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      Expression of alternative developmental pathways in the cabbage butterfly, Pieris melete and their differences in life history traits

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          Abstract

          The seasonal life cycle of the cabbage butterfly, Pieris melete is complicated because there are three options for pupal development: summer diapause, winter diapause, and nondiapause. In the present study, we tested the influence of temperature, day length, and seasonality on the expression of alternative developmental pathways and compared the differences in life history traits between diapausing and directly developing individuals under laboratory and field conditions. The expression of developmental pathway strongly depended on temperature, day length, and seasonality. Low temperatures induced almost all individuals to enter diapause regardless of day length; relatively high temperatures combined with intermediate and longer day lengths resulted in most individuals developing without diapause in the laboratory. The field data revealed that the degree of phenotypic plasticity in relation to developmental pathway was much higher in autumn than in spring. Directly developing individuals showed shorter development times and higher growth rates than did diapausing individuals. The pupal and adult weights for both diapausing and directly developing individuals gradually decreased as rearing temperature increased, with the diapausing individuals being slightly heavier than the directly developing individuals at each temperature. Female body weight was slightly lower than male body weight. The proportional weight losses from pupa to adult were almost the same in diapausing individuals and in directly developing individuals, suggesting that diapause did not affect weight loss at metamorphosis. Our results highlight the importance of the expression of alternative developmental pathways, which not only synchronizes this butterfly's development and reproduction with the growth seasons of the host plants but also exhibits the bet‐hedging tactic against unpredictable risks due to a dynamic environment.

          Abstract

          The expression of developmental pathways in the cabbage butterfly, Pieris melete highly depended on temperature, day length, and seasonality and exhibited a bet‐hedging tactic. Directly developing individuals showed shorter development time, higher growth rate, and relatively lower body weight than diapausing individuals. There were no significant differences in weight losses between the direct and diapause development individuals. Our results highlight the importance of the expression of alternative developmental pathways in the evolution of life history.

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          Most cited references43

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          Intraspecific Variation in Body Size and Fecundity in Insects: A General Relationship

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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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              Risk-spreading and bet-hedging in insect population biology.

              K R Hopper (1999)
              In evolutionary ecology, risk-spreading (i.e. bet-hedging) is the idea that unpredictably variable environments favor genotypes with lower variance in fitness at the cost of lower arithmetic mean fitness. Variance in fitness can be reduced by physiology or behavior that spreads risk of encountering an unfavorable environment over time or space. Such risk-spreading can be achieved by a single phenotype that avoids risks (conservative risk-spreading) or by phenotypic variation expressed by a single genotype (diversified risk-spreading). Across these categories, three types of risk-spreading can be usefully distinguished: temporal, metapopulation, and within-generation. Theory suggests that temporal and metapopulation risk-spreading may work under a broad range of population sizes, but within-generation risk-spreading appears to work only when populations are small. Although genetic polymorphisms have sometimes been treated as risk-spreading, the underlying mechanisms are different, and they often require different conditions for their evolution and thus are better treated separately. I review the types of evidence that could be used to test for risk-spreading and discuss evidence for risk-spreading in facultative diapause, migration polyphenism, spatial distribution of oviposition, egg size, and other miscellaneous traits. Although risk-spreading theory is voluminous and well developed in some ways, rarely has it been used to generate detailed, testable hypotheses about the evolution of risk-spreading. Furthermore, although there is evidence for risk-spreading, particularly in facultative diapause, I have been unable to find any definitive tests with unequivocal results showing that risk-spreading has been a major factor in the evolution of insect behaviors or life histories. To advance our understanding of risk-spreading in the wild, we need (a) explicit empirical models that predict levels of diversifying risk-spreading for several insect populations in several environments that vary in uncertainty, and (b) tests of these models using measurements of phenotypes and their fitnesses over several generations in each environment.
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                Author and article information

                Contributors
                185859707@qq.com
                Journal
                Ecol Evol
                Ecol Evol
                10.1002/(ISSN)2045-7758
                ECE3
                Ecology and Evolution
                John Wiley and Sons Inc. (Hoboken )
                2045-7758
                21 October 2019
                November 2019
                : 9
                : 21 ( doiID: 10.1002/ece3.v9.21 )
                : 12311-12321
                Affiliations
                [ 1 ] College of Computer and Information Engineering Jiangxi Agricultural University Nanchang China
                [ 2 ] Institute of Entomology Jiangxi Agricultural University Nanchang China
                [ 3 ] Department of Entomology University of Georgia Tifton GA USA
                [ 4 ] School of Education Huazhong University of Science and Technology Wuhan China
                [ 5 ] Foreign Language School Jiangxi Agricultural University Nanchang China
                Author notes
                [*] [* ] Correspondence

                Lan Xiao, School of Education, Huazhong University of Science and Technology, Wuhan 430074, China and Foreign Language School, Jiangxi Agricultural University, Nanchang 330045, China.

                Email: 185859707@ 123456qq.com

                Author information
                https://orcid.org/0000-0003-0756-4135
                https://orcid.org/0000-0002-7609-7414
                https://orcid.org/0000-0003-4665-4440
                Article
                ECE35731
                10.1002/ece3.5731
                6854102
                bf4da9d6-e41d-4d36-9658-01c6be4b8021
                © 2019 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
                : 08 July 2019
                : 06 September 2019
                : 11 September 2019
                Page count
                Figures: 4, Tables: 4, Pages: 11, Words: 7529
                Funding
                Funded by: National Natural Science Foundation of the People's Republic of China
                Award ID: 31260430
                Categories
                Original Research
                Original Research
                Custom metadata
                2.0
                November 2019
                Converter:WILEY_ML3GV2_TO_JATSPMC version:5.7.1 mode:remove_FC converted:14.11.2019

                Evolutionary Biology
                bet‐hedging,diapause,direct development,life history trait,pieris melete,temperature

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