The behavioural and physiological ecology of embryos: responding to the challenges of life inside an egg.

Adaptations of post-hatching animals have attracted far more study than have embryonic responses to environmental challenges, but recent research suggests that we have underestimated the complexity and flexibility of embryos. We advocate a dynamic view of embryos as organisms capable of responding - on both ecological and evolutionary timescales - to their developmental environments. By viewing embryos in this way, rather than assuming an inability of pre-hatching stages to adapt and respond, we can broaden the ontogenetic breadth of evolutionary and ecological research. Both biotic and abiotic factors affect embryogenesis, and embryos exhibit a broad range of behavioural and physiological responses that enable them to deal with changes in their developmental environments in the course of interactions with their parents, with other embryos, with predators, and with the physical environment. Such plasticity may profoundly affect offspring phenotypes and fitness, and in turn influence the temporal and spatial dynamics of populations and communities. Future research in this field could benefit from an integrated framework that combines multiple approaches (field investigations, manipulative experiments, ecological modelling) to clarify the mechanisms and consequences of embryonic adaptations and plasticity.