A TRIBUTE TO


Colin Allen Wraight (1945-2014)

A Dear Friend and a Brilliant Scientist

by

Govindjee

ColinWraight

Colin Allen Wraight passed away on July 10, 2014 shortly after retiring from his position as professor in Biochemistry, Biophysics & Quantitative Biology, and Plant Biology at the University of Illinois. Colin was also affiliated with the program in Molecular & Integrative Physiology. Above all, Colin was a very good friend to me, and Rajni. He was a wonderful human being and a brilliant scientist. He was always forthright and fearless, honest and upfront. I have always had the greatest respect for him. He was a very likeable person both in and out of the lab, one who always said what he meant. His thoughts were ahead of his time and I learned a lot from him. We miss him dearly.

Colin was born in 1945 in London, England. He studied at the University of Bristol, earning his BSc in 1967, and his PhD in 1971. There, he was one of Antony (Tony) R. Crofts’s first PhD students. Tony pointed out to me that each of the three papers from Colin’s Ph.D. research was seminal in establishing the chemiosmotic framework for photosynthetic energy conversion. They established (1) the dependence of quenching of chlorophyll (Chl) a fluorescence, associated with photosystem II (PSII; the water oxidizing system) on the pH gradient, leading to the important area of non-photochemical quenching (NPQ) of the excited state of Chl a, which is the basis for how plants protect themselves against excess light, This is an area that has interested me greatly and Volume 40 on NPQ in my series, Advances in Photosynthesis and Respiration, is now in press; (2) the dependence of delayed fluorescence on the proton gradient, studied through the kinetics of onset, again, an area very dear to my own research; and (3) the importance of vectorial photochemistry in generating the proton gradient, bringing the immediate energy yield of photosynthesis close to the theoretical thermodynamic limit (a la Nicolas Léonard Sadi Carnot in 1824). Together with Baz Jackson, who joined the Crofts lab at the same time, he represented the spearhead of the photosynthetic wing of the “Bristol mafia”. The successful assault on the then established chemical-coupling hypothesis, led to a much earlier acceptance in the photosynthesis community of Peter Mitchell’s chemiosmotic hypothesis as a general mechanism for electron-transfer linked ATP synthesis than in bioenergetics in the broader context.

Colin1976BrookhavenColin did postdoctoral research during 1971-1972 at the University of Leiden (in Louis (Lou) N.M. Duysens’ lab), continuing his interests in PSII linked fluorescence phenomena, and, during 1972-1974, at Cornell University (with Roderick (Rod) K. Clayton), where he pioneered biophysical exploitation of the newly isolated bacterial reaction center. He also briefly held a faculty position at the University of California at Santa Barbara before joining the University of Illinois as an Assistant Professor in the Departments of Plant Biology and Physiology & Biophysics in 1975.

Colin's many accomplishments were recognized well beyond our campus. He was plenary speaker at the 7th International Congress on Photosynthesis (1986), chaired a Gordon Conference on Photosynthesis (1988), and held a Guggenheim Fellowship (1988). At Illinois, Colin was appointed to the Center of Advanced Studies at UIUC (1988-1989); served as Director of the Center for Biophysics and Computational Biology (1995-1999), and as Head of the Department of Biochemistry (2004-2009), which he joined in 1995.

Colin was in the Department of Botany (Plant Biology) for 20 years before moving to Biochemistry. His laboratory was on the north side of the 2nd floor of Morrill Hall in space currently occupied by the Plant Biology office. I had the other half of that side of the 2nd floor. I had fun collaborating with Colin, discussing and writing papers together. He was not only brilliant and open in his thinking, but was a “hands on” person, and did experiments with us, turning and adjusting knobs, and sometimes even swearing. We collaborated on three topics, publishing four research papers: (1) With my PhD student Paul Jursinic, Colin and I showed, in 1978, a detailed and important relationship of membrane potential with microsecond delayed fluorescence from plants and algae, a topic he had earlier contributed to with Tony Crofts. (2) With several of our PhD students, Colin and I showed, in 1989 and 1992, that bicarbonate, which my Lab had shown to be essential in the reduction and protonation of mobile quinone molecules in plants, algae and cyanobacteria, was not needed at all in a similar reaction in anoxygenic photosynthetic bacteria! (3) With Vladimir Shinkarev, Colin’s long-time brilliant associate, Colin and I succeeded, in 1997, in obtaining new information on the kinetics of oxygen evolution via chlorophyll a fluorescence measurements.

WraightStructureColin’s research topics included Bioenergetics, Photosynthesis, Enzymology, Membrane Biology, Structure and Dynamics of Proteins. Among many early discoveries he made during his life, I note that he was the first to determine the high efficiency of bacterial reaction centers (shown at right), revealing that they converted light energy to chemical energy with a quantum yield of ~98%. He is perhaps best known for his work on the (electron) acceptor side of the reaction center of bacterial photosynthesis; he was the first to establish the two-electron gate mechanism for reduction of ubiquinone by bacterial reaction centers, shortly after the similar reaction in PSII of green plant photosynthesis was first shown in The Netherlands and France. He went on to show that certain herbicides function by displacing bound quinone from the specific site (now called the QB site) catalyzing the two-electron gate reaction. His interests in this area continued until the end, leading to major contributions to our detailed understanding of this important process, and the intimate coupling between electron and proton transfer leading to reduction of quinone.

Colin's publications, which span 44 years, are extensive and varied, showing his versatility, thoroughness, and originality in research. Although he worked with many systems, his most extensive contributions have been with his favorite organism, Rhodobacter sphaeroides. This photosynthetic bacterium has provided a playground for biophysical research, taking advantage of photoactivation of photosynthesis to probe reactions over 15 orders of magnitude in rate. Colin was always at the forefront in recognizing how this range could be exploited in the understanding of fundamental processes of proton-coupled electron transfer, leading to general lessons applicable in a much wider context.

Colin1999GovRetirementI remember many things about Colin, but three personal recollections come to mind. The first was when Colin gave his first seminar at Illinois before he was hired in 1975: he kept talking long after the allotted time had passed, but everyone was glued to his engaging presentation, rather unusual for the department. The second was when he was going out to celebrate his first wedding anniversary. He was indeed wearing a tie (a rare thing), and Mary asked Rajni to tie an Indian sari on her. The couple looked like Bollywood movie stars. The third was when I transferred to Plant Biology when the “Biophysics section” in “Physiology & Biophysics” was being closed in favor of a Center of Biophysics. A condition was placed on me that I would have to teach a large introductory course, which quite upset me until I found out that it was Colin who had suggested it, and then I accepted it right away. Later, I thanked him because I discovered that I really loved to teach those 700+ students.

The most recent work from his lab -- three papers with his PhD student, Alexander (Alex) T. Taguchi, who graduated this summer -- has appeared over the last year, reporting collaborative research with Sergei Dikanov exploring atomistic details of the two-electron gate mechanism in photosynthetic bacteria through use of high-resolution Electron Paragmagnetic Resonance (EPR) and specific isotopic labelling of the protein and of the quinone. Two other students, Aidas Mattis and Charles Sun, are completing their PhD degrees, and we can expect several more papers from each of them. The last poster (from a 2013 Gordon Conference) by Colin details his brilliant thoughts on “Design features of long distance proton transport in biology”. It is currently hanging outside 671 Morrill Hall for anyone to see and study.

An incomplete list of his collaborators (in alphabetical order) is: Rod Clayton, Tony Crofts, Sergei Dikanov, P. Leslie (Les) Dutton, Govindjee, Paul Jursinic, Jack S. Leigh, Peter Maroti, James (Jim) R.Norris, Robert (Bob) E. Overfield, David (Dave) R. Paterson, Vlad Shinkarev, Robert (Bob) J. Shopes, Alex Taguchi, Eiji Takahashi, and Xutong Wong. In 2013, he wrote a beautiful Tribute, in Photosynthesis Research, to his postdoc advisor “Roderick K. Clayton: a life, and some personal recollections.” (See PubMed for his publications.)

GCGovBookAfter Les Dutton’s talk at the 2014 Gordon Conference on Photosynthesis (Chair: David Kramer), on Colin's life and enormously insightful and brilliant research contributions, I honored Colin by presenting four volumes from the Series “Advances in Photosynthesis and Respiration” to four of the best young researchers “In Memory of Colin Wraight” (photo at right). There is now a plan to recognize Colin Wraight at forthcoming Gordon Conferences on Photosynthesis by awarding cash prizes.

In addition to his brilliant research contributions, Colin was a passionate teacher and mentor of many students and colleagues; he always gave unselfishly to others. He had unmatched breadth and depth, and quick wit; we will all remember the gracious hospitality that he and his wife, Mary, extended to all of us in SIB and MCB. We will remember his dedication to teaching and graduate training even during his illness. This was an inspiration to all who knew him. Throughout his long battle with the cancer, Colin maintained an active life in research and in departmental affairs. Even in the darkest periods, his fortitude, self-deprecation and wry sense of humor showed the real Colin, always a force to be reckoned with.

Mary, and their children, Lydia, Tristan and Sebastian, remember Colin fondly as the most loved person on earth. Neither they, nor I, nor his many friends, can accept that he is no longer here with us.

- Govindjee