Ursula Bellugi, a pioneer in the study of the biological foundations of language who was among the first to demonstrate that sign language was just as complex, abstract and systematic as spoken language, died on Sunday in San Diego. She died at the age of 91.

Rob Klima confirmed her death at the assisted living facility.

Dr. Bellugi was the director of the laboratory for cognitive neuroscience at the Salk Institute for Biological Studies in San Diego for much of the time he was there. She made significant contributions to three areas: the development of language in children, the linguistic structure and neurological basis of American Sign Language, and the social behavior and language abilities of people with a rare genetic disorder.

She leaves an indelible legacy of showing how humans communicate and socialize with each other.

Dr. Bellugi and her husband, Edward S. Klima, advanced understanding of the brain and the origins of language.

In 1960, William C. Stokoe Jr. described American Sign Language as a true language. He was ridiculed and attacked for that claim.

Dr. Bellugi and Dr. Klima were the first to show that the world's signed languages were actually languages in their own right.

The linguistic systems that were passed down from one generation to the next were established by Dr. Bellugi. She is considered the founder of the neurobiology of American Sign Language by the scientific community.

The left hemisphere of the brain has an innate tendency for language, even if you are not a linguist. Scientists were able to understand how the brain learns, interprets and forgets language.

This was a critical discovery for the deafness community, as it verified that our language is treated equally by the brain, just as we must be treated equally by society.

Until then, sign languages were seen as crude pantomime, with no rules, or as broken English, and children were discouraged from learning to sign. The work of the couple contributed to the acceptance of A.S.L. as a language of instruction and to the development of the Deaf Pride movement.

ImageDr. Bellugi with the celebrated deaf actress Marlee Matlin in 2009.
Dr. Bellugi with the celebrated deaf actress Marlee Matlin in 2009.Credit...Family photo
Dr. Bellugi with the celebrated deaf actress Marlee Matlin in 2009.

Williams syndrome was studied by Dr. Bellugi and her husband. She wanted to understand how the disorder, in which a set of about 20 genes is missing from one copy of a chromosomes, changed the brain and ultimately shaped behavior.

She helped paint a picture of the biology humans use to interact with the world around us.

In central Germany, Ursula Herzberger was born on Feb. 21, 1931, a center of science and technology. After fleeing Germany in 1934, her family settled in Rochester, New York, where her father became head of Kodak's optical research laboratories.

A special lens was developed by Mr. Herzberger that solved the color distortion in glass. Edith Herzberger was Ursula's mother.

Ursula majored in psychology at Antioch College and graduated in 1952. She married Piero Bellugi, an Italian composer and conductor, in 1953 and they had two sons before divorcing in 1959.

She moved to Cambridge, Mass., where she worked as a research assistant to Roger Brown, a psychologist at Harvard and the Massachusetts Institute of Technology who was studying how young children acquire language. She earned a doctor of education degree at Harvard in 1967, while raising her two sons as a single mother. One of her teachers was Dr. Klima.

She legally changed her name to Bellugi-Klima after they married. He began teaching at the University of California, San Diego. She taught at the campus of her husband's alma mater, the Salk Institute. She taught at San Diego State University.

She attracted a parade of research assistants and made a point of hiring many who were deaf.

Dr. Bellugi received many awards. She was elected to the National Academy of Sciences. She retired from her job at the age of 86.

She co-authored hundreds of papers and books with her husband. The signs of language was written by 10 associates. The Association of American Publishers hailed it the most outstanding book in the behavioral sciences.

Her brother, Hans Herzberger, her sister, Ruth Rosenberg, and her son, Rob, are all dead. The son of David Bellugi died.