A.B., Columbia University, 1973; M.Phil., Yale University, 1975; Ph.D., 1977; D.h.c., Ecole Normale Superieure; D.Sc.h.c., Yale University. Assistant Professor, Caltech, 1980-85; Associate Professor, 1985-89; Professor, 1989-2002; Beadle Professor, 2002-; HHMI Investigator, 2013-; Executive Officer, 1995-2000; Chair, Division of Biology, 2000-10.

Biological Engineering; Developmental Biology and Genetics; Systems Biology; Evolutionary and Organismal Biology