The evolution of cooperation often depends upon population structure, yet nearly all models of cooperation implicitly assume that this structure remains static. This is a simplifying assumption, because most organisms possess genetic traits that affect their population structure to some degree. These traits, such as a group size preference, affect the relatedness of interacting individuals and hence the opportunity for kin or group selection. We argue that models that do not explicitly consider their evolution cannot provide a satisfactory account of the origin of cooperation, because they cannot explain how the prerequisite population structures arise. Here, we consider the concurrent evolution of genetic traits that affect population structure, with those that affect social behavior. We show that not only does population structure drive social evolution, as in previous models, but that the opportunity for cooperation can in turn drive the creation of population structures that support it. This occurs through the generation of linkage disequilibrium between socio-behavioral and population-structuring traits, such that direct kin selection on social behavior creates indirect selection pressure on population structure. We illustrate our argument with a model of the concurrent evolution of group size preference and social behavior.
While worldwide census data provide statistical evidence that firmly link the population density with several indicators of social welfare, the precise mechanisms underlying these observations are largely unknown. Here we study the impact of population density on the evolution of public cooperation in structured populations and find that the optimal density is uniquely related to the percolation threshold of the host graph irrespective of its topological details. We explain our observations by showing that spatial reciprocity peaks in the vicinity of the percolation threshold, when the emergen…
Evolution of cooperation in the prisoner's dilemma and the public goods game is studied, where initially players belong to two independent structured populations. Simultaneously with the strategy evolution, players whose current utility exceeds a threshold are rewarded by an external link to a player belonging to the other population. Yet as soon as the utility drops below the threshold, the external link is terminated. The rewarding of current evolutionary fitness thus introduces a time-varying interdependence between the two populations. We show that, regardless of the details of the evoluti…
Societies consisting of cooperative individuals seem to require for their continuing success that defectors be policed. The precise connection between punishers and benefits, population structure, and division of labour, however, remains ill-understood. Many models assume costly "peer punishment" to enforce cooperation, but results in the economics literature suggest that this assumption may not be generally valid. In many human and animal societies, there is a division of labour between a purely supportive majority and a dedicated minority of police-like enforcers. Here we present several ext…
In the study of the evolution of cooperation, resource limitations are usually assumed just to provide a finite population size. Recently, however, it has been pointed out that resource limitation may also generate dynamical payoffs able to modify the original structure of the games. Here we study analytically a phase transition from a homogeneous population of defectors when resources are abundant to the survival of unconditional cooperators when resources reduce below a threshold. To this end, we introduce a model of simple agents, with no memory or ability of recognition, interacting in wel…
Indirect reciprocity in which players cooperate with unacquainted other players having good reputations is a mechanism for cooperation in relatively large populations subjected to social dilemma situations. When the population has group structure, as is often found in social networks, players in experiments are considered to show behavior that deviates from existing theoretical models of indirect reciprocity. First, players often show ingroup favoritism (i.e., cooperation only within the group) rather than full cooperation (i.e., cooperation within and across groups), even though the latter is…
The emergence and promotion of cooperation are two of the main issues in evolutionary game theory, as cooperation is amenable to exploitation by defectors, which take advantage of cooperative individuals at no cost, dooming them to extinction. It has been recently shown that the existence of purely destructive agents (termed jokers) acting on the common enterprises (public goods games) can induce stable limit cycles among cooperation, defection, and destruction when infinite populations are considered. These cycles allow for time lapses in which cooperators represent a relevant fraction of the…
Cooperation is a difficult proposition in the face of Darwinian selection. Those that defect have an evolutionary advantage over cooperators who should therefore die out. However, spatial structure enables cooperators to survive through the formation of homogeneous clusters, which is the hallmark of network reciprocity. Here we go beyond this traditional setup and study the spatiotemporal dynamics of cooperation in a population of populations. We use the prisoner's dilemma game as the mathematical model and show that considering several populations simultaneously give rise to fascinating spati…
Here we study the effects of adopting different strategies against different opponent instead of adopting the same strategy against all of them in the prisoner dilemma structured in well-mixed populations. We consider an evolutionary process in which strategies that provide reproductive success are imitated and players replace one of their worst interactions by the new one. We set individuals in a well-mixed population so that network reciprocity effect is excluded and we analyze both synchronous and asynchronous updates. As a consequence of the replacement rule, we show that mutual cooperatio…
We study the evolution of cooperation as a birth-death process in spatially extended populations. The benefit from the altruistic behavior of a cooperator is implemented by decreasing the death rate of its direct neighbors. The cost of cooperation is the increase of a cooperator's death rate proportional to the number of its neighbors. When cooperation has higher cost than benefit, cooperators disappear. Then the dynamics reduces to the contact process with defectors as the single particle type. Increasing the benefit-cost ratio above 1, the extinction transition of the contact process splits …
Microbes providing public goods are widespread in nature despite running the risk of being exploited by free-riders. However, the precise ecological factors supporting cooperation are still puzzling. Following recent experiments, we consider the role of population growth and the repetitive fragmentation of populations into new colonies mimicking simple microbial life-cycles. Individual-based modeling reveals that demographic fluctuations, which lead to a large variance in the composition of colonies, promote cooperation. Biased by population dynamics these fluctuations result in two qualitativ…
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 combined influence of selection and random fluctuations on the evolutionary dynamics of two-strategy ("cooperation" and "defection") games in populations comprising cooperation facilitators. The latter are individuals that support cooperation by enhancing the reproductive potential of cooperators relative to the fitness of defectors. By computing the fixation probability of a single cooperator in finite and well-mixed populations that include a fixed number of facilitators, and by using mean field analysis, we determine when selection promotes cooperation in the important classes …
Beijing University of Posts and Telecommunications（北京邮电大学）；Beihang University（北京航空航天大学）；Dartmouth College（达特茅斯学院）；Geisel School of Medicine at Dartmouth（达特茅斯盖泽尔医学院）；Yale University（耶鲁大学）
Cooperation is ubiquitous in every level of living organisms. It is known that spatial (network) structure is a viable mechanism for cooperation to evolve. Until recently, it has been difficult to predict whether cooperation can evolve at a network (population) level. To address this problem, Pinheiro et al. proposed a numerical metric, called Average Gradient of Selection (AGoS) in 2012. AGoS can characterize and forecast the evolutionary fate of cooperation at a population level. However, stochastic mutation of strategies was not considered in the analysis of AGoS. Here we analyzed the evolu…
Evolutionary games on networks traditionally involve the same game at each interaction. Here we depart from this assumption by considering mixed games, where the game played at each interaction is drawn uniformly at random from a set of two different games. While in well-mixed populations the random mixture of the two games is always equivalent to the average single game, in structured populations this is not always the case. We show that the outcome is in fact strongly dependent on the distance of separation of the two games in the parameter space. Effectively, this distance introduces payoff…
This paper investigates the evolution of strategic play where players drawn from a finite well-mixed population are offered the opportunity to play in a public goods game. All players accept the offer. However, due to the possibility of unforeseen circumstances, each player has a fixed probability of being unable to participate in the game, unlike similar models which assume voluntary participation. We first study how prescribed stochastic opting-out affects cooperation in finite populations. Moreover, in the model, cooperation is favored by natural selection over both neutral drift and defect…
Recent experimental results with humans involved in social dilemma games suggest that cooperation may be a contagious phenomenon and that the selection pressure operating on evolutionary dynamics (i.e., mimicry) is relatively weak. I propose an evolutionary dynamics model that links these experimental findings and evolution of cooperation. By assuming a small fraction of (imperfect) zealous cooperators, I show that a large fraction of cooperation emerges in evolutionary dynamics of social dilemma games. Even if defection is more lucrative than cooperation for most individuals, they often mimic…
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…
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 unex…
Ethnocentrism is a behavioral strategy seen on every scale of social interaction. Game-theory models demonstrate that evolution selects ethnocentrism because it boosts cooperation, which increases reproductive fitness. However, some believe that interethnic unions have the potential to foster universal cooperation and overcome in-group biases in humans. Here, we use agent-based computer simulations to test this hypothesis. Cooperative intergroup mating does lend an advantage to a universal cooperation strategy when the cost/benefit ratio of cooperation is low and local population diversity is …
How cooperation emerges in human societies is both an evolutionary enigma, and a practical problem with tangible implications for societal health. Population structure has long been recognized as a catalyst for cooperation because local interactions enable reciprocity. Analysis of this phenomenon typically assumes bi-directional social interactions, even though real-world interactions are often uni-directional. Uni-directional interactions -- where one individual has the opportunity to contribute altruistically to another, but not conversely -- arise in real-world populations as the result of …
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…
By challenging E. coli with sublethal norfloxacin for 10 days, Henry Lee and James Collins suggests the bacterial altruism leads to the population-wide resistance. By detailedly analyzing experiment data, we suggest that bacterial cooperation leads to population-wide resistance under norfloxacin pressure and simultaneously propose the bacteria shield is the possible feedback mechanism of less resistant bacteria. The bacteria shield is that the less resistant bacteria sacrifice the large number of themselves to consume norfloxacin and then to relieve the norfloxacin burden from highly resistant…
Bacteria populations rely on mechanisms such as quorum sensing to coordinate complex tasks that cannot be achieved by a single bacterium. Quorum sensing is used to measure the local bacteria population density, and it controls cooperation by ensuring that a bacterium only commits the resources for cooperation when it expects its neighbors to reciprocate. This paper proposes a simple model for sharing a resource in a bacterial environment, where knowledge of the population influences each bacterium's behavior. Game theory is used to model the behavioral dynamics, where the net payoff (i.e., uti…
We present a novel approach allowing the study of rare events like fixation under fluctuating environments, modeled as extrinsic noise, in evolutionary processes characterized by the dominance of one species. Our treatment consists of mapping the system onto an auxiliary model, exhibiting metastable species coexistence, that can be analyzed semiclassically. This approach enables us to study the interplay between extrinsic and demographic noise on the statistics of interest. We illustrate our theory by considering the paradigmatic prisoner's dilemma game whose evolution is described by the prob…
Dynamics of a social population is analyzed taking into account some physical constraints on individual behavior and decision making abilities. The model, based on Evolutionary Game Theory, predicts that a population has to pass through a series of different games, e.g as a consequence of environmental fluctuations, in order to develop social cooperation and communication skills. It differs from the general assumption that evolution of cooperation, the so called Cooperation Paradox, can be explained by a single set of rules for intra-population competitions. The developed methods, potentially,…
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 …
Cooperative behavior is widespread in nature, even though cooperating individuals always run the risk to be exploited by free-riders. Population structure effectively promotes cooperation given that a threshold in the level of cooperation was already reached. However, the question how cooperation can emerge from a single mutant, which cannot rely on a benefit provided by other cooperators, is still puzzling. Here, we investigate this question for a well-defined but generic situation based on typical life-cycles of microbial populations where individuals regularly form new colonies followed by …
In a world of hardening borders, nations may deprive themselves of enjoying the benefits of cooperative immigrants. Here, we analyze the effect of efficient cooperative immigrants on a population playing public goods games. We considered a population structured on a square lattice with individuals playing public goods games with their neighbors. The demographics are determined by stochastic birth, death, and migration. The strategies spread through imitation dynamics. Our model shows that cooperation among natives can emerge due to social contagion of good role-model agents that can provide be…
I study two mechanisms based on punishment to promote cooperation in the two-population snowdrift game. The first mechanism follows the traditional approach in the literature and is based on the inclusion of a third additional strategy in the payoff matrix of the stage-game. The second mechanism consists of letting cooperators to punish defectors with a given exogenous frequency. While both mechanisms share the same result regarding the minimum required level of punishment in order to eliminate defectors in both populations, stability in the mechanism following the second approach is more robu…
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…
King Fahd University of Petroleum and Minerals（国王法赫德国立石油与矿产大学）
Classical studies on aspiration-based dynamics suggest that a dissatisfied individual changes strategy without taking into account the success of others. This promotes defection spreading. The imitation-based dynamics allow individuals to imitate successful strategies without taking into account their own-satisfactions. In this article, we propose to study a dynamic based on aspiration which takes into account imitation of successful strategies for dissatisfied individuals. This helps cooperative members to resist. Individuals compare their success to their desired satisfaction level before ma…
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 …
Cooperation among individuals has been key to sustaining societies. However, natural selection favors defection over cooperation. Cooperation can be favored when the mobility of individuals allows cooperators to form a cluster (or group). Mobility patterns of animals sometimes follow a Lévy flight. A Lévy flight is a kind of random walk but it is composed of many small movements with a few big movements. The role of Lévy flights for cooperation has been studied by Antonioni and Tomassini. They showed that Lévy flights promoted cooperation combined with conditional movements triggered by neighb…
Cooperation is ubiquitous ranging from multicellular organisms to human societies. Population structures indicating individuals' limited interaction ranges are crucial to understand this issue. But it is still at large to what extend multiple interactions involving nonlinearity in payoff play a role on cooperation in structured populations. Here we show a rule, which determines the emergence and stabilization of cooperation, under multiple discounted, linear, and synergistic interactions. The rule is validated by simulations in homogenous and heterogenous structured populations. We find that t…
People tend to have their social interactions with members of their own community. Such group-structured interactions can have a profound impact on the behaviors that evolve. Group structure affects the way people cooperate, and how they reciprocate each other's cooperative actions. Past work has shown that population structure and reciprocity can both promote the evolution of cooperation. Yet the impact of these mechanisms has been typically studied in isolation. In this work, we study how the two mechanisms interact. Using a game-theoretic model, we explore how people engage in reciprocal co…
Cooperation is prevalent in nature, not only in the context of social interactions within the animal kingdom, but also on the cellular level. In cancer for example, tumour cells can cooperate by producing growth factors. The evolution of cooperation has traditionally been studied for well-mixed populations under the framework of evolutionary game theory, and more recently for structured populations using evolutionary graph theory. The population structures arising due to cellular arrangement in tissues however are dynamic and thus cannot be accurately represented by either of these frameworks.…
Pro-social punishment, whereby cooperators punish defectors, is often suggested as a mechanism that maintains cooperation in large human groups. Importantly, models that support this idea have to date only allowed defectors to be the target of punishment. However, recent empirical work has demonstrated the existence of anti-social punishment in public goods games. That is, individuals that defect have been found to also punish cooperators. Some recent theoretical studies have found that such anti-social punishment can prevent the evolution of pro-social punishment and cooperation. However, the…
We study the evolution of cooperation in a finite population interacting according to a simple model of like-with-like assortment. Evolution proceeds as a Moran process, and payoffs from the underlying cooperator-defector game are translated to positive fitnesses by an exponential transformation. These evolutionary dynamics can arise, for example, in a nest-structured population with rare migration. The use of the exponential transformation, rather than the usual linear one, is appropriate when interactions have multiplicative fitness effects, and allows for a tractable characterization of the…
The evolution of cooperation has been a perennial problem for evolutionary biology because cooperation is undermined by selfish cheaters (or "free riders") that profit from cooperators but do not invest any resources themselves. In a purely "selfish" view of evolution, those cheaters should be favored. Evolutionary game theory has been able to show that under certain conditions, cooperation nonetheless evolves stably. One of these scenarios utilizes the power of punishment to suppress free riders, but only if players interact in a structured population where cooperators are likely to be surrou…
The emergence and maintenance of cooperation has attracted intensive scholarly interest and has been analysed within the framework of evolutionary game theory. The role of innovation, which introduces novel strategies into the population, is a relatively understudied aspect of evolutionary game theory. Here, we investigate the effects of two sources of innovation---mutation and heterogeneous updating rules. These mechanisms allow agents to adopt strategies that do not rely on the imitation of other individuals. The model introduces---in addition to canonical imitation-based strategy updating--…
We consider a model for the evolution of cooperation in a population where individuals may have one of a number of different heritable and distinguishable markers or tags. Individuals interact with each of their neighbours on a square lattice by either cooperating by donating some benefit at a cost to themselves or defecting by doing nothing. The decision to cooperate or defect is contingent on each individual's perception of its interacting partner's tag. Unlike in other tag-based models individuals do not compare their own tag to that of their interaction partner. That is, there is no {\em s…
Emergence of cooperation in nonlinear higher-order public goods games
Institute for Cross-disciplinary Physics and Complex Systems IFISC (CSIC-UIB)（交叉物理与复杂系统研究所）；Department of Network and Data Science, Central European University Vienna（中欧大学维也纳网络与数据科学系）；Human Evolutionary Ecology Group, Department of Evolutionary Anthropology, University of Zurich（苏黎世大学生物人类学系人类进化生态组）；Center for Networks and Complexity, Luiss University of Rome（罗马LUISS大学网络与复杂性中心）；Department of AI, Data and Decision Sciences, Luiss University of Rome（罗马LUISS大学人工智能、数据与决策科学系）
