He obtained his PhD from New York University Medical Center, studying the roles of fibroblast growth factors in midbrain and cerebellum development. He then received postdoctoral training under the supervision of Drs. Lee Niswander and Kathryn Anderson at the Sloan-Kettering Institute and made the discovery that the primary cilium regulates Gli transcription factor activation and proteolytic processing in the mouse. The research in his lab is now focused on the molecular mechanisms of cilia biogenesis and the regulation of the Hedgehog signal transduction in mammals.