Reframing developmental biology and building evolutionary theory's new synthesis.

Gilbert and Epel present a new approach to developmental biology: embryogenesis must be understood within the full context of the organism's environment. Instead of an insular embryo following a genetic blueprint, this revised program maintains that embryogenesis is subject to inputs from the environment that generate novel genetic variation with dynamic consequences for development. Beyond allelic variation of structural genes and of regulatory loci, plasticity-derived epigenetic variation completes the triad of the major types of variation required for evolution. Developmental biology and ecology, disciplines that have previously been regarded as distinct, are presented here as fully integrated under the rubric of "eco-devo," and from this perspective, which highlights how the environment not only selects variation, it helps construct it, another synthesis with evolutionary biology must also be made, "eco-evo-devo." This second integration has enormous implications for expanding evolution theory, inasmuch as the Modern Synthesis (Provine 1971), which combined classical genetics and Darwinism in the mid-20th century, did not account for the role of development in evolution. The eco-evo-devo synthesis thus portends a major theoretical inflection in evolutionary biology. Following a description of these scientific developments, comment is offered as to how this new integrated approach might be understood within the larger shifts in contemporary biology.