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• BSc: B.A. Oregon State College, 1943-1944, 1946-1947 (Zoology) 
• Msc: Oregon State College, 1948-1949 (Zoology). 
• PhD: University of Pennsylvania, 1949-1952 (Zoology) 
• Postdoctoral Fellow of the Carnegie Inst. of Washington at Cold Spring Harbor, 1952-1953.