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Plant biologist Paul Green dies at age 67

Paul B. Green spent much of the last decade trying to answer a single question: How do biological patterns develop out of nothing? The medium for his exploration was the curve of a rose petal, the right-angle alternation of flower buds on the snapdragon, the double-spiral in the face of a sunflower and the scales of a pine cone.

Although he was an expert in plant physiology and development, a field that is currently dominated by molecular biology and genetics, Green held the unorthodox opinion that the basic patterns that are repeated over and over in the plant world are not due solely to gene action but to the mechanical properties of the tissues involved.

In a special symposium honoring him and his work that took place about two weeks before his death on Aug. 18 from pancreatic cancer, Green was surprised and pleased to learn that his research, which spans 40 years at the University of Pennsylvania and Stanford, has had a lasting impact on this field of study, termed plant morphogenesis.

"Paul Green has powered the field of plant morphogenesis and developmental biology to a new level of insight from which we cannot retreat, and which will affect our teaching of the subject for years to come, perhaps forever," commented Phil Lintilhac of the University of Vermont.

Hans Kende, an eminent plant biochemist at the Michigan State University Department of Energy Plant Research Laboratory, who wasn't able to attend the hastily scheduled meeting, wrote the symposium organizers that the uniqueness of Green's scientific contributions are well recognized. "There are many Hans Kendes in this world, i.e., biochemically wired people who work on hormone biosynthesis and action. There is one Paul Green."

Peter Ray and Virginia Walbot, friends and fellow faculty members in the biological sciences department, organized the symposium "Mechanisms of Plant Development" in his honor. About 80 scientists from around the country attended.

"Even I was surprised at what a major influence he has had," said Ray, a longtime friend and plant biologist. "Speaker after speaker stood up and shared how Paul had influenced them and what they have done as a result. Paul himself was taken by surprise, because he often felt that he was a lone voice. He did not realize how many people he had affected."

The extent of his influence was formally recognized the week following the symposium when the Botanical Society of America gave Green its prestigious award of merit for "his innovative, technically brilliant and analytically fundamental work on plant morphogenesis."

As Dorota Kwiatkowska, a visiting scholar in Green's lab from Wroclaw University in Poland, put it, "His models are now the basic textbook models. I haven't seen a recent book on phyllotaxis [the pattern of arrangement of leaves on a stem] that doesn't include his theories."

According to Ray, Green did not mentor an exceptionally large number of students, but he had a profound effect on those that he did. When she heard that he had cancer, one of his previous students, Dina Mandoli, now at the University of Washington, set up a "Paul Green Fan Club" that kept former students and colleagues informed of his condition by e-mail. Letters and e-mails of support poured in.

One of those who responded to the news was Zac Cande, now a member of the Department of Molecular and Cell Biology at the University of California-Berkeley. "I just wanted to say what an important influence you were with regards to my scientific life - as the twig is bent and so forth. . . . From you I got the realization that it was important to pay attention to the big picture as well as the small things, to think about and develop models that you should try and test, to look for the offbeat as well as the tried and true," he wrote.

Green's influence on his graduate students went well beyond the academic. Many of them lived in the big white stucco house that soon gained the nickname "Ma Green's International Hotel." Many discussions of plant development that started in the laboratory were continued over the kitchen table, where Green did much of his thinking and writing. An avid camper and mountaineer since childhood, Green and his family introduced many of his students, particularly those from other countries, to the pleasures of hiking, cross-country skiing and camping.

As Anne Sylvester, now a professor at the University of Idaho, wrote to the Greens, "Some people influence others and bring knowledge and joy to the world and you are both such people to me. I remember the time spent in Paul's lab and in your home as being turning points in my life, when I learned to think deeply, to live fully."

In addition to his research and work with students, Green was an active campus citizen. He served on a number of departmental committees, including facilities, admissions, undergraduate studies and graduate programs. At the campus level he served on the Faculty Senate, the Committee on Academic Appraisal and Achievement, and the Committee on Undergraduate Admissions and Financial Aids.

Green was a favorite of the staff in the biological sciences department. "He will really be missed," said Bettye Price, the financial manager. "I've never worked with such a kind man. The staff felt very highly of him. He was just wonderful."

In a memo to department staff upon learning of his death, Walbot spoke for many when she wrote, "Paul's insights, humanity and goodwill toward us all will be sorely missed. Paul had singular insights into the biophysics of plant growth and rigorous standards of proof. No matter what the topic, his quips and immediate puns made all talks memorable. Paul was a gracious human being, a real gentleman of science."

Paul Barnett Green was born in Philadelphia on Feb. 15, 1931. He died at his Stanford home on Aug. 18 at the age of 67. He was educated at the University of Pennsylvania and received his doctoral degree from Princeton. He was then elected to the Society of Fellows at Harvard. He spent his three-year tenure as a fellow at Harvard and the Swiss Federal Technical Institute working with the giant-celled freshwater alga Nitella. Using this as a model system, he was the first person to measure the pressure within a plant cell. At the end of the fellowship, Green joined the faculty at the University of Pennsylvania, where he taught for 12 years before moving to Stanford in 1971.

Green's professional honors include the Darbaker Prize and Pelton Award given by the Botanical Society of America, a Guggenheim Fellowship, a foreign membership in the Belgian Royal Academy of Sciences and Fine Arts and a fellowship in the Japanese Society for the Promotion of Science.

He is survived by his wife, Margaret, and three married children: Robert Green and Ann McKeown and their three daughters, Nancy, Stephanie and Amy, of Peaceful Valley, Calif.; Peter Green and Lorraine Hwang and their son, Stewart, and daughter, Jessie, also of Peaceful Valley; Kate Green and Wally Beaton of Ottawa, Canada; and his sister, Elenor Wise, of Butler, Pa.

A memorial service will be held Sept. 1 at 3 p.m. at the First Congregational Church of Palo Alto (Louis and Embarcadero Roads) with a reception to follow at the church. The family requests that those who wish to send memorial gifts do so to the Jasper Ridge Biological Preserve (Stanford University, Stanford CA 94305) or the Unitarian Service Committee of Canada (56 Sparks Street, Suite 705, Ottawa, Ontario, Canada K1P 5B1).


By David F. Salisbury

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