Efficiency and resilience of cooperation in asymmetric social dilemmas.

Direct reciprocity is a powerful mechanism for cooperation in social dilemmas. The very logic of reciprocity, however, seems to require that individuals are symmetric, and that everyone has the same means to influence each others' payoffs. Yet in many applications, individuals are asymmetric. Herein, we study the effect of asymmetry in linear public good games. Individuals may differ in their endowments (their ability to contribute to a public good) and in their productivities (how effective their contributions are). Given the individuals' productivities, we ask which allocation of endowments is optimal for cooperation. To this end, we consider two notions of optimality. The first notion focuses on the resilience of cooperation. The respective endowment distribution ensures that full cooperation is feasible even under the most adverse conditions. The second notion focuses on efficiency. The corresponding endowment distribution maximizes group welfare. Using analytical methods, we fully characterize these two endowment distributions. This analysis reveals that both optimality notions favor some endowment inequality: More productive players ought to get higher endowments. Yet the two notions disagree on how unequal endowments are supposed to be. A focus on resilience results in less inequality. With additional simulations, we show that the optimal endowment allocation needs to account for both the resilience and the efficiency of cooperation.

Mutation enhances cooperation in direct reciprocity.

Direct reciprocity is a powerful mechanism for the evolution of cooperation based on repeated interactions between the same individuals. But high levels of cooperation evolve only if the benefit-to-cost ratio exceeds a certain threshold that depends on memory length. For the best-explored case of one-round memory, that threshold is two. Here, we report that intermediate mutation rates lead to high levels of cooperation, even if the benefit-to-cost ratio is only marginally above one, and even if individuals only use a minimum of past information. This surprising observation is caused by two effects. First, mutation generates diversity which undermines the evolutionary stability of defectors. Second, mutation leads to diverse communities of cooperators that are more resilient than homogeneous ones. This finding is relevant because many real-world opportunities for cooperation have small benefit-to-cost ratios, which are between one and two, and we describe how direct reciprocity can attain cooperation in such settings. Our result can be interpreted as showing that diversity, rather than uniformity, promotes evolution of cooperation.

Social dilemmas among unequals.

Direct reciprocity is a powerful mechanism for the evolution of cooperation on the basis of repeated interactions1-4. It requires that interacting individuals are sufficiently equal, such that everyone faces similar consequences when they cooperate or defect. Yet inequality is ubiquitous among humans5,6 and is generally considered to undermine cooperation and welfare7-10. Most previous models of reciprocity do not include inequality11-15. These models assume that individuals are the same in all relevant aspects. Here we introduce a general framework to study direct reciprocity among unequal individuals. Our model allows for multiple sources of inequality. Subjects can differ in their endowments, their productivities and in how much they benefit from public goods. We find that extreme inequality prevents cooperation. But if subjects differ in productivity, some endowment inequality can be necessary for cooperation to prevail. Our mathematical predictions are supported by a behavioural experiment in which we vary the endowments and productivities of the subjects. We observe that overall welfare is maximized when the two sources of heterogeneity are aligned, such that more productive individuals receive higher endowments. By contrast, when endowments and productivities are misaligned, cooperation quickly breaks down. Our findings have implications for policy-makers concerned with equity, efficiency and the provisioning of public goods.

Evolution of reciprocity with limited payoff memory.

Direct reciprocity is a mechanism for the evolution of cooperation in repeated social interactions. According to the literature, individuals naturally learn to adopt conditionally cooperative strategies if they have multiple encounters with their partner. Corresponding models have greatly facilitated our understanding of cooperation, yet they often make strong assumptions on how individuals remember and process payoff information. For example, when strategies are updated through social learning, it is commonly assumed that individuals compare their average payoffs. This would require them to compute (or remember) their payoffs against everyone else in the population. To understand how more realistic constraints influence direct reciprocity, we consider the evolution of conditional behaviours when individuals learn based on more recent experiences. Even in the most extreme case that they only take into account their very last interaction, we find that cooperation can still evolve. However, such individuals adopt less generous strategies, and they cooperate less often than in the classical setup with average payoffs. Interestingly, once individuals remember the payoffs of two or three recent interactions, cooperation rates quickly approach the classical limit. These findings contribute to a literature that explores which kind of cognitive capabilities are required for reciprocal cooperation. While our results suggest that some rudimentary form of payoff memory is necessary, it suffices to remember a few interactions.

Indirect reciprocity with stochastic and dual reputation updates.

Cooperation is a crucial aspect of social life, yet understanding the nature of cooperation and how it can be promoted is an ongoing challenge. One mechanism for cooperation is indirect reciprocity. According to this mechanism, individuals cooperate to maintain a good reputation. This idea is embodied in a set of social norms called the "leading eight". When all information is publicly available, these norms have two major properties. Populations that employ these norms are fully cooperative, and they are stable against invasion by alternative norms. In this paper, we extend the framework of the leading eight in two directions. First, we include norms with 'dual' reputation updates. These norms do not only assign new reputations to an acting donor; they also allow to update the reputation of the passive recipient. Second, we allow social norms to be stochastic. Such norms allow individuals to evaluate others with certain probabilities. Using this framework, we characterize all evolutionarily stable norms that lead to full cooperation in the public information regime. When only the donor's reputation is updated, and all updates are deterministic, we recover the conventional model. In that case, we find two classes of stable norms: the leading eight and the 'secondary sixteen'. Stochasticity can further help to stabilize cooperation when the benefit of cooperation is comparably small. Moreover, updating the recipients' reputations can help populations to recover more quickly from errors. Overall, our study highlights a remarkable trade-off between the evolutionary stability of a norm and its robustness with respect to errors. Norms that correct errors quickly require higher benefits of cooperation to be stable.

Adaptive dynamics of memory-one strategies in the repeated donation game.

Human interactions can take the form of social dilemmas: collectively, people fare best if all cooperate but each individual is tempted to free ride. Social dilemmas can be resolved when individuals interact repeatedly. Repetition allows them to adopt reciprocal strategies which incentivize cooperation. The most basic model for direct reciprocity is the repeated donation game, a variant of the prisoner's dilemma. Two players interact over many rounds; in each round they decide whether to cooperate or to defect. Strategies take into account the history of the play. Memory-one strategies depend only on the previous round. Even though they are among the most elementary strategies of direct reciprocity, their evolutionary dynamics has been difficult to study analytically. As a result, much previous work has relied on simulations. Here, we derive and analyze their adaptive dynamics. We show that the four-dimensional space of memory-one strategies has an invariant three-dimensional subspace, generated by the memory-one counting strategies. Counting strategies record how many players cooperated in the previous round, without considering who cooperated. We give a partial characterization of adaptive dynamics for memory-one strategies and a full characterization for memory-one counting strategies.

Evolution of cooperation in stochastic games.

Social dilemmas occur when incentives for individuals are misaligned with group interests1-7. According to the 'tragedy of the commons', these misalignments can lead to overexploitation and collapse of public resources. The resulting behaviours can be analysed with the tools of game theory8. The theory of direct reciprocity9-15 suggests that repeated interactions can alleviate such dilemmas, but previous work has assumed that the public resource remains constant over time. Here we introduce the idea that the public resource is instead changeable and depends on the strategic choices of individuals. An intuitive scenario is that cooperation increases the public resource, whereas defection decreases it. Thus, cooperation allows the possibility of playing a more valuable game with higher payoffs, whereas defection leads to a less valuable game. We analyse this idea using the theory of stochastic games16-19 and evolutionary game theory. We find that the dependence of the public resource on previous interactions can greatly enhance the propensity for cooperation. For these results, the interaction between reciprocity and payoff feedback is crucial: neither repeated interactions in a constant environment nor single interactions in a changing environment yield similar cooperation rates. Our framework shows which feedbacks between exploitation and environment-either naturally occurring or designed-help to overcome social dilemmas.

Vacancies in growing habitats promote the evolution of cooperation.

We study evolutionary game dynamics in a growing habitat with vacancies. Fitness is determined by the global effect of the environment and a local prisoner's dilemma among neighbors. We study population growth on a one-dimensional lattice and analyze how the environment affects evolutionary competition. As the environment becomes harsh, an absorbing phase transition from growing populations to extinction occurs. The transition point depends on which strategies are present in the population. In particular, we find a 'cooperative window' in parameter space, where only cooperators can survive. A mutant defector in a cooperative community might briefly proliferate, but over time naturally occurring vacancies separate cooperators from defectors, thereby driving defectors to extinction. Our model reveals that vacancies provide a strong boost for cooperation by spatial selection.

Direct reciprocity between individuals that use different strategy spaces.

In repeated interactions, players can use strategies that respond to the outcome of previous rounds. Much of the existing literature on direct reciprocity assumes that all competing individuals use the same strategy space. Here, we study both learning and evolutionary dynamics of players that differ in the strategy space they explore. We focus on the infinitely repeated donation game and compare three natural strategy spaces: memory-1 strategies, which consider the last moves of both players, reactive strategies, which respond to the last move of the co-player, and unconditional strategies. These three strategy spaces differ in the memory capacity that is needed. We compute the long term average payoff that is achieved in a pairwise learning process. We find that smaller strategy spaces can dominate larger ones. For weak selection, unconditional players dominate both reactive and memory-1 players. For intermediate selection, reactive players dominate memory-1 players. Only for strong selection and low cost-to-benefit ratio, memory-1 players dominate the others. We observe that the supergame between strategy spaces can be a social dilemma: maximum payoff is achieved if both players explore a larger strategy space, but smaller strategy spaces dominate.

Indirect reciprocity with private, noisy, and incomplete information.

Indirect reciprocity is a mechanism for cooperation based on shared moral systems and individual reputations. It assumes that members of a community routinely observe and assess each other and that they use this information to decide who is good or bad, and who deserves cooperation. When information is transmitted publicly, such that all community members agree on each other's reputation, previous research has highlighted eight crucial moral systems. These "leading-eight" strategies can maintain cooperation and resist invasion by defectors. However, in real populations individuals often hold their own private views of others. Once two individuals disagree about their opinion of some third party, they may also see its subsequent actions in a different light. Their opinions may further diverge over time. Herein, we explore indirect reciprocity when information transmission is private and noisy. We find that in the presence of perception errors, most leading-eight strategies cease to be stable. Even if a leading-eight strategy evolves, cooperation rates may drop considerably when errors are common. Our research highlights the role of reliable information and synchronized reputations to maintain stable moral systems.

Social immunity modulates competition between coinfecting pathogens.

Coinfections with multiple pathogens can result in complex within-host dynamics affecting virulence and transmission. While multiple infections are intensively studied in solitary hosts, it is so far unresolved how social host interactions interfere with pathogen competition, and if this depends on coinfection diversity. We studied how the collective disease defences of ants - their social immunity - influence pathogen competition in coinfections of same or different fungal pathogen species. Social immunity reduced virulence for all pathogen combinations, but interfered with spore production only in different-species coinfections. Here, it decreased overall pathogen sporulation success while increasing co-sporulation on individual cadavers and maintaining a higher pathogen diversity at the community level. Mathematical modelling revealed that host sanitary care alone can modulate competitive outcomes between pathogens, giving advantage to fast-germinating, thus less grooming-sensitive ones. Host social interactions can hence modulate infection dynamics in coinfected group members, thereby altering pathogen communities at the host level and population level.

The Red Queen and King in finite populations.

In antagonistic symbioses, such as host-parasite interactions, one population's success is the other's loss. In mutualistic symbioses, such as division of labor, both parties can gain, but they might have different preferences over the possible mutualistic arrangements. The rates of evolution of the two populations in a symbiosis are important determinants of which population will be more successful: Faster evolution is thought to be favored in antagonistic symbioses (the "Red Queen effect"), but disfavored in certain mutualistic symbioses (the "Red King effect"). However, it remains unclear which biological parameters drive these effects. Here, we analyze the effects of the various determinants of evolutionary rate: generation time, mutation rate, population size, and the intensity of natural selection. Our main results hold for the case where mutation is infrequent. Slower evolution causes a long-term advantage in an important class of mutualistic interactions. Surprisingly, less intense selection is the strongest driver of this Red King effect, whereas relative mutation rates and generation times have little effect. In antagonistic interactions, faster evolution by any means is beneficial. Our results provide insight into the demographic evolution of symbionts.

Memory-n strategies of direct reciprocity.

Humans routinely use conditionally cooperative strategies when interacting in repeated social dilemmas. They are more likely to cooperate if others cooperated before, and are ready to retaliate if others defected. To capture the emergence of reciprocity, most previous models consider subjects who can only choose from a restricted set of representative strategies, or who react to the outcome of the very last round only. As players memorize more rounds, the dimension of the strategy space increases exponentially. This increasing computational complexity renders simulations for individuals with higher cognitive abilities infeasible, especially if multiplayer interactions are taken into account. Here, we take an axiomatic approach instead. We propose several properties that a robust cooperative strategy for a repeated multiplayer dilemma should have. These properties naturally lead to a unique class of cooperative strategies, which contains the classical Win-Stay Lose-Shift rule as a special case. A comprehensive numerical analysis for the prisoner's dilemma and for the public goods game suggests that strategies of this class readily evolve across various memory-n spaces. Our results reveal that successful strategies depend not only on how cooperative others were in the past but also on the respective context of cooperation.

Dynamics of prebiotic RNA reproduction illuminated by chemical game theory.

Many origins-of-life scenarios depict a situation in which there are common and potentially scarce resources needed by molecules that compete for survival and reproduction. The dynamics of RNA assembly in a complex mixture of sequences is a frequency-dependent process and mimics such scenarios. By synthesizing Azoarcus ribozyme genotypes that differ in their single-nucleotide interactions with other genotypes, we can create molecules that interact among each other to reproduce. Pairwise interplays between RNAs involve both cooperation and selfishness, quantifiable in a 2 × 2 payoff matrix. We show that a simple model of differential equations based on chemical kinetics accurately predicts the outcomes of these molecular competitions using simple rate inputs into these matrices. In some cases, we find that mixtures of different RNAs reproduce much better than each RNA type alone, reflecting a molecular form of reciprocal cooperation. We also demonstrate that three RNA genotypes can stably coexist in a rock-paper-scissors analog. Our experiments suggest a new type of evolutionary game dynamics, called prelife game dynamics or chemical game dynamics. These operate without template-directed replication, illustrating how small networks of RNAs could have developed and evolved in an RNA world.

Democratic decisions establish stable authorities that overcome the paradox of second-order punishment.

Individuals usually punish free riders but refuse to sanction those who cooperate but do not punish. This missing second-order peer punishment is a fundamental problem for the stabilization of cooperation. To solve this problem, most societies today have implemented central authorities that punish free riders and tax evaders alike, such that second-order punishment is fully established. The emergence of such stable authorities from individual decisions, however, creates a new paradox: it seems absurd to expect individuals who do not engage in second-order punishment to strive for an authority that does. Herein, we provide a mathematical model and experimental results from a public goods game where subjects can choose between a community with and without second-order punishment in two different ways. When subjects can migrate continuously to either community, we identify a bias toward institutions that do not punish tax evaders. When subjects have to vote once for all rounds of the game and have to accept the decision of the majority, they prefer a society with second-order punishment. These findings uncover the existence of a democracy premium. The majority-voting rule allows subjects to commit themselves and to implement institutions that eventually lead to a higher welfare for all.

Cooperation and control in multiplayer social dilemmas.

Direct reciprocity and conditional cooperation are important mechanisms to prevent free riding in social dilemmas. However, in large groups, these mechanisms may become ineffective because they require single individuals to have a substantial influence on their peers. However, the recent discovery of zero-determinant strategies in the iterated prisoner's dilemma suggests that we may have underestimated the degree of control that a single player can exert. Here, we develop a theory for zero-determinant strategies for iterated multiplayer social dilemmas, with any number of involved players. We distinguish several particularly interesting subclasses of strategies: fair strategies ensure that the own payoff matches the average payoff of the group; extortionate strategies allow a player to perform above average; and generous strategies let a player perform below average. We use this theory to describe strategies that sustain cooperation, including generalized variants of Tit-for-Tat and Win-Stay Lose-Shift. Moreover, we explore two models that show how individuals can further enhance their strategic options by coordinating their play with others. Our results highlight the importance of individual control and coordination to succeed in large groups.

Evolution of extortion in Iterated Prisoner's Dilemma games.

Iterated games are a fundamental component of economic and evolutionary game theory. They describe situations where two players interact repeatedly and have the ability to use conditional strategies that depend on the outcome of previous interactions, thus allowing for reciprocation. Recently, a new class of strategies has been proposed, so-called "zero-determinant" strategies. These strategies enforce a fixed linear relationship between one's own payoff and that of the other player. A subset of those strategies allows "extortioners" to ensure that any increase in one player's own payoff exceeds that of the other player by a fixed percentage. Here, we analyze the evolutionary performance of this new class of strategies. We show that in reasonably large populations, they can act as catalysts for the evolution of cooperation, similar to tit-for-tat, but that they are not the stable outcome of natural selection. In very small populations, however, extortioners hold their ground. Extortion strategies do particularly well in coevolutionary arms races between two distinct populations. Significantly, they benefit the population that evolves at the slower rate, an example of the so-called "Red King" effect. This may affect the evolution of interactions between host species and their endosymbionts.

Evolutionary performance of zero-determinant strategies in multiplayer games.

Repetition is one of the key mechanisms to maintain cooperation. In long-term relationships, in which individuals can react to their peers׳ past actions, evolution can promote cooperative strategies that would not be stable in one-shot encounters. The iterated prisoner׳s dilemma illustrates the power of repetition. Many of the key strategies for this game, such as ALLD, ALLC, Tit-for-Tat, or generous Tit-for-Tat, share a common property: players using these strategies enforce a linear relationship between their own payoff and their co-player׳s payoff. Such strategies have been termed zero-determinant (ZD). Recently, it was shown that ZD strategies also exist for multiplayer social dilemmas, and here we explore their evolutionary performance. For small group sizes, ZD strategies play a similar role as for the repeated prisoner׳s dilemma: extortionate ZD strategies are critical for the emergence of cooperation, whereas generous ZD strategies are important to maintain cooperation. In large groups, however, generous strategies tend to become unstable and selfish behaviors gain the upper hand. Our results suggest that repeated interactions alone are not sufficient to maintain large-scale cooperation. Instead, large groups require further mechanisms to sustain cooperation, such as the formation of alliances or institutions, or additional pairwise interactions between group members.

Sanctions as honest signals--the evolution of pool punishment by public sanctioning institutions.

In many species, mutual cooperation is stabilized by forms of policing and peer punishment: if cheaters are punished, there is a strong selective pressure to cooperate. Most human societies have complemented, and sometimes even replaced, such peer punishment mechanisms with pool punishment, where punishment is outsourced to central institutions such as the police. Even before free-riding occurs, such institutions require investments, which could serve as costly signals. Here, we show with a game theoretical model that this signaling effect in turn can be crucial for the evolution of punishment institutions: In the absence of such signals, pool punishment is only stable with second-order punishment and can only evolve when individuals have the freedom not to take part in any interaction. With such signals, individuals can opportunistically adjust their behavior, which promotes the evolution of stable pool punishment even in situations where no one can stand aside. Thus, the human propensity to react opportunistically to credible punishment threats is often sufficient to establish stable punishment institutions and to maintain high levels of cooperation.

Quantitative assessment can stabilize indirect reciprocity under imperfect information.

The field of indirect reciprocity investigates how social norms can foster cooperation when individuals continuously monitor and assess each other's social interactions. By adhering to certain social norms, cooperating individuals can improve their reputation and, in turn, receive benefits from others. Eight social norms, known as the "leading eight," have been shown to effectively promote the evolution of cooperation as long as information is public and reliable. These norms categorize group members as either 'good' or 'bad'. In this study, we examine a scenario where individuals instead assign nuanced reputation scores to each other, and only cooperate with those whose reputation exceeds a certain threshold. We find both analytically and through simulations that such quantitative assessments are error-correcting, thus facilitating cooperation in situations where information is private and unreliable. Moreover, our results identify four specific norms that are robust to such conditions, and may be relevant for helping to sustain cooperation in natural populations.