His PhD training was with Andrew W. Murray at the University of California, San Francisco, studying the process of chromosome segregation. He went on to join Tim Mitchison’s group in the Department of Cell Biology and the Institute for Chemistry and Cell Biology at Harvard Medical School studying cytokinesis. His research at Stanford has focused on chromosome segregation mechanisms and, in particular, the formation and function of centromeres. Using diverse model organisms and approaches, his group has developed new systems to understand, at a mechanistic level, the epigenetic control of centromere establishment and assembly. Current efforts in his group are exploring the multiscale organization of eukaryotic genomes, the control of zygotic genome activation, and the regulation of heterochromatin formation. Photo courtesy of Whitney Johnson.