The reputation of punishers

Punishment is a potential mechanism to stabilise cooperation between self-regarding agents. Theoretical and empirical studies on the importance of a punitive reputation have yielded conflicting results. Here, we propose that a variety of factors interact to explain why a punitive reputation is sometimes beneficial and sometimes harmful. We predict that benefits are most likely to occur in forced play scenarios and in situations where punishment is the only means to convey an individual's cooperative intent and willingness to uphold fairness norms. By contrast, if partner choice is possible and an individual's cooperative intent can be inferred directly, then individuals with a nonpunishing cooperative reputation should typically be preferred over punishing cooperators

Power asymmetries and punishment in a prisoner's dilemma with variable cooperative investment

In many two-player games, players that invest in punishment finish with lower payoffs than those who abstain from punishing. These results question the effectiveness of punishment at promoting cooperation, especially when retaliation is possible. It has been suggested that these findings may stem from the unrealistic assumption that all players are equal in terms of power. However, a previous empirical study which incorporated power asymmetries into an iterated prisoner's dilemma (IPD) game failed to show that power asymmetries stabilize cooperation when punishment is possible. Instead, players cooperated in response to their partner cooperating, and punishment did not yield any additional increase in tendency to cooperate. Nevertheless, this previous study only allowed an all-or-nothing–rather than a variable–cooperation investment. It is possible that power asymmetries increase the effectiveness of punishment from strong players only when players are able to vary their investment in cooperation. We tested this hypothesis using a modified IPD game which allowed players to vary their investment in cooperation in response to being punished. As in the previous study, punishment from strong players did not increase cooperation under any circumstances. Thus, in two-player games with symmetric strategy sets, punishment does not appear to increase cooperation

Do cleaning organisms reduce the stress response of client reef fish?

Marine cleaning interactions in which cleaner fish or shrimps remove parasites from visiting 'client' reef fish are a textbook example of mutualism. However, there is yet no conclusive evidence that cleaning organisms significantly improve the health of their clients. We tested the stress response of wild caught individuals of two client species, Chromis dimidiata and Pseudanthias squamipinnis, that had either access to a cleaner wrasse Labroides dimidiatus, or to cleaner shrimps Stenopus hispidus and Periclimenes longicarpus, or no access to cleaning organisms. For both client species, we found an association between the presence of cleaner organisms and a reduction in the short term stress response of client fish to capture, transport and one hour confinement in small aquaria, as measured with cortisol levels. It is conceivable that individuals who are more easily stressed than others pay a fitness cost in the long run. Thus, our data suggest that marine cleaning mutualisms are indeed mutualistic. More generally, measures of stress responses or basal levels may provide a useful tool to assess the impact of interspecific interactions on the partner species

The role of serotonin in the modulation of cooperative behavior

We examined the modulatory role of serotonin in cooperation between unrelated individuals. The identification of the neurohormonal candidates that may modulate levels of cooperation in marine cleaning mutualisms has been a major aim in recent years. Our results provide evidence that serotonin is a driver of cooperative behavioral activities and contribute to the understanding of neural pathways of cooperation, which aim to unravel the basic drive of animal tendencies to cooperate with other

Punishment and cooperation in nature.

Humans use punishment to promote cooperation in laboratory experiments but evidence that punishment plays a similar role in non-human animals is comparatively rare. In this article, we examine why this may be the case by reviewing evidence from both laboratory experiments on humans and ecologically relevant studies on non-human animals. Generally, punishment appears to be most probable if players differ in strength or strategic options. Although these conditions are common in nature, punishment (unlike other forms of aggression) involves immediate payoff reductions to both punisher and target, with net benefits to punishers contingent on cheats behaving more cooperatively in future interactions. In many cases, aggression yielding immediate benefits may suffice to deter cheats and might explain the relative scarcity of punishment in nature

A reinforcement learning model for grooming up the hierarchy in primates

Primates spend a considerable amount of time grooming each other. Grooming is regularly found to be traded reciprocally (for grooming) or for rank-related benefits in the presence of food competition. It has been suggested that if food sources are clustered and monopolizable, then lower-ranked individuals will groom higher-ranked ones in order to be tolerated on food patches. This leads to grooming being directed up the hierarchy. However, the conditions where this is expected to occur are based on verbal reasoning alone, and no quantitative analysis of the conditions favouring grooming up the hierarchy appear in the literature. Here, we develop a quantitative model investigating when food competition can result in grooming up the hierarchy. Individuals are assumed to take actions pertaining to whom to groom, where to feed and whom to tolerate on food patches. By allowing individuals to choose actions according to reinforcement learning, we delineate conditions where groups of individuals will express reciprocal grooming and grooming up the hierarchy depending on environmental conditions (e.g. quality, number of food patches). In particular, we show that conditions of intense food competition may lead to less grooming up the hierarchy. The predictions of our model could guide future comparative studies and meta-analyses investigating social relationships in primates

Coevolution between positive reciprocity, punishment, and partner switching in repeated interactions.

Cooperation based on mutual investments can occur between unrelated individuals when they are engaged in repeated interactions. Individuals then need to use a conditional strategy to deter their interaction partners from defecting. Responding to defection such that the future payoff of a defector is reduced relative to cooperating with it is called a partner control mechanism. Three main partner control mechanisms are (i) to switch from cooperation to defection when being defected ('positive reciprocity'), (ii) to actively reduce the payoff of a defecting partner ('punishment'), or (iii) to stop interacting and switch partner ('partner switching'). However, such mechanisms to stabilize cooperation are often studied in isolation from each other. In order to better understand the conditions under which each partner control mechanism tends to be favoured by selection, we here analyse by way of individual-based simulations the coevolution between positive reciprocity, punishment, and partner switching. We show that random interactions in an unstructured population and a high number of rounds increase the likelihood that selection favours partner switching. In contrast, interactions localized in small groups (without genetic structure) increase the likelihood that selection favours punishment and/or positive reciprocity. This study thus highlights the importance of comparing different control mechanisms for cooperation under different conditions

The benefits of being seen to help others: indirect reciprocity and reputation-based partner choice

When one individual helps another, it benefits the recipient and may also gain a reputation for being cooperative. This may induce others to favour the helper in subsequent interactions, so investing in being seen to help others may be adaptive. The best-known mechanism for this is indirect reciprocity (IR), in which the profit comes from an observer who pays a cost to benefit the original helper. IR has attracted considerable theoretical and empirical interest, but it is not the only way in which cooperative reputations can bring benefits. Signalling theory proposes that paying a cost to benefit others is a strategic investment which benefits the signaller through changing receiver behaviour, in particular by being more likely to choose the signaller as a partner. This reputation-based partner choice can result in competitive helping whereby those who help are favoured as partners. These theories have been confused in the literature. We therefore set out the assumptions, the mechanisms and the predictions of each theory for how developing a cooperative reputation can be adaptive. The benefits of being seen to be cooperative may have been a major driver of sociality, especially in humans. This article is part of the theme issue ‘The language of cooperation: reputation and honest signalling’

The psychological foundations of reputation-based cooperation

Humans care about having a positive reputation, which may prompt them to help in scenarios where the return benefits are not obvious. Various game-theoretical models support the hypothesis that concern for reputation may stabilize cooperation beyond kin, pairs or small groups. However, such models are not explicit about the underlying psychological mechanisms that support reputation-based cooperation. These models therefore cannot account for the apparent rarity of reputation-based cooperation in other species. Here, we identify the cognitive mechanisms that may support reputation-based cooperation in the absence of language. We argue that a large working memory enhances the ability to delay gratification, to understand others' mental states (which allows for perspective-taking and attribution of intentions) and to create and follow norms, which are key building blocks for increasingly complex reputation-based cooperation. We review the existing evidence for the appearance of these processes during human ontogeny as well as their presence in non-human apes and other vertebrates. Based on this review, we predict that most non-human species are cognitively constrained to show only simple forms of reputation-based cooperation. This article is part of the theme issue ‘The language of cooperation: reputation and honest signalling’