Richard Lewontin, a geneticist and evolutionary biologist at Harvard University, died on July 4 at the age of 92, according to an obituary. Mary Jane Lewontin, his wife of more than 70 years, died three days prior on July 1. Lewontin studied genetic diversity within populations and helped develop the use of protein gel electrophoresis to examine this at a molecular level. 

“He’s considered one of the evolutionary biology greats,” Adriana Briscoe, an evolutionary biologist at the University of California, Irvine, who was a graduate student in Lewontin’s lab from 1993 until 1998, tells The Scientist. “He’s considered a giant in his field.” 

Born in New York City in 1929, Lewontin graduated with a bachelor’s degree in biology from Harvard University in 1951 and then went to Columbia University to study fruit fly population density with his graduate advisor, Theodosius Dobzhansky, according to The New York Times. He graduated with a master’s degree in 1952 and a doctorate in zoology in 1954. 

In 1966, at the University of Chicago, Lewontin and John Hubby published two papers that pioneered the use of protein gel electrophoresis to study genetic variation within populations of wild fruit flies. Not only did the technique lay the groundwork for the field of molecular genetics, but it revealed a surprising amount of genetic diversity within the population. 

He joined Harvard in 1973 as a professor, and remained there until his retirement in 2003, according to a memoriam written by Harvard’s Museum of Comparative Zoology. 

Lewontin was well-known for his critiques of adaptationist programs—the idea that all organismal traits have been “optimized” due to natural selection. Rather, he argued that genetic variation within a population could also be the product of random chance, or due to selection on linked loci on the genome.

He also wrote a seminal 1972 paper in which he argued there is more genetic variation within members of a population of humans than there is between members of different groups, undermining the idea that there is a genetic basis for the idea of race.

“Richard ‘Dick’ Lewontin was [a] foundational scholar in the field of evolutionary genetics and evolution writ large whose impact on the field is hard to over-estimate,” writes Elena Kramer, the department chair in Harvard’s Department of Organismic and Evolutionary Biology, in an email to The Scientist. “He is recognized as a phenomenal scholar but also a talented communicator, both as a teacher and writer, whose fluency in communicating science was underpinned by a deep understanding of his material and the practice of teaching.”

Lewontin won numerous awards and honors, including a the Sewall Wright Award in 1994, honorary lifetime membership in the American Society of Naturalists, the 2015 Crafoord Prize in Biosciences, and the 2017 Thomas Hunt Morgan Medal from the Genetics Society of America. 

Briscoe says Lewontin expected his graduate students to come up with their own ideas, which could be intimidating. “But that also meant that at the end of your PhD, you really own your intellectual labor,” she adds, noting that Lewontin was concerned about how some academics would appropriate their students’ work. She says he wouldn’t even put his name on one of his student’s papers unless he felt he had contributed to the study. 

“He was blunt and gruff, the kind of person who did not suffer fools gladly, but he also had a sense of humor, a lot of fondness for his students, and he told entertaining stories,” says Briscoe. “Scientists, philosophers, and historians of science flocked to his lab to participate in the lab’s lively seminars, and of course, to speak with him about ideas.

Lewontin and his wife are survived by four sons, seven grandchildren, and one great-grandchild.