Divergence in DNA photorepair efficiency among genotypes from contrasting UV radiation environments in nature

Populations of organisms routinely face abiotic selection pressures, and a central goal of evolutionary biology is to understand the mechanistic underpinnings of adaptive phenotypes. Ultraviolet radiation (UVR) is one of earth's most pervasive environmental stressors, potentially damaging DNA in any organism exposed to solar radiation. We explored mechanisms underlying differential survival following UVR exposure in genotypes of the water flea Daphnia melanica derived from natural ponds of differing UVR intensity. The UVR tolerance of a D. melanica genotype from a high‐UVR habitat depended on the presence of visible and UV‐A light wavelengths necessary for photoenzymatic repair of DNA damage, a repair pathway widely shared across the tree of life. We then measured the acquisition and repair of cyclobutane pyrimidine dimers, the primary form of UVR‐caused DNA damage, in D. melanica DNA following experimental UVR exposure. We demonstrate that genotypes from high‐UVR habitats repair DNA damage faster than genotypes from low‐UVR habitats in the presence of visible and UV‐A radiation necessary for photoenzymatic repair, but not in dark treatments. Because differences in repair rate only occurred in the presence of visible and UV‐A radiation, we conclude that differing rates of DNA repair, and therefore differential UVR tolerance, are a consequence of variation in photoenzymatic repair efficiency. We then rule out a simple gene expression hypothesis for the molecular basis of differing repair efficiency, as expression of the CPD photolyase gene photorepair did not differ among D. melanica lineages, in both the presence and absence of UVR.     

Fitness cost from fluctuating ultraviolet radiation in Daphnia magna

Solar ultraviolet radiation (UVR) is an important environmental threat for organisms in aquatic systems, but its temporally variable nature makes the understanding of its effects ambiguous. The aim of our study was to assess potential fitness costs associated with fluctuating UVR in the aquatic zooplankter Daphnia magna. We investigated individual survival, reproduction and behaviour when exposed to different UVR treatments. Individuals exposed to fluctuating UVR, resembling natural variations in cloud cover, had the lowest fitness (measured as the number of offspring produced during their lifespan). By contrast, individuals exposed to the same, but constant UVR dose had similar fitness to control individuals (not exposed to UVR), but they showed a significant reduction in daily movement. The re-occurring threat response to the fluctuating UVR treatment thus had strong fitness costs for D. magna, and we found no evidence for plastic behavioural responses when continually being exposed to UVR, despite the regular, predictable exposure schedule. In a broader context, our results imply that depending on how variable a stressor is in nature, populations may respond with alternative strategies, a framework that could promote rapid population differentiation and local adaptation.     

Embryogenesis plasticity and the transmission of maternal effects in Daphnia pulex

Understanding how genetic, nongenetic, and environmental cues are integrated during development may be critical in understanding if, and how, organisms will respond to rapid environmental change. Normally, only post‐embryonic studies are possible. But in this study, we developed a real‐time, high‐throughput confocal microscope assay that allowed us to link Daphnia embryogenesis to offspring life history variation at the individual level. Our assay identified eight clear developmental phenotypes linked by seven developmental stages, the duration of which were correlated with the expression of specific offspring life history traits. Daphnia embryogenesis varied not only between clones reared in the same environment, but also within a single clone when mothers were of different ages or reared in different food environments. Our results support the hypothesis that Daphnia embryogenesis is plastic and can be altered by changes in maternal state or maternal environment. As well as furthering our understanding of the mechanisms underpinning parental effects, our assay may also have an industrial application if it can be used as a rapid ecotoxicological prescreen for testing the effect that pollutant doses have on offspring life histories traditionally assayed with a 21‐day Daphnia reproduction test.     

The oxidative costs of territory quality and offspring provisioning

The costs of reproduction are an important constraint that shapes the evolution of life histories, yet our understanding of the proximate mechanism(s) leading to such life‐history trade‐offs is not well understood. Oxidative stress is a strong candidate measure thought to mediate the costs of reproduction, yet empirical evidence supporting that increased reproductive investment leads to oxidative stress is equivocal. We investigated whether territory quality and offspring provisioning increase oxidative stress in male snow buntings (Plectrophenax nivalis) using a repeated sampling design. We show that arrival oxidative stress is not a constraint on territory quality or the number of offspring fledged. Nevertheless, owners of higher‐quality territories experienced an oxidative cost, with this cost increasing more rapidly in younger males. Males that provisioned offspring at a high rate also experienced increased oxidative stress. Together, these findings support the potential role of oxidative stress in mediating life‐history trade‐offs. Future work should consider that reproductive workload is not limited to offspring care, and other activities – including territory defence – may contribute significantly to the costs of reproduction.     

EVOLUTION OF TRADE-OFFS BETWEEN SEXUAL AND ASEXUAL PHASES AND THE ROLE OF REPRODUCTIVE PLASTICITY IN THE GENETIC ARCHITECTURE OF APHID LIFE HISTORIES

Life‐history theory postulates that evolution is constrained by trade‐offs (i.e., negative genetic correlations) among traits that contribute to fitness. However, in organisms with complex life cycles, trade‐offs may drastically differ between phases, putatively leading to different evolutionary trajectories. Here, we tested this possibility by examining changes in life‐history traits in an aphid species that alternates asexual and sexual reproduction in its life cycle. The quantitative genetics of reproductive and dispersal traits was studied in 23 lineages (genotypes) of the bird cherry‐oat aphid Rhopalosiphum padi, during both the sexual and asexual phases, which were induced experimentally under specific environmental conditions. We found large and significant heritabilities (broad‐sense) for all traits and several negative genetic correlations between traits (trade‐offs), which are related to reproduction (i.e., numbers of the various sexual or asexual morphs) or dispersal (i.e., numbers of winged or wingless morphs). These results suggest that R. padi exhibits lineage specialization both in reproductive and dispersal strategies. In addition, we found important differences in the structure of genetic variance–covariance matrices ( G ) between phases. These differences were due to two large, negative genetic correlations detected during the asexual phase only: (1) between fecundity and age at maturity and (2) between the production of wingless and winged parthenogenetic females. We propose that this differential expression in genetic architecture results from a reallocation scheme during the asexual phase, when sexual morphs are not produced. We also found significant G × E interaction and nonsignificant genetic correlations across phases, indicating that genotypes could respond independently to selection in each phase. Our results reveal a rather unique situation in which the same population and even the same genotypes express different genetic (co)variation under different environmental conditions, driven by optimal resource allocation criteria.     

Oxidative shielding and the cost of reproduction

Life‐history theory assumes that reproduction and lifespan are constrained by trade‐offs which prevent their simultaneous increase. Recently, there has been considerable interest in the possibility that this cost of reproduction is mediated by oxidative stress. However, empirical tests of this theory have yielded equivocal support. We carried out a meta‐analysis to examine associations between reproduction and oxidative damage across markers and tissues. We show that oxidative damage is positively associated with reproductive effort across females of various species. Yet paradoxically, categorical comparisons of breeders versus non‐breeders reveal that transition to the reproductive state is associated with a step‐change reduction in oxidative damage in certain tissues and markers. Developing offspring may be particularly sensitive to harm caused by oxidative damage in mothers. Therefore, such reductions could potentially function to shield reproducing mothers, gametes and developing offspring from oxidative insults that inevitably increase as a consequence of reproductive effort. According to this perspective, we hypothesise that the cost of reproduction is mediated by dual impacts of maternally‐derived oxidative damage on mothers and offspring, and that mothers may be selected to diminish such damage. Such oxidative shielding may explain why many existing studies have concluded that reproduction has little or no oxidative cost. Future advance in life‐history theory therefore needs to take account of potential transgenerational impacts of the mechanisms underlying life‐history trade‐offs.     

An examination of the potential effects of food limitation on the ultraviolet radiation (UVR) tolerance of Diaptomus minutus

• 1 A series of field and laboratory experiments was performed to test the possible influence of food limitation on UVR tolerance in the calanoid copepod Diaptomusminutus.
• 2 Food‐limited animals incubated at 0.5 m in a clear‐water lake for over three days showed no evidence for increased mortality compared to food‐replete animals when exposed to UVR.
• 3 Animals exposed to artificial UVR in the laboratory showed high mortality. There was no evidence that feeding improved UVR tolerance.
• 4 The results of these experiments show that UVR tolerance is not affected by a short‐term lack of energy or a lack of essential nutrients. However, effects of non‐nutritive dietary components such as carotenoids and mycosporine‐like amino acids cannot be excluded.

     

Context‐dependent effects of cold stress on behavioral,physiological, and life‐history traits of the red flour beetle

Animals are exposed in nature to a variety of stressors. While stress is generally harmful, mild stress can also be beneficial and contribute to reproduction and survival. We studied the effect of five cold shock events versus a single cold shock and a control group, representing three levels of stress (harsh, mild, and no stress), on behavioral, physiological, and life‐history traits of the red flour beetle (Tribolium castaneum, Herbst 1797). Beetles exposed to harsh cold stress were less active than a control group: they moved less and failed more frequently to detect a food patch. Their probability to mate was also lower. Beetle pairs exposed to harsh cold stress frequently failed to reproduce at all, and if reproducing, females laid fewer eggs, which were, as larvae in mid‐development, smaller than those in the control group. However, harsh cold stress led to improved female starvation tolerance, probably due to enhanced lipid accumulation. Harsh cold shock also improved tolerance to an additional cold shock compared to the control. Finally, a single cold shock event negatively affected fewer measured response variables than the harsh cold stress, but also enhanced neither starvation tolerance nor tolerance to an additional cold shock. The consequences of a harsher cold stress are thus not solely detrimental but might even enhance survival under stressful conditions. Under benign conditions, nevertheless, harsh stress impedes beetle performance. The harsh stress probably shifted the balance point of the survival‐reproduction trade‐off, a shift that did not take place following exposure to mild stress.     

Demographic processes underlying fitness restoration in bdelloid rotifers emerging from dehydration

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Contrasting patterns of density‐dependent selection at different life stages can create more than one fast–slow axis of life‐history variation

There has been much recent research interest in the existence of a major axis of life‐history variation along a fast–slow continuum within almost all major taxonomic groups. Eco‐evolutionary models of density‐dependent selection provide a general explanation for such observations of interspecific variation in the "pace of life." One issue, however, is that some large‐bodied long‐lived “slow” species (e.g., trees and large fish) often show an explosive “fast” type of reproduction with many small offspring, and species with “fast” adult life stages can have comparatively “slow” offspring life stages (e.g., mayflies). We attempt to explain such life‐history evolution using the same eco‐evolutionary modeling approach but with two life stages, separating adult reproductive strategies from offspring survival strategies. When the population dynamics in the two life stages are closely linked and affect each other, density‐dependent selection occurs in parallel on both reproduction and survival, producing the usual one‐dimensional fast–slow continuum (e.g., houseflies to blue whales). However, strong density dependence at either the adult reproduction or offspring survival life stage creates quasi‐independent population dynamics, allowing fast‐type reproduction alongside slow‐type survival (e.g., trees and large fish), or the perhaps rarer slow‐type reproduction alongside fast‐type survival (e.g., mayflies—short‐lived adults producing few long‐lived offspring). Therefore, most types of species life histories in nature can potentially be explained via the eco‐evolutionary consequences of density‐dependent selection given the possible separation of demographic effects at different life stages.     

A captive breeding experiment reveals no evidence of reproductive isolation among lineages of a polytypic poison frog

Reproductive isolation is central to the generation of biodiversity, yet a clear understanding of the contributions of alternative reproductive barriers to this process remains elusive. Studies of young lineages that have diverged in ecologically important traits can offer insights into the chronology and relative importance of various isolating mechanisms during speciation. In poison frogs (Dendrobatidae), within‐species lineages often differ dramatically in coloration, a trait subject to natural and sexual selection. Coloration in the strawberry poison frog (Oophaga pumilio) is particularly diverse and previous work suggests the potential for reproductive isolation. We used a captive breeding experiment to assess the extent of reproductive isolation among three allopatric, genetically distinct O. pumilio lineages that differ in coloration. We compared reproduction of within‐ and between‐lineage pairs, predicting that if lineages are isolated, within‐lineage pairs would be most successful. We also examined the fertility and productivity of F1 backcrosses of admixed offspring. We found no evidence suggesting behavioural pre‐zygotic or post‐zygotic reproductive isolation, indicating that isolation would not be maintained by intrinsic mechanisms in the event of secondary contact. Future work should address costs of between‐lineage matings exerted by extrinsic natural and/or sexual selection against admixed offspring.     

How clonal are clones? A quest for loss of heterozygosity during asexual reproduction in Daphnia magna

Due to the lack of recombination, asexual organisms are predicted to accumulate mutations and show high levels of within‐individual allelic divergence (heterozygosity); however, empirical evidence for this prediction is largely missing. Instead, evidence of genome homogenization during asexual reproduction is accumulating. Ameiotic crossover recombination is a mechanism that could lead to long genomic stretches of loss of heterozygosity (LOH) and unmasking of mutations that have little or no effect in heterozygous state. Therefore, LOH might be an important force for inducing variation among asexual offspring and may contribute to the limited longevity of asexual lineages. To investigate the genetic consequences of asexuality, here we used high‐throughput sequencing of Daphnia magna for assessing the rate of LOH over a single generation of asexual reproduction. Comparing parthenogenetic daughters with their mothers at several thousand genetic markers generated by restriction site‐associated DNA (RAD) sequencing resulted in high LOH rate estimation that largely overlapped with our estimates for the error rate. To distinguish these two, we Sanger re‐sequenced the top 17 candidate RAD‐loci for LOH, and all of them proved to be false positives. Hence, even though we cannot exclude the possibility that short stretches of LOH occur in genomic regions not covered by our markers, we conclude that LOH does not occur frequently during asexual reproduction in D. magna and ameiotic crossovers are very rare or absent. This finding suggests that clonal lineages of D. magna will remain genetically homogeneous at least over time periods typically relevant for experimental work.     

Competition and facilitation among fungal plant parasites affect their life-history traits

Multi-infections may result in either competitive exclusion or coexistence on the same host of pathogen genotypes belonging to the same or different species. Epidemiological consequences of multiple infections, particularly how the development and transmission of a pathogen can be modified by the presence of another pathogen, are well documented. However, understanding how life history strategies of each pathogen modulate co-infection outcomes remains quite elusive. To analyze how co-infection drives changes in life history traits and affects co-existence in epidemic pathogens, we infected detached pea stipules with two fungal species, Peyronellaea pinodes and Phoma medicaginis var. pinodella (considering two strains per species), part of the ascochyta blight complex but presenting different life history strategies. All pairwise combinations (including self-pairs) between two strains of each species were tested. Strains were inoculated simultaneously, but apart from one another on the stipule. For each strain, four life history traits were measured: incubation period, necrosis area six days after inoculation, latent period and offspring production. Results show that, in co-infection, when resources are highly allocated to lesion development, the time between inoculation and the appearance of reproduction structures (latent period) and offspring production decreased, and vice-versa relative to single infections. The direction and/or magnitude of these responses to co-infection depend on the co-infecting strains. Moreover, these changes were always higher in self-pairs than in mixed co-infections. These results suggest facilitation between co-infecting strains, resulting in the selection of an intermediate level of virulence (here measured as the lesion development) at the expense of pathogen offspring production. This strategy allows the development and reproduction of each co-infecting strain when sharing limited resources. However, the direction and strength of these life history traits variations in co-infection depend on the life history strategy of the co-infecting strains, with a clear difference between ‘opportunists', ‘scavengers' and ‘pioneer colonisers'.     

Interaction of COP1 and UVR8 regulates UV‐B‐induced photomorphogenesis and stress acclimation in Arabidopsis

The ultraviolet‐B (UV‐B) portion of the solar radiation functions as an environmental signal for which plants have evolved specific and sensitive UV‐B perception systems. The UV‐B‐specific UV RESPONSE LOCUS 8 (UVR8) and the multifunctional E3 ubiquitin ligase CONSTITUTIVELY PHOTOMORPHOGENIC 1 (COP1) are key regulators of the UV‐B response. We show here that uvr8‐null mutants are deficient in UV‐B‐induced photomorphogenesis and hypersensitive to UV‐B stress, whereas overexpression of UVR8 results in enhanced UV‐B photomorphogenesis, acclimation and tolerance to UV‐B stress. By using sun simulators, we provide evidence at the physiological level that UV‐B acclimation mediated by the UV‐B‐specific photoregulatory pathway is indeed required for survival in sunlight. At the molecular level, we demonstrate that the wild type but not the mutant UVR8 and COP1 proteins directly interact in a UV‐B‐dependent, rapid manner in planta. These data collectively suggest that UV‐B‐specific interaction of COP1 and UVR8 in the nucleus is a very early step in signalling and responsible for the plant's coordinated response to UV‐B ensuring UV‐B acclimation and protection in the natural environment.     

Thermal plasticity is independent of environmental history in an intertidal seaweed

Organisms inhabiting the intertidal zone have been used to study natural ecophysiological responses and adaptations to thermal stress because these organisms are routinely exposed to high‐temperature conditions for hours at a time. While intertidal organisms may be inherently better at withstanding temperature stress due to regular exposure and acclimation, they could be more vulnerable to temperature stress, already living near the edge of their thermal limits. Strong gradients in thermal stress across the intertidal zone present an opportunity to test whether thermal tolerance is a plastic or canalized trait in intertidal organisms. Here, we studied the intertidal pool‐dwelling calcified alga, Ellisolandia elongata, under near‐future temperature regimes, and the dependence of its thermal acclimatization response on environmental history. Two timescales of environmental history were tested during this experiment. The intertidal pool of origin was representative of long‐term environmental history over the alga's life (including settlement and development), while the pool it was transplanted into accounted for recent environmental history (acclimation over many months). Unexpectedly, neither long‐term nor short‐term environmental history, nor ambient conditions, affected photosynthetic rates in E. elongata. Individuals were plastic in their photosynthetic response to laboratory temperature treatments (mean 13.2°C, 15.7°C, and 17.7°C). Further, replicate ramets from the same individual were not always consistent in their photosynthetic performance from one experimental time point to another or between treatments and exhibited no clear trend in variability over experimental time. High variability in climate change responses between individuals may indicate the potential for resilience to future conditions and, thus, may play a compensatory role at the population or species level over time.     

Manipulating reproductive effort leads to changes in female reproductive scheduling but not oxidative stress

The trade‐off between reproductive investment and lifespan is the single most important concept in life‐history theory. A variety of sources of evidence support the existence of this trade‐off, but the physiological costs of reproduction that underlie this relationship remain poorly understood. The Free Radical Theory of Ageing suggests that oxidative stress, which occurs when there is an imbalance between the production of damaging Reactive Oxygen Species (ROS) and protective antioxidants, may be an important mediator of this trade‐off. We sought to test this theory by manipulating the reproductive investment of female mice (Mus musculus domesticus) and measuring the effects on a number of life history and oxidative stress variables. Females with a greater reproductive load showed no consistent increase in oxidative damage above females who had a smaller reproductive load. The groups differed, however, in their food consumption, reproductive scheduling and mean offspring mass. Of particular note, females with a very high reproductive load delayed blastocyst implantation of their second litter, potentially mitigating the costs of energetically costly reproductive periods. Our results highlight that females use strategies to offset particularly costly periods of reproduction and illustrate the absence of a simple relationship between oxidative stress and reproduction.     

Recovery and immune priming modulate the evolutionary trajectory of infection‐induced reproductive strategies

In response to parasite exposure, organisms from a variety of taxa undergo a shift in reproductive investment that may trade off with other life‐history traits including survival and immunity. By suppressing reproduction in favour of somatic and immunological maintenance, hosts can enhance the probability of survival and recovery from infection. By plastically enhancing reproduction through terminal investment, on the other hand, hosts under the threat of disease‐induced mortality could enhance their lifetime reproductive fitness through reproduction rather than survival. However, we know little about the evolution of these strategies, particularly when hosts can recover and even bequeath protection to their offspring. In this study, we develop a stochastic agent‐based model that competes somatic maintenance and terminal investment strategies as they trade off differentially with lifespan, parasite resistance, recovery and transgenerational immune priming. Our results suggest that a trade‐off between reproduction and recovery can drive directional selection for either terminal investment or somatic maintenance, depending on the cost of reproduction to lifespan. However, some conditions, such as low virulence with a high cost of reproduction to lifespan, can favour diversifying selection for the coexistence of both strategies. The introduction of transgenerational priming into the model favours terminal investment when all strategies are equally likely to produce primed offspring, but favours somatic maintenance if it confers even a slight priming advantage over terminal investment. Our results suggest that both immune priming and recovery may modulate the evolution of reproductive shift diversity and magnitude upon exposure to parasites.     

Heterogeneity in individual quality in birds: overall patterns and insights from a study on common terns

While life‐history theory predicts a tradeoff between reproduction and survival, positive covariance, indicative of heterogeneity in individual quality, is often reported among individuals from natural populations. We review longitudinal studies of wild bird populations that test the relationship between annual reproductive success and lifespan and find the majority to report a positive correlation, while none reports a negative correlation. Heterogeneity in individual quality in resource acquisition, masking resource‐based tradeoffs, therefore appears to be common in birds. Considering that there is little evidence for heritable variation in fitness, heterogeneity in individual quality among adults may be due to life‐long effects of developmental conditions. In a 20‐year case study on common terns Sterna hirundo, we test for life‐long effects of cohort quality and within‐cohort nest quality, but find no significant effects on long‐term proxies of quality. Since other studies do find strong life‐long effects of developmental conditions, we suggest that the brood reduction strategy adopted by common terns, causing the majority of offspring to die rapidly after hatching, efficiently reduces variation in offspring quality at independence. As such, a brood reduction strategy may contribute to reduced heterogeneity in adult survival in stochastic environments, both suggested to be more common and adaptive in long‐lived species. Further study is required to assess heterogeneity in individual reproduction, especially in relation to environmental stochasticity and species’ life‐history strategies, in order to assess whether the relative strength of selection in early and late life may indeed affect the magnitude of heterogeneity in individual quality over life, and how this is mediated by parent–offspring conflict.     

Revisiting the link between breeding effort and oxidative balance through field evaluation of two sympatric sibling insect species

The idea that oxidative stress could be a major force governing evolutionary trade‐offs has recently been challenged by experimental approaches in laboratory conditions, triggering extensive debates centered on theoretical and methodological issues. Here, we revisited the link between oxidative stress and reproduction by measuring multiple antioxidant and oxidative damages in wild‐caught females of two sibling weevil species (Curculio elephas, C. glandium). The strength of our study arised from (1) studied species that were sympatric and exploited similar resource, but displayed contrasting reproductive strategies and (2) individuals were sampled throughout adult life so as to relate oxidative status to breeding effort. We found that the short‐lived C. elephas sacrifices red‐ox homeostasis for immediate reproduction upon emergence as characterized by low antioxidant defenses and elevated oxidative damage. Comparatively, C. glandium massively invests in antioxidant and maintains low oxidative damage, which may contribute to their extended prereproductive period. Intriguingly, we also reveal, for the first time in a field study, an unexpected reactivation of antioxidant defenses with the onset of reproduction. Our results thus support the existence of a strong, but complex relationship between oxidative stress and life‐history evolution and highlight the need for a finer‐scale picture of antioxidant strategies.     

INTERACTIONS BETWEEN UV RADIATION AND TEMPERATURE LIMIT INFERENCES FROM SINGLE‐FACTOR EXPERIMENTS1

The global environment is changing. Substantial shifts in temperature, rainfall, cloud cover, and UV radiation (UVR) are all predicted as a result of anthropogenic activity. Although the actual and potential effects of changes in single environmental variables are being studied intensively, the interactive effects of multiple stressors have received little attention. Here we offer the first experimental evidence of interactive effects between UVR and temperature on germination and growth in multicellular organisms. To address the question of how temperature affects survival and growth of organisms in the presence of UVR, we exposed early life stages of two species of intertidal algae, Alaria marginata Postels et Ruprecht and Fucus gardneri Silva, to four levels of UVR at three temperatures for 56 h. PAR and day length (12:12‐h light:dark) were held constant across all treatments. UVR levels bracketed natural levels, and temperatures were within the range of ambient temperatures. Designated endpoints were germination rate and cell number, and we recorded mortality where survival was nil. Our results support the hypothesis that temperature mediates the net biological effect of UVR and vice versa. For instance, spores of A. marginata were able to survive and grow at 15° C at all UV levels and at 10° C in the absence of UVR but were unable to survive at 10° C in the presence of high levels of UVR. Our results suggest that the ability to predict the effects of global change hinges on understanding interactions among environmental variables, imposing strict limits on inferences made from single‐factor experiments.