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Maternal hormones are an excellent pathway for the mother to influence offspring development, and birds provide exceptional opportunities to study these hormone-mediated maternal effects. Two dominant hypotheses about the function of yolk androgens in avian eggs concern maternal manipulation of sibling competition and post hatching paternal care. In megapodes, however, neither sibling competition nor post hatching parental care exists. Eggs are incubated by external heat sources, and chicks dig themselves out of their underground nest and live independently of their parents and their siblings. In this first study on egg androgens of such a megapode, the Australian Brush-turkey Alectura lathami, we found nevertheless substantial amounts of maternal androgens. Since size of the incubation mound, incubation temperature, egg size and laying date greatly vary in this species, we analysed variation in testosterone (T), androstenedione (A4) and dihydrotestosterone (DHT) in relation to these factors. T concentrations were significantly higher in eggs from bigger mounds and laid at greater depth, which may compensate via anabolic effects for the longer duration and higher energetic requirements of chicks when digging themselves out. T concentrations were higher in smaller eggs, and both yolk A4 and T concentrations increased with laying date, perhaps as a compensatory measure, while DHT concentrations only varied across different mounds. These results indicate that maternal androgens may influence offspring development outside the contexts of sibling competition or parental care. (c) 2007 Elsevier Inc. All rights reserved.

Avian eggs contain substantial levels of various hormones of maternal origin and have recently received a lot of interest, mainly from behavioral ecologists. These studies strongly depend on the measurement of egg hormone levels, but the method of measuring these levels has received little attention. This paper describes the sampling, extraction, and assay of hormones in avian eggs and related methodological problems. The method of sampling is important because hormones are not homogeneously distributed within the egg, and after onset of embryonic development their levels may decrease and increase due to changes in egg structure and secretion or uptake of hormones by the embryo. The extraction of hormones from the yolk and chromatographic separation of different hormones for immunoassays can strongly influence the results because such procedures remove interfering substances such as proteins, lipids, and other hormones and their metabolites, which can cross-react with the antiserum used. Finally, the assay itself needs more validation than many studies report, especially with respect to the accuracy and specificity of the hormone measurements. We conclude that the addressed issues need more attention for the correct interpretation of differences in hormone levels within and between studies.

SUMMARY Within- and between-clutch variation in yolk titres of hormones of maternal origin has been found in many avian species. So far, experiments have revealed mainly beneficial effects of maternal androgens. This would also apply to black-headed gulls (Larus ridibundus). Previous experiments have shown that chicks benefit from these higher levels since their competitive abilities are improved and growth and survival probabilities thus enhanced. However, not all females show the same increase in yolk hormones from first to last egg or invest equally high amounts of androgens in their clutches. Possibly, there is a trade-off between the beneficial effects of high androgen levels and potential costs, such as increased metabolic rates. We studied possible metabolic costs of experimentally elevated yolk androgen levels for chicks of several age classes, starting three days prior to hatching until fledging at an age of approximately 30 days. Daily energy expenditure in the field, measured using the doubly labelled water technique, did not differ between treatments or between sexes. Oxygen consumption measured in birds at rest in the lab (RMR) did not vary between chicks hatched from androgen-injected (T) or oil-injected (Oil) control eggs at any age in thermo-neutral or below thermo-neutral conditions. Males showed a lower RMR than females towards the fledging age. We conclude that it is unlikely that the costs of high maternal androgen levels can be found in higher energy expenditure in the chick.