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.     

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.     

Do descriptive social norms drive peer punishment? Conditional punishment strategies and their impact on cooperation

Peer punishment is widely considered a key mechanism supporting cooperation in human groups. Although much research shows that human behavior is shaped by the prevailing social norms, little is known about how punishment decisions are impacted by the social context. We present a set of large-scale incentivized experiments in which participants (999 American participants recruited via Amazon Mechanical Turk) could punish their partner conditional on either the level of cooperation or the level of punishment displayed by others who previously interacted in the same setting. While many participants punish independently of levels of cooperation or punishment, a substantial portion punishes free riding more severely when cooperation is more common (‘norm enforcement’), or when free riding is more severely punished by others (‘conformist punishment’). With a dynamic model we demonstrate that conditional punishment strategies can substantially promote cooperation. In particular, conformist punishment helps cooperation to gain a foothold in a population, and norm enforcement helps to maintain cooperation at high levels. Our results provide solid empirical evidence of conditional punishment strategies and illustrate their possible implications for the dynamics of human cooperation.     

Conflict, sticks and carrots: war increases prosocial punishments and rewards

Unlike most species, humans cooperate extensively with group members who are not closely related to them, a pattern sustained in part by punishing non-cooperators and rewarding cooperators. Because internally cooperative groups prevail over less cooperative rival groups, it is thought that violent intergroup conflict played a key role in the evolution of human cooperation. Consequently, it is plausible that propensities to punish and reward will be elevated during intergroup conflict. Using experiments conducted before, during and after the 2006 Israel-Hezbollah war, we show that, during wartime, people are more willing to pay costs to punish non-cooperative group members and reward cooperative group members. Rather than simply increasing within-group solidarity, violent intergroup conflict thus elicits behaviours that, writ large, enhance cooperation within the group, thereby making victory more likely.     

Why people punish defectors. Weak conformist transmission can stabilize costly enforcement of norms in cooperative dilemmas

In this paper, we present a cultural evolutionary model in which norms for cooperation and punishment are acquired via two cognitive mechanisms: (1) payoff-biased transmission-a tendency to copy the most successful individual; and (2) conformist transmission-a tendency to copy the most frequent behavior in the population. We first show that if a finite number of punishment stages is permitted (e.g. two stages of punishment occur if some individuals punish people who fail to punish non-cooperators), then an arbitrarily small amount of conformist transmission will stabilize cooperative behavior by stabilizing punishment at some n -th stage. We then explain how, once cooperation is stabilized in one group, it may spread through a multi-group population via cultural group selection. Finally, once cooperation is prevalent, we show how prosocial genes favoring cooperation and punishment may invade in the wake of cultural group selection.     

Altruistic Punishment and Between-Group Competition

Collective action, or the large-scale cooperation in the pursuit of public goods, has been suggested to have evolved through cultural group selection. Previous research suggests that the costly punishment of group members who do not contribute to public goods plays an important role in the resolution of collective action dilemmas. If large-scale cooperation sustained by the punishment of defectors has evolved through the mechanism of cultural group selection, two implications regarding costly punishment follow: (1) that people are more willing to punish defecting group members in a situation of intergroup competition than in a single-group social dilemma game and (2) that levels of "perverse" punishment of cooperators are not affected by intergroup competition. We find confirmation for these hypotheses. However, we find that the effect of intergroup competition on the punishment of defectors is fully explained by the stronger conditionality of punishment on expected punishment levels in the competition condition.     

Towards a Biopsychological Understanding of Costly Punishment: The Role of Basal Cortisol

Recent findings have documented a negative relation of basal endogenous cortisol and aggression after a provocation (i.e., reactive aggression) in humans. We build on these findings and investigated the relation of endogenous cortisol and reactive aggression in a social dilemma situation, that is, costly punishment of individuals who did not appropriately contribute to a common group project. Specifically, we predicted that basal cortisol is negatively related to costly punishment of uncooperative individuals. In the present study, basal cortisol was assessed prior to a public goods game with the option to punish other group members. In line with previous research on reactive aggression and basal cortisol, we found that basal cortisol was indeed negatively related to costly punishment. The findings are important for understanding costly punishment because this tendency has been documented as a possible basis for the evolution of cooperation.     

Punishing free riders: direct and indirect promotion of cooperation

Human cooperation in a large group of genetically unrelated people is an evolutionary puzzle. Despite its costly nature, cooperative behavior is commonly found in all human societies—a fact that has interested researchers from a wide range of disciplines, including biology, economics, and psychology, to name a few. Many behavioral experiments have demonstrated that cooperation within a group can be sustained when free riders are punished. We argue that punishment has both a direct effect and an indirect effect on promoting cooperation. The direct effect of punishment alters the consequences of cooperation and defection in such a way as to make a rational person prefer cooperation. The indirect effect of punishment promotes cooperation among conditional cooperators by providing the condition necessary for their cooperation (i.e., the expectation that other members will also cooperate). Here we present data from two one-shot n-person prisoner's dilemma games, demonstrating that the indirect effect of punishment complements the direct effect to increase cooperation in the game. Furthermore, we show that direct and indirect effects are robust across two forms of punishment technology: either when punishment is voluntarily provided by game players themselves or when it is exogenously provided by the experimenter.     

Group bias in cooperative norm enforcement

A hallmark of human social cognition is the tendency for both adults and children to favour members of their own groups. Critically, this in-group bias exerts a strong influence on cooperative decision-making: people (i) tend to share more with members of their in-group and (ii) differentially enforce fairness norms depending on the group membership of their interaction partners. But why do people show these group biases in cooperation? One possibility is that the enforcement of cooperative norm violations is an evolved mechanism supporting within-group cooperation (Norms-Focused Hypothesis). Alternatively, group bias in cooperation could be a by-product of more general affective preferences for in-group members (Mere Preferences Hypothesis). Here, we appraise evidence from studies of both adults and children with the goal of understanding whether one of these two accounts is better supported by existing data. While the pattern of evidence is complex, much of it is broadly consistent with the Mere Preferences Hypothesis and little is uniquely supportive of the Norms-Focused Hypothesis. We highlight possible reasons for this complexity and suggest ways that future work can continue to help us understand the important relationship between group bias and cooperation.     

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.     

Female cleaner fish cooperate more with unfamiliar males

Joint group membership is of major importance for cooperation in humans, and close ties or familiarity with a partner are also thought to promote cooperation in other animals. Here, we present the opposite pattern: female cleaner fish, Labroides dimidiatus, behave more cooperatively (by feeding more against their preference) when paired with an unfamiliar male rather than with their social partner. We propose that cooperation based on asymmetric punishment causes this reversed pattern. Males are larger than and dominant to female partners and are more aggressive to unfamiliar than to familiar female partners. In response, females behave more cooperatively with unfamiliar male partners. Our data suggest that in asymmetric interactions, weaker players might behave more cooperatively with out-group members than with in-group members to avoid harsher punishment.     

More 'altruistic' punishment in larger societies

If individuals will cooperate with cooperators, and punish non-cooperators even at a cost to themselves, then this strong reciprocity could minimize the cheating that undermines cooperation. Based upon numerous economic experiments, some have proposed that human cooperation is explained by strong reciprocity and norm enforcement. Second-party punishment is when you punish someone who defected on you; third-party punishment is when you punish someone who defected on someone else. Third-party punishment is an effective way to enforce the norms of strong reciprocity and promote cooperation. Here we present new results that expand on a previous report from a large cross-cultural project. This project has already shown that there is considerable cross-cultural variation in punishment and cooperation. Here we test the hypothesis that population size (and complexity) predicts the level of third-party punishment. Our results show that people in larger, more complex societies engage in significantly more third-party punishment than people in small-scale societies.     

Cooperation‐mediated plasticity in dispersal and colonization

Kin selection theory predicts that costly cooperative behaviors evolve most readily when directed toward kin. Dispersal plays a controversial role in the evolution of cooperation: dispersal decreases local population relatedness and thus opposes the evolution of cooperation, but limited dispersal increases kin competition and can negate the benefits of cooperation. Theoretical work has suggested that plasticity of dispersal, where individuals can adjust their dispersal decisions according to the social context, might help resolve this paradox and promote the evolution of cooperation. Here, we experimentally tested the hypothesis that conditional dispersal decisions are mediated by a cooperative strategy: we quantified the density‐dependent dispersal decisions and subsequent colonization efficiency from single cells or groups of cells among six genetic strains of the unicellular Tetrahymena thermophila that differ in their aggregation level (high, medium, and low), a behavior associated with cooperation strategy. We found that the plastic reaction norms of dispersal rate relative to density differed according to aggregation level: highly aggregative genotypes showed negative density‐dependent dispersal, whereas low‐aggregation genotypes showed maximum dispersal rates at intermediate density, and medium‐aggregation genotypes showed density‐independent dispersal with intermediate dispersal rate. Dispersers from highly aggregative genotypes had specialized long‐distance dispersal phenotypes, contrary to low‐aggregation genotypes; medium‐aggregation genotypes showing intermediate dispersal phenotype. Moreover, highly aggregation genotypes showed evidence for beneficial kin‐cooperation during dispersal. Our experimental results should help to resolve the evolutionary conflict between cooperation and dispersal: cooperative individuals are expected to avoid kin‐competition by dispersing long distances, but maintain the benefits of cooperation by dispersing in small groups.     

Intergroup bias in third-party punishment stems from both ingroup favoritism and outgroup discrimination

Social norms pervade almost every aspect of social interaction. If they are violated, not only legal institutions, but other members of society as well, punish, i.e., inflict costs on the wrongdoer. Sanctioning occurs even when the punishers themselves were not harmed directly and even when it is costly for them. There is evidence for intergroup bias in this third-party punishment: third-parties, who share group membership with victims, punish outgroup perpetrators more harshly than ingroup perpetrators. However, it is unknown whether a discriminatory treatment of outgroup perpetrators (outgroup discrimination) or a preferential treatment of ingroup perpetrators (ingroup favoritism) drives this bias. To answer this question, the punishment of outgroup and ingroup perpetrators must be compared to a baseline, i.e., unaffiliated perpetrators. By applying a costly punishment game, we found stronger punishment of outgroup versus unaffiliated perpetrators and weaker punishment of ingroup versus unaffiliated perpetrators. This demonstrates that both ingroup favoritism and outgroup discrimination drive intergroup bias in third-party punishment of perpetrators that belong to distinct social groups.     

HUMAN COOPERATION BASED ON PUNISHMENT REPUTATION

The threat of punishment usually promotes cooperation. However, punishing itself is costly, rare in nonhuman animals, and humans who punish often finish with low payoffs in economic experiments. The evolution of punishment has therefore been unclear. Recent theoretical developments suggest that punishment has evolved in the context of reputation games. We tested this idea in a simple helping game with observers and with punishment and punishment reputation (experimentally controlling for other possible reputational effects). We show that punishers fully compensate their costs as they receive help more often. The more likely defection is punished within a group, the higher the level of within‐group cooperation. These beneficial effects perish if the punishment reputation is removed. We conclude that reputation is key to the evolution of punishment.     

The psychology of deterrence explains why group membership matters for third-party punishment

Humans regularly intervene in others' conflicts as third-parties. This has been studied using the third-party punishment game: A third-party can pay a cost to punish another player (the “dictator”) who treated someone else poorly. Because the game is anonymous and one-shot, punishers are thought to have no strategic reasons to intervene. Nonetheless, punishers often punish dictators who treat others poorly. This result is central to a controversy over human social evolution: Did third-party punishment evolve to maintain group norms or to deter others from acting against one's interests? This paper provides a critical test. We manipulate the ingroup/outgroup composition of the players while simultaneously measuring the inferences punishers make about how the dictator would treat them personally. The group norm predictions were falsified, as outgroup defectors were punished most harshly, not ingroup defectors (as predicted by ingroup fairness norms) and not outgroup members generally (as predicted by norms of parochialism). The deterrence predictions were validated: Punishers punished the most when they inferred that they would be treated the worst by dictators, especially when better treatment would be expected given ingroup/outgroup composition.     

Runaway selection for cooperation and strict-and-severe punishment

Punishing defectors is an important means of stabilizing cooperation. When levels of cooperation and punishment are continuous, individuals must employ suitable social standards for defining defectors and for determining punishment levels. Here we investigate the evolution of a social reaction norm, or psychological response function, for determining the punishment level meted out by individuals in dependence on the cooperation level exhibited by their neighbors in a lattice-structured population. We find that (1) cooperation and punishment can undergo runaway selection, with evolution towards enhanced cooperation and an ever more demanding punishment reaction norm mutually reinforcing each other; (2) this mechanism works best when punishment is strict, so that ambiguities in defining defectors are small; (3) when the strictness of punishment can adapt jointly with the threshold and severity of punishment, evolution favors the strict-and-severe punishment of individuals who offer slightly less than average cooperation levels; (4) strict-and-severe punishment naturally evolves and leads to much enhanced cooperation when cooperation without punishment would be weak and neither cooperation nor punishment are too costly; and (5) such evolutionary dynamics enable the bootstrapping of cooperation and punishment, through which defectors who never punish gradually and steadily evolve into cooperators who punish those they define as defectors.     

Influences of Relatedness,Food Deprivation,and Sex on Adult Behaviors in the Group‐living Insect Forficula auricularia

The evolution of group living is generally associated with the emergence of social behaviors that ensure fitness benefits to group members. However, the expression of these behaviors may depend on group composition, which can vary over time with respect to sex, starvation status, and relatedness. Here, we investigated (1) whether adults of the group‐living European earwig, Forficula auricularia, show cooperative behaviors toward conspecifics and (2) whether sex, food availability, and relatedness shape the nature and frequency of these behaviors. We conducted a full‐factorial experiment using 108 unisexual pairs of adults, in which we manipulated these three factors and video‐recorded the earwig behaviors for 45 min. Our results revealed that adults mostly expressed self‐directed and aggressive behaviors. Nevertheless, they also showed allogrooming, a social behavior that offers scope for cooperation. Pairs of males displayed longer bouts of aggression and allogrooming (when it occurred) than pairs of females. Food deprivation had no effect on male behaviors, but females spent less time self‐grooming and walking when they were food deprived. Finally, low relatedness between adults did not influence any of the measured behaviors, but exacerbated frass production, possibly due to social stress. Overall, these results indicate the limited role of cooperation among F. auricularia adults during their group‐living phase.     

Strong reciprocity, human cooperation, and the enforcement of social norms

This paper provides strong evidence challenging the self-interest assumption that dominates the behavioral sciences and much evolutionary thinking. The evidence indicates that many people have a tendency to voluntarily cooperate, if treated fairly, and to punish noncooperators. We call this behavioral propensity “strong reciprocity” and show empirically that it can lead to almost universal cooperation in circumstances in which purely self-interested behavior would cause a complete breakdown of cooperation. In addition, we show that people are willing to punish those who behaved unfairly towards a third person or who defected in a Prisoner’s Dilemma game with a third person. This suggests that strong reciprocity is a powerful device for the enforcement of social norms involving, for example, food sharing or collective action. Strong reciprocity cannot be rationalized as an adaptive trait by the leading evolutionary theories of human cooperation (in other words, kin selection, reciprocal altruism, indirect reciprocity, and costly signaling theory). However, multilevel selection theories of cultural evolution are consistent with strong reciprocity.     

Wrath of God: religious primes and punishment

Recent evidence indicates that priming participants with religious concepts promotes prosocial sharing behaviour. In the present study, we investigated whether religious priming also promotes the costly punishment of unfair behaviour. A total of 304 participants played a punishment game. Before the punishment stage began, participants were subliminally primed with religion primes, secular punishment primes or control primes. We found that religious primes strongly increased the costly punishment of unfair behaviours for a subset of our participants--those who had previously donated to a religious organization. We discuss two proximate mechanisms potentially underpinning this effect. The first is a 'supernatural watcher' mechanism, whereby religious participants punish unfair behaviours when primed because they sense that not doing so will enrage or disappoint an observing supernatural agent. The second is a 'behavioural priming' mechanism, whereby religious primes activate cultural norms pertaining to fairness and its enforcement and occasion behaviour consistent with those norms. We conclude that our results are consistent with dual inheritance proposals about religion and cooperation, whereby religions harness the byproducts of genetically inherited cognitive mechanisms in ways that enhance the survival prospects of their adherents.