Recent experimental evidence [Grujić et al., PLoS ONE 5, e13749 (2010)] on the spatial Prisoner's Dilemma suggests that players choosing to cooperate or not on the basis of their previous action and the actions of their neighbors coexist with steady defectors and cooperators. We here study the coexistence of these three strategies in the multiplayer iterated Prisoner's Dilemma by means of the replicator dynamics. We consider groups with n = 2, 3, 4 and 5 players and compute the payoffs to every type of player as the limit of a Markov chain where the transition probabilities between actions are found from the corresponding strategies. We show that for group sizes up to n = 4 there exists an interior point in which the three strategies coexist, the corresponding basin of attraction decreasing with increasing number of players, whereas we have not been able to locate such a point for n = 5. We analytically show that in the infinite n limit no interior points can arise. We conclude by discussing the implications of this theoretical approach on the behavior observed in experiments.
Do people discriminate between men and women when they have the option to punish defectors or reward cooperators? Here we report on four pre-registered experiments, that shed some light on this question. Study 1 (N=544) shows that people do not discriminate between genders when they have the option to punish (reward) defectors (cooperators) in a one-shot prisoner's dilemma with third-party punishment/reward. Study 2 (N=253) extends Study 1 to a different method of punishing/rewarding: participants are asked to rate the behaviour of a defector/cooperator on a scale of 1 to 5 stars. In this case…
Traditionally, resource limitation in evolutionary game theory is assumed just to impose a constant population size. Here we show that resource limitations may generate dynamical payoffs able to alter an original prisoner's dilemma, and to allow for the stable coexistence between unconditional cooperators and defectors in well-mixed populations. This is a consequence of a self-organizing process that turns the interaction payoff matrix into evolutionary neutral, and represents a resource-based control mechanism preventing the spread of defectors. To our knowledge, this is the first example of …
In the study of the evolution of cooperation, resource limitations are usually assumed just to provide a finite population size. Recently, however, agent-based models have pointed out that resource limitation may modify the original structure of the interactions and allow for the survival of unconditional cooperators in well-mixed populations. Here, we present analytical simplified versions of two types of agent-based models recently published: one in which the limiting resource constrains the ability of reproduction of individuals but not their survival, and a second one where the limiting re…
We study the evolution of cooperation in spatial public goods games with four competing strategies: cooperators, defectors, punishing cooperators, and punishing defectors. To explore the robustness of the cooperation-promoting effect of costly punishment, besides the usual strategy adoption dynamics we also apply strategy mutations. As expected, frequent mutations create kind of well-mixed conditions, which support the spreading of defectors. However, when the mutation rate is small, the final stationary state does not significantly differ from the state of the mutation-free model, independent…
Consider a cooperation game on a spatial network of habitat patches, where players can relocate between patches if they judge the local conditions to be unfavorable. In time, the relocation events may lead to a homogeneous state where all patches harbor the same relative densities of cooperators and defectors or they may lead to self-organized patterns, where some patches become safe havens that maintain an elevated cooperator density. Here we analyze the transition between these states mathematically. We show that safe havens form once a certain threshold in connectivity is crossed. This thre…
Despite the wealth of empirical and theoretical studies, the origin and maintenance of cooperation is still an evolutionary riddle. In this context, ecological life-history traits which affect the efficiency of selection may play a role, though these are often ignored. We consider here species such as bacteria, fungi, invertebrates and plants which exhibit resting stages in the form of a quiescent state or a seedbank. When quiescent, individuals are inactive and reproduce upon activation, while under seed bank parents produce offspring remaining dormant for different amount of time. We assume …
We study the conditions for persistent cooperation in an off-lattice model of mobile agents playing the Prisoner's Dilemma game with pure, unconditional strategies. Each agent has an exclusion radius rP, which accounts for the population viscosity, and an interaction radius rint, which defines the instantaneous contact network for the game dynamics. We show that, differently from the rP=0 case, the model with finite-sized agents presents a coexistence phase with both cooperators and defectors, besides the two absorbing phases, in which either cooperators or defectors dominate. We provide, in a…
Many mathematical frameworks of evolutionary game dynamics assume that the total population size is constant and that selection affects only the relative frequency of strategies. Here, we consider evolutionary game dynamics in an extended Wright-Fisher process with variable population size. In such a scenario, it is possible that the entire population becomes extinct. Survival of the population may depend on which strategy prevails in the game dynamics. Studying cooperative dilemmas, it is a natural feature of such a model that cooperators enable survival, while defectors drive extinction. Alt…
Cooperation is ubiquitous in nature, but explaining its existence remains a central interdisciplinary challenge. Cooperation is most difficult to explain in the Prisoner's Dilemma game, where cooperators always lose in direct competition with defectors despite increasing mean fitness. Here we demonstrate how spatial population expansion, a widespread natural phenomenon, promotes the evolution of cooperation. We engineer an experimental Prisoner's Dilemma game in the budding yeast Saccharomyces cerevisiae to show that, despite losing to defectors in nonexpanding conditions, cooperators increase…
Economic experiments have shown that punishment can increase public goods game contributions over time. However, the effectiveness of punishment is challenged by second-order freeriding and antisocial punishment. The latter implies that non-cooperators punish cooperators, while the former implies unwillingness to shoulder the cost of punishment. Here we extend the theory of cooperation in the spatial public goods game by considering four competing strategies, which are traditional cooperators and defectors, as well as cooperators who punish defectors and defectors who punish cooperators. We sh…
Social, biological and economic networks grow and decline with occasional fragmentation and re-formation, often explained in terms of external perturbations. We show that these phenomena can be a direct consequence of simple imitation and internal conflicts between 'cooperators' and 'defectors'. We employ a game-theoretic model of dynamic network formation where successful individuals are more likely to be imitated by newcomers who adopt their strategies and copy their social network. We find that, despite using the same mechanism, cooperators promote well-connected highly prosperous networks …
Stability of evolutionary dynamics of non-repeated Prisoner's Dilemma game with non-uniform interaction rates [1], via benefit and cost dilemma is studied . Moreover, the stability condition (b+c/b-c)2 < r1r3 is derived in case of coexistence between cooperators and defectors .If r1,r3 -> infinity cooperation is the dominant strategy and defectors can no longer exploit cooperators.
Finding ways to overcome the temptation to exploit one another is still a challenge in behavioural sciences. In the framework of evolutionary game theory, punishing strategies are frequently used to promote cooperation in competitive environments. Here, we introduce altruistic punishers in the spatial public goods game. This strategy acts as a cooperator in the absence of defectors, otherwise it will punish all defectors in their vicinity while bearing a cost to do so. We observe three distinct behaviours in our model: i) in the absence of punishers, cooperators (who don't punish defectors) ar…
In structured populations the spatial arrangement of cooperators and defectors on the interaction graph together with the structure of the graph itself determines the game dynamics and particularly whether or not fixation of cooperation (or defection) is favored. For a single cooperator (and a single defector) and a network described by a regular graph the question of fixation can be addressed by a single parameter, the structure coefficient. As this quantity is generic for any regular graph, we may call it the generic structure coefficient. For two and more cooperators (or several defectors) …
Dense cooperative networks are an essential element of social capital for a prosperous society. These networks enable individuals to overcome collective action dilemmas by enhancing trust. In many biological and social settings, network structures evolve endogenously as agents exit relationships and build new ones. However, the process by which evolutionary dynamics lead to self-organization of dense cooperative networks has not been explored. Our large group prisoner's dilemma experiments with exit and partner choice options show that core-periphery segregation of cooperators and defectors dr…
According to the standard protocol of spatial public goods game, a cooperator player invests not only into his own game but also into the games organized by neighboring partners. In this work, we relax this assumption by allowing cooperators to decide which neighboring group to prefer instead of supporting them uniformly. In particular, we assume that they select their most successful neighbor and focus external investments exclusively into the related group. We show that this very simple alteration of the dynamical rule results in a surprisingly positive evolutionary outcome -- cooperators pr…
We consider a stochastic model for evolution of group-structured populations in which interactions between group members correspond to the Prisoner's Dilemma or the Hawk-Dove game. Selection operates at two organization levels: individuals compete with peer group members based on individual payoff, while groups also compete with other groups based on average payoff of group members. In the Prisoner's Dilemma, this creates a tension between the two levels of selection, as defectors are favored at the individual level, whereas groups with at least some cooperators outperform groups of defectors …
Cooperation is beneficial for the species as a whole, but, at the level of an individual, defection pays off. Natural selection is then expected to favor defectors and eliminate cooperation. This prediction is in stark contrast with the abundance of cooperation at all levels of biological systems: from cells cooperating to form a biofilm or an organism to ecosystems and human societies. Several explanations have been proposed to resolve this paradox, including direct reciprocity, kin, and group selection. However, our work builds upon an observation that selection on cooperators might depend b…
Deliberate deceptiveness intended to gain an advantage is commonplace in human and animal societies. In a social dilemma, an individual may only pretend to be a cooperator to elicit cooperation from others, while in reality he is a defector. With this as motivation, we study a simple variant of the evolutionary prisoner's dilemma game entailing deceitful defectors and conditional cooperators that lifts the veil on the impact of such two-faced behavior. Defectors are able to hide their true intentions at a personal cost, while conditional cooperators are probabilistically successful at identify…
Ecology and evolution are inherently linked, and studying a mathematical model that considers both holds promise of insightful discoveries related to the dynamics of cooperation. In the present article, we use the prisoner's dilemma (PD) game as a basis for long-term apprehension of the essential social dilemma related to cooperation among unrelated individuals. We upgrade the contemporary PD game with an inclusion of evolution-induced act of punishment as a third competing strategy in addition to the traditional cooperators and defectors. In a population structure, the abundance of ecological…
Range expansion is a universal process in biological systems, and therefore plays a part in biological evolution. Using a quantitative individual-based method based on the stochastic process, we identify that enhancing the inherent self-proliferation advantage of cooperators relative to defectors is a more effective channel to promote the evolution of cooperation in range expansion than weakening the benefit acquisition of defectors from cooperators. With this self-proliferation advantage, cooperators can rapidly colonize virgin space and establish spatial segregation more readily, which acts …
Limitation of resources has been recently introduced as a mechanism for the survival and coexistence of cooperators with defectors in well-mixed populations. Here we examine the same model on a scale-free network. A prisoner's dilemma game on a scale-free network has shown coexistence of cooperators and defectors for the entire range of parameters. Our results show that by introducing the network to the limited resources model, the cooperator-dominated region in the parameter space expands comparing to the results of well-mixed population and the coexistence region becomes narrower. The effect…
Whether or not cooperation is favored over defection in evolutionary games can be assigned by structure coefficients for any arrangement of cooperators and defectors on any network modeled as a regular graph. We study how these structure coefficients relate to a scaling of dilemma strength in social dilemma games. It is shown that some graphs permit certain arrangements of cooperators and defectors to possess particularly large structure coefficients. Moreover, these large coefficients imply particularly large sections of a bounded parameter plane spanned by scaling gamble-intending and risk-a…
The functioning of animal as well as human societies fundamentally relies on cooperation. Yet, defection is often favorable for the selfish individual, and social dilemmas arise. Selection by individuals' fitness, usually the basic driving force of evolution, quickly eliminates cooperators. However, evolution is also governed by fluctuations that can be of greater importance than fitness differences, and can render evolution effectively neutral. Here, we investigate the effects of selection versus fluctuations in social dilemmas. By studying the mean extinction times of cooperators and defecto…
We study the evolutionary dynamics of the prisoner's dilemma game in which cooperators and defectors interact with another actor type called exiters. Rather than being exploited by defectors, exiters exit the game in favour of a small payoff. We find that this simple extension of the game allows cooperation to flourish in well-mixed populations when iterations or reputation are added. In networked populations, however, the exit option is less conducive to cooperation. Instead, it enables the coexistence of cooperators, defectors, and exiters through cyclic dominance. Other outcomes are also po…
The fact that individuals will most likely behave differently in different situations begets the introduction of conditional strategies. Inspired by this, we study the evolution of cooperation in the spatial public goods game, where besides unconditional cooperators and defectors, also different types of conditional cooperators compete for space. Conditional cooperators will contribute to the public good only if other players within the group are likely to cooperate as well, but will withhold their contribution otherwise. Depending on the number of other cooperators that are required to elicit…
When the interactions between cooperators (C) and defectors (D) can be partially avoided within a population, there may be an overall enhancement of cooperation. One example of such screening mechanism occurs in the presence of risk-averse agents (loners, L) that are neutral towards others, i.e., both L and its opponent, whatever its strategy, receive the same payoff. Their presence in the Prisoner's Dilemma (PD) game sustains the coexistence of cooperators and defectors far beyond the level attained in their absence. Another screening mechanism is a heterogeneous landscape obtained, for examp…
Synchronization, cooperation, and chaos are ubiquitous phenomena in nature. In a population composed of many distinct groups of individuals playing the prisoner's dilemma game, there exists a migration dilemma: No cooperator would migrate to a group playing the prisoner's dilemma game lest it should be exploited by a defector; but unless the migration takes place, there is no chance of the entire population's cooperator-fraction to increase. Employing a randomly rewired coupled map lattice of chaotic replicator maps, modelling replication-selection evolutionary game dynamics, we demonstrate th…
Here we consider a game theoretic model of multilevel selection in which individuals compete based on their payoff and groups also compete based on the average payoff of group members. Our focus is on multilevel social dilemmas: games in which individuals are best off cheating, while groups of individuals do best when composed of many cooperators. We analyze the dynamics of the two-level replicator dynamics, a nonlocal hyperbolic PDE describing deterministic birth-death dynamics for both individuals and groups. While past work on such multilevel dynamics has restricted attention to scenarios w…
Utilizing common resources is always a dilemma for community members. While cooperator players restrain themselves and consider the proper state of resources, defectors demand more than their supposed share for a higher payoff. To avoid the tragedy of the common state, punishing the latter group seems to be an adequate reaction. This conclusion, however, is less straightforward when we acknowledge the fact that resources are finite and even a renewable resource has limited growing capacity. To clarify the possible consequences, we consider a coevolutionary model where beside the payoff-driven …
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 single, randomly placed mutants. We find that for any configuration cooperation is never favored for birth-death (BD) updat…
One of the most direct human mechanisms of promoting cooperation is rewarding it. We study the effect of sharing a reward among cooperators in the most stringent form of social dilemma, namely the Prisoner's Dilemma. Specifically, for a group of players that collect payoffs by playing a pairwise Prisoner's Dilemma game with their partners, we consider an external entity that distributes a fixed reward equally among all cooperators. Thus, individuals confront a new dilemma: on the one hand, they may be inclined to choose the shared reward despite the possibility of being exploited by defectors;…
Inspired by the fact that people have diverse propensities to punish wrongdoers, we study a spatial public goods game with defectors and different types of punishing cooperators. During the game, cooperators punish defectors with class-specific probabilities and subsequently share the associated costs of sanctioning. We show that in the presence of different punishing cooperators the highest level of public cooperation is always attainable through a selection mechanism. Interestingly, the selection not necessarily favors the evolution of punishers who would be able to prevail on their own agai…
In the framework of the paradigmatic prisoner's dilemma, we investigate the evolutionary dynamics of social dilemmas in the presence of "cooperation facilitators". In our model, cooperators and defectors interact as in the classical prisoner's dilemma game, where selection favors defection. However, here the presence of a small number of cooperation facilitators enhances the fitness (reproductive potential) of cooperators, while it does not alter that of defectors. In a finite population of size N, the dynamics of the prisoner's dilemma with facilitators is characterized by the probability tha…
Diffusion-induced instabilities promote cooperation in eco-evolutionary networks
Bagchi School of Public Health, Ahmedabad University（艾哈迈达巴德大学巴吉公共卫生学院）；Department of Physical Sciences, Indian Institute of Science Education and Research Kolkata（印度科学教育与研究学院加尔各答分校物理科学系）；Department of Mathematics and Namur Institute for Complex Systems - naXys, University of Namur（南纳尔大学数学与南纳尔复杂系统研究所-naXys）；Faculty of Natural Sciences and Mathematics, University of Maribor（马里博尔大学自然科学与数学学院）；Community Healthcare Center Dr. Adolf Drolc Maribor（马里博尔阿道夫·德罗尔克博士社区健康中心）；Department of Physics, Kyung Hee University（庆熙大学物理系）；University College, Korea University（韩国大学综合研究生院）；Physics and Applied Mathematics Unit, Indian Statistical Institute（印度统计物理学与应用数学单元）
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 distingui…
Cooperation is the backbone of modern human societies, making it a priority to understand how successful cooperation-sustaining mechanisms operate. Cyclic dominance, a non-transitive setup comprising at least three strategies wherein the first strategy overrules the second which overrules the third which, in turn, overrules the first strategy, is known to maintain bio-diversity, drive competition between bacterial strains, and preserve cooperation in social dilemmas. Here, we present a novel route to cyclic dominance in voluntary social dilemmas by adding to the traditional mix of cooperators,…
Cooperators that refuse to participate in sanctioning defectors create the second-order free-rider problem. Such cooperators will not be punished because they contribute to the public good, but they also eschew the costs associated with punishing defectors. Altruistic punishers - those that cooperate and punish - are at a disadvantage, and it is puzzling how such behaviour has evolved. We show that sharing the responsibility to sanction defectors rather than relying on certain individuals to do so permanently can solve the problem of costly punishment. Inspired by the fact that humans have str…
The basic social dilemma is frequently captured by a public goods game where participants decide simultaneously whether to support a common pool or not and after the enhanced contributions are distributed uniformly among all competitors. What if the result of common efforts is {\it not} distributed immediately, but it is reinvested and added to the pool for a next round? This extension may not only result in an enhanced benefit for group members but also opens new strategies for involved players because they may act in distinct rounds differently. In this work we focus on the simplest case whe…
The emergence and maintenance of cooperative behavior is a fascinating topic in evolutionary biology and social science. The public goods game (PGG) is a paradigm for exploring cooperative behavior. In PGG, the total resulting payoff is divided equally among all participants. This feature still leads to the dominance of defection without substantially magnifying the public good by a multiplying factor. Much effort has been made to explain the evolution of cooperative strategies, including a recent model in which only a portion of the total benefit is shared by all the players through introduci…
We study the problem of the emergence of cooperation in the spatial Prisoner's Dilemma. The pioneering work by Nowak and May showed that large initial populations of cooperators can survive and sustain cooperation in a square lattice with imitate-the-best evolutionary dynamics. We revisit this problem in a cost-benefit formulation suitable for a number of biological applications. We show that if a fixed-amount reward is established for cooperators to share, a single cooperator can invade a population of defectors and form structures that are resilient to re-invasion even if the reward mechanis…
In the traditional setup of public goods game all players are involved in every available groups and the mutual benefit is shared among competing cooperator and defector strategies. But in real life situations the group formation of players could be more sophisticated because not all players are attractive enough for others to participate in a joint venture. What if only those players can initiate a group formation and establish a game who are successful enough to the neighbors? To elaborate this idea we employ a modified protocol and demonstrate that a carefully chosen threshold to establish …
Populations composed of a collection of subpopulations (demes) with random migration between them are quite common occurrences. The emergence and sustenance of cooperation in such a population is a highly researched topic in the evolutionary game theory. If the individuals in every deme are considered to be either cooperators or defectors, the migration dilemma can be envisaged: The cooperators would not want to migrate to a defector-rich deme as they fear of facing exploitation; but without migration, cooperation can not be established throughout the network of demes. With a view to studying …
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 th…
We present an extensive, systematic study of the Prisoner's Dilemma and Snowdrift games on a square lattice under a synchronous, noiseless imitation dynamics. We show that for both the occupancy of the network and the (random) mobility of the agents there are intermediate values that may increase the amount of cooperators in the system and new phases appear. We analytically determine the transition lines between these phases and compare with the mean field prediction and the observed behavior on a square lattice. We point out which are the more relevant microscopic processes that entitle coope…
Learning from a partner who collects higher payoff is a frequently used working hypothesis in evolutionary game theory. One of the alternative dynamical rules is when the focal player prefers to follow the strategy choice of the majority in the local neighborhood, which is often called as conformity-driven strategy update. In this work we assume that both strategy learning methods are present and compete for space within the framework of a coevolutionary model. Our results reveal that the presence of payoff-driven strategy learning method becomes exclusive for high succer's payoff and/or high …
Tolerance implies enduring trying circumstances with a fair and objective attitude. To determine whether evolutionary advantages might be stemming from diverse levels of tolerance in a population, we study a spatial public goods game, where in addition to cooperators, defectors, and loners, tolerant players are also present. Depending on the number of defectors within a group, a tolerant player can either cooperate in or abstain from a particular instance of the game. We show that the diversity of tolerance can give rise to synergistic effects, wherein players with a different threshold in ter…
Strategy changes are an essential part of evolutionary games. Here we introduce a simple rule that, depending on the value of a single parameter $w$, influences the selection of players that are considered as potential sources of the new strategy. For positive $w$ players with high payoffs will be considered more likely, while for negative $w$ the opposite holds. Setting $w$ equal to zero returns the frequently adopted random selection of the opponent. We find that increasing the probability of adopting the strategy from the fittest player within reach, i.e. setting $w$ positive, promotes the …
We have studied the evolution of strategies in spatial public goods games where both individual (peer) and institutional (pool) punishments are present beside unconditional defector and cooperator strategies. The evolution of strategy distribution is governed by imitation based on random sequential comparison of neighbors' payoff for a fixed level of noise. Using numerical simulations we have evaluated the strategy frequencies and phase diagrams when varying the synergy factor, punishment cost, and fine. Our attention is focused on two extreme cases describing all the relevant behaviors in suc…
Aging is always present, tailoring our interactions with others and postulating a finite lifespan during which we are able to exercise them. We consider the prisoner's dilemma game on a square lattice, and examine how quenched age distributions and different aging protocols influence the evolution of cooperation when taking the life experience and knowledge accumulation into account as time passes. In agreement with previous studies, we find that a quenched assignment of age to players, introducing heterogeneity to the game, substantially promotes cooperative behavior. Introduction of aging an…
Wealthy individuals may be less tempted to defect than those with comparatively low payoffs. To take this into consideration, we introduce coevolutionary success-driven multigames in structured populations. While the core game is always the weak prisoner's dilemma, players whose payoffs from the previous round exceed a threshold adopt only a minimally low temptation to defect in the next round. Along with the strategies, the perceived strength of the social dilemma thus coevolves with the success of each individual player. We show that the lower the threshold for using the small temptation to …
Punishment and partner switching are two well-studied mechanisms that support the evolution of cooperation. Observation of human behaviour suggests that the extent to which punishment is adopted depends on the usage of alternative mechanisms, including partner switching. In this study, we investigate the combined effect of punishment and partner switching in evolutionary prisoner's dilemma games conducted on a network. In the model, agents are located on the network and participate in the prisoner's dilemma games with punishment. In addition, they can opportunistically switch interaction partn…
