In Memoriam: Ann Silver

It may come as a surprise to many of us who regarded Ann as quintessentially English to learn that her family was from Scotland and that her great, great grandfather was a friend of Alexander Boswell (son of James Boswell, Samuel Johnson's biographer). Moreover, she was not born in Scotland or in England but in Meerut in the western part of Uttar Pradesh (where the ‘Indian Mutiny’ had started). To make it even more complicated her father was born in Grenada (West Indies) where his father, a Minister in the Church of Scotland, was working. Grenada had become independent in 1974 so when applying for a passport in 1974, Ann had to get her grandfather's birth and marriage certificates before she could become British again. Her family returned to England in 1930 and her father (who had been in a cavalry regiment) set up a riding school. She had a country upbringing in Suffolk with horses, animals and nature, and this, she believed, is why she became a physiologist. Her schooling was disrupted by World War II but at the end of the war she returned to St Felix's school in Suffolk to take her school certificate in physics, chemistry and biology. From there she went to the University of Edinburgh. In her first year she read zoology, chemistry and physics. In her second year she studied physiology with Catherine Hebb (who ran a physiology course for non-medics). In her final year Ann studied physiology under David Witteridge, graduating in Physiology in 1953. At about that time, Catherine Hebb moved to the Babraham Institute in Cambridge, but Ann began her PhD work in Edinburgh. Her formal supervisor was George John Romanes, but for day-to-day work she turned to Catherine Hebb. Ann spent her first year in Edinburgh but was funded by the Agricultural and Food Research Council (ARC) and later moved to Babraham, working on cholinergic transmission, investigating the effects of organophosphorous compounds. She was involved with several techniques, recordings, histology and histochemistry, working on choline acetyltransferase (choline acetylase) and axonal transport. Her work at that time was compromised by her need for orthopaedic surgery on her spine initially in Edinburgh and then in Cambridge which placed her in a great deal of pain. Despite this she completed her thesis work (with Catherine Hebb) largely on enzyme transport along lesioned nerves. Her PhD examiner was R. H. S. Thompson from Guy's and the internal examiner was David Whitteridge. She said her thesis was readable, but in fact it was beautifully written. In the early part of her career, Ann rubbed shoulders with a small community of female scientists at a time when most scientists were men. For example, apart from Catherine Hebb, a Canadian who was the sister of Donald Hebb, best known for his theory of Hebbian learning, she met and knew Marthe Vogt, who is remembered for her work on the role of neurotransmitters, especially adrenaline, and whose sister Marguerite, a cancer biologist and virologist, was noted for her research on polio at the Salk Institute for Biological Studies in California. After completing her PhD, Ann collaborated with many people at Babraham: for example, with Tim Biscoe on the carotid body and Kris Krnjević on the iontophoretic application of acetylcholine to neurons in the cerebral cortex. Later she went to Montreal and worked with Kris there. She also collaborated with Wilhelm Feldberg at Mill Hill, mostly working with Pedro Guertzenstein; she also did quite a lot of collaborative work with Charles Shute and Peter Lewis then in the Anatomy Department in Cambridge. Her collaborations were almost always related to the cholinergic nervous system. When asked sometime later what she was most proud of scientifically, she replied: ‘the experiments with Kris Krnjević; I enjoyed them very much. Some of the other work was rather bread and butter.’ These experiments involved all the techniques with which she was familiar, histochemistry of cholinergic systems and recording the chemical sensitivity of cortical neurons. They were published in the Journal of Physiology are reviewed clearly in her book The Biology of Cholinesterases (Frontiers in Biology vol. 36, North-Holland Publishing Co., Amsterdam, 1974). In the early 1970s, however, cholinesterases became less fashionable. This may have been because acetylcholine and the enzymes associated with it seemed to be found in numerous parts of the nervous system, many in the basal forebrain, and could not be associated with a particular neural system. Since then, it has become apparent that cholinergic neurons provide inputs to many regions including the cerebral cortex and make a complex contribution to the physiology of wakefulness and sleep. They have also become a focus of research in ageing and neural degeneration, including the pathological events in Alzheimer's disease. She continued for a while working with muscimol and GABA. However, at that time (1970−1978) she was much involved with the Physiological Society countering anti-vivisectionist activity. In 1974 Barry Cross became the Director at Babraham and she was persuaded to come out of the lab to become Information Officer at Babraham, answering Press enquiries relating to anti-vivisectionist activity and continued to work on her book. She found the writing rewarding but at the same time hard work, so much so that she said later, ‘I was glad when it was finished’. In 1985 Barry was forced to make a number of people redundant, a group that included Ann. At this point she moved to the Physiology Department in Cambridge and worked in the Physiology Library. Here she started editing and reviewing for the Physiological Society and began the part of her career for which she is perhaps best remembered by many people. She also started to teach, not so much physiology but how to present the physiological work of others. She started with Part II students (third-year undergraduates) helping them to write up their projects. As time went on this work expanded to include PhD students, eventually across the Faculty of Biology, and in other universities. For many years she gave courses on how to write a PhD thesis. The courses had three parts: first on organising the work, second the writing itself, and finally one presenting the figures, tables and references. She did not want to teach physiology but rather how to present it. In her words: ‘My problem is that I have doubts about things and think I would have possibly been a confusing lecturer. Instead of saying, “This is so, and this is so, and that is so”, I would have been trying to cover the bits that didn't quite fit. However, when teaching students how to write I did feel that I knew what I was talking about and so could give confident lectures, whereas I think I would have given rather wishy-washy physiology lectures.’ When asked later if she was a member of staff of the Department of Physiology in Cambridge, she said, ‘no. I've never been a member of staff. I've been a visitor for over 23 years’, but she clearly earned her keep working for the Physiological Society, of which she was a member of the Committee from 1978 to 1982, became a Trustee of the Benevolent Fund, and enjoyed many interactions with Andrew Huxley, Richard Adrian, Robert Comline, Pat Merton, Tony Angel and Denis Noble, and many of the editors of the Journal of Physiology. The 1970s were a busy time for the Silver family. There were three Silvers in Cambridge: Ann, her brother Ian (who later moved to Bristol) and Marian (who died tragically early). Ian told the story that he had a student at Churchill who said: ‘I've just read your book.’ So, Ian said, ‘No, that's my sister's book.’ So, he said, ‘Oh yes, I've just heard her lecture.’ So, Ian said, ‘No, that's my wife.’ Ann joined the Society in 1963. She saw many changes to the Society as time went on. When she joined the membership, it was considered a step up as the Society was regarded as elitist. Applications were submitted to the Committee and each member was given a bundle of applicant proposals to consider. It could sometimes take about 2 years from being proposed as a member to being elected. Big changes were made in the governance of the Society in the late 1990s. The largest was in centralisation. Before that the Society had been run by its members and senior members, in a way that would today be regarded as eccentric. For example, there was no central office. Society Officers did not have secretaries but took advantage of the services available in their own lab or department. Some were very dependent on family members. As soon as the Society was centralised, there developed a cadre of people who were managers, and who had views based on business management, reducing the influence of Members. Society meetings became less frequent, the pattern of meetings changed and voting on acceptance of Abstracts was stopped in 2004. Council meetings met independently of Society Meetings and were less frequent, perhaps only three times a year, and the numbers attending were larger. In 1980 the publications office was set up in Cambridge. When Julian Jack retired as Treasurer the first Administrative Office was set up, in Oxford, with Heather Dalitz in charge. It stayed in Oxford for about 6 years and then was re-established in London. Ann felt that moving the Publications Office to Cambridge has been a success and there has not been a very big turnover of staff. We owe a debt of gratitude to Linda Rimmer, Carol Huxley, Ann Watson, Diana Greenslade and Jonny Goodchild and the rest of those who worked for The Journal and were very loyal to it. In a later interview with Ann in 2008, Dafydd Walters asked her what happened when she left Babraham. He mentioned that she had said, ‘Comline rescued me’. Were you in employment by his office? Ann replied, ‘No, I've been unemployed since 1985.’ Ann's attitude to life can be summed up simply: ‘She always did what she considered to be right.’ Her book The Biology of Cholinesterases was described by one reviewer (R. B. Collier) as follows: ‘The book is so clearly written that it is a pleasure to read.’ This article draws heavily on an interview with Ann Silver that was conducted on 15 May 2008 by Dafydd Walters and Martin Rosenberg. None. R.D.: Conception or design of the work; Drafting the work or revising it critically for important intellectual content; Final approval of the version to be published; Agreement to be accountable for all aspects of the work. None. Please note: The publisher is not responsible for the content or functionality of any supporting information supplied by the authors. Any queries (other than missing content) should be directed to the corresponding author for the article.

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