Tracing exploitation of egg boons: an experimental study using fatty acids and stable isotopes.

Coordinated spawning of marine animals releases millions of planktonic eggs into the environment, known as egg boons. Eggs are rich in essential fatty acids and may be an important lipid subsidy to egg consumers. Our aim was to validate the application of fatty acid and stable isotope tracers of egg consumption to potential egg consumers and to confirm egg consumption by the selected species. We conducted feeding experiments with ctenophores, crustaceans and fishes. We fed these animals a common diet of Artemia or a commercial feed (Otohime) and simulated egg boons for half of them by intermittently supplementing the common diet with red drum (Sciaenops ocellatus) eggs for 10-94 days. Controls did not receive eggs. Fatty acid profiles of consumers fed eggs were significantly different from those of controls 24 h after the last egg-feeding event. Consumers took on fatty acid characteristics of eggs. In fishes and ctenophores, fatty acid markers of egg consumption did not persist 2-5 days after the last egg-feeding event, but markers of egg consumption persisted in crustaceans for at least 5-10 days. Additionally, consumption of eggs, which had high values of δ15N, led to δ15N enrichment in crustaceans and a fish. We conclude that fatty acids and nitrogen stable isotope can be used as biomarkers of recent egg consumption in marine animals, validating their use for assessing exploitation of egg boons in nature.

Climate driven disruption of transitional alpine bumble bee communities.

Pollinators at high elevations face multiple threats from climate change including heat stress, failure to phenological match advancing flower resources and competitive pressure from range-expanding species of lower elevations. We conducted long-term multi-site surveys of alpine bumble bees to determine how phenology of range-stable and range-expanding species is responding to climate change. We ask whether bumble bee responses generate mismatches with floral resources, and whether these mismatches in turn promote community disruption and potential species replacement. In alpine environments of the central Rocky Mountains, range-stable and range-expanding bumble bees exhibit phenological mismatches with flowering host plants due to earlier flowering of preferred resources under warmer spring temperatures. However, workers of range-stable species are more canalised in their foraging schedules, exploiting a relatively narrow portion of the flowering season. Specifically, range-stable species show less variance in phenology in response to temporally and spatially changing conditions than range-expanding ones. Because flowering duration drives the seasonal abundance of floral resources at the landscape scale, we hypothesize that canalisation of phenology in alpine bumble bees could reduce their access to earlier or later season flowers. Warmer conditions are decreasing abundances of range-stable alpine bumble bees above the timberline, increasing abundance of range-expanding species, and facilitating a novel and more species-diverse bumble bee community. However, this trend is not explained by greater phenological mismatch of range-stable bees. Results suggest that conversion of historic habitats for cold-adapted alpine bumble bee species into refugia for more heat-tolerant congeners is disrupting bumble bee communities at high elevations, though the precise mechanisms accounting for these changes are not yet known. If warming continues, we predict that the transient increase in diversity due to colonization by historically low-elevation species will likely give way to declines of alpine bumble bees in the central Rocky Mountains.

Indications for rapid evolution of trait means and thermal plasticity in range-expanding populations of a butterfly.

Currently, poleward range expansions are observed in many taxa, often in response to anthropogenic climate change. At the expanding front, populations likely face cooler and more variable temperature conditions, imposing thermal selection. This may result in changes in trait means or plasticity, the relative contribution of which is not well understood. We, here, investigate evolutionary change in range-expanding populations of the butterfly Pieris mannii, by comparing populations from the core and the newly established northern range under laboratory conditions. We observed both changes in trait means and in thermal reaction norms. Range-expanding populations showed a more rapid development, potentially indicative of counter-gradient variation and an increased cold tolerance compared with core populations. Genotype-environment interactions prevailed in all associated traits, such that the above differences were restricted to cooler environmental conditions. In range-expanding populations, plasticity was decreased in developmental traits enabling relatively rapid growth even under cooler conditions but increased in cold tolerance arguably promoting higher activity under thermally challenging conditions. Notably, these changes must have occurred within a time period of ca. 10 years only. Our results suggest, in line with contemporary theory, that the evolution of plasticity may play a hitherto underestimated role for adaptation to climatic variation. However, rather than generally increased or decreased levels of plasticity, our results indicate fine-tuned, trait-specific evolutionary responses to increase fitness in novel environments.

The relative importance of plasticity versus genetic differentiation in explaining between population differences; a meta-analysis.

Both plasticity and genetic differentiation can contribute to phenotypic differences between populations. Using data on non-fitness traits from reciprocal transplant studies, we show that approximately 60% of traits exhibit co-gradient variation whereby genetic differences and plasticity-induced differences between populations are the same sign. In these cases, plasticity is about twice as important as genetic differentiation in explaining phenotypic divergence. In contrast to fitness traits, the amount of genotype by environment interaction is small. Of the 40% of traits that exhibit counter-gradient variation the majority seem to be hyperplastic whereby non-native individuals express phenotypes that exceed those of native individuals. In about 20% of cases plasticity causes non-native phenotypes to diverge from the native phenotype to a greater extent than if plasticity was absent, consistent with maladaptive plasticity. The degree to which genetic differentiation versus plasticity can explain phenotypic divergence varies a lot between species, but our proxies for motility and migration explain little of this variation.

Drop it like it's hot: Interpopulation variation in thermal phenotypes shows counter-gradient pattern.

Ectotherms utilise a complex array of behavioural and physiological mechanisms to cope with variation in suboptimal thermal environments. However, these mechanisms may be insufficient for population persistence under contemporary climate change, resulting in a greater need to understand how local populations respond to geographic variation in climate. In this study, we explored the potential for local adaptation and acclimation in thermal traits and behaviours using wild and captive populations of a small agamid lizard (the jacky lizard, Amphibolurus muricatus). We predicted that wild lizards from a high elevation site would have cooler thermal preferences compared to those at low elevation sites to match the more restricted thermal resources at higher, cooler elevations. We additionally explored whether variation in thermal traits was due to recent acclimation or fixed population differences, such as due to developmental plasticity or local adaptation. In contrast to our predictions, we found high-elevation lizards began panting at higher temperatures and had higher thermal preferences relative to lower elevation lizards. When allowed to bask freely, there was no difference in the intensity of basking or daily duration of time spent basking between lizards from different elevations. Although the high-elevation lizards appeared to show stronger acclimation to recent air temperatures compared to low-elevation lizards, this difference was not significant. Similarly, captive lizards acclimated under long and short basking regimes showed no major differences in thermal traits or basking behaviour. Our results are consistent with the presence of counter-gradient variation in thermal phenotypes of lizards, and suggest that these are driven by local adaptive responses or developmental effects rather than recent acclimation.

Little plant, big city: a test of adaptation to urban environments in common ragweed (Ambrosia artemisiifolia).

A full understanding of how cities shape adaptation requires characterizing genetically-based phenotypic and fitness differences between urban and rural populations under field conditions. We used a reciprocal transplant experiment with the native plant common ragweed, (Ambrosia artemisiifolia), and found that urban and rural populations have diverged in flowering time, a trait that strongly affects fitness. Although urban populations flowered earlier than rural populations, plants growing in urban field sites flowered later than plants in rural field sites. This counter-gradient variation is consistent adaptive divergence between urban and rural populations. Also consistent with local adaptation, both urban and rural genotypes experienced stronger net selection in the foreign than in the local habitat, but this pattern was not significant for male fitness. Despite the evidence for local adaptation, rural populations had higher lifetime fitness at all sites, suggesting that selection has been stronger or more uniform in rural than urban populations. We also found that inter-population differences in both flowering time and fitness tended to be greater among urban than rural populations, which is consistent with greater drift or spatial variation in selection within urban environments. In summary, our results are consistent with adaptive divergence of urban and rural populations, but also suggest there may be greater environmental heterogeneity in urban environments which also affects evolution in urban landscapes.

Global patterns and clines in the growth of common carp Cyprinus carpio.

This review provides a meta-analytical assessment of the global patterns and clines in the growth of Cyprinus carpio as measured by length-at-age (Lt ) or von Bertalanffy growth function (VBGF) parameters, mass-length relationship (W-Lt ) and condition factor, based on literature data. In total, 284 studies were retrieved spanning 91 years of research and carried out on 381 waterbodies-locations in 50 countries in all five continents. Although native C. carpio achieved larger (asymptotic) size relative to its non-native counterpart, the latter grew faster during the first 7 years of life. Lentic populations (especially in natural lakes) also achieved larger sizes relative to lotic ones and the same was true for populations in cold and temperate v. arid climates. Unlike previous studies (on much more restricted datasets), only weak latitudinal clines in instantaneous growth rate, Lt at age 3 and mortality were observed globally and this was probably due to the presence of counter-gradient growth variation at all representative age classes (i.e. 1-10 years). Slightly negative allometry was revealed by the W-Lt and the related form factor tended to distinguish the more elongated and torpedo-shaped body typical of the wild form from the deeper body of feral-domesticated C. carpio. Existing population dynamics models for C. carpio will benefit from the comprehensive range of waterbody type × climate class-specific VBGF parameters provided in the present study; whereas, more studies are needed on the species' growth in tropical regions and to unravel the possibility of confounding effects on age estimation due to both historical and methodological reasons.

Disentangling plasticity of serotiny, a key adaptive trait in a Mediterranean conifer.

PREMISE OF THE STUDY: Serotiny, the maintenance of ripe seeds in closed fruits or cones until fire causes dehiscence, is a key adaptive trait of plants in fire-prone ecosystems, but knowledge of phenotypic plasticity for cone retention in woody plants is extremely scarce. On the basis of published literature and our field observations, we hypothesized that increased aridity might decrease the aerial seed bank as a plastic response, not necessarily adaptive. METHODS: We used a Pinus halepensis common garden replicated in three contrasted sites (mild, cold, and dry) to separate population differentiation from phenotypic plasticity of cone serotiny and canopy cone bank (CCB). Differences in growth among trees of the same provenance allowed us to include size effect as a proxy of ontogenetic age for the same chronological age of the trees. KEY RESULTS: Tree size had a strong negative effect on serotiny, but serotiny degree differed among trial sites even after accounting for size effects. As hypothesized, serotiny was lower at the harsh (dry and cold) sites compared with the mild site. Genetic variation for size-dependent cone serotiny and significant population × site interaction were confirmed, the latter implying different plasticity of serotiny among populations. Population differentiation for CCB showed an ecotypic trend, with positive correlation with temperature oscillation (continentality) and negative correlation with summer rainfall. CONCLUSIONS: Growth-limiting environments exacerbated the precocious release of seeds, contrary to the ecotypic trend found for the aerial cone bank, suggesting a counter-gradient plasticity. This plastic response is potentially maladaptive under a scenario of frequent wildfires.

Elevating Air Temperature May Enhance Future Epidemic Risk of the Plant Pathogen Phytophthora infestans.

Knowledge of pathogen adaptation to global warming is important for predicting future disease epidemics and food production in agricultural ecosystems; however, the patterns and mechanisms of such adaptation in many plant pathogens are poorly understood. Here, population genetics combined with physiological assays and common garden experiments were used to analyze the genetics, physiology, and thermal preference of pathogen aggressiveness in an evolutionary context using 140 Phytophthora infestans genotypes under five temperature regimes. Pathogens originating from warmer regions were more thermophilic and had a broader thermal niche than those from cooler regions. Phenotypic plasticity contributed ~10-fold more than heritability measured by genetic variance. Further, experimental temperatures altered the expression of genetic variation and the association of pathogen aggressiveness with the local temperature. Increasing experimental temperature enhanced the variation in aggressiveness. At low experimental temperatures, pathogens from warmer places produced less disease than those from cooler places; however, this pattern was reversed at higher experimental temperatures. These results suggest that geographic variation in the thermal preferences of pathogens should be included in modeling future disease epidemics in agricultural ecosystems in response to global warming, and greater attention should be paid to preventing the movement of pathogens from warmer to cooler places.

Evidence of hatch-time based growth compensation in the early life history of two salmonid fishes.

Initial body size can indicate quality within-species, with large size increasing the likelihood of survival. However, some populations or individuals may have body size disadvantages due to spatial/temporal differences in temperature, photoperiod, or food. Across-populations, animals often have locally adapted physiology to compensate for relatively poor environmental influences on development and growth, while within-population individual behavioral adjustments can increase food intake after periods of deprivation and provide opportunities to catch up (growth compensation). Previous work has shown that growth compensation should include within-population differences related to short growing seasons due to delayed hatch time. We tested the hypothesis that individual fish that hatch later grow faster than those that hatch earlier. The relative magnitude of such a response was compared with growth variation among populations. We sampled young of the year Arctic charr and brook trout from five rivers in northern Labrador. Daily increments from otoliths were used to back-calculate size to a common age and calculate growth rates. Supporting the hypothesis, older fish were not larger at capture than younger fish because animals that hatched later grew faster, which may indicate age-based growth compensation.

Life histories as mosaics: Plastic and genetic components differ among traits that underpin life-history strategies.

Life-history phenotypes emerge from clusters of traits that are the product of genes and phenotypic plasticity. If the impact of the environment differs substantially between traits, then life histories might not evolve as a cohesive whole. We quantified the sensitivity of components of the life history to food availability, a key environmental difference in the habitat occupied by contrasting ecotypes, for 36 traits in fast- and slow-reproducing Trinidadian guppies. Our dataset included six putatively independent origins of the slow-reproducing, derived ecotype. Traits varied substantially in plastic and genetic control. Twelve traits were influenced only by food availability (body lengths, body weights), five only by genetic differentiation (interbirth intervals, offspring sizes), 10 by both (litter sizes, reproductive timing), and nine by neither (fat contents, reproductive allotment). Ecotype-by-food interactions were negligible. The response to low food was aligned with the genetic difference between high- and low-food environments, suggesting that plasticity was adaptive. The heterogeneity among traits in environmental sensitivity and genetic differentiation reveals that the components of the life history may not evolve in concert. Ecotypes may instead represent mosaics of trait groups that differ in their rate of evolution.

Counter-gradient variation of reproductive effort in a widely distributed temperate oak (Quercus petraea).

The genetic and phenotypic variability of life history traits determines the demographic attributes of tree populations and, thus, their responses to anthropogenic climate change. Growth- and survival-related traits have been widely studied in forest ecology, but little is known about the determinism of reproductive traits.Using an elevation gradient experiment in the Pyrenees we assessed the degree to which variations in reproductive effort along climatic gradients are environmentally or genetically driven, by comparing oak populations (Quercus petraea) growing under field and common garden conditions.In situ monitoring revealed a decline in reproductive effort with increasing elevation and decreasing temperature. In common garden conditions, significant genetic differentiation was observed between provenances for reproduction and growth: trees from cold environments (high elevations) grew more slowly, and produced larger acorns in larger numbers. Our observations show that genetic and phenotypic clines for reproductive traits have opposite signs (counter-gradient) along the environmental gradient as opposed to growth, for which genetic variation parallels phenotypic variation (co-gradient).The counter-gradient found here for reproductive effort reveals that genetic variation partly counteracts the phenotypic effect of temperature, moderating the change in reproductive effort according to temperature. We consider the possible contribution to this counter-gradient in reproductive effort as an evolutionary trade-off between reproduction and growth.

Counter-Gradient Momentum Transport Through Subtropical Shallow Convection in ICON-LEM Simulations.

It is well known that subtropical shallow convection transports heat and water vapor upwards from the surface. It is less clear if it also transports horizontal momentum upwards to significantly affect the trade winds in which it is embedded. We utilize unique multiday large-eddy simulations run over the tropical Atlantic with ICON-LEM to investigate the character of shallow convective momentum transport (CMT). For a typical trade-wind profile during boreal winter, CMT acts as an apparent friction to decelerate the north-easterly flow. This effect maximizes below the cloud base while in the cloud layer, friction is very small, although present over a relatively deep layer. In the cloud layer, the zonal component of the momentum flux is counter-gradient and penetrates deeper than reported in traditional shallow cumulus LES cases. The transport through conditionally sampled convective updrafts and downdrafts explains a weak friction effect, but not the counter-gradient flux near the cloud tops. The analysis of the momentum flux budget reveals that, in the cloud layer, the counter-gradient flux is driven by convectively triggered nonhydrostatic pressure-gradients and horizontal circulations surrounding the clouds. A model set-up with large domain size and realistic boundary conditions is necessary to resolve these effects.

Behavioral constraints on local adaptation and counter-gradient variation: Implications for climate change.

Resource allocation to growth, reproduction, and body maintenance varies within species along latitudinal gradients. Two hypotheses explaining this variation are local adaptation and counter-gradient variation. The local adaptation hypothesis proposes that populations are adapted to local environmental conditions and are therefore less adapted to environmental conditions at other locations. The counter-gradient variation hypothesis proposes that one population out performs others across an environmental gradient because its source location has greater selective pressure than other locations. Our study had two goals. First, we tested the local adaptation and counter-gradient variation hypotheses by measuring effects of environmental temperature on phenotypic expression of reproductive traits in the burying beetle, Nicrophorus orbicollis Say, from three populations along a latitudinal gradient in a common garden experimental design. Second, we compared patterns of variation to evaluate whether traits covary or whether local adaptation of traits precludes adaptive responses by others. Across a latitudinal range, N. orbicollis exhibits variation in initiating reproduction and brood sizes. Consistent with local adaptation: (a) beetles were less likely to initiate breeding at extreme temperatures, especially when that temperature represents their source range; (b) once beetles initiate reproduction, source populations produce relatively larger broods at temperatures consistent with their local environment. Consistent with counter-gradient variation, lower latitude populations were more successful at producing offspring at lower temperatures. We found no evidence for adaptive variation in other adult or offspring performance traits. This suite of traits does not appear to coevolve along the latitudinal gradient. Rather, response to selection to breed within a narrow temperature range may preclude selection on other traits. Our study highlights that N. orbicollis uses temperature as an environmental cue to determine whether to initiate reproduction, providing insight into how behavior is modified to avoid costly reproductive attempts. Furthermore, our results suggest a temperature constraint that shapes reproductive behavior.

Where is the optimum? Predicting the variation of selection along climatic gradients and the adaptive value of plasticity. A case study on tree phenology.

Many theoretical models predict when genetic evolution and phenotypic plasticity allow adaptation to changing environmental conditions. These models generally assume stabilizing selection around some optimal phenotype. We however often ignore how optimal phenotypes change with the environment, which limit our understanding of the adaptive value of phenotypic plasticity. Here, we propose an approach based on our knowledge of the causal relationships between climate, adaptive traits, and fitness to further these questions. This approach relies on a sensitivity analysis of the process-based model phenofit, which mathematically formalizes these causal relationships, to predict fitness landscapes and optimal budburst dates along elevation gradients in three major European tree species. Variation in the overall shape of the fitness landscape and resulting directional selection gradients were found to be mainly driven by temperature variation. The optimal budburst date was delayed with elevation, while the range of dates allowing high fitness narrowed and the maximal fitness at the optimum decreased. We also found that the plasticity of the budburst date should allow tracking the spatial variation in the optimal date, but with variable mismatch depending on the species, ranging from negligible mismatch in fir, moderate in beech, to large in oak. Phenotypic plasticity would therefore be more adaptive in fir and beech than in oak. In all species, we predicted stronger directional selection for earlier budburst date at higher elevation. The weak selection on budburst date in fir should result in the evolution of negligible genetic divergence, while beech and oak would evolve counter-gradient variation, where genetic and environmental effects are in opposite directions. Our study suggests that theoretical models should consider how whole fitness landscapes change with the environment. The approach introduced here has the potential to be developed for other traits and species to explore how populations will adapt to climate change.

The evolution of phenotypic plasticity when environments fluctuate in time and space.

Most theoretical studies have explored the evolution of plasticity when the environment, and therefore the optimal trait value, varies in time or space. When the environment varies in time and space, we show that genetic adaptation to Markovian temporal fluctuations depends on the between-generation autocorrelation in the environment in exactly the same way that genetic adaptation to spatial fluctuations depends on the probability of philopatry. This is because both measure the correlation in parent-offspring environments and therefore the effectiveness of a genetic response to selection. If the capacity to genetically respond to selection is stronger in one dimension (e.g., space), then plasticity mainly evolves in response to fluctuations in the other dimension (e.g., time). If the relationships between the environments of development and selection are the same in time and space, the evolved plastic response to temporal fluctuations is useful in a spatial context and genetic differentiation in space is reduced. However, if the relationships between the environments of development and selection are different, the optimal level of plasticity is different in the two dimensions. In this case, the plastic response that evolves to cope with temporal fluctuations may actually be maladaptive in space, resulting in the evolution of hyperplasticity or negative plasticity. These effects can be mitigated by spatial genetic differentiation that acts in opposition to plasticity resulting in counter-gradient variation. These results highlight the difficulty of making space-for-time substitutions in empirical work but identify the key parameters that need to be measured in order to test whether space-for-time substitutions are likely to be valid.

Systems biology derived source-sink mechanism of BMP gradient formation.

A morphogen gradient of Bone Morphogenetic Protein (BMP) signaling patterns the dorsoventral embryonic axis of vertebrates and invertebrates. The prevailing view in vertebrates for BMP gradient formation is through a counter-gradient of BMP antagonists, often along with ligand shuttling to generate peak signaling levels. To delineate the mechanism in zebrafish, we precisely quantified the BMP activity gradient in wild-type and mutant embryos and combined these data with a mathematical model-based computational screen to test hypotheses for gradient formation. Our analysis ruled out a BMP shuttling mechanism and a bmp transcriptionally-informed gradient mechanism. Surprisingly, rather than supporting a counter-gradient mechanism, our analyses support a fourth model, a source-sink mechanism, which relies on a restricted BMP antagonist distribution acting as a sink that drives BMP flux dorsally and gradient formation. We measured Bmp2 diffusion and found that it supports the source-sink model, suggesting a new mechanism to shape BMP gradients during development.

Improving peak shapes with counter gradients in two-dimensional high performance liquid chromatography.

To achieve the greatest peak capacity in two-dimensional high performance liquid chromatography (2D-HPLC) a gradient should be operated in both separation dimensions. However, it is known that when an injection solvent that is stronger than the initial mobile phase composition is deleterious to peak performance, thus causing problems when cutting a portion from one gradient into another. This was overcome when coupling hydrophilic interaction with reversed phase chromatography by introducing a counter gradient that changed the solvent strength of the second dimension injection. It was found that an injection solvent composition of 20% acetonitrile in water gave acceptable results in one-dimensional simulations with an initial composition of 5% acetonitrile. When this was transferred to a 2D-HPLC separation of standards it was found that a marked improvement in peak shape was gained for the moderately retained analytes (phenol and dimethyl phthalate), some improvement for the weakly retained caffeine and very little change for the strongly retained n-propylbenzene and anthracene which already displayed good chromatographic profiles. This effect was transferred when applied to a 2D-HPLC separation of a coffee extract where the indecipherable retention profile was transformed to a successful application multidimensional chromatography with peaks occupying 71% of the separation space according to the geometric approach to factor analysis.