Index versus tandem selection after repeated generations of selection

Summary A theoretical comparison between two multiple-trait selection methods, index and tandem selection, after several generations of selection was carried out. An infinite number of loci determining the traits, directional and truncation selection, discrete generations and infinite population size were assumed. Under these assumptions, changes in genetic parameters over generations are due to linkage disequilibrium generated by selection. Changes continue for several generations until equilibrium is approached. Algebraic expressions for asymptotic responses from index selection can be derived if index weights are maintained constant across generations. Expressions at equilibrium for genetic parameters and responses are given for the index and its component traits. The loss in response by using initial index weights throughout all generations, instead of updating them to account for changes in genetic parameters, was analyzed. The benefit of using optimum weights was very small ranging from 0% to about 1.5% for all cases studied. Recurrence formulae to predict genetic parameters and responses at each generation of selection are given for both index and tandem selection. A comparison between expected response in the aggregate genotype at equilibrium from index and tandem selection is made considering two traits of economic importance. The results indicate that although index selection is more efficient for improving the aggregate breeding value, its relative efficiency with respect to tandem selection decreases after repeated cycles of selection. The reduction in relative efficiency is highest with the highest selection intensity and heritabilities and with negative correlations between the two traits. The advantage of index over tandem selection might be further reduced if changes in genetic parameters due to gene frequency changes produced by selection, random fluctuations due to the finite size of the population, and errors in estimation of parameters, were also considered.     

Selection for uniformity in livestock by exploiting genetic heterogeneity of residual variance

In some situations, it is worthwhile to change not only the mean, but also the variability of traits by selection. Genetic variation in residual variance may be utilised to improve uniformity in livestock populations by selection. The objective was to investigate the effects of genetic parameters, breeding goal, number of progeny per sire and breeding scheme on selection responses in mean and variance when applying index selection. Genetic parameters were obtained from the literature. Economic values for the mean and variance were derived for some standard non-linear profit equations, e.g. for traits with an intermediate optimum. The economic value of variance was in most situations negative, indicating that selection for reduced variance increases profit. Predicted responses in residual variance after one generation of selection were large, in some cases when the number of progeny per sire was at least 50, by more than 10% of the current residual variance. Progeny testing schemes were more efficient than sib-testing schemes in decreasing residual variance. With optimum traits, selection pressure shifts gradually from the mean to the variance when approaching the optimum. Genetic improvement of uniformity is particularly interesting for traits where the current population mean is near an intermediate optimum.     

Asymmetrical correlated responses to selection under an infinitesimal genetic model

Summary Asymmetry in correlated responses to selection is expected when more than one cycle of selection is practised due to changes in genetic parameters produced by selection. In large populations, under the infinitesimal model these changes are due to linkage disequilibrium generated by selection and not to gene frequency changes. This study examines the conditions under which asymmetrical correlated responses are to be expected when an infinitesimal model is considered. Asymmetrical correlated responses in two traits in respect to which trait is selected are expected if the two traits have different heritabilities. Predicted asymmetry increases with the absolute value of the genetic correlation between the two traits, the difference between the two heritabilities, the intensity of selection and the number of generations of selection. Linkage disequilibrium generated by selection should be taken into account in explaining asymmetrical correlated responses observed in selection experiments.     

Bayesian covariance selection in generalized linear mixed models

The generalized linear mixed model (GLMM), which extends the generalized linear model (GLM) to incorporate random effects characterizing heterogeneity among subjects, is widely used in analyzing correlated and longitudinal data. Although there is often interest in identifying the subset of predictors that have random effects, random effects selection can be challenging, particularly when outcome distributions are nonnormal. This article proposes a fully Bayesian approach to the problem of simultaneous selection of fixed and random effects in GLMMs. Integrating out the random effects induces a covariance structure on the multivariate outcome data, and an important problem that we also consider is that of covariance selection. Our approach relies on variable selection-type mixture priors for the components in a special Cholesky decomposition of the random effects covariance. A stochastic search MCMC algorithm is developed, which relies on Gibbs sampling, with Taylor series expansions used to approximate intractable integrals. Simulated data examples are presented for different exponential family distributions, and the approach is applied to discrete survival data from a time-to-pregnancy study.     

Translating insights from the seed metabolome into improved prediction for lipid-composition traits in oat (Avena sativa L.)

Oat (Avena sativa L.) seed is a rich resource of beneficial lipids, soluble fiber, protein, and antioxidants, and is considered a healthful food for humans. Little is known regarding the genetic controllers of variation for these compounds in oat seed. We characterized natural variation in the mature seed metabolome using untargeted metabolomics on 367 diverse lines and leveraged this information to improve prediction for seed quality traits. We used a latent factor approach to define unobserved variables that may drive covariance among metabolites. One hundred latent factors were identified, of which 21% were enriched for compounds associated with lipid metabolism. Through a combination of whole-genome regression and association mapping, we show that latent factors that generate covariance for many metabolites tend to have a complex genetic architecture. Nonetheless, we recovered significant associations for 23% of the latent factors. These associations were used to inform a multi-kernel genomic prediction model, which was used to predict seed lipid and protein traits in two independent studies. Predictions for 8 of the 12 traits were significantly improved compared to genomic best linear unbiased prediction when this prediction model was informed using associations from lipid-enriched factors. This study provides new insights into variation in the oat seed metabolome and provides genomic resources for breeders to improve selection for health-promoting seed quality traits. More broadly, we outline an approach to distill high-dimensional “omics” data to a set of biologically meaningful variables and translate inferences on these data into improved breeding decisions.     

Levels of selection on threshold characters

Threshold models are useful for understanding the evolution of dimorphic traits with polygenic bases. Selection for threshold characters on individuals is expected to be frequency dependent because of the peculiar way that selection views underlying genetic and environmental factors. Selection among individuals is inefficient because individual phenotypes fall into only two discrete categories that map imperfectly to the underlying genes. Incidence, however, can be continuously distributed among groups, making among-group selection relatively more efficient. Differently put, the group-mean phenotype can be a better predictor of an individual's genotype than that individual's own phenotype. Because evolution in group-structured populations is governed by the balance of selection within and between groups, we can expect threshold traits to evolve in fundamentally different ways when group mean fitness is a function of morph frequency. We extend the theory of selection on threshold traits to include group selection using contextual analysis. For the simple case of linear group-fitness functions, we show that the group-level component of selection, like the individual-level component, is frequency dependent. However, the conditions that determine which component dominates when levels of selection are in conflict (as described by Hamilton's rule) are not frequency dependent. Thus, enhanced group selection is not an inherent property of threshold characters. Nevertheless, we show that predicting the effects of multiple levels of selection on dimorphic traits requires special considerations of the threshold model.     

Pollinator-mediated natural selection in Penstemon digitalis

Measuring the agents of natural selection is important because it allows us to understand not only which traits are expected to evolve but also why they will evolve. Natural selection by pollinators on floral traits is often assumed because in outcrossing animal-pollinated plants flowers are generally thought to function as advertisements of rewards directed at pollinators. We tested the role of bee pollinators in selection on Penstemon digitalis and found that pollinators were driving selection for larger and more flowers. However, what makes our publication unique is the additional information we gained from reviewing the few other studies that also directly tested whether pollinators were agents of selection on floral traits. As we would expect if pollinators are important agents of selection, selection on floral traits was significantly stronger when pollinators were present than when their choices were experimentally removed. Taken together, these results suggest that pollinators can be important drivers of selection in contemporary populations.Key words: plant vigor, Penstemon, pollinators, natural selection, selection gradient     

Tests for the joint evolution of mating system and drought escape in Mimulus

Background and Aims

Self-fertilizing taxa are often found at the range margins of their progenitors, where sub-optimal habitats may select for alternative physiological strategies. The extent to which self-fertilization is favoured directly vs. arising indirectly through correlations with other adaptive life history traits is unclear. Trait responses to selection depend on genetic variation and covariation, as well as phenotypic and genetic responses to altered environmental conditions. We tested predictions of the hypothesis that self-fertilization in Mimulus arises through direct selection on physiological and developmental traits that allow seasonal drought escape.

Methods

Phenotypic selection on mating system and drought escape traits was estimated in field populations of M. guttatus. In addition, trait phenotype and phenotypic selection were compared between experimental wet and dry soil in two greenhouse populations each of M. guttatus and M. nasutus. Finally, genetic variation and covariation for traits were compared between wet and dry soil treatments in a greenhouse population of M. guttatus.

Key Results

Consistent with predictions, selection for early flowering was generally stronger than for mating system traits, and selection for early flowering was stronger in dry soil. Inconsistent with predictions, selection for water-use efficiency was largely absent; selection for large flowers was stronger than for drought escape in the field; and most drought escape and mating system traits were not genetically correlated. A positive genetic correlation between flowering time and flower size, which opposed the adaptive contour, emerged only in wet soil, suggesting that variation in water availability may maintain variation in these traits. Plastic responses to soil moisture treatments supported the idea that taxonomic divergence could have been facilitated by plasticity in flowering time and selfing.

Conclusions

The hypothesis that plant mating systems may evolve indirectly via selection on correlated life history characteristics is plausible and warrants increased attention.     

Testing for biases in selection on avian reproductive traits and partitioning direct and indirect selection using quantitative genetic models

Key life history traits such as breeding time and clutch size are frequently both heritable and under directional selection, yet many studies fail to document microevolutionary responses. One general explanation is that selection estimates are biased by the omission of correlated traits that have causal effects on fitness, but few valid tests of this exist. Here, we show, using a quantitative genetic framework and six decades of life‐history data on two free‐living populations of great tits Parus major, that selection estimates for egg‐laying date and clutch size are relatively unbiased. Predicted responses to selection based on the Robertson–Price Identity were similar to those based on the multivariate breeder's equation (MVBE), indicating that unmeasured covarying traits were not missing from the analysis. Changing patterns of phenotypic selection on these traits (for laying date, linked to climate change) therefore reflect changing selection on breeding values, and genetic constraints appear not to limit their independent evolution. Quantitative genetic analysis of correlational data from pedigreed populations can be a valuable complement to experimental approaches to help identify whether apparent associations between traits and fitness are biased by missing traits, and to parse the roles of direct versus indirect selection across a range of environments.     

Competition and facilitation among fungal plant parasites affect their life-history traits

Multi-infections may result in either competitive exclusion or coexistence on the same host of pathogen genotypes belonging to the same or different species. Epidemiological consequences of multiple infections, particularly how the development and transmission of a pathogen can be modified by the presence of another pathogen, are well documented. However, understanding how life history strategies of each pathogen modulate co-infection outcomes remains quite elusive. To analyze how co-infection drives changes in life history traits and affects co-existence in epidemic pathogens, we infected detached pea stipules with two fungal species, Peyronellaea pinodes and Phoma medicaginis var. pinodella (considering two strains per species), part of the ascochyta blight complex but presenting different life history strategies. All pairwise combinations (including self-pairs) between two strains of each species were tested. Strains were inoculated simultaneously, but apart from one another on the stipule. For each strain, four life history traits were measured: incubation period, necrosis area six days after inoculation, latent period and offspring production. Results show that, in co-infection, when resources are highly allocated to lesion development, the time between inoculation and the appearance of reproduction structures (latent period) and offspring production decreased, and vice-versa relative to single infections. The direction and/or magnitude of these responses to co-infection depend on the co-infecting strains. Moreover, these changes were always higher in self-pairs than in mixed co-infections. These results suggest facilitation between co-infecting strains, resulting in the selection of an intermediate level of virulence (here measured as the lesion development) at the expense of pathogen offspring production. This strategy allows the development and reproduction of each co-infecting strain when sharing limited resources. However, the direction and strength of these life history traits variations in co-infection depend on the life history strategy of the co-infecting strains, with a clear difference between ‘opportunists', ‘scavengers' and ‘pioneer colonisers'.     

Genetic architecture of survival and fitness-related traits in two populations of Atlantic salmon

The additive genetic effects of traits can be used to predict evolutionary trajectories, such as responses to selection. Non-additive genetic and maternal environmental effects can also change evolutionary trajectories and influence phenotypes, but these effects have received less attention by researchers. We partitioned the phenotypic variance of survival and fitness-related traits into additive genetic, non-additive genetic and maternal environmental effects using a full-factorial breeding design within two allopatric populations of Atlantic salmon (Salmo salar). Maternal environmental effects were large at early life stages, but decreased during development, with non-additive genetic effects being most significant at later juvenile stages (alevin and fry). Non-additive genetic effects were also, on average, larger than additive genetic effects. The populations, generally, did not differ in the trait values or inferred genetic architecture of the traits. Any differences between the populations for trait values could be explained by maternal environmental effects. We discuss whether the similarities in architectures of these populations is the result of natural selection across a common juvenile environment.     

Two Species with an Unusual Combination of Traits Dominate Responses of British Grasshoppers and Crickets to Environmental Change

There are large variations in the responses of species to the environmental changes of recent decades, heightening interest in whether their traits may explain inter-specific differences in range expansions and contractions. Using a long-term distributional dataset, we calculated range changes of grasshoppers and crickets in Britain between the 1980s and the 2000s and assessed whether their traits (resource use, life history, dispersal ability, geographic location) explain relative performance of different species. Our analysis showed large changes in the distributions of some species, and we found a positive relationship between three traits and range change: ranges tended to increase for habitat generalists, species that oviposit in the vegetation above ground, and for those with a southerly distribution. These findings accord well with the nature of environmental changes over this period (climatic warming; reductions in the diversity and increases in the height of vegetation). However, the trait effects applied mainly to just two species, Conocephalus discolor and Metrioptera roeselii, which had shown the greatest range increases. Once they were omitted from the analysis, trait effects were no longer statistically significant. Previous studies on these two species emphasised wing-length dimorphism as the key to their success, resulting in a high phenotypic plasticity of dispersal and evolutionary-ecological feedback at their expanding range margins. This, combined with our results, suggests that an unusual combination of traits have enabled these two species to undertake extremely rapid responses to recent environmental changes. The fact that our results are dominated by two species only became apparent through cautious testing of the results’ robustness, not through standard statistical checks. We conclude that trait-based analyses may contribute to the assessment of species responses to environmental change and provide insights into underlying mechanisms, but results need to be interpreted with caution and may have limited predictive power.     

Variation,selection and evolution of function-valued traits

We describe an emerging framework for understanding variation, selection and evolution of phenotypic traits that are mathematical functions. We use one specific empirical example – thermal performance curves (TPCs) for growth rates of caterpillars – to demonstrate how models for function-valued traits are natural extensions of more familiar, multivariate models for correlated, quantitative traits. We emphasize three main points. First, because function-valued traits are continuous functions, there are important constraints on their patterns of variation that are not captured by multivariate models. Phenotypic and genetic variation in function-valued traits can be quantified in terms of variance-covariance functions and their associated eigenfunctions: we illustrate how these are estimated as well as their biological interpretations for TPCs. Second, selection on a function-valued trait is itself a function, defined in terms of selection gradient functions. For TPCs, the selection gradient describes how the relationship between an organism's performance and its fitness varies as a function of its temperature. We show how the form of the selection gradient function for TPCs relates to the frequency distribution of environmental states (caterpillar temperatures) during selection. Third, we can predict evolutionary responses of function-valued traits in terms of the genetic variance-covariance and the selection gradient functions. We illustrate how non-linear evolutionary responses of TPCs may occur even when the mean phenotype and the selection gradient are themselves linear functions of temperature. Finally, we discuss some of the methodological and empirical challenges for future studies of the evolution of function-valued traits.     

Predicting evolutionary responses to selection on polyandry in the wild: additive genetic covariances with female extra-pair reproduction

The evolutionary forces that underlie polyandry, including extra-pair reproduction (EPR) by socially monogamous females, remain unclear. Selection on EPR and resulting evolution have rarely been explicitly estimated or predicted in wild populations, and evolutionary predictions are vulnerable to bias due to environmental covariances and correlated selection through unmeasured traits. However, evolutionary responses to (correlated) selection on any trait can be directly predicted as additive genetic covariances (cov_A) with appropriate components of relative fitness. I used comprehensive life-history, paternity and pedigree data from song sparrows (Melospiza melodia) to estimate cov_A between a female''s liability to produce extra-pair offspring and two specific fitness components: relative annual reproductive success (ARS) and survival to recruitment. All three traits showed non-zero additive genetic variance. Estimates of cov_A were positive, predicting evolution towards increased EPR, but 95% credible intervals overlapped zero. There was therefore no conclusive prediction of evolutionary change in EPR due to (correlated) selection through female ARS or recruitment. Negative environmental covariance between EPR and ARS would have impeded evolutionary prediction from phenotypic selection differentials. These analyses demonstrate an explicit quantitative genetic approach to predicting evolutionary responses to components of (correlated) selection on EPR that should be unbiased by environmental covariances and unmeasured traits.     

Lack of genetic structure among ecologically adapted populations of an Australian rainforest Drosophila species as indicated by microsatellite markers and mitochondrial DNA sequences

Although fragmented rainforest environments represent hotspots for invertebrate biodiversity, few genetic studies have been conducted on rainforest invertebrates. Thus, it is not known if invertebrate species in rainforests are highly genetically fragmented, with the potential for populations to show divergent selection responses, or if there are low levels of gene flow sufficient to maintain genetic homogeneity among fragmented populations. Here we use microsatellite markers and DNA sequences from the mitochondrial ND5 locus to investigate genetic differences among Drosophila birchii populations from tropical rainforests in Queensland, Australia. As found in a previous study, mitochondrial DNA diversity was low with no evidence for population differentiation among rainforest fragments. The pattern of mitochondrial haplotype variation was consistent with D. birchii having undergone substantial past population growth. Levels of nuclear genetic variation were high in all populations while F(ST) values were very low, even for flies from geographically isolated areas of rainforest. No significant differentiation was observed between populations on either side of the Burdekin Gap (a long-term dry corridor), although there was evidence for higher gene diversity in low-latitude populations. Spatial autocorrelation coefficients were low and did not differ significantly from random, except for one locus which revealed a clinal-like pattern. Comparisons of microsatellite differentiation contrasted with previously established clinal patterns in quantitative traits in D. birchii, and indicate that the patterns in quantitative traits are likely to be due to selection. These results suggest moderate gene flow in D. birchii over large distances. Limited population structure in this species appears to be due to recent range expansions or cycles of local extinctions followed by recolonizations/expansions. Nevertheless, patterns of local adaptation have developed in D. birchii that may result in populations showing different selection responses when faced with environmental change.     

The genome of the Xingu scale-backed antbird (Willisornis vidua nigrigula) reveals lineage-specific adaptations

Antbirds (Thamnophilidae) are a large neotropical family of passerine bird renowned for the ant-following foraging strategies of several members of this clade. The high diversity of antbirds provides ample opportunity for speciation studies, however these studies can be hindered by the lack of an annotated antbird reference genome. In this study, we produced a high-quality annotated reference genome for the Xingu Scale-backed Antbird (Willisornis vidua nigrigula) using 10X Genomics Chromium linked-reads technology. The assembly is 1.09 Gb, with a scaffold N50 of 12.1 Mb and 17,475 annotated protein coding genes. We compare the proteome of W. v. nigrigula to several other passerines, and produce annotations for two additional antbird genomes in order to identify genes under lineage-specific positive selection and gene families with evidence for significant expansions in antbirds. Several of these genes have functions potentially related to the lineage-specific traits of antbirds, including adaptations for thermoregulation in a humid tropical environment.     

A general model of the relation between phenotypic selection and genetic response

The phenotypic view of selection assumes that genetic responses can be predicted from selective forces and heritability — or in the classical quantitative genetic equation: R = h²S. However, data on selection in bird populations show that often no selection responses is found, despite consistent selective forces on phenotypes and significant heritable variation. Such discrepancies may arise due to the assumption that selection only acts on observed phenotypes. We derive a general selection equation that takes into account the possibility that some relevant (internal or external) traits are not measured. This equation shows that the classic equation applies if selection directly acts on the measured, phenotypic traits. This is not the case when, for instance, there are unknown internal genetic trade-offs, or unknown common environmental factors affecting both trait and fitness. In such cases, any relationship between phenotypic selection and genetic response is possible. Fortunately, the classical model can be tested by comparing phenotypic and genetic covariances between traits and fitness; an indication that important internal or external traits are missing can thus be obtained. Such an analysis was indeed found in the literature; for selection on fledging weight in Great Tits it yielded valuable extra information.     

Environmental and Parental Influences on Offspring Health and Growth in Great Tits (Parus major)

Sexual selection requires both that there is heritable variation in traits related to fitness, and that either some of this variation is linked to traits of the parents, and/or that there are direct benefits of choosing particular individuals as mates. This suggests that if direct benefits are important offspring performance should be predicted by traits of the rearing adults. But if indirect benefits are more significant offspring performance should be predicted by traits of the adults at the nest-of-origin. We conducted cross-fostering experiments in great tits (Parus major) over four years, in two of which we manipulated environmental conditions by providing supplemental food. In a third year, some nestlings were directly supplemented with carotenoids. Nestlings in broods whose rearing adults received supplemental food were heavier and had improved immune responses even when controlling for body mass. Nestling immune function was related to measures of the yellow plumage color of both the rearing male and the putative father. Nestling body mass was influenced by the coloration of both the rearing female and the genetic mother. Our results suggest that features of both their social and putative genetic parents influence nestling health and growth. From this it would appear that females could be gaining both direct and indirect benefits through mate choice of male plumage traits and that it would be possible for males to similarly gain through mate choice of female traits.