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.     

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 volunteer's dilemma and the optimal size of a social group

If one or few individuals are enough to perform an action that produces a collective good and if this action has a cost, living in group can be beneficial because the cost can be shared with other individuals. Without coordination, however, the production of a collective good by the contribution of one or few individuals is inefficient and can be modelled as a volunteer's dilemma. In the volunteer's dilemma the individuals that pay the cost for the production of the collective good benefit from their action if nobody else volunteers, but the cost is wasted if too many individuals volunteer. Increasing group size reduces the need of volunteering for each member of the group; the overall benefit for the group, however, decreases too because the larger the group is, the less likely it is that the collective good is produced. This problem persists even with a high degree of relatedness between group members; an optimal, intermediate group size exists that maximizes the probability to produce the collective good.     

The option to leave: Conditional dissociation in the evolution of cooperation

Conditional dissociation, i.e. the option to leave an interacting partner in response to his behaviour, is a mechanism that has been shown to promote cooperation in several settings, but the fundamental features that make conditional dissociation work in this way are not yet fully understood. This paper identifies some of the key conditions that make conditional dissociation lead to high levels of cooperation, explains how this mechanism can support the evolutionary coexistence of cooperative and non-cooperative behaviour typically observed in nature, and provides an analytical formula to estimate the expected degree of cooperation thus achieved. Our model involves a population of individuals who are paired to play an iterated prisoner’s dilemma. All individuals share the same capacity to react to the action previously chosen by their partner and, without any other a priori constraint or exclusion, they may use any behavioural rule that is compatible with this capacity. The dynamic evolution of the population eventually enters either a non-cooperative or a partially cooperative regime, depending mainly on the expected lifetime of individuals. Whenever the partially cooperative regime materializes, the cornerstone of its long-run stability is the coexistence of defectors and “Out-for-Tat” strategists, the latter being those who start cooperating and respond to defection by merely leaving. We find, therefore, that conditional dissociation is the essential disciplinary device supporting cooperation, whilst other conditional strategies (such as Tit-for-Tat) remain present only in small population shares. These conclusions are obtained both by extensive numerical simulations and through analytical mean-field methods that approximate the stochastic simulation dynamics and deliver accurate predictions for general parameter configurations.     

The face of another: anonymity and facial symmetry influence cooperation in social dilemmas

In the present paper, we study how the morphological features related to developmental physiology of other participants influence the decision to cooperate in a social dilemma. To that end, we let a large sample of men play a prisoner's dilemma game, both anonymously and against a series of counterparts whose photographs were shown. We focus on three characteristics already linked to cooperative behavior and with described effects on the adult facial shape: facial fluctuating asymmetry, a frequently employed but debated proxy for developmental instability; the degree of facial dimorphism, related to levels of testosterone during puberty; and the second to fourth digit ratio, related to relative prenatal testosterone exposure. We find significantly higher cooperation rates in anonymous interactions than in the non-anonymous round. We also find that individuals are more likely to cooperate with more asymmetric counterparts, and that more asymmetric participants were less likely to believe that their counterpart would cooperate. Variables related to exposure to testosterone during development do not display any effect. We conclude by discussing how out-of-the-game rewards can explain our results.     

Beyond sex,gender, and other dilemmas: Human pelvic morphology from an integrative context

Recent research on the pelvis has clarified the flexibility of pelvic bones to manage nearly infinite possibilities in terms of selection and drift, while still maintaining excellent bipedalism. Despite this work, and the studies outlining the diversity of pelvic morphology across the hominin lineage, conversations continue to be stymied by distractions related to purported trade-offs that the different functions the pelvis must either allow for (e.g., parturition) or directly perform (e.g., attachment sites of muscles). Here we show that tight constraints on morphology are not evident in the pelvic variation of multiple human populations. We thus provide further evidence that human pelves are not geometrically similar and that pelvic morphology successfully balances the intersection of population history, active selective, and drift.     

Wind energy production in forests conflicts with tree-roosting bats

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The dependency game: Multiperson reciprocal sharing leads to stable cooperation which can evolve into group formation

In the standard model for reciprocal collaboration, the repeated prisoner's dilemma (PD), it has proved difficult to establish collaboration in larger groups, necessitating the introduction of additional mechanisms such as reputation or assortedness. The problem is corroborated because current multiperson PDs model simultaneous player action, known as a common goods situation, whereas multiperson collaboration could be easier to obtain in a PD with alternate player action, a private goods situation. Here we present such a game, called a dependency game, and show that stable collaboration can be obtained in a 255 player simulation if only players are allowed to remember three previous benefactors, so they can play advanced tit-for-tat. Furthermore, we show that such a freely collaborating population is threatened by assorted strategies, which define groups that parasitize on independent tit-for-tat players. By excluding others, these groups engage in indirect reciprocal behaviour. Our model therefore combines many hitherto separate collaboration-enhancing concepts into one game, and suggests that group formation and collaboration are two separate social phenomena.