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The Rev. Federico Lombardi, the Vatican spokesman, praised Dr. Levi-Montalcini’s civil and moral efforts, saying she was an “inspiring” example for Italy and the world, the ANSA news agency said.

Pietro Calissano, an Italian scientist who worked for some 40 years with Dr. Levi-Montalcini, including in the United States, said the work the Nobel laureate did on nerve growth factor was continuing. The protein assists portions of the central nervous system that have been damaged by disease or injury.

“Over the years, this field of investigation has become ever more important in the world of neuroscience,” Mr. Calissano was quoted by ANSA as saying. Mr. Calissano began studying under Dr. Levi-Montalcini in 1965 and recalled her ability to relate to students on a very human level, with none of the elite airs that often characterize Italian professors.

“I remember we were in a closet with cell cultures when she offered me a fellowship,” Mr. Calissano said. He added that research building on Dr. Levi-Montalcini’s pioneering achievements continues. “We are working on a possible application in the treatment of Alzheimer’s,” he added.

The 1943 German invasion of Italy forced the Levi-Montalcini family to flee to Florence and live underground. After the Allies liberated the city, she worked as a doctor at a center for refugees.

In 1947 Dr. Levi-Montalcini was invited to the United States, where she remained for more than 20 years, which she called “the happiest and most productive” of her life. She held dual Italian-U.S. citizenship.

During her research at Washington University in St. Louis, she discovered nerve growth factor, the first substance known to regulate the growth of cells. She showed that when tumors from mice were transplanted to chicken embryos, they induced rapid growth of the embryonic nervous system. She concluded that the tumor released a nerve-growth-promoting factor that affected certain types of cells.

The research increased the understanding of many conditions, including tumors, developmental malformations and senile dementia. It also led to the discovery by Stanley Cohen of another substance, epidermal growth factor, which stimulates the proliferation of epithelial cells. The two shared the Nobel Prize for medicine in 1986.

Dr. Levi-Montalcini returned to Italy to become the director of the laboratory of cell biology of the National Council of Scientific Research in Rome in 1969.

After retiring in the late 1970s, she continued to work as a guest professor and wrote several books to popularize science. She created the Levi-Montalcini Foundation to grant scholarships and promote educational programs worldwide, particularly for women in Africa.

In 2001 Dr. Levi-Montalcini was made a senator-for-life, one of the country’s highest honors.

She then became active in parliament, especially between 2006 and 2008, when she and other life senators would cast their votes to back the thin majority of center-left Premier Romano Prodi.

Dr. Levi-Montalcini had no children and never married, fearing such ties would undercut her independence.

“I never had any hesitation or regrets in this sense,” she said in a 2006 interview. “My life has been enriched by excellent human relations, work and interests. I have never felt lonely.”

Italian mathematician Piergiorgio Odifreddi said he was always struck by the contrast of this “petite, frail woman and the power of her mind.” He recalled comments that Dr. Levi-Montalcini made when she turned 100. She mentioned that she would sleep no more than two or three hours a night because “I have no time to lose,” Mr. Odifreddi told Sky TG24.

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