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      The evolution of phenotypic plasticity under global change

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          Abstract

          Marine ecosystems are currently in a state of flux, with ocean warming and acidification occurring at unprecedented rates. Phenotypic plasticity underpins acclimatory responses by shifting the mean phenotype in a population, which may buffer the negative effects of global change. However, little is known about how phenotypic plasticity evolves across multiple generations. We tested this by reciprocally-transplanting the polychaete Ophryotrocha labronica between control and global change scenarios (ocean warming and acidification in isolation and combined) over five generations. By comparing the reaction norms of four life-history traits across generations, we show that juvenile developmental rate in the combined scenario was the only trait that changed its plastic response across generations when transplanted back to control conditions, and that adaptive plasticity was conserved in most traits, despite significant levels of selection and strong declines in individual fitness in the multi-generational exposure. We suggest the change in level of plasticity in the combined scenario is caused by differential allocation of energy between the mean and the plasticity of the trait along the multigenerational exposure. The ability to maintain within-generational levels of plasticity under global change scenarios has important eco-evolutionary and conservation implications, which are examined under the framework of assisted evolution programs.

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          Adaptive versus non-adaptive phenotypic plasticity and the potential for contemporary adaptation in new environments

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            Non-adaptive plasticity potentiates rapid adaptive evolution of gene expression in nature.

            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.
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              The evolution of growth trajectories: what limits growth rate?

              According to life-history theory, growth rates are subject to strong directional selection due to reproductive and survival advantages associated with large adult body size. Yet, growth is commonly observed to occur at rates lower than the maximum that is physiologically possible and intrinsic growth rates often vary among populations. This implies that slower growth is favoured under certain conditions. Realized growth rate is thus the result of a compromise between the costs and advantages of growing rapidly, and the optimal rate of growth is not equivalent to the fundamental maximum rate. The ecological and evolutionary factors influencing growth rate are reviewed, with particular emphasis on how growth might be constrained by direct fitness costs. Costs of accelerating growth might contribute to the variance in fitness that is not attributable to age or size at maturity, as well as to the variation in life-history strategies observed within and among species. Two main approaches have been taken to study the fitness trade-offs relating to growth rate. First, environmental manipulations can be used to produce treatment groups with different rates of growth. Second, common garden experiments can be used to compare fitness correlates among populations with different intrinsic growth rates. Data from these studies reveal a number of potential costs for growth over both the short and long term. In order to acquire the energy needed for faster growth, animals must increase food intake. Accordingly, in many taxa, the major constraint on growth rate appears to arise from the trade-off between predation risk and foraging effort. However, growth rates are also frequently observed to be submaximal in the absence of predation, suggesting that growth trajectories also impact fitness via other channels, such as the reallocation of finite resources between growth and other traits and functions. Despite the prevalence of submaximal growth, even when predators are absent, there is surprisingly little evidence to date demonstrating predator-independent costs of growth acceleration. Evidence that does exist indicates that such costs may be most apparent under stressful conditions. Future studies should examine more closely the link between patterns of resource allocation to traits in the adult organism and lifetime fitness. Changes in body composition at maturation, for example, may determine the outcome of trade-offs between reproduction and survival or between early and late reproduction. A number of design issues for studies investigating costs of growth that are imposed over the long term are discussed, along with suggestions for alternative approaches. Despite these issues, identifying costs of growth acceleration may fill a gap in our understanding of life-history evolution: the relationships between growth rate, the environment, and fitness may contribute substantially to the diversification of life histories in nature. © 2010 The Author. Biological Reviews © 2010 Cambridge Philosophical Society.
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                Author and article information

                Contributors
                emma.gibbin@epfl.ch
                Journal
                Sci Rep
                Sci Rep
                Scientific Reports
                Nature Publishing Group UK (London )
                2045-2322
                8 December 2017
                8 December 2017
                2017
                : 7
                : 17253
                Affiliations
                [1 ]ISNI 0000 0001 2185 197X, GRID grid.265702.4, Département de Biologie Chimie et Géographie, , Université du Québec à Rimouski, ; 300 Allée des Ursulines, Rimouski, Québec G5L 3A1 Canada
                [2 ]ISNI 0000000121839049, GRID grid.5333.6, Laboratory for Biological Geochemistry, , School of Architecture, Civil and Environmental Engineering, École Polytechnique Fédérale de Lausanne (EPFL), ; Lausanne, Switzerland
                [3 ]ISNI 0000 0004 0474 1797, GRID grid.1011.1, College of Science and Engineering, , James Cook University, ; Townsville, 4811 Queensland Australia
                [4 ]ISNI 0000 0001 2169 1275, GRID grid.433534.6, Centre d’Ecologie Fonctionnelle et Evolutive (CEFE- CNRS), ; Montpellier, Cedex 5 UMR 5175 France
                Author information
                http://orcid.org/0000-0003-4224-9683
                Article
                17554
                10.1038/s41598-017-17554-0
                5722875
                29222433
                e21a41af-5f91-44c2-9c0e-94f5ed8c17f4
                © The Author(s) 2017

                Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/.

                History
                : 8 March 2017
                : 28 November 2017
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