1861-1862 studies of medicine and of natural sciences at the University of Jena under K. Gegenbaur, then at Heidelberg, Göttingen and Leipzig; 1867 Dr.med. under A. von Bezold; 1867 assistant to F. C. Donders in Utrecht; 1871 associate professor of biology and histology, 1888 professor of physiology at Utrecht University in succession to Donders; 1897-1908 professor of physiology and head of the Physiologische Institut at the University of Berlin.
