Maternal effects as generators of evolutionary change: a reassessment.

Despite being a center of debate in biology for centuries, the connection between the generation of novel adaptive variation and its inheritance remains a contentious issue. In evolutionary and behavioral ecology, assigning natural and sexual selection a creative and anticipatory role unmasks the need to explicitly consider the link between a trait's functional importance and its inheritance and results in confusion about selection as an adaptive modifier of development versus selection as a passive filter of already produced forms. In developmental genetics, an emphasis on regulatory versus coding aspects of molecular evolutionary change overlooks the fundamental question of the origination of an inherited developmental toolkit and assignment of its regulatory functions. Because maternal effects, by definition, combine developmental induction of functionally important changes and their inheritance, they bridge the origin and evolution of organismal adaptability, at least on short time scales. The explosion of empirical studies of maternal effects raises a question--are maternal effects ubiquitous but short-term adjusters and fine-tuners of an evolved form with only secondary importance for evolutionary change? Or are they a particularly clear example of a stage in a continuum of inheritance systems that accumulates, internalizes, and passes on the most consistent and adaptive organism-environment interactions? Here I place recent empirical studies of avian maternal effects into the evolutionary framework of variation, selection, and inheritance to examine whether maternal effects provide a window into evolutionary processes.