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R. Porter, U. Proske and R. F. Mark This memoir was originally published in Historical Records of Australian Science, vol.15, no.1, 2004. When Archie McIntyre died peacefully in St Vincent’s Hospital in Launceston, Tasmania on 20 July 2002, Australia lost one of its most significant contributors to the development of modern neuroscience. Less well known, perhaps, because of his self-effacing manner, than eminent peers like Jack Eccles, he was nevertheless a major driving force behind Australia’s excellence in neurophysiological research. The power of his intellect, his creative abilities and his practical skills were recognised early and he was given the opportunity to use these to make innovative contributions to research, first while he was still a student at the University of Sydney, later as a young medical graduate and then in aviation medicine during the Second World War. His interest in nerves, sensory receptors, their reflex actions and their projection to the brain, developed after the war when he spent time at the Rockefeller Institute in New York and at Cambridge University in England where he was influenced by the leading scientists of the day. He brought back to the University of Otago in Dunedin, New Zealand and then to the foundation days at Monash University in Melbourne an enthusiasm for research and scholarship that established these as leading centres for neurophysiology internationally and Archie himself as a world authority on sensory receptors. He was a leader in the growth and the establishment of scientific societies (Australian Physiological and Pharmacological Society, Australian Neuroscience Society) and served Australian science and society through his work with the Australian Academy of Science, the Australian Research Grants Committee, and the Australian and New Zealand Association for the Advancement of Science. Family Background and Formative Years Archie was born on 1 May 1913 in Edinburgh, Scotland, the second of four children of Dr William Keverall (Bill) and Margaret (Madge, née David) McIntyre who had moved to Scotland from Australia for Bill to study medicine. Bill McIntyre was from Hobart, Tasmania. In talking about him, Archie described Bill as ‘a bit of an old warrior’, for he had volunteered for both the Boer War and the First World War. Bill’s mother, Adeline Janette, was said to have flown with Charles Kingsford Smith. Bill chose to study engineering at the University of Sydney with the intention of becoming a mining engineer. There he met and married the daughter of the Professor of Geology. Edgeworth David was a distinguished scientist who came to Australia in 1882 to the post of Assistant Geological Surveyor for the Government of New South Wales. David was subsequently appointed Foundation Professor of Geology at the University of Sydney in 1891 and, along with his student, Douglas Mawson, joined Shackleton’s Antarctic expedition on the Nimrod in 1907. During the summers of 1907/08 and 1908/09 he led parties that climbed the active volcano, Mt Erebus, and walked, pulling sleds, to the South Magnetic Pole and back in time to be picked up again by the Nimrod and returned to Australia. After discussions with his eminent father-in-law, and with his support, Bill abandoned engineering and transferred to medicine. Here a determining factor was his wife’s illness following the birth of their first-born child (Peggy). Madge developed puerperal fever from which she eventually recovered. The experience, however, led Bill to change careers. With financial assistance from Edgeworth David he travelled to Edinburgh to undertake medical studies, which explains how Archie came to be born in that city. Bill graduated in medicine in 1915 and joined the British 80th Field Ambulance, which saw service first on the Western Front and then in Macedonia. He returned to Edinburgh in 1919 and brought the family back to Launceston in 1920 where he remained for the rest of his life. Bill and Margaret were well known personalities in the lives of many Tasmanians. Margaret was involved in aspects of community work, especially in relation to drama, music and the status of women. She was elected to the upper house of the Tasmanian Parliament in 1948. Three months later she died in an airline crash on her way home to Launceston following a National Council of Women Conference in Brisbane. Bill was greatly loved for his life-long devotion to the health and well being of the people of Launceston, especially of the women and children. He seemed to have been present at almost everyone’s birth! Archie’s education began at home. He learned the three Rs and developed a love of poetry under the guidance of his highly cultivated and intellectual mother, before commencing formal schooling in Launceston at the age of 8. His early schooling was hampered by his shortsightedness so that these were not happy times for Archie who couldn’t see the blackboard from the back of the classroom. And, anyway, he already knew most of the things being taught. His parents therefore sent him to Sydney in 1925 to spend the last four years of secondary education at Barker College in Hornsby. During this time he got to know his famous grandparents well and became particularly fond of his aunt Mollie whom he subsequently referred to as his second mother. He did well in secondary school and won a University Exhibition in 1929 at the age of 16. In 1930 Archie enrolled at the University of Sydney as a BSc student with a special interest in Chemistry. During his first year he discovered the fascination of Biology, which had not been taught at his boys’ school, and he transferred to Medicine to follow this new interest. Although he said that the transition from Science to Medicine was at first difficult, he had a dedication to learning and an excellent memory that soon took him to the top of the class. He became one of the small group of students selected to be a Prosector in Anatomy in 1932. In a tape-recorded interview with his sister, conducted at his home in Launceston in November 1996, Archie recalled that he had transported partially dissected body parts home to Hornsby on the train for further study. When asked how his grandparents had reacted to having parts of a corpse in the house, his reply was ‘I don’t think I told them’. His experimental skills led him to enrol in 1933 for a BSc in Medical Science, which involved a research project and preparation of a thesis. He mastered venepuncture and blood gas analysis using the Haldane apparatus. The professor of physiology, H. Whitridge Davies, supervised his project. He had to master use of the complicated apparatus required to measure oxygen and CO2 tensions in the blood, as well as to devise methods of withdrawing blood samples from his own veins. Here Archie developed an ingenious method of attaching the plunger of a syringe to the gas lines at the back of the laboratory bench. He then inserted the needle into his own vein and carefully moved his arm back away from the bench to draw blood into the syringe. He rejoined the medical class in 1934 and studied pathology and pharmacology in fourth year. He spent most of his ‘clinical’ time in fifth and sixth year at the Royal Prince Alfred Hospital where ‘it was easier to keep in touch with the Profs, people like Harold Dew and Lambie’ (quoted from transcript of interview in November 1996). He graduated in 1936, when he was awarded the University Medal and shared top place in the graduating class with Ruthven Blackburn. During his early years in Tasmania, Archie had acquired a love for the bush. His father had taught him some of the basic survival skills as well as how to hunt and fish. Archie subsequently became an enthusiastic trout fisherman, a sport he continued to pursue until well into retirement. During the early 1930s, whenever he came home from Sydney for the summer vacation, Archie went bushwalking. Together with his sister Peggy, he began walking through the Tasmanian wilderness, at that time largely unexplored country (Sharman, 1998). Those early trips were done with a minimum of equipment — no tent, just a blanket and groundsheet, a few supplies and the fishing rod. They often headed into the Cradle Mountain area, in particular, and into what is today the Walls of Jerusalem National Park. This was all wild, uncharted country and as they travelled through it they named some of the lakes they passed, like Lake Tyre (from McIntyre) and Ahchees Lake. These experiences instilled in Archie a love of the natural world that contributed to his sense of humility over man’s place in nature and stimulated him to attempt to understand nature through science. Archie spent 1937 as a Resident Medical Officer at Royal Prince Alfred Hospital, Sydney. His gift for making things work attracted the attention of Sydney’s clinical leaders like Sir Harold Dew, Professor of Surgery. His accomplishments led to his being awarded a Research Fellowship at the University of Sydney and an annual research grant of five hundred pounds for the years 1938–1940. During this time he invented a technique of nystagmography using an electro-oculogram, and was astonished to find that, in nystagmus, the head and the eyes move in opposite directions. Archie’s interest in nerves and nerve functions had been stimulated by one of his teachers in the early years of his medical course, Martin Canny, a part-time lecturer who also worked at the Kanematsu Institute. At that time opportunities to engage in neurophysiological studies were decidedly limited. In his first major research project Archie had to adapt an ancient string galvanometer (Matthews oscillograph) to detect nerve action potentials in the 6th nerve. In another project, he had to utilise this primitive recording method in an attempt to measure contractions in uterine muscle, obtained during operations on patients at the Royal Hospital for Women (2). One of his first original scientific observations, published in the Journal of Physiology in 1939 (3) was on the quick component of nystagmus. At that time it remained unclear whether nystagmus arose as a result of proprioceptive signals from extrinsic eye muscles, or whether its control was entirely of central origin. Archie used cats as his experimental animal. After cutting the 3rd, 4th and 6th cranial nerves he found he was still able to record efferent activity from the central cut end of the sixth nerve — activity typical of that associated with nystagmus. The observation put beyond doubt the view that the nerve activity associated with nystagmus was entirely of central origin. While he was at Prince Alfred Hospital, Archie was befriended by Arthur Burkitt who had held the chair of anatomy at the University of Sydney since 1925, succeeding J.T. Wilson and J.I. Hunter in that position. The name J.T. Wilson was to come up again later in Archie’s life in association with his work on the platypus. Burkitt was a valuable ally since he had an extensive library that he made freely available for Archie’s use. Burkitt also arranged for Archie to give some of his first lectures. Archie had met his future wife Anne Williams while he was staging a play at his aunt Mollie’s. They became engaged in 1939 and married on 30 March 1940. Anne’s grandfather’s family had emigrated from England as free settlers in the convict days and the family were given land at Milson’s Point, Sydney. Anne has written that the children ‘were rowed across the harbour by a ticket-of-leave servant to Circular Quay and then they walked through the bush, where charcoal burners worked, to Macquarie Street to school’. Her mother’s family were missionaries by the name of Blomfield. Anne’s parents and the David family were friends, and when Archie was studying at the University of Sydney and was living with his Aunt Mollie, Anne got to know him. In her words: ‘We seemed to get on well from the word go’. Anne’s father was a solicitor in a well- established law firm in Sydney. Although he lost his job during the 1930s depression, he insisted that she stay at school and complete her Leaving Certificate. Anne was interested in art and architecture. She enrolled at East Sydney Technical College in Paddington to study drawing and painting. She was granted one of two scholarships that provided free tuition for two years while she was taught life drawing and painting by Douglas Dundas. Although Anne had hoped to spend some time living with her artist aunt in Paris, the war was looming. When she and Archie married, they were fortunate that Archie’s grandmother allowed them to live very cheaply in her cottage in Hunters Hill. Michael and Margaret were born there during the war years. At the outbreak of the war, Archie, who was already in the militia, joined the Air Force. Because of the relationship of the work he had done already on the vestibular system to some of the physiological problems in aviation medicine, Archie became involved in a number of air combat research projects during the years 1941–1946. He developed a method for detecting air-sickness-prone pilots — previously a major cause of dropout from flight training. Subsequently, he worked with Frank Cotton on the production of a G-suit that would keep pilots from blacking out, by preventing blood from pooling in the veins of the lower body when the plane pulled out of a steep dive. Testing of the suits was done on a human centrifuge. Archie used himself as a guinea pig in this and a number of other research projects. Anne reports that ‘the primitive centrifuge at the University of Sydney would break down frequently’ which meant that, from time to time, Archie’s body was subjected to excessive gravitational strains. It meant that in later years he was to suffer from varicose veins. Eventually, towards the end of the war, the Australian suits were used in Spitfires in Darwin. Archie’s aviation medicine work was of such significance that he was sent to the USA to visit centrifuge laboratories there and to Britain where he worked in the Physiological Aviation Medicine Unit near Farnborough, run by Sir Bryan Matthews. Here he worked on the development and testing of an ejection seat, required by the need to bail out of the faster aircraft being brought into service. Some of this work was quite dangerous as it involved trials to discover the amount of explosive necessary for effective ejection without injury to the pilot’s spinal column. Some volunteers suffered major back injuries, but again Archie continued to be a test subject. He later declared that it was his view that he couldn’t ask others to do what he was not prepared to do himself. He made other important discoveries about depth perception — so important for pilots during landing — and demonstrated that this was not correlated with visual acuity. Archie was demobilised in 1946 having reached the rank of squadron leader and, as he put it, having no squadron to lead. After the war Archie was offered the opportunity to join Roy (Pansy) Wright in the Physiology Department at the University of Melbourne. However, he decided that he really wanted to go overseas for further postgraduate experience. He was awarded a Rockefeller Fellowship and moved with his family to New York and the Rockefeller Institute where he worked during 1946–1948 in the company of major figures in neuroscience including Herbert Gasser, Lorente de No, Birdsey Renshaw and David Lloyd. His work with David Lloyd, who had been a Rhodes Scholar in Charles Sherrington’s laboratory in Oxford, provided Archie with the foundations for his future work in experimental neurophysiology. Lloyd had one of the most modern laboratories, equipped with the latest valve-operated electronics. Together, Lloyd and McIntyre studied long-spinal reflexes, the origin of dorsal root potentials and the central projection pathway for Group I afferents in peripheral nerves. Lloyd himself was a shy, sensitive person who prided himself on his dissection skills and on the care he took in assembling data before coming to any firm conclusion. Archie was deeply fond of Lloyd and in later years adopted some of Lloyd’s style, his skill in dissection and his care in designing experiments. Family Life on a Rockefeller Scholarship Anne McIntyre reports that the worst time in her life was undoubtedly the period of two years spent in New York while Archie worked at the Rockefeller Institute. ‘We knew nobody and the value of money changed overnight while we were paying $120 a month for a forty dollar apartment. We somehow managed to feed the kids. Archie and the other scholarship holders were given a square meal at the Institute at mid-day. My normal weight was nine and a half stone and when we left after two years I weighed seven stone’. Towards the end of his time in New York, Archie was awarded a Nuffield Scholarship to work in Cambridge, England, again with Bryan Matthews. In Cambridge Archie borrowed a bicycle from Alan Hodgkin to travel to the laboratory every day. He had to build his own equipment, a muscle-stretching device for frog muscle. He received a lot of help from Bryan Matthews, the Head of Department, and always retained a sense of gratitude towards him. While in Cambridge, Archie made frequent visits to London to see his friend Bernard Katz, whom he had met in Australia when Katz was working at the Kanematsu Institute with Eccles and Kuffler on neuromuscular transmission. They had become friends and when Bernard married Rita Penly shortly after the war, Archie was best man. Katz returned to London in 1946 where he began a series of experiments that laid the basis for our present-day understanding of synaptic transmission and which brought him the Nobel Prize in 1970. An important and revolutionary feature of Katz’s work in the post-war period was his use of microelectrodes to analyse details of the transmembrane events in neuromuscular transmission. Archie was deeply impressed with this technique and developed the idea that a similar approach could be used to analyse synaptic events within the central nervous system. Here he was combining his experience of whole- nerve recordings of reflex events acquired at the Rockefeller Institute under David Lloyd with Katz’s novel approach at the single-cell level. While in Cambridge, Archie received an offer from Jack Eccles to take up a senior lectureship in Eccles’ department in Dunedin, New Zealand. Archie felt that he had an obligation to go back to Sydney where Burkitt had promised him there would be a job — but not a senior lectureship. After some hesitation he opted for Dunedin where he had a definite offer. The vacancy arose because of the departure from Dunedin of Victor MacFarlane who had left to take up the Chair of Physiology at the University of Queensland. Archie had met Eccles some years previously while Eccles was still at the Kanematsu Institute. Eccles was interested in McIntyre because he knew Archie had worked with Lloyd, one of Eccles’ main competitors in the field of central synaptic action. In one of Lloyd’s exceedingly careful and painstaking studies, he had measured the latency of ‘direct’ inhibition of motoneurons in the spinal cord and found it to be the same as the latency for monosynaptic excitation. He therefore concluded that ‘direct’ inhibition was also monosynaptic. Eccles subsequently demonstrated the disynaptic tempo of ‘direct’ inhibition by means of microelectrode recordings. It turned out that because the excitatory post-synaptic potential had a finite rise time, an inhibitory potential of longer (disynaptic) latency could still block the excitation. Following this debate, Lloyd abandoned the subject of neurophysiology and began to work on sweat glands. When Archie arrived in Dunedin he assembled an electrophysiology recording set-up of the kind he had used in New York and began to explore the technique of pushing microelectrodes into the spinal cord of anaesthetised animals as a means of recording activity in central neurons, at the single-cell level. At the time, Patton and Woodbury, K. Frank and others in the USA were attempting similar recordings. Initially Eccles did not show much interest in what McIntyre was trying to do. He soon recognised the importance of this approach, however, and began to use Archie’s equipment in experiments on motoneurons with Jack Coombs and Lawrence Brock that would eventually bring him the Nobel Prize. During their time in Dunedin, Eccles and McIntyre published papers together on plasticity of the central nervous system (18) and chromatolysis in motoneurons (19). In these experiments Archie did most of the dissections because of the skill he had acquired under Lloyd. In writing up the experiments, Archie recalls, it was characteristic of Eccles to want to speculate further than, Archie thought, the evidence allowed, and the speculation was often declared as a firm conclusion. The move to Dunedin was a very happy change for the family. Anne was able to do some painting though she felt that most of the New Zealanders who shared her interests were based in Auckland. Splendid food was now available. The McIntyres became firm friends with the Mantons (subsequently Guy Manton became Dean of Arts at Monash) and the Thomsons, owners of Earnslaw Station. The two academic families camped on the station land during their first summer holiday in New Zealand. Later they built a hut below the mountains on the edge of a small lake, Lake Wakatipu, on the edge of the Thomsons’ property. The lake’s water came from the Earnslaw glacier. This hut and the summer holidays spent there left a lasting impression on all members of the family. The children felt themselves to be New Zealanders. Archie was able to get ‘away from the University’ and his family recall ‘that was the time when Dad played with us, taught us to fish, taught us to make things like a small, light boat, to find good stones for building and so on’. The company of a professor of classics and his family on these trips brought a particularly scholarly tone to their holidays. Anne recalls Guy Manton, on a small bridge over a rivulet, telling the children the story of Horatio defending the bridge to Rome. The McIntyres helped the Thomsons bring in the hay and were occasionally permitted to drive the small truck in low gear while others tossed the bales into the back. Otago Medical School — Dunedin With the appointment of Eccles to the chair of physiology in the John Curtin School of Medical Research in 1951, Archie became acting head of department and then was appointed professor of physiology at the University of Otago from 1952, a post he filled with great distinction for the next nine years. He built up a substantial research department and stimulated a large number of medical students to commence careers in physiological research. He passed on his enthusiasm for research as well as the attention to detail and dissection skills that were needed in order to ensure a successful outcome. Some of the most successful of Archie’s students were Richard Mark, Ian McDonald, Julian Jack, John Steiner, Colwyn Trevarthen, Ainslie Iggo, John Ludbrook, John Hubbard and Austin Doyle. Archie attracted a zoologist, Geoff Satchell, to his department, and John Veale, who had double degrees in physics and medicine, joined him from Auckland. Archie’s nine years in Dunedin were some of the happiest years of his career. Here are some fragments of reminiscences by some of Archie’s students in Dunedin. One of John Ludbrook’s recollections is of when he was an undergraduate in Dunedin. In third year Medicine, students were required to take a trip in an ex-RAF high altitude chamber. Archie sat in it (with an oxygen mask on) while the eight students ascended to around 20,000 feet. As each student collapsed in sequence, Archie would attach an oxygen mask to him/her. John dimly remembers noticing that his fingernails became blue at around 16,000 feet, just before he blacked out. Those were the heroic days of physiology! Richard Mark was an editor of the medical students’ annual at that time. He recollects a brief verse describing Archie:
Julian Jack in a piece published in the Otago Daily Times on 7 September 2002 described McIntyre as ‘very gentle and relatively permissive as a supervisor, but without making one feel he did not care’. Some doctoral supervisors, he noted, were often not so supportive, perhaps fearful of being overtaken by their ‘children’: ‘Archie was completely happy to be totally supportive and provided the kind of mentorship which resulted in many of his students going on to high positions — unfortunately, for New Zealand, often overseas’. What Jack recalled as a ‘piece of informed imagination’ led to work by an Otago contemporary, Ian McDonald, into trying to understand multiple sclerosis (MS), the disease where axons lose their myelin. ‘That research launched Ian on work which enabled him to show, for the first time, that these demyelinated axons don’t necessarily stop conducting impulses’. It was one of the major therapeutic hopes for treatment of MS and led to the realisation that it was not possible to relieve symptoms. In 1953 Archie gave a paper in Montreal on the chromatolysis work with Eccles. He used that occasion to spend a couple of months back in David Lloyd’s laboratory. Lloyd was working with Cuy Hunt at the time. The visit led to a collaboration between the three of them and publication in 1955 of a paper on the monosynaptic reflex (30). As a result of their work together, Hunt and McIntyre became friends and they remained close for the rest of Archie’s life. During his time at the Rockefeller, Archie had also become impressed by Herbert Gasser, the director of the Institute. Gasser used to wander into the laboratory rather casually and in the ensuing conversation often made, in passing, rather pertinent remarks about some problem or other. So it was with Archie — Gasser mentioned that little was known of the properties of the cutaneous sense organs supplied by the peripheral nerves that he, Gasser, had studied. This comment eventually led to three landmark papers on sensory receptors by Hunt and McIntyre (43, 44, 45). During 1959–1960 Archie took study leave, six months at University College, London, to visit Bernard Katz and then a second six months in Salt Lake City, Utah. In Utah, Archie and Cuy Hunt carried out the experiments that led to those three important papers. There he also met and became good friends with Ed Perl and Carlos Eyzaguirre. In 1961, Archie was approached to apply for the chair of physiology at the newly established Monash University in Melbourne. Because he wanted to return to Australia, he accepted the challenge of building up a new department from scratch. He made it one of his priorities to devote himself to the selection of staff who would generate an active programme of research. Some were old students of Archie’s. He brought with him from New Zealand members of staff like John Veale, as well as several technical assistants. Among his first Melbourne recruits were Mollie Holman, Geoff Bentley, Ian McCance and Ian R. McDonald. He brought Richard Mark back from California and recruited Laurie Geffen and Bob Porter from Oxford. His recollection of the early days at Monash was that everything seemed to change rapidly. He would plan buildings for a set number of students, then that number would be doubled. Because of his good judgement in the selection of staff, the department was soon a thriving place, establishing its reputation locally and overseas. Archie believed that the best approach to the teaching of neuroscience was an integrated one. However, many of his early attempts in this direction were frustrated by the lack of co-operation from other departments. Archie soon collected a group of local PhD students around him, much as he had done in Dunedin. This time, however, he was very busy establishing the department as well as contributing to the administration of Australian science, leaving him rather little time for student supervision. Nevertheless a number of successful candidatures were completed, including those of Pat Dorward, Uwe Proske, Poh- Tek Yeo, Ed Gregory, Paul Kenins and Paramsothy Subranarian. Uwe had completed an Honours year in Adelaide under Geoff Satchell. Geoff, a graduate of Leeds University had been working in the Zoology Department in Dunedin on sewage flies when he decided he really wanted to be a physiologist. Archie invited him across into Physiology as a lecturer and Geoff soon established himself as an authority on the cardiovascular system in sharks. When Archie moved to Monash, Geoff went to Adelaide. When Uwe professed his enthusiasm for neuroscience, Geoff recommended he transfer to Monash to do a PhD under Archie’s supervision. The McIntyres and the Satchells remained close friends. Archie and Anne were especially fond of Geoff’s wife Truda. For the first decade of its new life Archie’s department had to deal with both physiology and pharmacology. Archie strongly encouraged the development of BSc courses in both subjects. Staff members were required to develop modern courses and to design practical classes for both science and medical students. John Phillis, a neuropharmacologist, was one of the earliest appointments to the Department. He was energetic in his development of a BSc programme in physiology, including rather sophisticated experiments in practical classes involving whole animals, often sheep as well as cats. Other early staff appointments included Rod Westerman (1965) Gray Woolley (1965) and Colin Gibbs (1966). The staff were ably assisted by Jeff Robinson the laboratory manager. From the start Archie insisted that the Department incorporate an animal house and a mechanical workshop. Archie was innovative in his approach to teaching and utilised the available technologies of video recording, primitive as they were at the time, to support didactic teaching. Who amongst the medical students of those days could have failed to be impressed by videotaped recordings of classical neurophysiological experiments being performed by the Professor in a white coat and a bow-tie? He emphasised the importance of the practical class in physiological education and insisted that students come to appreciate the experimental evidence on which knowledge is based. He felt it was especially important to have some understanding of the history of science ‘to appreciate that in our current efforts we are standing on the shoulders of others’. In every aspect of the work of his Department and the Faculty he was personally committed and involved. Using the same approach as he had used when he subjected himself to rides on the human centrifuge, he provided leadership by being the most actively involved member of the team. That he succeeded, in a very few years, in developing one of the most highly regarded departments of physiology in the world is testimony to the effectiveness of his approach and to the wisdom with which he managed his responsibilities as a Head of Department. Archie resigned from the headship in 1974, four years before his retirement. Scientific Accomplishments in Neurophysiology Archie’s grounding in classical Sherringtonian methods for the study of the nervous system came from his experiences in David Lloyd’s laboratory. Here he utilised preparations like the decapitate cat to examine conduction in spinal cord pathways and the co-ordination of hindlimb and forelimb reflexes. In addition, he studied conduction in afferent pathways like the dorsal columns when these pathways were activated by stimulation of peripheral nerves. Once he had set up his recording apparatus in Dunedin, he was able to examine motoneurons directly and he commenced studies on plasticity (use and disuse) and on chromatolysis (degeneration), resulting from section of motor nerves. This was work that was later taken much further by Eccles and his colleagues. His seminal paper (with Lawrence Brock) on the responses of motoneurons to stimulation by microelectrodes was published in the Proceedings of the University of Otago Medical School in 1953 (27) and, in the same year, in the same journal, he published a paper on cortical projections of afferent impulses in muscle nerves (26), opening up a field in which he retained an interest throughout the rest of his life and which continues to challenge, even today, those involved in the cortical control of muscle contraction. The landmark papers by Hunt and McIntyre (43, 44, 45), based on the prompting of Herbert Gasser, identified the properties of cutaneous receptors in the cat. The approach taken here is reminiscent of a similar technique employed by Gasser himself. The preparation, the anaesthetized cat, was familiar to both Hunt and McIntyre. The technique of dissecting fragments of dorsal roots had been established earlier by Hunt (Kuffler, Hunt & Quillian, 1951). In this series of experiments Hunt and McIntyre systematically measured the conduction velocities of all myelinated afferents arising from receptors in the skin of the ankle, the interosseous membrane and the flexor digitorum longus muscle. Receptor properties were matched with axonal conduction velocities which were then converted to estimates of axon diameters. These were compared with histological nerve profiles, as Hunt had done earlier in 1954. At the time, these studies represented the most comprehensive and systematic analysis of cutaneous receptors ever carried out. They opened up a whole new field of study. One subsequent series of experiments that deserves particular mention was Archie’s work on the cortical projection of Pacinian afferents, published jointly with Mollie Holman and John Veale in 1967 (62). Archie already knew that there was a strong projection pathway to the cortex by Pacinian afferents. He and Cuy Hunt had learned earlier to expose the interosseous membrane of the cat hind limb and the associated population of Pacinian corpuscles which could be individually identified under the dissecting microscope. He managed to stimulate a single corpuscle with a fine glass stylus, monitor its action potential in the tiny interosseous nerve and record the cortical evoked potential from that one impulse. It is generally agreed that a cortical representation is a necessary prerequisite for conscious sensation. It was already known at that time that stimulation of Pacinian corpuscles produced distinct sensations of touch/vibration. Archie’s work demonstrated that one impulse in a single afferent fibre was sufficient to engage the machinery for cortical representation and therefore most probably for conscious sensation. It supported the view that whatever cortical circuits were involved in generating sensations, they were not likely to be very elaborate or widely distributed. Archie’s style in scientific research was essentially curiosity-based, but unlike most people, he wanted to know more about things in depth. A common theme that recurred again and again in his writings was his fascination with and awe of the brain. Here his strong sense of humility emerged. He kept emphasising that despite the growing sophistication of our knowledge, we were still only scratching the surface. He abhorred the idea that we already knew all that there was to be known about a particular topic. He shunned scientific dogmatism and complacency. Because of his curiosity and his intellectual fascination with mechanism, Archie’s interests knew few boundaries. Certainly he wasn’t interested in anything predictable and boring. Because of his breadth of interest and because some of his students at Monash were trained zoologists, he embarked on studies in comparative neurophysiology, always leaning on techniques and methods of analysis that he had acquired in previous cat experiments. Here it was Archie’s view that what was a complex and sophisticated situation in higher mammals was often represented in a simplified and more readily studied form in the lower vertebrates. This led Pat Dorward and Ed Gregory to study the mysterious Grandry and Herbst corpuscles in birds. Uwe Proske studied muscle spindles in reptiles where each receptor has only a single intrafusal fibre compared with the dozen or more fibres in mammalian spindles. The reflex action of these spindles was subsequently pursued by Paul Kenins. Archie had a real interest in psychophysics, in particular in the proprioceptive senses. During the 1960s there was an ongoing controversy over how joint position and movement were signalled. It was known that blindfolded human subjects could unhesitatingly touch the tips of their noses, indicative of a precise sense of limb position and body image. Work on the cat had suggested that joint receptors were primarily responsible for this. The data all derived from studies on the medial articular nerve of the cat’s knee (Skoglund, 1973). Archie knew that this nerve was not a pure joint nerve but was sometimes contaminated with afferents from the popliteus muscle. This led to a study with David Tracey and Uwe Proske in which he demonstrated, unambiguously, that the so- called mid-range joint afferents were in fact muscle afferents. This conclusion provided supporting evidence for the view that muscle receptors were primarily responsible for position sense. Archie was particularly interested in Australian native animals, especially the platypus and the echidna. As a keen trout fisherman he had often seen platypus as he waded up a stream. His interests in these animals led to a study in 1985 with his ex- students Ainsley Iggo and Uwe Proske on the sensory innervation of skin of the echidna snout (99). At this time Archie was already officially retired and living in Launceston, Tasmania. He would come back to Melbourne for a few weeks at a time to carry out these collaborative efforts. Another memorable series of experiments carried out at this time with John Rawson and Uwe Proske used a technique of raising the electrical thresholds of muscle spindle afferents by vibration (established by another student, Julian Jack) to show, for the first time, that tendon organ afferents projected to the cerebral cortex (100). In 1986 the story broke of an electric sense in the platypus. Immediately Archie asked, ‘but what are the receptors involved?’. Answering that question led to yet another collaboration with Ed Gregory, Ainsley Iggo and Uwe Proske. It produced three major papers on electroreception in the platypus (101, 104, 109) and a further paper demonstrating the existence of an electric sense in the echidna (113). Archie McIntyre was elected to Fellowship of the Australian Academy of Science in 1963. He was a member of the Council of the Academy during 1968–1974, being Secretary, Biological Sciences during 1970–1974. He had been a founding member of the Australian Physiological and Pharmacological Society (APPS) and he was one of the driving forces behind the establishment of the Australian Neuroscience Society. In recognition of Archie’s contributions to APPS, the society set up the A.K. McIntyre Prize in 1994 for members of APPS who have made significant contributions in their pre-doctoral and immediate post-doctoral years. The prize has been awarded each year since then. Archie was very active in the promotion of physiology and neuroscience throughout Australia. He served on ANZAAS committees and was a member of both the ARGC and NH&MRC research support agencies. He served on the Program Committee of IUPS and was Chair of the National Committee for Physiological Sciences. Archie retired in 1978 and he and Anne moved to Launceston, Tasmania where Anne designed and supervised the construction of their home, ‘Montacute’, on the outskirts of the town, overlooking the valley of the river Esk, with the slopes of the Ben Lomond visible in the distance. Here they were close to Archie’s brother and sister and other members of family. Nevertheless, Archie continued to remain in touch with his scientific interests and, until about 1990, regularly visited colleagues at Monash to engage in further experiments on sense organs — some of them involving through-the-night recordings, exploring the electric sense organs of the platypus. In retirement Archie was able to plant a vineyard with about fifty vines and he became seriously involved in wine- making. His early interest in chemistry was reflected in his establishment of a fully- functional oenological analysis laboratory. The end product was of high quality and enjoyed by all visitors to Montacute. As Archie’s health began to deteriorate, he and Anne found it necessary to leave ‘Montacute’ and move into a retirement village close to the centre of Launceston. Archie is survived by his wife Anne and his three children, Michael, Margaret and Richard. A fragment of poetry written by Archie sometime in the later years of his life:
List of Awards and Affiliations
The authors are grateful to Mrs Anne McIntyre for her helpful advice and for the comments she provided in conversations and in letters. We have had access to the transcripts of tape recordings of interviews conducted by Archie’s sister in November 1996 and March 2001 in Launceston. There are also transcripts of conversations between Archie and Uwe Proske recorded on 2 and 3 June 1995 in Launceston. These transcripts are all held by the Australian Academy of Science. References
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R. Porter, Faculty of Medicine, Health & Molecular Sciences, James Cook University, Townsville. U. Proske, Department of Physiology, Monash University, Melbourne. R. F. Mark, Research School of Biological Sciences, Australian National University. (Richard Mark died during the period of preparation of this memoir. He did see a preliminary draft and was in general agreement with its contents.)
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