Systems Evolutionary Biology of Waddington’s Canalization and Genetic Assimilation

In recent years, there has been growing interest in computer modeling of the evolution of gene and cell regulatory networks, in general, and in computational studies of the classic ideas of Baldwin, Schmalhausen, Waddington, and followers, in particular. Two related aspects of Waddington’s evolutionary theories are the concepts of canalization and of genetic assimilation. Canalization is associated with the robust development of an individual to diverse perturbations and noise, though, when fluctuations in developmental factors exceed a particular limit, the normal developmental trajectory can be “thrown out” of the robust canal, resulting in an altered phenotype. If selective pressure favors the new phenotype, an initial individual loss of canalization can lead to phenotypic changes in the population (with canalization then becoming established for the new phenotype). Genetic assimilation is the subsequent genetic fixing of the new trait in the population. Recent experimental and theoretical works have established a quantitative basis for these classic concepts of Waddington; this chapter will review these new developments in systems evolutionary biology