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1.  Cyclic dominance in evolutionary games: a review 
Journal of the Royal Society Interface  2014;11(100):20140735.
Rock is wrapped by paper, paper is cut by scissors and scissors are crushed by rock. This simple game is popular among children and adults to decide on trivial disputes that have no obvious winner, but cyclic dominance is also at the heart of predator–prey interactions, the mating strategy of side-blotched lizards, the overgrowth of marine sessile organisms and competition in microbial populations. Cyclical interactions also emerge spontaneously in evolutionary games entailing volunteering, reward, punishment, and in fact are common when the competing strategies are three or more, regardless of the particularities of the game. Here, we review recent advances on the rock–paper–scissors (RPS) and related evolutionary games, focusing, in particular, on pattern formation, the impact of mobility and the spontaneous emergence of cyclic dominance. We also review mean-field and zero-dimensional RPS models and the application of the complex Ginzburg–Landau equation, and we highlight the importance and usefulness of statistical physics for the successful study of large-scale ecological systems. Directions for future research, related, for example, to dynamical effects of coevolutionary rules and invasion reversals owing to multi-point interactions, are also outlined.
PMCID: PMC4191105  PMID: 25232048
cyclical interactions; pattern formation; phase transitions; mobility; coevolution; self-organization
2.  Defection and extortion as unexpected catalysts of unconditional cooperation in structured populations 
Scientific Reports  2014;4:5496.
We study the evolution of cooperation in the spatial prisoner's dilemma game, where besides unconditional cooperation and defection, tit-for-tat, win-stay-lose-shift and extortion are the five competing strategies. While pairwise imitation fails to sustain unconditional cooperation and extortion regardless of game parametrization, myopic updating gives rise to the coexistence of all five strategies if the temptation to defect is sufficiently large or if the degree distribution of the interaction network is heterogeneous. This counterintuitive evolutionary outcome emerges as a result of an unexpected chain of strategy invasions. Firstly, defectors emerge and coarsen spontaneously among players adopting win-stay-lose-shift. Secondly, extortioners and players adopting tit-for-tat emerge and spread via neutral drift among the emerged defectors. And lastly, among the extortioners, cooperators become viable too. These recurrent evolutionary invasions yield a five-strategy phase that is stable irrespective of the system size and the structure of the interaction network, and they reveal the most unexpected mechanism that stabilizes extortion and cooperation in an evolutionary setting.
PMCID: PMC4074784  PMID: 24975112
3.  Optimal interdependence between networks for the evolution of cooperation 
Scientific Reports  2013;3:2470.
Recent research has identified interactions between networks as crucial for the outcome of evolutionary games taking place on them. While the consensus is that interdependence does promote cooperation by means of organizational complexity and enhanced reciprocity that is out of reach on isolated networks, we here address the question just how much interdependence there should be. Intuitively, one might assume the more the better. However, we show that in fact only an intermediate density of sufficiently strong interactions between networks warrants an optimal resolution of social dilemmas. This is due to an intricate interplay between the heterogeneity that causes an asymmetric strategy flow because of the additional links between the networks, and the independent formation of cooperative patterns on each individual network. Presented results are robust to variations of the strategy updating rule, the topology of interdependent networks, and the governing social dilemma, thus suggesting a high degree of universality.
PMCID: PMC3747507  PMID: 23959086
4.  If Cooperation Is Likely Punish Mildly: Insights from Economic Experiments Based on the Snowdrift Game 
PLoS ONE  2013;8(5):e64677.
Punishment may deter antisocial behavior. Yet to punish is costly, and the costs often do not offset the gains that are due to elevated levels of cooperation. However, the effectiveness of punishment depends not only on how costly it is, but also on the circumstances defining the social dilemma. Using the snowdrift game as the basis, we have conducted a series of economic experiments to determine whether severe punishment is more effective than mild punishment. We have observed that severe punishment is not necessarily more effective, even if the cost of punishment is identical in both cases. The benefits of severe punishment become evident only under extremely adverse conditions, when to cooperate is highly improbable in the absence of sanctions. If cooperation is likely, mild punishment is not less effective and leads to higher average payoffs, and is thus the much preferred alternative. Presented results suggest that the positive effects of punishment stem not only from imposed fines, but may also have a psychological background. Small fines can do wonders in motivating us to chose cooperation over defection, but without the paralyzing effect that may be brought about by large fines. The later should be utilized only when absolutely necessary.
PMCID: PMC3669423  PMID: 23741367
5.  Evolutionary dynamics of group interactions on structured populations: a review 
Interactions among living organisms, from bacteria colonies to human societies, are inherently more complex than interactions among particles and non-living matter. Group interactions are a particularly important and widespread class, representative of which is the public goods game. In addition, methods of statistical physics have proved valuable for studying pattern formation, equilibrium selection and self-organization in evolutionary games. Here, we review recent advances in the study of evolutionary dynamics of group interactions on top of structured populations, including lattices, complex networks and coevolutionary models. We also compare these results with those obtained on well-mixed populations. The review particularly highlights that the study of the dynamics of group interactions, like several other important equilibrium and non-equilibrium dynamical processes in biological, economical and social sciences, benefits from the synergy between statistical physics, network science and evolutionary game theory.
PMCID: PMC3565747  PMID: 23303223
cooperation; public goods; pattern formation; self-organization; coevolution
6.  Interdependent network reciprocity in evolutionary games 
Scientific Reports  2013;3:1183.
Besides the structure of interactions within networks, also the interactions between networks are of the outmost importance. We therefore study the outcome of the public goods game on two interdependent networks that are connected by means of a utility function, which determines how payoffs on both networks jointly influence the success of players in each individual network. We show that an unbiased coupling allows the spontaneous emergence of interdependent network reciprocity, which is capable to maintain healthy levels of public cooperation even in extremely adverse conditions. The mechanism, however, requires simultaneous formation of correlated cooperator clusters on both networks. If this does not emerge or if the coordination process is disturbed, network reciprocity fails, resulting in the total collapse of cooperation. Network interdependence can thus be exploited effectively to promote cooperation past the limits imposed by isolated networks, but only if the coordination between the interdependent networks is not disturbed.
PMCID: PMC3560361  PMID: 23378915
7.  Wisdom of groups promotes cooperation in evolutionary social dilemmas 
Scientific Reports  2012;2:576.
Whether or not to change strategy depends not only on the personal success of each individual, but also on the success of others. Using this as motivation, we study the evolution of cooperation in games that describe social dilemmas, where the propensity to adopt a different strategy depends both on individual fitness as well as on the strategies of neighbors. Regardless of whether the evolutionary process is governed by pairwise or group interactions, we show that plugging into the “wisdom of groups” strongly promotes cooperative behavior. The more the wider knowledge is taken into account the more the evolution of defectors is impaired. We explain this by revealing a dynamically decelerated invasion process, by means of which interfaces separating different domains remain smooth and defectors therefore become unable to efficiently invade cooperators. This in turn invigorates spatial reciprocity and establishes decentralized decision making as very beneficial for resolving social dilemmas.
PMCID: PMC3418638  PMID: 22893854
8.  If players are sparse social dilemmas are too: Importance of percolation for evolution of cooperation 
Scientific Reports  2012;2:369.
Spatial reciprocity is a well known tour de force of cooperation promotion. A thorough understanding of the effects of different population densities is therefore crucial. Here we study the evolution of cooperation in social dilemmas on different interaction graphs with a certain fraction of vacant nodes. We find that sparsity may favor the resolution of social dilemmas, especially if the population density is close to the percolation threshold of the underlying graph. Regardless of the type of the governing social dilemma as well as particularities of the interaction graph, we show that under pairwise imitation the percolation threshold is a universal indicator of how dense the occupancy ought to be for cooperation to be optimally promoted. We also demonstrate that myopic updating, due to the lack of efficient spread of information via imitation, renders the reported mechanism dysfunctional, which in turn further strengthens its foundations.
PMCID: PMC3328045  PMID: 22511999
9.  Evolutionary Establishment of Moral and Double Moral Standards through Spatial Interactions 
PLoS Computational Biology  2010;6(4):e1000758.
Situations where individuals have to contribute to joint efforts or share scarce resources are ubiquitous. Yet, without proper mechanisms to ensure cooperation, the evolutionary pressure to maximize individual success tends to create a tragedy of the commons (such as over-fishing or the destruction of our environment). This contribution addresses a number of related puzzles of human behavior with an evolutionary game theoretical approach as it has been successfully used to explain the behavior of other biological species many times, from bacteria to vertebrates. Our agent-based model distinguishes individuals applying four different behavioral strategies: non-cooperative individuals (“defectors”), cooperative individuals abstaining from punishment efforts (called “cooperators” or “second-order free-riders”), cooperators who punish non-cooperative behavior (“moralists”), and defectors, who punish other defectors despite being non-cooperative themselves (“immoralists”). By considering spatial interactions with neighboring individuals, our model reveals several interesting effects: First, moralists can fully eliminate cooperators. This spreading of punishing behavior requires a segregation of behavioral strategies and solves the “second-order free-rider problem”. Second, the system behavior changes its character significantly even after very long times (“who laughs last laughs best effect”). Third, the presence of a number of defectors can largely accelerate the victory of moralists over non-punishing cooperators. Fourth, in order to succeed, moralists may profit from immoralists in a way that appears like an “unholy collaboration”. Our findings suggest that the consideration of punishment strategies allows one to understand the establishment and spreading of “moral behavior” by means of game-theoretical concepts. This demonstrates that quantitative biological modeling approaches are powerful even in domains that have been addressed with non-mathematical concepts so far. The complex dynamics of certain social behaviors become understandable as the result of an evolutionary competition between different behavioral strategies.
Author Summary
Why do friends spontaneously come up with mutually accepted rules, cooperation, and solidarity, while the creation of shared moral standards often fails in large communities? In a “global village”, where everybody may interact with anybody else, it is not worthwhile to punish people who cheat. Moralists (cooperative individuals who undertake punishment efforts) disappear because of their disadvantage compared to cooperators who do not punish (so-called “second-order free-riders”). However, cooperators are exploited by free-riders. This creates a “tragedy of the commons”, where everybody is uncooperative in the end. Yet, when people interact with friends or local neighbors, as most people do, moralists can escape the direct competition with non-punishing cooperators by separating from them. Moreover, in the competition with free-riders, moralists can defend their interests better than non-punishing cooperators. Therefore, while seriously depleted in the beginning, moralists can finally spread all over the world (“who laughs last laughs best effect”). Strikingly, the presence of a few non-cooperative individuals (“deviant behavior”) can accelerate the victory of moralists. In order to spread, moralists may also form an “unholy cooperation” with people having double moral standards, i.e., free-riders who punish non-cooperative behavior, while being uncooperative themselves.
PMCID: PMC2861625  PMID: 20454464

Results 1-9 (9)