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      The strength of the association between heterozygosity and probability of interannual local recruitment increases with environmental harshness in blue tits

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          Abstract

          The extent of inbreeding depression and the magnitude of heterozygosity–fitness correlations ( HFC) have been suggested to depend on the environmental context in which they are assayed, but little evidence is available for wild populations. We combine extensive molecular and capture–mark–recapture data from a blue tit ( Cyanistes caeruleus) population to (1) analyze the relationship between heterozygosity and probability of interannual adult local recruitment and (2) test whether environmental stress imposed by physiologically suboptimal temperatures and rainfall influence the magnitude of HFC. To address these questions, we used two different arrays of microsatellite markers: 14 loci classified as neutral and 12 loci classified as putatively functional. We found significant relationships between heterozygosity and probability of interannual local recruitment that were most likely explained by variation in genomewide heterozygosity. The strength of the association between heterozygosity and probability of interannual local recruitment was positively associated with annual accumulated precipitation. Annual mean heterozygosity increased over time, which may have resulted from an overall positive selection on heterozygosity over the course of the study period. Finally, neutral and putatively functional loci showed similar trends, but the former had stronger effect sizes and seemed to better reflect genomewide heterozygosity. Overall, our results show that HFC can be context dependent, emphasizing the need to consider the role of environmental heterogeneity as a key factor when exploring the consequences of individual genetic diversity on fitness in natural populations.

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          Most cited references33

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          Effect size, confidence interval and statistical significance: a practical guide for biologists.

          Null hypothesis significance testing (NHST) is the dominant statistical approach in biology, although it has many, frequently unappreciated, problems. Most importantly, NHST does not provide us with two crucial pieces of information: (1) the magnitude of an effect of interest, and (2) the precision of the estimate of the magnitude of that effect. All biologists should be ultimately interested in biological importance, which may be assessed using the magnitude of an effect, but not its statistical significance. Therefore, we advocate presentation of measures of the magnitude of effects (i.e. effect size statistics) and their confidence intervals (CIs) in all biological journals. Combined use of an effect size and its CIs enables one to assess the relationships within data more effectively than the use of p values, regardless of statistical significance. In addition, routine presentation of effect sizes will encourage researchers to view their results in the context of previous research and facilitate the incorporation of results into future meta-analysis, which has been increasingly used as the standard method of quantitative review in biology. In this article, we extensively discuss two dimensionless (and thus standardised) classes of effect size statistics: d statistics (standardised mean difference) and r statistics (correlation coefficient), because these can be calculated from almost all study designs and also because their calculations are essential for meta-analysis. However, our focus on these standardised effect size statistics does not mean unstandardised effect size statistics (e.g. mean difference and regression coefficient) are less important. We provide potential solutions for four main technical problems researchers may encounter when calculating effect size and CIs: (1) when covariates exist, (2) when bias in estimating effect size is possible, (3) when data have non-normal error structure and/or variances, and (4) when data are non-independent. Although interpretations of effect sizes are often difficult, we provide some pointers to help researchers. This paper serves both as a beginner's instruction manual and a stimulus for changing statistical practice for the better in the biological sciences.
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            On the correlation between heterozygosity and fitness in natural populations.

            Three primary hypotheses currently prevail for correlations between heterozygosity at a set of molecular markers and fitness in natural populations. First, multilocus heterozygosity-fitness correlations might result from selection acting directly on the scored loci, such as at particular allozyme loci. Second, significant levels of linkage disequilibrium, as in recently bottlenecked-and-expanded populations, might cause associations between the markers and fitness loci in the local chromosomal vicinity. Third, in partially inbred populations, heterozygosity at the markers might reflect variation in the inbreeding coefficient and might associate with fitness as a result of effects of homozygosity at genome-wide distributed loci. Despite years of research, the relative importance of these hypotheses remains unclear. The screening of heterozygosity at polymorphic DNA markers offers an opportunity to resolve this issue, and relevant empirical studies have now emerged. We provide an account of the recent progress on the subject, and give suggestions on how to distinguish between the three hypotheses in future studies.
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              Females increase offspring heterozygosity and fitness through extra-pair matings.

              Females in a variety of species commonly mate with multiple males, and there is evidence that they benefit by producing offspring of higher genetic quality; however, the nature of these genetic benefits is debated. Enhanced offspring survival or quality can result from intrinsic effects of paternal genes---'good genes'--or from interactions between the maternal and paternal genomes--'compatible genes'. Evidence for the latter process is accumulating: matings between relatives lead to decreased reproductive success, and the individual level of inbreeding--measured as average heterozygosity--is a strong fitness predictor. Females should thus benefit from mating with genetically dissimilar males. In many birds, social monogamy restricts mate choice, but females may circumvent this by pursuing extra-pair copulations. Here we show that female blue tits, Parus caeruleus, increase the heterozygosity of their progeny through extra-pair matings. Females thereby produce offspring of higher reproductive value, because less inbred individuals have increased survival chances, a more elaborate male secondary sexual trait (crown colour) and higher reproductive success. The cost of inbreeding may therefore be an important factor driving the evolution of female extra-pair mating.
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                Author and article information

                Contributors
                esperanza.sferrer@gmail.com
                Journal
                Ecol Evol
                Ecol Evol
                10.1002/(ISSN)2045-7758
                ECE3
                Ecology and Evolution
                John Wiley and Sons Inc. (Hoboken )
                2045-7758
                21 November 2016
                December 2016
                : 6
                : 24 ( doiID: 10.1002/ece3.2016.6.issue-24 )
                : 8857-8869
                Affiliations
                [ 1 ] Grupo de Investigación de la Biodiversidad Genética y CulturalInstituto de Investigación en Recursos Cinegéticos (CSIC‐UCLM‐JCCM) Ciudad RealSpain
                [ 2 ] Departamento de Ciencias Ambientales Facultad de Ciencias Ambientales y BioquímicaUniversidad de Castilla‐La Mancha ToledoSpain
                [ 3 ] Institute of Evolutionary Biology and Environmental StudiesUniversity of Zurich ZurichSwitzerland
                [ 4 ] Department of Integrative EcologyEstación Biológica de Doñana (EBD‐CSIC) SevilleSpain
                [ 5 ] Departamento de Ecología EvolutivaMuseo Nacional de Ciencias Naturales (CSIC) MadridSpain
                Author notes
                [*] [* ] Correspondence

                Esperanza S. Ferrer, Departamento de Ciencias Ambientales, Facultad de Ciencias Ambientales y Bioquímica, Universidad de Castilla‐La Mancha, Toledo, Spain.

                Email: esperanza.sferrer@ 123456gmail.com

                Article
                ECE32591
                10.1002/ece3.2591
                5192745
                7dec5a92-e377-40e4-9c1d-664b35efeab9
                © 2016 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.

                This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

                History
                : 17 August 2016
                : 10 October 2016
                : 17 October 2016
                Page count
                Figures: 4, Tables: 3, Pages: 13, Words: 10653
                Funding
                Funded by: Ministerio de Educación y Ciencia
                Award ID: GCL2007‐61395
                Funded by: Ministerio de Ciencia e Innovación
                Award ID: CGL2010-21933-C02-01
                Funded by: Junta de Comunidades de Castilla‐La Mancha‐European Social Fund
                Award ID: POIC10‐0269‐7632
                Funded by: Forschungskredit of the University of Zurich
                Award ID: FK‐14‐103
                Funded by: Ramón y Cajal Fellowship
                Award ID: RYC‐2013‐12501
                Funded by: Severo Ochoa Program
                Award ID: SEV‐2012‐0262
                Categories
                Original Research
                Original Research
                Custom metadata
                2.0
                ece32591
                December 2016
                Converter:WILEY_ML3GV2_TO_NLMPMC version:5.0.0 mode:remove_FC converted:25.12.2016

                Evolutionary Biology
                cyanistes caeruleus,functional and neutral markers,genotype‐by‐environment interaction,heterozygosity,hfc,selection differential

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