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Clinton Edward Ballou
In Memoriam

Clinton Edward Ballou

Professor of Biochemistry and Molecular Biology, Emeritus

UC Berkeley

Clinton E. Ballou was a Professor of Biochemistry at UC Berkeley from 1955 to 1989, and, after reorganization of the biological sciences, Professor of Biochemistry and Molecular Biology until his retirement in 1991. He remained active until his laboratory closed in 1995. He died on March 8, 2021, having led a remarkable life.

Clint was born on June 18, 1923 in King Hill, Idaho, one of five children. His father and mother came from Nebraska, but moved to Idaho in 1912. In an unpublished memoir, As I Recall: A Personal History (2004), Clint recounted growing up in that small farming community during the Depression. The family moved to Boise, Idaho, in 1939, which opened new horizons for Clint. After finishing high school in 1940, he studied chemistry and biology at Boise Junior College (now Boise State University). His instructor in analytical chemistry, impressed with Clint's aptitude, recommended him for a scholarship to study Chemistry at Oregon State College (now Oregon State University). Clint went to Corvallis, Oregon, in fall 1942 and completed his final two undergraduate years there, including publishing his first two scientific papers in 1944.

The US had entered World War II in December 1941. Clint enrolled in a Navy program through which he could retain his college deferment from the military draft, finish his degree, and then be eligible for an officer's commission in the Navy upon completing boot camp. After receiving his B.S. in Chemistry, Clint was inducted into the Navy, finished boot camp, and was commissioned as an Ensign in the Naval Reserve in July 1944. After officer training and instruction in ship-board radar, he was assigned to the troop carrier USS Garrard as Assistant Communications Officer. Clint was granted a short leave to visit his parents after his older brother Bill, a gunnery sergeant, died in April 1945 when his B17 bomber crashed returning from a raid on Germany. One week after Clint's return to the USS Garrard on May 1, 1945, the war in Europe ended, but the Pacific conflict continued unabated. The USS Garrard was sent to sea for the invasion of Japan and ordered to Okinawa, where the crew engaged enemy airplanes with anti-aircraft fire. However, invasions of the main islands were averted after Hiroshima and Nagasaki were bombed, and Japan  surrendered unconditionally  on August 15, 1945. Thereafter, the USS Garrard shuttled prisoners of war from elsewhere in Japan to hospital facilities in Yokohama, and then shuttled returning troops from the Philippines back to the USA. In January 1946, Clint was promoted from Ensign to Lieutenant  and then released from active duty in July. Because of his service, Clint received educational benefits under the GI Bill, which supported his graduate education.

Clint was advised to apply to the University of Wisconsin, Madison, to study Biochemistry under Professor Karl P.G. Link, discoverer of the potent anti-coagulant dicoumarol. Clint was accepted and arrived in Madison,  only to find Link in a tuberculosis sanatorium near the campus. Nonetheless, for his Master's thesis, Clint investigated biodegradation of dicoumarol in rabbits. For his doctoral dissertation, he chose a project in synthetic carbohydrate chemistry aimed at understanding the structural basis for the alkali-sensitivity of the glycoside-coumarin conjugates excreted as biological breakdown products of dicoumarol.  He received his Ph.D. in 1950.

Two years earlier, in Fall 1948, Link had admitted to his laboratory his very first female graduate student, Dorothy Lun Wu. When Lun left China, her father told her, "Please do not marry an American." Lun and Clint were married in December 1949. In that era, a marriage involving a woman of Asian ethnicity so closely following the end of WW II was extraordinary. One can only marvel at the strength of the bonds of love that Lun and Clint shared. Indeed, they were an inseparable couple until Lun's death on June 2, 2017, at age 91.

Clint conducted one year of postdoctoral research in Edinburgh under Professor Edmund L. Hirst, followed by a second year with Professor Hermann O. L. Fischer at UC Berkeley, all supported by an NIH Fellowship. Lun and Clint traveled by ship both to and from Scotland. After finally arriving in Berkeley in early 1951, and securing a small apartment, their first child (daughter Linda) was born in January 1952 and their second child (son Philip) was born in June 1954.

When Clint started his work with Fischer, the Biochemistry Department Chair was Wendell M. Stanley, who came to UC Berkeley from the Rockefeller Institute (now Rockefeller University) two years after winning the 1946 Nobel Prize in Chemistry. The faculty was crowded onto the 3rd floor of Mulford Hall, so Clint was sent to the Forest Products Laboratory elsewhere in Mulford; consequently, he first collaborated with a colleague there to characterize what turned out to be inositol and related compounds in sugar pine heartwood.

In late 1951, the Biochemistry faculty moved into a newly constructed 4-story building on Gayley Road, designated the Biochemistry and Virus Laboratory, but later renamed Stanley Hall (demolished in 2005, and replaced in 2007). In the Fischer Lab, Clint worked closely with Donald L. MacDonald, an Instructor in Biochemistry. They devised novel syntheses for several phosphorylated sugars, like D-erythrose-4-phosphate, that were known or suspected intermediates in carbohydrate metabolism.

In 1954, Clint commenced exploring the academic job market. In the end, he accepted an appointment at Berkeley as an Assistant Professor of Biochemistry, effective July 1, 1955. With his tenure-track position, Clint was able to accept graduate students. He inherited two exceptional students from Fischer: Finn Wold, later a notable pioneer in the design and use of affinity labels and transition state analogues to elucidate enzyme structure and function; and Lewis I. Pizer, later recognized for path-finding work on a herpesvirus (HSV-2) that causes cervical cancer and on the varicella zoster virus that causes chickenpox and shingles.  

In December 1956, Clint and Lun moved their family into a new home they had built on Creston Road in the Berkeley hills, which they occupied for the ensuing half-century, until they moved to assisted living facilities for seniors.

Clint and his students were highly productive right off the bat. As a result, Clint was promoted to Associate Professor with tenure on July 1, 1957, just two years after assuming his initial appointment, a remarkable accomplishment, especially given that during the same period he served as an Assistant Dean in the College of Letters and Science. He was promoted to full Professor in 1962 and was Biochemistry Department Chair from 1964-1968. Clint also served as a member of several committees in the Academic Senate, including the Committee on Courses of Instruction and the Budget Committee. His service spanned over a decade.

In 1964, certain faculty in the Virus Laboratory formed the Department of Molecular Biology; and, in fall of the same year, those residents who were formally faculty in the Biochemistry Department moved into the new Biochemistry Building (later renamed Barker Hall). The unnatural rift between Biochemistry and Molecular Biology was not healed until July 1, 1989, with the creation of the Division of Biochemistry and Molecular Biology (now Division of Biochemistry, Biophysics and Structural Biology) in the current Department of Molecular and Cell Biology, all established through reorganization of the biological sciences, led by UC Berkeley biochemist Professor Daniel E. Koshland, Jr.

During the four decades that followed Clint's initial faculty appointment, and with students, postdocs, collaborators, and visiting scientists, Clint made ground-breaking contributions in several major areas of carbohydrate and polysaccharide, glycolipid, and glycoprotein biochemistry. He maintained an interest in sugar-phosphate synthesis and metabolism. He conducted path-finding work on phosphoinositides and inositol-polyphosphates. Clint also made novel discoveries about mycobacterial polymethylpolysaccharides and mycobacterial glycolipids because of their relevance to tuberculosis.

In the late 1960s, Clint initiated analysis of the cell walls of budding yeast Saccharomyces cerevisiae, especially the outermost mannoprotein layer. Appreciating the advantages of applying genetic analysis to his studies, Clint was one of ten students accepted into the inaugural Molecular Biology and Genetics of Yeast course at the Cold Spring Harbor Laboratory in summer 1970. Over the course of the next 20+ years, often with Lun as his collaborator (in 16 publications), Clint used a productive blend of genetic and biochemical analysis to elucidate the structures and functions of the glycosyl chains in yeast mannans.

Clint received prestigious accolades for his research accomplishments, including an NSF Senior Fellowship (1961), a Guggenheim Foundation Fellowship (1968), election to the US National Academy of Sciences (1975), named lectureships at the University of Notre Dame (1976), Duke University (1977), and elsewhere, receipt of the Claude Hudson Award in Carbohydrate Chemistry from the American Chemical Society (1981), and the Berkeley Citation from the University of California (1992). In 1994, an entire volume of the journal Carbohydrate Research was dedicated to papers written in Clint's honor.

There was something about Clint's countenance or demeanor that gave a misleading impression that he was remote, reserved, or unapproachable; his nickname among the undergraduates in his courses was "The Iceman." However, nothing could be further from the truth. To a person, his junior colleagues knew how kind, generous, and helpful Clint could be. Although Clint was always solicitous about not intruding on their business, he was incredibly helpful any time his guidance or assistance was asked for. Likewise, other members of the Biochemistry Dept. remember Clint's and Lun's warmth and generosity. On many Fridays at 4 PM, Clint and Lun hosted a floor-wide get-together that featured home-made ice cream prepared in an old-fashioned hand-cranked machine. Similarly, Clint and Lun generously shared with friends their enjoyment of good food and fine wines, and experimented with gourmet cooking on many memorable evenings in their home on Creston Road.

Lun and Clint greatly enjoyed international travel, and Clint took full advantage of his sabbatical leaves. In 1961, he took the family to Paris; for his second sabbatical in 1968, they made extended stays in Kyoto and again in Paris. In 1972, after Nixon thawed relations between the USA and China, Clint and Lun took their children to meet Lun's side of the family. In 1975, Clint conducted his third sabbatical in Basel, Switzerland; and, for his last in 1982-83, Clint spent six months in Grenoble, France, and then six months at Imperial College, London.

At a dinner held in honor of Clint and Lun on the occasion of his retirement, he read aloud a moving essay on his life, his formative influences, and how they meshed with the early years of the department and the past and current personalities within it. It was clear that this was Clint's way of eliciting respect and understanding for the department's history and of inculcating in those gathered there an enduring appreciation for the legacy of excellence we shared and whose continuity we were all charged with preserving into the future.

Jeremy Thorner, Professor Emeritus, BBS Div., MCB Dept., UC Berkeley
Jasper Rine, Professor, GGD Div., MCB Dept., UC Berkeley
Giovanna Ferro-Luzzi Ames, Professor Emerita, BBS Div., MCB Dept., UC Berkeley
Bruce N. Ames, Professor Emeritus, BBS Div., MCB Dept., UC Berkeley