"We human beings do not fall in love by telepathy: We have to move into proximity with each other," Dr. Perper wrote in Forum magazine in 1987.
"It is behavior, vivacity, that attracts people, not looks, beauty, not elegance of dress," he told L.A. Life in 1995.
Later, on learning that Japanese manga comics depicted courtship and sexuality differently than did most American comics at the time, Dr. Perper began to study and write about manga and anime. Those two modes, cartoons and animation with a science fiction or fantasy theme, he said, "provide ways to connect with young people and initiate conversations about sexuality."
He served as book review editor for the Journal of Sex Research, the Journal of Sex Education and Therapy, and Mechademia: An Annual Forum for Anime, Manga, and the Fan Arts.
Dr. Perper also wrote quirky fiction. He delighted in creating oddball comebacks to spam e-mail and devising humorous wordplay.
In recent years, his passion had been creating the adventure Web comic The Adventures of Princess Adele of Utopia ( www.princessadele.com) in collaboration with Martha Cornog, his wife of nearly 30 years, and the artist Jamar Nicholas.
After growing up in Greenwich Village, Dr. Perper earned undergraduate and doctoral degrees in biology and genetics, both from the City College of New York.
He spent four years in the pharmaceutical industry before joining the faculty at Rutgers, where he taught biology and did research from 1972 to 1979. His research with rats and gerbils involved observing reproductive and parenting behavior.
After obtaining the Guggenheim grant in 1980, he did independent research from home with his wife. He said he never tired of watching people flirt in singles bars. "If the magic is less mysterious than we thought, it is no less entrancing," he told Forum in 1987.
Surviving besides his wife is a nephew, Robert Daniel Ullmann.
A memorial will be at 5 p.m. March 14 at the Fleisher Art Memorial, 719 Catharine St. Burial is private.
Contributions may be made to the Fleisher Art Memorial via www.fleisher.org, or the Comic Book Legal Defense Fund via http://cbldf.org/.