An alternative mechanism for skeletal muscle dysfunction in long-term post-viral lung disease.

Chronic lung disease is often accompanied by disabling extrapulmonary symptoms, notably skeletal muscle dysfunction and atrophy. Moreover, the severity of respiratory symptoms correlates with decreased muscle mass and in turn lowered physical activity and survival rates. Previous models of muscle atrophy in chronic lung disease often modeled chronic obstructive pulmonary disease (COPD) and relied on cigarette smoke exposure and LPS stimulation, but these conditions independently affect skeletal muscle even without accompanying lung disease. Moreover, there is an emerging and pressing need to understand the extrapulmonary manifestations of long-term post-viral lung disease (PVLD) as found in COVID-19. Here, we examine the development of skeletal muscle dysfunction in the setting of chronic pulmonary disease caused by infection due to the natural pathogen Sendai virus using a mouse model of PVLD. We identify a significant decrease in myofiber size when PVLD is maximal at 49 days after infection. We find no change in the relative types of myofibers, but the greatest decrease in fiber size is localized to fast-twitch-type IIB myofibers based on myosin heavy chain immunostaining. Remarkably, all biomarkers of myocyte protein synthesis and degradation (total RNA, ribosomal abundance, and ubiquitin-proteasome expression) were stable throughout the acute infectious illness and chronic post-viral disease process. Together, the results demonstrate a distinct pattern of skeletal muscle dysfunction in a mouse model of long-term PVLD. The findings thereby provide new insights into prolonged limitations in exercise capacity in patients with chronic lung disease after viral infections and perhaps other types of lung injury.NEW & NOTEWORTHY Our study used a mouse model of post-viral lung disease to study the impact of chronic lung disease on skeletal muscle. The model reveals a decrease in myofiber size that is selective for specific types of myofibers and an alternative mechanism for muscle atrophy that might be independent of the usual markers of protein synthesis and degradation. The findings provide a basis for new therapeutic strategies to correct skeletal muscle dysfunction in chronic respiratory disease.

Keep your cool: Overwintering physiology in response to urbanization in the acorn ant, Temnothorax curvispinosus.

Winter presents a challenge for survival, yet temperate ectotherms have remarkable physiological adaptations to cope with low-temperature conditions. Under recent climate change, rather than strictly relaxing pressure on overwintering survival, warmer winters can instead disrupt these low-temperature trait-environment associations, with negative consequences for populations. While there is increasing evidence of physiological adaptation to contemporary warming during the growing season, the effects of winter warming on physiological traits are less clear. To address this knowledge gap, we performed a common garden experiment using relatively warm-adapted versus cold-adapted populations of the acorn ant, Temnothorax curvispinosus, sampled across an urban heat island gradient, to explore the effects of winter conditions on plasticity and evolution of physiological traits. We found no evidence of evolutionary divergence in chill coma recovery nor in metabolic rate at either of two test temperatures (4 and 10 °C). Although we found the expected plastic response of increased metabolic rate under the 10 °C acute test temperature as compared with the 4 °C test temperature, this plastic response, (i.e., the acute thermal sensitivity of metabolic rate), was not different across populations. Surprisingly, we found that winter-acclimated urban ant populations exhibited higher heat tolerance compared with rural ant populations, and that the magnitude of divergence was comparable to that observed among growing-season acclimated ants. Finally, we found no evidence of differences between populations with respect to changes in colony size from the beginning to the end of the overwintering experiment. Together, these findings indicate that despite the evolution of higher heat tolerance that is often accompanied by losses in low-temperature tolerance, urban acorn ants have retained several components of low-temperature physiological performance when assessed under ecologically relevant overwintering conditions. Our study suggests the importance of measuring physiological traits under seasonally-relevant conditions to understand the causes and consequences of evolutionary responses to contemporary warming.

Natural Selection on Adults Has Trait-Dependent Consequences for Juvenile Evolution in Dragonflies.

AbstractAlthough natural selection often fluctuates across ontogeny, it remains unclear what conditions enable selection in one life-cycle stage to shape evolution in others. Organisms that undergo metamorphosis are useful for addressing this topic because their highly specialized life-cycle stages cannot always evolve independently despite their dramatic life-history transition. Using a comparative study of dragonflies, we examined three conditions that are hypothesized to allow selection in one stage to affect evolution in others. First, we tested whether lineages with less dramatic metamorphosis (e.g., hemimetabolous insects) lack the capacity for stage-specific evolution. Rejecting this hypothesis, we found that larval body shape evolves independently from selection on adult shape. Next, we evaluated whether stage-specific evolution is limited for homologous and/or coadapted structures. Indeed, we found that selection for larger wings is associated with the evolution of coadapted larval sheaths that store developing wing tissue. Finally, we assessed whether stage-specific evolution is restricted for traits linked to a single biochemical pathway. Supporting this hypothesis, we found that species with more wing melanization in the adult stage have evolved weaker melanin immune defenses in the larval stage. Thus, our results collectively show that natural selection in one stage imposes trait-dependent constraints on evolution in others.

Myogenic Cell Expression of Intercellular Adhesion Molecule-1 Contributes to Muscle Regeneration after Injury.

This study investigated intercellular adhesion molecule-1 (ICAM-1), a membrane protein that mediates cell-to-cell adhesion and communication, as a mechanism through which the inflammatory response facilitates muscle regeneration after injury. Toxin-induced muscle injury to tibialis anterior muscles of wild-type mice caused ICAM-1 to be expressed by a population of satellite cells/myoblasts and myofibers. Myogenic cell expression of ICAM-1 contributed to the restoration of muscle structure after injury, as regenerating myofibers were more abundant and myofiber size was larger for wild-type compared with Icam1-/- mice during 28 days of recovery. Contrastingly, restoration of muscle function after injury was similar between the genotypes. ICAM-1 facilitated the restoration of muscle structure after injury through mechanisms involving the regulation of myofiber branching, protein synthesis, and the organization of nuclei within myofibers after myogenic cell fusion. These findings provide support for a paradigm in which ICAM-1 expressed by myogenic cells after muscle injury augments their adhesive and fusogenic properties, which, in turn, facilitates regenerative and hypertrophic processes that restore structure to injured muscle.

Physiological adaptation to cities as a proxy to forecast global-scale responses to climate change.

Cities are emerging as a new venue to overcome the challenges of obtaining data on compensatory responses to climatic warming through phenotypic plasticity and evolutionary change. In this Review, we highlight how cities can be used to explore physiological trait responses to experimental warming, and also how cities can be used as human-made space-for-time substitutions. We assessed the current literature and found evidence for significant plasticity and evolution in thermal tolerance trait responses to urban heat islands. For those studies that reported both plastic and evolved components of thermal tolerance, we found evidence that both mechanisms contributed to phenotypic shifts in thermal tolerance, rather than plastic responses precluding or limiting evolved responses. Interestingly though, for a broader range of studies, we found that the magnitude of evolved shifts in thermal tolerance was not significantly different from the magnitude of shift in those studies that only reported phenotypic results, which could be a product of evolution, plasticity, or both. Regardless, the magnitude of shifts in urban thermal tolerance phenotypes was comparable to more traditional space-for-time substitutions across latitudinal and altitudinal clines in environmental temperature. We conclude by considering how urban-derived estimates of plasticity and evolution of thermal tolerance traits can be used to improve forecasting methods, including macrophysiological models and species distribution modelling approaches. Finally, we consider areas for further exploration including sub-lethal performance traits and thermal performance curves, assessing the adaptive nature of trait shifts, and taking full advantage of the environmental thermal variation that cities generate.

A mother's legacy: the strength of maternal effects in animal populations.

Although mothers influence the traits of their offspring in many ways beyond the transmission of genes, it remains unclear how important such 'maternal effects' are to phenotypic differences among individuals. Synthesizing estimates derived from detailed pedigrees, we evaluated the amount of phenotypic variation determined by maternal effects in animal populations. Maternal effects account for half as much phenotypic variation within populations as do additive genetic effects. Maternal effects most greatly affect morphology and phenology but, surprisingly, are not stronger in species with prolonged maternal care than in species without. While maternal effects influence juvenile traits more than adult traits on average, they do not decline across ontogeny for behaviour or physiology, and they do not weaken across the life cycle in species without maternal care. These findings underscore maternal effects as an important source of phenotypic variation and emphasise their potential to affect many ecological and evolutionary processes.

Temperature shapes the costs, benefits and geographic diversification of sexual coloration in a dragonfly.

The environment shapes the evolution of secondary sexual traits by determining how their costs and benefits vary across the landscape. Given the thermal properties of dark coloration generally, temperature should crucially influence the costs, benefits and geographic diversification of many secondary sexual colour patterns. We tested this hypothesis using sexually selected wing coloration in a dragonfly. We find that greater wing coloration heats males - the magnitude of which improves flight performance under cool conditions but dramatically reduces it under warm conditions. In a colder region of the species' range, behavioural observations of a wild population show that these thermal effects translate into greater territorial acquisition on thermally variable days. Finally, geo-referenced photographs taken by citizen scientists reveal that this sexually selected wing coloration is dramatically reduced in the hottest portions of the species' range. Collectively, our results underscore temperature's capacity to promote and constrain the evolution of sexual coloration.

On the evolution of carry-over effects.

The environment experienced early in life often affects the traits that are developed after an individual has transitioned into new life stages and environments. Because the phenotypes induced by earlier environments are then screened by later ones, these 'carry-over effects' influence fitness outcomes across the entire life cycle. While the last two decades have witnessed an explosion of studies documenting the occurrence of carry-over effects, little attention has been given to how they adapt and diversify. To aid future research in this area, we present a framework for the evolution of carry-over effects. Carry-over effects can evolve in two ways. First, the expression of traits later in life may become more or less dependent on the developmental processes of earlier stages (e.g., 'adaptive decoupling'). Genetic correlations between life stages then either strengthen or weaken. Alternatively, those influential developmental processes that begin early in life may become more or less sensitive to that earlier environment. Here, plasticity changes in all the traits that share those developmental pathways across the whole life cycle. Adaptive evolution of a carry-over effect is governed by selection on the induced phenotypes in the later stage, and also by selection on any developmentally linked traits in the earlier life stage. When these selective pressures conflict, the evolution of the carry-over effect will be biased towards maximizing performance in the life stage with stronger selection. Because life stages often contribute unequally to total fitness, the strength of selection in any one stage depends on: (a) the relationship between the traits and the stage-specific fitness components (e.g., juvenile survival, adult mating success), and (b) the reproductive value of the life stage. Considering the evolution of carry-over effects reveals several intriguing features of the evolution of life histories and phenotypic plasticity more generally. For instance, carry-over effects that manifest as maladaptive plasticity in one life stage may represent an adaptive strategy for maximizing fitness in stages with stronger selection. Additionally, adaptation to novel environments encountered early in the life cycle may be faster in the presence of carry-over effects that influence sexually selected traits.

Obstructive sleep apnea negatively impacts objectively measured physical activity.

PURPOSE: Obesity and obstructive sleep apnea (OSA) are frequent comorbid conditions. The impact of OSA on objectively measured physical activity (PA), independent of obesity, is not clear. The purpose of this study is to examine the effect of OSA on PA measured via accelerometer. METHODS: Overweight-to-obese individuals were recruited and screened for the presence of OSA via portable diagnostic device and divided into an OSA (n = 35) and control group (n = 24). Daytime sleepiness was assessed with the Epworth Sleepiness Scale. Body composition was assessed with dual-energy X-ray absorptiometry. Subjects wore an accelerometer (Actigraph GT3X+, Actigraph Corp., Pensacola, FL) for a minimum of 4 and maximum of 7 days, including at least one weekend day. RESULTS: There were no group differences in body mass index (BMI) or daytime sleepiness. Waist and neck circumference were higher in the OSA group. The OSA group was significantly older than the control group. The OSA group had fewer steps, moderate intensity minutes, moderate-to-vigorous minutes, number of PA bouts per day (≥ moderate intensity PA for ≥ 10 consecutive minutes), and total number of PA bouts. When adjusted for age, the PA bout data was no longer significant. CONCLUSION: Individuals screened as likely possessing OSA were less physically active than individuals without OSA when measured through objective means. We found no group differences in daytime sleepiness, BMI, or percent fat, suggesting other mechanisms than obesity and sleepiness for this difference.

Urban heat islands advance the timing of reproduction in a social insect.

For many species, the timing of life cycle events is advancing under contemporary global climate change. However, much less is known regarding phenological shifts as a result of other sources of anthropogenic change, such as urban warming. In both cases, progress has been hampered by a focus on phenological traits such as the timing of emergence, rather than the phenology of more directly related fitness traits such as the timing of reproduction. Here we explore how urban heat island effects shape the timing of reproduction in an acorn-dwelling ant. We used a common garden experiment with acorn ants collected from three cities in the eastern United States along a latitudinal gradient and reared long-term in the laboratory under five temperature treatments. This allowed us to quantify the effects of temperature on reproductive phenology across three scales-a biogeographic temperature cline, three urban vs. rural temperature comparisons, and five laboratory rearing temperatures. At our northernmost and southernmost cities (spanning 6° of latitude), we found both urbanization and warmer laboratory rearing temperature significantly advanced reproductive phenology; ants from the lowest latitude city also had earlier reproductive phenology compared with the higher latitude cities. In the field, the differences in urban versus rural acorn ant reproductive phenology translate to approximately one month earlier reproduction in the urban populations. For insects with synchronous mating events, such as ants, shifts in the already short window of time to reproduce could limit mating across environments, potentially leading to reproductive isolation between urban and rural populations.

Remarkable insensitivity of acorn ant morphology to temperature decouples the evolution of physiological tolerance from body size under urban heat islands.

Environmental temperature can alter body size and thermal tolerance, yet the effects of temperature rise on the size-tolerance relationship remain unclear. Terrestrial ectotherms with larger body sizes typically exhibit greater tolerance of high (and low) temperatures. However, while warming tends to increase tolerance of high temperatures through phenotypic plasticity and evolutionary change, warming tends to decrease body size through these mechanisms and thus might indirectly contribute to worse tolerance of high temperatures. These contrasting effects of warming on body size, thermal tolerance, and their relationship are increasingly important in light of global climate change. Here, we used replicated urban heat islands to explore the size-tolerance relationship in response to warming. We performed a common garden experiment with a small acorn-dwelling ant species collected from urban and rural populations across three different cities and reared under five laboratory rearing temperatures from 21 to 29 °C. We found that acorn ant body size was remarkably insensitive to laboratory rearing temperature (ant workers exhibited no phenotypic plasticity in body size across rearing temperature) and among populations experiencing cooler rural versus warmer urban environmental temperatures (no evolved differences in body size between urban and rural populations). Further, this insensitivity of body size to temperature was highly consistent across each of the three cities we examined. Because body size was robust to temperature variation, previously described plastic and evolved shifts in heat (and cold) tolerance in acorn ant responses to urbanization were shown to be independent of shifts in body size. Indeed, genetic (colony-level) correlations between heat and cold tolerance traits and body size revealed no significant association between size and tolerance. Our results show how typical trait correlations, such as between size and thermal tolerance, might be decoupled as populations respond to contemporary environmental change.

Evolution of thermal tolerance and its fitness consequences: parallel and non-parallel responses to urban heat islands across three cities.

The question of parallel evolution-what causes it, and how common it is-has long captured the interest of evolutionary biologists. Widespread urban development over the last century has driven rapid evolutionary responses on contemporary time scales, presenting a unique opportunity to test the predictability and parallelism of evolutionary change. Here we examine urban evolution in an acorn-dwelling ant species, focusing on the urban heat island signal and the ant's tolerance of these altered urban temperature regimes. Using a common-garden experimental design with acorn ant colonies collected from urban and rural populations in three cities and reared under five temperature treatments in the laboratory, we assessed plastic and evolutionary shifts in the heat and cold tolerance of F1 offspring worker ants. In two of three cities, we found evolved losses of cold tolerance, and compression of thermal tolerance breadth. Results for heat tolerance were more complex: in one city, we found evidence of simple evolved shifts in heat tolerance in urban populations, though in another, the difference in urban and rural population heat tolerance depended on laboratory rearing temperature, and only became weakly apparent at the warmest rearing temperatures. The shifts in tolerance appeared to be adaptive, as our analysis of the fitness consequences of warming revealed that while urban populations produced more sexual reproductives under warmer laboratory rearing temperatures, rural populations produced fewer. Patterns of natural selection on thermal tolerances supported our findings of fitness trade-offs and local adaptation across urban and rural acorn ant populations, as selection on thermal tolerance acted in opposite directions between the warmest and coldest rearing temperatures. Our study provides mixed support for parallel evolution of thermal tolerance under urban temperature rise, and, importantly, suggests the promising use of cities to examine parallel and non-parallel evolution on contemporary time scales.

Larval body condition regulates predator-induced life-history variation in a dragonfly.

Organisms with complex life cycles commonly exhibit adaptive plasticity in the timing of transitions between life stages. While the threat of predation is predicted to induce earlier transitions, empirical support has been equivocal. When predation risk affects both the propensity to transition to the next life stage and the ability to reach the energetic thresholds necessary to complete the transition, only those individuals in the best physiological condition may be able to accelerate development and emerge earlier. To test this hypothesis, we followed uniquely marked dragonfly larvae (Pachydiplax longipennis) through emergence in pools where we factorially manipulated the presence of a large heterospecific predator (Anax junius) and cannibalism risk via conspecific size variation. Consistent with our hypothesis, high-condition larvae were more likely to emerge in the presence of the heterospecific predator than in its absence, and low-condition larvae were more likely to emerge in its absence than in its presence. Moreover, high-condition larvae emerged earlier when cannibalism risk was high than when it was low. Predation risk therefore has condition-dependent effects on emergence. As predation risk frequently affects resource accumulation, similar mechanisms across taxa could commonly underlie the incongruence between empirical results and theoretical expectations for predator-induced life-history variation.

Immune deployment increases larval vulnerability to predators and inhibits adult life-history traits in a dragonfly.

While deploying immune defences early in ontogeny can trade-off with the production and maintenance of other important traits across the entire life cycle, it remains largely unexplored how features of the environment shape the magnitude or presence of these lifetime costs. Greater predation risk during the juvenile stage may particularly influence such costs by (1) magnifying the survival costs that arise from any handicap of juvenile avoidance traits and/or (2) intensifying allocation trade-offs with important adult traits. Here, we tested for predator-dependent costs of immune deployment within and across life stages using the dragonfly, Pachydiplax longipennis. We first examined how larval immune deployment affected two traits associated with larval vulnerability to predators: escape distance and foraging under predation risk. Larvae that were induced to mount an immune response had shorter escape distances but lower foraging activity in the presence of predator cues. We also induced immune responses in larvae and reared them through emergence in mesocosms that differed in the presence of large predatory dragonfly larvae (Aeshnidae spp.). Immune-challenged larvae had later emergence overall and lower survival in pools with predators. Immune-challenged males were also smaller at emergence and developed less sexually selected melanin wing coloration, but these effects were independent of predator treatment. Overall, these results highlight how mounting an immune defence early in ontogeny can have substantial ecological and physiological costs that manifest both within and across life stages.

Trade-offs between larval survival and adult ornament development depend on predator regime in a territorial dragonfly.

Trade-offs between juvenile survival and the development of sexually selected traits can cause ontogenetic conflict between life stages that constrains adaptive evolution. However, the potential for ecological interactions to alter the presence or strength of these trade-offs remains largely unexplored. Antagonistic selection over the accumulation and storage of resources could be one common cause of environment-specific trade-offs between life stages: higher condition may simultaneously enhance adult ornament development and increase juvenile vulnerability to predators. We tested this hypothesis in an ornamented dragonfly (Pachydiplax longipennis). Higher larval body condition indeed enhanced the initial development of its intrasexually selected wing coloration, but was opposed by viability selection in the presence of large aeshnid predators. In contrast, viability selection did not oppose larval body condition in pools when aeshnids were absent, and was not affected when we manipulated cannibalism risk. Trade-offs between larval survival and ornament development, mediated through the conflicting effects of body condition, therefore occurred only under high predation risk. We additionally characterized how body condition influences several traits associated with predator avoidance. Although body condition did not affect burst distance, it did increase larval abdomen size, potentially making larvae easier targets for aeshnid predators. As high body condition similarly increases vulnerability to predators in many other animals, predator-mediated costs of juvenile resource accumulation could be a common, environment-specific limitation on the elaboration of sexually selected traits.

What Are the Environmental Determinants of Phenotypic Selection? A Meta-analysis of Experimental Studies.

Although many selection estimates have been published, the environmental factors that cause selection to vary in space and time have rarely been identified. One way to identify these factors is by experimentally manipulating the environment and measuring selection in each treatment. We compiled and analyzed selection estimates from experimental studies. First, we tested whether the effect of manipulating the environment on selection gradients depends on taxon, trait type, or fitness component. We found that the effect of manipulating the environment was larger when selection was measured on life-history traits or via survival. Second, we tested two predictions about the environmental factors that cause variation in selection. We found support for the prediction that variation in selection is more likely to be caused by environmental factors that have a large effect on mean fitness but not for the prediction that variation is more likely to be caused by biotic factors. Third, we compared selection gradients from experimental and observational studies. We found that selection varied more among treatments in experimental studies than among spatial and temporal replicates in observational studies, suggesting that experimental studies can detect relationships between environmental factors and selection that would not be apparent in observational studies.

The effect of acute aerobic exercise on hemostasis in obstructive sleep apnea.

PURPOSE: Individuals with obstructive sleep apnea (OSA) have an altered hemostatic balance; however, the exercise response is less described. The purpose of this study is to determine the hemostatic response after acute aerobic exercise in obstructive sleep apnea. METHODS: Eighteen males (nine OSA vs. nine controls) were recruited from the university and local community. Individuals with evidence of cardiovascular, pulmonary, or metabolic disease were excluded. An apnea-hypopnea index (AHI) of >5 was a criterion for OSA. Subjects performed a treadmill exercise test at 35 and 70% predicted VO2 reserve during the morning hours. Pre-exercise blood samples were obtained after 15 min supine rest and within 2 min following exercise. Repeated measures ANOVA were performed for factor VIII antigen, tissue plasminogen activator (tPA) antigen, tPA activity, and PAI-1 activity. Correlational analysis compared resting and post-exercise hemostatic factors with age, BMI, and AHI. RESULTS: Mean AHI was 13.00 ± 12.6. No exercise × condition interactions were observed for hemostatic markers. There was a main effect for exercise in factor VIII, tPA antigen, and tPA activity in both groups. PAI-1 activity tended to be elevated in OSA (145%) compared to controls which remained after exercise (205%) (P = 0.05). Post-exercise FVIII/Ag correlated with BMI (r = 0.52), while resting tPA/Ag correlated with AHI (r = 0.49) and age (r = 0.50). CONCLUSION: The hemostatic response after acute aerobic exercise is unaffected in mild OSA, although PAI-1 activity seems to be elevated, reducing fibrinolytic potential. BMI seems to correlate with FVIII/Ag, while tPA/Ag is associated with AHI and age.

The predictability and magnitude of life-history divergence to ecological agents of selection: a meta-analysis in livebearing fishes.

Environments causing variation in age-specific mortality - ecological agents of selection - mediate the evolution of reproductive life-history traits. However, the relative magnitude of life-history divergence across selective agents, whether divergence in response to specific selective agents is consistent across taxa and whether it occurs as predicted by theory, remains largely unexplored. We evaluated divergence in offspring size, offspring number, and the trade-off between these traits using a meta-analysis in livebearing fishes (Poeciliidae). Life-history divergence was consistent and predictable to some (predation, hydrogen sulphide) but not all (density, food limitation, salinity) selective agents. In contrast, magnitudes of divergence among selective agents were similar. Finally, there was a negative, asymmetric relationship between offspring-number and offspring-size divergence, suggesting greater costs of increasing offspring size than number. Ultimately, these results provide strong evidence for predictable and consistent patterns of reproductive life-history divergence and highlight the importance of comparing phenotypic divergence across species and ecological selective agents.

Choosy Cannibals Preferentially Consume Siblings with Relatively Low Fitness Prospects.

When an individual can selfishly cannibalize a relative or altruistically set it free, the benefits of altruism will be positively associated with the relative's fitness prospects (the benefits it receives from altruism). We tested the prediction that altruism should be preferentially directed toward high-quality relatives using larvae of the New Mexican spadefoot toad (Spea multiplicata), a species in which tadpoles plastically express omnivore and carnivore ecomorphs. In a no-choice design, we presented carnivores with sibling or nonsibling omnivores varying in developmental stage, which is positively associated with survival in this toad's ephemeral larval environment. There was a significant interaction between relatedness and developmental stage on the probability of cannibalism: carnivores were overall more likely to cannibalize less developed omnivores, but this effect was exaggerated when the potential victim was a sibling. This evidence that altruists favor relatives with high fitness prospects highlights the numerous factors shaping altruism's payoffs.

Sexual selection's impacts on ecological specialization: an experimental test.

In many species, individuals specialize on different resources, thereby reducing competition. Such ecological specialization can promote the evolution of alternative ecomorphs-distinct phenotypes adapted for particular resources. Elucidating whether and how this process is influenced by sexual selection is crucial for understanding how ecological specialization promotes the evolution of novel traits and, potentially, speciation between ecomorphs. We evaluated the population-level effects of sexual selection (as mediated by mate choice) on ecological specialization in spadefoot toad tadpoles that express alternative ecomorphs. We manipulated whether sexual selection was present or reversed by mating females to their preferred versus non-preferred males, respectively. We then exposed their tadpoles to resource competition in experimental mesocosms. The resulting distribution of ecomorphs was similar between treatments, but sexual selection generated poorer trait integration in, and lower fitness of, the more specialized carnivore morph. Moreover, disruptive and directional natural selection were weaker in the sexual selection present treatment. Nevertheless, this effect on disruptive selection was smaller than previously documented effects of ecological opportunity and competitor density. Thus, sexual selection can inhibit adaptation to resource competition and thereby hinder ecological specialization, particularly when females obtain fitness benefits from mate choice that offset the cost of producing competitively inferior offspring.