His death, at a residential care facility, was caused by complications of the new coronavirus, his wife, Robin Kennedy, said. He had suffered a severe stroke in 2015.

Stanford had been Kennedy’s life since 1960, when, not yet 30, he joined its faculty as an assistant professor of biology. And except for a stint in the late 1970s as head of the FDA under President Jimmy Carter, he remained wedded to the university, becoming provost and then president in 1980, beginning an 11-year tenure.

It was a productive one. During his presidency, the university opened the Stanford Humanities Center and campuses in Oxford, England; Kyoto, Japan; and Washington; diversified the Western culture curriculum; and raised $1.2 billion in a five-year centennial campaign, although by the end of the decade the university was facing deficits.

His tenure also coincided with fiery debates over anti-war protests and academic freedom by both professors and students, divestiture of the university’s holdings in companies doing business in South Africa, and $160 million in damage inflicted by the Loma Prieta Earthquake in 1989.

A would-be writer who had become a neurobiologist in college adventitiously, Kennedy found his leadership under the microscope in the early 1990s, when the university was accused — and later cleared — of improperly billing the Navy for research expenses.

The accusations were aired by federal auditors and Rep. John D. Dingell Jr., a tenacious Michigan Democrat, who said that Stanford may have billed the government for as much as $200 million in improper expenses on research contracts for more than a decade.

By 1994, Stanford had agreed that a total of about $3 million had been inadvertently billed to the government, but the federal auditors concluded that there was no evidence of misrepresentation by the university.

Still, the damage was done to Stanford’s reputation, and Kennedy resigned in 1991, attributing the government accusations to political and personal vendettas and acknowledging that they had contributed to his decision to step down.

“It is very difficult, I have concluded, for a person identified with a problem to be the spokesman for its solution,” he said in announcing his resignation. He went on to edit the journal Science.

But he had his ardent supporters on the Bay Area campus, where he was known to bike to work and engage with students. Among them was his protégé Cory Booker, the future senator from New Jersey, whom Kennedy had encouraged to apply for a Rhodes Scholarship.

“To watch him lead through the indirect cost crisis, through professional and personal attacks, under tremendous stress and strain, with clouds amassed over his head and challenges raining on him," Booker wrote in the foreword to Kennedy’s memoir, “A Place in the Sun” (2017), “was a study in leadership, character, and discipline, always better shown in times of crisis than when all is going well.”

Donald Kennedy was born Aug. 18, 1931, in Manhattan to William and Barbara (Bean) Kennedy. His father was a writer, an editor and an assistant dean of the Harvard Business School. His mother was a teacher and journalist.

As his father repeatedly switched jobs, Donald was raised in about a half-dozen locales, including Greenwich, Connecticut, by the time he was 15.

After graduating from the Dublin School in New Hampshire, he enrolled in Harvard University intending to major in English and be a writer; at one point he received an A on a 5,000-word final paper in creative writing. But, as he recalled in his memoir, his professor, perhaps pointing him toward a more profitable profession, asked him over sherry one night: “ Tell me, Don. What else interests you?’”

“Surprised by the question, I gathered my wits and responded, ‘Well, biology and natural history, I guess.’ ”

“ ‘Biology,’ he said. ‘That sounds like a wonderful choice.’ ”

He earned his bachelor’s degree in 1952, followed by a master’s and a doctorate, all three from Harvard. And all in biology.

Kennedy was recruited to the FDA in 1977 by Joseph A. Califano Jr., the secretary of Health, Education and Welfare. During his tenure there the agency’s proposed ban on saccharin, the artificial sweetener, was defeated, but overall his record won plaudits from industry representatives and consumer advocates alike.

He returned to Stanford briefly as provost before he was named president.

Kennedy was a familiar presence on campus, not only biking to the quadrangle but also inviting students to join him on his morning runs up to the Dish, the radio antenna in the foothills of the campus.

“Kennedy is not someone whom students hear once when they arrive and then once when they graduate,” The Stanford Daily, the student newspaper, editorialized in 1991.

A former student, Ingrid Schwontes Jackoway, was quoted as saying in an alumni publication: “I will never forget Donald Kennedy getting up on the lab table at the front of the lecture hall and assuming a quadruped position to demonstrate to us the concepts of dorsal, ventral, cephalo and caudal. His first concern was always with teaching effectively, not preserving his dignity.”

Kennedy’s marriage to Jeanne Dewey ended in divorce. In addition to his wife, Robin Hamill, who was associate counsel at Stanford when they married in 1987, he is survived by two daughters from his first marriage, Page Kennedy Rochon and Julia Kennedy Tussing; two stepchildren, Cameron Kennedy and Jamie Hamill; his brother, Dorsey; and nine grandchildren.

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Kennedy was the editor-in-chief of Science, the weekly journal of the American Association for the Advancement of Science, from 2000-08. But even there he was not immune to controversy. Researchers had fabricated their findings in several articles, and a reported sighting of an extinct ivory-billed woodpecker appeared to have been mistaken.

Among his other books were “The Cold and the Dark: The World After Nuclear War” (1984) with Carl Sagan and Paul R. Ehrlich, and “Academic Duty” (1997). At his death he was Bing professor for environmental science emeritus at Stanford.

Shortly after he became president, Kennedy told the student radio station, KZSU, that he intended to keep his perspective despite the pressures of the job.

“The president is ultimately the person to whom the problems come,” he said. “What you need then is to walk around, or visit a dormitory, or to give a class, or to meet a student who wants to come in and talk about a career choice. I find those occasions very uplifting because they’re not automatically negative. They’re not the kind of problems that are programmed for the president’s desk because they haven’t been solved by anybody else. Instead, they’re the kinds of things that go on around here day by day, and make this a terrific place.”

This article originally appeared in The New York Times .