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 evolutionary game and the interaction structure, the self-organization of fitness and reward gives rise to distinguished players that act as strong catalysts of cooperative behavior. However, there also exist critical utility thresholds beyond which distinguished players are no longer able to percolate. The interdependence between the two populations then vanishes, and cooperators are forced to rely on traditional network reciprocity alone. We thus demonstrate that a simple strategy-independent form of rewarding may significantly expand the scope of cooperation on structured populations. The formation of links outside the immediate community seems particularly applicable in human societies, where an individual is typically member in many different social networks.
Rewarding cooperation is in many ways expected behaviour from social players. However, strategies that promote antisocial behaviour are also surprisingly common, not just in human societies, but also among eusocial insects and bacteria. Examples include sanctioning of individuals who behave prosocially, or rewarding of freeriders who do not contribute to collective enterprises. We therefore study the public goods game with antisocial and prosocial pool rewarding in order to determine the potential negative consequences on the effectiveness of positive incentives to promote cooperation. Contrar…
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…
It is well known that cooperation cannot be an evolutionary stable strategy for a non-iterative game in a well-mixed population. In contrast, structured populations favor cooperation since cooperators can benefit each other by forming local clusters. Previous studies have shown that scale-free networks strongly promote cooperation. However, little is known about the invasion mechanism of cooperation in scale-free networks. To study microscopic and macroscopic behaviors of cooperators' invasion, we conducted computational experiments of the evolution of cooperation in scale-free networks where,…
Our wellbeing depends as much on our personal success, as it does on the success of our society. The realization of this fact makes cooperation a very much needed trait. Experiments have shown that rewards can elevate our readiness to cooperate, but since giving a reward inevitably entails paying a cost for it, the emergence and stability of such behavior remain elusive. Here we show that allowing for the act of rewarding to self-organize in dependence on the success of cooperation creates several evolutionary advantages that instill new ways through which collaborative efforts are promoted. R…
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 spatial arrangement of individuals is thought to overcome the dilemma of cooperation: When cooperators engage in clusters they might share the benefit of cooperation while being more protected against non-cooperating individuals, which benefit from cooperation but save the cost of cooperation. This is paradigmatically shown by the spatial prisoner's dilemma model. Here, we study this model in one and two spatial dimensions, but explicitly take into account that in biological setups fitness collection and selection are separated processes occurring mostly on vastly different time scales. Th…
Inclusive Fitness Theory (IFT) was proposed half a century ago by W.D. Hamilton to explain the emergence and maintenance of cooperation between individuals that allows the existence of society. Contemporary evolutionary ecology identified several factors that increase inclusive fitness, in addition to kin-selection, such as assortation or homophily, and social synergies triggered by cooperation. Here we propose an Extend Inclusive Fitness Theory (EIFT) that includes in the fitness calculation all direct and indirect benefits an agent obtains by its own actions, and through interactions with ki…
Evolution depends on the fitness of organisms, the expected rate of reproducing. Directly getting offspring is the most basic form of fitness, but fitness can also be increased indirectly by helping genetically related individuals (such as kin) to increase their fitness. The combined effect is known as inclusive fitness. Here it is argued that a further elaboration of fitness has evolved, particularly in humans. It is called extensive fitness and it incorporates producing organisms that are merely similar in phenotype. The evolvability of this mechanism is illustrated by computations on a simp…
In spatial evolutionary games the fitness of each individual is traditionally determined by the payoffs it obtains upon playing the game with its neighbors. Since defection yields the highest individual benefits, the outlook for cooperators is gloomy. While network reciprocity promotes collaborative efforts, chances of averting the impending social decline are slim if the temptation to defect is strong. It is therefore of interest to identify viable mechanisms that provide additional support for the evolution of cooperation. Inspired by the fact that the environment may be just as important as…
Cooperation is a persistent behavioral pattern of entities pooling and sharing resources. Its ubiquity in nature poses a conundrum. Whenever two entities cooperate, one must willingly relinquish something of value to the other. Why is this apparent altruism favored in evolution? Classical solutions assume a net fitness gain in a cooperative transaction which, through reciprocity or relatedness, finds its way back from recipient to donor. We seek the source of this fitness gain. Our analysis rests on the insight that evolutionary processes are typically multiplicative and noisy. Fluctuations ha…
We compare the likelihood of different socially relevant features to allow the evolutionary emergence and maintenance of cooperation in a generalized variant of the iterated Prisoners Dilemma game. Results show that the average costs/benefit balance of cooperation is the primary constraint for its establishment and maintenance. Behavior increasing inclusive fitness such as assortation, homophily, kin-selection and tagging of individuals, is second in importance. Networks characteristics were the least important in favoring the establishment and maintenance of cooperation, despite being the mos…
Here, we study the evolution of specialization using realistic computer simulations of bacteria that secrete two public goods in a dynamic fluid. Through this first principles approach, we find physical factors such as diffusion, flow patterns, and decay rates are as influential as fitness economics in governing the evolution of community structure, to the extent that when mechanical factors are taken into account, (1) Generalist communities can resist becoming specialists, despite the invasion fitness of specialization, (2) Generalist and specialists can both resist cheaters despite the invas…
Without contributing, defectors take more benefit from social resources than cooperators which is the reflection of a specific character of individuals. However, natural physical mechanisms of our society promote cooperation. Thus, in the long run, the evolution about genetic variation is something more than the social evolution about fitness. The loci of evolutionary paths of the cooperation and the defection are correlated, but not a full complement of each other. Yet, the only single specific mechanism which is operated by some rules explains the enhancement of cooperation where the indepen…
The hypothesis that living systems can benefit from operating at the vicinity of critical points has gained momentum in recent years. Criticality may confer an optimal balance between exceedingly ordered and too noisy states. We here present a model, based on information theory and statistical mechanics, illustrating how and why a community of agents aimed at understanding and communicating with each other converges to a globally coherent state in which all individuals are close to an internal critical state, i.e. at the borderline between order and disorder. We study --both analytically and c…
Evolutionary games on graphs have been extensively studied. A variety of graph structures, graph dynamics, and behaviours of replicators have been explored. These models have primarily been studied in the framework of facilitation of cooperation, and much previous research has shed light on this field of study. However, there has been little attention devoted to truncation selection as most models employ proportional selection (such as in the replicator equation) or `imitate the best.' Here we systematically explore truncation selection on periodic square lattices, where replicators below a fi…
Working together in groups may be beneficial if compared to isolated efforts. Yet this is true only if all group members contribute to the success. If not, group efforts may act detrimentally on the fitness of their members. Here we study the evolution of cooperation in public goods games on scale-free networks that are subject to deletion of links that are connected to the highest-degree individuals, i.e., on networks that are under attack. We focus on the case where all groups a player belongs to are considered for the determination of payoffs; the so-called multi-group public goods games. W…
Multilevel selection is an important organizing principle that crucially underlies evolutionary processes from the emergence of cells to eusociality and the economics of nations. Previous studies on multilevel selection assumed that the effective higher-level selection emerges from lower-level reproduction. This leads to selection among groups, although only individuals reproduce. We introduce selective group extinction, where groups die with a probability inversely proportional to their group fitness. When accounting for this the critical benefit-to-cost ratio is substantially lowered. Becaus…
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）
Economic experiments reveal that humans value cooperation and fairness. Punishing unfair behavior is therefore common, and according to the theory of strong reciprocity, it is also directly related to rewarding cooperative behavior. However, empirical data fail to confirm that positive and negative reciprocity are correlated. Inspired by this disagreement, we determine whether the combined application of reward and punishment is evolutionary advantageous. We study a spatial public goods game, where in addition to the three elementary strategies of defection, rewarding and punishment, a fourth …
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…
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 …
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 …
This paper discusses the role of opportunistic punisher who may act selfishly to free-ride cooperators or not to be exploited by defectors. To consider opportunistic punisher, we make a change to the sequence of one-shot public good game; instead of putting action choice first before punishment, the commitment of punishment is declared first before choosing the action of each participant. In this commitment-first setting, punisher may use information about her team, and may defect to increase her fitness in the team. Reversing sequence of public good game can induce different behavior of punis…
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…
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…
Both conventional wisdom and empirical evidence suggests that arranging a prior commitment or agreement before an interaction enhances the chance of reaching mutual cooperation. Yet it is not clear what mechanisms can promote the participation in and compliance with such a commitment, especially when the former is costly and deviating from the latter is profitable. Prior work either considers regimented commitments where compensation is assumed enforceable from dishonest committers, or assume implicit commitments from every individual (so they are all in and thus being treated as such). Here w…
According to the evolutionary death-birth protocol, a player is chosen randomly to die and neighbors compete for the available position proportional to their fitness. Hence, the status of the focal player is completely ignored and has no impact on the strategy update. In this work, we revisit and generalize this rule by introducing a weight factor to compare the payoff values of the focal and invading neighbors. By means of evolutionary graph theory, we analyze the model on joint transitive graphs to explore the possible consequences of the presence of a weight factor. We find that focal weigh…
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 …
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 …
Starting with Darwin, biologists have asked how populations evolve from a low fitness state that is evolutionarily stable to a high fitness state that is not. Specifically of interest is the emergence of cooperation and multicellularity where the fitness of individuals often appears in conflict with that of the population. Theories of social evolution and evolutionary game theory have produced a number of fruitful results employing two-state two-body frameworks. In this study we depart from this tradition and instead consider a multi-player, multi-state evolutionary game, in which the fitness …
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…
Situations involving cooperative behaviour are widespread among animals and humans alike. Game theory and evolutionary dynamics have provided the theoretical and computational grounds to understand what are the mechanisms that allow for such cooperation. Studies in this area usually take into consideration different behavioural strategies and investigate how they can be fixed in the population under evolving rules. However, how those strategies emerged from basic evolutionary mechanisms continues to be not fully understood. To address this issue, here we study the emergence of cooperative stra…
Cancer cells obtain mutations which rely on the production of diffusible growth factors to confer a fitness benefit. These mutations can be considered cooperative, and studied as public goods games within the framework of evolutionary game theory. The population structure, benefit function and update rule all influence the evolutionary success of cooperators. We model the evolution of cooperation in epithelial cells using the Voronoi tessellation model. Unlike traditional evolutionary graph theory, this allows us to implement global updating, for which birth and death events are spatially deco…
The study of the evolution of cooperative behaviours --which provide benefits to others-- and altruism --which provides benefits to others at a cost to oneself-- has been on the core of the evolutionary game theoretical framework since its foundation. The fast development of the theory during the last years has improved our knowledge of the issue, but carried attached a diversification of concepts which affected communication between scientists. Furthermore, the main root of conflict in the struggle for life identified by Darwin, the limited amount of resources present in any ecosystem, which …
Existing theoretical models of evolution focus on the relative fitness advantages of different mutants in a population while the dynamic behavior of the population size is mostly left unconsidered. We here present a generic stochastic model which combines the growth dynamics of the population and its internal evolution. Our model thereby accounts for the fact that both evolutionary and growth dynamics are based on individual reproduction events and hence are highly coupled and stochastic in nature. We exemplify our approach by studying the dilemma of cooperation in growing populations and show…
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 so…
The concept of fitness is central to evolution, but it quantifies only the expected number of offspring an individual will produce. The actual number of offspring is also subject to noise, arising from environmental or demographic stochasticity. In nature, individuals who are more fecund tend to have greater variance in their offspring number -- sometimes far greater than the Poisson variance assumed in classical models of population genetics. Here, we develop a model for the evolution of two types reproducing in a population of non-constant size. The frequency-dependent fitness of each type i…
Relatedness and synergy affect the selection pressure on cooperation and altruism. Although early work investigated the effect of these factors independently of each other, recent efforts have been aimed at exploring their interplay. Here, we contribute to this ongoing synthesis in two distinct but complementary ways. First, we integrate models of $n$-player matrix games into the direct fitness approach of inclusive fitness theory, hence providing a framework to consider synergistic social interactions between relatives in family and spatially structured populations. Second, we illustrate the …
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 cooperat…
Through the lens of game theory, cooperation is frequently considered an unsustainable strategy: if an entire population is cooperating, each indi- vidual can increase its overall fitness by choosing not to cooperate, thereby still receiving all the benefit of its cooperating neighbors while no longer expending its own energy. Observable cooperation in naturally-occurring public goods games is consequently of great interest, as such systems offer insight into both the emergence and sustainability of cooperation. Here we consider a population that obeys a public goods game on a network of discr…
Cooperation is ubiquitous across all levels of biological systems ranging from microbial communities to human societies. It, however, seemingly contradicts the evolutionary theory, since cooperators are exploited by free-riders and thus are disfavored by natural selection. Many studies based on evolutionary game theory have tried to solve the puzzle and figure out the reason why cooperation exists and how it emerges. Network reciprocity is one of the mechanisms to promote cooperation, where nodes refer to individuals and links refer to social relationships. The spatial arrangement of mutant in…
Cooperation and competition between pathogens can alter the amount of individuals affected by a co-infection. Nonetheless, the evolution of the pathogens' behavior has been overlooked. Here, we consider a co-evolutionary model where the simultaneous spreading is described by a two-pathogen susceptible-infected-recovered model in an either synergistic or competitive manner. At the end of each epidemic season, the pathogens species reproduce according to their fitness that, in turn, depends on the payoff accumulated during the spreading season in a hawk-and-dove game. This co-evolutionary model …
Punishment is a popular tool when governing commons in situations where free riders would otherwise take over. It is well known that sanctioning systems, such as the police and courts, are costly and thus can suffer from those who free ride on other's efforts to maintain the sanctioning systems (second-order free riders). Previous game-theory studies showed that if populations are very large, pool punishment rarely emerges in public good games, even when participation is optional, because of second-order free riders. Here we show that a matching fund for rewarding cooperation leads to the emer…
