Comparisons of positive assortative mating and sexual selection models

A series of theoretical models of positive assortative mating and sexual selection are contrasted. It is established that for a dominant trait partial positive assortative mating generally implies some fixation, whereas sexual selection exhibits a unique globally stable polymorphism exhibiting Hardy-Weinberg proportions. The effects of monogamy against polygamy do not qualitatively alter the equilibrium outcomes, although the rate of evolutionary change is generally slowed with monogamy vis-à-vis polygamy. For sexual selection the influence of timing of random mating as against preferential mating causes no change in the equilibrium states, although the rates of convergence can be slowed if sexual selection occurs late in the breeding season. Under assortative mating the timing can alter the equilibrium outcomes. The amount of heterozygosity is always deficient in cases of assortative mating, but always exhibits Hardy-Weinberg ratios under a sexual selection mechanism. This suggests that observations consistent with Hardy-Weinberg equilibrium states cannot preclude ipso facto certain forms of selection forces, including mating patterns and some natural selection structures.     

Ultraviolet sexual dimorphism and assortative mating in blue tits

In spite of strong evidence for viability-based sexual selection and sex ratio adjustments, the blue tit, Parus caeruleus, is regarded as nearly sexually monomorphic and no epigamic signals have been found. The plumage coloration has not, however, been studied in relation to bird vision, which extends to the UV-A waveband (320 to 400 nm). Using molecular sex determination and UV/VIS spectrometry, we report here that blue tits are sexually dichromatic in UV/blue spectral purity (chroma) of the brilliant crown patch. It is displayed in courtship by horizontal posturing and erected nape feathers. A previously undescribed sexual dimorphism in crown size (controlling for body size) further supports its role as an epigamic ornament. Against grey-brown leaf litter and bark during pair formation in early spring, but also against green vegetation, UV contributes strongly to conspicuousness and sexual dimorphism. This should be further enhanced by the UV/bluish early morning skylight (''woodland shade'') in which blue tits display. Among 18 breeding pairs, there was strong assortative mating with respect to UV chroma, but not size, of the crown ornament. We conclude that blue tits are markedly sex dimorphic in their own visual world, and that UV/violet coloration probably plays a role in blue tit mate acquisition.     

Two-trait selection response with marker-based assortative mating

 Marker-based assortative mating (MAM) – the mating of individuals that have similar genotypes at random marker loci – can increase selection response for a single trait by 3–8% over random mating (RM). Genetic gain is usually desired for multiple traits rather than for a single trait. My objectives in this study were to (1) compare MAM, phenotypic assortative mating (PAM), and RM of selected individuals for improving two traits and (2) determine when MAM will be most useful for improving two traits. I simulated 20 generations of selecting 32 out of 200 individuals in an F₂ population. The individuals were selected based on an index (SI) of two traits and were intermated by MAM, PAM, or RM. I studied eight genetic models that differed in three contrasts: (1) weight, number of quantitative trait loci (QTL), and heritability (h ²) for each trait; (2) linkage of QTL for each trait; and (3) trait means of the inbred parents of the F₂. For SI and the two component traits, MAM increased short-term selection response by 5–8% in six out of the eight genetic models. The MAM procedure was least effective in two genetic models, wherein the QTL for one trait were unlinked to the QTL for the other trait and the parents of the F₂ had divergent means for each trait. The loss of QTL heterozygosity was much greater with MAM than with PAM or RM. Consequently, the advantage of MAM over RM dissipated after 5–7 generations. Differences were small between selection responses with PAM and RM. The MAM procedure can enhance short-term selection response for two traits when selection is not stringent, h ² is low, and the means of the parents of the F₂ are equal for each trait. Received: 10 June 1998 / Accepted: 5 August 1998     

Polymorphic equilibria with assortative mating and selection in subdivided populations

Two modes of assortative mating, partial selfing and assorting by phenotypic classes, are studied in a subdivided population. Differential viability is allowed and the selection intensities and assorting tendencies are permitted to vary among the habitats. There exists a symmetric polymorphism; we delimit its level of heterozygosity and stability nature (dependent on the selection intensities and assorting propensities). This complements studies of the fixation states and thereby provides further insight into the global equilibrium structure in subdivided populations. Circumstances are given where the fixation states and symmetric polymorphism comprise the global equilibrium structure. Examples are also given where migration engenders stable polymorphic equilibria and stable polymorphic equilibrium cycles which are absent in single demes without migration.     

Substrate selection and assortative mating in Gammarus pulex L.

We reject the published proposition that assortative mating for size in Gammarus is brought about purely as a result of spatial heterogeneity in substrate particle size. We confirm that, in general, large Gammarus prefer substrates with larger particle size than small Gammarus. However, in artificial streams in the laboratory, in choice experiments Gammarus males and females showed positive assortative mating for size in each of three substrates differing in mean particle size. We propose an explanation for assortative mating based on sexual selection.     

Models of cultural niche construction with selection and assortative mating

Niche construction is a process through which organisms modify their environment and, as a result, alter the selection pressures on themselves and other species. In cultural niche construction, one or more cultural traits can influence the evolution of other cultural or biological traits by affecting the social environment in which the latter traits may evolve. Cultural niche construction may include either gene-culture or culture-culture interactions. Here we develop a model of this process and suggest some applications of this model. We examine the interactions between cultural transmission, selection, and assorting, paying particular attention to the complexities that arise when selection and assorting are both present, in which case stable polymorphisms of all cultural phenotypes are possible. We compare our model to a recent model for the joint evolution of religion and fertility and discuss other potential applications of cultural niche construction theory, including the evolution and maintenance of large-scale human conflict and the relationship between sex ratio bias and marriage customs. The evolutionary framework we introduce begins to address complexities that arise in the quantitative analysis of multiple interacting cultural traits.     

Precopulatory mating behavior and sexual dimorphism in the amphipod Crustacea

Accounts in the literature of precopulatory mate-guarding in gammaridean amphipods are that males use one of two strategies for mating: either they mate-guard by carrying or attending their mates until they are ready to molt and be fertilized, or they do not guard, instead searching benthically or swarming pelagically at the time that females are ready to molt. Mate-guarding by carrying has been documented for species of the superfamilies Gammaroidea, Talitroidea, and Hadzioidea. Mate-guarding by attending has been found in the more sedentary Corophioidea and Caprellidea. Non-mate-guarders that search pelagically are species of Ampeliscoidea, Lysianassoidea, Phoxocephaloidea, Oedicerotoidea, and Pontoporeioidea. Non-mate-guarders that mate-search benthically are species of Eusiroidea, Crangonyctoidea, and Haustorioidea. Mate-guarding and non-mate-guarding males develop different secondary sex characters at maturity. Mate-guarding males have enhancements for fighting and signalling. These alterations are more elaborate in males that attend their mates than in males that carry their mates. Non-mate-guarders that search pelagically develop enhancements for swimming and sensing. Non-mate-guarders that remain benthic exhibit little change at maturity. Most mate-guarding males develop their secondary sexual characters over several molts and mate over more than one instar. Pelagic mate-searchers develop their secondary sexual characters at the last molt and mating is confined to the last instar. Females of most mate-guarding species are iteroparous, while fewer than half of non-mate-guarding species are so. It is hypothesized that mate-guarding arose more than once in the evolutionary history of amphipod Crustacea.     

Effect of assortative mating on genetic change due to selection

Summary Two mathematical models (A and B) were used to study joint effects of selection and assortative mating on genetic change. Computer simulation was used to verify and extend the results. In each model, the genotype was additive with equal effects at each of N loci and the environmental distribution was N(0, ²). In Model A, each locus had two alleles; in Model B, allelic effects at each locus followed a normal distribution. Using Model A, genetic change with assortative or random mating of selected parents was evaluated for combinations of number of loci (N = 1, 2, 3), heritability in base population (H^[0] = 0.2, 0.5, 0.8), allelic frequency in base population (p = 0.1, 0.5), and proportion selected ( = 0.20, 0.85). Using Model B, genetic change with or without assortative mating was calculated for combinations of N (1, 2, 3, 5, 10, 100, H^[0] (0.2, 0.5, 0.8) and (0.20, 0.85). Response to selection under both mating systems in a finite population was estimated using Model A from 200 replications of a computer simulation; this was done for all combinations of N (1,2, 3, 5, 10) and (0.20, 0.85), with H^[0] = 0.5 and p = 0.1. Results obtained with both models indicate that the effect of assortative mating on genetic change increases with H^[0] and , and decreases with p. With Model A, the relationship between N and the effect of assortative mating on genetic change was not clear; with Model B, however, the advantage of assortative over random mating increased with N, as expected. Simulation results were in agreement with theory of Model A. This study indicates that selection with assortative mating can have a sizable (10 to 20%) long-term advantage over selection with random mating of parents when H^[0] is high, p is low and is large.     

The joint effects of selection and assortative mating on multiple polygenic characters

Communicated by A. Robertson     

Rules for assortative mating in relation to selection for linear merit functions

Summary This study examined how assortative mating (without selection) based on linear combinations of two traits could be used to change genetic parameters so as to increase efficiency of selection. The efficiency of the Smith-Hazel index for improvement of multiple traits is a function of phenotypic and genetic variances and covariances, and of the relative economic values of the traits involved. Assortative mating is known to change genetic variances and covariances. Recursive formulae were derived to obtain these variances and covariances after t generations of assortative mating on linear combinations (mating rules) of phenotypic values for two traits, with a given correlation between mates. Selection efficiency after t generations of assortative mating without selection was expressed as a function of random mating genetic parameters, economic values, the mating rule, and the correlation between mates. Selection efficiency was maximized with respect to the coefficients in the mating rule. Because the objective function was nonlinear, a computer routine was used for maximizing it. Two cases were considered. When random mating heritabilities for the two traits were h_X²=0.25 and h_Y²=0.50, the genetic correlation r_XY=-0.60, and the economic values were a_X=3 and a_Y=1, continued assortative mating based on the optimal mating rule for 31 generations (with a correlation of 0.80 between mates) increased selection efficiency by 29%. Heritabilities changed to 0.38 and 0.66, respectively, and the genetic correlation became – 0.79. When h_X²=0.60, h_Y²=0.60, r_XY=– 0.20, a₁=1 and a₂=1, 36 generations of continued assortative mating with the optimal mating rule increased the efficiency of selection by 17%, heritabilities became h_X²= h_Y²=0.71, and the genetic correlation changed to 0.25. Only three generations of assortative mating were required to change the sign of the genetic correlation.     

Loading constraints,body size and mating preference in Gammarus species

Size relationships among pairing Gammarus were examined with reference to two hypotheses for sexual size dimorphism and assortative mating among aquatic Peracarida (Crustacea). The sizes of pairing and non-pairing animals were compared in different experimental conditions where the size of one or both sexes was controlled. The experimental results present a complex picture which suggests that both sexual selection and loading constraints are likely to play a role in determining mating decisions in these animals.Department of Adult & Continuing Education, University of Durham     

Analysis of partial assortative mating and sexual selection models for a polygamous species involving a trait based on levels of heterozygosity

The consequences of preferential mating in the presence of partial assortative and sexual selection mechanisms are ascertained for a two-allele one-locus trait involving two phenotype classes C₁ = {all homozygotes} and C₂ = {heterozygotes}. Relevant biological cases may include Burley (1977, Proc. Nat. Acad. Sci. USA74, 3476–3479), Wilbur et al. (1978, Evolution32, 264–270), and Singh and Zouros (1978, Evolution32, 342–353). When the preference rate for the heterozygote exceeds that for homozygotes, it is established that the unique stable state is the central Hardy-Weinberg equilibrium. The rate of approach is faster with sexual selection than for the corresponding model of assortative mating. When the preference rates favor the homozygotes then in this symmetric model of sexual selection two asymmetric Hardy-Weinberg polymorphisms can evolve, and which succeeds depends on initial conditions. The models are also analyzed with natural selection acting on phenotypes superimposed on assortative mating. In this case we can have up to three coexisting stable states involving both fixation alternatives and a central polymorphism. The corresponding model with sexual selection maintains either the central equilibrium as in assortative mating or two asymmetric polymorphic equilibria.     

Ancestry-related assortative mating in Latino populations

Background

While spouse correlations have been documented for numerous traits, no prior studies have assessed assortative mating for genetic ancestry in admixed populations.

Results

Using 104 ancestry informative markers, we examined spouse correlations in genetic ancestry for Mexican spouse pairs recruited from Mexico City and the San Francisco Bay Area, and Puerto Rican spouse pairs recruited from Puerto Rico and New York City. In the Mexican pairs, we found strong spouse correlations for European and Native American ancestry, but no correlation in African ancestry. In the Puerto Rican pairs, we found significant spouse correlations for African ancestry and European ancestry but not Native American ancestry. Correlations were not attributable to variation in socioeconomic status or geographic heterogeneity. Past evidence of spouse correlation was also seen in the strong evidence of linkage disequilibrium between unlinked markers, which was accounted for in regression analysis by ancestral allele frequency difference at the pair of markers (European versus Native American for Mexicans, European versus African for Puerto Ricans). We also observed an excess of homozygosity at individual markers within the spouses, but this provided weaker evidence, as expected, of spouse correlation. Ancestry variance is predicted to decline in each generation, but less so under assortative mating. We used the current observed variances of ancestry to infer even stronger patterns of spouse ancestry correlation in previous generations.

Conclusions

Assortative mating related to genetic ancestry persists in Latino populations to the current day, and has impacted on the genomic structure in these populations.     

Reinforcement and divergence under assortative mating

Traits that cause assortative mating such as the flowering time in plants and body size in animals can produce reproductive isolation between hybridizing populations. Can selection against unfit hybrids cause two populations to diverge in their mean values for these kinds of traits? Here I present a haploid analytical model of one population that receives gene flow from another. The partial pre-zygotic isolation between the two populations is caused by assortative mating for a trait that is influenced by any number of genes with additive effects. The post-zygotic isolation is caused by selection against genetic incompatibilities that can involve any form of selection on individual genes and gene combinations (epistasis). The analysis assumes that the introgression rate and selection coefficients are small. The results show that the assortment trait mean will not diverge from the immigrants unless there is direct selection on the trait favouring it to do so or there are genes of very large effect. The amount of divergence at equilibrium is determined by a balance between direct selection on the assortment trait and introgression from the other population. Additional selection against hybrid genetic incompatibilities reduces the effective migration rate and allows greater divergence. The role of assortment in speciation is discussed in the light of these results.