Richard Lewontin  Richard Charles "Dick" Lewontin (born March 29, 1929) is an American evolutionary biologist, mathematician, geneticist, and social commentator. A leader in developing the mathematical basis of population genetics and evolutionary theory, he pioneered the application of techniques from molecular biology, such as gel electrophoresis, to questions of genetic variation and evolution. In a pair of seminal 1966 papers co-authored with J.L. Hubby in the journal "Genetics", Lewontin helped set the stage for the modern field of molecular evolution. In 1979 he and Stephen Jay Gould introduced the term "spandrel" into evolutionary theory. From 1973 to 1998, he held an endowed chair in zoology and biology at Harvard University, and since 2003 has been a research professor there. Lewontin opposes genetic determinism. Lewontin was born in New York City, to parents descended from late 19th-century Eastern European Jewish immigrants. He attended Forest Hills High School and the École Libre des Hautes Études in New York. In 1951 he graduated from Harvard College (BS, biology). In 1952, Lewontin received a master's degree in mathematical statistics, followed by a doctorate in zoology in 1954, both from Columbia University, where he was a student of Theodosius Dobzhansky. He held faculty positions at North Carolina State University, the University of Rochester, and the University of Chicago. In 1973 Lewontin was appointed as Alexander Agassiz Professor of Zoology and Professor of Biology at Harvard University, holding the position until 1998. Lewontin has worked in both theoretical and experimental population genetics. A hallmark of his work has been an interest in new technology. He was the first person to do a computer simulation of the behavior of a single gene locus (previous simulation work having been of models with multiple loci). In 1960 he and Ken-Ichi Kojima were the first population geneticists to give the equations for change of haplotype frequencies with interacting natural selection at two loci. This set off a wave of theoretical work on two-locus selection in the 1960s and 1970s. Their paper gave a theoretical derivation of the equilibria expected, and also investigated the dynamics of the model by computer iteration. Lewontin later introduced the D' measure of linkage disequilibrium. (He also introduced the term "linkage disequilibrium", about which many population geneticists have been unenthusiastic.) In 1966, he and Jack Hubby published a paper that revolutionized population genetics.