Evolution of learned strategy choice in a frequency-dependent game

In frequency-dependent games, strategy choice may be innate or learned. While experimental evidence in the producer-scrounger game suggests that learned strategy choice may be common, a recent theoretical analysis demonstrated that learning by only some individuals prevents learning from evolving in others. Here, however, we model learning explicitly, and demonstrate that learning can easily evolve in the whole population. We used an agent-based evolutionary simulation of the producer-scrounger game to test the success of two general learning rules for strategy choice. We found that learning was eventually acquired by all individuals under a sufficient degree of environmental fluctuation, and when players were phenotypically asymmetric. In the absence of sufficient environmental change or phenotypic asymmetries, the correct target for learning seems to be confounded by game dynamics, and innate strategy choice is likely to be fixed in the population. The results demonstrate that under biologically plausible conditions, learning can easily evolve in the whole population and that phenotypic asymmetry is important for the evolution of learned strategy choice, especially in a stable or mildly changing environment.     

How does hunger affect convergence on prey patches in a social forager?

Internal state, in this case hunger, is known to influence both the organisation of animal groups and the social foraging interactions that occur within them. In this study, we investigated the effects of hunger upon the time taken to locate and converge upon hidden simulated prey patches in a socially foraging fish, the threespine stickleback (Gasterosteus aculeatus). We predicted that groups of food‐deprived fish would find and recruit to prey patches faster than recently fed groups, reasoning that they might search more rapidly and be more attentive to inadvertent social information produced by other foragers. Instead we saw no difference between the two groups in the time taken to find the patches and found that in fact, once prey patches had been discovered, it was the recently fed fish that converged on them most rapidly. This finding is likely due to the fact that recently fed fish tend to organise themselves into fewer but larger subgroups, which arrived at the food patch together. Hunger has a significant impact upon the social organisation of the fish shoals, and it appears that this has a stronger effect upon the rate at which they converged upon the food patches than does internal state itself.     

Individual differences in plasticity and sampling when playing behavioural games

When engaged in behavioural games, animals can adjust their use of alternative tactics until groups reach stable equilibria. Recent theory on behavioural plasticity in games predicts that individuals should differ in their plasticity or responsiveness and hence in their degree of behavioural adjustment. Moreover, individuals are predicted to be consistent in their plasticity within and across biological contexts. These predictions have yet to be tested empirically and so we examine the behavioural adjustment of individual nutmeg mannikins (Lonchura punctulata), gregarious ground-feeding passerines, when playing two different social foraging games: producer-scrounger (PS) and patch-choice (PC) games. We found: (i) significant individual differences in plasticity and sampling behaviour in each of the two games, (ii) individual differences in sampling behaviour were consistent over different test conditions within a game (PC) and over a six month period (PS), (iii) but neither individual plasticity nor sampling behaviour was correlated from one social foraging game to another. The rate at which birds sampled alternative tactics was positively associated with seed intake in PS trials but negatively associated in PC trials. These results suggest that games with frequency dependence of pay-offs can maintain differences in behavioural plasticity but that an important component of this plasticity is group- and/or context-specific.     

领燕鸻繁殖群体的集群觅食

集合点假说（Assembly—point hypothesis）认为：动物局部地区聚群数量的增加有助于不同的繁殖地的个体聚集成群体,然后一起寻找食物斑块觅食,这种聚群有助于个体从集群觅食中获得利益。本文通过领燕鸻（Glareola pratincola）离开和回到繁殖地的先后顺序来验证以上假说。我们在29天的观察中,发现有4天该群中个体离开繁殖地的先后顺序不是随机分布的,尽管这种策略会随着环境条件而改变,但仍然表明领燕鸻的觅食群体在繁殖地就已经聚集形成了。食物资源（飞虫）的短暂性和波动性以及由繁殖地到觅食地的距离均与所验证的假说的假设相一致[动物学报51（6）：1141—1145,2005]。     

Synergy and discounting of cooperation in social dilemmas

The emergence and maintenance of cooperation by natural selection is an enduring conundrum in evolutionary biology, which has been studied using a variety of game theoretical models inspired by different biological situations. The most widely studied games are the Prisoner's Dilemma, the Snowdrift game and by-product mutualism for pairwise interactions, as well as Public Goods games in larger groups of interacting individuals. Here, we present a general framework for cooperation in social dilemmas in which all the traditional scenarios can be recovered as special cases. In social dilemmas, cooperators provide a benefit to the group at some cost, while defectors exploit the group by reaping the benefits without bearing the costs of cooperation. Using the concepts of discounting and synergy for describing how benefits accumulate when more than one cooperator is present in a group of interacting individuals, we recover the four basic scenarios of evolutionary dynamics given by (i) dominating defection, (ii) coexistence of defectors and cooperators, (iii) dominating cooperation and (iv) bi-stability, in which cooperators and defectors cannot invade each other. Generically, for groups of three or more interacting individuals further, more complex, dynamics can occur. Our framework provides the first unifying approach to model cooperation in different kinds of social dilemmas.     

Pavlovian Prisoner's Dilemma—Analytical results, the quasi-regular phase and spatio-temporal patterns

The Prisoner's Dilemma (PD) game is applied in several research fields due to the emergence of cooperation among selfish players. In this work the PD is studied in a one-dimensional lattice, where each cell represents a player, which in turn can interact with the neighbors playing the PD (cooperate or defect). The update of states adopts the Pavlovian Evolutionary Strategy (PES) or Darwinian Evolutionary Strategy (DES). Adopting PES, if a player receives a positive payoff greater than his/her aspiration level, he/she keeps the current state, and switches otherwise. Adopting DES, player compares his/her payoff with payoff of opponents. If it is not the highest, he/she copies the state of fittest player, switching the state if it is different of his/her current state. The critical temptation values obtained analytically are reported, and the cluster patterns that emerge from the interactions among the players are shown. Also we defined analytical functions that calculate the maximum/minimum size of defective/cooperative clusters. Also, the parameter space is explored with exhaustive computational simulations, which confirm the analytical results and reinforce that Pavlovian strategy foments cooperation among players. In steady state, system can reach the cooperative or quasi-regular phases, when adopting the PES, and cooperative, defective or chaotic phases, adopting the DES. The new quasi-regular phase occurs when several players switch their states in each round, but the proportion of cooperators does not show significant variation. Additionally, the present work shows that the lowest temptation level (T=1) may be considered a trivial case only for the particular case where the players interact with only one neighbor, otherwise system presents the same features that for higher temptation values.     

Oxytocin and cooperation under conditions of uncertainty: The modulating role of incentives and social information

The neuropeptide Oxytocin (OT) has been implicated in many aspects of mammalian social behavior. This study investigates how OT interacts with two well-studied determinants of cooperative behavior: incentives and social information. Participants received OT or a placebo and played two economic games: a Coordination Game (with strong incentives to cooperate) and a Prisoner's Dilemma (with weak cooperative incentives). OT enhanced cooperation only when social information was present, and this effect was significantly more pronounced in the Coordination Game. When social information was lacking, OT surprisingly decreased cooperation. Consistent with the well-established role of OT in trust-building and in social cognition, social information appears to be crucial for OT to boost cooperative expectations in an interdependent social interaction that provides incentives to cooperate. When these cues are absent, OT appears to instead elicit a risk-averse strategy.     

Fairness evolution in the ultimatum game is a function of reward size

I formulate a simple model of the ultimatum game, in which a proposer and a responder can receive a reward if they agree on how to divide this reward between them. The model is easy to analyse and shows that strong tendencies to fair division are expected when evolution of strategy frequencies follow the traditional gradient dynamics assumed in evolutionary models. The mean stable offer is typically around 20-40% although this depends on the maximum payoff and if rejection thresholds can evolve independently from proposals. The stable proportion offered at evolutionary equilibrium increases with the maximum payoff, if proposal and acceptance thresholds are dictated by the same strategy and cannot evolve independently. If proposal and acceptance evolve independently, the stable proportion instead decreases with the maximum payoff. The stable outcome may also show substantial variation.     

How inconsistency between attitude and behavior persists through cultural transmission

Individuals tend to conform their behavior to that of the majority. Consequently, an individual's behavior is not always consistent with his or her attitude, and such inconsistency sometimes causes mental distress. Understanding the mechanism of sustaining inconsistency between attitude and behavior is a challenging problem from the viewpoint of evolutionary theory. We constructed an evolutionary game theory model in which each player has an attitude and behavior toward a single social norm, and the players' attitudes and behaviors are affected by three types of cultural transmission: vertical, oblique, and horizontal. We assumed that strategy is a combination of attitude and behavior and that the process of learning or transmitting the social norm depends on the life stage of each player. Adults play a coordination game in which players whose behaviors match those of the majority obtain a high payoff, which is diminished by any inconsistency between attitude and behavior. The adults' strategies are passed to newborns via vertical transmission, and the frequency of a newborn's replication of strategy is proportional to the corresponding adult's payoff. Newborns imitate behaviors of unrelated adults via oblique transmission. Juveniles change their attitudes or behaviors by observing other juveniles' behaviors or inferring other juveniles' attitudes (horizontal transmission). We conclude that the key factor for sustaining inconsistency between attitude and behavior is the ability of players to infer and imitate others' attitudes, and that oblique transmission promotes inconsistency.     

Temporal partitioning and aggression among foragers: modeling the effects of stochasticity and individual state

In many natural systems, individuals compete with conspecificsand heterospecifics for food and in some cases, individualshave been observed to partition their foraging times or fightover food. In this study, I investigated when it is optimalfor a consumer to partition time and be aggressive. I formulatedan individual-based model of foraging and used game theoryto find evolutionarily stable strategies (ESSs) that maximizethe probability that consumers survive each day and acquiretheir daily food requirements. Consumers choose when to forageand when to behave aggressively during confrontations overfood. Consumers are each associated with a state variable,representing the amount of food eaten, and a dominance ranking,which describes how likely they are to forage and fight forfood. The ESS is sensitive to food abundance, consumer state,and the dominance ranking. When food is abundant, temporalpartitioning is often an ESS where the dominant consumer foragesfirst; however, partitioning is unlikely to be an ESS whenfood abundance is low. Fights over food are typically avoidedbut may be part of an ESS when food abundance is low, bothconsumers are hungry, or the time available for foraging eachday is drawing to a close. Because the ESS is sensitive toconsumer state, the stochastic nature of finding food oftenresults in considerable variation in observed foraging dynamicsfrom one day to the next, even when consumers adopt the samestate-dependent strategy each day. Results are compared withempirical observations, and I discuss implications for consumercoexistence.     

Evolutionary Psychology, Human Universals, and the Standard Social Science Model

Proponents of evolutionary psychology take the existence of humanuniversals to constitute decisive evidence in favor of their view. Ifthe same social norms are found in culture after culture, we have goodreason to believe that they are innate, they argue. In this paper Ipropose an alternative explanation for the existence of humanuniversals, which does not depend on them being the product of inbuiltpsychological adaptations. Following the work of Brian Skyrms, I suggestthat if a particular convention possesses even a very small advantageover competitors, whatever the reason for that advantage, we shouldexpect it to become the norm almost everywhere. Tiny advantages aretranslated into very large basins of attraction, in the language of gametheory. If this is so, universal norms are not evidence for innatepsychological adaptations at all. Having shown that the existence ofuniversals is consistent with the so-called Standard Social ScienceModel, I turn to a consideration of the evidence, to show that thisstyle of explanation is preferable to the evolutionary explanation, atleast with regard to patterns of gender inequality.     

Neuroethology of reward and decision making

Ethology, the evolutionary science of behaviour, assumes that natural selection shapes behaviour and its neural substrates in humans and other animals. In this view, the nervous system of any animal comprises a suite of morphological and behavioural adaptations for solving specific information processing problems posed by the physical or social environment. Since the allocation of behaviour often reflects economic optimization of evolutionary fitness subject to physical and cognitive constraints, neurobiological studies of reward, punishment, motivation and decision making will profit from an appreciation of the information processing problems confronted by animals in their natural physical and social environments.