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      Watching the days go by: Asymmetric regulation of caterpillar development by changes in photoperiod

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          Abstract

          Many insects possess the plastic ability to either develop directly to adulthood, or enter diapause and postpone reproduction until the next year, depending on environmental cues (primarily photoperiod) that signal the amount of time remaining until the end of the growth season. These two alternative pathways often differ in co‐adapted life‐history traits, for example, with slower development and larger size in individuals headed for diapause. The developmental timing of these differences may be of adaptive importance: If traits diverge early, the potential for phenotypic differences between the pathways is greater, whereas if traits diverge late, the risk may be lower of expressing a maladaptive phenotype if the selective environment changes during development. Here, we explore the effects of changes in photoperiodic information during life on pupal diapause and associated life‐history traits in the butterfly Pararge aegeria. We find that both pupal diapause and larval development rate are asymmetrically regulated: While exposure to long days late in life (regardless of earlier experiences) was sufficient to produce nondiapause development and accelerate larval development accordingly, more prolonged exposure to short days was required to induce diapause and slow down prediapause larval development. While the two developmental pathways diverged early in development, development rates could be partially reversed by altered environmental cues. Meanwhile, pathway differences in body size were more inflexible, despite emerging late in development. These results show how several traits may be shaped by the same environmental cue (photoperiod), but along subtly different ontogenies, into an integrated phenotype.

          Abstract

          In many insects, winter diapause and nondiapause development comprise two plastically induced, alternative life‐history strategies. By subjecting butterfly larvae to controlled daylength regimes, we show that the co‐adapted traits comprising these strategies (diapause decision, body size, and growth rate) follow different ontogenies, and are differentially sensitive to changes in information during development.

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          Meeting the energetic demands of insect diapause: nutrient storage and utilization.

          Insects in diapause characteristically feed very little or not at all, thus they are largely or totally dependent on energy reserves sequestered prior to the entry into diapause. Fats are the dominant reserve used during this period, but non-fat reserves are also important for some species, especially during certain phases of diapause. Metabolic depression, coupled with the low temperatures of winter, facilitates the economic utilization of reserves during the many months typical of most diapauses. Though many insects store additional lipid prior to the entry into diapause, our review of the literature indicates that this is not always the case. We provide evidence that interactions between nutrient storage and metabolism can influence the decision to enter diapause and determine how long to remain in diapause. In addition, the energy reserves expended during diapause have a profound effect on post-diapause fitness. Though the physiological and biochemical mechanisms that regulate nutrient homeostasis prior to and during diapause remain poorly known, we propose several mechanisms that have the potential to contribute to diapause-associated nutrient homeostasis. Potential players include insulin signaling, neuropeptide F, cGMP-kinase, AMP-activated protein kinase, and adipokinetic hormone.
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            Costs and limits of phenotypic plasticity

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              Evolution of Animal Photoperiodism

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                Author and article information

                Contributors
                olle.lindestad@gmail.com
                karl.gotthard@zoologi.su.se
                Journal
                Ecol Evol
                Ecol Evol
                10.1002/(ISSN)2045-7758
                ECE3
                Ecology and Evolution
                John Wiley and Sons Inc. (Hoboken )
                2045-7758
                18 March 2021
                May 2021
                : 11
                : 10 ( doiID: 10.1002/ece3.v11.10 )
                : 5402-5412
                Affiliations
                [ 1 ] Department of Zoology Stockholm University Stockholm Sweden
                Author notes
                [*] [* ] Correspondence

                Olle Lindestad and Karl Gotthard, Department of Zoology, Stockholm University, Stockholm, Sweden.

                Emails: olle.lindestad@ 123456gmail.com (O. L.); karl.gotthard@ 123456zoologi.su.se (K. G)

                Author information
                https://orcid.org/0000-0002-8226-3167
                Article
                ECE37433
                10.1002/ece3.7433
                8131801
                c48704e5-f456-443f-8836-1f2bdb0daa4f
                © 2021 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.

                This is an open access article under the terms of the http://creativecommons.org/licenses/by/4.0/ License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

                History
                : 17 February 2021
                : 16 December 2020
                : 18 February 2021
                Page count
                Figures: 6, Tables: 0, Pages: 11, Words: 8162
                Funding
                Funded by: Vetenskapsrådet , open-funder-registry 10.13039/501100004359;
                Award ID: 2010‐5341
                Award ID: 2017‐04500
                Categories
                Original Research
                Original Research
                Custom metadata
                2.0
                May 2021
                Converter:WILEY_ML3GV2_TO_JATSPMC version:6.0.2 mode:remove_FC converted:19.05.2021

                Evolutionary Biology
                alternative life‐history strategies,butterfly,developmental plasticity,diapause,phenotypic plasticity,photoperiodism

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