Coexistence of cooperation and defection in public goods games

The production of public goods by the contribution of individual volunteers is a social dilemma because an individual that does not volunteer can benefit from the public good produced by the contributions of others. Therefore it is generally believed that public goods can be produced only in the presence of repeated interactions (which allow reciprocation, reputation effects and punishment) or relatedness (kin selection). Cooperation, however, often occurs in the absence of iterations and relatedness. We show that when the production of a public good is a Volunteer's Dilemma, in which a fixed number of cooperators is necessary to produce the public good, cooperators and defectors persist in a mixed equilibrium, without iterations and without relatedness. This mixed equilibrium is absent in the N-person Prisoner's Dilemma, in which the public good is a linear function of the individual contributions. We also show that the Prisoner's Dilemma and the Volunteer's Dilemma are the two opposite extremes of a general public goods game, and that all intermediate cases can have a mixed equilibrium like the Volunteer's Dilemma. The coexistence of cooperators and defectors, therefore, is a typical outcome of most social dilemmas, which requires neither relatedness nor iterations.     

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.     

Individual heterogeneity and costly punishment: a volunteer's dilemma

Social control and the enforcement of social norms glue a society together. It has been shown theoretically and empirically that informal punishment of wrongdoers fosters cooperation in human groups. Most of this research has focused on voluntary and uncoordinated punishment carried out by individual group members. However, as punishment is costly, it is an open question as to why humans engage in the punishment of wrongdoers even in one-time-only encounters. While evolved punitive preferences have been advocated as proximate explanations for such behaviour, the strategic nature of the punishment situation has remained underexplored. It has been suggested to conceive of the punishment situation as a volunteer''s dilemma (VOD), where only one individual''s action is necessary and sufficient to punish the wrongdoer. Here, we show experimentally that implementing the punishment situation as a VOD sustains cooperation in an environment where punishers and non-punishers coexist. Moreover, we show that punishment-cost heterogeneity allows individuals to tacitly agree on only the strongest group member carrying out the punishment, thereby increasing the effectiveness and efficiency of social norm enforcement. Our results corroborate that costly peer punishment can be explained without assuming punitive preferences and show that centralized sanctioning institutions can emerge from arbitrary individual differences.     

Why mutual helping in most natural systems is neither conflict-free nor based on maximal conflict

Mutual helping for direct benefits can be explained by various game theoretical models, which differ mainly in terms of the underlying conflict of interest between two partners. Conflict is minimal if helping is self-serving and the partner benefits as a by-product. In contrast, conflict is maximal if partners are in a prisoner''s dilemma with both having the pay-off-dominant option of not returning the other''s investment. Here, we provide evolutionary and ecological arguments for why these two extremes are often unstable under natural conditions and propose that interactions with intermediate levels of conflict are frequent evolutionary endpoints. We argue that by-product helping is prone to becoming an asymmetric investment game since even small variation in by-product benefits will lead to the evolution of partner choice, leading to investments by the chosen class. Second, iterated prisoner''s dilemmas tend to take place in stable social groups where the fitness of partners is interdependent, with the effect that a certain level of helping is self-serving. In sum, intermediate levels of mutual helping are expected in nature, while efficient partner monitoring may allow reaching higher levels.     

Human cooperation in social dilemmas: comparing the Snowdrift game with the Prisoner's Dilemma

Explaining the evolution of cooperation among non-relatives is one of the major challenges for evolutionary biology. In this study, we experimentally examined human cooperation in the iterated Snowdrift game (ISD), which has received little attention so far, and compared it with human cooperation in the iterated Prisoner's Dilemma (IPD), which has become the paradigm for the evolution of cooperation. We show that iteration in the ISD leads to consistently higher levels of cooperation than in the IPD. We further demonstrate that the most successful strategies known for the IPD (generous Tit-for-Tat and Pavlov) were also successfully used in the ISD. Interestingly, we found that female players cooperated significantly more often than male players in the IPD but not in the ISD. Moreover, female players in the IPD applied Tit-for-Tat-like or Pavlovian strategies significantly more often than male players, thereby achieving significantly higher pay-offs than male players did. These data demonstrate that the willingness to cooperate does not only depend on the type of the social dilemma, but also on the class of individuals involved. Altogether, our study shows that the ISD can potentially explain high levels of cooperation among non-relatives in humans. In addition, the ISD seems to reflect the social dilemma more realistically than the IPD because individuals obtain immediate direct benefits from the cooperative acts they perform and costs of cooperation are shared between cooperators.     

Prisoner's dilemma posed by fitness-associated recombination strategies

Genetic recombination is a central and repeated topic of study in the evolution of life. However, along with the influence of recombination on evolution, we understand surprisingly little of how selection shapes the nature of recombination. One explanation for recombination is that it allows organisms to escape from perilous situations where they experience very low fitness. As a corollary, it has been suggested that selection should favor recombination at low fitness and not at high fitness (fitness-associated recombination, FAR), and theory suggests that such strategies can indeed be selected. Here we develop models to further investigate the evolution of FAR. Consistent with previous works, we find that FAR can invade and dominate over a strategy of uniform recombination that is independent of fitness. However, our simulation results suggest that extreme FAR strategies, known as group-elitism, are not necessarily superior to other FAR strategies. Moreover, we argue that FAR domination will often occur with a net loss of mean population fitness. Interestingly, this suggests that the strategy of not recombining at high fitness will sometimes be analogous to a defector strategy from the famous "prisoner's dilemma" game: a selfish strategy that is selected but leads to a loss of mean fitness for all players.     

Evolutionary escape from the prisoner's dilemma

The classic prisoner's dilemma model of game theory is modified by introducing occasional variations on the options available to players. Mutation and selection of game options reliably change the game matrix, gradually, from a prisoner's dilemma game into a byproduct mutualism one, in which cooperation is stable, and "temptation to defect" is replaced by temptation to cooperate. This result suggests that when there are many different potential ways of interacting, exploring those possibilities may make escape from prisoner's dilemmas a common outcome in the world. A consequence is that persistent prisoner's dilemma structures may be less common than one might otherwise expect.     

The evolution of altruism: game theory in multilevel selection and inclusive fitness

Although the prisoner's dilemma (PD) has been used extensively to study reciprocal altruism, here we show that the n-player prisoner's dilemma (NPD) is also central to two other prominent theories of the evolution of altruism: inclusive fitness and multilevel selection. An NPD model captures the essential factors for the evolution of altruism directly in its parameters and integrates important aspects of these two theories such as Hamilton's rule, Simpson's paradox, and the Price covariance equation. The model also suggests a simple interpretation of the Price selection decomposition and an alternative decomposition that is symmetrical and complementary to it. In some situations this alternative shows the temporal changes in within- and between-group selection more clearly than the Price equation. In addition, we provide a new perspective on strong vs. weak altruism by identifying their different underlying game structures (based on absolute fitness) and showing how their evolutionary dynamics are nevertheless similar under selection (based on relative fitness). In contrast to conventional wisdom, the model shows that both strong and weak altruism can evolve in periodically formed random groups of non-conditional strategies if groups are multigenerational. An integrative approach based on the NPD helps unify different perspectives on the evolution of altruism.     

Molecular systematics for taxonomic placement of a gorilla of uncertain origin

An adult male gorilla was donated from private ownership in 1994 to Zoo Atlanta and became part of the American Zoo and Aquarium Association's Gorilla Species Survival Plan. This animal, Ivan, was captured as an infant in Africa in 1964. Ambiguity regarding origin and concomitant sub‐species designation was resolved by analyzing the cytochrome oxidase II mitochondrial gene known to contain eight diagnostic sites for gorilla sub‐species. Ivan has the diagnostic profile that characterizes the western clade Gorilla gorilla gorilla. Zoo Biol 18:429–432, 1999. © 1999 Wiley‐Liss, Inc.