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Beldade, P.

Publications and source records attributed to Beldade, P..

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Predicting and Analyzing the Response to Selection on Correlated Characters

The breeders equation generally provides robust predictions for the short-term evolution of single characters. When selection targets two or more characters simultaneously, there are often large discrepancies between predicted and observed responses. We assessed how well this standard model predicts responses to bivariate selection on wing color pattern characteristics in the tropical butterfly Bicyclus anynana. In separate laboratory selection experiments, two sets of serially repeated eyespots were subjected to ten generations of concerted and antagonistic selection for either size or color composition. We compared predicted and actual selection responses over successive generations, using the phenotypic data, selection differentials, and estimates of the genetic variance-covariance matrix G. We found differences in the precision of predictions between directions of selection but did not find any evidence of systematic biases in our predictions depending on the direction of selection. Our investigation revealed significant environmental effects on trait evolution across generations. When these were accounted for, predictions using the standard model improved considerably. In the experiment on eyespot size, secondary splitting of selection lines allowed the estimation of changes in G after nine generations of selection. Changes were not in general agreement with expectations from the breeders equation. A contour plot of prediction errors across trait space suggests that directional epistasis in the eyespot genotype-phenotype map might occur but estimates of changes in G are too model-dependent to verify whether they agree with that hypothesis. Altogether, our results underscore the need for quantitative genetics to investigate and estimate potential effects of multivariate non-linear genotype-phenotype maps and of environmental effects on G.

evolutionary biology

Genetic basis of thermal plasticity variation in Drosophila melanogaster body size

Body size is a quantitative trait that is closely associated to fitness and under the control of both genetic and environmental factors. While developmental plasticity for this and other traits is heritable and under selection, little is known about the genetic basis for variation in plasticity that can provide the raw material for its evolution. We quantified genetic variation for body size plasticity in Drosophila melanogaster by measuring thorax and abdomen length of females reared at two temperatures from a panel representing naturally segregating alleles, the Drosophila Genetic Reference Panel (DGRP). We found variation between genotypes for the levels and direction of thermal plasticity in size of both body parts. We then used a Genome-Wide Association Study (GWAS) approach to unravel the genetic basis of inter-genotype variation in body size plasticity, and used different approaches to validate selected QTLs and to explore potential pleiotropic effects. We found mostly \"private QTLs\", with little overlap between the candidate loci underlying variation in plasticity for thorax versus abdomen size, for different properties of the plastic response, and for size versus size plasticity. We also found that the putative functions of plasticity QTLs were diverse and that alleles for higher plasticity were found at lower frequencies in the target population. Importantly, a number of our plasticity QTLs have been targets of selection in other populations. Our data sheds light onto the genetic basis of inter-genotype variation in size plasticity that is necessary for its evolution.\n\nSignificance StatementThe environmental conditions under which development takes place can affect developmental outcomes and lead to the production of phenotypes adjusted to the environment adults will live in. This developmental plasticity, which can help organisms cope with environmental heterogeneity, is heritable and under selection. Plasticity can itself evolve, a process that will be partly dependent on the available genetic variation for this trait. Using a wild-derived D. melanogaster panel, we identified DNA sequence variants associated to variation in thermal plasticity for body size. We found that these variants correspond to a diverse set of gene functions. Furthermore, their effects differ between body parts and properties of the thermal response, which can, therefore, evolve independently. Our results shed new light onto a number of key questions about the long discussed genes for plasticity.

evolutionary biology

Seasonal plasticity for anti-predatory strategies: matching colour and colour preference for effective crypsis

Effective anti-predatory strategies typically require matching appearance and behavior in prey, and there are many compelling examples of behavioral repertoires that enhance the effectiveness of morphological defenses. When protective adult morphology is induced by developmental environmental conditions predictive of future predation risk, adult behavior should be adjusted accordingly to maximize predator avoidance. While behavior is typically strongly affected by the adult environment, developmental plasticity in adult behavior -- mediated by the same pre-adult environmental cues that affect morphology -- could ensure an effective match between anti-predatory morphology and behavior. The coordination of environmentally-induced responses may be especially important in populations exposed to predictable environmental fluctuations (e.g. seasonality). Here, we studied early and late life environmental effects on a suite of traits expected to work together for effective crypsis. We focused on wing color and background color preference in Bicyclus anynana, a model of developmental plasticity that relies on crypsis as a seasonal strategy for predator avoidance. Using a full-factorial design, we disentangled effects of developmental and adult ambient temperature on both appearance and behavior. We showed that developmental conditions affect both adult color and color preference, with temperatures that simulate natural dry season conditions leading to browner butterflies with a perching preference for brown backgrounds. This effect was stronger in females, especially when butterflies were tested at lower ambient temperatures. In contrast to the expectation that motionlessness enhances crypsis, we found no support for our hypothesis that the browner dry-season butterflies would be less active. We argue that the integration of developmental plasticity for morphological and behavioral traits might improve the effectiveness of seasonal anti-predatory strategies. IMPACT SUMMARY To avoid predation, prey rely on strategies that typically include a variety of morphological and behavioral characteristics working together to deceive or scare predators. While some protective traits are a constitutive property of the prey species (e.g. hedgehog spines), others are produced only when the risk of encountering predators is high. For example, Daphnia crustaceans develop protective helmets and spines when exposed to cues that signal predator presence. When anti-predator morphologies are environmentally-induced, or plastic, the responses of associated behavioral traits should also be environmentally-dependent to ensure that animals exhibit behavioral repertoires that match their appearance. We used the tropical butterfly Bicyclus anynana, the squinting bush brown, to study these coordinated responses. In its natural habitat, with alternating dry and wet seasons, this thermally plastic butterfly alternates between seasonal forms with distinct wing color patterns related to distinct strategies to avoid predation. Dry season individuals, which develop under cooler temperatures, have small pattern elements on their wings and are cryptic against the brown background of dry foliage. In contrast, wet season individuals have large ornamental pattern elements that deflect attacks from predators towards the wing margin and away from their vulnerable bodies. We tested whether the cooler temperatures of the dry season also influenced other traits that can presumably improve the effectiveness of camouflage. In particular, we found that both the ornamental colors (appearance) and the choice of resting background colors (behavior) were affected by the temperature experienced during development. We found that dry season temperatures lead to browner butterflies with a stronger preference to rest on brown backgrounds. While behavior is typically very flexible in relation to changes in the adult environment, a developmental imprint on adult behavior can help ensure an effective match between the cryptic appearance and background choice behavior.

evolutionary biology

Complex effects of day and night temperature fluctuations on thermally plastic traits in an experimental model of adaptive seasonal plasticity.

BackgroundChanges in development in response to seasonally variable environments can produce phenotypes adjusted to fluctuating seasonal conditions and help organisms cope with temporal heterogeneity. In contrast to what happens in natural situations, experimental studies of developmental plasticity typically use environmental factors held constant during development, precluding assessment of potential environment-by-environment interaction effects.\n\nResultsWe tested effects of circadian fluctuations in temperature on a series of thermally plastic traits in a model of adaptive seasonal plasticity, the butterfly Bicyclus anynana. Comparing phenotypes from individuals reared under two types of fluctuations (warmer days with cooler nights, and cooler days with warmer nights) and those reared under a constant temperature of the same daily average allowed us to identify complex patterns of response to day and night temperatures. We found evidence of additive-like effects (for body size), but also different types of \"dominance\"-type effects where one particular period of the light cycle (for development time) or one particular extreme temperature (for eyespot size) had a relatively larger contribution to phenotype expression. We also gathered evidence against the hypothesis that thermal plasticity in development time drives thermal plasticity in other traits.\n\nConclusionsCombined effects of fluctuating day and night temperatures include additive-like effects as well as different types of environmental-dominance interaction effects. Differences between plastic traits reveal independent responses to temperature, and possible independent assessment of temperature conditions. Our study underscores the importance of understanding how organisms integrate complex environmental information towards a complete understanding of natural phenotypic variation and of the potential impact of environmental change thereon.

evolutionary biology