Miguel dos Santos

The evolution of punishment through reputation

Punishment of non-cooperators has been observed to promote cooperation. Such punishment is an evolutionary puzzle because it is costly to the punisher while beneficial to others, for example, through increased social cohesion. Recent studies have concluded that punishing strategies usually pay less than some non-punishing strategies. These findings suggest that punishment could not have directly evolved to promote cooperation. However, while it is well established that reputation plays a key role in human cooperation, the simple threat from a reputation of being a punisher may not have been sufficiently explored yet in order to explain the evolution of costly punishment. Here, we first show analytically that punishment can lead to long-term benefits if it influences ones reputation and thereby makes the punisher more likely to receive help in future interactions. Then, in computer simulations, we incorporate up to 40 more complex strategies that use different kinds of reputations (e.g. from generous actions), or strategies that not only include punitive behaviours directed towards defectors but also towards cooperators for example. Our findings demonstrate that punishment can directly evolve through a simple reputation system. We conclude that reputation is crucial for the evolution of punishment by making a punisher more likely to receive help in future interactions, and that experiments investigating the beneficial effects of punishment in humans should include reputation as an explicit feature.

The evolutionary significance of costly punishment is still to be demonstrated.

Costly punishment represents an evolutionary puzzle because it involves an individual paying a cost to harm another individual (1, 2). A recent study by Wu et al. (3) confirmed earlier conclusions (4) that costly punishment is mostly maladaptive and may have evolved for other reasons. We welcome their emphasis on cultural differences in experimental games (5), but we argue that these first tests (3, 4) are not yet conclusive with regard to the functional significance of costly punishment. There are three problems: (i) the definition of “defection” in their Prisoners Dilemma is problematic for studies on punishment, (ii) a closer examination of their payoff matrices reveals them to be nonconducive to punishment, and (iii) their specific game structure imposes an additional opportunity cost that further reduces the effectiveness of punishment.

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 evolution of anti-social rewarding and its countermeasures in public goods games

Cooperation in joint enterprises can easily break down when self-interests are in conflict with collective benefits, causing a tragedy of the commons. In such social dilemmas, the possibility for contributors to invest in a common pool-rewards fund, which will be shared exclusively among contributors, can be powerful for averting the tragedy, as long as the second-order dilemma (i.e. withdrawing contribution to reward funds) can be overcome (e.g. with second-order sanctions). However, the present paper reveals the vulnerability of such pool-rewarding mechanisms to the presence of reward funds raised by defectors and shared among them (i.e. anti-social rewarding), as it causes a cooperation breakdown, even when second-order sanctions are possible. I demonstrate that escaping this social trap requires the additional condition that coalitions of defectors fare poorly compared with pro-socials, with either (i) better rewarding abilities for the latter or (ii) reward funds that are contingent upon the public good produced beforehand, allowing groups of contributors to invest more in reward funds than groups of defectors. These results suggest that the establishment of cooperation through a collective positive incentive mechanism is highly vulnerable to anti-social rewarding and requires additional countermeasures to act in combination with second-order sanctions.

Examining punishment at different explanatory levels

Experimental studies on punishment have sometimes been over-interpreted not only for the reasons Guala lists, but also because of a frequent conflation of proximate and ultimate explanatory levels that Gualas review perpetuates. Moreover, for future analyses we may need a clearer classification of different kinds of punishment.

Antisocial rewarding in structured populations

Cooperation in collective action dilemmas usually breaks down in the absence of additional incentive mechanisms. This tragedy can be escaped if cooperators have the possibility to invest in reward funds that are shared exclusively among cooperators (prosocial rewarding). Yet, the presence of defectors who do not contribute to the public good but do reward themselves (antisocial rewarding) deters cooperation in the absence of additional countermeasures. A recent simulation study suggests that spatial structure is sufficient to prevent antisocial rewarding from deterring cooperation. Here we reinvestigate this issue assuming mixed strategies and weak selection on a game-theoretic model of social interactions, which we also validate using individual-based simulations. We show that increasing reward funds facilitates the maintenance of prosocial rewarding but prevents its invasion, and that spatial structure can sometimes select against the evolution of prosocial rewarding. Our results suggest that, even in spatially structured populations, additional mechanisms are required to prevent antisocial rewarding from deterring cooperation in public goods dilemmas.

Exposure to superfluous information reduces cooperation and increases antisocial punishment in reputation-based interactions

Human cooperation is often based on reputation gained from previous interactions with third parties. Such reputation can be built on generous or punitive actions, and both, one’s own reputation and the reputation of others have been shown to influence decision making in experimental games that control for confounding variables. Here we test how reputation-based cooperation and punishment react to disruption of the cognitive processing in different kinds of helping games with observers. Saying a few superfluous words before each interaction was used to possibly interfere with working memory. In a first set of experiments, where reputation could only be based on generosity, the disruption reduced the frequency of cooperation and lowered mean final payoffs. In a second set of experiments where reputation could only be based on punishment, the disruption increased the frequency of antisocial punishment (i.e. of punishing those who helped) and reduced the frequency of punishing defectors. Our findings suggest that working memory can easily be constraining in reputation-based interactions within experimental games, even if these games are based on a few simple rules with a visual display that provides all the information the subjects need to play the strategies predicted from current theory. Our findings also highlight a weakness of experimental games, namely that they can be very sensitive to environmental variation and that quantitative conclusions about antisocial punishment or other behavioral strategies can easily be misleading.

The coevolution of cooperation and cognition in humans

Cooperative behaviours in archaic hunter–gatherers could have been maintained partly due to the gains from cooperation being shared with kin. However, the question arises as to how cooperation was maintained after early humans transitioned to larger groups of unrelated individuals. We hypothesize that after cooperation had evolved via benefits to kin, the consecutive evolution of cognition increased the returns from cooperating, to the point where benefits to self were sufficient for cooperation to remain stable when group size increased and relatedness decreased. We investigate the theoretical plausibility of this hypothesis, with both analytical modelling and simulations. We examine situations where cognition either (i) increases the benefits of cooperation, (ii) leads to synergistic benefits between cognitively enhanced cooperators, (iii) allows the exploitation of less intelligent partners, and (iv) the combination of these effects. We find that cooperation and cognition can coevolve—cooperation initially evolves, favouring enhanced cognition, which favours enhanced cooperation, and stabilizes cooperation against a drop in relatedness. These results suggest that enhanced cognition could have transformed the nature of cooperative dilemmas faced by early humans, thereby explaining the maintenance of cooperation between unrelated partners.

Pleiotropy, cooperation, and the social evolution of genetic architecture

Pleiotropy has been suggested as a novel mechanism for stabilising cooperation in bacteria and other microbes. The hypothesis is that linking cooperation with a trait that provides a personal (private) benefit can outweigh the cost of cooperation in situations when cooperation would not be favoured by mechanisms such as kin selection. We analysed the theoretical plausibility of this hypothesis, with analytical models and individual-based simulations. We found that (1) pleiotropy does not stabilise cooperation, unless the cooperative and private traits are linked via a genetic architecture that cannot evolve (mutational constraint); (2) if the genetic architecture is constrained in this way, then pleiotropy favours any type of trait and not especially cooperation; (3) if the genetic architecture can evolve, then pleiotropy does not favour cooperation; and (4) there are several alternative explanations for why traits may be linked, and causality can even be predicted in the opposite direction, with cooperation favouring pleiotropy. Our results suggest that pleiotropy could only explain cooperation under restrictive conditions and instead show how social evolution can shape the genetic architecture.

Stochasticity in economic losses increases the value of reputation in indirect reciprocity.

Recent theory predicts harsh and stochastic conditions to generally promote the evolution of cooperation. Here, we test experimentally whether stochasticity in economic losses also affects the value of reputation in indirect reciprocity, a type of cooperation that is very typical for humans. We used a repeated helping game with observers. One subject (the “Unlucky”) lost some money, another one (the “Passer-by”) could reduce this loss by accepting a cost to herself, thereby building up a reputation that could be used by others in later interactions. The losses were either stable or stochastic, but the average loss over time and the average efficiency gains of helping were kept constant in both treatments. We found that players with a reputation of being generous were generally more likely to receive help by others, such that investing into a good reputation generated long-term benefits that compensated for the immediate costs of helping. Helping frequencies were similar in both treatments, but players with a reputation to be selfish lost more resources under stochastic conditions. Hence, returns on investment were steeper when losses varied than when they did not. We conclude that this type of stochasticity increases the value of reputation in indirect reciprocity.