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In 2010, he turned against inclusive fitness, publishing a paper attacking the concept with Martin A. Nowak of Harvard and Corina E. Tarnita, now at Princeton.

“The basic foundations of inclusive fitness theory are unsound,” Dr. Wilson said in a 2012 interview.

Richard Dawkins, a leading exponent of inclusive fitness, ruthlessly attacked Dr. Wilson in a review of “The Social Conquest of Earth,” which he said was full of “downright perverse misunderstandings.”

Mr. Hamilton described “inclusive fitness,” as this concept came to be known, in a 1963 paper.

Many people have posed explanations for this confusing genetic inheritance, such as the idea from Sabine Eggers and her colleagues that childless women have inclusive fitness because they help raise other women’s children.