Spotting altruistic dictator game players and mingling with them: the elective assortation of classmates

Altruism can evolve through assortation if the selfish advantage of egoistic individuals is outcompeted by the benefits of mutual cooperation between altruists. This selection process is possible if (a) individuals can distinguish altruists from egoists and (b) altruists cooperate electively with other altruists, leaving egoists no chance but to mingle with each other. This study investigates whether these two conditions are fulfilled in a natural setting. One hundred twenty-two students of six secondary school classes (age 10 to 19 years) played an anonymous dictator game, which functioned as a measure of altruism. Afterwards and unannounced, the students had to estimate their classmates' decisions and did so better than chance. Sociometry revealed that the accuracy of predictions depended on social closeness. Friends and disliked classmates were judged more accurately than liked classmates or those met with indifference. Moreover, altruists were friends with more altruistic persons than were egoists. The results confirm the existence of the two prerequisites for the evolution of altruism through assortation: the predictability of altruistic behavior and the association of altruists.     

Evolution,altruism and cognitive architecture: a critique of Sober and Wilson’s argument for psychological altruism

Sober and Wilson have propose a cluster of arguments for the conclusion that “natural selection is unlikely to have given us purely egoistic motives” and thus that psychological altruism is true. I maintain that none of these arguments is convincing. However, the most powerful of their arguments raises deep issues about what egoists and altruists are claiming and about the assumptions they make concerning the cognitive architecture underlying human motivation.     

Can altruism evolve in purely viscous populations?

Summary Limited dispersal is often thought to facilitate the evolution of altruism by increasing the degree of relatedness among interacting individuals. Limited dispersal can have additional effects, however, such as local population regulation, that inhibits the evolution of altruism. Many models of structured populations assume that a viscous stage of the life cycle alternates with a global mixing stage, which allows the advantages of interactions among close relatives without the disadvantages of local population regulation. Here we analyse a computer simulation model of pure population viscosity, in which offspring are always deposited close to parents and no global mixing stage exists. As expected, limited dispersal generates a high coefficient of relatedness among interacting individuals. Patches of altruists, however, are unable to export their productivity to other regions of the landscape and are easily invaded by selfish types from neighbouring patches. Unlike models of alternating viscosity, in which high relatedness and local population regulation can be decoupled, these two opposing effects are inextricably linked in purely viscous populations, which therefore are not conducive to the evolution of altruistic traits.     

Altruism, cheating, and anticheater adaptations in cellular slime molds

Cellular slime molds (CSMs) possess a remarkable life cycle that encompasses an extreme act of altruism. CSM cells live as individual amoebae until starved, then aggregate and ultimately transform themselves into a multicellular fruiting body. This fruiting body consists of stalk cells (altruists that eventually die) and spores (the beneficiaries of this sacrifice). Altruistic systems such as this are vulnerable to cheaters, which are individuals unrelated to the altruists that obtain the benefits provided by them without reciprocating. Here, we investigate two forces that can maintain CSM altruism despite cheating: kin selection and anticheater adaptations. First, we present new kinship-based models based on CSM developmental biology to evaluate the efficacy of kin selection. These models show that stalk-making genotypes can still be maintained when aggregations are initiated by multiple "founder" spores, provided that spores of stalkless fruiting bodies have low rates of dispersal and dispersal success is a concave function of stalk height. Second, we review proposals that several features of CSM development, such as the chemical suppression of the redifferentiation of prestalk cells into prespores, act as anticheater adaptations.     

Genetic relatedness in viscous populations

Summary Hamilton's inclusive fitness rule shows that the evolution of altruism is facilitated by high genetic relatedness of altruists to their beneficiaries. But the evolution of altruism is inhibited when the beneficiaries are also close competitors of the altruist, as will often be true in structured or viscous populations. However, Hamilton's rule still gives the correct condition for the evolution of altruism if relatedness is measured with respect to the local competitive neighbourhood.     

Nice natives and mean migrants: the evolution of dispersal-dependent social behaviour in viscous populations

There has been much interest in the evolution of social behaviour in viscous populations. While low dispersal increases the relatedness of neighbours, which tends to promote the evolution of indiscriminate helping behaviour, it can also increase competition between neighbours, which tends to inhibit the evolution of helping and may even favour harming behaviour. In the simplest scenario, these two effects exactly cancel, so that dispersal rate has no impact on the evolution of helping or harming. Here, we show that dispersal rate does matter when individuals can adjust their social behaviour conditional on whether they have dispersed or whether they have remained close to their place of origin. We find that nondispersing individuals are weakly favoured to indiscriminately help their neighbours, whereas dispersing individuals are more readily favoured to indiscriminately harm their neighbours.     

Mate finding, dispersal distances and population growth in invading species: a spatially explicit model

The expansion of populations into new areas is dependent upon dispersal distances and the ability of colonists to find mates. These factors interact through the spatial distribution of individuals. We develop a mechanistic, spatially explicit model to investigate the interaction between dispersal distances and mate finding in expanding populations. At high dispersal distances and low mate finding abilities, population growth was constrained by the inverse density dependent inability of adults to find mates (an Allee effect). In contrast, at low dispersal distances and high mate finding abilities, growth was constrained by the density dependent inability of dispersers to find vacant territories. Population growth was highest in between these extremes. We suggest that these spatial interactions play an important role in the expansion of populations into new areas and that this methodology provides a useful tool for investigating them.     

Natural selection of altruism in inelastic viscous homogeneous populations

Biological explanations are given of three main uninterpreted theoretical results on the selection of altruism in inelastic viscous homogeneous populations, namely that non-overlapping generations hinder the evolution of altruism, fecundity effects are more conducive to altruism than survival effects, and one demographic regime (so-called death-birth) permits altruism whereas another (so-called birth-death) does not. The central idea is ‘circles of compensation’, which measure how far the effects of density dependence extend from a focal individual. Relatednesses can then be calculated that compensate for density dependence. There is very generally a ‘balancing circle of compensation’, at which the viscosity of the population slows up selection of altruism, but does not affect its direction, and this holds for altruism towards any individual, not just immediate neighbours. These explanations are possible because of recent advances in the theory of inclusive fitness on graphs. The assumption of node bitransitivity in that recent theory is relaxed to node transitivity and symmetry of the dispersal matrix, and new formulae show how to calculate relatedness from dispersal and vice versa.     

Lower group productivity under kin-selected reproductive altruism

Abstract Hamilton's rule provides the foundation for understanding the genetic evolution of social behavior, showing that altruism is favored by increased relatedness and increased productivity of altruists. But how likely is it that a new altruistic mutation will satisfy Hamilton's rule by increasing the reproductive efficiency of the group? Altruism per se does not improve efficiency, and hence we would not expect a typical altruistic mutation to increase the mean productivity of the population. We examined the conditions under which a mutation causing reproductive altruism can spread when it does not increase productivity. We considered a population divided into temporary groups of genetically similar individuals (typically family groups). We show that the spread of altruism requires a pleiotropic link between altruism and enhanced productivity in diploid organisms, but not in haplodiploid organisms such as Hymenoptera. This result provides a novel biological understanding of the barrier to the spread of reproductive altruism in diploids. In haplodiploid organisms, altruism within families that lowers productivity may spread, provided daughters sacrifice their own reproduction to raise full‐sisters. We verified our results using three single‐locus genetic models that explore a range of the possible reproductive costs of helping. The advantage of female‐to‐female altruism in haplodiploids is a well‐known prediction of Hamilton's rule, but its importance in relaxing the linkage between altruism and efficiency has not been explored. We discuss the possible role of such unproductive altruism in the origins of sociality. We also note that each model predicts a large region of parameter space were polymorphism between altruism and selfishness is maintained, a pattern independent of dominance.     

Role of male dispersal in the evolution of female altruistic behavior in viscous populations

A computer simulation was conducted to examine the effect of differential dispersal of sexes on the evolution of altruism in viscous populations. First, a basic model, which was regarded as a purely viscous population model, was constructed. The model was assumed to be the same as the simulation model of Wilson etal. (1992), except that it assumed sexual reproduction and that only females show altruistic behavior toward females. For the basic model, altruism could not evolve when b/Nc, where b is the benefit of the altruism to the recipient, c is the cost to the altruist, and N is the number of interacting neighbors. The male dispersal model I assumed that females disperse to nine neighboring sites including the natal site, but males disperse to eight sites farther than females do. For this model, altruistic alleles could evolve when b/N was equal to c or b/N was slightly smaller than c only when the male dispersal distance was slightly larger than those of females. The male dispersal model II assumed that the male dispersal distance follows a normal probability distribution. The Vole model was based on actual data of the gray-sided vole, Clethironomys rufocanus bedfordiae, whose frequency distribution of dispersal distance was similar to a normal distribution. For these models, altruism could evolve under the condition that b/N was slightly smaller than c when the dispersal distances of males were larger than those of females. The results indicate that the differential dispersal of sexes, in which females are philopatric and males disperse farther than females, can somewhat increase the probability of spreading altruistic alleles in viscous populations.     

Multilevel selection and human altruism

Views on the evolution of altruism based upon multilevel selection on structured populations pay little attention to the difference between fortuitous and deliberate processes leading to assortative grouping. Altruism may evolve when assortative grouping is fortuitously produced by forces external to the organism. But when it is deliberately produced by the same proximate mechanism that controls altruistic responses, as in humans, exploitation of altruists by selfish individuals is unlikely and altruism evolves as an individually advantageous trait. Groups formed with altruists of this sort are special, because they are not affected by subversion from within. A synergistic process where altruism is selected both at the individual and at the group level can take place.     

Not only states but traits — Humans can identify permanent altruistic dispositions in 20 s

Humans behave altruistically in one-shot interactions under total anonymity. In search of explanations for such behavior, it has been argued that at least some individuals have a general tendency to behave altruistically independent of profitability. In fact, a stable altruistic trait would be adaptive if it were recognizable. Then, altruists could choose each other in order to retain benefits through mutual cooperation. Previous research has shown that individuals can predict the degree of altruistic behavior of strangers by reading signs of emotions evoked in significant social decisions. However, the identification of benevolent emotional states is no guarantee of the existence of permanent altruistic traits, though permanent traits are the preferable criterion for selection of good interaction partners. In this study, we tested whether individuals are able to identify altruistic traits. Judges watched 20-s silent video clips of unacquainted target persons and were asked to estimate the behavior of these target persons in a money-sharing task. As the videotapes of the target persons had been recorded in a setting unrelated to altruistic behavior, the judges could not base their estimates on situational cues related to the money-sharing task but instead had to draw on stable signals of altruism. Estimates were significantly better than chance, indicating that individuals can identify permanent altruistic traits in others. As this mechanism raises opportunities for selective interactions between altruists, our findings are discussed with respect to their relevance for explaining the evolution of altruism through assortment.     

Population viscosity and the evolution of altruism

The term population viscosity refers to limited dispersal, which increases the genetic relatedness of neighbors. This effect both supports the evolution of altruism by focusing the altruists' gifts on relatives of the altruist, and also limits the extent to which altruism may emerge by exposing clusters of altruists to stiffer local competition. Previous analyses have emphasized the way in which these two effects can cancel, limiting the viability of altruism. These papers were based on models in which total population density was held fixed. We present here a class of models in which population density is permitted to fluctuate, so that patches of altruists are supported at a higher density than patches of non-altruists. Under these conditions, population viscosity can support the selection of both weak and strong altruism.     

Dispersal of introduced house sparrows Passer domesticus: an experiment

An important issue concerning the introduction of non-indigenous organisms into local populations is the potential of the introduced individuals to spread and interfere both demographically and genetically with the local population. Accordingly, the potential of spatial dispersal among introduced individuals compared with local individuals is a key parameter to understand the spatial and temporal dynamics of populations after an introduction event. In addition, if the variance in dispersal rate and distance is linked to individual characteristics, this may further affect the population dynamics. We conducted a large-scale experiment where we introduced 123 house sparrows from a distant population into 18 local populations without changing population density or sex ratio. Introduced individuals dispersed more frequently and over longer distances than residents. Furthermore, females had higher probability of dispersal than males. In females, there was also a positive relationship between the wing length and the probability of dispersal and dispersal distance. These results suggest that the distribution and frequency of introduced individuals may be predicted by their sex ratio as well as their phenotypic characteristics.     

Competitive altruism: from reciprocity to the handicap principle

Current work on cooperation is focused on the theory of reciprocal altruism. However, reciprocity is just one way of getting a return on an investment in altruism and is difficult to apply to many examples. Reciprocity theory addresses how animals respond dynamically to others so as to cooperate without being exploited. I discuss how introducing differences in individual generosity together with partner choice into models of reciprocity can lead to an escalation in altruistic behaviour. Individuals may compete for the most altruistic partners and non-altruists may become ostracized. I refer to this phenomenon as competitive altruism and propose that it can represent a move away from the dynamic responsiveness of reciprocity. Altruism may be rewarded in kind, but rewards may be indirectly accrued or may not involve the return of altruism at all, for example if altruists tend to be chosen as mates. This variety makes the idea of competitive altruism relevant to behaviours which cannot be explained by reciprocity. I consider whether altruism might act as a signal of quality, as proposed by the handicap principle. I suggest that altruistic acts could make particularly effective signals because of the inherent benefits to receivers. I consider how reciprocity and competitive altruism are related and how they may be distinguished.     

WHEN HAWKS GIVE RISE TO DOVES: THE EVOLUTION AND TRANSITION OF ENFORCEMENT STRATEGIES

The question of how altruism can evolve despite its local disadvantage to selfishness has produced a wealth of theoretical and empirical research capturing the attention of scientists across disciplines for decades. One feature that has remained consistent through this outpouring of knowledge has been that researchers have looked to the altruists themselves for mechanisms by which altruism can curtail selfishness. An alternative perspective may be that just as altruists want to limit selfishness in the population, so may the selfish individuals themselves. These alternative perspectives have been most evident in the fairly recent development of enforcement strategies. Punishment can effectively limit selfishness in the population, but it is not free. Thus, when punishment evolves among altruists, the double costs of exploitation from cheaters and punishment make the evolution of punishment problematic. Here we show that punishment can more readily invade selfish populations when associated with selfishness, whereas altruistic punishers cannot. Thereafter, the establishment of altruism because of enforcement by selfish punishers provides the ideal invasion conditions for altruistic punishment, effectively creating a transition of punishment from selfishness to altruistic. Thus, from chaotic beginnings, a little hypocrisy may go a long way in the evolution and maintenance of altruism.     

Using Genealogical Mapping and Genetic Neighborhood Sizes to Quantify Dispersal Distances in the Neotropical Passerine,the Black-Capped Vireo

Dispersal is a key demographic process, ultimately responsible for genetic connectivity among populations. Despite its importance, quantifying dispersal within and between populations has proven difficult for many taxa. Even in passerines, which are among the most intensely studied, individual movement and its relation to gene flow remains poorly understood. In this study we used two parallel genetic approaches to quantify natal dispersal distances in a Neotropical migratory passerine, the black-capped vireo. First, we employed a strategy of sampling evenly across the landscape coupled with parentage assignment to map the genealogical relationships of individuals across the landscape, and estimate dispersal distances; next, we calculated Wright’s neighborhood size to estimate gene dispersal distances. We found that a high percentage of captured individuals were assigned at short distances within the natal population, and males were assigned to the natal population more often than females, confirming sex-biased dispersal. Parentage-based dispersal estimates averaged 2400m, whereas gene dispersal estimates indicated dispersal distances ranging from 1600–4200 m. Our study was successful in quantifying natal dispersal distances, linking individual movement to gene dispersal distances, while also providing a detailed look into the dispersal biology of Neotropical passerines. The high-resolution information was obtained with much reduced effort (sampling only 20% of breeding population) compared to mark-resight approaches, demonstrating the potential applicability of parentage-based approaches for quantifying dispersal in other vagile passerine species.     

Measures of gene flow in the Columbian ground squirrel

From analyses of published data and a review of the literature, I studied indirect and direct measures of gene flow among populations of Columbian ground squirrels, Spermophilus columbianus. New analyses were used to examine an allozyme data set (seven polymorphic loci) that had been collected by Zammuto and Millar (1985a) from six populations of ground squirrels that were spread over 183 km. G-tests indicated significant variation in allele frequencies among populations, but F-statistics revealed relatively little population differentiation (average F _ST=0.026). F _ST values were used to estimate rates of gene flow indirectly and indicated fairly high rates of gene flow (average N _e m=13.5). Recorded dispersal distances of individual ground squirrels were fairly short (most<4 km, maximum recorded distance was 8.5 km), and the minimum distance between populations used to create the allozyme data set was about 25 km. Thus, direct dispersal among the populations in the allozyme data set was highly unlikely. Small genetically effective populations may have experienced high rates of migration over short distances (about 43% of adults in local populations were immigrants), however, resulting in homogeneous allele frequencies over the geographic range. This explanation provides an alternative to invoking gene flow in the recent past to explain discrepancies between dispersal distances in the field and homogenization of allele frequencies over large ranges, Mammalian species that have virtually complete dispersal of subadult males from the natal area might be expected to exhibit relatively high rates of gene flow, regardless of actual dispersal distances. Genetically effective populations may be much smaller than more extensive ecological populations and experience higher rates of gene flow.     

The evolution of generalized reciprocity on social interaction networks

Generalized reciprocity (help anyone, if helped by someone) is a minimal strategy capable of supporting cooperation between unrelated individuals. Its simplicity makes it an attractive model to explain the evolution of reciprocal altruism in animals that lack the information or cognitive skills needed for other types of reciprocity. Yet, generalized reciprocity is anonymous and thus defenseless against exploitation by defectors. Recognizing that animals hardly ever interact randomly, we investigate whether social network structure can mitigate this vulnerability. Our results show that heterogeneous interaction patterns strongly support the evolution of generalized reciprocity. The future probability of being rewarded for an altruistic act is inversely proportional to the average connectivity of the social network when cooperators are rare. Accordingly, sparse networks are conducive to the invasion of reciprocal altruism. Moreover, the evolutionary stability of cooperation is enhanced by a modular network structure. Communities of reciprocal altruists are protected against exploitation, because modularity increases the mean access time, that is, the average number of steps that it takes for a random walk on the network to reach a defector. Sparseness and community structure are characteristic properties of vertebrate social interaction patterns, as illustrated by network data from natural populations ranging from fish to primates.     

浙江天台山甜槠种群遗传结构的空间自相关分析

采用空间自相关分析方法对浙江天台山亚热带常绿阔叶林优势种甜槠种群全部个体及不同年龄级个体的小尺度空间遗传结构进行了分析,以探讨甜槠种群内遗传变异的分布特征及其形成机制。根据11个ISSR引物所提供的多态位点,经GenAlEx 6软件计算地理坐标和遗传距离矩阵在10个距离等级下的空间自相关系数。在样地内,甜槠种群内个体在空间距离小于10 m时存在显著的正空间遗传结构,其X-轴截矩为9.945。甜槠种群的空间遗传结构与其种子短距离传播和广泛的花粉传播有关。Ⅰ年龄级、Ⅱ年龄级和III年龄级个体在空间距离小于10 m时存在显著的正空间遗传结构,其X-轴截矩分别为11.820、9.746和9.792。当距离等级为5 m时,其空间自相关系数r分别为0.068、0.054和0.070。Ⅳ年龄级个体在所有空间距离等级中均不存在显著的空间遗传结构。甜槠是多年生、长寿命植物,自疏作用是导致IV年龄级个体空间遗传结构消失的主要原因。