This chapter reappraises Waddington’s processual theory of epigenetics and examines its implications for contemporary evolutionary biology. It focuses in particular on the ontological difference between two conflicting assumptions that have been conflated in the recent debate over the nature of cryptic variability: a substance view that is consistent with the modern synthesis and construes variability as a preexisting pool of random genetic variation; and a processual view, which derives from Waddington’s conception of developmental canalization and understands variability as an epigenetic process. The chapter also discusses how these opposing interpretations fare in their capacity to explain the genetic assimilation of acquired characters.