His Nobel Prize–winning studies with Drosophila, (including the discovery of the Drosophila Bithorax complex of homeotic genes, and elucidation of its function), founded the field of evolutionary developmental biology and laid the groundwork for our current understanding of the universal, evolutionarily conserved strategies controlling animal development. He is credited with development of the complementation test. His key publications in the fields of genetics, developmental biology, radiation and cancer are presented in the book Genes, Development and Cancer, which was released in 2004.