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The death of Patricia Lund on 13 December 2023, marks the end of an era in metabolic research defined by Hans Krebs in his memoirs as follows: "When examining a biological phenomenon, it is always important to examine the whole process and not merely a fragment in a damaged tissue" HA Krebs, Reminiscences and Reflections, Oxford University Press (1981); p63. Pat was the last of the five original members of the Metabolic Research Laboratory (MRL), formed after Krebs's mandated retirement from the Whitley Chair of Biochemistry at the University of Oxford in 1967. The MRL was housed in a wing at the back of the Radcliffe Infirmary, next to a medical ward, and administratively a part of the Nuffield Department of Clinical Medicine. Besides Professor Hans Krebs (1900–1981), aka 'The Prof', and Pat, other core members of the MRL were Leonard Eggleston (1920–1972), Reginald (Reg) Hems (1926–2005) and Dermot Hedley (Derek) Williamson (1920–1998). The MRL was a bustling place, with about 15 members orderly distributed among benches and desks in one large room; it was a uniquely happy place and productive scientific environment, quietly, but strongly, led by Krebs and ably supported by the very capable scientists he surrounded himself with, including Pat. Almost unique to the lab was a plethora of visiting researchers from all over the world, many of whom later went on to great eminence. All of them relished a highly positive atmosphere, due largely to the welcome, hospitality and stimulation of the permanent members: Pat Lund was highly regarded by all for her wit, warmth and wisdom. And for her unfailing willingness to help others, at the bench and in the library.Patricia Lund was born in 1933, in North Yorkshire, England, to a farming family. She attended Malton Grammar School, attaining A levels in physics, chemistry and biology, unusual choices for girls at that time. She opted not to attend university as she believed her family would be unable to support her financially, in spite of her interest in agricultural science and in pharmacy. Instead, she chose to work as a laboratory technician for the nearby York-based confectioners Rowntree, a major employer in the area. Subsequently, Pat worked for a small company (J 182: 1430] – what a start for a career in science! After several years in Oxford, in 1960 she moved to the USA and worked with Boris Magasanik in microbiology, initially at the Harvard Medical School, subsequently moving with his lab to the MIT in Cambridge, MA. At this time, she was considered more like a research associate than a technician and was beginning to find her own scientific voice. Pat returned to Oxford in 1962 when Krebs (on the recommendation of Kornberg – himself also originally a lab technician, who had worked with Krebs at Sheffield University and who by then had moved to Leicester) invited her to become a graduate student in his laboratory. Professor Hans (later Sir Hans) Krebs had been awarded the Nobel prize in 1953 and was now the Whitley Professor of Biochemistry at Oxford University and founder of the MRC Unit for Research in Cell Metabolism in that department. She accepted (and declined an offer by Fred Sanger to work as his personal research assistant in Cambridge, UK!) and moved back to Oxford. Nobody believed that Pat was 'just a farmer's daughter from Yorkshire' and that in her research she simply addressed 'just the questions she saw in front of her'. Both are, of course, true statements, but also profound understatements. Pat was admitted to Linacre College, Oxford University, where she obtained her undergraduate degree in 1965. When Krebs officially retired and established the MRL across the road in the Radcliffe Infirmary in 1967, Pat moved with him, gaining her DPhil in 1969, supervised by Krebs.Her early work centred on not only renal glucose synthesis, the glyoxylate cycle (described shortly before in 1957 by Kornberg and Krebs) and the urea cycle, but also hepatic metabolism and the redox state. Her 1967 paper in the Biochemical Journal co-authored by Krebs and Derek Williamson Williamson DH, Lund P, Krebs HA. Biochem J (1966) 103: 514 on the mammalian hepatic NAD+/NADH ratio in cytosol and mitochondria became one of the most cited papers in the field of oxygen-mediated respiration (2150 citations to date), a Science Citation Classic. Marion Stubbs (1940–2012; latterly of St. George's Hospital Medical School) was a fellow DPhil student in the MRL and a life-long friend of Pat's, sadly pre-deceasing her. Later, Pat's interests turned to mammalian amino acid metabolism. She described a radiochemical assay for glutamine synthetase which transformed research in this entire field. Some decades later, Pat's original observations on glutamine metabolism and the unique metabolic profile of rapidly dividing enterocytes laid the foundation for many unexpected new models in cancer cell metabolism. By a twist of fate, Pat should die from cancer, bravely borne.In a personal letter to one of us (HT), 2 years ago, Pat commented on a piece I had been asked to write about the MRL: "Your article brought back happy memories of our days in the MRL. Sad that so many colleagues from those days are now dead. Our recollections are obviously slightly different. We used to dread the Prof 'helping' us with an experiment. It usually ended in disaster. I remember him causing a fire when demonstrating how to insert a stick of phosphorus into the center well of a manometer cup. On another occasion, he let some mice escape from a cage and one shot up his trouser leg. It was a case of 'Do as I say and not as I do.' The Prof. was quite naive and unworldly in some ways. Reg's feeble jokes left him doubled up with laughter. The Prof. didn't drink alcohol, and one day, after lunch, he encountered Brian Ross, who said having half a pint of lager had made him feel sleepy. Prof's response: 'Couldn't you ask them for a quarter of a pint.' I myself once asked for a few hours off, promising to make up the time. 'Pat, you can never make up the time' and refused my request. I owe the Prof. a lot. He took me on – also Derek Dermot Williamson – to do a DPhil when I didn't even have a first degree yet – bending all the Oxford rules. I now support the Sir Hans Krebs Trust, which son John, now Lord Krebs, initiated after the family sold Prof's Nobel Prize Medal. It made a fraction of Watson's! I'm afraid I haven't kept up with research on glutamine or anything else and had to look at my old papers to remind myself about gut cells. I'm amazed all that stuff is still relevant."When the leadership of the MRL passed on from the Prof to Derek Williamson and thence to Geoff Gibbons, Pat's involvement in scientific research and teaching at the bench continued in the MRL. She was an editor of the Biochemical Journal. She continued to make seminal contributions with David Wiggins and Geoffrey Livesey, elucidating branched-chain amino acid metabolism and inter-tissue nitrogen fluxes and leaving a rich legacy in the field of intermediary metabolism.Towards the end of her working life, Pat eventually found her way back to the Oxford Science Area, on South Parks Road, working with Kieran Clarke and George Radda on cardiac metabolism, initially returning to the Biochemistry building, where she had started, and latterly moving to the Physiology building, with the same group.Pat was a distinguished scientist who succeeded in surmounting not only the 'glass ceiling' but also the toughened 'glass door' through which women had to pass to enter a male-dominated environment, by her talent and determination. Driving through Oxford in her dark green MG sports car, she was a lady with predictably positive poise who gave elegance to this environment – always immaculately turned out, she alone in the lab managed to make a white lab coat look like a fashion accessory.Pat moved to France following her retirement, but in later years moved back to the family farm in North Yorkshire where she had been born, with her husband Keith Maddox, who survives her. She had two younger brothers, Richard (who pre-deceased her) and John (who survives her).It was our unforgettable privilege to have worked with her.
Taegtmeyer et al. (Fri,) studied this question.