The origins of variation: evolutionary insights from developmental science.

Evidence from contemporary epigenetic research indicates that it is not biologically meaningful to discuss genes without reference to the molecular, cellular, organismal, and environmental context within which they are activated and expressed. Genetic and nongenetic factors, including those beyond the organism, constitute a dynamic relational developmental system. This insight highlights the importance of bringing together genetics, development, and ecology into one explanatory framework for a more complete understanding of the emergence and maintenance of phenotypic stability and variability. In this Chapter, I review some examples of this integrative effort and explore its implications for developmental and evolutionary science, with a particular emphasis on the origins of phenotypic novelty. I argue that developmental science is critical to this integrative effort, in that evolutionary explanation cannot be complete without developmental explanation. This is the case because the process of development generates the phenotypic variation on which natural selection can act.