indicate that spatial reciprocity promotes the evolution of cooperation in TD game and the spatial TD game model can interpret the anomalous behavior observed in previous behavioral experiments

Evolutionary games on graphs

Game theory is one of the key paradigms behind many scientific disciplines from biology to behavioral sciences to economics. In its evolutionary form and especially when the interacting agents are linked in a specific social network the underlying solution concepts and methods are very similar to those applied in non-equilibrium statistical physics. This review gives a tutorial-type overview of the field for physicists. The first three sections introduce the necessary background in classical and evolutionary game theory from the basic definitions to the most important results. The fourth section surveys the topological complications implied by non-mean-field-type social network structures in general. The last three sections discuss in detail the dynamic behavior of three prominent classes of models: the Prisoner's Dilemma, the Rock-Scissors-Paper game, and Competing Associations. The major theme of the review is in what sense and how the graph structure of interactions can modify and enrich the picture of long term behavioral patterns emerging in evolutionary games.Comment: Review, final version, 133 pages, 65 figure

Evolutionary consequences of behavioral diversity

Iterated games provide a framework to describe social interactions among groups of individuals. Recent work stimulated by the discovery of "zero-determinant" strategies has rapidly expanded our ability to analyze such interactions. This body of work has primarily focused on games in which players face a simple binary choice, to "cooperate" or "defect". Real individuals, however, often exhibit behavioral diversity, varying their input to a social interaction both qualitatively and quantitatively. Here we explore how access to a greater diversity of behavioral choices impacts the evolution of social dynamics in finite populations. We show that, in public goods games, some two-choice strategies can nonetheless resist invasion by all possible multi-choice invaders, even while engaging in relatively little punishment. We also show that access to greater behavioral choice results in more "rugged " fitness landscapes, with populations able to stabilize cooperation at multiple levels of investment, such that choice facilitates cooperation when returns on investments are low, but hinders cooperation when returns on investments are high. Finally, we analyze iterated rock-paper-scissors games, whose non-transitive payoff structure means unilateral control is difficult and zero-determinant strategies do not exist in general. Despite this, we find that a large portion of multi-choice strategies can invade and resist invasion by strategies that lack behavioral diversity -- so that even well-mixed populations will tend to evolve behavioral diversity.Comment: 26 pages, 4 figure

Small games and long memories promote cooperation

Complex social behaviors lie at the heart of many of the challenges facing evolutionary biology, sociology, economics, and beyond. For evolutionary biologists in particular the question is often how such behaviors can arise \textit{de novo} in a simple evolving system. How can group behaviors such as collective action, or decision making that accounts for memories of past experience, emerge and persist? Evolutionary game theory provides a framework for formalizing these questions and admitting them to rigorous study. Here we develop such a framework to study the evolution of sustained collective action in multi-player public-goods games, in which players have arbitrarily long memories of prior rounds of play and can react to their experience in an arbitrary way. To study this problem we construct a coordinate system for memory-$m$ strategies in iterated $n$-player games that permits us to characterize all the cooperative strategies that resist invasion by any mutant strategy, and thus stabilize cooperative behavior. We show that while larger games inevitably make cooperation harder to evolve, there nevertheless always exists a positive volume of strategies that stabilize cooperation provided the population size is large enough. We also show that, when games are small, longer-memory strategies make cooperation easier to evolve, by increasing the number of ways to stabilize cooperation. Finally we explore the co-evolution of behavior and memory capacity, and we find that longer-memory strategies tend to evolve in small games, which in turn drives the evolution of cooperation even when the benefits for cooperation are low