Joel E. Keizer

Keizer joined the UC Davis faculty in 1971 as an assistant professor of chemistry and by 1978 had achieved the level of professor. In the following years, his interest turned to mathematical modeling of biological processes, and in 1993 he was given a joint appointment as professor of biology. Highly unusual for a theoretical biologist, his biological modeling was very much appreciated by experimental biologists. 

Keizer was an affectionate man with a deep love for his family and friends. He embraced all aspects of life with a particular passion for the outdoors, whether he was fly fishing with his son on the McCloud and Deschutes Rivers, poking in tidepools for nudibranchs, or hunting for Indian arrowheads in the Warner Mountains. He regularly returned to his beloved Oregon Coast to spend time fishing, hiking and walking on the beach with his wife and daughter. His annual pilgrimage to Oregon allowed him to return to his roots with family and friends. 

Keizer was born Aug. 31, 1942, in North Bend, Ore. He attended Beaverton High School in Beaverton. Ore., just outside of Portland, and graduated as president of his class in 1960. Aspiring to be a historian, he nevertheless excelled at science. When he entered Reed College in Portland, he declared as a chemistry major. He graduated Phi Beta Kappa with a BA degree in chemistry in 1964. Having won a National Science Foundation Graduate Fellowship, he went to Princeton University to study biochemistry. Unhappy with the program at Princeton, he soon returned to Oregon to attend the University of Oregon and to marry Susan Swank, a classmate from Reed. 

He first worked with Sidney Bernhard in experimental biochemistry and then switched to theoretical chemistry, working with Terrell Hill. He received his Ph.D. degree in chemical physics in 1969. While publishing two experimental papers with Bernhard in the journal Biochemistry in 1966, his thesis work was highly mathematical and appeared in two papers in the Journal of Statistical Physics in 1970. This counterplay between experiment and theory would characterize his work throughout his career. As a postdoctoral fellow at the Battelle Memorial Institute in Columbus, Ohio, from 1969 to 1971, he developed his theoretical skills and began a long series of papers, many in the Journal of Chemical Physics. He began his academic career at UC Davis in 1971 in the chemistry department and rapidly rose in rank to full professor in 1978, a post held until 1993. 

While most of his work was the theoretical, there were occasional collaborative excursions into experiment. Throughout the late '70s and early '80s, his interest was in fluctuations and non-equilibrium thermodynamics. He pioneered an approach to the thermodynamics of non-equilibrium steady states and collaborated on experiments to justify his reasoning. This stage of his scientific career culminated in his very successful monograph: "Statistical Thermodynamics of Nonequlibrium Processes" (Springer-Verlag, New York 1987), which was later translated into Russian. By the time of its publication, he had more than 60 journal publications to his credit. 

In 1983, 1985 and 1988, papers appeared in which he collaborated on research concerning pancreatic beta-cell oscillations. Keizer was beginning to reinvent himself, this time as a mathematical biologist. During the '90s, this type of research dominated his activities, so much so that he felt he had to associated himself with biologists on a regular basis. In 1993 he left the chemistry department and became a professor of biology. His model for insulin secretion published in The Proceedings of the National Academy of Sciences in 1992 has been cited almost 100 times, most often by experimentalists. His work on calcium oscillations and diffusion in cells resulted in a burst of papers during the '90s, which have already had a major impact on the research community. Keizer was at the peak of his powers in this work when cancer overcame him.

 Dissatisfied with the usual working environment for research in the university setting, Keizer established the Institute of Theoretical Dynamics (ITD) at UC Davis in 1986 and was its director until his death. This interdisciplinary think tank brought together UC Davis faculty from many different departments, visiting from around the world and was a training ground for young scientists. 

The Biology Division of the National Science Foundation selected ITD for a research training grant in 1997. Together with the matching funds from UC Davis this grant amounts to over $3 million for five years. Keizer's own research on calcium waves in frog eggs was funded by the NIH at the million-dollar level when he died. Keizer held a prestigious John Simon Guggenheim Memorial fellowship in 1986-1987 and was named a Fellow of the American Association for the Advancement of Science (AAAS) in 1995. He was an invited speaker at many international meetings throughout his career. He published more than 120 journal articles and book chapters, and his research grants over his career totaled millions of dollars. He had roughly two dozen graduate students and post-doctoral associates. He will be sorely missed by the scientific community and by his many personal friends. 

He is survived by his wife of 35 years, Susan Swank Keizer; his children, Sidney Jacob Keizer and Sarah Rebecca Keizer, both of San Francisco; his parents, Dr. and Mrs. John Keizer of King City, Ore.; and his siblings, Mary Lynn Duvall and Lewis Keizer. 

The family requests that any contributions be made to UC Davis Foundation/ Joel Keizer Memorial Fund, University of California, One Shields Avenue, Davis, CA 95616. 

Originally published in The Davis Enterprise on May 23, 1999.