Rainer Walter Guillery FRS, 28^th August 1929–7^th April 2017

Ray Guillery was a major figure in the field of neuroanatomy, particularly of the visual system. His ancestry, early life and role models during his undergraduate and graduate training all provided influences that strengthened his innate bias towards this research career. His approach was rooted in descriptive anatomy and was driven by a desire to define and describe the pathways underlying visual function. In his autobiography (1998) he defended this approach as follows: ‘The important roots of neuroscience in accurate descriptive accounts are often overlooked, and the joy of arriving at a reasonably accurate and lasting description of a structural relationship is not as widely appreciated as perhaps it should be’. More prosaically, he was fond of saying: ‘If you can't draw it, you don't understand it’. This strongly defined aspect of his character led to him choosing to take a BSc degree in Anatomy at University College, London, instead of continuing his medical training.

Ray was born on 28 August 1929, in Greifswald, Pomerania, on the Baltic coast of Germany. His father was a pathologist who, while training at the Charité Hospital, Berlin, met and married a Russian Jewish refugee who was a histology technician there. His paternal grandfather was a medically qualified ophthalmologist who specialised in visual acuity. His paternal grandmother's uncle, Otto Deiters, was an eminent neuroscientist whose name lives on in the lateral vestibular ‘Deiter's’ nucleus; the name was also originally applied to dendrites (Deiter's processes). There were medical relations on his mother's side, too; however, his maternal grandfather, an apothecary in St Petersburg before the First World War, had been forbidden by Nazi policies to practise by the time Ray knew him, and used to take the young boy to the zoo and on nature walks.

Ray's parents divorced when he was quite young. The combination of this and the political situation meant that his family became scattered among different relatives, so his childhood was very disrupted. He attended a Rudolf Steiner school in Berlin (1935–38), followed by brief periods in Switzerland and Holland. At the end of the summer term 1939, he and his sister went to stay with his godmother in North London, where they were also reunited with their mother. The two siblings were booked to return to Holland on 3 September, but the outbreak of war that very day left them stranded in England. Their mother was acting as au pair to a London family who then evacuated their children to Oxford, where Ray attended a ‘rather shabby’ preparatory school before moving to Sibford School, a Quaker boarding school in the Cotswolds (1940–46). He felt as an adult that his moral outlook had been influenced by the first and last of these schools. His holidays were largely spent in the North Oxford home of Professor Wilfred Le Gros Clark and his wife, where his sister was based. Le Gros Clark was at that time Dr Lee's Professor of Anatomy (see obituary by Weddell, 1972); Ray described him as ‘rather silent and distant’ but enjoyed cycling with him in the countryside at weekends, and was introduced to his laboratory, including the primates in the animal house. Through Ernst Chain, who had known Ray's godmother's family in Berlin, his mother was enabled to pick up her professional skills after the war and work as a pathology technician. Being keen for him to become a doctor, she prepared slides with serial sections of a guinea pig embryo for his Christmas present one year, to go with the microscope he also received.

Two years at a grammar school, and much independent work, resulted in a scholarship to study medicine at University College, London. There he was taught by J. Z Young and Bernard Katz among others and quickly became convinced that he wanted to aim for a career in research rather than medicine. He was awarded a scholarship to do an intercalated BSc in Anatomy, a new course started 2 years earlier, with P. K. Thomas as the only pupil that year (see obituary by King, 2008). In addition to the regular course of lectures and tutorials, chiefly by J. Z. Young, there were ‘intercollegiate’ lectures from other anatomists. Several of these were people who made important contributions to both Journal of Anatomy and the Anatomical Society, e.g. Frank Goldby, W. J. Hamilton, J. D. Boyd and E. C. Amoroso.

After completing his BSc in 1951, Ray stayed on at UCL as a PhD student with a 2‐year scholarship, under J. Z. Young's supervision. This was his first foray into neuroscience: a quantitative study of the axons of the fornix, from the hippocampus to the mammillary bodies of the hypothalamus. At the suggestion of his sister, he visited Oxford to re‐establish contact with Le Gros Clark. There he discovered to his horror that Daitz, one of Le Gros Clark's research associates (the other was Tom Powell), was also working on the fornix. Fortunately it was a complementary pathway, so Ray's research plans were not affected; on the contrary, after Daitz's sudden death, he continued to collaborate with Tom, and the two of them, later joined by Max Cowan, had many productive discussions and joint publications over the years, several of which were published in this Journal.

Ray remained at UCL as Assistant Lecturer, then Lecturer. He built on his thesis work by using the then new Nauta staining technique to trace the fibres from the fornix to the anterior thalamus. Like many of us early on in our research careers, he discovered that he had a competitor: Nauta himself had carried out the same study and had already submitted it to the Journal of Comparative Neurology. Ray nevertheless submitted his own paper to the Journal of Anatomy, which had a shorter publication delay, and both papers appeared in 1956. More pathway‐tracing work using the Nauta method followed, and a paper in the Journal of Anatomy on the hypothalamic connections published in 1957 became a citation classic. Soon after this, his desire to understand the fine structure of synapses under different conditions drew him into electron microscopy. This early research was largely carried out without collaborators; obviously this changed as his career progressed, and it is not possible to give due credit to all of them in the account that follows.

A year's sabbatical leave at the University of Wisconsin at Madison (1960–61) was not as productive from the research point of view as he had hoped but had the great advantage of widening his horizons in terms of methods of critical thinking as well as new contacts. On returning to London he began to work with Peter Ralston, successfully applying the Nauta stain to electron microscopy preparations. Technical developments at that time included the introduction of aldehyde fixation, which provided improved ultrastructural definition compared with that following osmium tetroxide fixation alone. Promotion to a Readership at UCL in 1963 and several job offers led him to think seriously about the future, and the following year he moved back to Madison, where he remained until 1977. In Madison he enjoying teaching undergraduate courses in neuroanatomy, and found his research very productive and rewarding. Colleagues and visitors included Max Cowan, Semir Zeki and Peter Ralston. Work on the visual system led to an interesting discovery about the fibre pathways from the retina to the lateral geniculate nucleus in Siamese cats: in this species, and (he later discovered) in all albinos, many of the fibres take an uncrossed instead of a crossed path. This means that the retinal input to the lateral geniculate nucleus is abnormal, and the animals lack binocular vision.

An offer from the University of Chicago enabled Ray to take up a challenge he had long desired, that of setting up a graduate neurobiology programme; he ran this successfully and with much enjoyment from 1977 to 1984. His work on the visual pathways continued and was extended to pigmented and albino ferrets: these have the advantage of being born at a less advanced stage of visual development than are cats, so can be studied postnatally at stages that are prenatal in cats.

In 1984, Ray was appointed to the Dr Lee's Chair of Anatomy at Oxford. This chair was founded in 1919 with the appointment of Arthur Thomson, and held by Wilfred Le Gros Clark from 1934 to 1962. After the initial difficulties he inherited with the poor relations between academic and technical staff, things ran smoothly. He was very supportive of his research‐active academic staff, one of whom was his old collaborator Tom Powell. He was also very keen to ensure that all the teaching was done to the highest standards and, not surprisingly, he took a very active interest in the neuroanatomy classes. Although he encouraged junior staff to take the lead, he was always there to demonstrate in the practical sessions, where his expertise was much sought after not only by the students but also by the staff. His research group was active and productive; in addition to his work on the optic chiasm, he investigated the thalamic reticular nucleus. This was the renewal of a project begun in Madison; it led to the discovery of an adjacent group of cells, the perireticular nucleus, whose functional significance is still not clearly understood.

While at Oxford he became the founding Editor‐in‐Chief of the European Journal of Neuroscience, a position he greatly enjoyed. He served as President of the Anatomical Society from 1993 to 1995, was Treasurer of the Society for Neuroscience and was on the editorial board of several neuroscience journals. Retirement in 1996 did not bring Ray's research career to an end: he returned to Madison with the title of Visiting Professor and Senior Scientist, and continued active experimental work and publication there for another 10 years.

In 1954, Ray married Margot Pepper, at that time a medical student at St Mary's; they had four children. Sadly, what had been a happy marriage for 30 years did not survive the move from Chicago in 1984, where she had a staff position in Dermatology. The promised position for her in Oxford did not materialise, and she returned to the USA. Their youngest child, Jane, married a Turk and works as a teacher and translator in Istanbul. Ray moved there from Wisconsin in 2006 as visiting professor in the Anatomy Department of Marmara University, to be closer to her and his grandchildren. By 2010, growing deafness and an inability to speak Turkish made him feel increasingly isolated, and he returned to Oxford, where he lived until his death. For most of those last years he was again a very welcome demonstrator in the neuroanatomy classes, and always enjoyed chatting about some of the latest neuroanatomical findings in the mid‐session coffee break. He welcomed visitors to his compact house with its well‐kept vegetable garden, where he gave G.M.‐K. some very helpful information on the historical aspects of neuroanatomy for her articles on the history of the Journal of Anatomy. Ray's last paper (Dieters and Guillery, 2013), co‐authored with his cousin, was a review of the work of their great‐great uncle, the neuroscientist Otto Deiters (1834–63). This was a very appropriate way to tie together his family history and his neuroanatomical research, and a fitting end to a lifetime of scientific publication.