1857-1864 studied medicine at the University of Heidelberg, at that time medical assistant of Friedreich; 1864 M.D. University of Heidelberg; 1865 Habilitation; 1869 associate professor, University of Heidelberg; 1880 tenured professor at the University of Leipzig; 1883 returned to Heidelberg; 1887 was elected to membership in the Deutsche Akademie der Naturforscher Leopoldina; 1907 professor emeritus.