Manipulation of primary sex ratio in birds: lessons from the homing pigeon (Columba livia domestica).

Across various animal taxa not only the secondary sex ratio but also the primary sex ratio (at conception) shows significant deviations from the expected equal proportions of sons and daughters. Birds are especially intriguing to study this phenomenon as avian females are the heterogametic sex (ZW); therefore sex determination might be under direct control of the mother. Avian sex ratios vary in relation to environmental or maternal condition, which can also affect the production of maternal steroids that in turn are involved in reproduction and accumulate in the developing follicle before meiosis. As the proximate mechanisms underlying biased primary sex ratio are largely elusive, we explored how, and to what extent, maternal steroid hormones may be involved in affecting primary or secondary sex ratio in clutches of various species of pigeons. First we demonstrated a clear case of seasonal change in sex ratio in first eggs both in the Rock Pigeon (Columba livia) and in a related species, the Wood Pigeon (Columba palumbus), both producing clutches of two eggs. In the Homing Pigeon (Columba livia domestica), domesticated from the Rock Pigeon, testosterone treatment of breeding females induced a clear male bias, while corticosterone induced a female bias in first eggs and we argue that this is in line with sex allocation theory. We next analyzed treatment effects on follicle formation, yolk mass, and yolk hormones, the latter both pre- and post-ovulatory, in order to test a diversity of potential mechanisms related to both primary and secondary sex ratio manipulation. We conclude that maternal plasma hormone levels may affect several pre-ovulatory mechanisms affecting primary sex ratio, whereas egg hormones are probably involved in secondary sex ratio manipulation only.