The evolution of strong reciprocity: cooperation in heterogeneous populations

How do human groups maintain a high level of cooperation despite a low level of genetic relatedness among group members? We suggest that many humans have a predisposition to punish those who violate group-beneficial norms, even when this imposes a fitness cost on the punisher. Such altruistic punishment is widely observed to sustain high levels of cooperation in behavioral experiments and in natural settings. We offer a model of cooperation and punishment that we call STRONG RECIPROCITY: where members of a group benefit from mutual adherence to a social norm, strong reciprocators obey the norm and punish its violators, even though as a result they receive lower payoffs than other group members, such as selfish agents who violate the norm and do not punish, and pure cooperators who adhere to the norm but free-ride by never punishing. Our agent-based simulations show that, under assumptions approximating likely human environments over the 100000 years prior to the domestication of animals and plants, the proliferation of strong reciprocators when initially rare is highly likely, and that substantial frequencies of all three behavioral types can be sustained in a population. As a result, high levels of cooperation are sustained. Our results do not require that group members be related or that group extinctions occur.     

Hadza Cooperation

Strong reciprocity is an effective way to promote cooperation. This is especially true when one not only cooperates with cooperators and defects on defectors (second-party punishment) but even punishes those who defect on others (third-party, “altruistic” punishment). Some suggest we humans have a taste for such altruistic punishment and that this was important in the evolution of human cooperation. To assess this we need to look across a wide range of cultures. As part of a cross-cultural project, I played three experimental economics games with the Hadza, who are hunter-gatherers in Tanzania. The Hadza frequently engaged in second-party punishment but they rarely engaged in third-party punishment. Other small-scale societies engaged in less third-party punishment as well. I suggest third-party punishment only became more important in large, complex societies to solve more pressing collective-action problems.     

Rare but severe concerted punishment that favors cooperation

As one of the mechanisms that are supposed to explain the evolution of cooperation among unrelated individuals, costly punishment, in which altruistic individuals privately bear the cost to punish defection, suffers from such drawbacks as decreasing individuals’ welfare, inducing second-order free riding, the difficulty of catching defection, and the possibility of triggering retaliation. To improve this promising mechanism, here we propose an extended Public Goods game with rare but severe concerted punishment, in which once a defector is caught punishment is triggered and the cost of punishment is equally shared among the remainder of the group. Analytical results show that, when the probability for concerted punishment is above a threshold, cooperating is, while defecting is not, an evolutionarily stable strategy in finite populations, and that this way of punishment can considerably decrease the total cost of inhibiting defection, especially in large populations.     

Evolution of cooperation and altruistic punishment when retaliation is possible

Altruistic punishment is suggested to explain observed high levels of cooperation among non-kin related humans. However, laboratory experiments as well as ethnographic evidence suggest that people might retaliate if being punished, and that this reduces the level of cooperation. Building on existing models on the evolution of cooperation and altruistic punishment, we explore the consequences of the option of retaliation. We find that cooperation and altruistic punishment does not evolve with larger population levels if the option of retaliation is included.     

Persistent cooperators in nature

The evolution and maintenance of cooperation fascinated researchers for several decades. Recently, theoretical models and experimental evidence show that costly punishment may facilitate cooperation in human societies. The puzzle how the costly punishment behaviour evolves can be solved under voluntary participation. Could the punishers emerge if participation is compulsory? Is the punishment inevitably a selfish behaviour or an altruistic behaviour? The motivations behind punishment are still an enigma. Based on public goods interactions, we present a model in which just a certain portion of the public good is divided equally among all members. The other portion is distributed to contributors when paying a second cost. The contributors who are willing to pay a second cost are called the persistent cooperators (PC), indicating their desire to retrieve the proportion of the payoff derived from their own contributions with persistent efforts. We show that the persistent cooperators can be costly punishers, which may account for the origin of human costly punishment behaviour under compulsory participation. In this sense our models may show theoretically that the original motivation behind punishment is to retrieve deserved payoff from their own contributions, a selfish incentive. But the persistent cooperators can also flourish or dominate the population in other situations. We list many real examples in which contributors are the persistent cooperators, and they benefit. This indicates a simple norm promoting cooperation: contributing more and gaining more.     

Reward from Punishment Does Not Emerge at All Costs

The conundrum of cooperation has received increasing attention during the last decade. In this quest, the role of altruistic punishment has been identified as a mechanism promoting cooperation. Here we investigate the role of altruistic punishment on the emergence and maintenance of cooperation in structured populations exhibiting connectivity patterns recently identified as key elements of social networks. We do so in the framework of Evolutionary Game Theory, employing the Prisoner''s Dilemma and the Stag-Hunt metaphors to model the conflict between individual and collective interests regarding cooperation. We find that the impact of altruistic punishment strongly depends on the ratio q/p between the cost of punishing a defecting partner (q) and the actual punishment incurred by the partner (p). We show that whenever q/p<1, altruistic punishment turns out to be detrimental for cooperation for a wide range of payoff parameters, when compared to the scenario without punishment. The results imply that while locally, the introduction of peer punishment may seem to reduce the chances of free-riding, realistic population structure may drive the population towards the opposite scenario. Hence, structured populations effectively reduce the expected beneficial contribution of punishment to the emergence of cooperation which, if not carefully dosed, may in fact hinder the chances of widespread cooperation.     

The evolution of punishment in n-player public goods games: a volunteer's dilemma

The evolution of punishment to stabilize cooperation in n-player games has been treated as a second-order social dilemma, where contributions to punishment of free-riders are altruistic. Hence it may only evolve under highly restricted conditions. Here, we build on recent insights using the volunteer's dilemma as an alternative payoff matrix for the evolution of cooperation. The key feature of a volunteer's dilemma is that the benefits of cooperation are a nonlinear function of the number of contributors, meaning that cooperation is negatively frequency dependent. We propose that nonlinear returns are also an inherent feature of punishment and that this insight allows for a simple and novel explanation of how punishment evolves in groups.     

Human altruism: economic, neural, and evolutionary perspectives

Human cooperation represents a spectacular outlier in the animal world. Unlike other creatures, humans frequently cooperate with genetically unrelated strangers, often in large groups, with people they will never meet again, and when reputation gains are small or absent. Experimental evidence and evolutionary models suggest that strong reciprocity, the behavioral propensity for altruistic punishment and altruistic rewarding, is of key importance for human cooperation. Here, we review both evidence documenting altruistic punishment and altruistic cooperation and recent brain imaging studies that combine the powerful tools of behavioral game theory with neuroimaging techniques. These studies show that mutual cooperation and the punishment of defectors activate reward related neural circuits, suggesting that evolution has endowed humans with proximate mechanisms that render altruistic behavior psychologically rewarding.     

Does dishonesty really invite third-party punishment? Results of a more stringent test

Many experiments have demonstrated that people are willing to incur cost to punish norm violators even when they are not directly harmed by the violation. Such altruistic third-party punishment is often considered an evolutionary underpinning of large-scale human cooperation. However, some scholars argue that previously demonstrated altruistic third-party punishment against fairness-norm violations may be an experimental artefact. For example, envy-driven retaliatory behaviour (i.e. spite) towards better-off unfair game players may be misidentified as altruistic punishment. Indeed, a recent experiment demonstrated that participants ceased to inflict third-party punishment against an unfair player once a series of key methodological problems were systematically controlled for. Noticing that a previous finding regarding apparently altruistic third-party punishment against honesty-norm violations may have been subject to methodological issues, we used a different and what we consider to be a more sound design to evaluate these findings. Third-party punishment against dishonest players withstood this more stringent test.     

Altruism, Altruistic Punishment and Social Investment

The concept of altruism is used in very different forms by computer scientists,economists, philosophers, social scientists, psychologists and biologists. Yet, in order to be useful in social simulations, the concept "altruism" requires a more precise meaning. A quantitative formulation is proposed here, based on the cost/benefit analysis of the altruist and of society at large. This formulation is applied in the analysis of the social dynamic working of behaviors that have been called "altruistic punishments", using the agent based computer model Sociodynamica. The simulations suggest that "altruistic punishment" on its own cannot maintain altruistic behaviors. "Altruistic behavior" is sustainable in the long term only if these behaviors trigger synergetic forces in society that eventually make them produce benefits to most individuals. The simulations suggest however that "altruistic punishment" may work as a "social investment", and is thus better called "decentralized social punishment". This behavior is very efficient in enforcing social norms. The efficiency of decentralized social punishment in enforcing norms was dependent on the type of labor structured of the virtual society. I conclude that what is called "altruistic punishment" emerges as a type of social investment that can evolve either through individual and/or group selection, as a successful device for changing or enforcing norms in a society. Social simulations will help us in better understanding the underlying dynamic working of such devices.     

The evolution of punishment and apology: an iterated prisoner's dilemma model

Recently, behaviors that seem to function as punishment or apology have been reported among non-human primates as well as humans. Such behaviors appear to play an important role in maintaining cooperation between individuals. Therefore, the evolution of these behaviors should be examined from the viewpoint of the evolution of cooperation. The iterated prisoner's dilemma (IPD) game is generally considered to be a standard model for the evolution of cooperation. In the present study, strategies accompanied by punishment-like attacks or apology-like behavior were introduced into the common IPD simulation. Punishment and apology were represented by the P signal and the AS signal given immediately after defection. A strategy with the P and AS signals, named the pPAS strategy, was proved to be an evolutionarily stable strategy under certain conditions. Numerical simulations were carried out according to different assigned values of the costs of punishment and apology. The simulations showed that pPAS could dominate the population (1) when the cost of giving P is relatively small, (2) when the cost of receiving P is relatively large, or (3) when the cost of giving AS is relatively large. The relative cost of giving AS had the clearest effect on the success of pPAS. pPAS can dominate the population even when a dominance asymmetry of the costs between two players was introduced. The present results suggest the possible evolution of social behaviors like punishment or apology as a means of maintaining cooperation. This revised version was published online in November 2006 with corrections to the Cover Date.     

Explaining the evolution of parochial punishment in humans

Humans usually favour members of their own group, ethnicity or culture (parochial cooperation), and punish out-group wrongdoers more harshly (parochial punishment). The evolution of parochial cooperation is mainly explained by intergroup conflict, as restricting cooperation to in-groups can provide a relative advantage during conflict. However, explanations for the evolution of parochial punishment are still lacking. It is unclear whether conflict can also explain parochial punishment, because conflict is expected to lead to full hostility towards out-groups, irrespective of their behaviour. Here, we use an agent-based simulation to explore which conditions favour the evolution of parochial third-party punishment. We show that when groups interact and then engage in conflict with each other, third-party punishment is not parochial but spiteful, and directed towards all out-groups. A parochial bias in punishment decisions evolves (i) without conflict, when groups compete against nature and enforcing cooperation requires many punitive actions, and (ii) with conflict, when groups come into conflict with a group other than one they previously interacted with. Our findings suggest that intergroup conflict does not always lead to parochial punishment, and that stable collaborative relations between groups is a key factor promoting third-party parochial punishment. Our findings also provide novel predictions on how punishment and intergroup conflict influence in-group bias in human societies.     

High strength-of-ties and low mobility enable the evolution of third-party punishment

As punishment can be essential to cooperation and norm maintenance but costly to the punisher, many evolutionary game-theoretic studies have explored how direct punishment can evolve in populations. Compared to direct punishment, in which an agent acts to punish another for an interaction in which both parties were involved, the evolution of third-party punishment (3PP) is even more puzzling, because the punishing agent itself was not involved in the original interaction. Despite significant empirical studies of 3PP, little is known about the conditions under which it can evolve. We find that punishment reputation is not, by itself, sufficient for the evolution of 3PP. Drawing on research streams in sociology and psychology, we implement a structured population model and show that high strength-of-ties and low mobility are critical for the evolution of responsible 3PP. Only in such settings of high social-structural constraint are punishers able to induce self-interested agents toward cooperation, making responsible 3PP ultimately beneficial to individuals as well as the collective. Our results illuminate the conditions under which 3PP is evolutionarily adaptive in populations. Responsible 3PP can evolve and induce cooperation in cases where other mechanisms alone fail to do so.     

Power and corruption

Cooperation is ubiquitous in the natural world. What seems nonsensical is why natural selection favors a behavior whereby individuals would lose out by benefiting their competitor. This conundrum, for almost half a century, has puzzled scientists and remains a fundamental problem in biology, psychology, and economics. In recent years, the explanation that punishment can maintain cooperation has received much attention. Individuals who punish noncooperators thrive when punishment does not entail a cost to the punisher. However when punishment is costly, cooperation cannot be preserved. Most literature on punishment fails to consider that punishers may act corruptly by not cooperating when punishing noncooperators. No research has considered that there might be power asymmetries between punishers and nonpunishers that turn one of these type of individuals more or less susceptible to experiencing punishment. Here, we formulate a general game allowing corruption and power asymmetries between punishers and nonpunishers. We show that cooperation can persist if punishers possess power and use it to act corruptly. This result provides a new interpretation of recent data on corrupt policing in social insects and the psychology of power and hypocrisy in humans. These results suggest that corruption may play an important role in maintaining cooperation in insects and human societies. In contrast with previous research, we contend that costly punishment can be beneficial for social groups. This work allows us to identify ways in which corruption can be used to the advantage of a society.     

The Effect of Power Asymmetries on Cooperation and Punishment in a Prisoner’s Dilemma Game

Recent work has suggested that punishment is detrimental because punishment provokes retaliation, not cooperation, resulting in lower overall payoffs. These findings may stem from the unrealistic assumption that all players are equal: in reality individuals are expected to vary in the power with which they can punish defectors. Here, we allowed strong players to interact with weak players in an iterated prisoner''s dilemma game with punishment. Defecting players were most likely to switch to cooperation if the partner cooperated: adding punishment yielded no additional benefit and, under some circumstances, increased the chance that the partner would both defect and retaliate against the punisher. Our findings show that, in a two-player game, cooperation begets cooperation and that punishment does not seem to yield any additional benefits. Further work should explore whether strong punishers might prevail in multi-player games.     

WHEN HAWKS GIVE RISE TO DOVES: THE EVOLUTION AND TRANSITION OF ENFORCEMENT STRATEGIES

The question of how altruism can evolve despite its local disadvantage to selfishness has produced a wealth of theoretical and empirical research capturing the attention of scientists across disciplines for decades. One feature that has remained consistent through this outpouring of knowledge has been that researchers have looked to the altruists themselves for mechanisms by which altruism can curtail selfishness. An alternative perspective may be that just as altruists want to limit selfishness in the population, so may the selfish individuals themselves. These alternative perspectives have been most evident in the fairly recent development of enforcement strategies. Punishment can effectively limit selfishness in the population, but it is not free. Thus, when punishment evolves among altruists, the double costs of exploitation from cheaters and punishment make the evolution of punishment problematic. Here we show that punishment can more readily invade selfish populations when associated with selfishness, whereas altruistic punishers cannot. Thereafter, the establishment of altruism because of enforcement by selfish punishers provides the ideal invasion conditions for altruistic punishment, effectively creating a transition of punishment from selfishness to altruistic. Thus, from chaotic beginnings, a little hypocrisy may go a long way in the evolution and maintenance of altruism.     

The structure of mutations and the evolution of cooperation

Evolutionary game dynamics in finite populations assumes that all mutations are equally likely, i.e., if there are n strategies a single mutation can result in any strategy with probability 1/n. However, in biological systems it seems natural that not all mutations can arise from a given state. Certain mutations may be far away, or even be unreachable given the current composition of an evolving population. These distances between strategies (or genotypes) define a topology of mutations that so far has been neglected in evolutionary game theory. In this paper we re-evaluate classic results in the evolution of cooperation departing from the assumption of uniform mutations. We examine two cases: the evolution of reciprocal strategies in a repeated prisoner's dilemma, and the evolution of altruistic punishment in a public goods game. In both cases, alternative but reasonable mutation kernels shift known results in the direction of less cooperation. We therefore show that assuming uniform mutations has a substantial impact on the fate of an evolving population. Our results call for a reassessment of the "model-less" approach to mutations in evolutionary dynamics.     

Reciprocity,culture and human cooperation: previous insights and a new cross-cultural experiment

Understanding the proximate and ultimate sources of human cooperation is a fundamental issue in all behavioural sciences. In this paper, we review the experimental evidence on how people solve cooperation problems. Existing studies show without doubt that direct and indirect reciprocity are important determinants of successful cooperation. We also discuss the insights from a large literature on the role of peer punishment in sustaining cooperation. The experiments demonstrate that many people are ‘strong reciprocators’ who are willing to cooperate and punish others even if there are no gains from future cooperation or any other reputational gains. We document this in new one-shot experiments, which we conducted in four cities in Russia and Switzerland. Our cross-cultural approach allows us furthermore to investigate how the cultural background influences strong reciprocity. Our results show that culture has a strong influence on positive and in especially strong negative reciprocity. In particular, we find large cross-cultural differences in ‘antisocial punishment’ of pro-social cooperators. Further cross-cultural research and experiments involving different socio-demographic groups document that the antisocial punishment is much more widespread than previously assumed. Understanding antisocial punishment is an important task for future research because antisocial punishment is a strong inhibitor of cooperation.     

The cost of cowardice: punitive sentiments towards free riders in Turkana raids

Models indicate that large-scale cooperation can be sustained by indirect reciprocity or direct punishment, but the relative importance of these mechanisms is unresolved. Unlike direct punishment, indirect sanctions can be meted out without cost to the sanctioner, but direct punishment is advantageous when the scale of cooperation exceeds the network size of individuals. It is of great interest to assess the importance of these mechanisms in small-scale acephalous groups in which people have lived for most of our evolutionary history. Here we evaluate sentiments towards free riders in combat among the Turkana, an acephalous nomadic pastoral society in East Africa who periodically mobilize for cattle-raids against neighboring ethnic groups. Using vignette studies, we probed participants’ motivation to sanction fictitious warriors who were cowards or deserters in a raid and compared it respectively to their reactions to an unskilled warrior or a warrior who turns back due to illness. Our results indicate that the Turkana are motivated to impose both indirect and direct sanctions on cowards consistent with indirect reciprocity and punishment models of cooperation. Our findings imply that both these mechanisms have shaped human cooperative psychology, and sheds light on how prestate societies solve the collective action problem in warfare.     

False friends are worse than bitter enemies: “Altruistic” punishment of in-group members

One of the most critical features of human society is the pervasiveness of cooperation in social and economic exchanges. Moreover, social scientists have found overwhelming evidence that such cooperative behavior is likely to be directed toward in-group members. We propose that the group-based nature of cooperation includes punishment behavior. Punishment behavior is used to maintain cooperation within systems of social exchange and, thus, is directed towards members of an exchange system. Because social exchanges often take place within groups, we predict that punishment behavior is used to maintain cooperation in the punisher's group. Specifically, punishment behavior is directed toward in-group members who are found to be noncooperators. To examine this, we conducted a gift-giving game experiment with third-party punishment. The results of the experiment (N=90) support the following hypothesis: Participants who are cooperative in a gift-giving game punish noncooperative in-group members more severely than they punish noncooperative out-group members.