Frank Ruddle, a trailblazer in genetic research and former professor in both the Biology and Genetics departments, died March 10 at Yale-New Haven Hospital. He was 83.

Ruddle’s lab at Yale was the site of many scientific milestones beginning in the 1970s, including the first insertion of foreign genes into the mouse genome in 1980, which created the first transgenic animal and opened the way for scientific research on genetically modified organisms. Ruddle is credited with organizing the first human genome mapping workshop at Yale in 1973 and developing gene-mapping technology that helped lead to the establishment and success of the Human Genome Project. Ruddle’s friends, students and colleagues remember him as a quiet, generous man with a wonderful sense of humor and a passion for science.

“I always felt that Frank was a pioneer in the field and really directed the project that eventually led to the mapping and sequencing of the human genome,” said Raju Kucherlapati, a genetics professor at Harvard Medical School who worked as a fellow in Ruddle’s lab in the 1970s. “They are not given posthumously, but he deserves to win a Nobel Prize for that effort.”

Born in 1929 in New Jersey to British parents, Ruddle grew up in Ohio and left high school early to join the U.S. Army Air Forces in Japan in 1946. He attended Wayne State University before receiving his master’s degree jointly from Wayne State and the Children’s Hospital of Detroit and earning a Ph.D. in zoology from the University of California, Berkeley. He joined the Yale faculty in 1961 after conducting postdoctoral research at Glasgow University.

During his 41 years at Yale, Ruddle served two terms as chair of the Biology Department for a total of 10 years. He mentored dozens of undergraduates, supervised roughly 52 postdoctoral fellows and guided 30 graduate students to their Ph.D.s.

Just as genomics began gaining international fame, Ruddle withdrew from traditional genetics and focused on developmental genetics, studying how a handful of similar genes control the development of multicellular organisms. During a sabbatical from Yale, he joined a team of scientists at the University of Basel in Switzerland and, together with William McGinnis, cloned the first mouse homeobox gene in 1983.

“The whole idea of transgenesis took off from his work,” said Cooduvalli Shashikant, a biology professor at Penn State University who worked in Ruddle’s lab for nearly a decade in the 1980s and 1990s.

Ruddle’s research on the evolution and expression of genes in animals expanded the scientific community’s understanding of how humans develop and how genes vary between species, biology professor Ronald Breaker said in a March 15 email. He added that Ruddle’s lab was a model for modern biology research labs because Ruddle used large teams to work on “truly exciting research with profound implications.”

Ruddle was a thoughtful, methodical man who knew exactly how much guidance and how much freedom to give researchers in his lab to help them thrive, former Yale postdoctoral fellows said.

Jon Gordon GRD ’78 MED ’80, the postdoctoral fellow who created the first transgenic mouse in Ruddle’s lab in 1980, said Ruddle always prioritized the advancement of human understanding of the world rather than emphasizing fame and career success, and he held his students to the same standard.

Ruddle played an influential role in the careers of many of his postdoctoral fellows and often invited them to his home, either individually or as a group. Kucherlapati recalled one afternoon when Ruddle took him to lunch in one of Yale’s residential colleges, which was a “thrill” for Kucherlapati. Ruddle and Kucherlapati — whose heights measured 6-foot-7 and 5-foot-2, respectively — earned themselves the nicknames “Mutt and Jeff” around the lab, a reference to the classic comic strip.

Ruddle loved his family, dogs and cooking, and he often took his 18-foot sailboat, Shamrock, out for day sails. After he retired, he started taking singing lessons at the Neighborhood Music School in New Haven. He had a “rich, baritone voice,” said New Haven resident Rita Umile, and Jane Jervis, a family friend, said he “brought down the house” with his rendition of “The Girl from Ipanema” at an annual talent show last year.

An active member of the New Haven community, Ruddle helped found the nonprofit now known as HomeHaven, which organizes activities, assistance and other services for New Haven residents who choose to continue living in their homes as they get older. The first president of the board of directors for Science Park, established as a small business incubator on the site of an abandoned factory, Ruddle co-founded and helped launch several biotechnology companies. In 2000, he won the Connecticut Innovations Special Achievement Award for his contributions to the biotechnology industry in Connecticut.

He is survived by his wife of 48 years, Nancy Ruddle GRD ’68 — a professor emeritus at the School of Public Health and the School of Medicine — two daughters and three grandchildren.

This article was updated to reflect the  version published in print March 25.