Obituary

Andrew A. Benson 1917 – 2015

ndrew A. Benson (called When Andy left Berkeley in Andy by his friends) 1955, he was appointed to the Adied on January 16, 2015, faculty at Pennsylvania State in La Jolla, California; he was University and turned his atten- born in Modesto, California, on tion to plant lipids. In 1957, he September 24, 1917. He gradu- discovered the major membrane ated from Modesto High School phospholipid phosphatidyl glyc- in 1935 and studied chemistry erol, and in 1961, the sulfolipid at the University of California, sulfoquinovosyl diglyceride. Both Berkeley, obtaining a B.S. in are essential to the formation and 1939. He performed graduate function of photosynthetic chloro- work at the California Institute of plast membranes. In recognition Technology under Carl Niemann, of his outstanding contributions to receiving his doctorate in 1942, the lipid field, the book Lipids in and then returned to Berkeley as Dee and Andy Benson at the dinner held at the La Jolla Country Club on Photosynthesis, edited by Hajime SOURCE: BUCHANAN an instructor in the Department of September 24, 2014, commemorating his 97th birthday. Wada and Norio Murata, was AND DOUCE, 2015. Chemistry. There, he was exposed dedicated to him in 2009 (Volume to photosynthesis and radioisotope 30 in the series “Advances in research by Samuel G. Ruben and times, collected in hot methanol, with other members of the Calvin Photosynthesis and Respiration,” Martin Kamen, two pioneers who and analyzed by two-dimensional group. The compounds identified Springer, Dordrecht). had discovered 14Carbon in 1940. paper chromatography. The in lollipop experiments served In 1962, Andy moved to His research came to a halt in 1942 labeled compounds were local- as the basis for formulating the the Marine Biology Research with the onset of World War II. ized on the chromatograms and carbon dioxide reduction cycle Division, Scripps Institution During that period Andy served identified by autoradiography. of photosynthesis. Although of Oceanography, in La Jolla, in various functions in the civil These techniques, introduced by Calvin had proposed the origi- where he spent the remainder of service. Andy to the Calvin group, subse- nal concept of a cycle, Andy his career, becoming emeritus In 1946, Andy was appointed quently became the gold standard was pivotal in bringing the idea in 1989. At Scripps he initially a research associate in the for 14C-labeling experiments by to fruition. He developed tech- continued his research on plant Radiation Laboratory at Berkeley laboratories worldwide. niques for degrading 14C-labeled lipids, but with time turned his in the photosynthesis group that In 1950, Andy chemically sugar phosphate intermediates to attention to marine biology. With was being assembled by Melvin identified ribulose 1,5-bisphos- locate the 14C-label in individual his coworkers he recognized Calvin. By feeding 14C-labeled phate as an essential component C-atoms, innovations central to the importance of calcitonin in CO2 to suspensions of the green of the photosynthetic carbon deciphering the cycle. By late calcium regulation in salmon, alga Scenedesmus, he identified reduction cycle. This 5-carbon 1953, the photosynthetic carbon studied the role of arsenic in 3-phophoglyceric acid (PGA) as sugar phosphate proved to be reduction cycle was more or less oceans, and identified previously the first stable product of photo- the long-sought missing link in worked out, and evidence for the unknown intermediates of arsenic synthesis in joint work carried the cycle that enables photosyn- complete cycle was published the metabolism in aquatic plants as out with Calvin. For these stud- thetic organisms to fix CO2 and following year. Working in 1954 well as the highest concentration ies, he designed the “lollipop,” a form 3-phosphoglycerate. Other with Jacques Mayaudon, Andy of arsenic in kidneys of giant flattened glass vessel that could intermediates of the carbon showed independently of Samuel clams of the Great Barrier Reef, be illuminated from both sides. cycle, including sedoheptulose Wildman that the highly abun- Australia. In 1992, he and Arthur 14 After the introduction of CO2, 7-phosphate (a 7-carbon sugar) dant “fraction I protein” of leaves Nonomura discovered the abil- samples of the illuminated and pentose monophosphates catalyzed the incorporation of ity of methanol to stimulate the Scenedesmus suspension were (5-carbon sugars), were identi- CO2 into ribulose 1,-5-bisphos- removed after short exposure fied by Andy in collaboration phate to yield 3-phosphoglycerate. continued on page 26 ASPB NEWS | VOLUME 42, NUMBER 2  25 Obituary

ANDREW BENSON butions to photosynthesis, this one of us (BBB; see https://www. Roland Douce continued from page 25 metabolic pathway is increasingly youtube.com/watch?v=c4jiYk- Université de Grenoble and CEA growth of crop plants. 13C studies referred to as the “Calvin-Benson W_30). The following year he Govindjee on this effect led to the detection cycle.” Many scientists and histori- celebrated his last birthday, his University of Illinois, Urbana- of an unexpected glycoside, a ans believe that the Nobel commit- 97th, with a dinner held at the Champaign lectin substrate, and a previously tee should have given the Nobel La Jolla Country Club that was unrecognized plant pathway. Prize to both Calvin and Benson. attended by his wife, Dee, and A more complete version of this obit- For 60 years, the work of One of us (Govindjee) persuaded other family members, close uary can be found in Lichtenthaler, Andrew Benson on the discovery Benson to present details of his friends, and former collaborators. H. K., Buchanan, B. B., Douce, R., of ribulose bisphosphate and the time with the Calvin group. This Then, as throughout his life, he and Govindjee (2015). Andrew A. identification of intermediates he later discreetly described in served as a model for all of us. A two personal retrospectives, Celebration of Life ceremony was Benson, 1917–2015. Photosynthesis involved in CO2 fixation has been Research. doi 10.1007/s11120-015- “Following the Path of Carbon held for him on February 6, 2015, at the cutting edge of plant science. 0119-8. The breadth of his approach, in Photosynthesis” and “Paving at the La Jolla Country Club. the quality of his work, and the the Path” (Benson, 2002 a,b). His References contributions were also described Awards advances in understanding that Benson, A. A. (2002a). Following in a television series (Walker, 2012) his discoveries brought about are Andy received numerous honors the path of carbon in photosynthe- simply superb. He captivated the and a popular book (Morton, and awards during his career, sis: A personal story. Photosynthesis enthusiasm of all scientists studying 2007). The breadth and depth of including the Sugar Research Research 73: 29–49. photosynthetic organisms. Rather his contributions to photosynthesis Foundation Award, 1950; Ernest Benson, A. A. (2002b). Paving the than work in the usual competitive became better known in the scien- Orlando Lawrence Memorial path. Annu Rev Plant Biol 53: 1–25. tific community when he made manner, he encouraged others to Award of the Atomic Energy Buchanan, B. B., and Douce, R. become involved and take up his a video with one of us (BBB) in Commission, 1962; Phil.D. ho- (2015). Andrew Benson honored studies where he left off. 2012. The video has been posted noris causa, University of Oslo, on birthday No. 97. Photosynthesis Key to Andy’s success was on YouTube (http://youtu.be/ Norway, 1965; fellow of AAAS, Research 123: 115–116. his habit of working at the bench GfQQJ2vR_xE), and the transcript 1965; Stephen Hales Prize of Buchanan, B. B., Douce, R., and almost every day and his dissat- has been published (Buchanan ASPB, 1972; elected member of Lichtenthaler, H. K. (2007). Andrew isfaction with imprecise results. and Wong, 2013). Filming of the the National Academy of Sciences, A. Benson. Photosynthesis Research 92: He always asked basic, essential video followed the publication of 1973; elected fellow of the 143–144. questions as the starting point for a special issue of Photosynthesis American Academy of Arts and Buchanan, B. B., and Wong, J. H. new research endeavors, always Research organized to commemo- Sciences, 1981; elected member of (2013). A conversation with Andrew approached scientific inquiries rate Andy’s 90th birthday in 2007 the Norwegian Society of Science Benson: Reflections on the discov- ery of the Calvin–Benson cycle. with enthusiasm, and expressed (Photosynthesis Research, Vol. 92; and Letters, 1984; Supelco/AOCS Photosynthesis Research 114: 207–214. great satisfaction in his work. He see also Buchanan et al., 2007). Research Award, American Oil was also known for his inspira- The special issue was presented Chemists Society, 1987; and Govindjee (2010). Celebrating Andrew Alm Benson’s 93rd birthday. tional personality. He would listen to Andy at a memorable dinner Lifetime Achievement Award of Photosynthesis Research 105: 201–208. carefully to the results and ideas held at the historic restaurant Le the Rebeiz Foundation for Basic Harkewicz, L. (2006). Oral history of of others, and then offer solid Procope in Paris that was orga- Biology, 2008. On the occasion Andrew Alm Benson. http://libraries. advice on additional experimental nized to honor him on his 90th th of his 97 birthday, he was hon- ucsd.edu/speccoll/siooralhistories/ birthday (Lichtenthaler et al., approaches. In this way, he intro- ored with the first Andrew A. Benson.pdf duced countless young and visiting 2008). Detailed information on Benson Award for “Conferring Lichtenthaler, H. K., Buchanan, B, scientists to plant biology, motivat- his biography and research activi- the Greatest Benefit on Mankind” and Douce, R. (2008). Honoring A. ing them to perform at the highest ties can also be found in an article for his recent work on the role of Benson. Photosynthesis Research 96: level. He hosted numerous scien- honoring him on his 93rd birthday lectins in improving crop produc- 181–183. (Govindjee, 2010) and in an earlier tists in his laboratory, especially tivity. The award is sponsored by Morton, O. (2007). Eating the sun: postdoctoral scholars. oral history interview (Harkewicz, BRANDT iHammer. How plants power the planet. New In the eight years he spent 2006). York: Harper Perennial. in Berkeley, Andy was the key Andy retained full mental Hartmut K. Lichtenthaler Karlsruhe Institute of Technology Walker, T. (2012). Botany: A bloom- contributor in elucidating the vigor until the very end of his ing history: Photosynthesis. http:// essentials of the carbon reduction life. In his 96th year, he made a Bob B. Buchanan www.bbc.co.uk/programmes/ cycle. In recognition of his contri- congratulatory video on behalf of University of California, Berkeley p011lymf. 26  ASPB NEWS | MARCH/APRIL 2015