Quantitative genetics and sex-specific selection on sexually dimorphic traits in bighorn sheep

Sexual conflict at loci influencing traits shared between the sexes occurs when sex-specific selection pressures are antagonistic relative to the genetic correlation between the sexes. To assess whether there is sexual conflict over shared traits, we estimated heritability and intersexual genetic correlations for highly sexually dimorphic traits (horn volume and body mass) in a wild population of bighorn sheep (Ovis canadensis) and quantified sex-specific selection using estimates of longevity and lifetime reproductive success. Body mass and horn volume showed significant additive genetic variance in both sexes, and intersexual genetic correlations were 0.24+/-0.28 for horn volume and 0.63+/-0.30 for body mass. For horn volume, selection coefficients did not significantly differ from zero in either sex. For body weight, selection coefficients were positive in females but did not differ from zero in males. The absence of detectable sexually antagonistic selection suggests that currently there are no sexual conflicts at loci influencing horn volume and body mass.     

Divergent selection on,but no genetic conflict over,female and male timing and rate of reproduction in a human population

The sexes often have different phenotypic optima for important life-history traits, and because of a largely shared genome this can lead to a conflict over trait expression. In mammals, the obligate costs of reproduction are higher for females, making reproductive timing and rate especially liable to conflict between the sexes. While studies from wild vertebrates support such sexual conflict, it remains unexplored in humans. We used a pedigreed human population from preindustrial Finland to estimate sexual conflict over age at first and last reproduction, reproductive lifespan and reproductive rate. We found that the phenotypic selection gradients differed between the sexes. We next established significant heritabilities in both sexes for all traits. All traits, except reproductive rate, showed strongly positive intersexual genetic correlations and were strongly genetically correlated with fitness in both sexes. Moreover, the genetic correlations with fitness were almost identical in men and women. For reproductive rate, the intersexual correlation and the correlation with fitness were weaker but again similar between the sexes. Thus, in this population, an apparent sexual conflict at the phenotypic level did not reflect an underlying genetic conflict over the studied reproductive traits. These findings emphasize the need for incorporating genetic perspectives into studies of human life-history evolution.     

Natural selection and non-metric traits in skeletal populations

The hypothesis of this article is that the frequency of non-metric traits varies at different reproductive periods because of differential mortality associated with non-metric traits. 450 male and 203 female crania were examined. Mortality in the reproduction period is connected with the elimination of individuals with certain variants of non-metric traits. It seems that key genes which perform the stimulating function in the appearance of traits in a given category of formation are under selective pressure. It is assumed that natural selection acts both on the morphology of organisms and on developmental process through environmental conditions. Selection not only optimises the fitness of morphological features but also the processes leading to the formation of features during ontogeny. Both these mechanisms influence the interpopulational variability of non-metric traits.     

Natural selection and the traits of individual organisms

It has recently been argued that origin essentialism regarding individual organisms entails that natural selection does not explain why individual organisms have the traits that they do. This paper defends this and related theses against Mohan Matthens recent objections.     

Responses in flight to selection on wing length in non-migratory milkweed bugs, Oncopeltus fasciatus

A sample of nonmigratory Oncopeltus fasciatus (Dallas) (Hemiptera: Lygaeidae) from Puerto Rico was subjected to bi-directional selection for wing length for 20 generations. Response to selection was strong in both directions. In the 8th and 14th generations samples were drawn from the selected lines and tested for tethered flight. There was little flight observed and no correlation between wing length and flight duration. There was also no correlation between wing length and fecundity. These results contrast with results from a migratory population from Iowa, U.S.A. where there were positive genetic correlations between wing length and both flight and fecundity. A migration-life history syndrome was thus apparent in the migratory Iowa population, but was not present in the non-migratory population from Puerto Rico.

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Résumé L'étude de la relation entre la longueur de l'aile, le vol et la fécondité a été faite sur une population sédentaire d'O. fasciatus de Porto Rico; les résultat ont été comparés à ceux d'une population migratrice de L'Iowa, U.S.A. Les punaises de Porto Rico ont été sélectionnées pour leurs ailes longues ou countes pendant 20 générations, en utilisant des lignées répétitives. Une forte réponse a été obtenue par rapport aux témoines des lignées non sélectionnées. Aux 8ème et 14ème générations de sélection, des punaises ont été prélevées pour examen de leur vol entravé. Très peu de vols ont été obtenus et il n'y avait aucune corrélation entre la durée du vol et la longueur de l'aile. Parallèlement, la fécondité pendant les 5 premiers jours de reproduction a été mesurée, et aucune corrélation n'a été mise en évidence, mais il n'y avait pas non plus de corrélation entre longueur de l'aile et fécondité. Ces résultats différent de ceux de la population migratrice de l'Iowa, pout laquelle il y avait une corrélation génétique positive entre longueur de l'aile, vol et fécondité. Chez les punaises de l'Iowa, il y aurait ainsi un syndrome biologique de migration avec des ailes longues, une fécondité élevée précoce et des vols de longue durée. II n'y avait pas de manifestation de ce syndrome chez les populations sédentaires de Porto Rico. Les O. fasciatus migrateurs et sédentaires différent évidemment par la base génétique de leur biologie.

     

Natural selection and the genetics of adaptation in threespine stickleback

Growing knowledge of the molecular basis of adaptation in wild populations is expanding the study of natural selection. We summarize ongoing efforts to infer three aspects of natural selection—mechanism, form and history—from the genetics of adaptive evolution in threespine stickleback that colonized freshwater after the last ice age. We tested a mechanism of selection for reduced bony armour in freshwater by tracking genotype and allele frequency changes at an underlying major locus (Ectodysplasin) in transplanted stickleback populations. We inferred disruptive selection on genotypes at the same locus in a population polymorphic for bony armour. Finally, we compared the distribution of phenotypic effect sizes of genes underlying changes in body shape with that predicted by models of adaptive peak shifts following colonization of freshwater. Studies of the effects of selection on genes complement efforts to identify the molecular basis of adaptive differences, and improve our understanding of phenotypic evolution.     

Natural selection on body size traits in a long‐lived bird,the snow petrel Pagodroma nivea

I consider the possibility of selection favouring large body size in a population of snow petrels (Pagodroma nivea), a long‐lived seabird species. I measured natural selection on body size traits in a population from 1987 to 1998. There was evidence of selection on body size associated with fecundity and survival. Directional selection on bill length and stabilizing selection on tarsus length associated with reproductive success were detected among males. Selection associated with survival favoured males with longer bills. However, selection was weak in all cases. No evidence of selection acting on female body size traits was detected. Offspring–parents regression suggested that bill length and tarsus length were heritable. Although I was able to identify the targets of selection in this population, I could not demonstrate the ecological implications of both tarsus length and bill length variation. The selection on male, but not on female, body size traits suggests factors such as intrasexual competition for nests and/or mates rather than factors such as feeding efficiency as mechanisms of selection on bill size.     

Antagonistic and reinforcing pleiotropy: a study of differences in development time in wing dimorphic insects

Morphological dimorphisms are found in many different taxa. Wing dimorphism in insects, in which some individuals possess wings and associated flight muscles and are thus volant while others lack a functional flight apparatus and are thus flightless, is a typical example of such types of dimorphisms. It has been extensively studied and such studies have demonstrated that the volant form, although possessing the advantage of flight capability, suffers a fitness cost in a delay in the onset of reproduction after emergence into the adult form and a reduced fecundity. Previous comparative analyses have suggested that there is no consistent trend for development time (hatching to adult) to differ between the two morphs. The present study analyses the phenotypic and genetic correlations between development time and wing morph in the cricket Gryllus firmus. It is shown that the macropterous (volant) morph develops faster than the micropterous (flightless morph). This trade-off is manifested at both thephenotypic and genetic level. Further, a comparative analysis shows that the same phenotypic trade-off is generally found in other Orthopteran species so far studied, but in other orders the micropterous morph develops faster. Provided that the phenotypic trade-off is genetically based, in the Orthoptera the fitness advantage of the earlier onset of reproduction in micropterous females is offset by the extended development time (antagonistic pleiotropy). However, in other orders there is reinforcing pleiotropy in that the micropterous females develop faster and reproduce sooner than the macropterous morph. These results highlight the complexity of fitness interactions and the need to study a phenomenon across several taxa.     

Natural selection hampers divergence of reproductive traits in a seed beetle

Speciation is thought to often result from indirect selection for reproductive isolation. This will occur when reproductive traits that cause reproductive isolation evolve (i) as a by‐product of natural selection on traits with which they are genetically correlated or (ii) as an indirect result of diversifying sexual selection. Here, we use experimental evolution to study the degree of divergent evolution of reproductive traits by manipulating the intensity of natural and sexual selection in replicated selection lines of seed beetles. Following 40 generations of selection, we assayed the degree of divergent evolution of reproductive traits between replicate selection lines experiencing the same selection regime. The evolution of reproductive traits was significantly divergent across selection lines within treatments. The evolution of reproductive traits was both slower and, more importantly, significantly less divergent among lines experiencing stronger directional natural selection. This suggests that reproductive traits did not evolve as an indirect by‐product of adaptation. We discuss several ways in which natural selection may hamper divergent evolution among allopatric populations.     

Predicting evolution of floral traits associated with mating system in a natural plant population

Evolution of floral traits requires that they are heritable, that they affect fitness, and that they are not constrained by genetic correlations. These prerequisites have only rarely been examined in natural populations. For Mimulus guttatus, we found by using the Riska-method that corolla width, anther length, ovary length and number of red dots on the corolla were heritable in a natural population. Seed production (maternal fitness) was directly positively affected by corolla width and anther size, and indirectly so by ovary length and number of red dots on the corolla. The siring success (paternal fitness), as estimated from allozyme data, was directly negatively affected by anther-stigma separation, and indirectly so by the corolla length-width ratio. Genetic correlations, estimated with the Lynch-method, were positive between floral size measures. We predict that larger flowers with larger reproductive organs, which generally favour outcrossing, will evolve in this natural population of M. guttatus.     

Natural selection. III. Selection versus transmission and the levels of selection

George Williams defined an evolutionary unit as hereditary information for which the selection bias between competing units dominates the informational decay caused by imperfect transmission. In this article, I extend Williams' approach to show that the ratio of selection bias to transmission bias provides a unifying framework for diverse biological problems. Specific examples include Haldane and Lande's mutation-selection balance, Eigen's error threshold and quasispecies, Van Valen's clade selection, Price's multilevel formulation of group selection, Szathmáry and Demeter's evolutionary origin of primitive cells, Levin and Bull's short-sighted evolution of HIV virulence, Frank's timescale analysis of microbial metabolism and Maynard Smith and Szathmáry's major transitions in evolution. The insights from these diverse applications lead to a deeper understanding of kin selection, group selection, multilevel evolutionary analysis and the philosophical problems of evolutionary units and individuality.     

Quantitative genetics of sexually dimorphic traits and capture of genetic variance by a sexually-selected condition-dependent ornament in red junglefowl (Gallus gallus)

We studied the quantitative genetics of sexually selected traits in a captive population of red junglefowl (Gallus gallus L.) using a multi-generational 'animal model' approach. We found significant heritability of mass, tarsus length (both strongly sexually dimorphic), residual mass, and male comb (a fleshy head ornament) length. Residual mass has a genetic correlation between the sexes smaller than unity and so could show partially independent responses to selection in the two sexes. In males, tarsus length and mass were not genetically correlated, and this produced a negative genetic correlation between tarsus length and residual mass. The male red junglefowl's comb, an ornament influencing female choice, is highly condition dependent. We show that expression of this ornament is heritable, however, and shows strong genetic correlation with a condition index, residual mass. Because residual mass is partly influenced by various aspects of condition, it appears that comb size has 'captured' genetic variability in condition.     

Predictions of response to selection caused by angling in a wild population of Atlantic salmon (Salmo salar)

1. Recreational angling activities in wild populations of Atlantic salmon may induce a selection pressure towards a reduction in body size and length if the angling season coincides with the return of the largest sea age fish class. 2. Using estimates of heritability for growth traits and estimates of the selection pressure from angling operating on growth, we predicted the response to selection expected to occur in a wild population of Atlantic salmon. 3. The dataset used here comprised individuals from two consecutive generations (parents and offspring) from the River Bidasoa (NW Spain). Offspring were assigned to parents using six highly polymorphic microsatellite loci. Use of restricted maximum likelihood methodology and the animal model allowed us to estimate the heritability for body length and body weight as well as their genetic correlation. 4. Estimated heritabilities (0.32 ± 0.12 for length and 0.32 ± 0.11 for weight) and selection pressure caused by angling were used to obtain predictions of response to selection because of angling. Our results suggested a decline of 1.9 mm in body length and 103.3 g in body weight per generation because of angling pressure. 5. The results derived from this study suggest that the angling season should be annually delayed in order to avoid selective angling of the multi‐year class and further reductions in body weight and length.     

Associations between environmental stress, selection history, and quantitative genetic variation in Drosophila melanogaster

Stressful environments may increase quantitative genetic variation in populations by promoting the expression of genetic variation that has not previously been eliminated or canalized by natural selection. This “selection history” hypothesis predicts that novel stressors will increase quantitative genetic variation, and that the magnitude of this effect will decrease following continued stress exposure. We tested these predictions using Drosophila melanogaster and sternopleural bristle number as a model system. In particular, we examined the effect of high temperature stress (31°C) on quantitative genetic variation before and after our study population had been reared at 31°C for 15 generations. High temperature stress was found to increase both additive genetic variance and heritability, but contrary to the selection history hypothesis prediction, the magnitude of this effect significantly increased after the study population had been reared for 15 generations under high temperature stress. These results demonstrate that high temperature stress increases quantitative genetic variation for bristle number, but do not support the selection history hypothesis as an explanation for this effect.     

Heritability estimates of life history traits in small white butterflyPieris rapae crucivora

Heritabilities and genetic correlations of life history characters (pupal weight, age-specific fecundities, and egg weight) of small white butterfly Pieris rapae crucivora are estimated by a quantitative genetic method (sib analysis). The results indicate moderate or high heritabilities and a largely negative genetic correaltion in age-specific fecundities.     

Using human demographic history to infer natural selection reveals contrasting patterns on different families of immune genes

Detecting regions of the human genome that are, or have been, influenced by natural selection remains an important goal for geneticists. Many methods are used to infer selection, but there is a general reliance on an accurate understanding of how mutation and recombination events are distributed, and the well-known link between these processes and their evolutionary transience introduces uncertainty into inferences. Here, we present and apply two new, independent approaches; one based on single nucleotide polymorphisms (SNPs) that exploits geographical patterns in how humans lost variability as we colonized the world, the other based on the relationship between microsatellite repeat number and heterozygosity. We show that the two methods give concordant results. Of these, the SNP-based method is both widely applicable and detects selection over a well-defined time interval, the last 50 000 years. Analysis of all human genes by their Gene Ontology codes reveals how accelerated and decelerated loss of variability are both preferentially associated with immune genes. Applied to 168 immune genes used as the focus of a previous study, we show that members of the same gene family tend to yield similar indices of selection, even when located on different chromosomes. We hope our approach will provide a useful tool with which to infer where selection has acted to shape the human genome.     

Heritability of corticosterone response and changes in life history traits during selection in the zebra finch

Vertebrates respond to environmental stressors through the neuro-endocrine stress response, which involves the production of glucocorticoids. We have selected independent, duplicate divergent lines of zebra finches for high, low and control corticosterone responses to a mild stressor. This experiment has shown that over the first four generations, the high lines have demonstrated a significant realized heritability of about 20%. However, the low lines have apparently not changed significantly from controls. This asymmetry in response is potentially because of the fact that all birds appear to be showing increased adaptation to the environment in which they are housed, with significant declines in corticosterone response in control lines as well as low lines. Despite the existence of two- to threefold difference in mean corticosterone titre between high and low lines, there were no observed differences in testosterone titre in adult male birds from the different groups. In addition, there were no consistent, significant differences between the lines in any of the life history variables measured--number of eggs laid per clutch, number of clutches or broods produced per pair, number of fledglings produced per breeding attempt, nor in any of egg, nestling and fledgling mortality. These results highlight the fact that the mechanisms that underlie variation in the avian physiological system can be modified to respond to differences between environments through selection. This adds an additional level of flexibility to the avian physiological system, which will allow it to respond to environmental circumstances.     

Sex-specific genetic variances in life-history and morphological traits of the seed beetle Callosobruchus maculatus

Knowledge of heritability and genetic correlations are of central importance in the study of adaptive trait evolution and genetic constraints. We use a paternal half-sib-full-sib breeding design to investigate the genetic architecture of three life-history and morphological traits in the seed beetle, Callosobruchus maculatus. Heritability was significant for all traits under observation and genetic correlations between traits (r(A)) were low. Interestingly, we found substantial sex-specific genetic effects and low genetic correlations between sexes (r(MF)) in traits that are only moderately (weight at emergence) to slightly (longevity) sexually dimorphic. Furthermore, we found an increased sire ([Formula: see text]) compared to dam ([Formula: see text]) variance component within trait and sex. Our results highlight that the genetic architecture even of the same trait should not be assumed to be the same for males and females. Furthermore, it raises the issue of the presence of unnoticed environmental effects that may inflate estimates of heritability. Overall, our study stresses the fact that estimates of quantitative genetic parameters are not only population, time, environment, but also sex specific. Thus, extrapolation between sexes and studies should be treated with caution.