Richard Lane Calendar

Richard Calendar, who was Professor of Molecular Biology (1968-1989) and, after a reorganization of the biological sciences, Professor of Biochemistry, Biophysics and Structural Biology (1989-2009), and then Professor of the Graduate School (2009-2020), passed away on October 10, 2020.

Rich was born on August 2, 1940, in Hackensack, New Jersey, the eldest of three brothers. He received a B.S. in chemistry from Duke University in 1962 and a Ph.D. in biochemistry from Stanford University in 1967, where his Ph.D. mentor was Professor Paul Berg, who was awarded the 1980 Nobel Prize in Chemistry. With Berg, Rich purified and characterized tyrosyl-tRNA synthetases. For Rich's postdoctoral training, having been inspired by the work on bacterial viruses (bacteriophages) by Professors A. Dale Kaiser and David S. Hogness in the same department at Stanford, he joined the lab of Giuseppe Bertani in the Department of Microbial Genetics at the Karolinska Institute in Stockholm, Sweden. There, Rich began his lifelong devotion to bacteriophage biology. Importantly, he took a course to learn Swedish, and ended up marrying his instructor, Gunilla. During their marriage (1969-1982), they had a son, Hugo, and a daughter, Johanna. There are now three grandchildren in the U.S. and Sweden: Sean, Sonya and Samuel.

Rich became an international leader in the study of bacteriophages in Escherichia coli, Listeria monocytogenes, and Bacillus anthracis. He wrote comprehensive reviews that spurred others to enter the field (e.g, his 1970 article in Annual Review of Microbiology). He literally “wrote the book” on this subject: The Bacteriophages, Oxford Univ. Press, 2^nd ed., 2006. Rich was a dedicated researcher who relished working with his hands throughout his career and continued to work at the bench until a month before his death. His final paper, describing work carried out in collaboration with colleagues at the Lawrence Berkeley National Laboratory and colleagues on the UC Berkeley campus, was posted on PubMed three days after his death.

For many years, Rich’s primary focus was on understanding the complex interactions between the bacteriophage P2, which infects Escherichia coli and can either insert itself into the bacterial genome or lyse the host cell, and the “satellite” bacteriophage P4, which can integrate into the host genome by itself but cannot engage in lytic growth without the presence of P2 as a “helper” phage. During his extensive studies of bacteriophages P2 and P4, he discovered that the genomes of these DNA viruses are concatenated, being interlinked like paper circles in a decorative chain. His Berkeley colleague Professor Nicholas R. Cozzarelli and others used these phage DNA molecules to understand DNA three-dimensional architecture (DNA topology) and to isolate and characterize enzymes that catalyze DNA breakage and rejoining reactions to unlink catenated molecules (DNA topoisomerases). Rich’s former graduate student, Brian Sauer, in his own lab, showed that the Cre recombinase encoded by another E. coli bacteriophage (P1), could be expressed in mammalian cells to direct site-specific recombination of introduced DNA into the host cell genome, which heralded the use of the Cre-loxP system, now a standard tool for mouse geneticists world-wide. These examples and many others illustrate how the discoveries Rich made, and the Ph.D. students he trained to appreciate the wonders of bacterial viruses, have had a profound impact on the progress of molecular biology. Rich’s most highly cited paper, “Construction, characterization, and use of two Listeria monocytogenes site-specific integration vectors,” J. Bacteriol. 184 (2002), a collaboration with other Berkeley colleagues, describes the development of phage integration vectors in the human pathogen Listeria monocytogenes for a potentially important biomedical application. Specifically, these vectors allowed for the rapid construction and stable expression of human tumor antigens in these bacterial cells for the development of Listeria-based vaccines now being evaluated for anti-cancer immunotherapy.

Rich was unrelentingly dedicated to undergraduate education, and he continued to teach for many years after his formal retirement. He created the first course in molecular biology at UC Berkeley and helped to establish a molecular biology major. Thereafter, he updated the extremely influential textbook Molecular Genetics: An Introductory Narrative (WH Freeman & Co., 2^nd ed., 1978), originally written by another late colleague, Professor Gunther S. Stent. Most recently, as an instructor in the largest class of the division’s curriculum, MCB 102 (Survey of the Principles of Biochemistry and Molecular Biology), he taught any one of its three distinct segments as needed. Rich was an innovator long before that became a buzzword; to engage the students, he would devise and play them songs to help them remember the essential facts. In his research laboratory, Rich trained over a hundred undergraduates, many of whom have gone on to prominent academic careers, including geneticists Susan L. Forsburg and Steven E. Finkel (both now at the University of Southern California) and Nobel Prize Laureate Andrew Z. Fire (now at Stanford University). Rich went out of his way to familiarize students, both graduate and undergraduate, with the fine arts, encouraging and even hosting them to attend the Berkeley Repertory Theater and Cal Performances, in addition to engaging them in discussions of research papers and science in general.

Rich's diverse interests were reflected in his extensive service to the Academic Senate. His twenty years of service started in 1972 with four years on the Committee on University Welfare and ended in 2017 after two years on the Committee on Academic Planning and Resource Allocation. In between, he served multiple terms on the Graduate Council and the committees on Research, Courses of Instruction, and Admissions, Enrollment, and Preparatory Education.

Rich will also be remembered for his wonderful sense of humor: he had a joke, pun, or hilarious cartoon for all occasions, in person and by email. Multiple collaborators and colleagues have expressed their gratitude for Rich’s intelligence and expertise as well as his genial and magnanimous nature. Rich regularly provided fellow scientists with phages and critical insights into phage biology. Since his recent death, numerous colleagues have contacted us with their condolences but also to request a visit to obtain phages from his personal collection. Rich was known by many for his annual winter Santa Lucia celebration where he played his guitar, sang Swedish songs, and provided ample refreshments for all, including his famous (and potent) version of the traditional quaff, known as glögg. Rich was a man who always enjoyed a good party! And he cared to share this joie de vivre with others, in an uplifting spirit that will not be forgotten.

Rich was very active outside of the lab. He started doing whitewater kayaking in the mid-1970s, making his own fiberglass boats. He also coached and refereed youth soccer in the '70s and '80s in support of his children. He performed Scottish country dancing for pleasure at local events such as the Solano Avenue Stroll, and he volunteered to help run the local Royal Scottish Country Dance Society. Through that society he met his partner Nancy Page, who was his constant companion for the last 25 years of his life.

We as a community remember how fortunate we are to have benefited from Rich’s research acumen, commitment to education, generous spirit, and good humor. We join Rich’s family, scientific brood, and close friends in commemorating him.

Daniel Portnoy
Jeremy Thorner
Kathleen Collins
2021