Boveri was born in Bamberg, Germany, and graduated in medicine from Munich in 1885. He remained at Munich to do cytological research until his appointment as professor of zoology and comparative anatomy at Würzburg in 1893. In 1888, he coined the term ‘centrosome’ for the region of the cell that contains the centriole, first discovered by Edouard van Beneden. Boveri also proved Beneden's theory that equal numbers of chromosomes are contributed by the egg and the sperm to the zygote. Boveri accurately described the formation of the polar bodies following meiosis in the egg cell, and made pioneering studies of sperm formation (spermatogenesis), introducing a diagrammatic representation of the process (1892), which is still in use today.