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PubMed2002

Spatial prisoner's dilemma games with dynamic payoff matrices.

The effects of dynamic payoff matrices on evolution of cooperation are studied based on the prisoner's dilemma game on a two-dimensional square lattice. The study is conducted by simulation and an analytical theory based on mean-field approximation. Payoff matrices are designed to evolve depending on a ratio of defectors (or cooperators) to the whole population. Dynamic payoff matrices are necessary to describe evolution of a society whose payoff may be affected by the results of actions of the members in the society. Introducing such payoff matrices helps to model dynamic aspects of societies.

Journal Article↗
PubMed1997

Evolutionary transitions and artificial life.

A major challenge for artificial life is to synthesize the evolutionary transitions that have repeatedly formed differentiated higher-level entities from cooperative organizations of lower-level entities, producing the nested hierarchical structure of living processes. This article identifies the key elements and relationships that must be incorporated or synthesized in an artificial life system if these transitions are to emerge. The processes currently included in artificial life systems are unable to provide an adequate basis for the emergence of the complex cooperative organization that is essential to the transitions. A new theory of the evolution of cooperative organization is developed that points to the additional processes that must be included in artificial life systems to underpin the emergence of the transitions.

Biological Evolution↗
PubMed2006

Cheating can stabilize cooperation in mutualisms.

Mutualisms present a challenge for evolutionary theory. How is cooperation maintained in the face of selection for selfishness and cheating? Both theory and data suggest that partner choice, where one species preferentially directs aid to the more cooperative members of the other species, is central to cooperation in many mutualisms. However, the theory has only so far considered the evolutionary effects of partner choice on one of the species in a mutualism in isolation. Here, we investigate the co-evolution of cooperation and choice in a choosy host and its symbiont. Our model reveals that even though choice and cooperation may be initially selected, it will often be unstable. This is because choice reduces variation in the symbiont and, therefore, tends to remove the selective incentive for its own maintenance (a scenario paralleled in the lek paradox in female choice and policing in within-species cooperation). However, we also show that when variability is reintroduced into symbionts each generation, in the form of less cooperative individuals, choice is maintained. This suggests that the presence of cheaters and cheater species in many mutualisms is central to the maintenance of partner choice and, paradoxically, cooperation itself.

Animals↗
PubMed2006

The evolution of function-valued traits for conditional cooperation.

In this paper we study the evolution of function-valued traits for cooperation in environments that display varying degrees of population viscosity. Traits measure an individual's intrinsic propensity to cooperate in a standard bilateral Prisoner's dilemma and can be increasing, decreasing or constant functions of the probability to interact with individuals of ones own genotype. We first analyse adaptation to homogenous environments (with constant degree of viscosity). Comparing environments characterized by different degrees of viscosity, we find that the relation between viscosity and the equilibrium type distribution is not monotone. In fact, it is possible that in fluid populations (no viscosity) there is more cooperation in equilibrium than in populations with intermediate degrees of viscosity. In a second step we analyse heterogenous environments (with varying degrees of viscosity). We find that under very weak assumptions on the distribution of the viscosity parameter strictly increasing functions are always selected and under some parameter constellations they are uniquely so.

Animals↗
PubMed1993

A strategy of win-stay, lose-shift that outperforms tit-for-tat in the Prisoner's Dilemma game.

The Prisoner's Dilemma is the leading metaphor for the evolution of cooperative behaviour in populations of selfish agents, especially since the well-known computer tournaments of Axelrod and their application to biological communities. In Axelrod's simulations, the simple strategy tit-for-tat did outstandingly well and subsequently became the major paradigm for reciprocal altruism. Here we present extended evolutionary simulations of heterogeneous ensembles of probabilistic strategies including mutation and selection, and report the unexpected success of another protagonist: Pavlov. This strategy is as simple as tit-for-tat and embodies the fundamental behavioural mechanism win-stay, lose-shift, which seems to be a widespread rule. Pavlov's success is based on two important advantages over tit-for-tat: it can correct occasional mistakes and exploit unconditional cooperators. This second feature prevents Pavlov populations from being undermined by unconditional cooperators, which in turn invite defectors. Pavlov seems to be more robust than tit-for-tat, suggesting that cooperative behaviour in natural situations may often be based on win-stay, lose-shift.

Biological Evolution↗
PubMed2006

Evolution of contingent altruism when cooperation is expensive.

The ubiquity of cooperation has motivated a major research program over the last 50 years to discover ever more minimal conditions for the evolution of altruism. One important line of work is based on favoritism toward those who appear to be close relatives. Another important line is based on continuing interactions, whether between individuals (e.g., reciprocity) or between lines of descent in a viscous population. Here, we use an agent-based model to demonstrate a new mechanism that combines both lines of work to show when and how favoritism toward apparently similar others can evolve in the first place. The mechanism is the joint operation of viscosity and of tags (heritable, observable, and initially arbitrary characteristics), which serve as weak and potentially deceptive indicators of relatedness. Although tags are insufficient to support cooperation alone, we show that this joint mechanism vastly increases the range of environments in which contingent altruism can evolve in viscous populations. Even though our model is quite simple, the subtle dynamics underlying our results are not tractable using formal analytic tools (such as analysis of evolutionarily stable strategies), but are amenable to agent-based simulation.

Altruism↗
PubMed2001

Cooperation and conflict in the evolution of multicellularity.

Multicellular organisms probably originated as groups of cells formed in several ways, including cell proliferation from a group of founder cells and aggregation. Cooperation among cells benefits the group, but may be costly (altruistic) or beneficial (synergistic) to individual cooperating cells. In this paper, we study conflict mediation, the process by which genetic modifiers evolve that enhance cooperation by altering the parameters of development or rules of formation of cell groups. We are particularly interested in the conditions under which these modifiers lead to a new higher-level unit of selection with increased cooperation among group members and heritable variation in fitness at the group level. By sculpting the fitness variation and opportunity for selection at the two levels, conflict modifiers create new functions at the organism level. An organism is more than a group of cooperating cells related by common descent; organisms require adaptations that regulate conflict within. Otherwise their continued evolution is frustrated by the creation of within-organism variation and conflict between levels of selection. The evolution of conflict modifiers is a necessary prerequisite to the emergence of individuality and the continued well being of the organism. Conflict leads--through the evolution of adaptations that reduce i--to greater individuality and harmony for the organism.

Animals↗
PubMed1992

The evolution of delayed dispersal in cooperative breeders.

Why do the young of cooperative breeders--species in which more than two individuals help raise offspring at a single nest--delay dispersal and live in groups? Answering this deceptively simple question involves examining the costs and benefits of three alternative strategies: (1) dispersal and attempting to breed, (2) dispersal and floating, and (3) delayed dispersal and helping. If, all other things being equal, the fitness of individuals that delay dispersal is greater than the fitness of individuals that disperse and breed on their own, intrinsic benefits are paramount to the current maintenance of delayed dispersal. Intrinsic benefits are directly due to living with others and may include enhanced foraging efficiency and reduced susceptibility to predation. However, if individuals that disperse and attempt to breed in high-quality habitat achieve the highest fitness, extrinsic constraints on the ability of offspring to obtain such high-quality breeding opportunities force offspring to either delay dispersal or float. The relevant constraint to independent reproduction has frequently been termed habitat saturation. This concept, of itself, fails to explain the evolution of delayed dispersal. Instead, we propose the delayed-dispersal threshold model as a guide for organizing and evaluating the ecological factors potentially responsible for this phenomenon. We identify five parameters critical to the probability of delayed dispersal: relative population density, the fitness differential between early dispersal/breeding and delayed dispersal, the observed or hypothetical fitness of floaters, the distribution of territory quality, and spatiotemporal environmental variability. A key conclusion from the model is that no one factor by itself causes delayed dispersal and cooperative breeding. However, a difference in the dispersal patterns between two closely related species or populations (or between individuals in the same population in different years) may be attributable to one or a small set of factors. Much remains to be done to pinpoint the relative importance of different ecological factors in promoting delayed dispersal. This is underscored by our current inability to explain satisfactorily several patterns including the relative significance of floating, geographic biases in the incidence of cooperative breeding, sexual asymmetries in delayed dispersal, the relationship between delayed dispersal leading to helping behavior and cooperative polygamy, and the rarity of the co-occurrence of helpers and floaters within the same population. Advances in this field remain to be made along several fronts.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed2006

Global analyses of evolutionary dynamics and exhaustive search for social norms that maintain cooperation by reputation.

Reputation formation is a key to understanding indirect reciprocity. In particular, the way to assign reputation to each individual, namely a norm that describes who is good and who is bad, greatly affects the possibility of sustained cooperation in the population. Previously, we have exhaustively studied reputation dynamics that are able to maintain a high level of cooperation at the ESS. However, this analysis examined the stability of monomorphic population and did not investigate polymorphic population where several strategies coexist. Here, we study the evolutionary dynamics of multiple behavioral strategies by replicator dynamics. We exhaustively study all 16 possible norms under which the reputation of a player in the next round is determined by the action of the self and the reputation of the opponent. For each norm, we explore evolutionary dynamics of three strategies: unconditional cooperators, unconditional defectors, and conditional cooperators. We find that only three norms, simple-standing, Kandori, and shunning, can make conditional cooperation evolutionarily stable, hence, realize sustained cooperation. The other 13 norms, including scoring, ultimately lead to the invasion by defectors. Also, we study the model in which private reputation errors exist to a small extent. In this case, we find the stable coexistence of unconditional and conditional cooperators under the three norms.

Biological Evolution↗
PubMed2005

Human behaviour: Egalitarian motive and altruistic punishment.

Altruistic punishment is a behaviour in which individuals punish others at a cost to themselves in order to provide a public good. Fehr and Gächter present experimental evidence in humans indicating that negative emotions towards non-cooperators motivate punishment, which, in turn, provokes a high degree of cooperation. Using Fehr and Gächter's original data, we provide an alternative analysis of their experiment that suggests that egalitarian motives are more important than motives for punishing non-cooperative behaviour. This finding is consistent with evidence that humans may have an evolutionary incentive to punish the highest earners in order to promote equality, rather than cooperation.

Altruism↗
PubMed2003

Punishment and reputation in spatial public goods games.

The puzzle of the emergence of cooperation between unrelated individuals is shared across diverse fields of behavioural sciences and economics. In this article we combine the public goods game originating in economics with evolutionary approaches traditionally used in biology. Instead of pairwise encounters, we consider the more complex case of groups of three interacting individuals. We show that territoriality is capable of promoting cooperative behaviour, as in the case of the Prisoner's Dilemma. Moreover, by adding punishment opportunities, the readiness to cooperate is greatly enhanced and asocial strategies can be largely suppressed. Finally, as soon as players carry a reputation for being willing or unwilling to punish, highly cooperative and fair outcomes are achieved. This group-beneficial result is obtained, intriguingly, by making individuals more likely to exploit their co-players if they can get away with it. Thus, less-cooperative individuals make more-cooperative societies.

Animals↗
PubMed2004

The selfish nature of generosity: harassment and food sharing in primates.

Animals may share food to gain immediate or delayed fitness benefits. Previous studies of sharing have concentrated on delayed benefits such as reciprocity, trade and punishment. This study tests an alternative model (the harassment or sharing-under-pressure hypothesis) in which a food owner immediately benefits because sharing avoids costly harassment from a beggar. I present an experiment that varies the potential ability of the beggar to harass, and of the owner to defend the food, to examine the effects of harassment on food sharing in two primate species: chimpanzees (Pan troglodytes) and squirrel monkeys (Saimiri boliviensis). For both species, high levels of harassment potential significantly increased both beggar harassment and sharing by the owner. Food defensibility did not affect harassment or sharing. Interestingly, squirrel monkeys and chimpanzees shared equally frequently with conspecifics despite a much higher natural sharing rate in chimpanzees. These results suggest that harassment can play a significant role in primate food sharing, providing a simple alternative to reciprocity. The selfish nature of harassment has implications for economic, psychological and evolutionary studies of cooperative systems.

Analysis of Variance↗
PubMed1998

The evolution of generosity and choosiness in cooperative exchanges.

In this paper we present a resource-explicit Donor-Receiver model for reciprocally altruistic interactions that obeys the defining inequalities of the Prisoner's Dilemma. In our model, individuals vary in the quantity of resource they invest when cooperating (termed "generosity") and they have the freedom to opt out of interactions with potential partners on the basis of their past experiences with these players (termed "choosiness"). Dynamic optimal solutions were found using a genetic algorithm in which the decision rules (cooperate or defect), generosity when cooperating, and choosiness exhibited by individuals when deciding to opt out, were all coded on genes held on two separate chromosomes. Through this genetic algorithm, individuals that had alleles which resulted in greatest success at playing our modified Prisoner's Dilemma left more offspring. When the benefit of receiving a unit resource exceeded the cost of giving, then generous cooperative behaviour tended to emerge within the population, even when the alleles of all the individuals in the starting population were set to defect. When the probability of individuals re-encountering one another was increased, individuals not only cooperated more, but they developed greater generosity. However, as the ratio of the benefits received to costs expended increased above 1, individuals in this model remained highly cooperative but their median generosity decreased significantly. In contrast to earlier studies using genetic algorithms, the extra potential for cheating afforded by asymmetrical degrees of generosity meant that genuinely cooperative behaviour did not emerge in the equivalent round-robin tournament in which individuals were not able to exercise partner preference.

Animals↗