Levels Of Additive Genetic Variance Research Articles

Quantitative genetic insights into the coevolutionary dynamics of male and female genitalia

The spectacular variability that typically characterizes male genital traits has largely been attributed to the role of sexual selection. Among the evolutionary mechanisms proposed to account for this diversity, two processes in particular have generated considerable interest. On the one hand, females may exploit postcopulatory mechanisms of selection to favour males with preferred genital traits (cryptic female choice; CFC), while on the other hand females may evolve structures or behaviours that mitigate the direct costs imposed by male genitalia (sexual conflict; SC). A critical but rarely explored assumption underlying both processes is that male and female reproductive traits coevolve, either via the classic Fisherian model of preference-trait coevolution (CFC) or through sexually antagonistic selection (SC). Here, we provide evidence for this prediction in the guppy (Poecilia reticulata), a polyandrous livebearing fish in which males transfer sperm internally to females via consensual and forced matings. Our results from a paternal half-sibling breeding design reveal substantial levels of additive genetic variation underlying male genital size and morphology-two traits known to predict mating success during non-consensual matings. Our subsequent finding that physically interacting female genital traits exhibit corresponding levels of genetic (co)variation reveals the potential intersexual coevolutionary dynamics of male and female genitalia, thereby fulfilling a fundamental assumption underlying CFC and SC theory.

Screening Corymbia populations for resistance to Puccinia psidii

The exotic rust pathogen Puccinia psidii is now widespread along the east coast of Australia from temperate Victoria to tropical far north Queensland, with a current host range exceeding 200 species from 37 myrtaceous genera. To determine the threat P. psidii poses to plantation and native eucalypts, artificial inoculation was used to screen germplasm of spotted gum (Corymbia spp.) for resistance to the biotype of P. psidii that has become established in Australia. The objective was to characterize resistance to P. psidii within the Corymbia species complex so that management strategies for the deployment of germplasm from existing breeding programmes of these spotted gum species could be developed. Symptom development initiated 7 days after inoculation, with resistant and susceptible seedlings identified within all species, provenances and families. Inter‐ and intraspecific variability in rust resistance was observed among spotted gum species. There was no apparent relationship between climatic conditions at the provenance origin and disease resistance. The heritability estimates for all assessments are moderate to high and indicate a significant level of additive genetic variance for rust resistance within the populations. The results of this study clearly identify potential to select for resistance at the family level within the tested populations. While the potential for P. psidii to detrimentally impact upon Corymbia in the nursery and in young plantations was demonstrated, estimations of the heritability of resistance suggest that efforts to enhance this trait through breeding have reasonable prospects for success.

The Evolution of Canalization and Evolvability in Stable and Fluctuating Environments

Using a multilinear model of epistasis we explore the evolution of canalization (reduced mutational effects) and evolvability (levels of additive genetic variance) under different forms of stabilizing and fluctuating selection. We show that the total selection acting on an allele can be divided into a component deriving from adaptation of the trait mean, a component of canalizing selection favoring alleles that epistatically reduce the effects of other allele substitutions, and a component of conservative selection disfavoring rare alleles. While canalizing selection operates in both stable and fluctuating environments, it may not typically maximize canalization, because it gets less efficient with increasing canalization, and reaches a balance with drift, mutation and indirect selection. Fluctuating selection leads to less canalized equilibria than stabilizing selection of comparable strength, because canalization then becomes influenced by erratic correlated responses to shifting trait adaptation. We conclude that epistatic systems under bounded fluctuating selection will become less canalized than under stabilizing selection and may support moderately increased evolvability if the amplitude of fluctuations is large, but canalization is still stronger and evolvability lower than expected under neutral evolution or under patterns of selection that shift the trait in directions of positive (reinforcing) epistasis.

Speeding Up Microevolution: The Effects of Increasing Temperature on Selection and Genetic Variance in a Wild Bird Population

Author SummaryThe speed of evolutionary change in a phenotypic trait is determined by two key components: the amount of genetic variance underlying the trait and the strength of selection acting on it. Many studies have shown that both selection and expression of genetic variance may depend on the environmental conditions the population experiences. However, the possibility that the strength of selection and the expression of genetic variance become positively or negatively associated as a result of this environmental covariance, so as to speed up or hamper an evolutionary response, has been largely ignored. Here we show that, in a wild bird population, the annual strength of selection on and the expression of genetic variance in timing of breeding (a key life history trait) are positively associated due to changing environmental conditions (warmer temperatures). Such a positive association should potentially speed up any microevolutionary response to selection (such as that imposed by climate warming). Our results illustrate the existence of substantial temporal variation in response to environmental heterogeneity, and thus highlight a so far neglected mechanism that may be important in determining the evolutionary dynamics in natural populations.

Good Genes and Sexual Selection in Dung Beetles (Onthophagus taurus): Genetic Variance in Egg-to-Adult and Adult Viability

Whether species exhibit significant heritable variation in fitness is central for sexual selection. According to good genes models there must be genetic variation in males leading to variation in offspring fitness if females are to obtain genetic benefits from exercising mate preferences, or by mating multiply. However, sexual selection based on genetic benefits is controversial, and there is limited unambiguous support for the notion that choosy or polyandrous females can increase the chances of producing offspring with high viability. Here we examine the levels of additive genetic variance in two fitness components in the dung beetle Onthophagus taurus. We found significant sire effects on egg-to-adult viability and on son, but not daughter, survival to sexual maturity, as well as moderate coefficients of additive variance in these traits. Moreover, we do not find evidence for sexual antagonism influencing genetic variation for fitness. Our results are consistent with good genes sexual selection, and suggest that both pre- and postcopulatory mate choice, and male competition could provide indirect benefits to females.

The heritability of multiple male mating in a promiscuous mammal.

The tendency of females to mate with multiple males is often explained by direct and indirect benefits that could outweigh the many potential costs of multiple mating. However, behaviour can only evolve in response to costs and benefits if there is sufficient genetic variation on which selection can act. We followed 108 mating chases of 85 North American red squirrels (Tamiasciurus hudsonicus) during 4 years, to measure each female's degree of multiple male mating (MMM), and used an animal model analysis of our multi-generational pedigree to provide what we believe is the first estimate of the heritability of MMM in the wild. Female red squirrels were highly polyandrous, mating with an average of 7.0 ± 0.2 males on their day of oestrus. Although we found evidence for moderate levels of additive genetic variation (CV(A) = 5.1), environmental variation was very high (CV(E) = 32.3), which resulted in a very low heritability estimate (h(2) < 0.01). So, while there is genetic variation in this trait, the large environmental variation suggests that any costs or benefits associated with differences among females in MMM are primarily owing to environmental and not genetic differences, which could constrain the evolutionary response to natural selection on this trait.

Quantitative genetic parameters for wild stream‐living brown trout: heritability and parental effects

Adaptability depends on the presence of additive genetic variance for important traits. Yet few estimates of additive genetic variance and heritability are available for wild populations, particularly so for fishes. Here, we estimate heritability of length-at-age for wild-living brown trout (Salmo trutta), based on long-term mark-recapture data and pedigree reconstruction based on large-scale genotyping at 15 microsatellite loci. We also tested for the presence of maternal and paternal effects using a Bayesian version of the Animal model. Heritability varied between 0.16 and 0.31, with reasonable narrow confidence bands, and the total phenotypic variance increased with age. When introducing dam as an additional random effect (accounting for c. 7% of total phenotypic variance), the level of additive genetic variance and heritability decreased (0.12-0.21). Parental size (both for sires and for dams) positively influenced length-at-age for juvenile trout--either through direct parental effects or through genotype-environment correlations. Length-at-age is a complex trait reflecting the effects of a number of physiological, behavioural and ecological processes. Our data show that fitness-related traits such as length-at-age can retain high levels of additive genetic variance even when total phenotypic variance is high.

Quantitative genetic variation in courtship song and its covariation with immune function and sperm quality in the field cricket Teleogryllus oceanicus

Acoustic signals used by males to attract females are among the most prominent examples of secondary sexual traits, yet we have only limited understanding of their genetic architecture. Male crickets produce a calling song to attract females and then switch to a courtship song that incites mounting by females once they are at close range. Although we know much about the genetics of cricket calling song, no study has yet examined the quantitative genetics of courtship song. Here, we conduct a quantitative genetic analysis of courtship song using the Australian cricket Teleogryllus oceanicus. We find substantial levels of additive genetic variation in courtship song parameters and genetic covariances between courtship song parameters that reflect female preferences. We also found evidence for a negative relationship between the amount of trill in the courtship song and the ability of males to mount an immune response and to produce an ejaculate containing a high proportion of viable sperm. Trade-offs between immunity and sexual signaling are a prerequisite for immunocompetence handicap models of preference evolution, whereas a trade-off between gaining matings and fertilizations is a fundamental assumption underlying sperm competition theory. These life-history trade-offs are likely to constrain evolutionary responses to selection from female choice, and to maintain additive genetic variance in courtship song.

ASYMMETRY OF GENETIC VARIATION IN FITNESS-RELATED TRAITS: APPARENT STABILIZING SELECTION ONgmax

The maintenance of genetic variation in traits closely associated with fitness remains a key unresolved issue in evolutionary genetics. One important qualification on the observation of genetic variation in fitness-related traits is that such traits respond asymmetrically to selection, evolving to a greater extent in the direction of lower fitness. Here we test the hypothesis that standing genetic variation in fitness-related traits is principally maintained for unfit phenotypes. Male Drosophila bunnanda vary in mating success (the primary determinant of male fitness) due to female mate choice. We used competitive mating success to partitioning males into two groups: successful (high fitness) and unsuccessful (low fitness). Relative to successful males, unsuccessful males harbored considerably greater levels of additive genetic variation for sexual signaling traits. This genetic asymmetry was detected for a multivariate trait that we demonstrated was not directly under stabilizing sexual selection, leading us to conclude the trait was under apparent stabilizing selection. Consequently, our results suggest genetic variance might be biased toward low fitness even for traits that are not themselves the direct targets of selection. Simple metrics of genetic variance are unlikely to be adequate descriptors of the complex nature of the genetic basis of traits under selection.

Quantitative genetic variation and selection of enzyme activities in Drosophila melanogaster

The nature of the genetic variation for the activity of three enzymes (α-GPD, ME, and SOD) was studied by means of analyses of variance among full-sib and half-sib families. The results presented here indicate that the genetic variation of activity of these enzymes consist primarily of non-additive genetic variance. A moderate level of additive genetic variation was found only for α-GPD activity. We also examined the question whether an association exists between enzyme activities and selection for preadult developmental time. Using the method developed by Lande and Arnold (1983), significant directional selection was observed for α-GPD activity.

Genetic parameters for growth, wood density and pulp yield in Eucalyptus globulus

Genetic variation and co-variation among the key pulpwood selection traits for Eucalyptus globulus were estimated for a range of sites in Portugal, with the aim of improving genetic parameters used to predict breeding values and correlated response to selection. The trials comprised clonally replicated full-sib families (eight trials) and unrelated clones (17 trials), and exhibited varying levels of pedigree connectivity. The traits studied were stem diameter at breast height, Pilodyn penetration (an indirect measure of wood basic density) and near infrared reflectance predicted pulp yield. Univariate and multivariate linear mixed models were fitted within and across sites, and estimates of additive genetic, total genetic, environmental and phenotypic variances and covariances were obtained. All traits studied exhibited significant levels of additive genetic variation. The average estimated within-site narrow-sense heritability was 0.19 ± 0.03 for diameter and 0.29 ± 0.03 for Pilodyn penetration, and the pooled estimate for predicted pulp yield was 0.42 ± 0.14. When they could be tested, dominance and epistatic effects were generally not statistically significant, although broad-sense heritability estimates were slightly higher than narrow-sense heritability estimates. Averaged across trials, positive additive (0.64 ± 0.08), total genetic (0.58 ± 0.04), environmental (0.38 ± 0.03) and phenotypic (0.43 ± 0.02) correlation estimates were consistently obtained between diameter and Pilodyn penetration. This data argues for at least some form of pleiotropic relationship between these two traits and that selection for fast growth will adversely affect wood density in this population. Estimates of the across-site genetic correlations for diameter and Pilodyn penetration were high, indicating that the genotype by environment interaction is low across the range of sites tested. This result supports the use of single aggregated selection criteria for growth and wood density across planting environments in Portugal, as opposed to having to select for performance in different environments.

Genetic variation in Eucalyptus nitens pulpwood and wood shrinkage traits

Eucalyptus nitens plantations are generally established for pulpwood production but an increasing area is being managed for solid wood. Genetic variation in, and correlations among, three Kraft pulpwood traits (diameter at breast height, basic density and near-infrared-predicted cellulose content) and three 12-mm wood-core shrinkage traits (recoverable collapse, net shrinkage and gross shrinkage) were examined, utilising data from two 9-year-old first-generation progeny trials in Tasmania. These trials contained approximately 400 open-pollinated families (over 100 of which were sampled for wood properties) representing three central-Victorian E. nitens races. Significant genetic variation at the race and/or within-race level was identified in all traits. Within races, relative levels of additive genetic variation were higher for shrinkage traits, although narrow-sense heritabilities were lower and the expression of genetic variation less stable across sites than for other wood property traits. Heterogeneous intertrait genetic correlations were identified across sites between growth and some wood property traits. However, where significant, genetic correlations indicated that within-race selection for growth would adversely affect core basic density and all core shrinkage traits. Furthermore, results based on cores suggested that within-race selection for higher basic density would favourably impact on cellulose content and collapse but selection for either higher basic density or cellulose content would adversely affect net shrinkage. Most within-race genetic variation in gross shrinkage appeared to be due to genetic variation in collapse. The implications of these results for sawn timber breeding will depend on the strength of genetic correlations between core traits and rotation-age objective traits and objective trait economic weights.

Population Bottlenecks Increase Additive Genetic Variance But Do Not Break a Selection Limit in Rain Forest Drosophila

According to neutral quantitative genetic theory, population bottlenecks are expected to decrease standing levels of additive genetic variance of quantitative traits. However, some empirical and theoretical results suggest that, if nonadditive genetic effects influence the trait, bottlenecks may actually increase additive genetic variance. This has been an important issue in conservation genetics where it has been suggested that small population size might actually experience an increase rather than a decrease in the rate of adaptation. Here we test if bottlenecks can break a selection limit for desiccation resistance in the rain forest-restricted fly Drosophila bunnanda. After one generation of single-pair mating, additive genetic variance for desiccation resistance increased to a significant level, on average higher than for the control lines. Line crosses revealed that both dominance and epistatic effects were responsible for the divergence in desiccation resistance between the original control and a bottlenecked line exhibiting increased additive genetic variance for desiccation resistance. However, when bottlenecked lines were selected for increased desiccation resistance, there was only a small shift in resistance, much less than predicted by the released additive genetic variance. The small selection response in the bottlenecked lines was no greater than that observed in the control lines. Thus bottlenecks might produce a statistically detectable change in additive genetic variance but this change has no impact on the response to selection.

Selective breeding can increase resistance of Atlantic salmon to furunculosis, infectious salmon anaemia and infectious pancreatic necrosis

We reasoned that by challenging large numbers of Atlantic salmon families with the causative agents of furunculosis, infectious salmon anaemia (ISA) and infectious pancreatic necrosis (IPN), we could show unequivocally that resistance to these diseases expresses moderate-to-high levels of additive genetic variation, and that the resistances are weakly correlated genetically. We tested this reasoning by challenging Atlantic salmon from 920 (approximately) full-sib families with the causative agents of furunculosis and ISA, and fish from 265 of these families with the causative agent of IPN. Additive genetic variation and genetic correlations were estimated by fitting a threshold liability model to the resistances assessed as binary traits. Resistance to furunculosis, ISA and IPN was moderate –to highly heritable. The marginal posterior means for heritability on the underlying liability scale were 0.37 for resistance to ISA, and 0.55 and 0.62 for resistance to IPN and furunculosis. Genetic correlations between the resistances were weak (−0.11 to 0.07). These levels of additive genetic variation indicate that resistance to furunculosis, ISA and IPN will respond to selection. The weak genetic correlations indicate that it should be relatively easy to improve resistance to the diseases simultaneously. We believe that there is now strong evidence that selectively breeding Atlantic salmon for resistance can be highly successful.

Genetic parameters of intra- and inter-specific hybrids of Eucalyptus globulus and E. nitens

Intra-specific hybrids within Eucalyptus nitens and E. globulus were compared directly with inter-specific E. nitens × globulus using common parents. Diameter (age 2, 4, 6, 10 years) and Pilodyn (age 6 years) were used as indirect measures of growth and wood density, respectively. Genetic parameters were estimated for all cross types and traits. A direct comparison of the general combining ability with the general hybridising ability was made. Heterosis was estimated for intra- and inter-specific hybrids. The intra-specific hybrids in both species exhibit intermediate heritabilities and levels of additive genetic variance. The inter-specific F1 E. nitens × globulus exhibited high incidence of abnormalities at young ages and high levels of later age mortality. The mean performance of surviving inter-specific F1 hybrids was generally intermediate for all traits, to varying degrees, between the parental intra-specific crosses, and there is little evidence for significant heterosis. Different provenances of E. globulus may exhibit different responses to hybridisation. In this case, the inter-specific F1 hybrids, using Taranna E. globulus, appear to perform worse on average than those using King Island provenance for growth traits. There is evidence that specific hybrid families are produced that outperform most of the pure species families for one or other of the traits examined; however, there is no reliable quantitative genetic method of predicting which parents should be used.

Variation in the peacock’s train shows a genetic component

Female peafowl (Pavo cristatus) show a strong mating preference for males with elaborate trains. This, however, poses something of a paradox because intense directional selection should erode genetic variation in the males' trains, so that females will no longer benefit by discriminating among males on the basis of these traits. This situation is known as the 'lek paradox', and leads to the theoretical expectation of low heritability in the peacock's train. We used two independent breeding experiments, involving a total of 42 sires and 86 of their male offspring, to estimate the narrow sense heritabilities of male ornaments and other morphometric traits. Contrary to expectation, we found significant levels of heritability in a trait known to be used by females during mate choice (train length), while no significant heritabilities were evident for other, non-fitness related morphological traits (tarsus length, body weight or spur length). This study adds to the building body of evidence that high levels of additive genetic variance can exist in secondary sexual traits under directional selection, but further emphasizes the main problem of what maintains this variation.

Estimation of genetic parameters in cultured Pacific oysters (Crassostrea gigas) grown in Tasmania, based on animal models

Selective breeding of Pacific oysters in Tasmania is undertaken by the commercial research company — Australian Seafood Industries Pty. Ltd. (ASI). As part of their research program, we estimated the genetic parameters for several economic traits with a view to refining their ongoing breeding objectives. In 2003, strip spawning of 11 dams and 14 sires produced offspring across 19 families of which 14 were half sibs (either maternal or paternal) and 5 were full sib families. These offspring were individually tagged, placed into family mesh bags and grown out for 25 months under commercial conditions. Genetic parameters were estimated using restricted maximum likelihood methods in ASReml as applied to univariate animal models. Heritability estimates for total wet weight, length, width, depth, wetmeat weight andwet shell weight were medium to high. Overall there was no evidence for sire or dam effects. Significant positive genetic and phenotypic correlations between weight and length, width and depth were observed. Length, width and depth traits were also highly positively correlated with each other (both genetically and phenotypically). Width and depth function index traits (calculated from corresponding trait measurements) however, displayed negative genetic correlations with all other traits although phenotypic correlations were positive. While our genetic parameters will not be applicable to all Pacific oyster grow-out conditions or breeding applications, the demonstrated levels of additive genetic variation in most traits shown in this study indicate that these traits should respond well to selection employed in ASI's oyster breeding program.

Genetic Variation in Cynodon transvaalensis Burtt‐Davy

Cynodon transvaalensis Burtt‐Davy (African bermudagrass) is used as a turfgrass and in interspecific hybridization to produce turfgrass cultivars. Information is lacking on the magnitude of intra‐specific genetic variation for traits related to turfgrass performance. A Design II mating population comprised of 320 F1 plants (4 parental sets, 16 crosses set−1, 5 F1 hybrids cross−1) was used to estimate genetic parameters for 21 traits. The F1 plants were evaluated in replicated field (13 traits) and greenhouse (8 traits) experiments in Stillwater, OK during 2002–2003. Genetic variation was detected for 17 of the 21 traits as indicated by significant (P &lt; 0.05) differences among families within sets. Both additive and dominance genetic effects were detected for most of the 17 traits, but dominance effects usually prevailed over additive effects. Broad sense heritability estimates varied from 0.42 to 0.96. Population improvement via recurrent selection techniques would be possible but difficult as indicated by low levels of additive genetic variation for genetic color, raceme number, seed number, and percent seed set. Dominance effects might be exploited to select clonally propagated F1 hybrid cultivars with enhanced sensor‐rated color, density, turf quality, spring greenup, fall dormancy, percent living cover, raceme number, raceme length, number of florets per inflorescence, plant height, stolon length, number of internodes, internode length, and leaf length.

Contrasting Levels of Variation in Neutral and Quantitative Genetic Loci on Island Populations of Moor Frogs (Rana arvalis)

Reduced levels of genetic variability and a prominent differentiation in both neutral marker genes and phenotypic traits are typical for many island populations as compared to their mainland conspecifics. However, whether genetic diversity in neutral marker genes reflects genetic variability in quantitative traits, and thus, their evolutionary potential, remains typically unclear. Moreover, the phenotypic differentiation on islands could be attributable to phenotypic plasticity, selection or drift; something which seldom has been tested. Using eight polymorphic microsatellite loci and quantitative genetic breeding experiments we conducted a detailed comparison on genetic variability and differentiation between Nordic islands (viz. Gotland, Oland and Laeso) and neighbouring mainland populations of moor frogs (Rana arvalis). As expected, the neutral variation was generally lower in island than in mainland populations. But as opposed to this, higher levels of additive genetic variation (VA) in body size and tibia length were found on the island of Gotland as compared to the mainland population. When comparing the differentiation seen in neutral marker genes (FST) with the differentiation in genes coding quantitative traits (QST) two different evolutionary scenarios were found: while selection might explain a smaller size of moor frogs on Gotland, the differentiation seen in tibia length could be explained by genetic drift. These results highlight the limited utility of microsatellite loci alone in inferring the causes behind an observed phenotypic differentiation, or in predicting the amount of genetic variation in ecologically important quantitative traits.

VERY LOW ADDITIVE GENETIC VARIANCE AND EVOLUTIONARY POTENTIAL IN MULTIPLE POPULATIONS OF TWO RAINFORESTDROSOPHILASPECIES

Most quantitative traits are thought to exhibit high levels of genetic variance and evolutionary potential. However, this conclusion may be biased by a lack of studies on nonmodel organisms and may not generalize to restricted species. A recent study on a single, southern population of the rainforest-restricted Drosophila birchii failed to find significant additive genetic variance for the desiccation resistance trait; however, it is unclear whether this pattern extends to other D. birchii populations or to other rainforest species. Here we use an animal model design to show very low levels of additive genetic variance for desiccation resistance in multiple populations of two highly sensitive rainforest species of Drosophila from tropical northeastern Australia. In contrast, relatively high levels of genetic variance were found for morphological traits in all populations of the species tested. This indicates limited evolutionary potential for evolving increased desiccation resistance in these rainforest restricted species.