A MODEL FOR GENOMIC IMPRINTING IN THE SOCIAL BRAIN: JUVENILES

What are imprinted genes doing in the adult brain? Genomic imprinting is when a gene's expression depends upon parent of origin. According to the prevailing view, the “kinship theory” of genomic imprinting, this effect is driven by evolutionary conflicts between genes inherited via sperm versus egg. This theory emphasizes conflicts over the allocation of maternal resources, and focuses upon genes that are expressed in the placenta and infant brain. However, there is growing evidence that imprinted genes are also expressed in the juvenile and adult brain, after cessation of parental care. These genes have recently been suggested to underpin neurological disorders of the social brain such as psychosis and autism. Here we advance the kinship theory by developing an evolutionary model of genomic imprinting for social behavior beyond the nuclear family. We consider the role of demography and mating system, emphasizing the importance of sex differences in dispersal and variance in reproductive success. We predict that, in hominids and birds, altruism will be promoted by paternally inherited genes and egoism will be promoted by maternally inherited genes. In nonhominid mammals we predict more diversity, with some mammals showing the same pattern and other showing the reverse. We discuss the implications for the evolution of psychotic and autistic spectrum disorders in human populations with different social structures.     

A MODEL FOR GENOMIC IMPRINTING IN THE SOCIAL BRAIN: ADULTS

Genomic imprinting refers to genes that are silenced when inherited via sperm or via egg. The silencing of genes conditional upon their parental origin requires an evolutionary explanation. The most widely accepted theory for the evolution of genomic imprinting—the kinship theory—argues that conflict between maternally inherited and paternally inherited genes over phenotypes with asymmetric effects on matrilineal and patrilineal kin results in self‐imposed silencing of one of the copies. This theory has been applied to imprinting of genes expressed in the placenta, and infant brain determining the allocation of parental resources being the source of conflict parental promiscuity. However, there is growing evidence that imprinted genes are expressed in the postinfant brain where parental promiscuity per se is no longer a source of conflict. Here, we advance the kinship theory by developing an evolutionary model of genomic imprinting in adults, driven by intragenomic conflict over allocation to parental versus communal care. We consider the role of sex differences in dispersal and variance in reproductive success as sources of conflict. We predict that, in hominids and birds, parental care will be expressed by maternally inherited genes. In nonhominid mammals, we predict more diversity, with some mammals showing the same pattern and other showing the reverse. We use the model to interpret experimental data on imprinted genes in the house mouse: specifically, paternally expressed Peg1 and Peg3 genes, underlying maternal care, and maternally expressed Gnas and paternally expressed Gnasxl genes, underlying communal care. We also use the model to relate ancestral demography to contemporary imprinting disorders of adults, in humans and other taxa.     

THE EVOLUTION OF PHENOTYPIC PLASTICITY IN LIFE-HISTORY TRAITS: PREDICTIONS OF REACTION NORMS FOR AGE AND SIZE AT MATURITY

We used life-history theory to predict reaction norms for age and size at maturation. We assumed that fecundity increases with size and that juvenile mortality rates of offspring decrease as ages-at-maturity of parents increase, then calculated the reaction norm by varying growth rate and calculating an optimal age at maturity for each growth rate. The reaction norm for maturation should take one of at least four shapes that depend on specific relations between changes in growth rates and changes in adult mortality rates, juvenile mortality rates, or both. Most organisms should mature neither at a fixed size nor at a fixed age, but along an age-size trajectory. The model makes possible a clear distinction between the genetic and phenotypic components of variation. The evolved response to selection is reflected in the shape and position of the reaction norm. The phenotypic response of a single organism to rapid or slow growth is defined by the location of its maturation event as a point on the reaction norm. A quantitative test with data from 19 populations and species of fish showed that predictions were in good agreement with observations (r = 0.93, P < 0.0001). The predictions of the model also agreed qualitatively with observed phenotypic variation in age and size at maturity in humans, platyfish, fruit flies, and red deer. This preliminary success suggests that experiments designed to test the predictions directly will be worthwhile.     

FEMALE-BIASED SEX RATIOS IN A FACULTATIVELY SOCIAL BEE AND THEIR IMPLICATIONS FOR SOCIAL EVOLUTION

Montane populations of the Australian allodapine bee, Exoneura bicolor, are characterized by high levels of cooperative nesting and strongly female-biased sex ratios. A conspecific population from heathland shows much lower levels of cooperative nesting and lower levels of female bias. In both habitats, sex-ratio bias is greatest in the smallest brood sizes and becomes successively less biased in larger broods. Parity is approached in the largest heathland colonies, but not for any brood-size category in montane areas. Adult intracolony relatedness is moderately high for colonies in both reused and newly founded nests in the montane habitat, but probably low or zero for newly founded nests in heathland. Colony efficiency, measured as the number of brood per adult, increases with colony size in both habitats, suggesting that cooperation between females increases mean female fitness. It is argued that patterns of sex allocation are consistent with nonlinear fitness-return models, in which the mean reproductive value of daughters increases with the number of daughters produced in a brood. Such increases probably arise from a number of social interactions, including cooperative brood defense, increased task efficiency, and lower per capita costs in nest construction. The term “local fitness enhancement” is introduced here to describe these effects collectively. The female-biased ratios should lower selective thresholds for sib-directed altruism, at least in the earlier stages of colony development. It is argued that local fitness enhancement facilitates eusociality in allodapine bees and could also play a role in other haplodiploid taxa, provided cooperative nesting largely involves sisters, colony efficiency increases with colony size, and optimal colony sizes are only achieved after two or more generations after founding.     

THE SOCIAL ORGANISM: CONGRESSES,PARTIES, AND COMMITTEES

We propose that what makes an organism is nearly complete cooperation, with strong control of intraorganism conflicts, and no affiliations above the level of the organism as unified as those at the organism level. Organisms can be made up of like units, which we call fraternal organisms, or different units, making them egalitarian organisms. Previous definitions have concentrated on the factors that favor high cooperation and low conflict, or on the adapted outcomes of organismality. Our approach brings these definitions together, conceptually unifying our understanding of organismality. Although the organism is a concerted cluster of adaptations, nearly all directed toward the same end, some conflict may remain. To understand such conflict, we extend Leigh's metaphor of the parliament of genes to include parties with different interests and committees that work on particular tasks.     

GnRH及其受体在性成熟前后鲇、黄颡鱼脑、垂体和卵巢中的免疫细胞化学定位

用链霉亲和素 -生物素化过氧化物酶复合物 (StreptAvidinBiotin peroxidaseComplex ,SABC)免疫细胞化学方法 ,使用促性腺激素释放激素 (Gonadotropin releasinghormone ,GnRH)以及促性腺激素释放激素受体 (GnRHR) 2种抗血清对性成熟前后的黄颡鱼 (Pelteobagrusfulvidraco)和鲇鱼 (Silurusasotus)的脑、垂体、卵巢中的免疫活性内分泌细胞进行了免疫细胞化学定位。结果表明GnRH和GnRHR免疫活性在两种鱼的各脑区、垂体、卵巢中均有分布 ;两种鱼在性成熟时它们的下丘脑、垂体和卵巢中的GnRH和GnRHR免疫反应细胞数目和免疫反应强度明显高于性成熟前。本文讨论了GnRH、GnRHR直接或间接参与黄颡鱼和鲇鱼性腺发育成熟调节的可能性及形态学证据。可为下丘脑 垂体 性腺轴、神经 内分泌、GnRH功能的多样性等研究领域提供新的形态学依据。     

SEX-INVESTMENT RATIOS AND RELATEDNESS IN THE MONOGYNOUS ANT LASIUS NIGER (L.)

The genetic variation at two marker loci in three populations of the monogynous ant Lasius niger was used to analyze the importance of relatedness structure to sex-investment ratios in sexuals produced by colonies living in different resource conditions. From a resource-rich monoculture to a population in a resource-limited environment, dry weight investment in queens decreased from female-biased (0.76) to equality with male investment (Boomsma et al., 1982). The investment ratios in sexuals expected from the estimated relatedness ratios, resulting from kin-selection theory (Trivers and Hare, 1976), were in good agreement with the observed ratios in all populations. This indicated that the workers can capitalize on the asymmetry in relatedness, opposing the queen's interest, despite the contrast in environmental conditions in the different populations. This asymmetry, however, almost disappeared in the marginal population, due to a high frequency of double mating and worker production of males, resulting in a much reduced queen-worker conflict. We suggest that different levels of polyandry might be favored at different points of the resource gradient, with the sex ratio secondarily depending on these polyandry levels. As there was no population subdivision or spatial structure within the populations, group-selection and local-mate-competition models cannot account for the observed female-biased ratios, whereas they were rather accurately predicted by kin-selection theory.