Disruptive viability selection on a black plumage trait associated with dominance

Traits used in communication, such as colour signals, are expected to have positive consequences for reproductive success, but their associations with survival are little understood. Previous studies have mainly investigated linear relationships between signals and survival, but both hump‐shaped and U‐shaped relationships can also be predicted, depending on the main costs involved in trait expression. Furthermore, few studies have taken the plasticity of signals into account in viability selection analyses. The relationship between signal expression and survival is of particular interest in melanin‐based traits, because their main costs are still debated. Here, we first determined the main factors explaining variability in a melanin‐based trait linked to dominance: the bib size of a colonial bird, the sociable weaver Philetairus socius. We then used these analyses to obtain a measure representative of the individual mean expression of bib size. Finally, we used capture–recapture models to study how survival varied in relation to bib size. Variation in bib size was strongly affected by year and moderately affected by age, body condition and colony size. In addition, individuals bearing small and large bibs had higher survival than those with intermediate bibs, and this U‐shaped relationship between survival and bib size appeared to be more pronounced in some years than others. These results constitute a rare example of disruptive viability selection, and point towards the potential importance of social costs incurred by the dominance signalling function of badges of status.     

Caught in the middle: Asymmetric competition causes high variance in intermediate trait abundances

In asymmetric competition between two individuals of the same or different species, one individual has a distinct advantage over the other due to a particular beneficial trait. An important trait that induces asymmetric competition is size (body size in animals, height in plants). There is usually a trade-off between fecundity and the trait that leads to competitive superiority (e.g. seed number vs seed size), enabling coexistence of populations with different trait values. These predictions on coexistence are based on classic deterministic models. Here, we explore the behaviour of a stochastic model of asymmetric competition where stochasticity is assumed to be demographic. We derive approximations for the temporal variance and covariance of the population sizes of the coexisting species. The derivations highlight that the variability of the population size of a species strongly depends on the stochastic fluctuations of species with higher trait values, while they are less influenced by species with lower trait values. Particularly, species with intermediate trait values are strongly affected resulting in relatively high variability. As a result these species have a relative high probability of extinction even though they have a larger population size than species with high trait values. We confirm these approximations with individual-based simulations. Thus, our analysis can explain gaps in size distributions as an emergent property of systems with a fecundity–competition trade-off.     

Cross-generational effects of climate change on expression of a sexually selected trait

Sexually selected traits and early breeding are often correlated with quality in birds: individuals that breed earlier in the season have more elaborate traits and raise more surviving offspring [1, 2]. As global climate warms, breeding date for many temperate birds is advancing [3, 4], but we lack corresponding information on climate-induced variation in sexual selection. Here, we investigated influences of climate on a sexually selected plumage trait in a Himalayan warbler (Phylloscopus humei). We found that when spring is warm, birds breed early. Subsequent to an early-breeding year, adults express relatively large sexually selected traits and rear offspring that also develop large traits. The positive effects of early breeding, plus the across-year correlation between parent and offspring cohorts, predict that warmer climates should lead to increases in trait size. However, trait size has not increased over the past 25 years, even though mean breeding date has advanced. We show that whereas warm springs have positive effects on trait size, warm summers have negative effects due to increased feather wear. Apparent stasis in the size of a sexually selected trait thus masks large, conflicting influences of climate change. Continued climate warming has the potential to affect the honesty of sexual signals, as trait expression and condition become increasingly disassociated.     

Metamerism,morphogenesis, and the expression of carabelli and other dental traits in humans

The patterning cascade model of tooth morphogenesis has emerged as a useful tool in explaining how tooth shape develops and how tooth evolution may occur. Enamel knots, specialized areas of dental epithelium where cusps initiate, act as signaling centers that direct the growth of surrounding tissues. For a new cusp to form, an enamel knot must form beyond the inhibition fields of other enamel knots. The model predicts that the number and size of cusps depends on the spacing between enamel knots, reflected in the spacing between cusps. Recently, work by our group demonstrated that the model predicted Carabelli trait expression in human first molars. Here we test whether differences in Carabelli trait expression along the molar row can also be predicted by the model. Crown areas and intercusp distances were measured from dental casts of 316 individuals with a digital microscope. Although absolute cusp spacing is similar in first and second molars, the smaller size and more triangular shape of second molars results in larger cusp spacing relative to size and, likely, less opportunity for the Carabelli trait to form. The presence and size of the hypocone (HY) and a range of small accessory cusps in a larger sample of 340 individuals were also found to covary with the Carabelli trait in a complex way. The results of this study lend further support to the view that the dentition develops, varies, and evolves as a single functional complex. Am J Phys Anthropol, 2013. © 2013 Wiley Periodicals, Inc.     

The Results of Sable (<Emphasis Type="Italic">Martes zibellina</Emphasis>) Reintroduction Demonstrate the Founder Effect

The relative abundance of intrapopulation groups with different parameters of skull size, coat color, and expression of an epigenetic cranial trait was compared in autochthonous, reintroduced, and donor populations of sable. Recovery of the species resources and broad variability of the phenotypic trait complex in the newly fomned populations were observed. A large proportion of the animals had the phenotype that included large size, dark coat color, and pronounced expression of a specific phene trait (foramen in the condylar fossa) and was not characteristic of the neighboring autochthonous populations. It is reasonable to attribute the presence of individuals with an unusual morphology in the newly formed populations of animals to a manifestation of the founder principle, because the effect of this principle was promoted by spatial isolation of the primary foci of translocated animals.     

Carabelli's trait and tooth size of human maxillary first molars

Carabelli's trait is a morphological feature that can occur on the protocone of human maxillary molars. This study tests the hypothesis that Carabelli's trait is correlated statistically with the dimensions of the crown's four principal cusps or whether, as a cingular feature, the trait truly accretes onto an otherwise unaffected crown. Computer-assisted image analysis was used to measure the 6 intercusp distances and 12 angular relationships among cusp tips on the permanent first molar of 300 young adult American whites. Carabelli's complex was scored using an 8-grade ordinal scheme. Crown size was quantified in three ways, namely as 1) maximum mesiodistal and buccolingual diameters, 2) the 6 intercusp distances, and 3) the 12 angular cusp arrangements. There was no sex difference in the morphological expression of Carabelli's trait in this sample. Overall crown size and intercusp distances were significantly and progressively larger in molars with larger Carabelli's trait expressions. There are graded size responses between crown size (mesiodistal and buccolingual diameters), sizes of the four principal cusps, and morphological stage of Carabelli's complex, though the statistical relationships are appreciably stronger in males than females. Carabelli's trait occurs preferentially in larger molars. In contrast, angular (shape) relationships among cusp tips are not discernibly affected by trait size in either sex. There is the situation, then, that Carabelli's trait is developmentally correlated with crown size, but with no apparent alteration of cusp arrangements, suggesting that the increases are isometric across the occlusal table. Why the association is much weaker in females remains speculative, but these data provide yet another line of evidence that, within a population, tooth size is associated in a positive fashion with crown complexity.     

The evolution of indicator traits for parental quality: the role of maternal and paternal effects

In systems where individuals provide material resources to their mates or offspring, mate choice based on traits that are phenotypically correlated with the quality of resources provided is expected to be adaptive. Several models have explored the evolution of mating preference where there are direct benefits to choice, but few have addressed how a phenotypic correlation can be established between a male indicator trait and the degree of parental investment. We present a model with three quantitative traits: male and female parental investment and a potential male indicator trait. In our model, the expression of the "indicator" trait in offspring is affected by parental investment. These effects are referred to as maternal or paternal effects, or as "indirect genetic effects" when parental investment is heritable. With genetic variation for degree of parental investment, offspring harbor genes for parental investment that are unexpressed before mating but will affect the investment that they provide when expressed. Because the investment received from the parents affects the expression of the indicator trait, there will be a correlation between the genes for parental investment inherited and the degree of expression of the indicator trait in the offspring. The indicator trait is thus an "honest" signal for the degree of paternal investment.     

Parasitism and the expression of sexual dimorphism

Although a negative covariance between parasite load and sexually selected trait expression is a requirement of few sexual selection models, such a covariance may be a general result of life‐history allocation trade‐offs. If both allocation to sexually selected traits and to somatic maintenance (immunocompetence) are condition dependent, then in populations where individuals vary in condition, a positive covariance between trait expression and immunocompetence, and thus a negative covariance between trait and parasite load, is expected. We test the prediction that parasite load is generally related to the expression of sexual dimorphism across two breeding seasons in a wild salamander population and show that males have higher trematode parasite loads for their body size than females and that a key sexually selected trait covaries negatively with parasite load in males. We found evidence of a weaker negative relationship between the analogous female trait and parasite infection. These results underscore that parasite infection may covary with expression of sexually selected traits, both within and among species, regardless of the model of sexual selection, and also suggest that the evolution of condition dependence in males may affect the evolution of female trait expression.     

Eye development in the Cape dune mole rat

Studies on mammalian species with naturally reduced eyes can provide valuable insights into the evolutionary developmental mechanisms underlying the reduction of the eye structures. Because few naturally microphthalmic animals have been studied and eye reduction must have evolved independently in many of the modern groups, novel evolutionary developmental models for eye research have to be sought. Here, we present a first report on embryonic eye development in the Cape dune mole rat, Bathyergus suillus. The eyes of these animals contain all the internal structures characteristic of the normal eye but exhibit abnormalities in the anterior chamber structures. The lens is small but develops normally and exhibits a normal expression of α- and γ-crystallins. One of the interesting features of these animals is an extremely enlarged and highly pigmented ciliary body. In order to understand the molecular basis of this unusual feature, the expression pattern of an early marker of the ciliary zone, Ptmb4, was investigated in this animal. Surprisingly, in situ hybridization results revealed that Ptmb4 expression was absent from the ciliary body zone of the developing Bathyergus eye.     

Prediction of breeding values for group-recorded traits including genomic information and an individually recorded correlated trait

Records on groups of individuals could be valuable for predicting breeding values when a trait is difficult or costly to measure on single individuals, such as feed intake and egg production. Adding genomic information has shown improvement in the accuracy of genetic evaluation of quantitative traits with individual records. Here, we investigated the value of genomic information for traits with group records. Besides, we investigated the improvement in accuracy of genetic evaluation for group-recorded traits when including information on a correlated trait with individual records. The study was based on a simulated pig population, including three scenarios of group structure and size. The results showed that both the genomic information and a correlated trait increased the accuracy of estimated breeding values (EBVs) for traits with group records. The accuracies of EBV obtained from group records with a size 24 were much lower than those with a size 12. Random assignment of animals to pens led to lower accuracy due to the weaker relationship between individuals within each group. It suggests that group records are valuable for genetic evaluation of a trait that is difficult to record on individuals, and the accuracy of genetic evaluation can be considerably increased using genomic information. Moreover, the genetic evaluation for a trait with group records can be greatly improved using a bivariate model, including correlated traits that are recorded individually. For efficient use of group records in genetic evaluation, relatively small group size and close relationships between individuals within one group are recommended.Subject terms: Genetic markers, Animal breeding     

Individual variability in the stress response of C57BL/6J male mice correlates with trait anxiety

Stress strongly alters the physiology and behavior of some individuals, while others are little or not affected. The causes of this individual variability have remained unknown. Here, we hypothesize that epigenetically induced levels of trait anxiety predict the stress response of individual mice in a genetically homogeneous population. Inbred C57BL/6 male mice were selected for their latency to freely enter from their home cage into an unfamiliar arena and classified as having high or low levels of trait anxiety. Mice were then exposed to acute stress (1-h olfactory contact with a rat) or control conditions. After 24 h, acute stress enhanced state anxiety measured in the elevated-plus maze test only in mice previously classified as having high levels of trait anxiety. This anxiogenic effect of acute stress was paralleled by enhanced novelty-induced plasma corticosterone secretion and increased messenger RNA (mRNA) expression for glucocorticoid and mineralocorticoid receptors in the hippocampus. No effects of acute stress were observed in mice classified as having low levels of trait anxiety. Under unstressed control conditions, mice only differed in basal levels of hippocampal mRNA for the glucocorticoid receptor, which were higher in mice with high trait anxiety than in mice with low trait anxiety. In summary, inbred C57BL/6 mice display a remarkably high interindividual variability in their trait anxiety that predicts the behavioral and neuroendocrine response to an acute stressor, indicating that expression of extremely different coping strategies can develop also between genetically identical individuals.     

Identity-by-descent estimation and mapping of qualitative traits in large, complex pedigrees

Computing identity-by-descent sharing between individuals connected through a large, complex pedigree is a computationally demanding task that often cannot be done using exact methods. What I present here is a rapid computational method for estimating, in large complex pedigrees, the probability that pairs of alleles are IBD given the single-point genotype data at that marker for all individuals. The method can be used on pedigrees of essentially arbitrary size and complexity without the need to divide the individuals into separate subpedigrees. I apply the method to do qualitative trait linkage mapping using the nonparametric sharing statistic S(pairs). The validity of the method is demonstrated via simulation studies on a 13-generation 3028-person pedigree with 700 genotyped individuals. An analysis of an asthma data set of individuals in this pedigree finds four loci with P-values <10(-3) that were not detected in prior analyses. The mapping method is fast and can complete analyses of approximately 150 affected individuals within this pedigree for thousands of markers in a matter of hours.     

THE VARIABILITY IS IN THE SEX CHROMOSOMES

Sex differences in the mean trait expression are well documented, not only for traits that are directly associated with reproduction. Less is known about how the variability of traits differs between males and females. In species with sex chromosomes and dosage compensation, the heterogametic sex is expected to show larger trait variability (“sex‐chromosome hypothesis”), yet this central prediction, based on fundamental genetic principles, has never been evaluated in detail. Here we show that in species with heterogametic males, male variability in body size is significantly larger than in females, whereas the opposite can be shown for species with heterogametic females. These results support the prediction of the sex‐chromosome hypothesis that individuals of the heterogametic sex should be more variable. We argue that the pattern demonstrated here for sex‐specific body size variability is likely to apply to any trait and needs to be considered when testing predictions about sex‐specific variability and sexual selection.     

Functional genetics of intraspecific ecological interactions in Arabidopsis thaliana

Studying the genetic basis of traits involved in ecological interactions is a fundamental part of elucidating the connections between evolutionary and ecological processes. Such knowledge allows one to link genetic models of trait evolution with ecological models describing interactions within and between species. Previous work has shown that connections between genetic and ecological processes in Arabidopsis thaliana may be mediated by the fact that quantitative trait loci (QTL) with 'direct' effects on traits of individuals also have pleiotropic 'indirect' effects on traits expressed in neighbouring plants. Here, we further explore these connections by examining functional relationships between traits affected directly and indirectly by the same QTL. We develop a novel approach using structural equation models (SEMs) to determine whether observed pleiotropic effects result from traits directly affected by the QTL in focal individuals causing the changes in the neighbours' phenotypes. This hypothesis was assessed using SEMs to test whether focal plant phenotypes appear to mediate the connection between the focal plants' genotypes and the phenotypes of their neighbours, or alternatively, whether the connection between the focal plants' genotypes and the neighbours' phenotypes is mediated by unmeasured traits. We implement this analysis using a QTL of major effect that maps to the well-characterized flowering locus, FRIGIDA. The SEMs support the hypothesis that the pleiotropic indirect effects of this locus arise from size and developmental timing-related traits in focal plants affecting the expression of developmental traits in their neighbours. Our findings provide empirical insights into the genetics and nature of intraspecific ecological interactions. Our technique holds promise in directing future work into the genetic basis and functional relationship of traits mediating and responding to ecological interactions.     

Sex-specific trait architecture in a spider with sexual size dimorphism

Sexual dimorphism, or sex-specific trait expression, may evolve when selection favours different optima for the same trait between sexes, that is, under antagonistic selection. Intra-locus sexual conflict exists when the sexually dimorphic trait under antagonistic selection is based on genes shared between sexes. A common assumption is that the presence of sexual-size dimorphism (SSD) indicates that sexual conflict has been, at least partly, resolved via decoupling of the trait architecture between sexes. However, whether and how decoupling of the trait architecture between sexes has been realized often remains unknown. We tested for differences in architecture of adult body size between sexes in a species with extreme SSD, the African hermit spider (Nephilingis cruentata), where adult female body size greatly exceeds that of males. Specifically, we estimated the sex-specific importance of genetic and maternal effects on adult body size among individuals that we laboratory-reared for up to eight generations. Quantitative genetic model estimates indicated that size variation in females is to a larger extent explained by direct genetic effects than by maternal effects, but in males to a larger extent by maternal than by genetic effects. We conclude that this sex-specific body-size architecture enables body-size evolution to proceed much more independently than under a common architecture to both sexes.     

Body size and social dominance influence breeding dispersal in male Pachydiplax longipennis (Odonata)

1. Dispersal behaviour can be affected by an individual's phenotype, by the environmental or social context they experience, and by interactions between these factors. Differential dispersal propensities between individuals may also be an important modifier of functional connectivity between populations. To assess how a key trait, body size, affected both social interactions and dispersal behaviour, this study examined the relationship between body size, antagonistic interactions, and breeding dispersal in male dragonflies (Pachydiplax longipennis) across a seasonal decline in adult body size. 2. During a seasonal peak in male body size in this study, dispersers were smaller than non‐dispersers. Later in the season, the body size of dispersers and non‐dispersers did not differ. 3. Focal observations found that body size was related to competitive dominance, large males engaged in aggressive chases more often and smaller males were more frequently pursued. 4. These results indicate that when large males were present, small males were more likely to disperse suggesting that dispersal is a tactic adopted by social subordinates in this context. If breeding dispersal is typically undertaken by subordinate males, functional connectivity between populations may be less than estimated from absolute dispersal rates.     

The genetics of phenotypic plasticity. XI. Joint evolution of plasticity and dispersal rate

In a spatially heterogeneous environment, the rate at which individuals move among habitats affects whether selection favors phenotypic plasticity or genetic differentiation, with high dispersal rates favoring trait plasticity. Until now, in theoretical explorations of plasticity evolution, dispersal rate has been treated as a fixed, albeit probabilistic, characteristic of a population, raising the question of what happens when the propensity to disperse and trait plasticity are allowed to evolve jointly. We examined the effects of their joint evolution on selection for plasticity using an individual-based computer simulation model. In the model, the environment consisted of a linear gradient of 50 demes with dispersal occurring either before or after selection. Individuals consisted of loci whose phenotypic expression either are affected by the environment (plastic) or are not affected (nonplastic), plus a locus determining the propensity to disperse. When dispersal rate and trait plasticity evolve jointly, the system tends to dichotomous outcomes of either high trait plasticity and high dispersal, or low trait plasticity and low dispersal. The outcome strongly depended on starting conditions, with high trait plasticity and dispersal favored when the system started at high values for either trait plasticity or dispersal rate (or both). Adding a cost of plasticity tended to drive the system to genetic differentiation, although this effect also depended on initial conditions. Genetic linkage between trait plasticity loci and dispersal loci further enhanced this strong dichotomy in evolutionary outcomes. All of these effects depended on organismal life history pattern, and in particular whether selection occurred before or after dispersal. These results can explain why adaptive trait plasticity is less common than might be expected.     

Pest control by the introduction of a conditional lethal trait on multiple loci: potential, limitations, and optimal strategies

Advances in genetics have made it feasible to genetically engineer insect strains carrying a conditional lethal trait on multiple loci. We model the release into a target pest population of insects carrying a dominant and fully penetrant conditional lethal trait on 1-20 loci. Delaying the lethality for several generations after release allows the trait to become widely spread in the target population before being activated. To determine effectiveness and optimal strategies for such releases, we vary release size, number of generations until the conditional lethality, nonconditional fitness cost resulting from gene insertions, and fitness reduction associated with laboratory rearing. We show that conditional lethal releases are potentially orders of magnitude more effective than sterile male releases of equal size, and that far smaller release sizes may be required for this approach than necessary with sterile males. For example, a release of male insects carrying a conditional lethal allele that is activated in the F4 generation on 10 loci reduces the target populatioin to 10(-4) of no-release size if there are initially two released males for every wild male. We show how the effectiveness of conditional lethal releases decreases as the nonconditional fitness reduction (i.e., fitness reduction before the trait becomes lethal) associated with the conditional lethal genes increases. For example, if there is a 5% nonconditional fitness cost per conditional lethal allele, then a 2:1 (released male:wild male) release with conditional lethal alleles that are activated in the F4 generation reduces the population to 2-5% (depending on the degree of density dependence) of the no-release size. If there is a per-allele reduction in fitness, then as the number of loci is increased there is a trade-off between the fraction of offspring carrying at least one conditional lethal allele and the fitness of the released insects. We calculate the optimal number of loci on which to insert the conditional lethal gene given various conditions. In addition, we show how laboratory-rearing fitness costs, density-dependence, and all-male versus male-female releases affect the efficiency of conditional lethal releases.