Maternally derived yolk testosterone enhances the development of the hatching muscle in the red-winged blackbirdAgelaius phoeniceus

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Abstract
Hatching asynchrony in avian species often leads to the formation of a size hierarchy that places last–hatched nestlings at a significant disadvantage. The hatching muscle (musculus complexus) is responsible for breaking the shell during hatching and for dorsal flexion of the neck during begging. An increase in its strength in last–hatched nestlings could mitigate the effects of hatching asynchrony by reducing the time required for hatching or enhancing the effectiveness of begging for parentally delivered food or both. We have previously found that yolk testosterone concentration increases with laying order in the red–winged blackbird Agelaius phoeniceus. In this study, we investigated the hypothesis that yolk testosterone has anabolic effects on the development of the complexus, thereby influencing competition among asynchronously hatched nestlings. We found that both yolk testosterone concentration and relative complexus mass (complexus mass/nestling body mass) increased with laying order and that these two variables were positively correlated in both newly hatched nestlings and in two–day–old broods. Moreover, direct injections of testosterone into egg yolks resulted in an increase in relative complexus mass, while injections of flutamide, a testosterone antagonist, resulted in a decrease in relative complexus mass. Neither yolk testosterone concentration nor relative complexus mass differed between male and female nestlings.