Kin selection in age-structured populations

There have been several discussions in the literature as to how to weight interactions between individuals of different ages in models of kin selection. It has commonly been assumed that the reproductive value of a given age is the most appropriate weight, for the purpose of calculating its contribution to inclusive fitness. This paper analyses a model of kin selection in an age-structured population. It is shown that reproductive value is relevant to behavioural interactions involving effects on survival, although the reproductive value of a given age does not provide an exact weighting of its fitness contribution in either discrete- or continuous-time populations. Reproductive value is not relevant to interactions involving effects on fecundity. The results are discussed in relation to observations on behavioural asymmetries involving age differences.     

Kin competition,natal dispersal and the moulding of senescence by natural selection

Most theoretical models for the evolution of senescence have assumed a very large, well mixed population. Here, we investigate how limited dispersal and kin competition might influence the evolution of ageing by deriving indicators of the force of selection, similar to Hamilton (Hamilton 1966 J. Theor. Biol. 12, 12–45). Our analytical model describes how the strength of selection on survival and fecundity changes with age in a patchy population, where adults are territorial and a fraction of juveniles disperse between territories. Both parent–offspring competition and sib competition then affect selection on age-specific life-history traits. Kin competition reduces the strength of selection on survival. Mutations increasing mortality in some age classes can even be favoured by selection, but only when fecundity deteriorates rapidly with age. Population structure arising from limited dispersal however selects for a broader distribution of reproduction over the lifetime, potentially slowing down reproductive senescence. The antagonistic effects of limited dispersal on age schedules of fecundity and mortality cast doubts on the generality of conditions allowing the evolution of ‘suicide genes’ that increase mortality rates without other direct pleiotropic effects. More generally, our model illustrates how limited dispersal and social interactions can indirectly produce patterns of antagonistic pleiotropy affecting vital rates at different ages.     

Intense selection in an age-structured population

In a population with overlapping generations, intense selection can perturb the age distribution and thus affect the rate of increase of an advantageous allele. We found that the age-specific nature of intense selection, such as that generated by many diseases, can affect the outcome of selection on loci, such as those conferring disease resistance. We also found that the temporal dynamics of selection alter the speed of evolution, particularly when selection is intense, and even more so when it is age-specific. We relate our model and results to selection for disease resistance, although the results have broader implications for inferences about past selection pressures in general.     

Philopatry and natal dispersal in a sedentary population of western marsh harrier

Dispersal is a key life-history trait because it influences population dynamics and population genetic structure. From a behavioural perspective, the study of natal dispersal requires some understanding of the mechanisms that affect individual movements, because movements of an animal form a path that is continuous throughout its life. Our aim was to investigate juvenile dispersal strategies in the western marsh harrier Circus aeruginosus , between fledging and first breeding attempt (from 1 to 4 years later, depending on the sex and individual). Using radio tracking, we monitored dispersing juvenile harriers and their home-range size variations within a sedentary population in central western France from 2001 to 2007. Juvenile dispersal strategy was mainly characterized by a very high natal philopatry (i.e. birds that remained within the study area) and short-distance dispersal. All but one bird (out of 39) remained within the study area between their first winter and their first spring, and 96.0% during their first spring. The distance moved at 2 years of age was significantly larger for males than for females (3800± sd 3593 m and 935± sd 481 m for seven males and six females, respectively), in contrast to most bird species studied so far. Home-range size was not sex biased and significantly decreased with age. In addition, non-breeding birds had larger home ranges (1603± sd 2128 ha) than breeders (349± sd 185 ha). Using data obtained from other populations, juvenile marsh harrier dispersal strategies appeared to be determined by migratory status (migratory birds dispersing farther) and demographic parameters (juvenile survival or fecundity).     

Evolution of cannibalism in an age-structured population

One of Bobisud's models for the evolution of cannibalism is reanalyzed by applying the method of finding evolutionarily stable strategies (or ESS's). It is demonstrated that ‘no cannibalism’ never will be an ESS if the initial rate of cannibalism is too large. It is further demonstrated that individual selection may even result in the evolution of cannibalism during food abundance. Some empirical case studies are briefly discussed in relation to this model.     

Optimal harvesting of an age-structured population

Here we investigate the optimal harvesting of an age-structured population. We use the McKendrick model of population dynamics, and optimize a discounted yield on an infinite time horizon. The harvesting function is allowed to depend arbitrarily on age and time and its magnitude is unconstrained. We obtain, in addition to existence, the qualitative result that an optimal harvesting policy consists of harvesting at no more than three distinct ages.     

Influential factors for natal dispersal in an avian island metapopulation

Dispersal is a key parameter in evolutionary, demographic and conservation theory, but the factors influencing dispersal between populations are rarely known, and the contribution of immigrants to population stability remains uncertain. Using dispersal data from nine island populations of song sparrows, we show that female and male immigrants responded differently to population structure: in females, immigration varied with adult sex ratio; whereas immigration by males was more influenced by population density. These patterns are consistent with the hypothesis that intra-sexual competition for breeding resources influenced recruitment patterns. Immigrants often constituted a substantial fraction of local population size, and in six cases immigration by females prevented the extirpation of that sex from the island. Breeding vacancies and extirpations may have been more likely in females because their apparent survival was lower than in males. Local recruitment and immigration varied markedly among islands, perhaps as consequence of island size and isolation. Overall, our results suggest that immigration varied with local demography in a sex-specific way, stabilized population numbers and reduced extinction rates in the smallest populations.     

Female-biased natal and breeding dispersal in an alpine lizard, Niveoscincus microlepidotus

We measured two aspects of dispersal in the alpine Australian scincid lizard, Niveoscincus micolepidotus : (1) natal dispersal, i.e. shift in home range over the lizard's first year of life, and (2) breeding dispersal, i.e. shifts of home ranges between breeding attempts as adults. On average, displacements were surprisingly small. Female neonates dispersed about twice as far as did males in the same cohort (means of 12 m vs. 6 m). A female's natal dispersal distance was not correlated with her body size or our estimate of physiological performance (sprint speed). However, larger, faster-running male neonates dispersed further than did smaller, slower males. As was the case for neonates, adult females moved significantly further between breeding seasons than did adult males (14.2 m vs. 9.6 m). Because of a female's long gestation period (more than 1 year), two groups of females occur simultaneously in the population, non-ovulated (i.e. with yolking folicles) and pregnant females (i.e. approaching parturition). Females that were not yet ovulated showed a markedly stronger dispersal in response to high reproductive effort (i.e. clutch size in relation to body condition) than did pregnant females. In adult males, body size was negatively correlated with dispersal distance, suggesting that although males have overlapping territories, they exhibit an increasing level of site tenacity with age and/or size. Thus, selection for the relatively more pronounced site tenacity in adult males may have resulted in the more marked philopatric behaviour compared to females also as neonates.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79 , 277–283.     

Senescence and antibiotic resistance in an age-structured population model

Different theories have been proposed to understand the growing problem of antibiotic resistance of microbial populations. Here we investigate a model that is based on the hypothesis that senescence is a possible explanation for the existence of so-called persister cells which are resistant to antibiotic treatment. We study a chemostat model with a microbial population which is age-structured and show that if the growth rates of cells in different age classes are sufficiently close to a scalar multiple of a common growth rate, then the population will globally stabilize at a coexistence steady state. This steady state persists under an antibiotic treatment if the level of antibiotics is below a certain threshold; if the level exceeds this threshold, the washout state becomes a globally attracting equilibrium.     

Habitat fragmentation shapes natal dispersal and sociality in an Afrotropical cooperative breeder

It remains poorly understood how effects of anthropogenic activity, such as large-scale habitat fragmentation, impact sociality in animals. In cooperatively breeding species, groups are mostly formed through delayed offspring dispersal, and habitat fragmentation can affect this process in two opposite directions. Increased habitat isolation may increase dispersal costs, promoting delayed dispersal. Alternatively, reduced patch size and quality may decrease benefits of philopatry, promoting dispersal. Here, we test both predictions in a cooperatively breeding bird (placid greenbul, Phyllastrephus placidus) from an Afrotropical cloud forest archipelago. Males born in fragmented forest dispersed about 1 year earlier than those born in continuous forest. Contrary to females, males also started to reproduce earlier and mostly settled within their natal patch. Females only rarely delayed their dispersal for more than 1 year, both in fragmented and continuous forests. Our results suggest that early male dispersal and reproduction is jointly driven by a decrease in the value of the natal territory and an increase in local breeding opportunities in fragmented forest. While plasticity in dispersal strategies of cooperative breeders in response to anthropogenic change is believed to optimize reproduction-survival trade-offs, to what extent it shapes the ability of species to respond to rapid environmental change remains to be studied.     

Kin selection and ethnic group selection

Ethnicity looks something like kinship on a larger scale. The same math can be used to measure genetic similarity within ethnic/racial groups and relatedness within families. For example, members of the same continental race are about as related (r = 0.18–0.26) as half-siblings (r = 0.25). However (contrary to some claims) the theory of kin selection does not apply straightforwardly to ethnicity, because inclusive fitness calculations based on Hamilton's rule break down when there are complicated social interactions within groups, and/or groups are large and long-lasting. A more promising approach is a theory of ethnic group selection, a special case of cultural group selection. An elementary model shows that the genetic assimilation of a socially enforced cultural regime can promote group solidarity and lead to the regulation of recruitment to groups, and to altruism between groups, based on genetic similarity – in short, to ethnic nepotism. Several lines of evidence, from historical population genetics and political psychology, are relevant here.     

植物的亲缘选择

亲缘选择是指在一个随机交配群体中的个体基于亲缘关系而以一种非随机性的方式相互作用,其作用结果是亲缘个体得到更大的广义适合度。综述了亲缘选择和亲缘竞争两种观点以及各自的试验支持证据;分析了导致亲缘选择试验结果出现分歧的原因,认为这主要是由于对亲缘选择理解上的模糊以及试验设计的不严谨所致。植物间的亲缘选择研究不仅相对较少,对亲缘选择的机制研究更为欠缺,这就造成了目前对此问题在科学认识上出现不少盲点。综合前期研究,提出今后对亲缘选择的研究应该首先界定"亲缘"程度,同时改良试验设计方案,选择多种不同生境下的物种对亲缘选择进行深入研究,并且考虑环境因子对植物亲缘选择的影响。同时,对植物亲缘识别机制的研究应该从生理生化方面出发,通过定性定量地分析探索植物根系分泌物在植物亲缘识别中的作用和作用途径。     

Kin groups and trait groups: population structure and epidemic disease selection

A Monte Carlo simulation based on the population structure of a small-scale human population, the Semai Senoi of Malaysia, has been developed to study the combined effects of group, kin, and individual selection. The population structure resembles D.S. Wilson's structured deme model in that local breeding populations (Semai settlements) are subdivided into trait groups (hamlets) that may be kin-structured and are not themselves demes. Additionally, settlement breeding populations are connected by two-dimensional stepping-stone migration approaching 30% per generation. Group and kin-structured group selection occur among hamlets the survivors of which then disperse to breed within the settlement population. Genetic drift is modeled by the process of hamlet formation; individual selection as a deterministic process, and stepping-stone migration as either random or kin-structured migrant groups. The mechanism for group selection is epidemics of infectious disease that can wipe out small hamlets particularly if most adults become sick and social life collapses. Genetic resistance to a disease is an individual attribute; however, hamlet groups with several resistant adults are less likely to disintegrate and experience high social mortality. A specific human gene, hemoglobin E, which confers resistance to malaria, is studied as an example of the process. The results of the simulations show that high genetic variance among hamlet groups may be generated by moderate degrees of kin-structuring. This strong microdifferentiation provides the potential for group selection. The effect of group selection in this case is rapid increase in gene frequencies among the total set of populations. In fact, group selection in concert with individual selection produced a faster rate of gene frequency increase among a set of 25 populations than the rate within a single unstructured population subject to deterministic individual selection. Such rapid evolution with plausible rates of extinction, individual selection, and migration and a population structure realistic in its general form, has implications for specific human polymorphisms such as hemoglobin variants and for the more general problem of the tempo of evolution as well.     

Demographic analysis from summaries of an age-structured population

Demographic analyses of age-structured populations typically rely on life history data for individuals, or when individual animals are not identified, on information about the numbers of individuals in each age class through time. While it is usually difficult to determine the age class of a randomly encountered individual, it is often the case that the individual can be readily and reliably assigned to one of a set of age classes. For example, it is often possible to distinguish first-year from older birds. In such cases, the population age structure can be regarded as a latent variable governed by a process prior, and the data as summaries of this latent structure. In this article, we consider the problem of uncovering the latent structure and estimating process parameters from summaries of age class information. We present a demographic analysis for the critically endangered migratory population of whooping cranes (Grus americana), based only on counts of first-year birds and of older birds. We estimate age and year-specific survival rates. We address the controversial issue of whether management action on the breeding grounds has influenced recruitment, relating recruitment rates to the number of seventh-year and older birds, and examining the pattern of variation through time in this rate.