Evolution of direct and indirect reciprocity

Indirect reciprocity (IR) occurs when individuals help those who help others. It is important as a potential explanation for why people might develop cooperative reputations. However, previous models of IR are based on the assumption that individuals never meet again. Yet humans and other animals often interact repeatedly within groups, thereby violating the fundamental basis of these models. Whenever re-meeting can occur, discriminating reciprocators can decide whether to help those who helped others (IR) or those who helped them (direct reciprocity, DR). Here I used simulation models to investigate the conditions in which we can expect the different forms of reciprocity to predominate. I show that IR through image scoring becomes unstable with respect to DR by experience scoring as the probability of re-meeting increases. However, using the standing strategy, which takes into account the context of observed defections, IR can be stable with respect to DR even when individuals interact with few partners many times. The findings are important in showing that IR cannot explain a concern for reputation in typical societies unless reputations provide as reliable a guide to cooperative behaviour as does experience.     

The evolution of cooperation and altruism--a general framework and a classification of models

One of the enduring puzzles in biology and the social sciences is the origin and persistence of intraspecific cooperation and altruism in humans and other species. Hundreds of theoretical models have been proposed and there is much confusion about the relationship between these models. To clarify the situation, we developed a synthetic conceptual framework that delineates the conditions necessary for the evolution of altruism and cooperation. We show that at least one of the four following conditions needs to be fulfilled: direct benefits to the focal individual performing a cooperative act; direct or indirect information allowing a better than random guess about whether a given individual will behave cooperatively in repeated reciprocal interactions; preferential interactions between related individuals; and genetic correlation between genes coding for altruism and phenotypic traits that can be identified. When one or more of these conditions are met, altruism or cooperation can evolve if the cost-to-benefit ratio of altruistic and cooperative acts is greater than a threshold value. The cost-to-benefit ratio can be altered by coercion, punishment and policing which therefore act as mechanisms facilitating the evolution of altruism and cooperation. All the models proposed so far are explicitly or implicitly built on these general principles, allowing us to classify them into four general categories.     

Upstream reciprocity and the evolution of gratitude

If someone is nice to you, you feel good and may be inclined to be nice to somebody else. This every day experience is borne out by experimental games: the recipients of an act of kindness are more likely to help in turn, even if the person who benefits from their generosity is somebody else. This behaviour, which has been called ‘upstream reciprocity’, appears to be a misdirected act of gratitude: you help somebody because somebody else has helped you. Does this make any sense from an evolutionary or a game theoretic perspective? In this paper, we show that upstream reciprocity alone does not lead to the evolution of cooperation, but it can evolve and increase the level of cooperation if it is linked to either direct or spatial reciprocity. We calculate the random walks of altruistic acts that are induced by upstream reciprocity. Our analysis shows that gratitude and other positive emotions, which increase the willingness to help others, can evolve in the competitive world of natural selection.     

Indirect reciprocity in asymmetric interactions: when apparent altruism facilitates profitable exploitation

Intraspecific cooperation and interspecific mutualism often feature an asymmetry in the scope for exploitation. We investigate the evolution of indirect reciprocity in an asymmetric game, loosely modelled on interactions between cleaner fishes and clients, in which 'actors' can choose to help or to exploit a 'recipient' that approaches them, while recipients can only choose whether or not to approach an actor (based on the observation of its behaviour towards others). We show that when actors vary in state over time, in a manner that influences the potential gains from exploitation, an equilibrium is possible at which recipients avoid actors whom they have observed exploiting others in the past, and actors help when the potential gains from exploitation are low but choose to exploit when the potential gains are high. In this context, helping is favoured not because it elicits reciprocal altruism ('help so that you may be helped'), but because it facilitates profitable exploitation ('help so that you may gain the opportunity to harm'). The cost of helping one recipient is thereby recouped through exploitation of another. Indirect reciprocity is thus possible even in asymmetric interactions in which one party cannot directly 'punish' exploitation or 'reward' helping by the other.     

Tag-based indirect reciprocity by incomplete social information

Evolution of altruistic behaviour in interacting individuals is accounted for by, for example, kin selection, direct reciprocity, spatially limited interaction and indirect reciprocity. Real social agents, particularly humans, often take actions based on similarity between themselves and others. Although tag-based indirect reciprocity in which altruism occurs exclusively among similar flocks is a natural expectation, its mechanism has not really been established. We propose a model of tag-based indirect reciprocity by assuming that each player may note strategies of others. We show that tag-based altruism can evolve to eradicate other strategies, including unconditional defectors for various initial strategy configurations and parameter sets. A prerequisite for altruism is that the strategy is sometimes, but not always, visible to others. Without visibility of strategies, policing does not take place and defection is optimal. With perfect visibility, what a player does is always witnessed by others and cooperation is optimal. In the intermediate regime, discriminators based on tag proximity, rather than mixture of generous players and defectors, are most likely to evolve. In this situation, altruism is realized based on homophily in which players are exclusively good to similar others.     

Indirect reciprocity can overcome free-rider problems on costly moral assessment

Indirect reciprocity is one of the major mechanisms of the evolution of cooperation. Because constant monitoring and accurate evaluation in moral assessments tend to be costly, indirect reciprocity can be exploited by cost evaders. A recent study crucially showed that a cooperative state achieved by indirect reciprocators is easily destabilized by cost evaders in the case with no supportive mechanism. Here, we present a simple and widely applicable solution that considers pre-assessment of cost evaders. In the pre-assessment, those who fail to pay for costly assessment systems are assigned a nasty image that leads to them being rejected by discriminators. We demonstrate that considering the pre-assessment can crucially stabilize reciprocal cooperation for a broad range of indirect reciprocity models. In particular for the most leading social norms, we analyse the conditions under which a prosocial state becomes locally stable.     

Constraining free riding in public goods games: designated solitary punishers can sustain human cooperation

Much of human cooperation remains an evolutionary riddle. Unlike other animals, people frequently cooperate with non-relatives in large groups. Evolutionary models of large-scale cooperation require not just incentives for cooperation, but also a credible disincentive for free riding. Various theoretical solutions have been proposed and experimentally explored, including reputation monitoring and diffuse punishment. Here, we empirically examine an alternative theoretical proposal: responsibility for punishment can be borne by one specific individual. This experiment shows that allowing a single individual to punish increases cooperation to the same level as allowing each group member to punish and results in greater group profits. These results suggest a potential key function of leadership in human groups and provides further evidence supporting that humans will readily and knowingly behave altruistically.     

Social semantics: how useful has group selection been?

In our social semantics review (J. Evol. Biol., 2007, 415–432), we discussed some of the misconceptions and sources of confusion associated with group selection. Wilson (2007, this issue) claims that we made three errors regarding group selection. Here, we aim to expand upon the relevant points from our review in order to refute this claim. The last 45 years of research provide clear evidence of the relative use of the kin and group selection approaches. Kin selection methodologies are more tractable, allowing the construction of models that can be applied more easily to specific biological examples, including those chosen by Wilson to illustrate the utility of the group selection approach. In contrast, the group selection approach is not only less useful, but also appears to frequently have negative consequences by fostering confusion that leads to wasted effort. More generally, kin selection theory allows the construction of a unified conceptual overview that can be applied across all taxa, whereas there is no formal theory of group selection.     

When is bigger better? The effects of group size on the evolution of helping behaviours

Understanding the evolution of sociality in humans and other species requires understanding how selection on social behaviour varies with group size. However, the effects of group size are frequently obscured in the theoretical literature, which often makes assumptions that are at odds with empirical findings. In particular, mechanisms are suggested as supporting large‐scale cooperation when they would in fact rapidly become ineffective with increasing group size. Here we review the literature on the evolution of helping behaviours (cooperation and altruism), and frame it using a simple synthetic model that allows us to delineate how the three main components of the selection pressure on helping must vary with increasing group size. The first component is the marginal benefit of helping to group members, which determines both direct fitness benefits to the actor and indirect fitness benefits to recipients. While this is often assumed to be independent of group size, marginal benefits are in practice likely to be maximal at intermediate group sizes for many types of collective action problems, and will eventually become very small in large groups due to the law of decreasing marginal returns. The second component is the response of social partners on the past play of an actor, which underlies conditional behaviour under repeated social interactions. We argue that under realistic conditions on the transmission of information in a population, this response on past play decreases rapidly with increasing group size so that reciprocity alone (whether direct, indirect, or generalised) cannot sustain cooperation in very large groups. The final component is the relatedness between actor and recipient, which, according to the rules of inheritance, again decreases rapidly with increasing group size. These results explain why helping behaviours in very large social groups are limited to cases where the number of reproducing individuals is small, as in social insects, or where there are social institutions that can promote (possibly through sanctioning) large‐scale cooperation, as in human societies. Finally, we discuss how individually devised institutions can foster the transition from small‐scale to large‐scale cooperative groups in human evolution.     

The 'spiteful' origins of human cooperation

We analyse generosity, second-party ('spiteful') punishment (2PP), and third-party ('altruistic') punishment (3PP) in a cross-cultural experimental economics project. We show that smaller societies are less generous in the Dictator Game but no less prone to 2PP in the Ultimatum Game. We might assume people everywhere would be more willing to punish someone who hurt them directly (2PP) than someone who hurt an anonymous third person (3PP). While this is true of small societies, people in large societies are actually more likely to engage in 3PP than 2PP. Strong reciprocity, including generous offers and 3PP, exists mostly in large, complex societies that face numerous challenging collective action problems. We argue that 'spiteful' 2PP, motivated by the basic emotion of anger, is more universal than 3PP and sufficient to explain the origins of human cooperation.     

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.