 He completed undergraduate studies at Pomona College in 1925 and was awarded a Ph.D. in botany from the University of California in 1930.From 1925 to 1950 he was based at the Carnegie Institute of Washington, at Stanford University, where he worked on plant species concepts with Jens Clausen and William Hiesey.In 1949, the three of them were co-recipients of the Mary Soper Pope Memorial Award in botany.