The general protected invasion theory: Sex biases in parental and alloparental care

The biases towards eusociality, female workers and maternal care in haplodiploid versus diploid insects may result from the relatively low probabilities that rare mutant, partially dominant alleles promoting these behaviours will be lost by genetic drift in haplodiploid populations (Reeve, 1993). A generalization of this 'protected invasion' theory also predicts that parental and alloparental care will tend to be associated with the homogametic sex in diploid populations if the Y chromosome of the heterogametic sex is absent or largely inert. Sex differences in (allo)parental care (i.e. either parental or alloparental care) should increase with increased asymmetry between the sexes in the fraction of behaviour-influencing loci occurring on their characteristic sex chromosomes. The theory explains the strong predisposition towards female (allo)parental care in mammals, a contrasting tendency towards male (allo)parental care in birds, the propensity for joint male and female (allo)parental care in termites, and biases towards female cooperation in social spiders. The theory also explains the apparent rarity or absence of alloparental care in marsupials, an intriguing consequence of preferential paternal X-chromosome inactivation in this taxon. Thus protected invasion theory possibly provides new insights into the relationship between social structure and the genetic system. The theory does not compete with ecological or kin-selective hypotheses for the advantages of (allo)parental care; indeed, such advantages must exist for protected-invasion biases to operate.     

Demography and the tragedy of the commons

Individual success in group‐structured populations has two components. First, an individual gains by outcompeting its neighbours for local resources. Second, an individual's share of group success must be weighted by the total productivity of the group. The essence of sociality arises from the tension between selfish gains against neighbours and the associated loss that selfishness imposes by degrading the efficiency of the group. Without some force to modulate selfishness, the natural tendencies of self interest typically degrade group performance to the detriment of all. This is the tragedy of the commons. Kin selection provides the most widely discussed way in which the tragedy is overcome in biology. Kin selection arises from behavioural associations within groups caused either by genetical kinship or by other processes that correlate the behaviours of group members. Here, I emphasize demography as a second factor that may also modulate the tragedy of the commons and favour cooperative integration of groups. Each act of selfishness or cooperation in a group often influences group survival and fecundity over many subsequent generations. For example, a cooperative act early in the growth cycle of a colony may enhance the future size and survival of the colony. This time‐dependent benefit can greatly increase the degree of cooperation favoured by natural selection, providing another way in which to overcome the tragedy of the commons and enhance the integration of group behaviour. I conclude that analyses of sociality must account for both the behavioural associations of kin selection theory and the demographic consequences of life history theory.     

A general model of the public goods dilemma

An individually costly act that benefits all group members is a public good. Natural selection favours individual contribution to public goods only when some benefit to the individual offsets the cost of contribution. Problems of sex ratio, parasite virulence, microbial metabolism, punishment of noncooperators, and nearly all aspects of sociality have been analysed as public goods shaped by kin and group selection. Here, I develop two general aspects of the public goods problem that have received relatively little attention. First, variation in individual resources favours selfish individuals to vary their allocation to public goods. Those individuals better endowed contribute their excess resources to public benefit, whereas those individuals with fewer resources contribute less to the public good. Thus, purely selfish behaviour causes individuals to stratify into upper classes that contribute greatly to public benefit and social cohesion and to lower classes that contribute little to the public good. Second, if group success absolutely requires production of the public good, then the pressure favouring production is relatively high. By contrast, if group success depends weakly on the public good, then the pressure favouring production is relatively weak. Stated in this way, it is obvious that the role of baseline success is important. However, discussions of public goods problems sometimes fail to emphasize this point sufficiently. The models here suggest simple tests for the roles of resource variation and baseline success. Given the widespread importance of public goods, better models and tests would greatly deepen our understanding of many processes in biology and sociality.     

Direct fitness for dynamic kin selection

The direct-fitness approach to modelling the evolution of social traits is an alternative to the classical inclusive-fitness-based approach. Despite both its utility and popularity, the direct-fitness approach has not yet been extended to include the analysis of dynamic traits, i.e. traits whose level of expression may vary over time. In this article, I apply the direct-fitness approach to cope with the evolution of a dynamic resource-allocation behaviour when this behaviour influences the fitness of relatives. I am able to implement the direct-fitness approach using components (reproductive value, fitness changes and measures of relatedness) found in standard, social-evolutionary models. I illustrate the modified direct-fitness model with an example studied by previous authors, and I show how the direct-fitness perspective can aid the validation of analytical results by means of a genetic algorithm.     

Individual and social discounting in a viscous population

Social discounting in economics involves applying a diminishing weight to community-wide benefits or costs into the future. It impacts on public policy decisions involving future positive or negative effects, but there is no consensus on the correct basis for determining the social discount rate. This study presents an evolutionary biological framework for social discounting. How an organism should value future benefits to its local community is governed by the extent to which members of the community in the future are likely to be its kin. Trade-offs between immediate and delayed benefits to an individual or to its community are analysed for a modelled patch-structured iteroparous population with limited dispersal. It is shown that the social discount rate is generally lower than the individual (private) discount rate. The difference in the two rates is most pronounced, in ratio terms, when the dispersal level is low and the hazard rate for patch destruction is much smaller than the individual mortality rate. When decisions involve enforced collective action rather than individuals acting independently, social investment increases but the social discount rate remains the same.     

植物亲缘识别的研究进展

植物的亲缘识别(kin recognition)指植物通过识别周边个体与自己的亲缘关系, 调整自身的生长生态策略、促进亲缘个体的生存与繁衍。研究表明, 植物主要通过特定的叶片挥发物、根系分泌物、感光载体等途径, 识别周边个体与自己的亲缘关系, 改变自身形态学策略(如根系大小、根冠比、种子数量等)或者生理代谢策略(次生代谢物质、防御蛋白等), 调整与周边个体的竞争强度, 缓和与近亲缘个体之间的竞争, 加强与远亲缘或非亲缘个体的竞争。同时亲缘识别的强度也受环境因子(养分等)的影响。结合目前的研究进展, 该文分析了导致亲缘识别的研究结果存在差异或争议的主要原因, 认为主要与实验材料的选择、亲缘关系的界定标准、环境条件及测定的指标不统一有关。将来的研究应重点从生理生化、分子、代谢水平上深入研究植物亲缘识别的机理。