Genetic assimilation, robustness and plasticity are key processes in the development and evolution of novel traits

This is a commentary on how C.H. Waddington's experiments in the 1950's, first published in 1953 in a provocatively titled paper "Genetic assimilation of an acquired character," laid the foundation for the field of phenotypic plasticity, and how the ideas he developed eventually led to new ways of understanding phenotypic robustness, plasticity, and how novel traits develop and evolve. The "acquired characters" that Waddington worked with were based on Goldschmidt's ideas of "phenocopies": new phenotypes that develop after an environmental stress that resemble the phenotypes of known mutations. The idea behind genetic assimilation, first outlined by Waddington in 1942, is that existing developmental pathways can be rearranged and redirected through selection to stabilize the phenocopy phenotype, without requiring new mutations. In the short term, Waddington's work led to the discovery of heat shock proteins and the role of Hsp90 in masking defective proteins and allowing the accumulation of cryptic genetic variation. Subsequent studies revealed a host of stabilizing systems that operate at all levels of biological organization that make phenotypes robust to genetic and environmental variation. Many of these resemble homeostatic mechanisms that don't require a stress shock but operate under normal physiological conditions and allow for the accumulation of large amounts of cryptic genetic variation. This cryptic genetic variation can be revealed by mutations or environmental factors that destabilize a homeostatic mechanism. Selection can then act on the phenotypic variants that are produced. This scenario corresponds to the modern phenotype-first hypothesis for the evolution of novel traits that was foreseen by Waddington as early as 1942.