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.     

Frequency-dependent selection and the evolution of assortative mating

A long-standing goal in evolutionary biology is to identify the conditions that promote the evolution of reproductive isolation and speciation. The factors promoting sympatric speciation have been of particular interest, both because it is notoriously difficult to prove empirically and because theoretical models have generated conflicting results, depending on the assumptions made. Here, we analyze the conditions under which selection favors the evolution of assortative mating, thereby reducing gene flow between sympatric groups, using a general model of selection, which allows fitness to be frequency dependent. Our analytical results are based on a two-locus diploid model, with one locus altering the trait under selection and the other locus controlling the strength of assortment (a "one-allele" model). Examining both equilibrium and nonequilibrium scenarios, we demonstrate that whenever heterozygotes are less fit, on average, than homozygotes at the trait locus, indirect selection for assortative mating is generated. While costs of assortative mating hinder the evolution of reproductive isolation, they do not prevent it unless they are sufficiently great. Assortative mating that arises because individuals mate within groups (formed in time or space) is most conducive to the evolution of complete assortative mating from random mating. Assortative mating based on female preferences is more restrictive, because the resulting sexual selection can lead to loss of the trait polymorphism and cause the relative fitness of heterozygotes to rise above homozygotes, eliminating the force favoring assortment. When assortative mating is already prevalent, however, sexual selection can itself cause low heterozygous fitness, promoting the evolution of complete reproductive isolation (akin to "reinforcement") regardless of the form of natural selection.     

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     

Sexual selection and assortative mating: an experimental test

Mate choice and mate competition can both influence the evolution of sexual isolation between populations. Assortative mating may arise if traits and preferences diverge in step, and, alternatively, mate competition may counteract mating preferences and decrease assortative mating. Here, we examine potential assortative mating between populations of Drosophila pseudoobscura that have experimentally evolved under either increased (‘polyandry’) or decreased (‘monogamy’) sexual selection intensity for 100 generations. These populations have evolved differences in numerous traits, including a male signal and female preference traits. We use a two males: one female design, allowing both mate choice and competition to influence mating outcomes, to test for assortative mating between our populations. Mating latency shows subtle effects of male and female interactions, with females from the monogamous populations appearing reluctant to mate with males from the polyandrous populations. However, males from the polyandrous populations have a significantly higher probability of mating regardless of the female's population. Our results suggest that if populations differ in the intensity of sexual selection, effects on mate competition may overcome mate choice.     

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.     

Intraguild interactions promote assortative mating and affect sexual attractiveness in a phytophagous fly

Changes in acoustic and substrate-borne sexual signals in phytophagous insects associated with host plant shifts are known to have the potential to promote assortative mating, reproductive isolation and speciation. In this article, we ask whether the switch between pure herbivory and intraguild predation (IGP), which is common amongst phytophagous insects, has similar potential. Male flies in the genus Lipara (Diptera: Chloropidae) search for females by vibrating reed stems and waiting for a reply. By kleptoparasitizing other phytophagous species in the genus (a form of IGP), Lipara rufitarsis can increase its nonsexual fitness considerably. We looked at the impact of IGP on the timing of hatching, body size and attractiveness of male calls in L. rufitarsis . L. rufitarsis males that had engaged in IGP hatched significantly earlier than purely phytophagous flies and were significantly larger, but their calls were less likely to elicit responses from females during playback experiments. We conclude that, although behavioural observations of females provided no evidence of 'like preferring like', changes in phenology associated with IGP are likely to promote assortative mating in this system. The general preference of females for the calls of smaller males is a phenomenon worthy of further study: it may have no adaptive significance, or it may indicate that mating with large males is associated with a fitness cost.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 171–180.     

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

Communicated by A. Robertson     

Continuous-genotype models and assortative mating

Feldman and Cavalli-Sforza (1979a,b) have argued that the convergence properties of classical models of assortative mating are not known, and that these models involve arbitrary assumptions which assume rather than derive the achievement of equilibrium. A careful consideration of all models shows that the classical models are well defined and seem to achieve their equilibria. The model used by Feldman and Cavalli-Sforza involves an arbitrary assumption. Consideration of the models of Wright, Fisher, Bulmer, and Lande in the context of assortative mating or of selection versus mutation shows that these models are consistent with each other. The treatment of the balance between mutation and normalizing selection by Cavalli-Sforza and Feldman comes to conclusions sharply different from those of other authors, apparently as a result of this same arbitrary assumption.     

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 _X ² =0.25 and h _Y ² =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 _X ² =0.60, h _Y ² =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 _X ² = h _Y ² =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.     

Estimating sexual selection and sexual isolation effects from mating frequencies

Abstract.— Sexual selection (defined as the change in genotypic or phenotypic frequencies of mated versus total population frequencies) and sexual isolation (defined as the deviation from random mating in mated individuals) show different evolutionary consequences and partially confounded causes. Traditionally, the cross-product estimator has been used to quantify sexual selection, whereas a variety of indexes, such as Yule V , Yule Q, YA , joint I , and others have been used to quantify sexual isolation. Because the two types of estimators use different scales, the effects of both processes cannot be monitored simultaneously. We describe three new related statistics that quantify both sexual selection ( PSS ) and sexual isolation ( PSI ) effects for every mating pair combination in polymorphic traits, as well as measure their combined effects ( PTI = PSI X PSS ). The new statistics have the advantage of providing information on every mating pair combination, quantifying the effects of sexual selection and isolation in the same units, and detecting asymmetry in sexual isolation. The ability of the new statistics to ascertain the biological causes of sexual selection and sexual isolation are investigated under different models involving distinct marginal frequencies, mate propensity, and mate choice coefficients. We also studied the use of classical isolation indexes applied on PSI coefficients, instead of on raw data. The use of the classical indexes applied to PSI coefficients considerably reduces the statistical bias of the estimates, revealing the good estimation properties of the new statistics.     

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