until 1942. His friends included Gregory Bateson, Walter Gropius, C. P. Snow, Solly Zuckerman, Joseph Needham, and John Desmond Bernal. His interests began with palaeontology but moved on to the heredity and development of living things. He also studied philosophy. During World War II he was involved in operational research with the Royal Air Force and became scientific advisor to the Commander in Chief of Coastal Command from 1944 to 1945. After the war, in 1947, he replaced Francis Albert Eley Crew as Professor of Animal Genetics at the University of Edinburgh. He would stay at Edinburgh for the rest of life with the exception of one year (1960–1961) when he was a Fellow on the faculty in the Center for Advanced Studies at Wesleyan University in Middletown, Connecticut. His personal papers are largely kept at the University of Edinburgh library. He died in Edinburgh on 26 September 1975. Family Waddington was married twice. His first marriage produced a son, C. Jake Waddington, professor of physics at the University of Minnesota, but ended in 1936. He then married Margaret Justin Blanco White, daughter of the writer Amber Reeves, with whom he had two daughters, the anthropologist Caroline Humphrey (1943–) and mathematician Dusa McDuff (1945–). Evolution In the early 1930s, Waddington and many other embryologists looked for the molecules that would induce the amphibian neural tube. The search was beyond the technology of that time, and most embryologists moved away from such deep problems. Waddington, however, came to the view that the answers to embryology lay in genetics, and in 1935 went to Thomas Hunt Morgan's Drosophila laboratory in California, even though this was a time when most embryologists felt that genes were unimportant and just played a role in minor phenomena such as eye colour. In the late 1930s, Waddington produced formal models about how gene regulatory products could generate developmental phenomena, showed how the mechanisms underpinning Drosophila development could be studied through a systematic analysis of mutations that affected the development of the Drosophila wing. In a period of great creativity at the end of the 1930s, he also discovered mutations that affected cell phenotypes and wrote his first textbook of "developmental epigenetics", a term that then meant the external manifestation of genetic activity. Waddington introduced the concept of canalisation, the ability of an organism to produce the same phenotype despite variation in genotype or environment. He also identified