 He was educated at the University of Pennsylvania (A.B., 1879; M.D., 1882), and studied biology at the universities of Leipzig and Jena in 1882–1883, and at Johns Hopkins for a year.In 1884 he was appointed professor of vertebrate morphology at The Wistar Institute of Anatomy and Biology, and became a director of the institute.He was a professor of zoölogy at the University of Pennsylvania from 1894 to 1905, secretary of Penn's biological faculty (1884–1889), and dean of Penn's college faculty (1889–1894).