Prosocial punishment has been proved to be a powerful mean to promote cooperation. Recent studies have found that social exclusion, which indeed can be regarded as a kind of punishment, can also support cooperation. However, if prosocial punishment and exclusion are both present, it is still unclear which strategy is more advantageous to curb free-riders. Here we first study the direct competition between different types of punishment and exclusion. We find that pool (peer) exclusion can always outperform pool (peer) punishment both in the optional and in the compulsory public goods game, no matter whether second-order sanctioning is considered or not. Furthermore, peer exclusion does better than pool exclusion both in the optional and in the compulsory game, but the situation is reversed in the presence of second-order exclusion. Finally, we extend the competition among all possible sanctioning strategies and find that peer exclusion can outperform all other strategies in the absence of second-order exclusion and punishment, while pool exclusion prevails when second-order sanctioning is possible. Our results demonstrate that exclusion is a more powerful strategy than punishment for the resolution of social dilemmas.

Pro-social punishment is a key driver of harmonious and stable society. However, this institution is vulnerable to corruption since law-violators can avoid sanctioning by paying bribes to corrupt law-enforcers. Consequently, to understand how altruistic behavior survives in a corrupt environment is an open question. To reveal potential explanations here we introduce corrupt enforcers and violators into the public goods game with pool punishment, and assume that punishers, as corrupt enforcers, may select defectors probabilistically to take a bribe from, and meanwhile defectors, as corrupt violators, may select punishers stochastically to be corrupted. By means of mathematical analysis, we aim to study the necessary conditions for the evolution of cooperation in such corrupt environment. We find that cooperation can be maintained in the population in two distinct ways. First, cooperators, defectors, and punishers can coexist by all keeping a steady fraction of the population. Second, these three strategies can form a cyclic dominance that resembles a rock-scissors-paper cycle or a heteroclinic cycle. We theoretically identify conditions when the competing strategies coexist in a stationary way or they dominate each other in a cyclic way. These predictions are confirmed numerically.

BackgroundMicrobial communities are susceptible to the public goods dilemma, whereby individuals can gain an advantage within a group by utilizing, but not sharing the cost of producing, public goods. In bacteria, the development of quorum sensing (QS) can establish a cooperation system in a population by coordinating the production of costly and sharable extracellular products (public goods). Cooperators with intact QS system and robust ability in producing public goods are vulnerable to being undermined by QS-deficient defectors that escape from QS but benefit from the cooperation of others. Although microorganisms have evolved several mechanisms to resist cheating invasion in the public goods game, it is not clear why cooperators frequently coexist with defectors and how they form a relatively stable equilibrium during evolution.ResultsWe show that in Pseudomonas aeruginosa, QS-directed social cooperation can select a conditional defection strategy prior to the emergence of QS-mutant defectors, depending on resource availability. Conditional defectors represent a QS-inactive state of wild type (cooperator) individual and can invade QS-activated cooperators by adopting a cheating strategy, and then revert to cooperating when there are abundant nutrient supplies irrespective of the exploitation of QS-mutant defector. Our mathematical modeling further demonstrates that the incorporation of conditional defection strategy into the framework of iterated public goods game with sound punishment mechanism can lead to the coexistence of cooperator, conditional defector, and defector in a rock-paper-scissors dynamics.ConclusionsThese findings highlight the importance of behavioral heterogeneity in stabilizing the population structure and provide a potential reasonable explanation for the maintenance and evolution of cooperation in microbial communities.Electronic supplementary materialThe online version of this article (10.1186/s12915-019-0639-3) contains supplementary material, which is available to authorized users.

Pro-social punishment and exclusion are common means to elevate the level of cooperation among unrelated individuals. Indeed, it is worth pointing out that the combined use of these two strategies is quite common across human societies. However, it is still not known how a combined strategy where punishment and exclusion are switched can promote cooperation from the theoretical perspective. In this paper, we thus propose two different switching strategies, namely peer switching that is based on peer punishment and peer exclusion, and pool switching that is based on pool punishment and pool exclusion. Individuals adopting the switching strategy will punish defectors when their numbers are below a threshold and exclude them otherwise. We study how the two switching strategies influence the evolutionary dynamics in the public goods game. We show that an intermediate value of the threshold leads to a stable coexistence of cooperators, defectors and players adopting the switching strategy in a well-mixed population, and this regardless of whether the pool-based or the peerbased switching strategy is introduced. Moreover, we show that the pure exclusion strategy alone is able to evoke a limit cycle attractor in the evolutionary dynamics, such that cooperation can coexist with other strategies. *

Summary Finding appropriate incentives to enforce collaborative efforts for governing the commons in risky situations is a long-lasting challenge. Previous works have demonstrated that both punishing free-riders and rewarding cooperators could be potential tools to reach this goal. Despite weak theoretical foundations, policy makers frequently impose a punishment-reward combination. Here, we consider the emergence of positive and negative incentives and analyze their simultaneous impact on sustaining risky commons. Importantly, we consider institutions with fixed and flexible incentives. We find that a local sanctioning scheme with pure reward is the optimal incentive strategy. It can drive the entire population toward a highly cooperative state in a broad range of parameters, independently of the type of institutions. We show that our finding is also valid for flexible incentives in the global sanctioning scheme, although the local arrangement works more effectively.