 He studied at the universities of Freiburg, Munich (under Richard Hertwig), Berlin and Strasbourg (under Friedrich Goltz and Ernst Julius Richard Ewald); receiving his PhD in 1895 at Munich.From 1896 to 1911 he worked at the institute of physiology in Strasbourg, where in 1898 he obtained his doctorate in medicine.In 1911 he became a professor of physiology at the University of Kiel, and four years later, relocated as a professor to the University of Frankfurt.