He completed his bachelor’s degree at Cornell and obtained his Ph.D. at the University of Michigan, where he studied mutations in the Hoxa13 gene that affect limb and urogenital patterning in mice and humans. From 1998–2002 he obtained postdoctoral training at Stanford the lab of David Kingsley where he initiated cis-regulatory studies of the Gdf6 gene. Since 2002 he has worked at Vanderbilt where his lab continues studies on Gdf6, Bmp2 and Bmp4 cis-regulation and function relating to skeletal development.