In Memoriam: Alexander N Glazer

Alexander N. Glazer (1935 – 2021)
Professor of the Graduate School, UC Berkeley
Past Director, University of California Natural Reserve System

Alexander N. Glazer first joined the University of California as a faculty member in the Dept. of Biological Chemistry (1964-1976) of the School of Medicine at UCLA and Associate Member of the UCLA Institute of Molecular Biology (1965-1976). He then moved to the Berkeley campus, where he was Professor (1976-1989) and Chair (1977-1982) of the Dept. of Microbiology and Immunology and, following reorganization of the biological sciences and establishment in 1989 of the Dept. of Molecular and Cell Biology, Professor (1989-1994) in its Division of Biochemistry and Molecular Biology (now Biochemistry, Biophysics and Structural Biology) and then Co-Chair (1994-1997), despite his retirement (as a Professor of the Graduate School) in 1994. To continue to contribute further to the University, and spurred by his own deep life-long appreciation for nature, he applied to be and served as Director of the UC Natural Reserve System (1998-2009). Alex passed away on July 18, 2021, in Orinda, CA, at age 86, having led a remarkable and impactful life.

Alex, an only child, was born on July 7, 1935, in Łódź, Poland. The family moved to Warsaw, where his father was head accountant for a bank and his mother a homemaker and esthetician. Nazi Germany invaded Poland on Sept. 1, 1939, and occupied the country. Alex’s father was killed during the war; but, hiding in the Polish countryside, Alex was liberated with his mother in 1945. Throughout his life, Alex always avoided talking about that horrific period. After the war’s end, Alex and his mother made their way to France, where they awaited transit visas for their eventual emigration to Australia, where Alex's aunt (his mother's sister) lived in Sydney. During that waiting period, Alex traveled to Israel where he stayed with other relatives who had emigrated there before the war, and then rejoined his mother in France. In 1946-47, Alex (at age 11) and his mother finally made their way to Australia via ship. It was on that voyage that Alex commenced learning English. He thrived in Australia. While completing high school with a superb academic record, he played soccer, was something of a chess prodigy, and, perhaps most notably, was awarded a prize for his scholastic achievement in English.

On the basis of his outstanding accomplishment, he was admitted to the Univ. of Sydney, where he obtained his B.Sc. with First Class Honors (1957) and M.Sc. (1958) degrees in Biochemistry. It was in his Master's thesis research under Hugh A. McKenzie (1923-2008), in which he used spectroscopy and other methods to monitor the urea-induced denaturation of serum albumin and ovalbumin [Nature 180: 1286-1287 (1957)] and other proteins, that Alex first immersed himself in the physico-chemical study of proteins, a subject that became the central focus of much of his subsequent life's work.

At the Univ. of Sydney, Alex met and married (1958) Eva (née Warner), whose family had emigrated to Australia from Germany in 1938, and who survives him. They have two children, Judy (born 1962) and John (born 1965), and four grandchildren—  Rebecca (21) and Danny (18) in Los Altos, CA, by Judy (and husband Brian Donahue); and, Jai (16) and Shaan (11) in New York, NY, by John ( and wife Nandita). 

While still a university student, Alex had attended a lecture by visiting Prof. Emil L. Smith (then at the Univ. of Utah) on structure-function relationships in a proteolytic enzyme, papain. Given his emerging fascination with proteins, Alex found the talk inspiring and decided, then and there, to go to Utah and do his graduate studies with Smith, which he did. He worked on various problems in protein chemistry, including determination of the amino sequence of peptides [ J. Biol. Chem. 235: 3159-3162 (1960)] and the ultraviolet spectra of proteins and polypeptides [J. Biol. Chem. 236: 2942-2947 (1961)]; however, he was awarded his Ph.D. (1960) with a dissertation entitled “The Sulfur Distribution of Papain and Related Studies." Alex's analysis, published subsequently [J. Biol. Chem. 240: 201-208 (1965)], indicated that papain had one free cysteine side chain in its active site and three disulfide bonds, a conclusion fully verified a decade later when Smith and his then-colleagues published the complete amino acid sequence of this protease [J. Biol. Chem. 245: 3485-3492 (1970)]. This remarkable accuracy is just one manifestation of the meticulous care that was always a hallmark of Alex's approach to research. Alex's experiences in Utah had an additional and equally important consequence for the remainder of his career—  the National Parks and other areas of astounding natural beauty in Utah, and elsewhere in the southwest, enthralled Alex, inculcating a passion for preservation of our environment.

To foster his further post-Ph.D. training, Alex was awarded a prestigious fellowship from the Jane Coffin Childs Memorial Fund for Medical Research. With this support, Alex and Eva went, first, to Israel where Alex conducted postdoctoral research (1961-62) with Ephriam Katchalski (1916-2009) at the Weizmann Institute of Science in Rehovoth, where he used N-carboxy-L-tyrosine anhydride to decorate the free amino groups in the protease trypsin [ J. Biol. Chem. 237: 1832-1838 (1962)]. Alex and Eva then moved to Cambridge, UK, where their daughter Judy was born and Alex spent another year (1962-63) with Fred Sanger (1918-2013) at the Medical Research Council (MRC) Laboratory of Molecular Biology, studying the iodination of residues in both serum albumin and chymotrypsin [Biochem. J. 90: 92-98 (1964)]. Chemical modification of proteins would continue to be a major theme in much of Alex's early independent research. Indeed, a decade later, with two biochemist colleagues from UCLA, David S. Sigman (1939-2001) and Robert J. DeLange (1937—), Alex literally wrote the book on this subject—  Chemical Modification of Proteins: Selected Methods and Analytical Procedures, North-Holland Publishing Co., Amsterdam / American Elsevier Publishing Co., New York, 1975, 205pp.

After completion of his postdoctoral training, Alex was offered an Asst. Prof appointment in 1964 by his former thesis preceptor Emil Smith (1911-2009), who had moved to UCLA the previous year to become Prof. and Chair of the Dept. of Biological Chemistry in the medical school there. After their move to Los Angeles, Eva gave birth to their son John. Alex rose rapidly through the ranks, receiving tenure just two years later (1966) and promotion to full Prof. just four years after that (1970). At UCLA, Glazer continued to work, for a brief time, with Smith on various aspects of protein sequence determination and analysis of certain naturally-occurring modified amino acids in some proteins, such as the Nε-trimethyl-lysine found in mitochondrial cytochrome c in all eukaryotes [ J. Biol. Chem. 244: 1385-1388 (1969)]. On his own, Alex continued studies of the properties of other major proteases (papain, subtilisin, trypsin, and chymotrypsin), and other hydrolases (ribonuclease, lysozyme) that were paradigms for enzymological research in that era of biochemistry. However, two experiences dramatically altered the direction of his research and life. First, he commenced a decade-long collaboration with other UCLA colleagues, in particular Bob DeLange and microbiologist Rafael J. "Ralph" Martinez (1928-2017), to determine the sequence and structure of bacterial flagellins, especially from Bacillus subtilis [J. Mol. Biol. 28: 45-51 (1967); J. Biol. Chem. 251: 701-704 (1976)]. Second, after his promotion to tenure in 1970, Alex took a sabbatical at UC Berkeley with the husband-and-wife scientific team of Roger Stanier (1916-1982) and Germaine Cohen-Bazire (1920-2001) in the then-Dept. of Bacteriology and Immunology, where Alex was introduced to the photosynthetic cyanobacteria (formerly blue-green algae) and became captivated by the challenge of deconvoluting the components, structure, and function of the large light-harvesting complexes, dubbed phycobilisomes, in these organisms [Proc. Natl. Acad. Sci. USA 68: 1398-1401 (1971)]. Unsettled by the campus unrest caused by the student anti-Vietnam War protests of that era, Stanier and Cohen-Bazire left UC Berkeley the very next year (1971) to direct the Microbial Physiology Unit at the Institute Pasteur in Paris, France; but, after Alex's return to UCLA, he still was able to continue a long-distance collaboration with Cohen-Bazire for several more years [Arch. Microbiol. 104: 29-32 (1975)]. At UCLA, additional collaborative studies on phycobilisomes were conducted with one of us (Eiserling) [J. Ultrastruct. Res. 47: 16-25 (1974); Arch. Microbiol. 110: 61-75 (1976)].

After these exposures to microbiology, when the opportunity arose to return to UC Berkeley and join the faculty in the by-then-renamed Dept. of Microbiology and Immunology originally founded by Roger Stanier, Alex took advantage and commenced his new appointment in 1976. In Berkeley, further dissecting the subunits (phycobiliproteins) and their chromophores in phycobilisomes, deciphering the molecular architecture and mechanism of assembly of phycobilisomes, and delineating the energy transfer pathways within phycobilisomes became the intense focus of Alex's research and it flourished for the next 30+ years [ Annu. Rev. Microbiol. 36: 173-198 (1982); Annu. Rev. Biochem. 52: 125-157 (1983); Annu. Rev. Biophys. Biophys. Chem. 14: 47-77 (1985); J. Biol. Chem. 264: 1-4 (1989); J. Biol. Chem. 284: e12-e14 (2009)]. These studies were often conducted in conjunction with collaborators there, like electron microscopist Robley C. Williams (1908-1995) and chemist Henry Rapoport (1918-2002), and elsewhere, like Cohen-Bazire, DeLange, J. Clark Lagarias (since 1980 a Prof. at UC Davis), and his former UCLA graduate student Donald A. Bryant (since 1981 a Prof. at Pennsylania State Univ.). The capacity to arrange for sustained and fruitful collaborations on such a frequent basis was another hallmark of Alex's abiity to spread his infectious joy in doing research.

Alex had an especially keen, creative and fertile mind. He took an expansive view of all of modern science. For example, with Stanford Prof. Lubert Stryer (1938—), a long-time friend since their postdoc days at the MRC, they envisioned practical applications of proteins as small, stable, and highly fluorescent as phycobiliproteins, and developed use of phycobiliproteins as covalent antibodies tags [ J. Cell Biol. 93: 981-986 (1982)], an application commercialized by Becton-Dickinson, Inc. As another example, with chemist and Berkeley colleague Richard A. Mathies (1946—), they devised a suite of fluorescent energy-transfer reagents for detection and sequence analysis of DNA [Curr. Opin. Biotechnol. 8: 94-102 (1997)], which also have been successfully commercialized. As a third example, and realizing the potential of "Big Data," Alex in collaboration with statistician and Berkeley colleague Peter J. Bickel, exploited the burgeoning protein sequence databases publicly available to explore the patterns of amino acid sequence conservation and change over both phylogeny and evolutionary time to gain new insights about factors responsible for the coupling of structure to function in protein families [Proc. Natl. Acad. Sci. USA 99: 14764-14771 (2002]. As a final example, with long-time colleague and bacteriologist Hiroshi Nikaido (1932—), Alex wrote a very influential textbook, Microbial Biotechnology: Fundamentals of Applied Microbiology, which went through two editions [W.H. Freeman, 1994, 640pp.; and, Cambridge Univ. Press, 2007, 576pp.].

Another facet of Alex's many talents was a gift for administration, and he never shied away from opportunities to serve in leadership roles. Aside from serving as Chair of the Dept. of Microbiology and Immunology, during the subsequent reorganization of the biological sciences on the Berkeley campus that was initiated in 1980-81 and led by biochemist Daniel E. Koshland Jr. (1920-2007), Alex was a pivotal member of its primary steering committee, which oversaw the successful establishment in 1989 of the new Dept. of Molecular and Cell Biology (MCB) and its two sisters, the Dept. of Integrative Biology and the Dept. of Plant Biology. As part of that process, there was also bureaucratic restructuring, and Alex assumed the position of Head  (1990-1994) of the Administrative Services Unit that serviced two of the buildings (Stanley Hall and Donner Laboratory) that housed MCB faculty. Thereafter, he became one of the two Co-Chairs of the entire MCB Dept. He was also able to bring in new resources to enhance the campus intellectual environment. For example, after Stanier's death in 1982, Alex persuaded Prentice-Hall, Inc., the original publisher of Stanier's revelatory microbiology textbook, The Microbial World, to provide an endowment for a Roger Y. Stanier Memorial Lecture to be given, initially, in alternate years at UC Berkeley and at the Institute Pasteur, which commenced in 1985 with a presentation by then-MIT Professor Salvador Luria (1912-1991) and continues at UC Berkeley, now on an annual basis, to this very day under the stewardship of microbiologist and Prof. Daniel A. Portnoy (1956—).

In the early 1990s, the State of California and the University suffered one of its periods of fiscal stringency. To reduce the campus operating budget, faculty were offered the Voluntary Early Retirement Incentive Program (VERIP or "Plus 5"), which provided the equivalent of an extra five years of service and other inducements to maximize retirement income and convince them to retire early, thereby moving them off the operating payroll and onto their UC pensions. For this reason, at age 59, Alex formally retired in 1994 and became a Prof. of the Graduate School. This choice opened up new freedom for Alex to continue to serve the University. Aside from his following three-year stint as MCB Dept. Co-Chair, Alex pursued the equivalent of a second career when he served for the subsequent eleven years as Director of the UC Systemwide Natural Reserve System, where he was able to apply his deep knowledge of and appreciation for the natural world and oversee the unspoiled areas of California (39 preserves distributed among 12 different regions) that wisely had been set aside for ecological, ethological, geological, and other study. Moreover, he turned the focus of his perceptive intellect to pressing issues in environmental science, such as the effects of anthropogenic fixed nitrogen, the lack of and contamination of freshwater sources, and the repercussions of natural gas and oil production and their use. These prescient concerns are now at the forefront of the global warming crisis that the nations of the world are now finally confronting. A more detailed and moving account of Alex's tenure as Director of the UC Natural Reserve System, and the profound impact that he had in that role, is available elsewhere (

Throughout his career, Alex received many accolades for his achievements and contributions, including:  the Endeavour Prize of the British Association for the Advancement of Science (1955); two Guggenheim Foundation Fellowships (1970-71 and 1982-83); the Darbaker Prize of the Botanical Society of America (1980); the Award for Excellence in Scientific Reviewing from the National Academy of Sciences (1991); election to the American Academy of Arts and Sciences (1996); and, election to the National Academy of Sciences (2001).

Equivalent to his impact as a research scientist and administrator, Alex was an accomplished and inspirational educator, truly respected and even beloved by his trainees. At the undergraduate level, Alex was a vocal advocate for equity in STEM education. After he stepped down from the directorship of the UC Natural Reserve System, he became an ardent supporter of and dedicated mentor in the UC Berkeley Biology Scholars Program (BSP), which challenges unwarranted assumptions about who is capable and qualified to do science. Established in 1992, BSP has supported approaching 4,000 students— 80% of whom are from low-income backgrounds and/or are the first in their family to attend college, 70% of whom are women, and 60% of whom are from ethnic groups traditionally underrepresented in the sciences.

Typical of the sentiments received from many of his former graduate students, Hays S. Rye (Texas A&M Univ.) commented about Alex:  "He’ll live on in the back of my mind, but I’m really going to miss knowing that he was in the world. I can’t think about any scientific problem without hearing his voice, thinking about how he might approach the issue and seeing that grin of his. I can only hope I have a fraction of the influence on my students that Alex had on me. Alex was quite unique. I don’t think I’ve ever been around anyone who could stand in the gale and chaos of the scientific process with such happy joy about just getting to solve great problems. It was infectious and brilliant and special. I know I don’t have his gifts, but I’ve tried to extract the best of what he taught me about doing science and made my best stab at trying to pass some of it on." Similarly, Don Bryant reflected:  "Among Alex’s many admirable qualities, a couple of things impressed me most. First, he was always kind, taking pains to clearly and calmly explain things, in a manner that led one to feel that you were speaking as colleagues and co-equals rather than individuals of different status, and where ideas and decisions were open to debate. Second, I believe that Alex could convince anyone, and maybe even inanimate objects, to be as excited about his science as he was, and, considering how enthusiastic he was all the time, that was a remarkable gift. These two things about Alex, together with his profound knowledge of protein chemistry, are what I learned and treasure the most from my time with him."

As his daughter Judy expressed so succinctly on behalf of her family, "Alex was a wonderful husband, father and grandfather, and a passionate lover and advocate for the outdoors. We miss that vibrant person and remember him with joy."

We offer our sincerest condolences to the Glazer family, and to all of Alex's scientific progeny, colleagues and friends.

Respectfully submitted,

Jeremy Thorner, Professor Emeritus
Division of Biochemistry, Biophysics and Structural Biology
Department of Molecular and Cell Biology
University of California, Berkeley

Frederick A. Eiserling, Professor Emeritus
Department of Microbiology, Immunology, and Molecular Genetics
David Geffen School of Medicine
University of California, Los Angeles