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      Sources of individual variation in plasma testosterone levels

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          Abstract

          The steroid hormone testosterone (T) plays a central role in the regulation of breeding in males, because many physiological, morphological and behavioural traits related to reproduction are T dependent. Moreover, in many seasonally breeding vertebrates, male plasma T levels typically show a pronounced peak during the breeding season. While such population-level patterns are fairly well worked out, the sources and the implications of the large variability in individual T levels within the seasonal cycle remain surprisingly little understood. Understanding the potential sources of individual variation in T levels is important for behavioural and evolutionary ecologists, for at least two reasons. First, in ‘honest signalling’ theory, T is hypothesized to play a critical role as the assumed factor that enforces honesty of the expression of sexually selected quality indicators. Second, T is often considered a key mediator of central life-history trade-offs, such as investment in survival versus reproduction or in mating versus parental care. Here, we discuss the patterns of within- and between-individual variation in male plasma T levels in free-living populations of birds. We argue that it is unclear whether this variability mainly reflects differences in underlying individual quality (intrinsic factors such as genetic or maternal effects) or in the environment (extrinsic factors including time of day, individual territorial status and past experience). Research in avian behavioural endocrinology has mainly focused on the effects of extrinsic factors, while other sources of variance are often ignored. We suggest that studies that use an integrative approach and investigate the relative importance of all potential sources of variation are essential for the interpretation of data on individual plasma T levels.

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          Maternal hormones as a tool to adjust offspring phenotype in avian species.

          Avian eggs contain substantial amounts of maternal hormones and so provide an excellent model to study hormone-mediated maternal effects. We review this new and rapidly evolving field, taking an ecological and evolutionary approach and focusing on effects and function of maternal androgens in offspring development. Manipulation of yolk levels of androgens within the physiological range indicates that maternal androgens affect behaviour, growth, morphology, immune function and survival of the offspring, in some cases even long after fledging. Descriptive and experimental studies show systematic variation in maternal androgen deposition both within and among clutches, as well as in relation to the sex of the embryo. We discuss the potential adaptive value of maternal androgen transfer at all these three levels. We conclude that maternal androgen deposition in avian eggs provides a flexible mechanism of non-genetic inheritance, by which the mother can favour some offspring over others, and adjust their developmental trajectories to prevailing environmental conditions, producing different phenotypes. However, the literature is less consistent than often assumed and at all three levels, the functional explanations need further experimental testing. The field would greatly benefit from an analysis of the underlying physiological mechanisms.
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            Yolk is a source of maternal testosterone for developing birds.

            H Schwabl (1993)
            The sex steroid hormones that affect development in birds have been thought to be produced exclusively by the embryo or neonate. I used radioimmunoassay to measure the amounts of androstenedione, 5 alpha-dihydrotestosterone, testosterone, 17 beta-estradiol, and corticosterone in the yolk of freshly laid canary (Serinus canaria) and zebra finch (Poephila guttata) eggs. Testosterone was found in both canary and zebra finch eggs, but its contents were much higher in the former than in the latter. The testosterone content of canary eggs in a same clutch increased with the order of laying, regardless of the genetic sex of the offspring that hatched from these eggs. Yolk testosterone was also present in the eggs of female canaries that were kept without a male, indicating that it is of maternal origin. The social rank of juvenile canaries was positively correlated with the concentration of yolk testosterone in the eggs from which they hatched, suggesting that the development of aggressive behavior of offspring might be subject to modification by maternal testosterone. These findings indicate that female songbirds can bestow upon their eggs a dose of hormone that modifies the behavior of offspring. Variable doses of these hormones might explain some of the individual variation in offspring behavior.
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              Sexual Selection

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                Author and article information

                Journal
                Philos Trans R Soc Lond B Biol Sci
                RSTB
                Philosophical Transactions of the Royal Society B: Biological Sciences
                The Royal Society (London )
                0962-8436
                1471-2970
                28 November 2007
                12 May 2008
                : 363
                : 1497 , Theme Issue ‘Integration of ecology and endocrinology in avian reproduction: a new synthesis’ compiled by John Wingfield, Tony Williams and Marcel Visser
                : 1711-1723
                Affiliations
                [1 ]Department of Behavioural Ecology and Evolutionary Genetics, Max Planck Institute for Ornithology PO Box 1564, 82305 Seewiesen, Germany
                [2 ]Research Group Evolution of Sexual Signals, Max Planck Institute for Ornithology Vogelwarte Radolfzell, 78315 Radolfzell am Bodensee, Germany
                Author notes
                [* ]Author for correspondence ( b.kempenaers@ 123456orn.mpg.de )
                Article
                rstb20070001
                10.1098/rstb.2007.0001
                2367619
                18048297
                3607eb73-217f-4ea1-934b-8e3d54e6f522
                Copyright © 2007 The Royal Society

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

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                Categories
                Review

                Philosophy of science
                genetic effects,plasticity,behavioural syndrome,hormone,maternal effects,heritability

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