AAS-Biographical memoirs-Macfarlane

With the death of Victor Macfarlane on 26 February 1982, Australia and New Zealand lost one of the most oustanding figures to have appeared in the Australasian scholastic world . Macfarlane' s remarkable reputation stemmed not only from his exceptional intelligence, originality of thought and the high quality of his many research activities, but particularly and uniquely because of the extraordinary breadth of his scientific achievements and cultural interests, ranging from studies of parasitic trematodes through clinical medicine and surgery, neurophysiology, adaptation to desert and arctic environments, ecophysiology of animals, including mankind, and nutrition, especially salt and water metabolism, to social interactions and the genesis of customs and rituals. Nor were these many research ventures trivial: whatever problems he chose to study, his approach was innovative and direct, and most often required a minimum of complex and sophisticated equipment. The task of doing justice to this polymath in writing about his career is indeed formidable.

Early days in New Zealand

Walter Victor Macfarlane was born in Christchurch, New Zealand, on 27 September 1913. His parents, Ada Constance Westerman and Walter Macfarlane, were first-generation New Zealanders, their families having migrated from Britain in the early days of settlement. Ada Westerman's parents came from Yorkshire, Walter Macfarlane's from Scotland. Victor was the first child of the marriage; two other children, both girls, made their appearance later. The first of these sisters, born about two years after Victor, died at the age of four from complications arising from pertussis (whooping cough), a notorious child-killer in those pre-antibiotic times. This tragedy must have been a shattering experience to the family. Victor's surviving sister, nine years his junior, recalls that Victor never spoke about the matter or even mentioned the younger sister, although from family photographs it was clear that he and this little girl had been devoted to one another.

The family lived in the beautiful Cashmere hills north of Christchurch, on the volcanic slopes of the Banks peninsula. Their house was high up, close to the upper steep, open, largely tussock-covered slopes. Looking south and west, the city of Christchurch could be seen below, and far to the west beyond the Canterbury plains was the magnificent backdrop of the Southern Alps. The Macfarlane family revelled in this clear, airy and open environment, and it would be difficult to imagine a better place in which to grow up. It must have played a major role in shaping the young Victor's developing interests and aspirations. At least of equal importance, however, was the domestic environment and the character and attitudes of the parents. According to Victor's sister Yvonne, their parents were not highly educated, but they both had great energy and vitality, and had wide interests. Their mother was well read in classical literature and poetry and had a remarkable memory, often quoting at length from her favourite poets . Their father, a successful builder, was quiet, strong and tireless. He was a fine husband and father, and he and his family were very popular in the district. The harmonious and tolerant home atmosphere meant that the children enjoyed a great deal of freedom within and outside the home. In addition to his own household obligations, Walter Macfarlane settled his widowed mother and two unmarried sisters on a farmlet of three and a half acres at the foot of the Cashmere hills, where they kept a cow and chickens, grew fruit of various kinds and quantities of vegetables and flowers.

Victor's sister writes: 'The big house and its produce was a Mecca for many during the depression. We adored this farmlet and the happy people there – not to mention the ever-present goodies to eat.'

From an early age, Victor was able to go, sometimes alone, sometimes with company, on extensive explorations of the adjoining steep tussocky hills and patches of native bush. Highly intelligent, with a lively curiosity, he loved these wanderings and observations of plant and animal life, wind and weather. His home and surroundings favoured this early development as a budding naturalist. Victor also became an avid reader and, although there is no record of when he began to read, there can be little doubt that this was early and that his mother played a big part in helping him to acquire this skill. But he was no self-effacing bookworm, and his life-long predilection for conversation on every conceivable topic developed early. These often lengthy conversations, especially during meals, are remembered by his sister as lively and provocative, and well sprinkled with puns. One topic brought up a number of times was comparative religion.

Although not narrowly religious, the Macfarlane family belonged to the Methodist Church, which was fortunate to have then amongst its New Zealand clergy a number of spritely, intelligent young men with a remarkably ecumenical outlook for the times. Amongst those attending the Cashmere Methodist Church was a group of lively and intelligent people who were the basis of much of the Macfarlane family's social life. One of these friends living nearby was Hugh Parton, a member of the staff of Canterbury University College's Department of Chemistry and later professor of chemistry at the University of Otago in Dunedin. Parton took a particular interest in the young Victor, who attended his Bible class at the church, and the pattern and direction of Victor's career was strongly influenced by this association.

Victor attended Cashmere Primary School and, from 1928 to 1931, Christchurch Boys' High School. His performance, though not spectacular, was sound and led to several prizes for general excellence. English, History, Geography and Chemistry were his main subjects. He showed a clear preference for the humanities, the subjects he intended to pursue at tertiary level being English and History, and this perhaps is not surprising in view of his extensive reading and love of language. That he chose to include Chemistry when enrolling for his BA at Canterbury University College in 1932 can be attributed to the influence of two people, his bible class convenor Hugh Parton and John Kidson, science teacher at Christchurch Boys' High School and also a member of the Cashmere Methodist Church. Little or no Biology was available in most boys' schools at the time.

Hugh Parton has provided an interesting sidelight on Victor's rather unusual manner of speech and accent. Everyone who knew him was intrigued by his manner of talking and by his accent, very different from that of most New Zealanders. He spoke in a very measured but sometimes hesitant way, with periodical pauses and occasional asides. Parton is convinced that these characteristics arose from elocution lessons that he received from the author and dramatist, Ngaio Marsh (later Dame Ngaio), who lived nearby in the Cashmere hills.

Victor's main outdoor recreation continued to be ever-wider exploration by bicycle and on foot of the surrounding country. These expeditions, mainly during holidays, often with a friend but sometimes alone, extended beyond the Banks peninsula and took him to the river-valleys, foothills and passes of the Southern Alps. He also engaged in the more usual sports and hobbies of school-age boys, including the making and flying of model aeroplanes, rugby football and rifle-shooting. He became a crack shot, and was a member of the school's Snowden Trophy shooting team. His passion for reading continued; his sister recalls that his room always seemed full of books. She also writes: 'I remember Victor in his teens sitting down for three days and reading the Bible from cover to cover!' At high school, he served very successfully as librarian for a year. At times the demands of his active physical life temporarily dampened his usual conversational sparkle. Like his father, he was physically strong and had great endurance. Each school day he bicycled to high school, five miles each way and with a stiff climb at the end of the day, at times in the teeth of a howling northwest wind. His sister writes that sometimes after rugby, 'a bruised "hooker" used to push his bicycle up the hill and fall into a bath for ages. On these occasions his usual lively meal conversations became monosyllabic.'

Victor's own words (1) summarise succintly this early phase of his life:

There were advantages to growing up in New Zealand. We lived on the andesitic lava flows that to James Cook had been Banks' Island. There was a great high playground of Poa tussock for sledging, and tangled native forest on the steep crater slopes for exploration. Across the plain we watched the southern Alps change from summer scree to winter snow. As an adolescent I began to explore them, often using a bicycle to get to the foothills. The rivers ran clear and fast and led to the Notofagus forests and these in turn, over the untracked moss, to gorges and the passes of the mountains. Then there were glaciers and snow peaks to climb. People were and still are the rarest fauna of the region. There could be few better places to grow up.

Civilisation from Europe came essentially in books. These I read avidly. In the routines of school, english, chemistry, geography and history provided the main interest and at the age of 13, 15 and 17 prizes for general excellence emerged. But there was no biological science in school work. I spent a vacation reading all the plays and prefaces of G.B. Shaw and found a framework for social understanding and criticism. Then the volumes of Wells, Huxley and Wells' Science of Life, consumed also in a vacation, was a new book of revelations. At 16 this provided a sweep of imaginative biology from the pre-Cambrian to human psychology. In spite of an invitation to return to school as head prefect I decided to go on to the University, Canterbury College in Christchurch.

Canterbury University College

In 1932, Victor Macfarlane enrolled in the Faculty of Arts, his first year subjects being English, French, History and Chemistry. His intention was to proceed to his MA degree in History. Because of his growing interest, through his own reading and the influence of Hugh Parton and John Kidson, in the broad sweep of science, he was considering, as the topic of his master's thesis, the history of science in New Zealand. However, these plans changed during the course of his first year as a result of the wider contacts and challenges encountered in this new, more varied, and liberal educational environment. Canterbury University College, with only about a thousand students, must have been a stimulating place for any bright young person fortunate enough to be able to study there despite the economic depression. Although staffing was meagre, the college was fortunate in having some excellent scholars and teachers in various departments, and the general atmosphere must have encouraged enquiry and learning. To young Macfarlane, seething with ideas and an unquenchable thirst for knowledge and understanding, the experience opened whole new vistas. In his own words:

There were a thousand gowned students. Frederick Sinclair was Professor of English and he roused my latent interest in etymology. Some of us accepted his invitation to spend lunchtime construing Dante, which led us not only to poetry but also to appraisal of human behaviour in a framework which categorised malice and treachery as the most human and least worthy forms of conduct. This extracurricular activity had much more lasting influence than many thousands of the formal hours of work. Periodically I walked up the hill with Edward Percival, Professor of Zoology, and we discussed the world and the sciences. He suggested that I might like to sample zoology in his course on evolution held on Friday nights between 7 and 8 p.m. In the next year I enrolled in zoology and followed it through to Masters level. With Edward Percival we pursued and discussed everything from genetics to ecology.

Thus Victor's second year subjects were English II, History II, Chemistry II and Zoology I. On returning in 1934 from a year overseas, Hugh Parton was surprised to find Victor in the Chemistry III class. In that year Victor also completed Zoology II as well as History III, qualifying for his BA. This highly unusual course continued the following year with Zoology III as prerequisite for his MA in zoology, which he completed in 1936.

In those depression years, jobs were few, and in the Department of Zoology there were no laboratory assistants. Victor offered his services as unpaid assistant during his second year, and continued these duties for the next two years. His autobiographical notes relate that he was able 'to combine work in four subjects with the duties of lab boy. This was a good way to learn how a laboratory works and also to acquire techniques like section-cutting, the culture of protozoa, and embryology. It gave a view of science from the inside while I was still a student.' Much of the stimulus and intellectual excitement came from personal and informal contacts and discussions out of formal class hours, such as those already mentioned with Sinclair and Percival. Victor's notes describe another informal activity: 'Another educational institution not mentioned in the calendar was a seat outside the Chemistry Department, in the quadrangle. There we discussed concept, percept, epistemology, sociology and the nature of things in a moderately sophisticated fashion. Hugh Parton was very much involved with this side of scientific thinking.'

Before enrolling at college, Victor through his reading and rural excursions and observations had become interested in the interactions between living organisms and their surroundings. But it was Edward Percival who really introduced him to ecology, which became in the broadest sense the main theme underlying his thinking and research from then on. This introduction was by way of studying the food supply in trout streams, which demonstrated that limitations of mean size and numbers in the trout population were determined by the predation of anglers rather than shortage of food.

Victor chose as the topic for his Master's thesis the study of two trematode parasites of eels that he had found in an eel brought into the laboratory for class purposes. This research 'meant a year of nights wading black rivers, tending fish traps, catching and counting 60,000 snails, small fish and trematodes. From this emerged four trematode life cycles, two new genera and a picture of the ecological impact of environment on the numbers and dispersion of parasites amongst their hosts'. On some of these watery expeditions, he took with him as assistant his young sister Yvonne.

The thesis was accepted, and he was awarded the degree of MA, Class II, in Zoology. Hugh Parton recalls Edward Percival's annoyance with the British external examiner. Percival was convinced that his failure to award a First for this admirable piece of research was the result of Victor's justifiably critical treatment of a question on which the examiner had dogmatic views. Be that as it may, closer to home the quality of the work was recognised and led to an invitation from the Wallaceville Veterinary Research Laboratory to join them as parasitologist and, in particular, to study the intermediate host of the New Zealand liver-fluke. For this work he was to receive £50.

Parasitologist

Victor accepted this offer, and set to work with his usual vigour and dedication to solve this important practical problem, the first with which he was faced as a professional zoologist. The host had been believed to be potamopygus, a very common fresh-water snail, but no proof had been forthcoming. Victor's research involved extensive field-work, often under severe climatic conditions, for which the determination and physical endurance that he had already demonstrated were essential. The task is well described in his biographical notes:

In the heavily dissected volcanic ash country of Hawkes Bay I looked for the host, riding horses along narrow tracks on near-precipices in search of molluscs. After about three months, likely looking rediae and cercariae were found in a swamp on the MacKinnon Station. For laboratory there was a table on the verandah of the great homestead and I saw the likely cercariae with the setting sun to illuminate the microscope. The peculiar pleasure of that sort of discovery was increased by talking of it over dinner at the vast polished table. Eighty years earlier this had seated 60 guests on their way north in convoy during the Maori wars. Now the table had only three occupants, but it retained the splendour of the patriarchal days.

The tiny fasciola larvae he had found were transmitted to the Wallaceville laboratory in small paraffin containers and there fed to rabbits. After two months, small liver-flukes were found in their livers and bile ducts, proving that the snail Limnaea truncatula was indeed the long-sought molluscan host in this district. Later, further search in other areas revealed that two other kinds of snail could also act as hosts for fasciola, one in the Buller region and one in Central Otago. The time spent in searching the Buller area included a week of black frost. 'The temperature did not rise above the freezing point of water at any time of the day or night and the mist froze continuously on the trees meshing them in ice. The other host was in Central Otago, where at least the ice conditions permitted some skating.' Thus the young parasitologist had solved his first problem, identifying the true mollusc hosts of the sheep liver-fluke and showing the earlier belief to be incorrect.

Subsequent activities included work in the Marlborough district on 'blowfly strike' in sheep that involved the setting and tending of many fly-traps in the hills and the collection of millions of flies. He devised experiments to determine the degree of fly-attractance of wool in different states of wetness from rain or urine contamination. This was done by setting out what Victor called a cafeteria of different samples of wool and counting the egg-deposition. Wet tip-wool proved to have the greatest fly-attractance. Similar tests were also carried out with larval nematode worms. Somehow during this busy couple of years Victor managed to fit in physics lessons with Frederick White, professor of physics at Canterbury University College (later Sir Frederick White, head of CSIRO). Victor performed the practical work alone in the laboratory on Saturdays. During his time attached to the Wallaceville laboratory, he became increasingly aware of gaps in his basic science training, especially in the rapidly growing fields of physiology and biochemistry. Furthermore, according to Hugh Parton, he also felt that the prospects for a graduate in classical zoology were not promising in a veterinary establishment. His father agreed to support him in obtaining the appropriate professional training, and suggested a veterinary school in Australia. However, Victor chose instead to tackle medicine in New Zealand's only medical school, at the University of Otago in Dunedin, the only establishment in the country then offering courses in physiology and biochemistry.

Dunedin and the University of Otago

To embark as a mature student on a course so exacting and prolonged as medicine requires exceptional determination as well as academic ability, attributes with which Victor had already shown he was amply endowed. The small city of Dunedin, founded by Scottish immigrants, was and is dominated by its university; indeed, the University of Otago was the first to be established in the whole country. Victor must once again have enjoyed working in an environment permeated with learning and enquiry, even though the excessive minutiae to be memorised in some subjects, especially gross human anatomy, can scarcely have been intellectually rewarding. But he endured it, even noting: 'The drilling in anatomy from William Gowland would have made FRCS candidates of us all, but the histology and embryology was not without its later use.' John Malcolm, nearing retirement, taught physiology well, with a nutritional bias; and Norman Edson very ably expounded to the class the biochemistry of the times. Nor did the winter climate of Dunedin, which has been euphemistically described as 'bracing', deter Victor's enthusiasm, despite living on short commons in 'digs' that were hardly luxurious. Victor characteristically wrote: 'The wet climate described in Hodge's The Wind and the Rain was useful encouragement to work, even at the level of working out problems in an unheated room at 2 or 3 o'clock in the morning with ice on the windows.' The physical problems of climate and accommodation apart, Victor, with his widespread interests, soon settled into the university community and became friendly not only with many students in medicine and other faculties, but also with staff, most of whom soon came to recognise and respect his remarkable talents. The city was well endowed with libraries, having a good lending library as well as those in the university. The medical school had its own library containing not only good runs of the major journals and a good supply of textbooks, but also a most valuable historical section obtained through a legacy, the Monro collection from Edinburgh of old books mainly on anatomy and physiology. Victor wrote:

There were 300 leather-bound volumes which recorded the thinking of Scarpa, Willis, Morgagni, Malpighi, Harvey, Sydenham, Addison and many others. As president of the Medical History Society, I had access to the books and enjoyed devising displays of their more pertinent contents. This reinforced the pleasures I had had in Canterbury of exploring Leeuwenhoek and reading the journals of explorers and biologists with Cook, d'Urville, Bellingshausen, Wilkes, or Vancouver who had passed the region up to 150 years previously.

Victor continued to read widely in all his subjects, in addition to the standard set texts and other material, and soon this was well known by the staff. It was usual for Horace Smirk, professor of medicine, to greet him in tutorials, clinics or oral exams with the words: 'Well, Macfarlane, what have you got to tell us today?' But his activities were by no means confined to medical academic study and history. He was able to draw very competently – his sister reports that 'his zoology illustrations were immaculate and beautiful to behold' – and in Dunedin he continued his interest in the visual arts. Doubtless this helped his friendship with Pamela Sinclair, a zoology student with outstanding artistic talent. He also enjoyed music, but was no performer and was thought to be tone-deaf. One new friend who opened up for him a wider musical world was Walter Griesbach, a physician who had been able to leave Hitler's Germany and settle in New Zealand. He gave lectures on endocrinology in the physiology course, as well as carrying out innovative research on pituitary-thyroid relationships. Griesbach was himself a competent pianist. Victor's notes relate:

I came to know him much better as his classic investigations of pituitary-thyroid cytology developed. He was a wise physician, a logically imaginative investigator and through him I came to know Brahms, Bach, Beethoven and Scarlatti much better.

Apart from social occasions with students or staff, such as his visits to Griesbach, Victor was also involved in other non-academic activities during the busy Dunedin days. These included taking up fencing, drilling with the Otago University Medical Corps, and producing several plays including a Greek tragedy.

The head of the medical school's Department of Public Health and Bacteriology was Charles (later Sir Charles) Hercus. He had carried out important pioneer work on goitre, a common condition in mountainous regions through iodine deficiency, and had been instrumental in largely preventing goitre by the use of iodised salt. One feature of his course was a requirement that each student undertake a study, out of formal class hours, of some problem of public health significance, and present a minor thesis on his or her work. Victor found a problem well suited to his earlier experience with parasites and schistosomes: to track down the cause of 'swimmer's itch' that affected many people after swimming in the country's many lakes. It seemed likely that the itchy rash might be an allergic reaction to some agent or organism in the water, and possible candidates included larval schistosomes. The work was done during vacations when Victor worked on a large property adjoining Lake Wanaka. Victor's notes vividly describe his activities:

I became fourth rouseabout on Wanaka Station working from 7am until 6pm for six days for £2 a week, mutton and potatoes and a sack bed in the shed. After work and on the seventh day I sought the nature of the itch and sat in the lake all day waiting to be attacked. Nothing happened. I set up a microscope in the undertaker's shed near the lake. There was a kerosene lamp for microscope work. I set it up on the carpenter's bench, sat on an upturned boat and was surrounded by empty coffins. Then one Sunday in some snails dredged from the lake I found fork-tailed cercariae and applied them to my arm. There was no effect. The coffin-maker's son, however, came by and some of the cercariae were applied to his skin. He complained of itching sensations and within 4 hours there were papules, forming the characteristic itching rash of swimmers in the lake. Ten days later I applied more of these organisms to my own skin and after an initial penetration itch, papules appeared with a 24-hour latency. I put some more organisms on my thigh and at 24 hours after invasion persuaded Archie Douglas, the general practitioner of the village, to undertake a biopsy. He took epidermis and dermis containing the papules. The scar is still present. But it took more than three months hunting serial sections of the tissue to find the lesion. Finally I realised that a small hole with debris in it was the invasion track, but the organisms had already been dissolved by 26 hours. Next season I made antigens from the snails, the cercariae and their secretions. Injections into exposed subjects showed that antibodies were present in all components. The cercarial body was the main antigen, and 10 days elapsed between the initial invasion and the build-up of adequate antibodies. This work on immunopathology was aided by the presence of a girls' fitness camp which yielded a supply of under-occupied girls, who obligingly and rather surprisingly allowed the larvae to invade the inner sides of their arms. From these I was able to take biopsies at timed intervals to follow the whole sequence of non-immune and immune responses to the organisms.

The phenomenon of habituation impressed him during this study. The glacier-fed lake's temperature stays much the same throughout the year (10-12°C), and initial immersion was painful and short. But after two weeks of daily immersion he was able to stay in the water all day without great discomfort and without shivering.

Late in 1944, Victor sat his final examinations. Not surprisingly he topped his year and won a number of awards – the McCallum, Colquhoun and Clinical Medicine medals and the Travelling Scholarship in Medicine. The degrees of MB, ChB were conferred early in 1945. For the first six months of that year he was the only resident house surgeon in the orthopedics unit of the Dunedin Hospital adjacent to the medical school buildings. He assisted with long operating lists and set innumerable fractures himself. After this introduction to the duties of an RMO, he became assistant to Murray Falconer in a newly established neurosurgical unit. Falconer was highly skilled in the special techniques necessary for exposure of the brain, spinal cord and nerve roots, and in the delicate procedures to be carried out on the nervous tissue. He was also completely dedicated to his craft and the welfare of his patients. Victor wrote:

There was little differentiation of night from day. Patients were studied in detail and the level of observation and logic applied in neurological studies were the highest I have encountered in biological work. Surgical craft was excellent and I enjoyed the skills acquired.

He and his chief collaborated in trying new procedures in their spinal nerve-root patients, using electrical stimulation and recording to define more clearly the defects in nerve conduction.

But Victor, who was becoming increasingly interested in the mechanisms underlying nerve and brain function, felt frustrated by the lack of time for analysis of the experiments and the limitations imposed by work on human subjects. During his final year as a student, a dynamic successor to John Malcolm took up his duties as professor of physiology. This was John Carew Eccles, an Australian Rhodes scholar and student of and collaborator with Sherrington for 12 years. He arrived fresh from some remarkably fruitful years of research in Sydney in which, with Bernard Katz and Stephen Kuffler, outstanding fundamental advances had been made in the mechanisms of transmission between nerve fibres and muscle cells, and between different nerve cells. During his work with Falconer, Victor used to cross Great King Street when duties allowed to attend Eccles' neurophysiological seminars. These contacts led to an offer by Eccles of a senior lectureship, beginning in 1947, which Victor accepted 'with a view to gaining more understanding of brain function'. He and Eccles must often have discussed philosophy as well as neurophysiology, both being greatly interested in the unorthodox views about scientific method of Karl Popper, who had come to Canterbury University College in 1937 and who lectured on his views in Dunedin in 1945 shortly before returning to Europe.

In the department of physiology, Victor was soon busy. As well as a heavy teaching load, he was actively engaged in research, and so began to acquire Eccles' electrophysiological expertise. During that year, the department was host to a Guggenheim Fellow from Johns Hopkins University, Chandler Brooks, who came to learn basic neurophysiological techniques to use in his studies of hypothalamic function. As those who have worked with Eccles well know, his experiments often last well into the early hours of the morning. Sandwiched between a busy teaching day and a 9am lecture the following morning, their conduct required considerable powers of endurance, no new thing for Victor. He wrote:

Spinal cord neurones and neuromuscular junctions filled days, nights and weekends. The old sub-culture of neurophysiology which was just entering its exponential phase opened up a tremendously lively intellectual groundbase. Chandler Brooks came from Johns Hopkins to try to learn to record from hypothalamic neurones. He learned a good deal about spinal cord neurones instead.

Victor's principal research was on the basic mechanism of neuromuscular transmission, involving electrical recording of the end-plate potential under various controlled conditions including the presence of pharmacological agents. The results provided further confirmation that release of acetylcholine from the nerve terminal is responsible for the transmission. Victor did not take up his travelling scholarship because, as he said,' it was unlikely that I could have found more valid and exclusive experience abroad than was obtainable with Murray Falconer and John Eccles'. In 1948, however, the chair of physiology at the University of Queensland was advertised. Victor applied and, at the age of 35, was appointed.

Brisbane

Victor arrived in Brisbane in January 1949 and discovered that it was a sauna. After the bracing physical and intellectual environment of Dunedin, he suffered considerable thermal and cultural shock in this large, brash and rapidly growing city with its sub-tropical climate. It took a long time for him to acclimatise and adjust, but he eventually did so.

The best way of making use of the day was to keep standing and working at some quite routine manipulation during the heat of the day. The temptation to sit and write or think led to coma...In time, I learned to sleep when it came, and then, waking at 2 or 3am, to use the cool hours from that time till dawn for writing, thinking and marking exam papers.

At the time, the Sir William Macgregor School of Physiology was in an old building situated in the city near the Houses of Parliament. There were two floors and a basement. The staff consisted of four graduates, three technical assistants and several typists. The teaching load was daunting: eight courses, with 700-800 students, and almost no modern equipment. The exception was a climatically-controlled 'hot room' and an adjoining laboratory in the basement, with some equipment including a balance for weighing human subjects. This was a legacy from Victor's predecessor, D.H.K. Lee. The heavy teaching burden was not helped by the situation of the lecture theatre which adjoined a steep road with noisy traffic. Victor's biographical notes relate:

Trucks carrying tons of gravel from the river ground up the road past the lecture room, taking 3 minutes for the journey. Up to 5 or 6 trucks an hour would pass this way. In autumn and spring if the windows were closed, the heat of 100 or more students was insupportable so that the windows were usually open and sound had its toll on communication.

By this time Victor had become engaged to his artistic Dunedin friend, Pamela Sinclair (daughter of New Zealand lawyer, Guy Sinclair) who, having completed her zoology course, was now working in the Department of Zoology at the University of Western Australia. Late in 1949, Victor crossed the continent, and they were married in Perth on 12 December.

The late Horace ('Harry') Waring, professor of zoology at the University of Western Australia, wrote on the occasion of Victor's retirement in 1978:

My first glimpse was of the pirate who, in the guise of a visitor, stole from me the woman who was not only my baby sitter, the department's official deflator of pompous professors, but whose artistic skill was to adorn my major opus.

While in the west, Victor took the opportunity of exploring the Wagin lakes south of Perth to look for schistosomes, which he had already found in swamps off the Brisbane river and in the Murray. He found them and again tested them on himself. He showed that they evoked the typical skin rash, and that applying dimethylpthalate (DMP) to the skin before immersion prevented invasion. He also obtained another biopsy of one of his lesions.

The young professor returned with his wife to Brisbane and resumed his many duties, but now with the support of a vivacious and talented helpmate. During the ten years of his chairmanship of the University of Queensland's department of physiology, courses were upgraded, modern teaching equipment installed, new staff recruited and a number of lines of his own research pursued. In addition to all this, he coped with a heavy administrative load, encouraged and collaborated with staff in their research endeavours, and trained and supervised research students. His sister Yvonne writes: 'One of the nice things about him was that he encouraged – almost bullied – people to use their talents.' This characteristic of the young professor in his first headship post contributed substantially to his success in rejuvenating the department, as did his iron determination and quiet obstinacy, especially in dealing with university committees. As if all these activities were not enough, Victor also undertook many other duties outside the university sphere, including provision of an electroencephalograph service for patients in Brisbane, membership of the CSIRO Council, chairmanship of the Queensland State Committee of CSIRO, Queensland representative on the Nuffield selection committee, and council member of the Queensland Institute of Medical Research.

After leaving New Zealand, Victor submitted to the University of Otago a thesis for the degree of MD, on the results of experiments carried out in Eccles' department on neuromuscular transmission. In 1950, the thesis was accepted cum laude. He continued with some further work on skeletal muscle in Brisbane after acquiring some basic electrophysiological equipment, and later he extended this research to cardiac muscle. However, other major research interests were developing. The sudden exposure to Brisbane's summer climate and the presence of a controlled climatic room combined to direct his attention to problems of thermal regulation and adaptation by animals and humans to different environments, including the associated mechanisms of water and salt metabolism. From that time on this became the central theme of his thinking and experimental activities, albeit with many forays into other areas of inquiry. In the laboratory, he carried out experiments on heat stress in sheep and other animals, with particular reference to salt and water balance and the regulation of renal function, and to the effects on reproductive function. The problems of thermal stress in humans was, of course, very much in his mind through personal experience. The RAAF was experiencing difficulty in manning its base on Manus Island, 2° from the equator, where the extreme humidity as well as heat made conditions very trying. They flew Victor to Manus to advise them about ways of improving the situation. He also visited the Commonwealth Building Research Unit at Ryde, NSW, to give advice on similar problems. He wrote:

I learned a good deal about the diversity of human reactions to thermal environments in the summer excursions to Sydney...The typists and indoor working subjects were much less tolerant to heat than carpenters or gardeners who were outdoors in the sun. The typists began to feel hot and sweaty at 26°C, whereas the outdoor subjects would sometimes not acknowledge thermal discomfort at 38°C.

The third thermally stressful environment studied was the very hot and arid central Australian Mitchell grass country at Toorak station, near Julia Creek. Here the agriculture department was setting up a tropical sheep research station, and Victor and his colleagues Ron Morris and Beth Howard worked there in summer vacations studying the water and electrolyte status of sheep, while themselves experiencing high temperatures and intense solar radiation.

A glance at the list of papers published by Victor and co-workers during his tenure of the Brisbane chair reveals the remarkable breadth and diversity of his research interests, the topics including schistosome dermatitis, properties of skeletal and cardiac muscle, effects of day-length on reproductive and other functions in ferrets, and pain-producing agents in the Queensland stinging bush Laportea, in addition to the main series of papers on the effects of different thermal environments on a range of body functions.

Victor's interests, always wide, were further extended during his first prolonged visit overseas in 1951. Chandler Brooks, who knew Victor from Dunedin days and had since become chairman of physiology in the State University of New York's Downtown Medical Center in Brooklyn, NY, invited Victor to work there as Visiting Professor. The Macfarlanes crossed the Pacific in the slow but luxurious, single-class propellor-driven aircraft of those days for their first glimpse of the bright new world of the United States, and soon settled in Brooklyn. Victor worked on several different problems, including the use of radioactive isotopes, a method which he used extensively in later years. In the research at Brooklyn, I¹³¹ was used to measure the rate of iodine uptake by the rat thyroid, and the effect of hypothalamic stimulation on this rate was examined. Victor also acquired the technique of making and using glass micropipette electrodes, developed by Graham, Gerard and Ling, for impaling single muscle and other excitable cells. Pam continued her artistic activities and studies, and they both enjoyed the cultural opportunities available in New York City, including the performing as well as the visual arts. Victor wrote:

New York was a relatively safe town in the early 1950s...It was still possible for Pam to work in the Art Students League at 57th Street and come home on the 11.30pm subway without any feeling of threat or disturbance The art, science, music (ancient and modern), Marthe Graham's dance and the steady stream of intelligence passing through New York made it a very civilised place from our point of view.

Victor took the opportunity of visiting other well known medical science laboratories in New York, including du Vigneaud's at Cornell – then on the verge of achieving the first synthesis of a polypeptide hormone – and the leaders in studies of kidney function, Homer Smith and Robert Pitts. Perhaps the most exciting event was the famous symposium at Cold Springs Harbour in 1951 at which Eccles and others described major breakthroughs in nerve cell physiology stemming from the intracellular microelectrode technique. Victor's words convey the excitement of the occasion:

There Hodgkin, Huxley, Katz and Eccles were in full flight, and Eccles had become converted to chemical neurotransmitter action. In 1951 when this symposium occurred the sodium-potassium permeability changes as a concept for action potentials in nerve and muscle had finally taken shape...and most of the essential stories for the Nobel prizes which Hodgkin and Huxley received for the nerves, Eccles for excitation and inhibition and Katz for micro-potentials, were there for those who cared to perceive them.

During this occasion Victor was able to converse at length with these giant figures in the world of cellular neurophysiology.

In the spring, the Macfarlanes explored more of the country, driving a small Austin to the west coast and back and meeting more leaders in physiological science while visiting medical schools and other institutions. In California, Victor met Magoun and Gerard, of the intracellular micro-electrodes, also Evans, Li and Chaikoff of endocrine fame. Altogether, fourteen US and four Canadian medical schools were visited to learn their organization, standards and general educational approaches. After these crowded months in North America, the return journey was made via Europe. There four months were spent visiting twelve of the main universities and medical schools in Britain and, on the continent, ten medical schools in France, Belgium, Holland, Denmark, Sweden, Germany and Italy. This intensive tour provided Victor with 'a rounded concept of what is done, what is possible and what is planned in research and teaching around this world'.

Back in Brisbane, Victor resumed once again the struggle against the heavy odds of excessive teaching commitments and inadequate support in those lean days before the Murray report. It is not surprising that there were times when he felt despondent, as noted by Hugh Parton after a visit. But he pressed on with his efforts to improve the courses and with his research, his zeal and techniques reinforced by the overseas experience. He also continued the summer visits to Julia Creek and his extra-curricular commitments in the community. The research on muscle cells was helped by acquisition of better electronic equipment and by the new techniques learned in New York, so that he was able to study the electrical and ionic events in both cardiac and skeletal muscle by means of intracellular microelectrodes. The events during recovery from activity in both these types of muscle interested him especially, and these were related to metabolic processes in the cells. The effects of differences in temperature on these processes were also examined, thereby linking this work to the major theme of thermal regulation and adaptation.

A symposium on 'Man and Animals in the Tropics' was planned by the Australian Academy of Science soon after receiving its Royal Charter in 1954. Eccles, a foundation Fellow and now heading the department of physiology in the John Curtin School of Medical Research at the Australian National University, arranged for Victor to join the organising committee, where he played a major role. The symposium was held in Brisbane during May 1956, and its success owed much to Victor's work on the committee, as well as to his paper delivered during the meeting on 'Mechanisms in heat adaptation'. 1956-57 were especially busy years for him, for in addition to the Academy symposium, he went on a UNESCO-sponsored science liaison visit to Indonesia, Malaya and India. He was also Australian representative on the International Standing Committee for Public Health and Medical Sciences of the Pacific Science Association, and in 1957 he attended the Pacific Science Congress at Bangkok as official representative of Australian medical sciences.

This was the same year in which Sputnik I gave its dramatic message to the world. In Australia it was followed by the 'Murray Report' on Australian universities and, shortly afterwards, the Universities Commission and a much-needed improvement in funding. As Victor put it: 'A strange astrological conjunction of Sputnik with Canberra set in motion funds for updating the old and for building new universities.' At about this time, Victor became convinced that Australian physiology, which had developed substantially since the war, needed its own professional association, and he began to take steps toward the founding of an Australian Physiological Society. En route to a CSIRO meeting in Melbourne, he went on a sort of missionary enterprise, as he put it, calling on several senior physiologists and putting the proposal to them. Reactions were mixed, mainly because of the distances and travel costs involved, especially in the case of Western Australia. The following year, with happier financial prospects after the Murray Report, general agreement was reached. The society's inaugural meeting was held in May 1960, appropriately at the University of Sydney, the institution which had established the nation's first department of physiology. Victor was elected as the society's first secretary and executive officer. More than any other individual, Victor was responsible for the society's genesis, and his influence during its formative years was profound. At its 21st birthday meeting at the University of New England in 1981, characteristically he gave a paper on the physiology of physiological societies. In a letter earlier that year discussing his plans for this talk, he wrote:

I thought I would talk about ethology, group behaviour and the limbic cortex...I think that group behaviour is about the most important piece of physiology there is, in terms of war and peace, food and drink, power and religion, politics and institutions. And as a gesture to the fusion of physiology with ethology I thought I'd talk about some of these tribal matters as they developed in the society.

By 1958, Victor was feeling the need to have more time for his multiple research activities, as well as relief from the relentless burden of teaching that he had shouldered for some nine years. Also, by this time, the Brisbane department was in good shape as a result of his efforts. The attraction of a senior non-teaching research post with Eccles in Canberra was strong, and during 1958 he made the decision to relinquish his chair and move to Canberra as Professorial Fellow. Also in 1958, the Macfarlanes set out on a second major overseas visit. Chandler Brooks had invited Victor to spend another period of work in his department, so he and Pam renewed their acquaintance with New York. Other visits in the New World were made as before, including one to Knut Schmidt-Nielsen, well known for work on desert animals, at Duke University, North Carolina; also a meeting in Syracuse, New York, at which Victor gave a well-received paper on his hot environment studies. Texas and Mexico were also visited before moving on to Europe, where Victor worked as Visiting Fellow in Professor Pickering's Regius Department of Medicine at the Radcliffe Infirmary, Oxford. Visits were also made to the Continent, this time including Spain and Portugal. Victor participated in an international meeting on tropical medicine held in Lisbon during September; his paper dealt with human water economy in the heat. Early in 1959, after returning to Australia, the Macfarlanes left Brisbane for Canberra, and Victor took up his new position at the Australian National University (ANU).

Canberra

While the physical and intellectual environment of Canberra was more bracing than that of Brisbane, Victor found that getting set up for his laboratory work, mainly on water and salt balance, was a slow business. There was also some physical isolation, his laboratory being on the first floor whereas the rest of the department, mostly Eccles' high-powered cellular neurophysiology group, operated on the fourth floor of the John Curtin building. Victor of course found plenty of things to do while establishing his laboratory, including more work on intracellularly-recorded activity of cardiac muscle, the equipment for which was available. He also became involved in studying the remarkable tolerance of rabbits to carbon dioxide and, in the reproductive field, observed that in Canberra the laboratory rat population produced an excess of males in their litters.

There seemed to be a similar disproportion in the human population of Canberra, at least for a time. Chandler Brooks writes that he believed Victor attributed this to changes in the water supply arising from clearing the native bush from catchments and replacing it with Pinus insignis. There seems to be no publication on this topic, in contrast to his work in Brisbane on the effects of ambient temperature and different latitudes on conception rates in animal and human populations. One topic which was closer to Eccles' field did engage his attention, namely, habituation of reflexes at purely spinal level. With Rod Westerman, a PhD student at the time, and W. Kozac from Poland, such plasticity was demonstrated. In cats surviving after complete spinal cord section, long-term changes in the pattern of simple spinal reflex responses were shown to follow repeated mechanical or thermal stimulation of the skin. Another sideline was further work on identifying the pain-producing toxins of the Queensland stinging plant Laportea, during which, according to Rod Westerman, Victor's nasal mucosa suffered from inhalation of the toxin-containing hairs from the leaves he was harvesting for analysis. However, his field work continued and intensified. Eventually the laboratory was ready and experimental work on water and salt mechanisms in animals – mainly sheep – resumed.

One unusual sideline of Victor's in Canberra was to solve a visual problem in the Academy's auditorium. The walls of this meeting hall were lined with pale, vertical timber studs with dark gaps between, and this background of alternating light and dark strips proved to evoke most uncomfortable visual effects, even to the point of vertigo in some people. Victor's solution was to reduce the strong light-dark contrast by filling the dark gaps with closely-spaced vertical strings, also of pale colour.

On the domestic and social front, the Macfarlanes made many new friends, such as Sir Keith Hancock and family, who were neighbours. They found the cultural life of the capital since the establishment of the ANU much to their liking. Pam's artistic activities flourished, and her work became well known and sought after, especially after a major exhibition of her paintings in 1961. And, to the parents' delight, a daughter (Ingereth) was born on 31 July, at the time of the exhibition in Melbourne. Yet it would appear that Victor's work in the John Curtin School fell somewhat short of his expectations. This may partly have resulted from the translation from chairmanship to readership level. He had believed he could be a professorial fellow and work much as he had done in Brisbane, but in 1978 he said: 'The facts are different. It is harder to work without a power-base.' Sir John Eccles wrote: 'He felt a second-class citizen on the 1st floor against the neuroscientists on the 4th floor...He became very indrawn, a great contrast to his ebullient character earlier.' Again, Victor was a gifted and dedicated educator, and it may be that lack of the stimulating, if often exhausting, contact with undergraduates contributed to his problems in the John Curtin School.

On the other hand, his field work flourished, and much was achieved, especially in the desert, during his six years in Canberra. He continued to attend and contribute to conferences in Australia and overseas, including the 1960 Arid Zone conference in Melbourne and the Xth Pan-Pacific Congress at Honolulu in 1961. As a result of a paper he gave in Hawaii on the stinging bush Laportea moroides, he obtained a substantial grant towards the studies of aborigines in the Australian desert and indigenous societies in New Guinea. In 1962, Victor attended an international symposium in Lucknow, India, on environmental physiology and psychology in arid conditions, contributing a paper on Merino sheep as desert animals. Afterwards, he was able to make his first visit to Africa, again with UNESCO support. Here Victor studied a range of African fauna in Kenya and elsewhere, including identical twin cattle, antelopes, Somali sheep and goats, and people. He measured their water turnover and ability to adapt to thermal stress.

The human environments studied included the deep mines of the Great Rand. He said:

...it was an astonishing experience too, to be up at dawn into little concrete boxes, where these people lived, go with them down the mines to these saturated environments of great heat down at the 8-12,000 foot depths. We measured various things, including the degree to which these people overheated as they tried to acclimatize to these mine environments and although they were tropical people (Bantu), many of them couldn't make the grade at temperatures like 32° dry bulb, 31° wet bulb, and those that overheated were the ones who didn't have much water turnover; those that didn't overheat had high water turnovers. This is the beginning of the sort of ecophysiological view of things – that turnover rates do determine adaptability and the amount of elasticity in the system.

In Queensland, Victor had measured water and salt turnover in adult Europeans, namely cane-cutters, working under hot and humid conditions, as well as in human subjects in the laboratory; but he was yet to study these functions in nomadic Australian aborigines in the desert. However, special opportunities for such work were shortly to be arranged.

After returning to Australia early in 1962, Victor planned more desert field trips. In the first of these, camels at Alice Springs were the subjects, and Knut Schmidt-Nielsen measured their metabolic rate while Victor studied their water turnover. The other venture aimed to study nomadic aborigines in the western desert during summer, and was a joint expedition with the anthropologist Norman Tindale, together with other Australian biologists and patrol officers from Woomera. The desert journey was delayed until 1963, but eventually the expedition set out in motor vehicles from Woomera. During the next four weeks, two field stations were set up. One was in the Rawlinson ranges in Western Australia, where aborigines of the Ngadadjura tribe, nomads who had little exposure to Europeans, were camped at a water hole. The other station was near Mt. Davies in the far north-west of South Australia, where members of the Pitjantjara and Nakako tribes were camped. Victor's biology group studied the Europeans in the party as well as the aborigines for water turnover and thermal balance, while Tindale recorded aboriginal songs and ceremonies on tape and film. In contrast to the expectation that desert-adapted humans might have a high, camel-like water economy, the measurements showed exactly the opposite. Like the African Bantu miners, the Australian aborigines are tropical in origin, having migrated from South-East Asia some 40,000 years ago, and their sweat-rate and water turnover are high. As Victor said: 'They have a higher water turnover by a factor of two than we Europeans do, even though they are in the howling desert.' They survive in summer by staying near waterholes and drinking a lot; they are 'great water drinkers' and can down 2 litres in 30-40 seconds; and of course they sweat profusely.

To carry out such field studies in animals or humans, ordinary laboratory procedures for measuring volumes of fluid input and output – 'by bucket', as Victor put it – are less than satisfactory, especially if the subjects are moving around in their usual activities. Periodical assays of electrolyte or hormone content of body fluids present little problem. Victor adopted tracer methods for measuring body fluid volumes and turnover, using tritium in small amounts (heavy water, deuterium oxide, needed mass spectrography, so was soon abandoned). Tritium (TOH) concentration is easily measured, and this proved ideal in field studies: 'with that it was possible to get nice answers, both on the amount of water in an animal and the rate at which it went through...It isn't a static pool, it is a flowing river'. One of the first applications of this method came during his first visit to New Guinea, also in 1963, in the highland Chimbu region, where he measured the throughput of water from mother's milk in babies. Victor had been greatly intrigued by reports that in the New Guinea mountains people didn't drink: he

was rather puzzled to think that there were people who did not drink, especially on latitude 6° from the equator. So we went up there into the mountains and lived in these rather nice thatched bamboo huts...on the top of a ridge. Somewhere about 300 feet below the ridge there was a stream which flowed down to the Chimbu river, but no Parian really felt that it was worthwhile walking down to the stream and dragging water up to the top. And the answer was that they had taken their water from the kar kar, the sweet potato, their staple diet. An adult male Chimbu eats about 4 kg a day...he gets over three litres of water a day out of just eating and burning the starch from this vegetable [of which 80% or more is water.]

Many more visits to other countries, including New Guinea, were to be made in Victor's pursuit of ecophysiology and palaeophysiology, terms he had introduced in 1961. But in Australia, another major event was brewing. In 1963, the University of Adelaide, on the advice of Dr Jim Melville, Director of its Waite Agricultural Research Institute, created a Chair of Animal Physiology. Victor Macfarlane was amongst the strong field of applicants, and was clearly the selection committee's first choice. Melville wrote in 1978 on the occasion of Victor 's formal retirement: 'I recall my relief and satisfaction when...the position brought a strong field of applicants. My relief and satisfaction were still further enhanced when Victor Macfarlane, the first choice of the Appointment committee, accepted the position.' So in 1964, Victor left Canberra for Adelaide to take up the position of Foundation Professor of Animal Physiology at the Waite Institute.

The Waite Institute

It can be surmised that this final academic appointment was the most satisfying position held by Victor during his professional career. Not only did he have once again a chairman's 'power-base', but the Institute's location in Adelaide made it ideally suited for his continuing field work in the dry two-thirds of the Australian continent Furthermore, once again he had undergraduate as well as postgraduate students to work with, though with a much more modest teaching load than had been his lot in Brisbane. He was also able to make many more visits abroad for ecophysiological studies of an increasingly wider range of animals. There is little in the record about Victor's teaching and other more routine activities during his 15 years in this appointment. However, there can be no doubt that the new department was highly successful, despite the limited resources available to it. Jim Melville wrote:

by whatever yardstick the success of the department is measured – the popularity of animal physiology as an option by third and fourth-year students, the number who have graduated with higher degrees, the number of mature scientists who have been accommodated, or the number and quality of scientific publications – there can be no doubt of the department's contribution to the Institute, the University and to science.

The Macfarlanes' house was beautifully situated at Crafers in the Adelaide hills, and looking south from it a panorama of Adelaide was spread out below. Visitors to the department from many countries enjoyed Macfarlane hospitality in this attractive setting, as well as in the laboratory. Some wrote their impressions, at the time of Victor's retirement, of their experiences at the Waite Institute. These included Hiroaki Shishido of the Japan National Institute of Animal Industry, Pulak Ghosh from the Central Arid Zone Institute at Jodhpur, Howard Haines of the University of Oklahoma's Department of Zoology and Hashim El Hadi from the Faculty of Veterinary Science at the University of Khartoum.

Shishido, who first met Victor during the Pan-Pacific Congress at Tokyo in 1966, writes warmly of his 'happy days at the Waite Institute' and the hospitality of the Macfarlane family. He failed, however, to make friends with the camel whose resistance to dehydration was under study, involving him in taking daily blood samples! 'She began to regard me as an enemy...Whenever she recognised me she roared, though I wanted to make a friend with her...In my living room there is hung a picture of the camel drawn by Mrs Macfarlane...' Pulak Ghosh met Victor during the 1962 UNESCO meeting at Lucknow, and in 1968 he worked at the Waite as UNESCO Fellow. Ghosh retains 'fond memories of warm hospitality...in the intellectually stimulating environment of his home'. Howard Haines 'liked the easy-going but efficient way that the business of the laboratory was run. This all reflects on Victor's abilities as the departmental head...There is no doubt that the best of my time with Victor MacEarlane was that in which ideas flowed. There were, invariably, challenging and interesting ideas or comments on whatever topics we were discussing...Ideas frequently ranged off into politics, society, literature and history.' From Khartoum, El Hadi wrote of his six months with Victor in 1976: 'I consider this as one of the most fruitful and bright periods of my scientific career...His coaching was close, keen and sincere.' He, too, was impressed by Victor's remarkable breadth of knowledge when he talked about 'the wider fields of science, architecture, history, arts, languages...I must say that I have learnt from him a few things about my own country.' Many other visitors from far and near enjoyed the stimulus and hospitality of the department and the Macfarlane family, and these quoted extracts convey something of the atmosphere and activities they encountered.

Victor's field work and overseas visits intensified during his chairmanship at the Waite. In December 1965, another expedition was made with Norman Tindale and an anthropologist from the University of Colorado into Pitjantjara country near the Musgrave Ranges. Once again, Victor's group carried out thermal and water balance studies on both white and aboriginal 'guinea pigs'. The following year Victor attended the Pan-Pacific Science Congress in Tokyo and made other visits, including one to Israel. In 1967, the New Guinea highlands were the scene of more work on water and salt metabolism in the indigenous mountain people.

Victor was amazed at the very low arterial blood pressure in this very fit, hardy population. He related this to the extremely low sodium intake in the mountain regions through prolonged leaching out of salt from the soil, with consequent low levels in their staple, vegetarian diet. The low sodium intake also, of course, is another factor helping to explain why the Chimbu do not need to drink water. Because of their low sodium intake, the hormonal (especially aldosterone from the adrenal cortex) and renal secretory mechanisms are adapted from birth to retain what sodium is needed for maintenance of blood volume, and to excrete the excess potassium from their staple vegetarian food. Ample blood flow is maintained through active tissues such as muscle despite the low driving pressure, because resistance to flow is low through relaxation of the muscular walls of arterioles; these are the vessels that become relatively rigid and narrowed in cases of hypertension. Victor also noted that, unlike the situation in Europeans, the blood pressures of these highlanders do not increase with age, but that exposure to European diets and culture soon changes the pattern to match that of most western societies.

These findings strengthened Victor's growing interest in the interaction between social factors – custom, tradition, and so on – the environment, and physiological function and adaptation. Group behaviour and the brain mechanisms upon which it depends – for many years one of his interests – began to loom as large as, or even larger than, his major theme of ecophysiology and climatic adaptation. High salt intake is a matter of social habit as well as addiction, and the figures obtained in the New Guinea highlands, at least for the non-Europeanised people, are very striking: 'these people have between 3 and 30 mEq per day of sodium going through them, as against the 100-200 mEq which wash through us daily for most of our lives, tighten up our arterioles, so that we get high blood pressure'. Later, another visit to New Guinea was made, to the Kuku Kuku country, the wildest part of New Guinea with only a year's contact with western ways. This time they measured thyroxine levels and turnover, and found remarkable correlations between thyroxine levels and social status.

In 1967, Victor also went to Britain again, including a visit to Aberdeen where one of his former lecturers in the medical school at Dunedin was now Professor Malcolm and chairman of physiology. Lawrence Malcolm notes that during this visit Victor introduced his department to the term 'ecophysiology'. 1968 was the year in which the 24th International Congress of Physiological Sciences was held in Washington, D.C., and Victor contributed to this, as well as visiting Japan again. Before and after the congress, Victor was once again a guest in Chandler Brooks' department in Nev York. Chandler writes: 'In this last visit I brought him to help me with the large flow of visitors before and after the IUPS Congress because Victor could talk with anyone about anything. I said to him, "Victor, you are a great help because if there is a period of silence you start to talk"...we got along very well, and so did our Russian, English, Yugoslav, Latin American, etc. visitors.' Further overseas visits were made over the next six years, including two more to Israel, another to Kenya, and an important period of work in Alaska where moose, musk oxen and reindeer were added to the animals investigated for water turnover. On his way back from Alaska in 1970, Victor stopped off in Japan once again, and renewed his acquaintance with Hiroaki Shishido.

In 1972 Victor Macfarlane was elected to Fellowship of the Australian Academy of Science, a well-deserved honour which should have been his several years earlier; it appears that the very extent of his interests and the range of topics on which he published were an obstacle to his election. In 1974, the 26th International Congress of Physiological Sciences was held in New Delhi, so Victor once again went to India and renewed contacts with colleagues in that country. After this he made his third visit to Israel and then, with Pam and Ingereth, worked in England at the University of Reading. During this time, his activities included some undergraduate teaching as well as advanced seminars. Geoffrey Waites wrote:

He brought all this rich experience to us in Reading in 1974 to 'weave patterns' for our students as he had done for so many others before. With his incomparable slides drawn from his travels on every continent, and others constructed with Pam's delicate artistry, he enchanted and educated. We still have the tapes of an excellent series of lectures on 'Brain and Behaviour', a model exposition in a difficult area.

In 1977, the Academy of Science arranged a one-day symposium on 'Water', which was held during its annual general meeting. Appropriately, Victor delivered a key address on 'the ecophysiology of water in desert organisms'. The paper based on this address sums up, perhaps better than any other, his findings and views about this topic and the broad role of water in biological systems. Amongst many other things, the paper demonstrates very clearly the genetic stability of physiological patterns over very long periods of time, and the importance of behavioural repertoires in adaptation to a wide range of environments. For example, the camelidae have the most efficient water economy, sheep are intermediate and cattle, originating in tropical regions, are the most wasteful; yet cattle can and do survive in very dry conditions, providing there is access to water. Camels, according to Victor, originated in the dry plains of North America; some went to Asia and Africa, others (the llamas) went south to non-desert regions. But their water economy has remained the same for some 3 million years. The Arctic musk ox, in terms of physiology, is also a camel. In addition to his studies of these large animals and humans, Victor also investigated different small native Australian animals such as dasyurid marsupials, and in these, too, wide differences in water economy became evident.

1977 was also the year of the 27th IUPS international congress, which was held in Paris. Victor attended and in addition to delivering papers, was chairman of a round table discussion on physiological adaptations to desert climates. In the same year, the one preceding his retirement, he went to Africa again, this time visiting Ghana, Ethiopia and the Sudan. Early in 1978, Victor was invited to participate in an international symposium on Arid Zone Research and Development held at Jodhpur, where he again renewed contact with Pulak Ghosh. He spent an evening with the Ghosh family and was in good form: 'He kept our young daughter and us glued to our chairs for more than two hours with a most vivid account of the fascinating history of the Scottish clans...The narration was generously seasoned with passing discourses on a host of subjects ranging from the Australian aborigines to Ikebana to Zebu cattle.'

Before his official retirement, Victor decided to make an experiment exploiting electronic technology as a new means of biographical recording. He was interviewed by Laurie Geffen, chairman of physiology at Flinders University, and this was videotaped by the Adelaide University's Advisory Centre for University Education. The tape is entitled Victor Macfarlane: the Genesis of a Ritual, and is sub-headed 'In airs, waters and places'. Victor had for long been interested in how rituals began, and thought that he might be pioneering a new kind of ritual for biographical purposes. The end of 1978 saw his official retirement and in 1979 he became Professor Emeritus of the University of Adelaide. In the same year, a symposium honouring Claude Bernard, organised by French physiologists, was held in Paris. Victor was pleased to have been invited to contribute to this special meeting, and according to Chandler Brooks, with whom he corresponded about his contribution, he worked very hard for this occasion. Chandler writes: 'One of Victor's last triumphs was participation in this Claude Bernard symposium in Paris.' His paper was an outstanding success.

Victor's retirement, clearly, was purely formal for, apart from shedding some administrative and routine teaching duties, he pursued research as actively as before, and continued to receive grants for its support and to write papers. He also undertook some undergraduate as well as advanced teaching, including lectures to undergraduate classes at the University of New England. These teaching sessions included some on the limbic system of the brain and its important role in behaviour. Professor Rogers recalls a seminar Victor gave on the limbic system. As he talked, he began sketching on the blackboard a diagram, 'which to say the least, had a wildly erotic content. A colleague of mine – always the fall guy – shouted, "Watch it, Victor." With a bland and rather sly look Victor said "What's that, Professor?" as he added a line which transformed the diagram to a well-recognised outline of part of the limbic system.' This harmless little trap illustrates one aspect of Victor's dry and sometimes elfish humour. During the busy but tragically few post-retirement years, in addition to pursuing research, giving seminars and lectures and participating in overseas and local scientific meetings, Victor also continued to play an active role in the affairs of a number of committees. These duties included chairmanship of the Academy of Science's National Committee for Nutrition. It was during a special meeting of this committee in Canberra that he suffered his sudden and final collapse.

Amongst Victor's many honours and distinctions, in addition to Fellowship of the Academy of Science, he received the Petersen Award in Animal Climatology, the Decennial Medal of the Negev Institute for Arid Zone Research, and the Mueller Medal of ANZAAS.

He was indeed a polymath, someone to whom all knowledge should be comprehensible. Chandler Brooks sums up the qualities of this remarkable man: 'An ability to do, an appreciation for the totality and freedom of the mind from conventional ways of thought is, to me, the Renaissance. That definition would characterise Victor Macfarlane.' But despite his many achievements and outstanding intellect, his breadth of knowledge, his iron determination and penetrating wit, he was a warm, modest and caring soul, with a deep concern for fellow humans as well as for the whole biosphere. This concern was manifest on many occasions when he used his medical training to help people in trouble. Sir Keith Hancock wrote of him in 1978: 'Had he so chosen, he would have rendered pre-eminent service in medical practice, for he possesses the two great gifts of healing – scientific knowledge and human understanding.'

Acknowledgements

Mrs Pamela Macfarlane, Adelaide; Mrs Yvonne Adams (née Macfarlane) of Dunedin, New Zealand; Miss Preethi Perera of the Waite Institute; Emeritus Professor Hugh Parton of Christchurch, New Zealand; Sir John Eccles, Switzerland; Professor Chandler McC. Brooks, of New York; Professor W.P. Rogers of Adelaide; Biographical notes for the Academy, by Victor Macfarlane; Collected tributes to Victor Macfarlane on the occasion of his retirement; Transcripts of ABC tape of interview of Victor Macfarlane by Michael Daley; Transcript of the University of Adelaide's tape: 'Victor Macfarlane – the Genesis of a Ritual'.

List of awards and affiliations

Awards
Marjorie McCallum Medal in Clinical Medicine
Colquhoun Medal in Medicine
Travelling Scholarship in Medicine, University of Otago
Nuffield Fellowship in Medicine
Fulbright Travelling Fellowship
Scholar in Residence, New York State University
Petersen Award in Animal Climatology
Decennial Medal, Negev Institute for Arid Zone Research
Mueller Medal of ANZAAS

Affiliations
Fellow of the Australian Academy of Science
Member of the Physiological Society of Great Britain
Foundation Secretary of the Australian Physiological and Pharmacological Society
Member of the Endocrine Society of Australia
Member of the Institute of Aboriginal Studies
Member of the International Biometeorological Society
Past President, Physiology Section, Australian and New Zealand Association for the Advancement of Science
Fellow of ANZAAS

Notes

(1) Biographical notes by Victor Macfarlane

A.K. McIntyre, Emeritus Professor of Monash University.

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