Egg antimicrobials, embryo sex and chick phenotype in the yellow-legged gull

Maternal effects through albumen quality are largely unexplored, despite the fundamental role that albumen exerts as source of proteins and water, as well as for antimicrobial defence of the embryo. We analysed the variation of two major albumen antimicrobials, avidin and lysozyme, by extracting samples from freshly laid eggs of the yellow-legged gull (Larus michahellis) and by correlating their levels to egg features. Lysozyme concentration increased along the laying sequence, while avidin concentration decreased. Both antimicrobials declined during the season. In addition, avidin concentration declined from first- to last-laid male eggs, whereas the opposite was true among the female eggs. We also analysed chick body mass and size and immune response, in relation to albumen antimicrobial levels in their original egg while controlling for potential covariation between egg quality and rearing conditions by cross-fostering eggs between nests. Tarsus length decreased with avidin concentration, particularly early in the season. Avidin concentration negatively predicted tarsus length of chicks and the phytohaemagglutinin response of females, but not males. However, chick phenotype did not covary with lysozyme albumen concentration. This is the first study where maternal effects mediated by albumen antimicrobials are investigated in relation to both sex and egg features in any wild bird species. Whether the observed patterns of variation in antimicrobial concentration are the by-product of maternal physiological constraints, or reflect adaptive allocation strategies, cannot be ascertained. The covariation between chick cell-mediated immunity and albumen avidin concentration might be causal, according to the documented effects of albumen proteins on immunity in other species.