Waddington's Legacy in Development and Evolution

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Abstract
This paper provides an overview of the life and works of Conrad Hal Waddington(1905-1975). After an early life spent apart from his parents pursuing ammonites, naturalhistory, geology and archaeology, Waddington took a degree in Geology at Cambridge (1926).Genetics and experimental embryology soon replaced palaeontology as he began his experimentalstudies on the chemical nature of the primary organizer discovered by Spemannand Mangold in 1924. It was during this period of collaboration with Joseph and DorothyNeedham that Waddington developed the concepts of evocation and individuation. Theestablishment of a Unit on Animal Breeding and Genetics in Edinburgh after the secondWorld War provided Waddington with his professional home for the rest of his career as he sought to integrate genetics and development into an evolutionarily relevant discipline.Our conception of embryonic development as a highly integrated series of canalized pathways owes much to Waddington's development of the concepts of canalization, chreods, epigenetics and the epigenotype. The metaphorical epigenetic landscape became the way that most developmental biologists “saw” the organization of embryonic development. The concept of supragenomic, integrated, heritable, epigenetic organization of embryonic development is arguably Waddington's lasting legacy to development and evolution. The integration of his epigenetic legacy into a quantitative developmental genetics model of the developmental and evolutionary origin of phenotypes is now being undertaken. It has still to be proven whether genetic assimilation, which Waddington demonstrated to be a real phenomenon in laboratory experiments, has been a force in evolutionary adaptation.

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