In Memoriam: Clinton Edward Ballou (1923–2021)

It is with great sadness that we share the sorrowful news about the passing on March 8, 2021, of our colleague and friend Clinton E. Ballou, Ph.D. Clint was Professor of Biochemistry (1955-1989), and, after our reorganization of the biological sciences, Professor of Biochemistry and Molecular Biology until his retirement in 1991, and then still active in research as a Professor Emeritus until he closed his laboratory in 1995, here at the University of California, Berkeley. He led a remarkable and remarkably impactful life.

Clint was born on June 18, 1923 in King Hill, Idaho, one of five children (himself, two brothers and two sisters). His father (William C. Ballou) and mother (Mollie Bernt) were born, raised and married in Nebraska, and then moved to Idaho in 1912, where William had been offered a position as a Section Foreman on the Union Pacific Oregon Short Line railroad. In a memoir Clint prepared for his own family [Ballou CE (2004) As I Recall: A Personal History], he recounted what it was like to grow up in a small-town farming community in the Depression era, hard by the Snake River, and the impact of his parents' decision in 1939, when he was sixteen, to move the family to Boise, Idaho, which Clint felt opened up a new world of opportunity for him. Clint finished high school in Boise and, in 1940, entered Boise Junior College (now, as of 1974, Boise State University), where he studied chemistry and biology in their Premedical Program. In his second year, Clint's instructor in analytical chemistry, who held a Ph.D. in Chemistry from Oregon State College (now, since 1961, Oregon State University), impressed with Clint's aptitude for the subject, recommended Clint for a scholarship to attend Oregon State College. Clint arrived in Corvallis, Oregon, to complete his final two years at Oregon State College in the fall of 1942. In his senior year, his Biochemistry Professor took Clint under his wing and the research he conducted there led to his first two scientific publications, both of which appeared in 1944 in Archives of Biochemistry (now, since 1952, Archives of Biochemistry and Biophysics).

The military draft had been introduced in September 1940 and, when Clint turned eighteen, he was required to register. The US entered World War II in December 1941, but Clint was granted a deferment because he was still in college. He learned of a Navy program by which he could retain his deferment, finish college through an accelerated program, then be inducted into the Navy as an able-bodied seaman, but be eligible for an officer's commission after completing boot camp. He completed all the requirements for his B.S. in Chemistry in March 1944 and then applied for a commission in the Naval Reserve. He was inducted into the Navy in Boise on April 1944, and reported to US Naval Training Station at Farragut, Idaho, on May 1944 for boot camp. The fact that Clint's given middle name at birth was actually Edgerton (a family name on his father's side) first came to light when he was required to provide a copy of his birth certificate upon his enlistment in the Navy; while growing up, he had been told, erroneously, that his middle name was Edward and, therefore, had used that on all official documents ever since.

Clint received his uniformed officer's commission (Ensign in the Naval Reserve) in July 1944, and reported to Naval Officer Training School in Tucson, AZ. Two months later, he was sent to the Naval Communications School in Hollywood, FL (near Miami Beach) for two months of training in the maintenance and use of ship-board radar. In January 1945, he was assigned to the attack transport ship USS Garrard (APA 84); APA ("Auxiliary Personnel, Attack") was the Navy's designation for the ships that carried troops into battle and from which separate landing craft were loaded. Before the USS Garrard was formally commissioned, Clint was sent to the Small Craft Training Center on Terminal Island, California (off the coast of Los Angeles). He moved aboard the USS Garrard in early March 1945, where his assignment was to serve as the Assistant Communications Officer and report to the Officer of the Deck. From southern California, the USS Garrard sailed to Seattle, Washington. Clint was granted a one-week leave to visit his parents in Boise when he learned that his older brother Bill, a gunnery sergeant on a B17 bomber, had died in April 1945 when his plane crashed in Belgium while returning from a raid on Germany. After reporting back to the USS Garrard, on May 1, 1945, the war in the European theater ended one week later, but the conflict in the Pacific continued unabated. Therefore, at the end of the first week in May 1945, the USS Garrard set to sea to be part of the plans for the invasion of Japan. The first port of call was Honolulu, Hawai'i, then Eniwetok in the Marshall Islands, then onward to Guam, and finally to anchorage in an isolated volcanic formation (Ulithi Atoll) in the Caroline Islands. Later, in reminiscing about his days on the ship, Clint recalled that the captain was prone to seasickness and frequently asked Clint to take the helm, so that the captain could go to bed. At the end of June, the USS Garrard was sent to Okinawa during the last stages of the invasion of that southernmost Japanese island, where Clint and the rest of the crew engaged enemy airplanes with anti-aircraft fire. In July 1945, the USS Garrard was sent to Leyte Island in the Philippines and then back to Eniwetok to prepare for an imminent ground invasion of mainland Japan. However, the wait for the invasion ended abruptly when the atomic bombs were dropped on Hiroshima (August 6, 1945) and Nagasaki (August 9, 1945), resulting in the unconditional surrender of Japan on August 15, 1945.

The USS Garrard was ordered to Tokyo and reached that harbor on August 30, 1945, and then sent up the east coast of Honshu to Sendai to take aboard American, British, Australian, New Zealand, Dutch and Chinese prisoners of war who had survived Japanese prison camps, whom they delivered back to Yokohama (at the southern end of Tokyo Bay), where there were hospitals set up to provide proper medical care. The USS Garrard then sailed from Japan on October 14, 1945, arriving at Portland, Oregon, on October 25, which they then departed on November 9, arriving in San Francisco on November 11. The USS Garrard then embarked for Manila in the Philippines on November 19, arriving on December 11 to pick up a load of returning troops; the ship departed on December 14 and returned to San Francisco on January 3, 1946. After two weeks of repairs at Mare Island Naval Shipyard in San Pablo Bay near Vallejo, CA (which closed in 1996), the USS Garrard docked at Hunters Point Naval Shipyard in San Francisco (which was decommissioned in 1974), and finally returned to Seattle on February 6, remained at anchor in Puget Sound until April 1, then docked in Bellingham, Washington, where the USS Garrard was officially decommissioned on April 16, 1946.

In January 1946, Clint was promoted from Ensign to Lieutenant (jg) and, on April 17, 1946, after the formal decommissioning of the USS Garrard, was given a month's leave and ordered to report to the Commandant, 12th Naval District, in San Francisco on May 19, 1946. He was assigned to do "busy work" in a Naval Office in the Ferry Building until he was released from active duty on July 16, 1946 (however, Clint's association with the Navy was not completely and formally terminated until June 18, 1956— his 33rd birthday —when, via a letter from the Acting Secretary of the Navy, he was informed that he had been recommended for and had received an Honorable Discharge from the Naval Reserve). Because of his military service, Clint became eligible for educational benefits under the GI Bill, which he used to very good advantage to support his graduate education.

Although Clint's tour of duty with the Navy placed his association with science on hold for two and a half years, on the advice of his old Biochemistry Professor at Oregon State College and others, Clint applied for admission to the University of Wisconsin, Madison, to study Biochemistry under the tutelage of Professor Karl P.G. Link (1901-1978), discoverer of the potent anti-coagulant dicoumarol [3,3'-methylenebis-(4-hydroxycoumarin)]. Clint was accepted and arrived in Madison, Wisconsin, in September, only to find Link in a tuberculosis sanatorium located on the north shore of Lake Mendota opposite the University campus. Nonetheless, for his Masters thesis, Clint investigated biodegradation of dicoumarol in rabbits. For his Doctoral Dissertation, Clint chose a project in synthetic carbohydrate chemistry aimed at understanding the structural basis of the alkali-sensitivity of the glycoside-coumarin conjugates excreted as biological breakdown products of dicoumarol [Ballou CE (1954) Alkali-sensitive glycosides. Adv. Carbohydr. Chem. 9: 59-95]. He was awarded his Ph.D. in Biochemistry in January 1950.

Two years before, in Fall 1948, Link admitted to his laboratory his very first female graduate student, Dorothy Lun Wu. Lun had come to America on a US government-sponsored scholarship that was part of a post-war program to retain good relations with China, which had been an ally against Japan. Ironically enough, when Lun left China, her father had told her, "Whatever you do, please do not marry an American." Lun and Clint were married in December 1949. Given that era, a bi-racial marriage, especially involving a woman of Asian ethnicity so closely following the end of WW II, was quite extraordinary. So, one can still marvel at the strength of the bonds of love that Lun and Clint shared and gave them the courage to wed under those circumstances. Indeed, they were an inseparable couple until Lun's death on June 2, 2017, at age 91.

With the completion of his doctoral dissertation looming, and with the desire to further expand his competence in carbohydrate chemistry and biochemistry, Clint applied to the National Institutes of Health for a postdoctoral fellowship to support his study for a year in Edinburgh, Scotland, with Professor Edmund L. Hirst (1898-1975), followed by a second year with Professor Hermann O. L. Fischer (1888-1960) at the University of California, Berkeley. His proposal was funded; so, Lun and Clint traveled by ship to Scotland in the summer of 1950. In the Hirst Laboratory, Clint learned and mastered the then-new and important methods of paper and column chromatography. By the time of their return sea voyage, Lun was pregnant with their first child (daughter Linda). Arriving in New York, they traveled westward by train, with stops in Madison and Boise, to reach California. After arriving in Berkeley in early 1951, and securing a small apartment, Linda was born in January 1952 and their second child (son Philip) was born in June 1954.

At the time Clint arrived for his work with Fischer, the then-Biochemistry Dept. was chaired by Wendell M. Stanley (1904-1971), who had come to UC Berkeley from the then-Rockefeller Institute (now, since 1965, Rockefeller University) just two years after winning the 1946 Nobel Prize in Chemistry; and, most of the faculty was crowded onto the 3rd floor of Mulford Hall. Given the space crunch, Clint was given his spot elsewhere in the building in the Forest Products Laboratory; consequently, he first collaborated with a colleague there to characterize what turned out to be inositol and related compounds in sugar pine heartwood [Ballou CE, Anderson AB (1953) On the cyclitols present in sugar pine. J. Amer. Chem. Soc. 75: 648–650 (1953); Ballou CE (1958) Inositol and related compounds (cyclitols). In The Metabolism of Secondary Plant Products (Encyclopedia of Plant Physiology, Vol. 10), Springer Verlag, Berlin, pp. 442-453].

In late 1951, all of the Biochemistry Dept. faculty moved into a newly constructed 4-story building on Gayley Road across from the Greek Theater on the east side of the campus, then designated the Biochemistry and Virus Laboratory, but later renamed Stanley Hall (demolished in 2005, and replaced by a much larger research and teaching edifice of the same name in 2007). There, in the Fischer Lab, Clint worked closely with Donald L. MacDonald (1922-2006), an Instructor in Biochemistry, who was also a member of the Fischer group. Together, they devised novel syntheses for several otherwise unavailable phosphorylated sugars that were known or suspected intermediates in various pathways of biological carbohydrate metabolism, in particular D-erythrose 4-phosphate [Ballou CE, Fischer HOL, MacDonald DL (1955) The synthesis and properties of D-erythrose 4-phosphate. J. Amer. Chem. Soc. 77: 5967-5970], which they provided freely to other investigators, including noteworthy biochemists Bernard L. Horecker (1914-2010) and Efraim Racker (1913-1991).

By 1954, Clint began to explore the academic job market. He was invited to apply for a position in the new Cellulose Research Institute at the State University of New York, Syracuse, and was also interviewed for a faculty position for a Biochemist in the Chemistry Dept. at Stanford University. Concomitantly, the UC Berkeley Biochemistry Dept. had instituted a search to replace Fischer, who was due to retire in 1956. Perhaps the interest shown by Stanford provoked his soon-to-be colleagues to act, and Clint was offered and accepted appointment as an Assistant Professor of Biochemistry, effective July 1, 1955. Not long after, with Fischer's retirement in 1956 and Don MacDonald moving to the National Institutes of Health in 1957 and then taking a regular faculty position in the Dept. of Biochemistry and Biophysics at Oregon State University in 1962, Clint was the only biochemist left at Berkeley pursuing research on carbohydrates.

With his tenure-track position secured, Clint was able to accept graduate students and commence his own independent research program, which he did by inheriting two exceptional students who were, in essence, abandoned by Fischer and who eventually became outstanding scientists in their own right: Finn Wold (1928-1997), who became 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 (1932-2006), recognized for his path-finding work on a herpesvirus (HSV-2) that causes cervical cancer and on the varicella zoster virus that causes chickenpox and shingles. With a stable income also secured, in December 1956, Clint and Lun moved their family of four out of their apartment and into a new home that they had built on Creston Road, which they then occupied for the ensuing half-century, until they moved to assisted living facilities for seniors in Oakland and later Alameda, CA.

In 1957, Wold and Ballou published two papers on the essential glycolytic enzyme enolase; and, not long after, Pizer and Ballou published a definitive study of another required glycolytic enzyme, phosphoglycerate mutase. Clint's third student Francis Lane (who married Pizer) and her spouse undertook studies of the inositol-phosphate component of animal cell phospholipids [Pizer FL, Ballou CE (1959) Studies on myo-inositol phosphates of natural origin. J. Amer. Chem. Soc. 81: 915-921; Ballou CE, Pizer LI (1959) Synthesis of an optically active myo-inositol 1-phosphate. J. Amer. Chem. Soc. 81: 4745], which were follow-ups, of sorts, to Clint's earlier work on the cyclitols in wood and began his on again-off again interest in the phosphoinositide lipids and inositol polyphosphates, especially myo-inositol 1,4,5-trisphosphate (IP3) and phytic acid (myo-inositol-hexakisphosphate or IP6) [Tomlinson RV, Ballou CE (1962) Myo-inositol polyphosphate intermediates in the dephosphorylation of phytic acid by phytase. Biochemistry 1: 166-171], about all of which he reminisced in a later retrospective essay [Ballou CE (2004) My brief encounter with the phosphoinositides and IP3. J. Biol. Chem. 279: 54975-54982].

Clint was promoted to Associate Professor with tenure, as of July 1, 1957, just two years after assuming his junior faculty appointment, a remarkable accomplishment, especially given the fact that during the same period he was also saddled with the additional obligation of serving as an Assistant Dean under two different Deans of the College of Letters and Science, botanist Lincoln Constance (1909-2001) and physicist William B. Fretter (1916-1991). He was promoted to full Professor in 1962 and served as Chair of the Biochemistry Dept. from 1964-1968.

In 1964, the Virus Laboratory in Stanley Hall and its main members became a new academic unit, the Department of Molecular Biology and the Virus Laboratory; and, in Fall of the same year, those members who were formally faculty in the Department of Biochemistry moved into a new six-story research and teaching facility, the Biochemistry Building [renamed Barker Hall in 1988 in honor of near-legendary UC Berkeley biochemist Horace A. Barker (1907-2000)], at the corner of Hearst & Oxford Streets on the west side of the campus. Clint secured a portion of the funds necessary to construct the Biochemistry Building through a grant request he wrote to the NIH for this purpose. In fact, Clint always had a good eye for those aspects of building facilities that would best serve the common good. For example, during planning for construction of Koshland Hall adjacent to Barker, Clint fought for preservation of adequate space for electrical, machine, and wood-working shops on the ground level. Likewise, during planning for the construction of the nearby Genetics and Plant Biology Building (GPBB), he was instrumental in making sure that its associated lecture hall (100 GPBB, now E.M. Cox Auditorium) had adequate capacity (204 seats), instead of an initially proposed space half that size. 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, as of 2011, the Division of Biochemistry, Biophysics and Structural Biology) in the current Department of Molecular and Cell Biology, all established as part of the reorganization of the biological sciences here, led by the wise, energetic and influential UC Berkeley biochemist Professor Daniel E. Koshland, Jr. (1920-2007).

In 1967, when Clint was Chair and the new Biochemistry Building was fully operational, he traveled to the NIH in Bethesda, Maryland, to meet with and recruit his soon-to-be colleague and now Professor of the Graduate School Bruce N. Ames (who remained active in research at the Children's Hospital of Oakland Research Institute until 2017). Ames, after coming to Berkeley and joining the Dept. of Biochemistry the very next year (1968), and much later when he himself was Chair (1983-89), often went to Clint for advice on matters ranging from scientific problems to how to deal with certain difficult personalities. Ames sought Clint's opinion and valued his judgment because Clint always responded in a deeply thoughtful way and was very straightforward, even when, on occasion, he admitted to being at a loss for what advice to give. Ames described Clint as “all stainless steel," not because he was hard on people, but because the strength of his character was so upright, trustworthy and reliable. Similarly, Giovanna Ferro- Luzzi Ames, now a Professor Emerita, upon first meeting Clint, and from her perspective as a fairly fresh immigrant from Italy (then, as now, a rather chaotic country), felt that he epitomized the “Ideal American"— a man from humble origins with high principles who embodied solidity, fairness and generosity. Subsequently, but in short order, she came to know that all of these characteristics were authentic, that he was a genuinely special person and, indeed, truly a "real" American. She recalled one incident that illustrated to her Clint's concern for always taking the high road. At one point, one of Clint's graduate students had been working on a project in conjunction with her lab; when it was completed, written up, and ready to be sent out for peer review, Ferro-Luzzi Ames told the student that her name should be left off the list of authors (not because she thought the results were faulty, but because she felt she had not contributed sufficiently to the work). Even though Clint was on sabbatical abroad, he telephoned back to the USA (in that era, a rather pricey call) to get her personal reassurance that his student was not trying to do something unethical, impressing Ferro-Luzzi Ames with how very seriously Clint took his responsibility to ensure his students behaved properly at all times and in every respect.

During the four decades that followed Clint's initial faculty appointment, and with students, postdocs, collaborators and visiting scientists too numerous to acknowledge individually here, Clint made ground-breaking contributions in several major areas of carbohydrate and polysaccharide, glycolipid, and glycoprotein biochemistry.

In addition to continued work on the phosphoinositides and inositol-polyphosphates already mentioned, Clint maintained an interest in sugar-phosphate synthesis and metabolism [Gillett JW, Ballou CE (1963) The synthesis of L-glycero-tetrulose 1-phosphate (L-erythrulose 1- phosphate) Biochemistry 2: 547-552; Tegge W, Ballou CE (1992) Syntheses of D-myo-inositol 1,4,5-trisphosphate affinity ligands. Carbohydr. Res. 230: 63-77]. Because of its relevance to tuberculosis, an affliction that his Ph.D. mentor Link suffered, Clint also invested considerable effort in studying the nature of the protective coatings that surround and protect mycobacteria and made many novel discoveries about mycobacterial polymethylpolysaccharides and mycobacterial glycolipids [Maloney DH, Ballou CE (1980) Polymethylpolysaccharide synthesis in an ethionine-resistant mutant of Mycobacterium smegmatis. J. Bacteriol. 141: 1217-1221; Kresge N, Simoni RD, Hill RL (2009) Mycobacterial glycophosphoinositides: the work of Clinton E. Ballou. J. Biol. Chem. 284: e13–e15].

In the late 1960s, Clint initiated what became perhaps his most well-recognized work— an analysis of the cell walls of the budding yeast Saccharomyces cerevisiae, especially the outermost mannoprotein layer and the nature of the glycosyl chains attached to those glycoproteins [Stewart TS, Mendershausen PB, Ballou CE (1968) Preparation of a manno- pentaose, mannohexaose, and mannoheptaose from Saccharomyces cerevisiae mannan. Biochemistry 7: 1843-1854; Stewart TS, Ballou CE (1968) A comparison of yeast mannans and phosphomannans by acetolysis. Biochemistry 7: 1855-1863; Jones GH, Ballou CE (1968) Isolation of an alpha-mannosidase which hydrolyzes yeast mannan. Structure of the backbone of yeast mannan. J. Biol. Chem. 243: 2442-2446]. Appreciating the advantages of applying genetic analysis to these studies because of the tractability of this yeast for that experimental approach, Clint applied for and was one of the ten students accepted into the inaugural Molecular Biology and Genetics of Yeast course at the Cold Spring Harbor Laboratory (June 14- July 4, 1970), which has been offered continuously every summer since. As related by one of the founding instructors, Prof. Gerald R. Fink (then at Cornell, now at the Whitehead Institute and MIT), Clint initially was having a very hard time understanding certain genetic concepts, especially tetrad analysis. Gerry was a bit perplexed and a bit embarrassed for his pupil because he knew what a renowned biochemist Clint was. But, very late one night, Gerry heard a knock at his door and opened it to find Clint, with an excited wide-eyed look, which actually scared Gerry a bit. However, Clint then blurted out, "I get it!! It's all binary!" and, then, turned on his heels and left. Thereafter, Clint was one of the stars in the course because he took something that he found confusing at first, but made it crystal clear to himself by boiling it down to its essence, which left his teacher Gerry with a big smile. Indeed, over the course of the next more than 20 years, Clint, often with Lun as his collaborator (in 16 publications), used a productive blend of genetic and biochemical analysis to elucidate the structures and functions of the glycosyl chains in yeast mannans [Ballou CE (1974) Some aspects of the structure, immunochemistry, and genetic control of yeast mannans. Adv. Enzymol. Relat. Areas Mol. Biol. 40: 239-270; Häusler A, Ballou L, Ballou CE, Robbins PW (1992) Yeast glycoprotein biosynthesis: MNT1 encodes an alpha-1,2-mannosyltransferase involved in O-glycosylation. Proc. Natl. Acad. Sci. USA 89: 6846-6850].

Like Wold and Pizer before them, a large number of Clint's UC Berkeley Ph.D. students and his postdoctoral trainees have since gone on to prominent independent careers in academic science. Clint himself was recognized for his research accomplishments by several prestigious accolades, including a National Science Foundation 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 others, receipt of the prestigious Claude Hudson Award in Carbohydrate Chemistry from the American Chemical Society (1981) and the Berkeley Citation from the University of California for "distinguished achievement and for notable service to the University" (1992). In 1994, an entire volume (251) of the journal Carbohydrate Research was dedicated solely to him and comprised papers all written in Clint's honor.

For persons who encountered Clint only on occasion, there was something about his countenance or demeanor that gave them the very misleading impression that he was remote, reserved and unapproachable. He was sometimes referred to as the 'Great Stone Face.' In this regard, his son Philip related that it was amusing to him how many of his acquaintances mentioned that his father seemed 'standoffish,' at least initially, and Phil recalled that when he was a student (a Music major) at UC Berkeley and Clint was still teaching, Phil was browsing the used books in Moe's Books (a Berkeley institution since 1959) and ran across a publication that comprised student reviews of Cal classes and Professors. The entry for Clint was generally favorable; but, as Phil recollected, the one thing he distinctly remembered was reading that the students' nickname for Clint was "The Iceman." However, nothing could be further from the truth.

To a person, those of his junior colleagues knew how kind, generous and helpful Clint could be. Jasper Rine recollects that, after the relative formality of his interview with Clint at the time Rine was being considered for a faculty position and then was hired, Clint was solicitous about not intruding in Jasper's business, yet was incredibly helpful any time Rine asked for his guidance. When he asked Clint for comments on the manuscript that he wanted to submit describing his first independent research findings, Clint gave him more extensive feedback and much better advice than he had ever received before. Randy Schekman recalls that his relationship with Clint was a little strained at first, but then Clint helped him too with his very first paper, especially through Clint's willingness to communicate it to the Proceedings of the National Academy of Sciences. Thereafter, Clint was quite supportive of his work and, on occasion, revealed his sense of humor. For example, at an early research retreat of the Biochemistry Dept. at the Asilomar Conference Center in Pacific Grove, CA, Professor Frederick H. Carpenter (1918- 1982) presented his group's work on a protein extracted from bovine eyes, and described the starting material— a bucket full of cow eyeballs collected from a slaughter house —whereupon Clint jumped up from his seat in the front row, hunched over in a mock gesture of gagging, and ran from the room, followed by a general chorus of laughter! Similarly, although initially an outsider in another department (Microbiology and Immunology) who eventually joined the Biochemistry Dept. officially in 1985, Jeremy Thorner recounted that, despite the rumors he had heard about Clint being reserved and unapproachable, he found Clint to be a very generous, thoughtful, and helpful mentor and Lun to always be warm, welcoming and cheerful.

Other members of the Biochemistry Dept. also remember other aspects of Clint's warmth and generosity. Carol Mimura, then a postdoc in the laboratory of Giovanna Ferro-Luzzi Ames, which was located at the time on the 3rd floor of Barker Hall immediately adjacent to the Ballou Lab, recalled with fondness, not only helpful consultations with Clint about her research work, but the fact that on occasional Fridays at 4 PM Clint and Lun hosted a floor-wide get-together that always featured home-made ice cream prepared in an old-fashioned hand-cranked model that required rock salt and a lot of elbow grease to chill down the ingredients. Indeed, such was just one among many community-building efforts that Clint and Lun fostered. Gary R. Gray, a postdoc from the University of Iowa who joined Clint's lab in 1969 and who was responsible for the 1994 issue of Carbohydrate Research dedicated solely to Clint, now a Professor Emeritus of Chemistry at the University of Minnesota, Twin Cities, recalled, as many of his other friends and colleagues can attest, how much Clint and Lun shared their enjoyment of good food and fine wines, and experimenting with gourmet cooking, with many memorable evenings spent sharing marvelous dinners prepared by Lun and Clint in their home on Creston Road. In fact, Clint and Lun were co-founders of a “good-eating club”, whose other members included Bruce and Giovanna Ames, and three other Biochemistry Dept. faculty, namely Jesse C. Rabinowitz (1925-2003), Stuart M. Linn, and Edward E. Penhoet; this group organized dinners in special restaurants throughout the year. Another cherished tradition of the club were fabulous New Year's Eve dinners, which took place in the members' homes, on a rotating basis; when it was their turn, Clint’s specialty was frequently (by general request) a delicious all-vegetable soup that took him an entire day to prepare and Lun's was a succulent duck imbued with the flavor of smoked tea leaves.

Everyone also saw the pride and joy that Clint took in driving and maintaining his vintage, champagne-yellow Porsche convertible, which remains a prized possession in the Ballou family to this day.  During summer vacations, Clint loved to spend time fly fishing and was enthralled with the challenges of hand-tying his own flies. After he retired, Clint read a great deal, and prided himself in successfully learning to recite “You Are Old, Father William” from Alice In Wonderland (chosen in honor of his own father William), as well as the opening monologue (in French!) of the character Alceste in Molière’s Le Misanthrope.”

Another important part of Lun's and Clint's shared interest was international travel. Clint always took full advantage of his sabbatical leaves. In 1961, he took the family to Paris, France. They lived in apartment on the Rue Pierre Curie (later renamed Rue Pierre et Marie Curie) near Jardin de Luxembourg, while Clint did research at the Centre National de Recherche Scientifique (CNRS) laboratories in Gif-sur-Yvette, a nearby Paris suburb. For his second sabbatical in 1968, Clint bought his entire family round-the-world airplane tickets, and they hopped their way around the globe with extended stays in Kyoto, where Clint worked with Osamu Hayaishi (1920–2015), and again in Paris, where Clint worked at the Institut Pasteur with Anne-Marie Staub (1914-2012). In 1972, shortly after Nixon thawed relations between the USA and the People's Republic of China, Clint and Lun took their children to meet Lun's side of the family— the first time Lun herself had returned to China since coming to the USA 24 years before. After waiting for approval of their visas in Hong Kong (then a British crown colony, which did not revert back to the Chinese government until 1997), they eventually were allowed to take a train to the mainland border; then, after walking across a bridge, they boarded a train on the Chinese side of the border and proceeded on their way, leading to a very emotional and memorable reunion with Lun's relatives. Clint spent his sabbatical leave in 1975 in the laboratory of Georg Friedrich "Fritz" Melchers, then in the Institute for Immunology in Basel, Switzerland (now, since 2003, a Senior Research Group Leader at the Max Planck Institute for Infectious Disease in Berlin, Germany). For his sabbatical in 1982-83, Clint spent six months in the CNRS facility in Grenoble, France, in the laboratory of Jacques Defaye and then six months at Imperial College, London, UK, with Howard R. Morris and Anne Dell. Perhaps Lun's and Clint's most pleasant trip to France was in 1990, when they (and their daughter Linda) attended the wedding of their son Philip to his wife Nanou in Pertuis, an ancient village about a dozen miles from Aix-en-Provence in the southeastern most part of France.

Those of us who were there recall that, at a dinner held by the Biochemistry Dept. at a fancy restaurant on the Berkeley waterfront in honor of Clint and Lun on the occasion of his retirement, he had prepared and read aloud a very 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.

In this time of loss, we as a community are privileged to remember how fortunate we are to have benefited from Clint’s research acumen, his community spirit, his skills as a teacher and mentor who set high standards, his generous spirit, and good humor. As his daughter Linda expressed it so succinctly "Clint was a fine man, and a scientist who made great contributions in the biochemistry field. He was also a remarkable husband, and a truly loving and supportive father and grandfather."

We offer our sincerest condolences to the Ballou family, and to all his scientific progeny, colleagues and friends.

Respectfully submitted (23 March 2021),

Jeremy Thorner, Professor Emeritus, BBS Div., MCB Dept., UC Berkeley
Randy Schekman, Professor, CDB 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