Alan Clarke was born on 5 June 1963 and studied cellular pathology at the 51¹ú²úÊÓƵ of Bristol (1981-84) before going on to a PhD at the 51¹ú²úÊÓƵ of Cambridge (1984-88). There he worked in the laboratory of Sir Martin Evans, who went on to share the 2007 Nobel Prize in Physiology or Medicine for his research in culturing embryonic stem cells and using them to cause genetic modifications in mice.

After completing his doctorate, Professor Clarke obtained a position as a research fellow (1988-94) ¨C later a Royal Society university research fellow (1994-99) ¨C in the 51¹ú²úÊÓƵ of Edinburgh¡¯s department of pathology.

He joined a team led by Andrew Wyllie, now emeritus professor of pathology at Cambridge, in exploring the new technology of gene targeting and produced a much-cited paper in Nature about how the deletion of the p53 gene in mice can prevent damaged cells from dying and hence lead them to form tumours.

Professor Wyllie recalls him as an ¡°inspirational, thoughtful, cheerful¡± colleague who ¡°quickly demonstrated his remarkable technical skills, his love of science and?his astonishing capacity for?work. The whiteboard in his office swiftly filled up with simultaneous ambitious projects ¨C completed, attempted and?contemplated¡­It was clear even from these early days that he was destined to contribute to the highest ranks of cancer science.¡±

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In 2000, Sir Martin ¨C the current chancellor of Cardiff 51¹ú²úÊÓƵ who was then professor of mammalian genetics ¨C was delighted to persuade Professor Clarke to move to Cardiff¡¯s School of Biosciences as a professor.

He would remain there for the rest of his life, developing mouse models for delivering gene mutations that were both tissue- and stage-specific and providing ever deeper insights into the early stages of tumour development. This had direct implications for indicating and assessing new therapeutic approaches, which he was able to pursue in close collaboration with a number of major pharmaceutical companies.

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A leading player in establishing and then heading the European Cancer Stem Cell Research Institute at Cardiff, Professor Clarke was proud that this rapidly created the first experimental anti-cancer stem-cell drug. He also served as director of the CRUK Cardiff?Centre as well as head of research for the School of Biosciences.

Professor Clarke was described by Sir Martin as ¡°an eternally upbeat, individualistic colleague and a superb scientist, teacher and leader¡±. He died unexpectedly on 28 December 2015 and is survived by his wife Kathryn and twin daughters Naomi and Lucy.

matthew.reisz@tesglobal.com