Phenotypic plasticity is the capacity for an individual genotype to produce different phenotypes in response to environmental variation. Most traits are plastic, but the degree to which plasticity is ...adaptive or non-adaptive depends on whether environmentally induced phenotypes are closer or further away from the local optimum. Existing theories make conflicting predictions about whether plasticity constrains or facilitates adaptive evolution. Debate persists because few empirical studies have tested the relationship between initial plasticity and subsequent adaptive evolution in natural populations. Here we show that the direction of plasticity in gene expression is generally opposite to the direction of adaptive evolution. We experimentally transplanted Trinidadian guppies (Poecilia reticulata) adapted to living with cichlid predators to cichlid-free streams, and tested for evolutionary divergence in brain gene expression patterns after three to four generations. We find 135 transcripts that evolved parallel changes in expression within the replicated introduction populations. These changes are in the same direction exhibited in a native cichlid-free population, suggesting rapid adaptive evolution. We find 89% of these transcripts exhibited non-adaptive plastic changes in expression when the source population was reared in the absence of predators, as they are in the opposite direction to the evolved changes. By contrast, the remaining transcripts exhibiting adaptive plasticity show reduced population divergence. Furthermore, the most plastic transcripts in the source population evolved reduced plasticity in the introduction populations, suggesting strong selection against non-adaptive plasticity. These results support models predicting that adaptive plasticity constrains evolution, whereas non-adaptive plasticity potentiates evolution by increasing the strength of directional selection. The role of non-adaptive plasticity in evolution has received relatively little attention; however, our results suggest that it may be an important mechanism that predicts evolutionary responses to new environments.
Celotno besedilo
Dostopno za:
DOBA, IJS, IZUM, KILJ, KISLJ, NUK, PILJ, PNG, SAZU, SBMB, SIK, UILJ, UKNU, UL, UM, UPUK
Body size has long been recognized to play a key role in shaping species interactions. For example, while small species thrive in a diversity of environments, they typically lose aggressive contests ...for resources with larger species. However, numerous examples exist of smaller species dominating larger species during aggressive interactions, suggesting that the evolution of traits can allow species to overcome the competitive disadvantage of small size. If these traits accumulate as lineages diverge, then the advantage of large size in interspecific aggressive interactions should decline with increased evolutionary distance. We tested this hypothesis using data on the outcomes of 23,362 aggressive interactions among 246 bird species pairs involving vultures at carcasses, hummingbirds at nectar sources, and antbirds and woodcreepers at army ant swarms. We found the advantage of large size declined as species became more evolutionarily divergent, and smaller species were more likely to dominate aggressive contests when interacting with more distantly-related species. These results appear to be caused by both the evolution of traits in smaller species that enhanced their abilities in aggressive contests, and the evolution of traits in larger species that were adaptive for other functions, but compromised their abilities to compete aggressively. Specific traits that may provide advantages to small species in aggressive interactions included well-developed leg musculature and talons, enhanced flight acceleration and maneuverability, novel fighting behaviors, and traits associated with aggression, such as testosterone and muscle development. Traits that may have hindered larger species in aggressive interactions included the evolution of morphologies for tree trunk foraging that compromised performance in aggressive contests away from trunks, and the evolution of migration. Overall, our results suggest that fundamental trade-offs, such as those associated with body size, are more likely to break down over evolutionary time, changing the rules that govern species interactions and structure ecological communities.
Celotno besedilo
Dostopno za:
DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
The impact of anthropogenic climate change on terrestrial organisms is often predicted to increase with latitude, in parallel with the rate of warming. Yet the biological impact of rising ...temperatures also depends on the physiological sensitivity of organisms to temperature change. We integrate empirical fitness curves describing the thermal tolerance of terrestrial insects from around the world with the projected geographic distribution of climate change for the next century to estimate the direct impact of warming on insect fitness across latitude. The results show that warming in the tropics, although relatively small in magnitude, is likely to have the most deleterious consequences because tropical insects are relatively sensitive to temperature change and are currently living very close to their optimal temperature. In contrast, species at higher latitudes have broader thermal tolerance and are living in climates that are currently cooler than their physiological optima, so that warming may even enhance their fitness. Available thermal tolerance data for several vertebrate taxa exhibit similar patterns, suggesting that these results are general for terrestrial ectotherms. Our analyses imply that, in the absence of ameliorating factors such as migration and adaptation, the greatest extinction risks from global warming may be in the tropics, where biological diversity is also greatest.
Species richness is greatest in the tropics, and much of this diversity is concentrated in mountains. Janzen proposed that reduced seasonal temperature variation selects for narrower thermal ...tolerances and limited dispersal along tropical elevation gradients Janzen DH (1967) Am Nat 101:233–249. These locally adapted traits should, in turn, promote reproductive isolation and higher speciation rates in tropical mountains compared with temperate ones. Here, we show that tropical and temperate montane stream insects have diverged in thermal tolerance and dispersal capacity, two key traits that are drivers of isolation in montane populations. Tropical species in each of three insect clades have markedly narrower thermal tolerances and lower dispersal than temperate species, resulting in significantly greater population divergence, higher cryptic diversity, higher tropical speciation rates, and greater accumulation of species over time. Our study also indicates that tropical montane species, with narrower thermal tolerance and reduced dispersal ability, will be especially vulnerable to rapid climate change.
Closely related, ecologically similar species often segregate their distributions along environmental gradients of time, space, and resources, but previous research suggests diverse underlying ...causes. Here, we review reciprocal removal studies in nature that experimentally test the role of interactions among species in determining their turnover along environmental gradients. We find consistent evidence for asymmetric exclusion coupled with differences in environmental tolerance causing the segregation of species pairs, where a dominant species excludes a subordinate from benign regions of the gradient but is unable to tolerate challenging regions to which the subordinate species is adapted. Subordinate species were consistently smaller and performed better in regions of the gradient typically occupied by the dominant species compared with their native distribution. These results extend previous ideas contrasting competitive ability with adaptation to abiotic stress to include a broader diversity of species interactions (intraguild predation, reproductive interference) and environmental gradients, including gradients of biotic challenge. Collectively, these findings suggest that adaptation to environmental challenge compromises performance in antagonistic interactions with ecologically similar species. The consistency of this pattern across diverse organisms, environments, and biomes suggests generalizable processes structuring the segregation of ecologically similar species along disparate environmental gradients, a phenomenon that we propose should be named the competitive exclusion–tolerance rule.
Species' thermal tolerances are used to estimate climate vulnerability, but few studies consider the role of the hydric environment in shaping thermal tolerances. As environments become hotter and ...drier, organisms often respond by limiting water loss to lower the risk of desiccation; however, reducing water loss may produce trade‐offs that lower thermal tolerances if respiration becomes inhibited. Here, we measured the sensitivity of water loss rate and critical thermal maximum (CTmax) to precipitation in nature and laboratory experiments that exposed click beetles (Coleoptera: Elateridae) to acute‐ and long‐term humidity treatments. We also took advantage of their unique clicking behavior to characterize subcritical thermal tolerances. We found higher water loss rates in the dry acclimation treatment compared to the humid, and water loss rates were 3.2‐fold higher for individuals that had experienced a recent precipitation event compared to individuals that had not. Acute humidity treatments did not affect CTmax, but precipitation indirectly affected CTmax through its effect on water loss rates. Contrary to our prediction, we found that CTmax was negatively associated with water loss rate, such that individuals with high water loss rate exhibited a lower CTmax. We then incorporated the observed variation of CTmax into a mechanistic niche model that coupled leaf and click beetle temperatures to predict climate vulnerability. The simulations indicated that indices of climate vulnerability can be sensitive to the effects of water loss physiology on thermal tolerances; moreover, exposure to temperatures above subcritical thermal thresholds is expected to increase by as much as 3.3‐fold under future warming scenarios. The correlation between water loss rate and CTmax identifies the need to study thermal tolerances from a “whole‐organism” perspective that considers relationships between physiological traits, and the population‐level variation in CTmax driven by water loss rate complicates using this metric as a straightforward proxy of climate vulnerability.
Estimates of climate variability are often developed from a single perspective without considering the integration of multiple traits at the organismal level. We studied interactions between thermal tolerance and water loss physiology in a high‐latitude beetle and found that water loss rates (and prior exposure to precipitation) played an important role in shaping climate vulnerability. We developed biophysical simulations of beetles on leaves under future climate warming scenarios to demonstrate that water loss rates influence thermal tolerance to future climate change.
Predation can be an important agent of natural selection shaping parental care behaviours, and can also favour behavioural plasticity. Parent birds often decrease the rate that they visit the nest to ...provision offspring when perceived risk is high. Yet, the plasticity of such responses may differ among species as a function of either their relative risk of predation, or the mean rate of provisioning. Here, we report parental provisioning responses to experimental increases in the perceived risk of predation. We tested responses of 10 species of bird in north temperate Arizona and subtropical Argentina that differed in their ambient risk of predation. All species decreased provisioning rates in response to the nest predator but not to a control. However, provisioning rates decreased more in species that had greater ambient risk of predation on natural nests. These results support theoretical predictions that the extent of plasticity of a trait that is sensitive to nest predation risk should vary among species in accordance with predation risk.
Synopsis
The concept of trade-offs permeates our thinking about adaptive evolution because they are exhibited at every level of biological organization, from molecular and cellular processes to ...organismal and ecological functions. Trade-offs inevitably arise because different traits do not occur in isolation, but instead are imbedded within complex, integrated systems that make up whole organisms. The genetic and mechanistic underpinning of trade-offs can be found in the pleiotropic nodes that occur in the biological pathways shared between traits. Yet, often trade-offs are only understood as statistical correlations, limiting the ability to evaluate the interplay between how selection and constraint interact during adaptive evolution. Here, we first review the classic paradigms in which physiologists and evolutionary biologists have studied trade-offs and highlight the ways in which network and molecular pathway approaches unify these paradigms. We discuss how these approaches allow researchers to evaluate why trade-offs arise and how selection can act to overcome trait correlations and evolutionary constraints. We argue that understanding how the conserved molecular pathways are shared between different traits and functions provides a conceptual framework for evolutionary biologists, physiologists, and molecular biologists to meaningfully work together toward the goal of understanding why correlations and trade-offs occur between traits. We briefly highlight the melanocortin system and the hormonal control of osmoregulation as two case studies where an understanding of shared molecular pathways reveals why trade-offs occur between seemingly unrelated traits. While we recognize that applying such approaches poses challenges and limitations particularly in the context of natural populations, we advocate for the view that focusing on the biological pathways responsible for trade-offs provides a unified conceptual context accessible to a broad range of integrative biologists.
Estimates of organismal thermal tolerance are frequently used to assess physiological risk from warming, yet the assumption that these estimates are predictive of mortality has been called into ...question. We tested this assumption in the cold-water-specialist frog,
. For seven populations, we used dynamic experimental assays to measure tadpole critical thermal maximum (CTmax) and measured mortality from chronic thermal stress for 3 days at different temperatures. We tested the relationship between previously estimated population CTmax and observed mortality, as well as the strength of CTmax as a predictor of mortality compared to local stream temperatures capturing varying timescales. Populations with higher CTmax experienced significantly less mortality in the warmest temperature treatment (25°C). We also found that population CTmax outperformed stream temperature metrics as the top predictor of observed mortality. These results demonstrate a clear link between CTmax and mortality from thermal stress, contributing evidence that CTmax is a relevant metric for physiological vulnerability assessments.
The empirical study of natural selection reveals that adaptations often involve trade‐offs between competing functions. Because natural selection acts on whole organisms rather than isolated traits, ...adaptive evolution may be constrained by the interaction between traits that are functionally integrated. Yet, few attempts have been made to characterize how and when such constraints are manifested or whether they limit the adaptive divergence of populations. Here we examine the consequences of adaptive life‐history evolution on locomotor performance in the live‐bearing guppy. In response to increased predation from piscivorous fish, Trinidadian guppies evolve an increased allocation of resources toward reproduction. These populations are also under strong selection for rapid fast‐start swimming performance to evade predators. Because embryo development increases a female’s wet mass as she approaches parturition, an increased investment in reproductive allocation should impede fast‐start performance. We find evidence for adaptive but constrained evolution of fast‐start swimming performance in laboratory trials conducted on second‐generation lab‐reared fish. Female guppies from high‐predation localities attain a faster acceleration and velocity and travel a greater distance during fast‐start swimming trials. However, velocity and distance traveled decline more rapidly over the course of pregnancy in these same females, thus reducing the magnitude of divergence in swimming performance between high‐ and low‐predation populations. This functional trade‐off between reproduction and swimming performance reveals how different aspects of the phenotype are integrated and highlights the complexity of adaptation at the whole‐organism level.
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