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. In this work we compare the conditions for cooperative success in an epithelium modelled using evolutionary graph theory, to those in a mechanical model of an epithelium =- the Voronoi tessellation model. Crucially, in this latter model cells are able to move, and birth and death are not spatially coupled. We calculate fixation probabilities in the Voronoi tessellation model through simulation and an approximate analytic technique and show that this leads to stronger promotion of cooperation in comparison with the evolutionary graph theory model.
This article considers a mechanism to explain the emergence and evolution of social cooperation. Selfish individuals tend to benefit themselves, which makes it hard for the maintenance of cooperation between unrelated individuals. We propose and validate that a smart group strategy can effectively facilitate the evolution of cooperation, provided cooperators spatially align whilst cooperating at a new level of alliance. The general evolutionary model presented here shows that a non-kin selection effect is a possible cause for cooperation between unrelated individuals and highlights that non-ki…
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…
We study the effect of contingent movement on the persistence of cooperation on complex networks with empty nodes. Each agent plays Prisoner's Dilemma game with its neighbors and then it either updates the strategy depending on the payoff difference with neighbors or it moves to another empty node if not satisfied with its own payoff. If no neighboring node is empty, each agent stays at the same site. By extensive evolutionary simulations, we show that the medium density of agents enhances cooperation where the network flow of mobile agents is also medium. Moreover, if the movements of agents …
This article aims to clarify the case and the mechanism where sanction or punishment by institution can deliver the evolution of cooperation. Compared to peer sanctioning, institutional sanctioning may be sensitive to players' attitude toward players who do not pre-commit punishment. Departed from former studies based on the punisher who always acts cooperatively, we assume that the punishing player is skeptical in that she cooperates in proportion to how many same types join in her team. Relying on stochastic adaptive dynamics, we show that institutional sanctioning coupled with skeptical pun…
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…
In a community-structured population, public goods games (PGG) occur both within and between communities. Such type of PGG is referred as multilevel public goods games (MPGG). We propose a minimalist evolutionary model of the MPGG and analytically study the evolution of cooperation. We demonstrate that in the case of sufficiently large community size and community number, if the imitation strength within community is weak, i.e., an individual imitates another one in the same community almost randomly, cooperation as well as punishment are more abundant than defection in the long run; if the im…
In biology, the evolution of increasingly cooperative groups has shaped the history of life. Genes collaborate in the control of cells; cells efficiently divide tasks to produce cohesive multicellular individuals; individual members of insect colonies cooperate in integrated societies. Biological cooperation provides a foundation on which to understand human behavior. Conceptually, the economics of efficient allocation and the game-like processes of strategy are well understood in biology; we find the same essential processes in many successful theories of human sociality. Historically, the tr…
Repeated interaction between individuals is the main mechanism for maintaining cooperation in social dilemma situations. Variants of tit-for-tat (repeating the previous action of the opponent) and the win-stay lose-shift strategy are known as strong competitors in iterated social dilemma games. On the other hand, real repeated interaction generally allows plasticity (i.e., learning) of individuals based on the experience of the past. Although plasticity is relevant to various biological phenomena, its role in repeated social dilemma games is relatively unexplored. In particular, if experience-…
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 …
Complex social behaviors lie at the heart of many of the challenges facing evolutionary biology, sociology, economics, and beyond. For evolutionary biologists in particular the question is often how such behaviors can arise \textit{de novo} in a simple evolving system. How can group behaviors such as collective action, or decision making that accounts for memories of past experience, emerge and persist? Evolutionary game theory provides a framework for formalizing these questions and admitting them to rigorous study. Here we develop such a framework to study the evolution of sustained collecti…
Configurational arrangement of network architecture and interaction character of individuals are two most influential factors on the mechanisms underlying the evolutionary outcome of cooperation, which is explained by the well-established framework of evolutionary game theory. In the current study, not only qualitatively but also quantitatively, we measure Moran-evolution of cooperation to support an analytical agreement based on the consequences of the replicator equation in a finite population. The validity of the measurement has been double-checked in the well-mixed network by the Langevin …
Migration is a fundamental trait in humans and animals. Recent studies investigated the effect of migration on the evolution of cooperation, showing that contingent migration favors cooperation in spatial structures. In those studies, only local migration to immediate neighbors was considered, while long-range migration has not been considered yet, partly because the long-range migration has been generally regarded as harmful for cooperation as it would bring the population to a well-mixed state that favors defection. Here, we studied the effects of adaptive long-range migration on the evoluti…
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…
The Weismann barrier, or the impossibility of inheritance of acquired traits, comprises a foundation of modern biology, and it has been a major obstacle in establishing the connection between evolution and ontogenesis. We propose the cooperative model based on the assumption that evolution is achieved by a cooperation between genetic mutations and acquired changes (phenotypic plasticity). It is also assumed in this model that natural selection operates on phenotypes, rather than genotypes, of individuals, and that the relationship between phenotypes and genotypes is one-to-many. In the simulat…
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…
The environment has a strong influence on a population's evolutionary dynamics. Driven by both intrinsic and external factors, the environment is subject to continual change in nature. To capture an ever-changing environment, we consider a model of evolutionary dynamics with game transitions, where individuals' behaviors together with the games they play in one time step influence the games to be played next time step. Within this model, we study the evolution of cooperation in structured populations and find a simple rule: weak selection favors cooperation over defection if the ratio of the b…
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…
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…
In recent years, there has been growing interest in studying evolutionary games with environmental feedback. Previous studies exclusively focus on two-player games. However, extension to multi-player game is needed to study problems such as microbial cooperation and crowdsourcing collaborations. Here, we study coevolutionary public goods games where strategies coevolve with the multiplication factors of group cooperation. Asymmetry can arise in such environmental feedback, where games organized by focal cooperators may have a different efficiency than the ones by defectors. Our analysis shows …
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,…
In this paper, we investigate the effect of "heterogeneity of link weight", heterogeneity of the frequency or amount of interactions among individuals, on the evolution of cooperation. Based on an analysis of the evolutionary prisoner's dilemma game on a weighted one-dimensional lattice network with "intra-individual heterogeneity", we confirm that moderate level of link-weight heterogeneity can facilitate cooperation. Furthermore, we identify two key mechanisms by which link-weight heterogeneity promotes the evolution of cooperation: mechanisms for spread and maintenance of cooperation. We al…
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…
Cooperation is ubiquitous at every level of living organisms. It is known that spatial (network) structure is a viable mechanism for cooperation to evolve. A recently proposed numerical metric, average gradient of selection (AGoS), a useful tool for interpreting and visualizing evolutionary dynamics on networks, allows simulation results to be visualized on a one-dimensional phase space. However, stochastic mutation of strategies was not considered in the analysis of AGoS. Here we extend AGoS so that it can analyze the evolution of cooperation where mutation may alter strategies of individuals…
Game theory research on the snowdrift game has showed that gradual evolution of the continuously varying level of cooperation in joint enterprises can demonstrate evolutionary merging as well as evolutionary branching. However, little is known about the consequences of changes in diversity at the cooperation level. In the present study I consider effects of costly rewards on the continuous snowdrift game. I show that not evolutionary merging but evolutionary branching can promote the emergence of pool reward, which can then enhance social welfare.
Cooperators forgo their interest to benefit others. Thus cooperation should not be favored by natural selection. It challenges the evolutionists, since cooperation is widespread. As one of the resolutions, information spreading has been revealed to play a key role in the emergence of cooperation. Individuals, however, are typically assumed to be passive in the information spreading. Here we assume that individuals are active to spread the information via self-recommendation. Individuals with higher intensities of self-recommendation are likely to have more neighbors. We find that i) eloquent c…
The structure of social networks is a key determinant in fostering cooperation and other altruistic behavior among naturally selfish individuals. However, most real social interactions are temporal, being both finite in duration and spread out over time. This raises the question of whether stable cooperation can form despite an intrinsically fragmented social fabric. Here we develop a framework to study the evolution of cooperation on temporal networks in the setting of the classic Prisoner's Dilemma. By analyzing both real and synthetic datasets, we find that temporal networks generally facil…
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 …
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）
We investigate the effect of randomness in both relationships and decisions on the evolution of cooperation. Simulation results show, in such randomness' presence, the system evolves to a more frequency cooperation state than in its absence. Specifically, there is an optimal amount of randomness, which can induce the highest level of cooperation. The mechanism of randomness promoting cooperation resembles a coherence-resonance-like fashion, which could be of particular interest in evolutionary game dynamics in economic, biological and social systems.
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…
The world in which we are living is a huge network of networks and should be described by interdependent networks. The interdependence between networks significantly affects the evolutionary dynamics of cooperation on them. Meanwhile, due to the diversity and complexity of social and biological systems, players on different networks may not interact with each other by the same way, which should be described by multiple models in evolutionary game theory, such as the Prisoner's Dilemma and Snowdrift Game. We therefore study the evolutionary dynamics of cooperation on two interdependent networks…
The theoretical description of the evolution of cooperation presented by Bergstrom based on assortative matching with partner choice allows to model the population dynamics in a game of Nonrepetitive Prisoners Dilemma. In this paper we present a short analysis of asymmetric effects brought into the game by self knowledge of the participants, that is the knowledge of one's own strategy. Within the same conceptual framework as introduced by Bergstrom inclusion of selfknowledge leads to different behaviour of the assortativity index and changes in payoffs for different strategies.
We study the emergence of cooperation in structured populations with any arrangement of cooperators and defectors on the evolutionary graph. Using structure coefficients defined for configurations describing such arrangements of any number of mutants, we provide results for weak selection to favor cooperation over defection on any regular graph with $N \leq 14$ vertices. Furthermore, the properties of graphs that particularly promote cooperation are analyzed. It is shown that the number of graph cycles of certain length is a good predictor for the values of the structure coefficient, and thus …
Rewards and penalties are common practical tools that can be used to promote cooperation in social institutions. The evolution of cooperation under reward and punishment incentives in joint enterprises has been formalized and investigated, mostly by using compulsory public good games. Recently, Sasaki et al. (2012, Proc Natl Acad Sci USA 109:1165-1169) considered optional participation as well as institutional incentives and described how the interplay between these mechanisms affects the evolution of cooperation in public good games. Here, we present a full classification of these evolutionar…
Spatial structure is one of the simplest and most studied ecological factors that affect the evolution of cooperation. It has been shown that spatial reciprocity promotes cooperation due to the formation of cooperative clusters, which provide mutual support against defectors. The usual assumption is that of constant population size, where no density-related effect is possible. Here, we extend the investigation of density-related effects on the evolution of cooperation. We integrate evolutionary game theory to the Lattice Lotka-Volterra Model. In our model, the birth rate depends on the local d…
Recent studies on the evolutionary dynamics of the Prisoner's Dilemma game in scale-free networks have demonstrated that the heterogeneity of the network interconnections enhances the evolutionary success of cooperation. In this paper we address the issue of how the characterization of the asymptotic states of the evolutionary dynamics depends on the initial concentration of cooperators. We find that the measure and the connectedness properties of the set of nodes where cooperation reaches fixation is largely independent of initial conditions, in contrast with the behavior of both the set of n…
The evolution of cooperation has remained an important problem in evolutionary theory and social sciences. In this regard, a curious question is why consistent cooperative and defective personalities exist and if they serve a role in the evolution of cooperation? To shed light on these questions, here, I consider a population of individuals who possibly play two consecutive rounds of public goods game, with different enhancement factors. Importantly, individuals have independent strategies in the two rounds. However, their strategy in the first round affects the game they play in the second ro…
Cooperators benefit others with paying costs. Evolution of cooperation crucially depends on the cost-benefit ratio of cooperation, denoted as $c$. In this work, we investigate the infinitely repeated prisoner's dilemma for various values of $c$ with four of the representative memory-one strategies, i.e., unconditional cooperation, unconditional defection, tit-for-tat, and win-stay-lose-shift. We consider replicator dynamics which deterministically describes how the fraction of each strategy evolves over time in an infinite-sized well-mixed population in the presence of implementation error and…
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 …
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…
In numerous contexts, individuals may decide whether they take actions to mitigate the spread of disease, or not. Mitigating the spread of disease requires an individual to change their routine behaviours to benefit others, resulting in a 'disease dilemma' similar to the seminal prisoner's dilemma. In the classical prisoner's dilemma, evolutionary game dynamics predict that all individuals evolve to 'defect.' We have discovered that when the rate of cooperation within a population is directly linked to the rate of spread of the disease, cooperation evolves under certain conditions. For disease…
We study the evolution of cooperation in the prisoner's dilemma and the snowdrift game on scale-free networks that are subjected to intentional and random removal of vertices. We show that, irrespective of the game type, cooperation on scale-free networks is extremely robust against random deletion of vertices, but declines fast if vertices with the maximal degree are targeted. In particular, attack tolerance is lowest if the temptation to defect is largest, whereby a small fraction of removed vertices suffices to decimate cooperators. The decline of cooperation can be directly linked to the d…
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…
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…
Evolution of cooperation is a widely studied problem in biology, social science, economics, and artificial intelligence. Most of the existing approaches that explain cooperation rely on some notion of direct or indirect reciprocity. These reciprocity based models assume agents recognize their partner and know their previous interactions, which requires advanced cognitive abilities. In this paper we are interested in developing a model that produces cooperation without requiring any explicit memory of previous game plays. Our model is based on the notion of, a concept introduced within behavior…
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…
Despite recent advances in reputation technologies, it is not clear how reputation systems can affect human cooperation in social networks. Although it is known that two of the major mechanisms in the evolution of cooperation are spatial selection and reputation-based reciprocity, theoretical study of the interplay between both mechanisms remains almost uncharted. Here, we present a new individual-based model for the evolution of reciprocal cooperation between reputation and networks. We comparatively analyze four of the leading moral assessment rules---shunning, image scoring, stern judging, …
