Search

Allometric analysis provides an insight into the function and diversifi cation mechanisms of body parts in organisms. The allometry and variability in size of various body parts of three hymenopteran species, Anisopteromalus calandrae (Howard), Anisopteromalus quinarius Gokhman & Baur, and Heterospilus prosopidis Viereck, which are solitary ectoparasitoids of storedproduct insect pests, were studied. The relationship between many of the traits measured and body size is negatively allometric, others were isometric, and none were positively allometric. The two species of Anisopteromalus were less variable in size than H. prosopidis and there were intersexual diff erences in both species of Anisopteromalus. Although the patterns in these diff erences are complex and diffi cult to interpret from an ecological perspective, based on information on their behaviour it is hypothesized that stabilizing selection is associated with the negative allometry of two traits: ovipositor length in at least two species and male leg length in both species of Anisopteromalus. This hypothesis is supported by the lower variability in the size of these body parts compared to other body parts examined in this study, which is typical of traits subject to stabilizing selection. [ABSTRACT FROM AUTHOR]

Background and Aims Leaf traits are known to be strong predictors of plant performance and can be expected to (co)vary along environmental gradients. We investigated the variation, integration, environmental relationships and evolutionary history of leaf functional traits in the genus Coffea , typically a rainforest understorey shrub, across Africa. A better understanding of the adaptive processes involved in leaf trait evolution can inform the use and conservation of coffee genetic resources in a changing climate. Methods We used phylogenetic comparative methods to investigate the evolution of six leaf traits measured from herbarium specimens of 58 African Coffea species. We added environmental data and data on maximum plant height for each species to test trait–environment correlations in various (sub)clades, and we compared continuous trait evolution models to identify variables driving trait diversification. Key Results Substantial leaf trait variation was detected across the genus Coffea in Africa, which was mostly interspecific. Of these traits, stomatal size and stomatal density exhibited a clear trade-off. We observed low densities of large stomata in early-branching lineages and higher densities of smaller stomata in more recent taxa, which we hypothesize to be related to declining CO2 levels since the mid-Miocene. Brownian motion evolution was rejected in favor of white noise or Ornstein–Uhlenbeck models for all traits, implying these traits are adaptively significant rather than driven by pure drift. The evolution of leaf area was likely driven by precipitation, with smaller leaves in drier climates across the genus. Conclusions Generally, Coffea leaf traits appear to be evolutionarily labile and governed by stabilizing selection, though evolutionary patterns and correlations differ depending on the traits and clades considered. Our study highlights the importance of a phylogenetic perspective when studying trait relationships across related taxa, as well as the consideration of various taxonomic ranges. [ABSTRACT FROM AUTHOR]

Understanding the drivers of community assembly is critical for predicting the future of biodiversity and ecosystem services. Ecological selection ubiquitously shapes communities by selecting for individuals with the most suitable trait combinations. Detecting selection types on key traits across environmental gradients and over time has the potential to reveal the underlying abiotic and biotic drivers of community dynamics. Here, we present a model‐based predictive framework to quantify the multidimensional trait distributions of communities (community trait spaces), which we use to identify ecological selection types shaping communities along environmental gradients. We apply the framework to over 3600 boreal forest understory plant communities with results indicating that directional, stabilizing, and divergent selection all modify community trait distributions and that the selection type acting on individual traits may change over time. Our results provide novel and rare empirical evidence for divergent selection within a natural system. Our approach provides a framework for identifying key traits under selection and facilitates the detection of processes underlying community dynamics. [ABSTRACT FROM AUTHOR]

Sperm competition is a potent mechanism of postcopulatory sexual selection that has been found to shape reproductive morphologies and behaviours in promiscuous animals. Especially sperm size has been argued to evolve in response to sperm competition through its effect on sperm longevity, sperm motility, the ability to displace competing sperm, and ultimately fertilization success. Additionally, sperm size has been observed to co-evolve with female reproductive morphology. Theoretical work predicts that sperm competition may select for longer sperm but may also favour shorter sperm if sperm size trades-off with number. In this study, we studied the relationship between sperm size and postmating success in the free-living flatworm, Macrostomum lignano. Specifically, we used inbred isolines of M. lignano that varied in sperm size to investigate how sperm size translated into the ability of worms to transfer and deposit sperm in a mating partner. Our results revealed a hump-shaped relationship with individuals producing sperm of intermediate size having the highest sperm competitiveness. This finding broadens our understanding of the evolution of sperm morphology by providing empirical support for stabilizing selection on sperm size under sperm competition. Graphical Abstract [ABSTRACT FROM AUTHOR]

Unveiling the ecological and evolutionary mechanisms that guide the differentiation of various plant growth forms into vertical forest strata during forest degradation continues to pose a challenge. We utilized the biogeochemical niche theory to decipher these mechanisms. We measured leaf concentrations of basal elements (C, N, P, K, Ca, and Mg) and used them to generate the biogeochemical niches in herbs, shrubs, trees, and climbers from different vertical strata within a degraded forest and its neighboring mature forest in subtropical montane zones. Our results showed that biogeochemical niches varied significantly among growth forms. Biogeochemical niches also differed significantly between species in the degraded and mature forests, as evidenced by the significantly lower leaf C concentration relative to other nutrients in degraded forest species. This could be attributable mainly to the expansive canopy openness in the degraded forest, facilitating the colonization of light-demanding self-supporters and climbers. The phylogenetic signals in leaf nutrient concentrations were significantly present across all species, suggesting the general similarities in biogeochemical niches among closely related species. Following evolutionary model comparisons, stabilizing selection towards varying optimal leaf nutrient concentrations for distinct growth forms in the degraded or mature forest emerged as the most suitable model. Our study thus highlights the important role of phylogenetic biogeochemical niche conservatism and stabilizing selection towards various optimal leaf nutrient compositions for diverse growth forms in both degraded and mature forests in the formation of vertical plant communities during subtropical forest degradation.

Given the many small-effect loci uncovered by genome-wide association studies (GWAS), polygenic scores have become central to genomic medicine, and have found application in diverse settings including evolutionary studies of adaptation. Despite their promise, polygenic scores have been found to suffer from limited portability across human populations. This at first seems in conflict with the observation that most common genetic variation is shared among populations. We investigate one potential cause of this discrepancy: stabilizing selection on complex traits. Counterintuitively, while stabilizing selection constrains phenotypic evolution, it accelerates the loss and fixation of alleles underlying trait variation within populations (GWAS loci). Thus even when populations share an optimum phenotype, stabilizing selection erodes the variance contributed by their shared GWAS loci, such that predictions from GWAS in one population explain less of the phenotypic variation in another. We develop theory to quantify how stabilizing selection is expected to reduce the prediction accuracy of polygenic scores in populations not represented in GWAS samples. In addition, we find that polygenic scores can substantially overstate average genetic differences of phenotypes among populations. We emphasize stabilizing selection around a common optimum as a useful null model to connect patterns of allele frequency and polygenic score differentiation. This article is part of the theme issue 'Celebrating 50 years since Lewontin's apportionment of human diversity'.

AbstractOrganismal traits often influence fitness via interactions with multiple species. That selection is not necessarily predictable from pairwise interactions, such as when interactions occur during different life cycle stages. Theoretically, directional selection during two sequential episodes (e.g., pollination and seed survival) can generate quadratic or correlational selection for a set of traits that passes both selective filters. We compared strength of selection during pollination versus seed predation in the field and tested whether interactions with multiple species give rise to nonlinear selection on floral traits. We planted common gardens with seeds of two species of Ipomopsis and hybrids at sites where pollination was primarily by hummingbirds or also included hawk moths. We examined selection on six floral traits, including corolla width, sepal width, color, nectar, and two scent compounds. Female fitness (seeds) was broken down into fitness during (1) pollination (seeds initiated) and (2) seed predation (proportion of seeds escaping fly predation). All traits showed evidence of selection. Directional and quadratic selection were stronger during seed initiation than during seed predation. Correlational selection occurred mostly during seed initiation rather than arising from combining species interactions at two points in the life cycle. These results underscore how multispecies interactions can combine to exert selection on trait combinations.

The article is a review of the book by P.A. Vodopyanov At the Turning Point of the Epochs: In Search of a Strategy for Creating the Future (Minsk: Belorussian Science; 2023. 469 p.). The monograph is of great theoretical and practical significance and outlines new horizons for the future sufficient development. The author argues that the humans’ embedment in nature and subordination to the laws of nature, the search for ways to overcome new challenges and risks caused by the negative consequences of scientific and technological achievements determine the new main directions of the coevolutionary strategy for sufficient development, which are associated with the extraordinary and unpopular measures: demographic regulation; reduction of industrial pressure on the biosphere through the introduction of nature-like technologies; economical use of natural resources and their replacement with artificial ones; increasing resource productivity through the introduction of new technologies; the use of alternative energy sources instead of hydrocarbon fuels; the changing vector of the social-economic development based on the laws of nature; a new type of morality and a new humanism; transition to the era of ‘new Enlightenment’ based on the ecological consciousness and thinking. The book identifies the main factors of maintaining the stability of natural ecosystems and the biosphere, which explain the need to use the laws of the biosphere organization to choose a strategy for a secure future. The author presents the key directions for ensuring the future based on bio-anthropocentrism as the main condition for preserving the biosphere and focuses on the main factors of human life under global instability and the need in appropriate ideological universals.

Stabilizing selection provides a challenge to molecular population genetics. Although stabilizing selection is ubiquitous, its genomic signature is difficult to distinguish from demographic signals. Experimental evolution provides a promising approach to characterize genomic regions exposed to stabilizing selection. A recent experimental evolution study of Aedes aegypti populations evolving either with or without sexual selection found a pattern of genetic differentiation suggestive of relaxed stabilizing selection. I argue that this study could not have detected the signal of relaxed stabilizing selection. I highlight why incorrect statistical methods resulted in a high number of false positive candidate single nucleotide polymorphism (SNPs) and discuss the fallacy of functional validation of candidate SNPs for polygenic traits by RNA-mediated knockdown. [ABSTRACT FROM AUTHOR]

Avian flush–pursuit insectivores typically use contrasting white plumage patches in their tails or wings to startle potential prey. Although experimental evidence indicates that the extent of white has been fine-tuned by natural selection to optimize foraging performance, the hypothesis that within-population plumage variation directly influences survival or lifetime reproduction and is subject to stabilizing selection has not been tested. Here, I provide such a test using data collected as part of a 14-year study of a colour-ringed breeding population of the hooded warbler (Setophaga citrina), a migratory flush–pursuit insectivore that shows inter-individual variation in the extent of white in the tail that is highly repeatable across moults and probably heritable. As expected under stabilizing selection, warblers with average-sized white tail patches achieved significantly higher apparent long-term survival than individuals with either a lesser or greater extent of white in the tail. Evidence of stabilizing selection was especially strong in males, an observation that is probably related to pronounced sexual habitat segregation on the wintering range. My results provide infrequently observed evidence of stabilizing selection operating in a natural population, and also illustrate how stabilizing selection can act on avian plumage traits outside the context of sexual and social signalling. [ABSTRACT FROM AUTHOR]

Background and Aims Morphological matching between flower and pollinator traits has been documented in diverse plant lineages. Indeed, the matching of corolla tube length and pollinator tongue length has been cited repeatedly as a classic case of coevolution. However, there are many possible evolutionary routes to trait matching. Our aim here is both to review the evolutionary mechanisms of plant–pollinator trait matching and to investigate a specific case of trait matching/mismatching in a genus of alpine gingers. Methods Roscoea gingers with long corolla tubes in the western Himalayas have pollinators with correspondingly long tongues, but the match between corolla tube and pollinator tongue lengths is not seen in the eastern Himalayas. Six floral traits were measured, including corolla tube depth, an internal trait controlling pollinator access to nectar. We calculated coefficients of variation and phylogenetically controlled correlation patterns of these traits in six Roscoea species in order to gain possible insights into stabilizing selection and modularization of these traits. Key Results The distal (nectar-containing) portion of the corolla tube exhibited lower coefficients of variations than did the basal portion. This is consistent with the hypothesis that pollinators mediate stabilizing selection on the distal, but not basal, portion of the corolla tube. This result, combined with phylogenetic data, suggests that the elevated liquid level of nectar in the distal tube evolved subsequent to dispersal into the eastern Himalayan region and loss of long-tongue pollinators. After accounting for phylogeny, corolla tube length, anther length, style length and labellum width were all intercorrelated. Corolla-tube depth was not part of this covariational module, however, suggesting separate adaptation to short-tongued pollinators. Conclusions The reduction in functional corolla tube depth in the Roscoea appears to be related to the loss of long-tongued pollinators associated with dispersal to the eastern Himalayas and pollination by short-tongued pollinators. The apparent mismatch between floral tubes and pollinator tongues is a case of cryptic trait matching between flowers and pollinators, underscoring the importance of combining floral anatomy with pollination ecology in assessing plant–pollinator trait matching. [ABSTRACT FROM AUTHOR]

Investigating gene expression evolution over micro- and macroevolutionary timescales will expand our understanding of the role of gene expression in adaptation and speciation. In this study, we characterized the evolutionary forces acting on gene expression levels in eye and brain tissue of five Heliconius butterflies with divergence times of ∼5-12 MYA. We developed and applied Brownian motion (BM) and Ornstein-Uhlenbeck (OU) models to identify genes whose expression levels are evolving through drift, stabilizing selection, or a lineage-specific shift. We found that 81% of the genes evolve under genetic drift. When testing for branch-specific shifts in gene expression, we detected 368 (16%) shift events. Genes showing a shift toward upregulation have significantly lower gene expression variance than those genes showing a shift leading toward downregulation. We hypothesize that directional selection is acting in shifts causing upregulation, since transcription is costly. We further uncovered through simulations that parameter estimation of OU models is biased when using small phylogenies and only becomes reliable with phylogenies having ≥ 50 taxa. Therefore, we developed a new statistical test based on BM to identify highly conserved genes (i.e., evolving under strong stabilizing selection), which comprised 3% of the orthoclusters. In conclusion, we found that drift is the dominant evolutionary force driving gene expression evolution in eye and brain tissue in Heliconius Nevertheless, the higher proportion of genes evolving under directional than under stabilizing selection might reflect species-specific selective pressures on vision and the brain that are necessary to fulfill species-specific requirements.

Morphological correlations can not only stem from developmental constraints but also from selective pressures. Butterfly eyespots are repeated wing color pattern elements, widespread across species. As developmental serial homologs, they are controlled by similar developmental pathways imposing correlations among eyespots: selection on a single eyespot may induce correlated responses in all eyespots. We study the variations in the ventral eyespots of Morpho telemachus , where two different selective regimes are likely to act: while most eyespots are always-visible, two eyespots are conditionally displayed: hidden at rest, they can be exposed when the butterflies are threatened, or during sexual interactions. We investigate how such contrasted selection across eyespots can alter the covariations imposed by their shared developmental origin. We quantified eyespots covariations within a large population of M. telemachus and compared the observed patterns to those found in M. helenor , where all eyespots are always-visible and thus probably affected by a similar selection regime. We found that M. telemachus conditionally displayed eyespots are less variable than always-visible eyespots and that these two eyespots form a separate variational module in this species, in contrast to M. helenor. Our results suggest that eyespots covariations were shaped by selection, highlighting how natural selection may promote the evolution of modularity. [ABSTRACT FROM AUTHOR]

The study of adaptation helps explain biodiversity and predict future evolution. Yet the process of adaptation can be difficult to observe due to limited phenotypic variation in contemporary populations. Furthermore, the scarcity of male fitness estimates has made it difficult to both understand adaptation and evaluate sexual conflict hypotheses. We addressed both issues in our study of two anther position traits in wild radish (Raphanus raphanistrum): anther exsertion (long filament − corolla tube lengths) and anther separation (long − short filament lengths). These traits affect pollination efficiency and are particularly interesting due to the unusually high correlations among their component traits. We measured selection through male and female fitness on wild radish plants from populations artificially selected to recreate ancestral variation in each anther trait. We found little evidence for conflicts between male and female function. We found strong evidence for stabilizing selection on anther exsertion and disruptive selection on anther separation, indicating positive and negative correlational selection on the component traits. Intermediate levels of exsertion are likely an adaptation to best contact small bees. The function of anther separation is less clear, but future studies might investigate pollen placement on pollinators and compare species possessing multiple stamen types. [ABSTRACT FROM AUTHOR]

Gene expression evolution is typically modeled with the stochastic Ornstein-Uhlenbeck (OU) process. It has been suggested that the estimation of within-species variations using replicated data can increase the predictive power of such models, but this hypothesis has not been fully tested. We developed EvoGeneX, a computationally efficient implementation of the OU-based method that models within-species variation. Using extensive simulations, we show that modeling within-species variations and appropriate selection of species improve the performance of the model. Further, to facilitate a comparative analysis of expression evolution, we introduce a formal measure of evolutionary expression divergence for a group of genes using the rate and the asymptotic level of divergence. With these tools in hand, we performed the first-ever analysis of the evolution of gene expression across different body-parts, species, and sexes of the Drosophila genus. We observed that genes with adaptive expression evolution tend to be body-part specific, whereas the genes with constrained evolution tend to be shared across body-parts. Among the neutrally evolving gene expression patterns, gonads in both sexes have higher expression divergence relative to other tissues and the male gonads have even higher divergence than the female gonads. Among the evolutionarily constrained genes, the gonads show different divergence patterns, where the male gonads, and not the female gonads, show less constrained divergence than other body-parts. Finally, we show interesting examples of adaptive expression evolution, including adaptation of odor binding proteins. [ABSTRACT FROM AUTHOR]

The dispersal of modern humans from Africa is now well documented with genetic data that track population history, as well as gene flow between populations. Phenetic skeletal data, such as cranial and pelvic morphologies, also exhibit a dispersal-from-Africa signal, which, however, tends to be blurred by the effects of local adaptation and in vivo phenotypic plasticity, and that is often deteriorated by postmortem damage to skeletal remains. These complexities raise the question of which skeletal structures most effectively track neutral population history. The cavity system of the inner ear (the so-called bony labyrinth) is a good candidate structure for such analyses. It is already fully formed by birth, which minimizes postnatal phenotypic plasticity, and it is generally well preserved in archaeological samples. Here we use morphometric data of the bony labyrinth to show that it is a surprisingly good marker of the global dispersal of modern humans from Africa. Labyrinthine morphology tracks genetic distances and geography in accordance with an isolation-by-distance model with dispersal from Africa. Our data further indicate that the neutral-like pattern of variation is compatible with stabilizing selection on labyrinth morphology. Given the increasingly important role of the petrous bone for ancient DNA recovery from archaeological specimens, we encourage researchers to acquire 3D morphological data of the inner ear structures before any invasive sampling. Such data will constitute an important archive of phenotypic variation in present and past populations, and will permit individual-based genotype-phenotype comparisons.

Modern molecular genetic datasets, primarily collected to study the biology of human health and disease, can be used to directly measure the action of natural selection and reveal important features of contemporary human evolution. Here we leverage the UK Biobank data to test for the presence of linear and nonlinear natural selection in a contemporary population of the United Kingdom. We obtain phenotypic and genetic evidence consistent with the action of linear/directional selection. Phenotypic evidence suggests that stabilizing selection, which acts to reduce variance in the population without necessarily modifying the population mean, is widespread and relatively weak in comparison with estimates from other species.

Mammals almost always have seven cervical vertebrae. The strong evolutionary constraint on changes in this number has been broken in sloths and manatees. We have proposed that the extremely low activity and metabolic rates of these species relax the stabilizing selection against changes in the cervical count. Our hypothesis is that strong stabilizing selection in other mammals is largely indirect and due to associated pleiotropic effects, including juvenile cancers. Additional direct selection can occur due to biomechanical problems (thoracic outlet syndrome). Low metabolic and activity rates are thought to diminish these direct and indirect effects. To test this hypothesis within the primates, we have compared the number of cervical vertebrae of three lorisid species with particularly low activity and metabolic rates with those of more active primate species, including with their phylogenetically closest active relatives, the galagids (bushbabies). In support of our hypothesis, we found that 37.6% of the lorisid specimens had an abnormal cervical count, which is a higher percentage than in the other nine primate families, in which the incidence varied from zero to 2.2%. We conclude that our data support the importance of internal selection in constraining evolvability and of a relaxed stabilizing selection for increasing evolvability. Additionally, we discuss that there is no support for a role of the muscularized diaphragm in the evolutionary constraint. [ABSTRACT FROM AUTHOR]

Deleterious genetic variation is maintained in populations at low frequencies. Under a model of stabilizing selection, rare (and presumably deleterious) genetic variants are associated with increase or decrease in gene expression from some intermediate optimum. We investigate this phenomenon in a population of largely Oryza sativa ssp. indica rice landraces under normal unstressed wet and stressful drought field conditions. We include single nucleotide polymorphisms, insertion/deletion mutations, and structural variants in our analysis and find a stronger association between rare variants and gene expression outliers under the stress condition. We also show an association of the strength of this rare variant effect with linkage, gene expression levels, network connectivity, local recombination rate, and fitness consequence scores, consistent with the stabilizing selection model of gene expression. [ABSTRACT FROM AUTHOR]

Abstract Background Natural selection can act on multiple genes in the same pathway, leading to polygenic adaptation. For example, adaptive changes were found to down-regulate six genes involved in ergosterol biosynthesis—an essential pathway targeted by many antifungal drugs—in some strains of the yeast Saccharomyces cerevisiae. However, the impact of this polygenic adaptation on metabolite levels was unknown. Here, we performed targeted mass spectrometry to measure the levels of eight metabolites in this pathway in 74 yeast strains from a genetic cross. Results Through quantitative trait locus (QTL) mapping we identified 19 loci affecting ergosterol pathway metabolite levels, many of which overlap loci that also impact gene expression within the pathway. We then used the recently developed v-test, which identified selection acting upon three metabolite levels within the pathway, none of which were predictable from the gene expression adaptation. Conclusions These data showed that effects of selection on metabolite levels were complex and not predictable from gene expression data. This suggests that a deeper understanding of metabolism is necessary before we can understand the impacts of even relatively straightforward gene expression adaptations on metabolic pathways.

Polygenic adaptation is thought to be ubiquitous, yet remains poorly understood. Here, we model this process analytically, in the plausible setting of a highly polygenic, quantitative trait that experiences a sudden shift in the fitness optimum. We show how the mean phenotype changes over time, depending on the effect sizes of loci that contribute to variance in the trait, and characterize the allele dynamics at these loci. Notably, we describe the two phases of the allele dynamics: The first is a rapid phase, in which directional selection introduces small frequency differences between alleles whose effects are aligned with or opposed to the shift, ultimately leading to small differences in their probability of fixation during a second, longer phase, governed by stabilizing selection. As we discuss, key results should hold in more general settings and have important implications for efforts to identify the genetic basis of adaptation in humans and other species.

• Biogeochemical niches vary among growth forms thriving in various forest strata. • Biogeochemical niches differ between degraded and mature forest species. • Biogeochemical niches are phylogenetically conserved during forest degradation. • Biogeochemical niches are converged for plant strata during forest degradation. Unveiling the ecological and evolutionary mechanisms that guide the differentiation of various plant growth forms into vertical forest strata during forest degradation continues to pose a challenge. We utilized the biogeochemical niche theory to decipher these mechanisms. We measured leaf concentrations of basal elements (C, N, P, K, Ca, and Mg) and used them to generate the biogeochemical niches in herbs, shrubs, trees, and climbers from different vertical strata within a degraded forest and its neighboring mature forest in subtropical montane zones. Our results showed that biogeochemical niches varied significantly among growth forms. Biogeochemical niches also differed significantly between species in the degraded and mature forests, as evidenced by the significantly lower leaf C concentration relative to other nutrients in degraded forest species. This could be attributable mainly to the expansive canopy openness in the degraded forest, facilitating the colonization of light-demanding self-supporters and climbers. The phylogenetic signals in leaf nutrient concentrations were significantly present across all species, suggesting the general similarities in biogeochemical niches among closely related species. Following evolutionary model comparisons, stabilizing selection towards varying optimal leaf nutrient concentrations for distinct growth forms in the degraded or mature forest emerged as the most suitable model. Our study thus highlights the important role of phylogenetic biogeochemical niche conservatism and stabilizing selection towards various optimal leaf nutrient compositions for diverse growth forms in both degraded and mature forests in the formation of vertical plant communities during subtropical forest degradation. [ABSTRACT FROM AUTHOR]

Andromonoecy, the presence of perfect and staminate flowers in the same individual, has evolved repeatedly in angiosperms. The staminate flowers are generally smaller than the perfect flowers in species that produce staminate flowers plastically when resources are limited. The smaller staminate flowers are expected to be less attractive to pollinators and have reduced size‐matching with pollinators than perfect flowers. We hypothesized that these potential disadvantages of staminate versus perfect flowers facilitate the evolution of sex‐specific floral morphology, such as allometric relationship between flower size and male reproductive organ.We compared six floral morphology traits, pollen production, pollinator visits and pollen removal from anthers between staminate and perfect flowers in several natural Commelina communis populations. Nectarless and zygomorphic C. communis flowers have polymorphic stamens with attracting, feeding and pollinating anthers and were visited by diverse pollinators.Staminate flowers were significantly smaller than perfect flowers, despite a large overlap in size between sexes. The lengths of pollinating stamens did not differ between staminate and perfect flowers, and staminate flowers produced significantly more pollen. We observed significantly more pollinator visits to perfect flowers than to staminate flowers. By contrast, pollen removal from pollinating stamens was significantly higher in staminate flowers than in perfect flowers.There is sexual dimorphism in flower morphology in C. communis. Staminate flowers with smaller attraction organs, similar pollinating stamens and higher pollen production assure higher pollen donor success relative to perfect flowers. Our results suggest that the morphological changes in staminate flowers enhance pollination success, even with limited resources. [ABSTRACT FROM AUTHOR]

Abstract The variance in phenotypic trait values is a product of environmental and genetic variation. The sensitivity of traits to environmental variation has a genetic component and is likely to be under selection. However, there are few studies investigating the evolution of this sensitivity, in part due to the challenges of estimating the environmental variance. The livestock literature provides a wealth of studies that accurately partition components of phenotypic variance, including the environmental variance, in well‐defined environments. These studies involve breeds that have been under strong selection on mean phenotype in optimal environments for many generations, and therefore represent an opportunity to study the potential evolution of trait sensitivity to environmental conditions. Here, we use literature on domestic cattle to examine the evolution of micro‐environmental variance (CVR—the coefficient of residual variance) by testing for differences in expression of CVR in animals from the same breed reared in different environments. Traits that have been under strong selection did not follow a null expectation of an increase in CVR in heterogenous environments (e.g., grazing), a pattern that may reflect evolution of increased uniformity in heterogeneous environments. When comparing CVR across environments of different levels of optimality, here measured by trait mean, we found a reduction in CVR in the more optimal environments for both life history and growth traits. Selection aimed at increasing trait means in livestock breeds typically occurs in the more optimal environments, and we therefore suspect that the decreased CVR is a consequence of evolution of the expression of micro‐environmental variance in this environment. Our results highlight the heterogeneity in micro‐environmental variance across environments and point to possible connections to the intensity of selection on trait means.

Ecological release is often attributed to the rapid adaptive diversification of phenotypic traits. However, it is not well understood how natural selection changes its strength and direction through the process of ecological release. Herein, we demonstrated how shell colour of the Japanese land snail Euhadra peliomphala simodae has diversified via a shift in natural selection due to ecological release after migration from the mainland to an island. This snail's shell colour diversified on the island due to disruptive selection after migration from the mainland. We used trail camera traps to identify the cause of natural selection on both the mainland and the island. We then conducted a mark-recapture experiment while collecting microhabitat use data. In total, we captured and marked around 1,700 snails on the mainland, some of which were preyed upon by an unknown predator. The trail camera traps showed that the predator is the large Japanese field mouse Apodemus speciosus, and the predatory frequency was higher on the mainland than on the island. However, this predation did not correlate with shell colour. Microhabitat use on the island was more extensive than on the mainland, with snails on the island using both ground and arboreal microhabitats. A Bayesian estimation showed that the stabilizing selection on shell colour came from factors other than predation. Our results suggest that the course of natural selection was modified due to ecological release after migration from the mainland, explaining one cause of the phenotypic diversification. [ABSTRACT FROM AUTHOR]

Background: Natural selection can act on multiple genes in the same pathway, leading to polygenic adaptation. For example, adaptive changes were found to down-regulate six genes involved in ergosterol biosynthesis--an essential pathway targeted by many antifungal drugs--in some strains of the yeast Saccharomyces cerevisiae. However, the impact of this polygenic adaptation on metabolite levels was unknown. Here, we performed targeted mass spectrometry to measure the levels of eight metabolites in this pathway in 74 yeast strains from a genetic cross. Results: Through quantitative trait locus (QTL) mapping we identified 19 loci affecting ergosterol pathway metabolite levels, many of which overlap loci that also impact gene expression within the pathway. We then used the recently developed v-test, which identified selection acting upon three metabolite levels within the pathway, none of which were predictable from the gene expression adaptation. Conclusions: These data showed that effects of selection on metabolite levels were complex and not predictable from gene expression data. This suggests that a deeper understanding of metabolism is necessary before we can understand the impacts of even relatively straightforward gene expression adaptations on metabolic pathways. [ABSTRACT FROM AUTHOR]

Spatial and temporal environmental variation can favor the evolution of adaptive phenotypic plasticity, such that genotypes alter their phenotypes in response to local conditions to maintain fitness across heterogeneous landscapes. When individuals show greater fitness in one habitat than another, asymmetric migration can restrict adaptation to the lower quality environment. In these cases, selection is predicted to favor traits that enhance fitness in the higher‐quality (source) habitat at the expense of fitness in the marginal (sink) habitat. Here, we test whether plasticity is adaptive in a system regulated by demographic source‐sink dynamics. Vaccinium elliottii (Ericaceae) occurs in dry upland and flood‐prone bottomland forests throughout the southeastern United States, but has larger populations and higher average individual fitness in upland sites. We conducted a multi‐year field experiment to evaluate whether plasticity in foliar morphology increases survival and lifespan. Both across and within habitats, selection favored plasticity in specific leaf area, stomatal density, and leaf size. Stabilizing selection acted on plasticity in stomatal density within habitats, suggesting that extreme levels of plasticity are disadvantageous. Thus, even in systems driven by source‐sink dynamics, temporal and spatial variation in conditions across the landscape and within habitat types can favor the evolution of plasticity. [ABSTRACT FROM AUTHOR]

Abstract Heritable variation in gene expression is common within species. Much of this variation is due to genetic differences outside of the gene with altered expression and is trans‐acting. This trans‐regulatory variation is often polygenic, with individual variants typically having small effects, making the genetic architecture and evolution of trans‐regulatory variation challenging to study. Consequently, key questions about trans‐regulatory variation remain, including the variability of trans‐regulatory variation within a species, how selection affects trans‐regulatory variation, and how trans‐regulatory variants are distributed throughout the genome and within a species. To address these questions, we isolated and measured trans‐regulatory differences affecting TDH3 promoter activity among 56 strains of Saccharomyces cerevisiae, finding that trans‐regulatory backgrounds varied approximately twofold in their effects on TDH3 promoter activity. Comparing this variation to neutral models of trans‐regulatory evolution based on empirical measures of mutational effects revealed that despite this variability in the effects of trans‐regulatory backgrounds, stabilizing selection has constrained trans‐regulatory differences within this species. Using a powerful quantitative trait locus mapping method, we identified ∼100 trans‐acting expression quantitative trait locus in each of three crosses to a common reference strain, indicating that regulatory variation is more polygenic than previous studies have suggested. Loci altering expression were located throughout the genome, and many loci were strain specific. This distribution and prevalence of alleles is consistent with recent theories about the genetic architecture of complex traits. In all mapping experiments, the nonreference strain alleles increased and decreased TDH3 promoter activity with similar frequencies, suggesting that stabilizing selection maintained many trans‐acting variants with opposing effects. This variation may provide the raw material for compensatory evolution and larger scale regulatory rewiring observed in developmental systems drift among species.

Abstract Reproductive synchrony is a widespread phenomenon that is predicted to be adaptive for prey with specialist predators but not for those with generalist ones. I tested this prediction in three polar seabird species characterized by different levels of predator specialization. In the Antarctic petrel, for which the only predator was highly specialized, hatching dates were highly synchronous and chicks that hatched close to the mean hatching date had a higher survival. In black‐legged kittiwakes and Brünnich's guillemots, whose predators were generalists, breeding was less synchronous and there was no fitness advantage in hatching close to the mean. This study emphasizes the potential importance of the relative timing of reproduction for individual fitness and supports the hypothesis that the adaptive value of breeding synchrony depends on the predator functional response.

Ultraconserved elements (UCEs) are stretches of hundreds of nucleotides with highly conserved cores flanked by variable regions. Although the selective forces responsible for the preservation of UCEs are unknown, they are nonetheless believed to contain phylogenetically meaningful information from deep to shallow divergence events. Phylogenetic applications of UCEs assume the same degree of rate heterogeneity applies across the entire locus, including variable flanking regions. We present a Wright–Fisher model of selection on nucleotides (SelON) which includes the effects of mutation, drift, and spatially varying, stabilizing selection for an optimal nucleotide sequence. The SelON model assumes the strength of stabilizing selection follows a position-dependent Gaussian function whose exact shape can vary between UCEs. We evaluate SelON by comparing its performance to a simpler and spatially invariant GTR+ Γ model using an empirical data set of 400 vertebrate UCEs used to determine the phylogenetic position of turtles. We observe much improvement in model fit of SelON over the GTR+ Γ model, and support for turtles as sister to lepidosaurs. Overall, the UCE-specific parameters SelON estimates provide a compact way of quantifying the strength and variation in selection within and across UCEs. SelON can also be extended to include more realistic mapping functions between sequence and stabilizing selection as well as allow for greater levels of rate heterogeneity. By more explicitly modeling the nature of selection on UCEs, SelON and similar approaches can be used to better understand the biological mechanisms responsible for their preservation across highly divergent taxa and long evolutionary time scales. [ABSTRACT FROM AUTHOR]

Phenotypic evolution through deep time is slower than expected from microevolutionary rates. This is the paradox of stasis. Previous models suggest stasis occurs because populations track adaptive peaks that remain relatively stable on million‐year intervals, raising the equally perplexing question of why these large changes are so rare. Here, we consider the possibility that peaks can move more rapidly than populations can adapt, resulting in extinction. We model peak movement with explicit population dynamics, parameterized with published microevolutionary estimates. Allowing extinction greatly increases the parameter space of peak movements that yield the appearance of stasis observed in real data through deep time. Extreme peak displacements, regardless of their frequency, will rarely result in an equivalent degree of trait evolution because of extinction. Thus, larger peak displacements will rarely be inferred using trait data from extant species or observed in fossil records. Our work highlights population ecology as an important contributor to macroevolutionary dynamics, presenting an alternative perspective on the paradox of stasis, where apparent constraint on phenotypic evolution in deep time reflects our restricted view of the subset of earth's lineages that were fortunate enough to reside on relatively stable peaks. [ABSTRACT FROM AUTHOR]

Stabilizing selection for lower variability of liveborn kits in a litter resulted in higher vitality and significantly (P < 0.05) higher mean number of weaned kits at 35 days of age. In the group of kits from selected mothers with higher variability per litter and coefficient of variation > 11 %, the mortality to weaning (35 days) and post-weaning (42 days) was 27.27 %. In the group from mothers with lower variability between litters and coefficient of variation < 11 %, the mortality within this period was zero. Females with lower variability of liveborn kits per litter and their kits at the 42nd day of age showed lower level of C-reactive protein (CRP) in blood plasma compared to the second group. Strong negative correlation (r = -0.795) was confirmed between the coefficient of variation of liveborn kits and the number of weaned kits with coefficient of determination r² = 0.633. In practice it means that higher balance or lower variability of the number of liveborn kits between the individual litters has positive influence on the number of weaned kits and this parameter was influenced on up to 63.3 %, while the rest of the influence (36.7 %) was entirely random. Strong negative correlation (r = -0.94) was noted also between the markers of milk yield in females up to the 21st day and the variation coefficient of liveborn kits. The determination coefficient in this case was r² = 0.884. These data suggest that higher stability of liveborn kits numbers between litters has a positive effect on the number of weaned kits of rabbits and also that targeted selection for lower levels of C-reactive protein in blood plasma can help to improve effective production through more effectively innate immunity against pathogens and therefore lower mortality of growing rabbits. [ABSTRACT FROM AUTHOR]

Populations can adapt to stressful environments through changes in gene expression. However, the role of gene regulation in mediating stress response and adaptation remains largely unexplored. Here, we use an integrative field dataset obtained from 780 plants of Oryza sativa ssp. indica (rice) grown in a field experiment under normal or moderate salt stress conditions to examine selection and evolution of gene expression variation under salinity stress conditions. We find that salinity stress induces increased selective pressure on gene expression. Further, we show that trans -eQTLs rather than cis -eQTLs are primarily associated with rice's gene expression under salinity stress, potentially via a few master-regulators. Importantly, and contrary to the expectations, we find that cis-trans reinforcement is more common than cis-trans compensation which may be reflective of rice diversification subsequent to domestication. We further identify genetic fixation as the likely mechanism underlying this compensation/reinforcement. Additionally, we show that cis - and trans -eQTLs are under different selection regimes, giving us insights into the evolutionary dynamics of gene expression variation. By examining genomic, transcriptomic, and phenotypic variation across a rice population, we gain insights into the molecular and genetic landscape underlying adaptive salinity stress responses, which is relevant for other crops and other stresses.

Inter-individual variation in gene expression has been shown to be heritable and is often associated with differences in disease susceptibility between individuals. Many studies focused on mapping associations between genetic and gene regulatory variation, yet much less attention has been paid to the evolutionary processes that shape the observed differences in gene regulation between individuals in humans or any other primate. To begin addressing this gap, we performed a comparative analysis of gene expression variability and expression quantitative trait loci (eQTLs) in humans and chimpanzees, using gene expression data from primary heart samples. We found that expression variability in both species is often determined by non-genetic sources, such as cell-type heterogeneity. However, we also provide evidence that inter-individual variation in gene regulation can be genetically controlled, and that the degree of such variability is generally conserved in humans and chimpanzees. In particular, we found a significant overlap of orthologous genes associated with eQTLs in both species. We conclude that gene expression variability in humans and chimpanzees often evolves under similar evolutionary pressures.

Among vertebrates, placental mammals are particularly variable in the covariance between cranial shape and body size (allometry), with rodents being a major exception. Australian murid rodents allow an assessment of the cause of this anomaly because they radiated on an ecologically diverse continent notably lacking other terrestrial placentals. Here, we use 3D geometric morphometrics to quantify species-level and evolutionary allometries in 38 species (317 crania) from all Australian murid genera. We ask whether ecological opportunity resulted in greater allometric diversity compared with other rodents or whether conserved allometry suggests intrinsic constraints and/or stabilizing selection. We also assess whether cranial shape variation follows the proposed rule of craniofacial evolutionary allometry (CREA), whereby larger species have relatively longer snouts and smaller braincases. To ensure we could differentiate parallel versus nonparallel species-level allometric slopes, we compared the slopes of rarefied samples across all clades. We found exceedingly conserved allometry and CREA-like patterns across the 10-million-year split between Mus and Australian murids. This could support both intrinsic-constraint and stabilizing-selection hypotheses for conserved allometry. Large-bodied frugivores evolved faster than other species along the allometric trajectory, which could suggest stabilizing selection on the shape of the masticatory apparatus as body size changes. [ABSTRACT FROM AUTHOR]

Genetic correlations between traits can strongly impact evolutionary responses to selection, and may thus impose constraints on adaptation. Theoretical and empirical work has made it clear that without strong linkage and with random mating, genetic correlations at evolutionary equilibrium result from an interplay of correlated pleiotropic effects of mutations, and correlational selection favoring combinations of trait values. However, it is not entirely clear how change in the overall strength of stabilizing selection across traits (breadth of the fitness peak, given its shape) influences this compromise between mutation and selection effects on genetic correlation. Here, we show that the answer to this question crucially depends on the intensity of genetic drift. In large, effectively infinite populations, genetic correlations are unaffected by the strength of selection, regardless of whether the genetic architecture involves common small‐effect mutations (Gaussian regime), or rare large‐effect mutations (House‐of‐Cards regime). In contrast in finite populations, the strength of selection does affect genetic correlations, by shifting the balance from drift‐dominated to selection‐dominated evolutionary dynamics. The transition between these domains depends on mutation parameters to some extent, but with a similar dependence of genetic correlation on the strength of selection. Our results are particularly relevant for understanding how senescence shapes patterns of genetic correlations across ages, and genetic constraints on adaptation during colonization of novel habitats. [ABSTRACT FROM AUTHOR]

Copyright of Revista Caatinga is the property of Revista Caatinga and its content may not be copied or emailed to multiple sites or posted to a listserv without the copyright holder's express written permission. However, users may print, download, or email articles for individual use. This abstract may be abridged. No warranty is given about the accuracy of the copy. Users should refer to the original published version of the material for the full abstract. (Copyright applies to all Abstracts.)

Aggression confers several fitness benefits including increased breeding opportunities and resource acquisition. Determining the relative contributions of genetic and environmental components to shaping aggression is essential for advancing our understanding of how selection affects the distribution of aggressive phenotypes in a population. In a From the Cover article in this issue of Molecular Ecology, vonHoldt et al. (2020) used RAD‐seq methods to obtain genome‐wide single nucleotide polymorphism (SNP) data to estimate heritability of interpack aggression of 141 North American grey wolves (Canis lupus) surveyed from 1995–2018. The authors inferred heritability using both a SNP‐based genetic relationship matrix (GRM) and a consensus pedigree informed by: (a) previously obtained microsatellite data; (b) past observations of parentage; and (c) statistical reconstruction of parent‐offspring pairs. SNP‐based (i.e., GRM) and pedigree‐based (i.e., consensus pedigree) heritability estimates were 37% and 14%, respectively, with an additional 14%–16% explained by natal pack effects. The study confirmed the previously discovered strong effects of relative pack size and breeding status on interpack aggression, illustrating how social dynamics and density‐dependent factors induce variance in aggressive behaviours. Finally, the authors found associations between average individual aggression scores (IAS) and specific candidate genes (MY09A and TRAK1). In sum, vonHoldt et al. (2020) provides an unprecedented and nuanced synthesis that not only suggests gene‐aggression associations, but also emphasizes how additive genetic variance and density‐dependent factors interact to maintain phenotypic variance in aggression over time. [ABSTRACT FROM AUTHOR]

The variance in phenotypic trait values is a product of environmental and genetic variation. The sensitivity of traits to environmental variation has a genetic component and is likely to be under selection. However, there are few studies investigating the evolution of this sensitivity, in part due to the challenges of estimating the environmental variance. The livestock literature provides a wealth of studies that accurately partition components of phenotypic variance, including the environmental variance, in well‐defined environments. These studies involve breeds that have been under strong selection on mean phenotype in optimal environments for many generations, and therefore represent an opportunity to study the potential evolution of trait sensitivity to environmental conditions. Here, we use literature on domestic cattle to examine the evolution of micro‐environmental variance (CVR—the coefficient of residual variance) by testing for differences in expression of CVR in animals from the same breed reared in different environments. Traits that have been under strong selection did not follow a null expectation of an increase in CVR in heterogenous environments (e.g., grazing), a pattern that may reflect evolution of increased uniformity in heterogeneous environments. When comparing CVR across environments of different levels of optimality, here measured by trait mean, we found a reduction in CVR in the more optimal environments for both life history and growth traits. Selection aimed at increasing trait means in livestock breeds typically occurs in the more optimal environments, and we therefore suspect that the decreased CVR is a consequence of evolution of the expression of micro‐environmental variance in this environment. Our results highlight the heterogeneity in micro‐environmental variance across environments and point to possible connections to the intensity of selection on trait means. [ABSTRACT FROM AUTHOR]

Pollinator-mediated selection might lead to among-trait differences in the degree and pattern of floral integration and intra-flower variation. To examine the patterns of intra-flower variation in floral traits, including nectar volume, we performed a field study using the zygomorphic flowers of Aconitum japonicum ssp. subcuneatum. We investigated (1) correlations between the sizes of the left and right sepals and petals, (2) variation in floral traits among plants, within plants and within flowers, (3) effects of sexual phases on floral integration variation in floral and nectar traits, and (4) the effect of size and intra-flower variation in traits of the left and right sepals and petals on pollen removal by pollinators. Lateral sepal area, but not lower sepal area, was highly correlated between the left and right sepals. Floral traits were more integrated during the male phase than during the female phase. Nectar standing crop in male-phase flowers correlated with helmet height and lateral and lower sepal area, but in female-phase flowers it only correlated with spur length. While intra-flower variance in lateral sepal area accounted for approximately 10% of the overall variance in these traits, the variance in lower sepal area accounted for 70% of the overall variance. Lateral sepal area had a negative effect on the number of pollen grains remaining after pollinator visits. Low variance in lateral sepals within flowers and measurements of pollen removal suggest that lateral sepals play a more important role in pollen export than the other traits. Left and right sepals may be the targets of selection for symmetry in zygomorphic flowers. [ABSTRACT FROM AUTHOR]

Background and aims: The drivers of variations in leaf nutrient concentrations in cave-dwelling plants remain poorly understood. We aimed to explore the effects of light, soil chemistry and phylogeny on leaf nutrient concentrations in cave-dwelling plants. Methods: We quantified light availability and sampled top-soils and leaves of the co-existing herbs and ferns in three caves. We used the traditional and phylogenetic comparative methods to determine the effects of light, soil chemistry and phylogeny on leaf nutrient concentrations and the cross-species correlations between leaf nutrients. Results: Leaf nutrient concentrations differed little among caves due to the non-significant relationships of leaf nutrient concentrations with light availability and soil nutrient concentrations across caves. The phylogenetic signals in leaf nutrient concentrations were significant for Ca, Mg and N but non-significant for the remaining nutrients. The evolutionary rates of leaf nutrient concentrations tended to increase with decreasing phylogenetic signals and were faster in herbs than ferns. These contrasting degrees of phylogenetic conservatism in leaf nutrient concentrations were best generated by Ornstein-Uhlenbeck models, i.e., stabilizing selection towards an optimum across species for P, K, S, Fe, Mn and Zn or higher optimal concentrations in herbs than ferns for Ca, Mg and N. Strong cross-species correlations between leaf nutrient concentrations such as Ca vs Mg and N vs P were found. Conclusions: Leaf nutrient concentrations in cave-dwelling plants showed convergent adaptations to cave environments and presented contrasting degrees of phylogenetic conservatism to produce leaf nutritional diversity for the co-existing herbs and ferns in caves. [ABSTRACT FROM AUTHOR]

Schizophrenia is a debilitating and heritable mental disorder associated with lower reproductive success. However, the prevalence of schizophrenia is stable over populations and time, resulting in an evolutionary puzzle: how is schizophrenia maintained in the population, given its apparent fitness costs? One possibility is that increased genetic liability for schizophrenia, in the absence of the disorder itself, may confer some reproductive advantage. We assessed the correlation and causal effect of genetic liability for schizophrenia with number of children, age at first birth and number of sexual partners using data from the Psychiatric Genomics Consortium and UK Biobank. Linkage disequilibrium score regression showed little evidence of genetic correlation between genetic liability for schizophrenia and number of children (rg = 0.002, p = 0.84), age at first birth (rg = −0.007, p = 0.45) or number of sexual partners (rg = 0.007, p = 0.42). Mendelian randomization indicated no robust evidence of a causal effect of genetic liability for schizophrenia on number of children (mean difference: 0.003 increase in number of children per doubling in the natural log odds ratio of schizophrenia risk, 95% confidence interval (CI): −0.003 to 0.009, p = 0.39) or age at first birth (−0.004 years lower age at first birth, 95% CI: −0.043 to 0.034, p = 0.82). We find some evidence of a positive effect of genetic liability for schizophrenia on number of sexual partners (0.165 increase in the number of sexual partners, 95% CI: 0.117–0.212, p = 5.30×10−10). These results suggest that increased genetic liability for schizophrenia does not confer a fitness advantage but does increase mating success.

Heritable variation in gene expression is common within species. Much of this variation is due to genetic differences outside of the gene with altered expression and is trans‐acting. This trans‐regulatory variation is often polygenic, with individual variants typically having small effects, making the genetic architecture and evolution of trans‐regulatory variation challenging to study. Consequently, key questions about trans‐regulatory variation remain, including the variability of trans‐regulatory variation within a species, how selection affects trans‐regulatory variation, and how trans‐regulatory variants are distributed throughout the genome and within a species. To address these questions, we isolated and measured trans‐regulatory differences affecting TDH3 promoter activity among 56 strains of Saccharomyces cerevisiae, finding that trans‐regulatory backgrounds varied approximately twofold in their effects on TDH3 promoter activity. Comparing this variation to neutral models of trans‐regulatory evolution based on empirical measures of mutational effects revealed that despite this variability in the effects of trans‐regulatory backgrounds, stabilizing selection has constrained trans‐regulatory differences within this species. Using a powerful quantitative trait locus mapping method, we identified ∼100 trans‐acting expression quantitative trait locus in each of three crosses to a common reference strain, indicating that regulatory variation is more polygenic than previous studies have suggested. Loci altering expression were located throughout the genome, and many loci were strain specific. This distribution and prevalence of alleles is consistent with recent theories about the genetic architecture of complex traits. In all mapping experiments, the nonreference strain alleles increased and decreased TDH3 promoter activity with similar frequencies, suggesting that stabilizing selection maintained many trans‐acting variants with opposing effects. This variation may provide the raw material for compensatory evolution and larger scale regulatory rewiring observed in developmental systems drift among species. [ABSTRACT FROM AUTHOR]