Selection in complex genetic systems IV. Multiple alleles and interactions between two loci

Summary A symmetric viability model for two loci with two alleles at one locus and m alleles at the other is suggested and analyzed. The analysis of the equilibria is complete if the two loci are absolutely linked, while if recombination is allowed the analysis is incomplete. The dynamics of the mode! resemble those of the two locus two allele model, namely that for loose linkage there will be no correlation between the loci and for tight linkage there may be strong correlation. The major caveats to this are: 1. The equilibria stable for tight linkage may belong to an array of different structures dependent on the selection and the number of alleles. 2. If both loci are overdominant in viability, the stable equilibria always contain all alleles segregating in the population; otherwise, the stable equilibria may only be two locus two allele high complementarity equilibria for tight linkage. 3. For intermediate linkage values and special selection values the boundary two locus two allele high complementarity equilibria may be stable simultaneously with the totally polymorphic central point at which there is no association between the loci.Dedicated to the memory of Ove Frydenberg.Research supported in part by a grant from the Danish Natural Science Research Council, a grant from National Science Foundation, U.S.A., and by USPHS grant NIH 10452-09-11.     

Selective sweeps for recessive alleles and for other modes of dominance

A selective sweep describes the reduction of linked genetic variation due to strong positive selection. If s is the fitness advantage of a homozygote for the beneficial allele and h its dominance coefficient, it is usually assumed that h = 1/2, i.e. the beneficial allele is co-dominant. We complement existing theory for selective sweeps by assuming that h is any value in [0, 1]. We show that genetic diversity patterns under selective sweeps with strength s and dominance 0 < h < 1 are similar to co-dominant sweeps with selection strength 2hs. Moreover, we focus on the case h = 0 of a completely recessive beneficial allele. We find that the length of the sweep, i.e. the time from occurrence until fixation of the beneficial allele, is of the order of \({\sqrt{N/s}}\) generations, if N is the population size. Simulations as well as our results show that genetic diversity patterns in the recessive case h = 0 greatly differ from all other cases.     

Kin selection and frequency dependence: a game theoretic approach

Game theory models show that the evolution of interactions between relatives is determined by two kinds of fitness effects: Hamilton's inclusive fitness effect, and a frequency-dependent synergistic effect. The latter arises when an individual's behaviour has different effects on the fitness of interactants, depending on whether or not they perform the same behaviour. Knowing the sign of the synergistic effect is sufficient to understand most of the qualitative features of genetic models that show departures from Hamilton's rule. Since this synergistic effect does not depend on the interactants being related, it is best viewed as something distinct from kin selection. In this view, Hamilton's rule is basically correct for describing kin selection, and most deviations from it are due to the distinct process of synergistic selection.     

Nonlinear frequency-dependent selection at a single locus with two alleles and two phenotypes

 The paper investigates the discrete frequency dynamics of two phenotype diploid models where genotypic fitness is an exponential function of the expected payoff in the matrix game. Phenotypic and genotypic equilibria are defined and their stability compared to frequency-dependent selection models based on linear fitness when there are two possible phenotypes in the population. In particular, it is shown that stable equilibria of both types can exist in the same nonlinear model. It is also shown that period-doubling bifurcations emerge when there is sufficient selection in favor of interactions between different phenotypes. Received: 22 October 1998     

Selection for costly sexual traits results in a vacant mating niche and male dimorphism

The expected strong directional selection for traits that increase a male's mating ability conflicts with the frequent observation that within species, males may show extreme variation in sexual traits. These male reproductive polymorphisms are usually attributed to direct male–male competition. It is currently unclear, however, how directional selection for sexually selected traits may convert into disruptive selection, and if female preference for elaborate traits may be an alternative mechanism driving the evolution of male polymorphism. Here, we explore this mechanism using the polyandric dwarf spider Oedothorax gibbosus as a model. We first show that males characterized by conspicuous cephalic structures serving as a nuptial feeding device (“gibbosus males”) significantly outperform other males in siring offspring of previously fertilized females. However, significant costs in terms of development time of gibbosus males open a mating niche for an alternative male type lacking expensive secondary sexual traits. These “tuberosus males” obtain virtually all fertilizations early in the breeding season. Individual‐based simulations demonstrate a hitherto unknown general principle, by which males selected for high investment to attract females suffer constrained mating opportunities. This creates a vacant mating niche of unmated females for noninvesting males and, consequently, disruptive selection on male secondary sexual traits.     

Environmental complexity and the purging of deleterious alleles

Sexual interactions among adults can generate selection on both males and females with genome‐wide consequences. Sexual selection through males is one component of this selection that has been argued to play an important role in purging deleterious alleles. A common technique to assess the influence of sexual selection is by a comparison of experimental evolution under enforced monogamy versus polygamy. Mixed results from past studies may be due to the use of highly simplified laboratory conditions that alter the nature of sexual interactions. Here, we examine the rate of purging of 22 gene disruption mutations in experimental polygamous populations of Drosophila melanogaster in each of two mating environments: a simple, high‐density environment (i.e., typical fly vials), and a lower density, more spatially complex environment. Based on past work, we expect sexual interactions in the latter environment to result in stronger selection in both sexes. Consistent with this, we find that mutations tend to be purged more quickly in populations evolving in complex environments. We discuss possible mechanisms by which environmental complexity might modulate the rate at which deleterious alleles are purged and putatively ascribe a role for sexual interactions in explaining the treatment differences in our experiment.     

Selection in a subdivided population with dominance or local frequency dependence

The interplay between population structure and natural selection is an area of great interest. It is known that certain types of population subdivision do not alter fixation probabilities of selected alleles under genic, frequency-independent selection. In the presence of dominance for fitness or frequency-dependent selection these same types of subdivision can have large effects on fixation probabilities. For example, the barrier to fixation of a fitter allele due to underdominance is reduced by subdivision. Analytic results presented here relate a subdivided population that conforms to a finite island model to an approximately equivalent panmictic population. The size of this equivalent population is different from (larger than) the actual size of the subdivided population. Selection parameters are also different in the hypothetical equivalent population. As expected, the degree of dominance is lower in the equivalent population. The results are not limited to dominance but cover any form of polynomial frequency dependence.     

Cloning and complete sequence of an HLA-A2 gene: analysis of two HLA-A alleles at the nucleotide level

The gene for the HLA-A2 antigen has been cloned from the human lymphoblastoid cell line 721. Comparison of this sequence with the published sequence for HLA-A3 permits the examination of two alleles at the extremely polymorphic HLA-A locus. A high degree of sequence conservation was seen in both coding and noncoding DNA, 97.2% and 94.5%, respectively. Interestingly, the 3' untranslated region was the most conserved, with 99.5% homology. The polymorphism of the HLA-A antigens results from a high proportion of amino acid substitutions relative to the total nucleotide changes in exons 2 and 3. Unlike the clustered differences seen in this region on comparison of two H-2K alleles of mouse, nucleotide substitutions between the HLA-A2 and A3 alleles are evenly distributed. Substitutions at silent sites and within introns were used to calculate an intra-allelic divergence time of at least 10 to 15 million years for these two HLA-A alleles.     

Selection,linkage, and dominance in small populations

Summary Linkage, dominance, and selection interact significantly to alter the mean coefficient of inbreeding. The effect of one is not predictable without the other two. Close linkage between adjacent loci in the presence of intense selection caused a different response with overdominant gene action from with partial dominance. When selection was random, effects of linkage and dominance on the coefficient of inbreeding were nonexistent; but when selection was by either phenotype or genotype, linkage and dominance became important. Joint effects between linkage, dominance, and selection are illustrated in specific simulated populations.     

Developmental variables and dominance rank in adolescent male mandrills (Mandrillus sphinx).

Previous research on semifree-ranging mandrills has shown that the degree of secondary sexual development differs among adult males. While some males are social, brightly colored, and have large testes and high levels of plasma testosterone, other males are peripheral or solitary, and lack fully developed secondary sexual features. In order to determine how these differences among males arise, and to investigate the influence of social factors, we examined the adolescent development of 13 semifree-ranging male mandrills of known age. Testicular volume began to increase markedly at 5.5 yr, and males began to develop secondary sexual adornments at the age of 6 yr. Males attained adult size and secondary sexual development at an average age of 9 yr. As males developed, they peripheralized, decreasing from 100% group-associated at 5 yr to 20% at 8 yr. At 9 yr some males reentered the social group and attained alpha rank, while others remained peripheral or solitary. Within this average development, there was marked variation among males in the timing of development. Adolescent males that were dominant for their age had higher testosterone levels, larger testes, and more advanced secondary sexual development than subordinate males. The implications of these findings are discussed in the light of differences that occur among adult males, male-male competition, and the evolution of secondary sexual adornments in this species.     

The large-male advantage in brown antechinuses: female choice, male dominance, and delayed male death

Male-biased dimorphism in body size is usually attributed tosexual selection acting on males, through either male competitionor female choice. Brown antechinuses (Antechinus stuartii) aresexually dimorphic in size, and heavier males are known to siremore offspring in the wild. We investigated four possible mechanismsthat might explain this large-male reproductive advantage. Wetested if there is a female preference for large males, a femalepreference for dominant males, if larger males compete moreeffectively for mates, and if there is a survival advantagefor large males during the mating season. We established nestinggroups of males in captivity and conducted mate choice trialsin which males from nesting groups either could or could notinteract. We assessed male dominance rank and recorded survivaltimes after mating. Females did not prefer larger males directly.The results suggest that the other three mechanisms of sexualselection tested account for the large-male advantage: largemales competed more successfully for mates, so were sociallydominant; females rejected subordinates (males they saw losingtwice in contests to previous mates); and dominant males survivedfor longer after their first mating. Females judged male rankbased on direct observation of male competitive interactionsat the time of mating and apparently could not distinguish rankfrom male scent. Effects of size and dominance on male reproductivesuccess are not confounded by age because male antechinusesare semelparous.     

布氏田鼠的社群气味选择和优势度的关系

化学信号中带有表明气味释放者社会等级等有关个体特征的信息（Halpin,1976）,许多来自于尿液、粪便以及一些腺体（如肛腺、胁腺）的分泌物中就携带有这些信号并可用于性别和个体识别,生理状况识别以及等级识别（Macdonaid,1985：Zhangeta1．,2005）。     

Female social dominance in two Eulemur species with different social organizations

Female social dominance is rare in mammals, but common in lemurs. We investigated social dominance in two Eulemur species; the polygynous crowned lemur (E. coronatus) and the monogamous red‐bellied lemur (E. rubriventer), using four and two social groups, respectively. We collected data on agonistic interactions and two types of affiliative behavior (grooming and maintaining spatial proximity). We used a combination of focal watches of individuals, instantaneous scan‐sampling of groups, and all‐occurrence of some behaviors in groups. We found that overall rates of agonistic interactions were higher in E. coronatus, and they also had more decided intersexual agonistic interactions than E. rubriventer. However, in both species the females won the vast majority of these agonistic interactions. E. coronatus females were groomed more often by males than vice versa, whereas no sex differences in grooming were observed in E. rubriventer. We found that males were responsible for maintaining spatial proximity in E. coronatus whereas in E. rubriventer, females were responsible. In one group of E. coronatus, the male was overweight and dominant to the female and this is the first observation of male dominance in a lemur species typically described as female dominant. We suggest that body weights in captivity be monitored for maintaining normal dominance relationships. Overall, agonistic behaviors were consistent with clear female social dominance in both E. coronatus and E. rubriventer. The affiliative behaviors also provided clear evidence for female dominance E. coronatus, but not for E. rubriventer. Zoo Biol 0: 1–14, 2007. © 2007 Wiley‐Liss, Inc.     

Masculine voices signal men's threat potential in forager and industrial societies

Humans and many non-human primates exhibit large sexual dimorphisms in vocalizations and vocal anatomy. In humans, same-sex competitors and potential mates attend to acoustic features of male vocalizations, but vocal masculinity especially increases perceptions of physical prowess. Yet, the information content of male vocalizations remains obscure. We therefore examined relationships between sexually dimorphic acoustic properties and men's threat potential. We first introduce a new measure of the structure of vocal formant frequencies, 'formant position' (Pf), which we show is more sexually dimorphic and more strongly related to height than is the most widely used measure of formant structure, 'formant dispersion', in both a US sample and a sample of Hadza foragers from Tanzania. We also show large sexual dimorphisms in the mean fundamental frequency (F0) and the within-utterance standard deviation in F0 (F0-s.d.) in both samples. We then explore relationships between these acoustic parameters and men's body size, strength, testosterone and physical aggressiveness. Each acoustic parameter was related to at least one measure of male threat potential. The most dimorphic parameters, F0 and Pf, were most strongly related to body size in both samples. In the US sample, F0 predicted testosterone levels, Pf predicted upper body strength and F0-s.d. predicted physical aggressiveness.     

Taxonomy and palaeobiology of the largest-ever marsupial, Diprotodon Owen, 1838 (Diprotodontidae, Marsupialia)

To determine accurately the rates of late Pleistocene megafaunal loss, it is fundamentally important to have accurate taxonomic information for every species. In Australia, accurate taxonomic information is lacking for several Pleistocene groups, including the largest marsupial ever to live, Diprotodon Owen, 1838. Diprotodon taxonomy has been complicated by early nomenclatural problems and by the occurrence of two distinct size classes of individuals that do not reflect an ontogenetic series. Traditionally, the two size classes have been regarded as separate species. However, a taxonomic investigation of large samples (> 1000 teeth) of Diprotodon material from several different fossil localities in Queensland, New South Wales, South Australia and Victoria suggests that there is little evidence for the discrimination of more than one morphospecies. Thus, Diprotodon is here considered a monotypic genus and the single morphospecies, D. optatum Owen, 1838 is considered to have been highly sexually dimorphic. By drawing analogy with extant sexually dimorphic megaherbivores and marsupials, the large form was probably male, and the small form was probably female. Diprotodon optatum probably moved in small, gender-segregated herds, and exhibited a polygynous breeding strategy. As a single morphospecies, D. optatum had a near-continental geographical distribution, similar to that of extant megaherbivores, possibly indicating its niche as a habitat generalist.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 153 , 389–417.