Bachelor, Brandeis University, Waltham, Massachusetts, 1971. Doctor of Philosophy, Harvard University, 1975.
Postdoctoral fellow Harvard Medical School, Boston, 1975-1978. Assistant professor biology University Pennsylvania, Philadelphia, 1978-1984, associate professor, 1984-1990, Patricia Williams professor biology, former chairman department biology, Charles and William L. Day distinguished professor biology, associate dean natural sciences.
Fellow American Association for the Advancement of Science.
Married Nicola Thomsen Neff, October 20, 1979.