A molecular biologist who made a pioneering contribution to the development of molecular biology. During the early days of molecular biology, he contributed greatly to the establishment of the genetic code including the discovery of termination codons, and also proved elegantly the existence of messenger RNA. These studies, together with numerous other works by him, led to the establishment of modern gene technology. He also developed an experimental system for a nematode, and demonstrated that the timing and location of cell division is completely programmed genetically.