Raymond John Stalker was born in Dimboola, Victoria on 6 August 1930 and died in Brisbane on 9 February 2014. He had a distinguished academic career at the Australian National University in Canberra and at the University of Queensland. His work on hypersonic flow was universally recognised, and the ‘Stalker Tube' facilities he pioneered were able to reach unprecedented flow speeds and were reproduced in many laboratories around the world.

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About this memoir

This memoir was originally published in Historical Records of Australian Science, vol. 27(1), 2016. It was written by Caroline Stalker, Richard Morgan and Roger I. Tanner.

Written by M.J. Aroney and A.D. Buckingham.

• Early life
• East London College
• University College London
• Le Fèvre's war work
• The early years in Sydney
• Research in Sydney
• Professional and community activities
• Personal
• About this memoir
  • Acknowledgements
  • Notes

Early life

Raymond James Wood Le Fèvre was born in North London on the first day of April, 1905. He was the eldest of three children of Raymond James Le Fèvre, the managing clerk of a firm of London solicitors, and his wife Ethel May (née Wood). Of his four grandparents, three had died before 1910. Only his father's mother, née Louise Darby, of Bath survived into his childhood. She was a strict, severe and religious person, always dressed in black as was then customary for widows. She, with her watch-maker husband, had, many years before, established a home in Richmond, Surrey, and there produced six children of whom Le Fèvre's father was the youngest boy. Her family practically formed the local church choir of St Elizabeth's Roman Catholic Church at Richmond. At eight years of age, Le Fèvre became an altar boy and he remained in close association with this church, eventually becoming the Master of Ceremonies in his 20s. Further, through his close friendship with many of the clergy, he found great interest in church and choral music, history, ritual and church vestments. This aspect of his education and life over the years ran in parallel with his secular life.

Le Fèvre's parents moved to St Margaret's, East Twickenham, in Middlesex, a district just across the Thames from Richmond. His schooling began in the infants' class of Gumley House Convent, Isleworth, and after about a year he was transferred to St James School, Twickenham, an elementary school staffed by Sisters of Mercy and lay teachers. About April 1915, he was transferred to the Salesian school at Farnborough, Hants, a boarding school which Le Fèvre has recorded as being 'austere though probably healthy'. Cricket and football were played but he showed no great aptitude for either. He was, however, a good swimmer; his family kept a punt on the Thames. He enjoyed visits to Farnborough Abbey and roaming relatively freely on Farnborough Common. The Abbey had been founded by the Empress Eugenie to house the tombs of the Emperor Napoleon III and his son the Prince Imperial, who had been killed by an assagai while fighting in Zululand in 1879, in the Crypt Chapel, which to the young Le Fèvre had a fascination as a sombre and mysterious place – an impression intensified by distant chanting from the church above.

In 1915 Farnborough Common was still open to the public. For small boys part of its attraction was a large pond in which on warm days they could swim, but a larger part arose from the use of the Common as an aerodrome for the adjoining Government Aircraft Factory that was then growing up around the old balloon and airship establishment. Here could be seen pusher-engined Farman biplanes, B.E.2c's, and other fragile-looking contraptions staggering shakily into or out of the air. Le Fèvre could not have known at the time that the G.A.F. had just gathered together a group of able men, mostly from Cambridge, destined to become leaders in various branches of pure and applied science – men such as F.A. Lindemann (later Lord Cherwell), B.M. Jones (later Sir Melville Jones), G.I. Taylor (later Sir Geoffrey Taylor), G.P. Thomson (later Sir George Thomson and a Nobel Laureate in physics), W.S. Farren (later Sir William Farren, Director of the Royal Aircraft Establishment during the Second World War), G.T.R. Hill (designer of the 'Pterodactyl' aeroplane), F.W. Aston (subsequently a Nobel Prize winner in chemistry), and Hermann Glauert (whose contributions to aerodynamical design, to aerofoil theory, to the analysis of auto-gyro flight, and so on, were to earn him election to the Royal Society of London in 1931; at the height of his powers he was unfortunately killed by a falling tree on the Common in mid-1934). In the course of his career, Le Fèvre was destined to have an official place in the Establishment which grew out of 'the muddle of huge shanties' that were there in his school days.

Educationally the Salesians were 'earnest and mostly encouraging'; their curriculum and methods were conventional. Latin, algebra, geometry and French were taught among other subjects and particular emphasis was given to religious knowledge and the catechism. In 1916 Le Fèvre's parents decided that he should live at home, so in the Spring of that year he was enrolled in the Isleworth County School in Middlesex. The ICS, although founded formally only in 1895, could trace its ancestry back to a Stuart Charity School founded ca. 1630. From 1715-1813 this became known as the Isleworth Charity School where, as recounted in later years by Le Fèvre, '40 boys and 20 girls were clothed and taught reverence for Protestant and Hanoverian authority, given principles but not opinions, and generally taught to be submissive, pious, industrious, and respectful citizens'. The school evolved into a nineteenth-century National School and an Upper Department was established that later merged into a new grammar school being formed by the British and Foreign School Society. Le Fèvre attended this school from 1916 to 1923, recalling that 'The atmosphere in all senses was pleasant – literally because we were often downwind either of Pears' soap works or Watney's Brewery, and figuratively because our small numbers made friendships with one another, and contacts with the staff, easier and freer than would have been the case in a larger establishment.'

Because of the war, a number of masters were in the services, their places being temporarily filled by women. One of the latter, Miss E.B. Murdoch, conveyed to Le Fèvre his first impressions of chemistry. She believed in practical experience as a method of learning; her introductory experiment was the examination of a mixture of iron filings and flowers of sulphur before and after heating. The vivid light emission accompanying combination and the altered behaviour towards a magnet underlined indelibly the differences between physical and chemical changes. From that moment, chemistry displaced history as Le Fèvre's favourite subject. Conversion was completed during the next annual session when physics and chemistry were taught by B.H. Walmsley, 'the finest teacher in all my pre-University experience'. Notable among his school activities were a strong interest in historical novels (graduating from Henty to Scott), photography (winning a competition in Isleworth for two enlargements, done with a home-made lantern, of architectural subjects at Verulamium and St Albans Cathedral), rifle shooting (twice winning the school championship), and listening to music both liturgical and orchestral.

With the passing of the matriculation examination, the question of a future career became important. His father would have liked him to study law but this prospect did not appeal as much as a livelihood involving science and preferably chemistry. It was B.H. Walmsley who convinced his parents that this was not a disastrous course and that it was essential that he should arm himself with a degree and, if possible, get some experience of research before being launched on to the labour market. He therefore went into the post-matriculation science stream with the object of taking the London Intermediate BSc examination during the two years of the sixth form. Four subjects were required by the regulations. He chose pure and applied mathematics, physics and chemistry. The Headmaster, W.T. Kenwood, a believer in general education, discipline, and the idea that improvement of boys came best from hard work, personally took the combined science and arts groups for four periods a week, using a procedure that could be termed 'seminar leading', in which one student would read some passage or other and the rest would discuss it. The students learned a lot in this way, for example, selected passages from the Old Testament easily led to 'facts of life'; Ruskin's Stones of Venice started talk about architecture, travel, painting, mosaics and the like; an elementary book on economics likewise gave some insight into the production, distribution and consumption of wealth, and probably reinforced the determination of many to make a university, rather than commerce, their goal. Although a martinet who inspired fear in many boys and who closely and severely supervised the staff, Kenwood insisted that the sixth-form timetables included some half-dozen hours each week for private study, on their own without any masters being present – excellent training for the self-learning later to be undertaken in the post-school years.

Walmsley took the class for physics and chemistry. His enthusiasm in the laboratory was genuine and infectious. He was always fair and just, never sarcastic or sharp-tongued. They worked extensively from J.W. Mellor's Modern Inorganic Chemistry, E. Edser's General Physics and Light, H.E. Hadley's Magnetism and Electricity and others of similar standard. His introduction to organic chemistry followed the plan upon which E.L. Lewis's Elements of Organic Chemistry was based; this was essentially practical. In the very first lesson they learnt about fermentation, brewing, enzymes, wines, and so forth, and made up an aqueous solution of glucose to which yeast was added. A few days later the mixture was distilled and the easy accessibility of alcohol made obvious. Its value as a raw material was then illustrated by preparing, from fresh alcohol, such compounds as ether, ethylene, acetaldehyde, ethyl acetate and acetamide. In this way they were introduced to the idea that organic chemistry dealt with families of inter-related and interconvertible molecules; moreover, they were acquiring enough personal experience to be able to appreciate the simplifying and systematising advantages of Kekulé's structure theory which, sixty years earlier, had brought logic and harmony into what might easily have become a vast catalogue of uncoordinated empirical facts. Opportunities were given also to try their hands at many of the reactions and procedures described by J.B. Cohen in the 1918 reprint of his Practical Organic Chemistry, an outstandingly popular book.

In 1922 Le Fèvre sat for the London Intermediate BSc and Higher School Certificates. The examinations were passed at a level sufficient to qualify him for a Middlesex Senior County Scholarship worth about £25 per annum, a sum that just covered the fees at East London College (now Queen Mary College). Application was made to this institution and admission granted after various interviews. In September 1922, therefore, he found himself an undergraduate looking towards a BSc degree of the University of London.

East London College

In 1922, East London College was young in years. It stood on a site originally provided by the Drapers' Company in 1887, and seemed to have grown around the People's Palace and other buildings that had once housed a technical school and facilities for popular entertainment. The College thus found itself the possessor of premises that included a large hall containing a pipe organ; nevertheless the area was architecturally very ordinary. During 1907, E.L.C. was recognised by the University of London as a School in the Faculties of Arts, Science, and Engineering. Chemistry established itself under the influence of Professor J.T. Hewitt, a student of Victor Meyer and a man whose standards and interests had been influenced by membership of St John's College, Cambridge and periods spent in the Universities of Berlin and Heidelberg. The research climate around Hewitt was immensely stimulating and favoured all branches of chemistry (1).

Chemists who benefited from such guidance included Sir J.C. Drummond (later professor of biochemistry, University College London), G.M. Bennett (later a professor at Sheffield, then at King's College London, and finally the Government Chemist), and E.E. Turner (of whom more later). Others in Hewitt's circle, either as staff colleagues or research collaborators, were F.G. Pope, A.D. Mitchell and J.J. Fox (later knighted, and in 1936 appointed to be Government Chemist). Fox had contacts with early pioneers of physical methods for the investigation of chemical problems and among his publications were some dealing with topics in the formative days of infrared spectroscopy.

In time, Le Fèvre was ushered into the presence of Professor J.R. Partington by S.K Tweedy, J.R.P.'s hard-working personal assistant. He was a little disappointed to find, not a bearded and reverend-looking professor – his ideas having been formed by pictures of Mendeleef – but instead a pink-faced, somewhat testy individual who expressed dissatisfaction at the extent of his mathematical experiences. He was given an extended booklist with many of the standard chemical texts of the time and including several items by Partington himself – it involved a not inconsiderable financial outlay.

In the laboratory, Le Fèvre worked among a group of friends that was to keep together for three or four years until the ends of their periods as research students. Among these were N. Hadley, H.F. Halliwell, M.A. Mayes, F.G. Soper, E.B. Evans, A. Brewin, H.C. Gull, D.D. Moir and T.B. Child. A.I. Vogel was a year senior to Le Fèvre and a research student, working with J.R. Partington on sulphur sesquioxide. Le Fèvre has put on record some of his impressions:

As a candidate for chemistry honours, with physics a subsidiary subject, I attended lectures by a range of staff: Partington on inorganic and historical chemistry (although J.R.P.'s vocal style was not exciting, it was redeemed by the fact that he, plus Farrow his lecture assistant, performed in front of the class most of the experiments and demonstrations described in Partington's 'Text Book of Inorganic Chemistry'); E.E. Turner on organic chemistry (Turner was always clear and precise, he wrote rapidly and neatly on the blackboard, and often carried out small-scale preparations, crystallisations, or distillations, to illustrate whatever was under discussion, E.E.T. showed more than anyone else I encountered during my student years, what could be done with the simplest apparatus used with understanding and experience). W.H. Paterson and D.C. Jones took us for physical chemistry, which in those days was almost non-instrumented, reasonably practical, and not highly mathematical. To me the most troublesome course was thermodynamics, the material of which seemed 'dry' in the extreme, especially when droned out by J.R.P. more-or-less verbatim from his book. For eloquence, I think that Dr Alan Ferguson, of the Department of Physics, talking on the properties of matter, surpassed all others we heard as undergraduates; to Ferguson I owe my especial interest in 'additive properties' – an interest which has persisted for 60 or more years. For enthusiasm, I rank E.E.T. above all others. He occupied a small room at one end of the large laboratory where we 2nd and 3rd year students were located. I was on the second bench from the door of his room from which he would often emerge and chat with whomsoever he encountered. E.E.T. had not long before returned from two years as a lecturer in the University of Sydney, where he had collaborated with G.J. Burrows in examining the optical resolvability of certain inorganic complexes having Fe, Al, As or Sb as central atoms. At E.L.C. he was continuing this programme, making alkaloidal ferrioxalates, aluminoxalates, and antimonoxalates, etc. I well remember being invited to admire a beautifully crystallised strychnine aluminoxalate spread out to dry. Through his ever-open door we could see that he worked constantly. Later on, I had many opportunities to admire Turner's general skills in the laboratory, at glass-blowing (he said he had been taught by his brother, a student under Ramsay at University College) and in all phases of preparative technique, an aspect of chemistry for which Turner had genuine enthusiasm.

Le Fèvre's interests extended well beyond his science studies. He availed himself fully of the range of cultural activities so readily accessible in London – theatres, museums, cathedrals and churches with outstanding choirs, cinemas, historic buildings.

In June 1925, he passed his final examinations with First Class Honours in Chemistry and at once began research with E.E. Turner. On the BSc results he was awarded a DSIR Scholarship of £140 a year. His initial effort in research was inauspicious, involving as it did a too vigorous reaction and subsequent fire, but despite Le Fèvre's apprehensions, most people laughed off the incident. He began a study of the orientations of substituents entering diphenyl compounds. This led him to investigate the general usefulness of piperidine when used as an agent in locating halogen atoms situated ortho or para to nitro groups in aromatics. The results consistently supported the view that piperidine was of great value as an agent. This work was summarised in his thesis for the MSc degree, awarded in 1927; it also formed part of one of the first papers published with Turner. The scission of diphenyl ethers by piperidine was also examined.

The major preoccupation of Le Fèvre and Turner at this time was with questions concerning the stereochemistry of diphenyl and its derivatives. Chemists were beginning to query the space formula proposed by Kaufler. Reinvestigations showed much of his evidence to be dependent on errors and mistakes. From the confusion, a simpler picture gradually developed. Le Fèvre and Turner were led to conclude that an unsubstituted diphenyl molecule will tend to be planar unless it is prevented by steric or other forces, in which case the structure as a whole would adopt the shape of a two-bladed propeller and become capable of optical resolution into dextro and laevo forms. This recognition of conformational isomerism in the aromatic series opened the way for developmental work in several laboratories in England and elsewhere.

University College London

In 1928, Le Fèvre became a lecturer in organic chemistry at University College London. His departmental duties were not numerous. Mainly these were to demonstrate in the organic laboratory under the direction of O.L. Brady and to help on Saturday mornings with the elementary chemistry classes for engineers that were being run by R.W. Lunt. His immediate research programme was concerned with finishing his PhD work; the thesis (QMC thesis 82) submitted that year incorporated material from a number of published papers. The nitration of 4,4'-difluorodiphenyl was to be his last contact with this type of work on the diphenyl series, although Turner and his colleagues continued with these compounds very productively for thirty or more years.

At University College, changes were occurring. Professor J.N. Collie retired in 1928 and was replaced by Robert Robinson (later Sir Robert) who was then entering one of his most active periods of research. Robinson had already established a brilliant reputation for his contributions to many separate regions of organic chemistry, notably to electronic theories of reactivity and structure and to knowledge of plant pigments, drugs, alkaloids and so on. Robinson rapidly built up a large following of research students, some coming with him to London from Manchester, others travelling from countries overseas. Thus in a short time the UCL Department of Organic Chemistry became very cosmopolitan and stimulating to its members in ways not always experienced in other university departments. The traditions of friendliness were fostered by daily meetings for tea, coffee and much informal conversation, and the fact that staff and postgraduate students behaved as near equals. Years before, a 'Seven Seas Club' had been established for research students who had travelled to London via one of the seven seas or who had been at UCL for more than three years; all this helped to create a happy and united social atmosphere.

Robinson was not a remote or distant departmental head but one always accessible to colleagues and students, as he usually worked in his private laboratory with its doors constantly open. His wife, Dr Gertrude Maude Robinson, joined him very frequently, thus adding her personal interests in plant pigments and her quiet and dignified charm to the other attractive qualities of the Department.

Le Fèvre's lectures have been described by Professor H.J.A. Dartnall as follows:

I was a Chemistry student at University College between the years 1931-34 and attended his lectures on Organic Chemistry. The theatre was always filled to overflowing, for his lectures were marvellously planned and were not only full of examination 'meat' from the student's point of view but full of interest too. Nearly all the statements he made were illustrated by experiments, which were carried out with breakneck speed, yet he would write the equations on the black-board very slowly so that we could get them all down in our note books. He was always immaculately groomed and attired (like most lecturers then) in morning dress. I suspect he was a little vain, but with good reason for he had a commanding presence and was very handsome. His entrance into the Lecture Theatre was always accompanied by stamping of feet (a sign of approval).

His interests to use his own phrase 'ranged from the sacramental to the excremental'! On one occasion he was well ahead of his syllabus and therefore treated us to a fascinating and learned discourse on the famous burial shroud of Christ at Turin, with a chemical interpretation of the mysterious markings on the shroud.

Le Fèvre became actively involved with questions concerning the orientations of groups entering aromatic structures – a hotly argued topic at the time. The orienting ability of oxonium oxygen was examined experimentally as oxygen was the last element requiring investigation in this connection in the 'onium' state. In discussion, Robinson had suggested that the phenylpyrylium salts might be looked at 'since it should be easy to get a positive oxonium pole conveniently situated to orient substitutions in that series'. From substitution experiments mainly upon derivatives of 2-phenylbenzopyrylium perchlorate, Le Fèvre was able to conclude that the positively charged oxonium pole forms one of the strongest meta-directive influences known. Another study was concerned with the variable electropolar properties of the nitroso-group. In aromatic systems the strongly activating effect of the nitroso-group on halogens situated in the ortho- or para-positions was noted and compared with the similar action of the nitro-group in analogous circumstances. The nitroso-group was behaving as a meta-directing group yet experiment showed that nitrosobenzene could be directly substituted by bromine to give 4-bromonitrosobenzene. The apparently anomalous character of the nitroso-radical in nitrosobenzene on substitution by electrophilic reagents was investigated and rationalized in terms of Robinson's mechanistic concepts. Supportive evidence was provided from dipole moment measurements of derivatives of nitrosobenzene. Other work of that period included studies of the dinitration of 1-phenylpiperidine from which it was concluded that the piperidine radical possesses an abnormally strong para-directing influence.

During 1932-34, a number of cases of orientation by alkyl groups were investigated where the results appeared to follow the steric rather than the electropolar nature of the subsitutents. In this connection the nitration, chlorination, bromination, iodination and sulphonation of p-cymene were investigated quantitatively. In parallel, polarisation and polarisability effects in aromatic hydro-carbons were examined by dipole moment measurements on p-ethyltoluene, p-cymene, p-tert-butyl toluene, etc, and some halogeno- and nitro-derivatives.

In 1931-32 the interaction of aqueous ammonium sulphide with formalin, begun in 1928 with a study of the period of induction, was carried a stage further by the isolation of the chief product in a pure state, thus enabling its constitution to be determined. Other work included colligative-property studies of solid terpenes.

These last two projects were to assume a special significance for Le Fèvre since they introduced to research a Miss Catherine Gunn Tideman who earlier had been a student in his practical chemistry classes. To quote Le Fèvre:

Among those taking organic chemistry during 1928-29 was one who seemed always cheerful, lively, ready with relevant comments on current or local affairs, full of conversational topics, of repartee, energy and vigour; a hockey player of enthusiasm (who had once knocked unconscious an opponent through an accidental head-to-head collision), a tireless dancer, a rider of horses (her grandmother had owned a riding school in Glasgow), not excessively teetotal but convivial with most of the women and men contemporary with her at UCL. She and I chatted in the lab. about many things not always scientific. She lived in Lambeth where she 'housekept' for her two brothers and knew all about the Old Vic to which later on, she and I went fairly frequently. She was then C.G. Tideman, later Mrs. Catherine G. Le Fèvre.

The marriage took place in Glasgow on 1 August 1931. They rented a house at Neasden, about twenty minutes by car from University College. Catherine Le Fèvre was to become her husband's most constant and most valued research colleague. Their subsequent work in physical organic chemistry was strongly encouraged by Professor C.K. (later Sir Christopher) Ingold who in 1930 had taken up a Chair of Chemistry at University College in succession to Robinson who had been elected to the Waynflete Chair of Chemistry at Oxford.

From 1933 Le Fèvre became increasingly interested in the applications of dipole moments to chemical problems. He had been introduced to the technique by J.W. Smith with whom he had collaborated in measuring the dielectric polarizations of nitroso-compounds and of quinoline and isoquinoline, and from whom he later inherited the apparatus when Smith left University College. Consequently, the structures of numerous other substances were examined by this method. An account of such structural studies as well as the use of polarisation measurements to probe inductive and mesomeric effects in molecules, is given in Le Fèvre's book, Dipole Moments, Their Measurement and Application in Chemistry (Methuen, London), which was first published in 1938. Problems of solute association and aggregation were also explored. A number of papers were published on the variation of molecular polarisation with the permittivity of the solvent and, as well, with changes of state. An equation connecting the true dipole moment of a gas with the apparent moment in solution was advanced and led to the starting of experimental work on the dipole moments in the vapour state of a number of compounds with negative Kerr constants. Referring to the English period of Le Fèvre's researches, Ingold was later to write:

He had become well known as a physical organic chemist of great power and originality. With several others in England he had cleared up a morass of confusion concerning optical activity which, in fact, arose from unrecognised confirmational causes. He had largely alone, made several important contributions on the distinctive roles of polarisation and polarisability in polarity. Dipole moments were a much-used tool: using it he showed quantitatively how polarity could be inverted by conjugation.

In March 1935, Le Fèvre was awarded the DSc degree; he was promoted to Reader in 1938. In that year also their son, Ian, was born. Soon after, Catherine was employed as a demonstrator at University College and she also taught chemistry at Queen's College, Harley Street.

About this time the Le Fèvres decided to set up apparatus for the measurement of the Kerr effect, i.e. electrically induced double refraction. In this they were encouraged by H.A. Stuart of the Universität Mainz and G. Szivessy of the Universität Bonn, and were helped by a grant from the Royal Society. By 1939, measurements on gases and on organic substances in solution were being undertaken. Other topics under investigation at the time included: the stereochemistry of l,2-diketones, the geometrical isomerism of diazo-, azoxy- and azo-compounds, the configurational relationships of certain anils, the associations of aliphatic acids and of aromatic nitroso-compounds, the dielectric polarizations of vapours, phototropy, and equilibria of the keto-enol type. The outbreak of war caused the temporary cessation of these activities. Prominent collaborators between 1928 and 1939 were C.G. Le Fèvre, J.W. Smith, J. Pearson, P.J. Markham, S.N. Ganguly, P. Russell, E.D. Hughes, H. Vine, V. de Gaouck, P.P. Hopf, G.S. Hartley, I. Dostrovsky and C.C. Caldwell.

Le Fèvre's war work

During September 1939, University College granted Le Fèvre temporary release and for a short time he was attached to the Ministry of Home Security for the training of Gas Identification Officers. About a dozen chemists were needed to help organise and train the GIOs giving them 'practical experience of the gases that might be encountered in time of war'. Special centres of instruction were arranged in London and the provinces. Le Fèvre was sent to one such centre at Battersea Polytechnic where he worked as an instructor until well into December.

In the early days of January 1940, Le Fèvre joined the Directorate of Scientific Research, Air Ministry (later the Ministry of Aircraft Production). There he was an adviser to RAF Commands on certain chemical aspects of armaments. He familiarised himself with the techniques of manufacturing, handling, storage and charging of toxic liquids and other materials. A careful watch was being maintained at the time for evidence of the deliberate use by the enemy of toxic chemicals for offensive purposes. Official apprehension was fuelled by intelligence reports of vesicant stocks held by the Germans, of records of mustard gas spraying by the Italians in their Abyssinian campaign, and of analogous activities of French and Spanish forces in North Africa. The Japanese had dropped gas bombs at Changsha and it was thought they might behave similarly if hostilities started in East Asia.

On 2 October 1940, a daughter, Nicolette, was born to the Le Fèvres in London at a time when air-raids were beginning to become regular nightly occurrences. Catherine and the children left soon after to return to Harrogate where they had been living.

Early in 1941, it was decided that a supply of chemical weapons should be sent to the RAF Command in Singapore and Le Fèvre was to go there as a chemical adviser, with the honorary rank of Wing Commander. He proceeded to Singapore through West Africa, Egypt, India, Burma and Malaya. The fortnight he spent in Cairo gave Le Fèvre the opportunity to see the arrangements made by F.B. Kipping (from St John's College, Cambridge) for the reception and storage of chemical stocks in the Middle East. In Singapore he was met and briefed by Wing Commander Ramsay Rae (later to become an Air Vice-Marshal), who was then the senior armament officer of the RAF Far Eastern Command.

Le Fèvre's first duties concerned the extant plans for anti-gas defence of all RAF areas and making arrangements for the receipt, storage and handling of chemical weapons that were expected to arrive in a few weeks' time. He found most buildings on airforce stations to be 'fantastically insecure, being constructed in tropical fashion of light-weight materials of various kinds...'. While contemplating the storage of gas weapons, he chanced to meet M.W.F. Tweedie, the Curator of the Raffles Museum, whose archaeological and speleological interests had given him a good knowledge of excavations and caves in Malaya. On Tweedie's advice, the Batu caves just outside Kuala Lumpur were examined. Those that were suitable were cleared of the bat dung that had lain undisturbed for years by offering it to the Malay inhabitants who gladly took it, regarding it as first-class fertiliser. Within a week, the first of the gas storage depots in Malaya was started, and soon stocked by train from Singapore.

The RAF Far Eastern Command had hurriedly constructed landing grounds at intervals along the Burma road, so Le Fèvre was sent to visit them, up as far as Lashio near the China-Burma frontier. Of special interest was a stop at Toungoo where Colonel Chennault and his Flying Tigers were based. They seemed knowledgeable about Japanese tactics and equipment, but when asked about earlier reports of chemical warfare activity in China they appeared genuinely ignorant.

On his return to Singapore, Le Fèvre was given instructions from London that he was to try to travel into China as a university lecturer, a civilian, suitably clothed at Air Force expense. He was to go to Changsha or wherever Japanese chemical weapons (or casualties therefrom) had been reported and to try to arrange that a few unexploded bombs, and other chemical warfare samples, be brought back to some place where analyses and examinations could be performed. It was thought that Singapore, which had an efficient Government Laboratory, would meet this requirement adequately. However, intelligence reports began to come in that a Japanese fleet was somewhere in the South China Sea and fears of an invasion attempt on the Kra Isthmus or the Malayan coast were becoming stronger and stronger. Le Fèvre's China trip was therefore suspended pro tem; instead, a somewhat ancient specimen of a Type 92 50-kg bomb was sent from (probably) the Air Attaché's Office in Chungking. The liquid contents seemed to be a fairly pure Lewisite/Mustard mixture, as already described in Japanese weapon manuals available at the time.

Pearl Harbour had been bombed on 7 December and air raids on Singapore were soon to be a regular occurrence. They were almost unopposed since the defenders lacked aircraft capable of dealing with the numbers and capabilities of the enemy. With the sinking of the Prince of Wales and the Repulse on 10 December and news of further Japanese successes it was thought advisable to bring back to Singapore the Batu caves stocks. Ultimately it was decided that the whole stock should be put into lighters and towed to one of the numerous small islands near Singapore, where they would be out of the way of operations but accessible if needed. St John's East, a once-time leper settlement, proved suitable. Seven lighters were loaded and towed by tugs; they were taken to the island, run up the beach and secured.

However, the war situation was rapidly worsening and as part of a general 'denial' scheme, Le Fèvre was ordered to plan the destruction of 'gas' stocks. With time running out, the lighters were taken out to sea and sunk. During the above operations, the S.S. Silver Larch arrived in Singapore bringing chemical and other cargo. It was re-routed to Oosthaven in south Sumatra. About a week before the capitulation, Le Fèvre left Singapore on a Yangtse river boat, the Whangpu, for Palembang, Sumatra, and from there he went by train to Oosthaven. Still pursuing the Silver Larch, he set out for Batavia on a 500-ton RAF auxiliary, the Tung Song, and from there went to Tjilatjap in south Java. A group of about 240, mostly RAF personnel, was taken on board the Tung Song, believed at the time to be the last friendly ship to leave Java before the Japanese occupation. The hazardous journey to Exmouth Gulf in northwest Australia took ten days or so. On 14 March 1942, they reached Fremantle. Thus, by chance, Le Fèvre made his first contact with Australia.

An interchange of signals took place between OHQ RAAF Kingsway and Air Force HQ Melbourne, and Le Fèvre found himself temporarily seconded to the RAAF at a time when fears were increasing of a Japanese invasion of Australia. He was given posts in the Directorates of Armament and Air Staff Policy. He remained in Australia nineteen months and travelled extensively about the continent and allied-occupied New Guinea, arranging for the storage and testing of mustard gas under tropical and semi-tropical conditions. It was.recognised that the physiological effects could be quite different from those known in Europe. Some of this work is described in a contribution by Le Fèvre to the 1983 Australian Department of Defence account, Mustard Gas Field Trials during World War II, by R.G. Gillis (2). He became a casualty when investigating a leakage on the Blue Funnel vessel Idomeneus, and was in Concord Hospital unable to see for six weeks. Although he regained his sight, he permanently lost his senses of taste and smell as a result of exposure to the vapours.

Le Fèvre visited most Australian universities, seeking to recruit Australian-trained chemists for war work involving chemical agents. In Sydney he met Professors J.C. Earl and C.E. Fawsitt separately, and recalled finding the Chemistry Department in a 'terrible state of disrepair' and the front lawn of the University dug up into air-raid trenches.

Having found an RAAF successor, he returned to the United Kingdom in December 1943, by air via the Pacific, USA, and the Atlantic. He resumed work at the Ministry of Aircraft Production headquarters as an Assistant Director (Research and Development, Armament Chemistry). In July 1945 he became Head of the Chemistry Department, Royal Aircraft Establishment, Farnborough. Since the authorities permitted a certain engagement in pure research, Le Fèvre took the opportunity to investigate aspects of diazocyanide chemistry that had been questioned in the chemical literature of the war period. One of his duties at Farnborough was to supervise the planning and construction of a new building for his department, an experience that would prove of great value a decade later when the opportunity came to build a new Chemistry School at the University of Sydney.

The early years in Sydney

After the war, Le Fèvre was approached in London by Professor Eric Ashby (later Lord Ashby), Chairman of the Professorial Board of the University of Sydney from 1942 to 1944, who invited him to take up the position of Director of Chemistry at that University.

Le Fèvre was attracted by Sydney as a city in which to live and to bring up a family far from the memories of a war-ravaged Europe. Moreover, the University of Sydney had a style somewhat reminiscent of the more traditional English universities and this too had its appeal. The teaching of chemistry there dated from the arrival in 1852 of Professor John Smith to take up the Chair of Chemistry and Experimental Philosophy. Robert Robinson held the Chair of Organic Chemistry, 1913-16. Another Nobel Laureate in the making was J.W. Cornforth who graduated MSc in 1939. G.J. Burrows, D.P. Mellor, F. Lions and F.P.J. Dwyer (who for a time had a very close collaboration with R.S. Nyholm) had made important contributions to coordination chemistry.

It was a regrettable fact that for some years prior to Le Fèvre's arrival, Chemistry at Sydney had become implacably divided into two groupings about Professors C.E. Fawsitt and J.C. Earl. It was to be Le Fèvre's task to attempt a reunification. He arrived in Sydney with his family in 1946 and in November of that year took up duties as Professor of Chemistry. The retirement of both Fawsitt and Earl allowed reforms in organisation to be made and at the beginning of 1948 the two separate departments of Chemistry and Organic Chemistry, Pure and Applied were fused into one Chemistry School. Le Fèvre was made Head of the School of Chemistry, a position he was to hold until his retirement in 1970. He was to be joined in 1952 by A.J. Birch who took up the Chair of Organic Chemistry and D.P. Craig who became the University's first Professor of Physical Chemistry. In the course of Le Fèvre's tenure, Birch would be replaced by C.W. Shoppee (office 1956-69) and Craig by A.E. Alexander (office 1956-70).

Le Fèvre's arrival in Sydney coincided with the great wave of post-war enrolments of ex-service and other new students. This highlighted the inadequacy of the chemistry building, something that was quite evident even before the war. A feature of the University's centenary celebrations in 1951 was a public appeal, and although the appeal did not reach anywhere near its target, it did produce an anonymous donation of £100,000 to be used for the building of a first wing of a new Chemistry School. The donor was subsequently disclosed to be the late Mrs. Brightie Phillips. With the promise of further funds from the government of New South Wales, construction began in 1955 with completion four years later, followed by an inauguration ceremony in 1960. Le Fèvre was intimately involved with the design and planning and with the supervision of construction of what was regarded by many as the finest chemistry building of any campus in Australia. D. Branagan and G. Holland pay homage to Le Fèvre's efforts in their history of science in the University of Sydney, Ever Reaping Something New (University of Sydney, 1985).

Le Fèvre's period as Head was characterised by the great impetus he gave to research. His active encouragement of research groups within the School is reflected in the statistic that during his first fourteen years in office the number of articles from the School published in learned journals amounted to 680, a major component of the University's entire research output. With the introduction of PhD programmes in the late 1940s, a rapid increase occurred in research student numbers and the Sydney School was to achieve a pre-eminence in research within the Australian university system. Le Fèvre himself had about one hundred research students and co-authors during his term of office. His publication list includes reviews, a book, and several hundred research papers.

Research in Sydney

Le Fèvre resumed his research work in Sydney, taking up and extending projects commenced in England but interrupted by the war. Much of the apparatus at University College had been destroyed in the bombing of London so new equipment had to be constructed for the Sydney laboratories. This was to be the beginning of more than two decades of highly productive and innovative research effort.

From the outset, a programme of investigation was undertaken of the structures and configurations of a variety of diazo-compounds, of photochemical and thermal transformations between isomeric forms, and of the kinetics of isomerisations in solution. These studies strongly supported Hantzsch's ideas on the structures of diazo-compounds and led (in the best traditions of the subject!) to controversy in the columns of Chemistry and Industry with H.H. Hodgson of Huddersfield (see, for example, Chem. and Ind, 1948, 270). Associated with Le Fèvre in this work were K.E. Calderbank, J. Northcott, J.D.C. Anderson, I.R. Wilson, A.A. Hukins, P. Souter, D.D. Brown, R.N. Whittem, H.C. Freeman, T.H. Liddicoet, I. Youhotsky and C.V. Worth. A number of infrared spectral studies were made with M.F. O'Dwyer, R.L. Werner, J.B. Sousa, W.T. Oh, I.H. Reece, R. Roper and M.J. Aroney, to specify the stretching frequency of the –N=N– group in a range of diazo-compounds and to identify infrared spectral features characteristic of the diazonium cation. Working with Sousa and Roper, Le Fèvre proceeded to investigate the kinetics of normal to iso-diazoate transformations in strongly alkaline solutions and to make various attempts to disentangle the complicated series of pH-dependent equilibria between diazoates and diazonium salts. The programme was effectively concluded in about 1963.

Early in the Sydney period, the first observations were made of the high polarities of J.C. Earl's newly discovered 'sydnones'. Information on molecular structures and the mesomeric effects operative in this new class of compounds was extracted from dipole moment and infrared spectral measurements. Interest in the sydnones led to an examination of 'model' structures such as antipyrin and phenylisooxazolone and to an understanding of electronic displacements in keten and some of its derivatives. C.L. Angyal, R.D. Brown, A.A. Hukins, E.M. Leake, G.A. Barclay and R.S. Armstrong contributed notably to this work.

Investigation of the effects of medium and state on the apparent dipole moments of substances, started with P. Russell and C.G. Le Fèvre before the war, was carried forward in collaboration with I.G. Ross, B.M. Smythe, G.A. Barclay, C.G. Le Fèvre, J.W. Mulley, C.L. Angyal, H.G. Holland, H.C. Freeman, A.D. Buckingham, B. Harris, E.P.A. Sullivan, D.A.A.S. Narayana Rao, J.Y.H. Chau, F. Maramba and J. Tardif. Dielectric polarisation measurements were made for a wide range of molecular substances in the gas, liquid or dissolved states primarily to assess existing theoretical treatments (Clausius-Mossotti-Debye, Onsager, Ross-Sack, and others) as well as various empirical relations by which the 'true' or gas-phase dipole moments of molecules could be predicted from experiments on solutions or on pure liquids. Correlations between solvent effects as observed from experiment and the sign of the Kerr constant attracted particular attention. In the 1953 edition of Dipole Moments (p.82), Le Fèvre concluded that:

None of the empirical or theoretical approaches...adequately embraces all the known facts concerning the effects of the medium in dipole moment measurements. Some seem valid for the differences between µ _solution and µ _gas but fail when tested on pure polar liquids, others – especially produced for polar liquids – are unable to cover all such liquids.

Such studies effectively culminated in empirical equations that fitted known data better than any others previously proposed. Le Fèvre was able to claim that 'a method, useful in practice, is now available for estimating µ _gas from either µ _solution or µ _liquid'. A postscript to this work was a study of polarisation effects in liquids carried out with Narayana Rao.

In parallel with the above programmes and substantially with colleagues already named, Le Fèvre continued his long-standing interest in the elucidation of problems of molecular structure and geometry using dipole moments, often in conjunction with infrared and electronic spectral data.

Examined were, inter alia, oxygen, sulphur and nitrogen containing heterocycles, monoterpenes, oximes, 'iodoxybenzene', substituted benzocinnoline 6-oxides, 2,2'-dipyridyl, 'diphenylmaleinitrile', substituted aryl nitro compounds, 1,4-dioxan and the temperature dependence of its structure, species with possible keto-enol tautomerism, and solute intermolecular interactions. Other topics that interested Le Fèvre in his first decade in Sydney included the thermotropy of spiro-pyrans and thermochromism of methyleneanthrones; the a priori calculation of atomic polarisations from spectroscopic data; and the influence of molecular shape and of medium characteristics on the dielectric relaxation times of dipolar solutes. About the mid-1960s, he was invited to write 'Dipole Moments (Electrical and Magnetic) for The Encyclopedia of Physics.

As mentioned earlier, among other investigations laid aside for the war was one concerning the Kerr effect and the uses that this phenomenon might have in chemistry. The effect, known since 1875, occurs when a voltage is applied to a dielectric causing it to become doubly refracting, i.e. the refractive index at a given wavelength is different in directions parallel and perpendicular to the applied field. The magnitude and sign of the measured effect depends on the 'Kerr constant' which is related to the structure of the molecules forming the medium and their electronic properties. Equipment for measurement of the Kerr effect was reconstructed in Sydney (incorporating remnants of the pre-war apparatus that had survived) and the programme was restarted in 1947. Financial help was forthcoming, mainly from Imperial Chemical Industries, Australia and New Zealand Limited. The UCL-Sydney activities were the subject of Le Fèvre's Presidential Address to Section B of ANZAAS at the 1955 Melbourne meeting, which was titled 'The Kerr Effect in Chemistry'. To quote from pp.41-42:

Experience before the war had shown two things: the difficulty of taking electric double refraction observations on vaporized materials, and the necessity of extracting useful information from measurements made on solutions. As the mathematical formulae for the Kerr effect were strictly applicable only to gaseous dielectrics, the full usefulness of this property to chemists was limited, since many interesting substances could not be vaporized without decomposition. In 1947 the whole situation had some analogies with that in which dipole moments were being measured about 1920. It seemed fundamentally necessary therefore that the possibilities of examining materials as solutes, and ultimately securing some value for these solutes at infinite dilution, should be explored. This was made the first objective of the Sydney work.

By 1953, Professor and Mrs Le Fèvre had devised and tested a method whereby the electric double refraction of a solute, expressed as the quantity _mK_solute (the solute molar Kerr constant at infinite dilution) could meaningfully be extracted from experiments in solution. Using a modified form of the classical theory of Langevin and Born, they were able to obtain, from _mK_solute, the anisotropy of polarisability of the solute molecule. As well, it was shown that for polar solutes the ratios _mK_solute / _mK_gas roughly resemble the corresponding µ²_solute / µ²_gas values, thus making it possible to calculate approximate gas-state Kerr constants from solution measurements. The Le Fèvre solution-state approach was regarded by H.A. Stuart and others in the field as an important achievement, especially since previous efforts had yielded indifferent results. It was to become clear with further work that the treatment should be restricted to observations made in non-polar media such as carbon tetrachloride, cyclohexane or dioxan. Nonetheless, wide fields of applications were opened. A full account detailing the theory, the technique of measurement, and results to hand at the time, appeared in 1955. Further results to 1959 were collated in 'The Kerr Effect', Chapter XXXVI of Physical Methods of Organic Chemistry, ed. A. Weissberger (with C.G. Le Fèvre), and in Le Fèvre's Liversidge Lecture. Progress up to the end of 1964 was summarised by Le Fèvre in 'Molecular Refractivity and Polarisability' in Advances in Physical Organic Chemistry, ed. V. Gold, wherein Le Fèvre placed particular emphasis on the application of the Kerr effect to stereo-structural problems. He illustrated this with the tabulation of a wide range of examples investigated in the Sydney laboratories. In a further account in 1970, 'Polarisation and Polarisability in Chemistry', Le Fèvre discussed the central nature of the polarisability concept in relation to molecular interaction, transition states, reaction pathways, energetics, and kinetics. His final overview was written, with C.G. Le Fèvre, about the time of his retirement: 'The Kerr Effect', Chapter VI of Physical Methods of Chemistry eds. A. Weissberger and B. Rossiter.

The Le Fèvres analysed Kerr constants from solution, together with solute dipole moments (in the case of polar substances) and molar electron polarisation data (the latter from molecular refractivity dispersion), to determine the molecular principal polarisabilities for solutes such as methyl and t-butyl halides, chloroform, benzene, a number of 1,3,5-trisubstituted and hexasubstituted benzenes which, because of their symmetry, are associated with polarisability ellipsoids of revolution. They were able to extend the method to molecules such as mono- and di-substituted methyl-, halogeno- and nitro-benzenes, pyridine, quinoline and acetone which, because of their lower symmetry, required additional data for specification of the polarisability tensors. This was provided, in the first instance, by recourse to early measurements in the literature of the depolarisation factor for light scattered transversely by a substance (from J. Cabannes, La diffusion moléculaire de la lumière, 1929). in this way the Le Fèvres were able to provide valuable information on solute polarisability, a measure of the electronic response to a perturbing field for particular directions within the molecular framework. A weakness in the procedure, however, was uncertainty as to the appropriateness of the light scattering data, which derived from measurements on gases or pure liquids. The problem was addressed by Le Fèvre, working with B. Purnachandra Rao, in the 1956-60 period. They developed a method of determining solute anisotropies from measurements of depolarisation factors of light scattered by solutions. With the aid of a grant from the Nuffield Foundation, they constructed apparatus based at first on the Cornu visual technique but modified later for photometric detection. This work placed on a firmer foundation the determination of anisotropic polarisabilities for solute molecules. Prior to this development, Le Fèvre had advanced a number of approximate methods of estimating polarisability along a specific molecular principal axis. These were based on: (a) the interpolation of values for members of a related group of molecules where regular trends in polarisability had been observed, as in the methyl-substituted benzenes; (b) a rough correlation which had been found between ratios of molecular dimensions and directional polarisabilities; or (c) the use of crystal-state refractive index and density measurements as was found appropriate for naphthalene.

With the determination of polarisability tensors for a substantial number of molecules of known geometry, the Le Fèvres proceeded to test the notion, suggested by E.H. Meyer and G. Otterbein (Physik.Z, 32, [1931] 290), that polarisability parameters can sensibly be ascribed to discrete molecular segments such as individual bonds. From dissection of the molecular values, Le Fèvre was able to set up an extensive scheme of anisotropic polarisabilities for bonds commonly encountered in organic molecules. A compilation of bond polarisabilities to 1965 was presented by Le Fèvre in his chapter in Advances in Physical Organic Chemistry. It was stated, however, that the anisotropy of a given bond should not be regarded as a 'universal' constant since it could be affected by the structural environment and particularly so by conjugation or mutual induction. Comparisons made between the C-X group polarisabilities (X=halogen, Me, NO₂ or CN) of aliphatic and aromatic structures showed small polarisability augmentations (exaltations) to occur along the C(aryl)-X bond axis with slight diminutions in directions perpendicular to that axis. The evidence, Le Fèvre wrote, correlated with 'the non-classical polarisability mechanisms long used in organic chemistry to formulate the temporary transmission of electrical effects from group to group or from substituent to reactive position (3)'.

A safe viewpoint, adopted by Le Fèvre, was that the apparent bond polarisabilities drawn from measurements on solutes should be regarded as empirical and that they may appropriately be applied to molecular situations analogous to those from which they were derived. A very considerable body of evidence justifying this was accumulated over the years, reference to which is made in the Le Fèvres' 1972 contribution to Physical Methods of Chemistry. Ancillary work involved devising empirical correlations between polarisability components and the dimensions of bonded atomic groupings, bond vibrational stretching frequencies and wavelengths of maximum absorption of the K-band in conjugated diphenylpolyenes, and the calculation from bond polarisabilities of optical rotatory powers.

The determination of bond polarisabilities as well as polarisability tensors for larger molecular segments such as phenyl, amide and other groups, though important, was regarded by Le Fèvre as the precursor phase to what can only be described as an enormous programme of study of the Kerr effect in relation to the stereo-structural analysis of molecules in solution. It had long been understood that the magnitude and sign of the Kerr effect could at times provide qualitative information about molecular geometry; for example thianthren, having a negative Kerr constant, must (from theory) be folded about a line joining the sulphur atoms. Phenazine, on the other hand, has a positive Kerr constant and is planar. The procedure most often used by Le Fèvre was to compute the molecular polarisability tensors (using component group parameters) and thence the molar Kerr constants for the various possible stereo-structures and to compare the latter with the molar Kerr constant from experiment. The power of the method, due in large measure to the fact that the Kerr constant is often greatly sensitive to variations of molecular geometry, was recognised early in the Sydney programme. Measured also, as a matter of routine, was the permanent dipole moment (as it came into the equations for the Kerr effect with polar molecules), and in many cases this assisted in the stereochemical analysis from dipole vector considerations. The techniques were used in a complementary manner. Apparent in Le Fèvre's later work of the 1960s is a greater use of other physical methods, notably nuclear magnetic resonance and infrared spectroscopy, to provide information that facilitated analysis of the Kerr effect, often through defining and/or limiting the range of confirmational possibilities. In the period 1955-71, Le Fèvre and his co-workers investigated the molecular stereo-structures in solution of a wide range of compounds.

It was Le Fèvre's grand design to establish the Kerr effect technique as a method of importance in molecular stereochemical analysis, and to prove this to be the case by applying it to a great number of substances to obtain information on solute geometry that was not readily accessible or not available at all from other techniques. In his 1965 chapter in Advances in Physical Organic Chemistry, he was able to conclude that:

The majority of the procedures currently being used in the confirmational analysis of solutes (e.g. infrared absorption differences between conformers, optical rotatory dispersion, NMR proton shifts, etc.) are qualitative and based upon empirical observations and analogies. It is therefore claimed that the present applications of anisotropic polarisabilities, built as they are on the theoretical arguments of Lorentz, Lorenz, Langevin, Born, Gans, Debye, and others, have – where solutes are concerned – advantages both in their foundations and in the quantitatively expressible natures of the conclusions they can provide.

In many of the 'confirmational papers' quoted above, evidence was presented for various electronic effects in molecules. As well, studies were made specifically to probe electromeric interactions, usually by seeking correlation between the observed polarisability exaltation and the direction of the delocalisation pathway, in aniline, toluene, t-butylbenzene, benzotrichloride and their para-substituted derivatives). In other work, Le Fèvre and J.M. Eckert used structure determination by the Kerr effect to examine steric courses for the replacement of hydroxyl by chlorine in cis-2-decalol. Also they were able to apply the Kerr effect to the conformational analysis of natural product derivatives such as those of cholesterol, tropine and y-pelletierine. Some aspects of this were followed up, with C.Y. Chen, in a series of ¹H NMR and IR spectral investigations of compounds with 6-membered heterocyclic systems. The observation in some instances of non-linear relationships between solute concentration and the Kerr effect or the dielectric polarisation led to investigations that showed solute-solute intermolecular associations of particular geometry to occur in solutions of benzyl alcohol, aniline and substituted anilines, normal alcohols, triisopropanolamine borate, tetrabutoxytitanium, carboxylic acids, and mercury(II) chloride. The Kerr effect technique was applied also to determining overall polarisability anisotropies and the morphologies of macromolecular species dissolved in non-polar media. Other parts of the programme involved studies of possible conformational changes in flexible molecules with variation of the dielectric characteristics of the medium; the influence of solvent on the apparent solute molar Kerr constant; wavelength dispersion of the Kerr constant; polarisabilities of non-bonding electron pairs; the temperature-independent component of the Kerr constant; and dispersion of dielectric absorption in the microwave region as applied to problems of 'anomalous atomic polarity', molecular structure and rigidity. Working with G.L.D. Ritchie, a more sensitive version of the apparatus for solutions was constructed and applied to measuring Kerr constants of gases and to deriving molecular and bond polarisabilities appropriate to this state. Other developments in the experimental technique, carried out with R.K. Pierens, involved the use of sinusoidal or of pulsed voltages – the latter found particular application in the measurement of the Kerr constants of weakly conducting solutions of substances such as keto-enol mixtures.

Other issues were to emerge from the main theme of Le Fèvre's research. One such was the application of the Kerr effect to investigating solvation. It had been noted in the earlier work that solute Kerr constants from benzene solutions sometimes differed markedly from those obtained from carbon tetrachloride or cyclohexane. This was eventually attributed to time-averaged non-random packing of the anisotropic benzene molecules about the solute, a notion supported by NMR evidence at the time. Le Fèvre and his colleagues used observed aromatic solvent-induced changes of the solute Kerr constant, with concomitant ¹H NMR, dipole moment and dielectric absorption data, to specify some stereochemical aspects of the interactions of polar molecules in these solvents. A somewhat similar approach was applied to studying p-hydrogen bonding as found for fluoroform in benzene, p-p-donor-acceptor complexes, and co-ordination of vanadylacetylacetonate with dioxan.

Concurrently with the above programmes, Le Fèvre launched a study in the early 1960s of the magnetic birefringence of molecular diamagnetic substances. The magnetic analogue of the Kerr effect, the Cotton-Mouton effect, was known to be related by theory to the molecular polarisability and magnetisability. A procedure was developed whereby Cotton-Mouton constants from solution could be used together with analogous Kerr effect data to obtain solute molecular magnetic anisotropies and, in many cases, the principal molecular magnetisabilities. The examination was undertaken of a large number of aromatics, primarily substituted benzenes, polynuclear hydrocarbons, heterocyclic compounds, and quinones. In cases where comparisons could be made, reasonable agreement was usually found with literature values of solid-state magnetic anisotropies obtained by crystal-torsion or crystal-oscillation methods. Indeed, one of the attractions of the technique to Le Fèvre was that it provided an independent and accessible physical property from which it was sometimes possible to derive polarisability data complementary to those from the Kerr effect. Trends in magnetisabilities were observed and interpreted in terms of a tentative bond magnetisability scheme.

A prime motivation was the hope that the magnetic anisotropies from experiment would provide a quantitative measure of electron delocalisation and, by inference, of aromatic character. Studied also was the dependence of solute Cotton-Mouton constants on benzene solvation. Le Fèvre's final publication in this series, before he retired from his Sydney post, reported the investigation of about forty pure aliphatic liquids and was concerned with the relationship between pure liquid and solution-state molar Cotton-Mouton constants.

More than seventy research students and colleagues were associated with Le Fèvre in studies of the Kerr and Cotton-Mouton effects and Rayleigh scattering. They included the following: C.G. Le Fèvre, M.J. Aroney, R.S. Armstrong, G.L.D. Ritchie, R.K. Pierens, J.M. Eckert, A.J. Williams, K.E. Calderbank, B. Purnachandra Rao, D.V. Radford, R. Bramley, L. Radom, P.J. Stiles, B.J. Orr, P.H. Cureton, E.P.A. Sullivan, D.S.N. Murthy, C.Y. Chen, J.D. Saxby, K.M.S. Sundaram, A. Sundaram, K.R. Skamp and L.H.L. Chia.

Le Fèvre's Sydney work of the 1950s and 1960s greatly enhanced his reputation as a physical-organic chemist of international standing. His election in 1959 as a Fellow of the Royal Society bears testimony to the distinction of 'his research in organic and physical organic chemistry'. It is pertinent to quote from Sir Christopher Ingold's preface to the Festschrift compiled in 1970 by colleagues of Professor Le Fèvre to celebrate his sixty-fifth birthday. Ingold says:

Le Fèvre's work on molecular electric anisotropy is so outstandingly original, so complete in its grand conception, such a major creation in physical organic chemistry...

It is remarkable that all that we now know of the electronic polarisability of molecules as a function of direction comes almost entirely from the work of the Sydney School. The unique quality of the record, as well as its massive character, is a matter for congratulations...

Professional and community activities

Le Fèvre's prodigious efforts in research appeared not to detract from his attention to other responsibilities. He was acknowledged to be a conscientious and able administrator. That he was a teacher of quality is undoubtedly true. His lectures and seminars were those of the enthusiast, yet presented with sincerity and style – a reflection of the man himself. Le Fèvre was philosophically committed to a specialist first-year teaching group (to be supplemented and assisted by other, more research-orientated, members of staff) to cater for the needs of between 1,500 and 2,100 first-year students about three-quarters of whom were in service courses for Faculties other than Science. The varied and complex needs of these students were for the period of Le Fèvre's tenure well catered for by the teaching group, initially under the directorship of J.J. Broe and later under A.J. Harle. Le Fèvre's administrative headship of the Chemistry School, spanning twenty-two years, encompassed a number of difficult periods, beginning with the tearing down of emotional and physical barriers to departmental amalgamation in 1948, surviving in the early years an era of great acrimony (some of it in the public arena), and negotiating periods of very rapid growth and of severe economic stringency. What cannot be denied is that within a few years of his arrival, Sydney was flourishing as a centre of chemical learning and postgraduate research. Perhaps this is the best testimony to Le Fèvre's achievements.

He found time also for participation in extra mural academic and professional activities. He was a Fellow of the Royal Institute of Chemistry and of the Royal Australian Chemical Institute, serving as Vice-President and New South Wales Branch President of the latter and being its Smith Medallist for 1952. He was a Foundation Fellow of the Australian Academy of Science (1954) and was one of the Petitioners to Her Majesty the Queen for a Royal Charter. Le Fèvre became a member of the first Council of the Academy. Before emigrating, he had served two periods of office on the Council of the Chemical Society in London. He twice served as a member of the Council of the Royal Society of New South Wales, 1948-51 and 1961-74; in 1960 he was the Liversidge Lecturer and in 1961 the President of this Society. Of historical importance was a chapter titled 'The Establishment of Chemistry within Australian Science – Contributions from New South Wales' in A Century of Scientific Progress, the centenary volume of the Royal Society of New South Wales, published in 1968, in which he traces the growth of chemistry in New South Wales from the establishment of the colony to the retirement of Archibald Liversidge in 1907 from the Chair of Chemistry and Mineralogy at Sydney. He was the Society's Medallist in 1969. At the Melbourne meetings of the Australian and New Zealand Association for the Advancement of Science in 1955 and 1967, Le Fèvre was respectively President of Section B and Masson Lecturer. Other distinctions include: Coronation Medallist (1953); Fellow, Queen Mary College, London (1962) Chemical Society of London Lecturer for Australia (1968). He was at various times a member of the following: Australian Journal of Chemistry editorial board, Society of Chemical Industry (Sydney Section) committee, Advisory Committee on Buildings for the Australian National University, Australian Broadcasting Commission Science Panel, Department of Supply and Development Chemical Sub-Committee, and the New South Wales Department of Public Health Pure Foods Advisory Committee. During 1947-48 he was a trustee of the Mitchell Library, Sydney, and a member of the Developmental Council set up by the Minister for Education to form an Institute (or a University) of Technology in New South Wales. He was a trustee of the Museum of Applied Arts and Sciences (1946-75) and a member of the Rotary Club of Sydney from 1948 till his retirement. Sir Mark Oliphant, a physicist and founding member of the Australian Academy of Science, sums up succinctly the feelings of many of Le Fèvre's colleagues:

Raymond was one of the earliest, and most valuable fellows of the Australian Academy of Science. He played a significant part in getting the young Academy off the ground, and in making it respectable, for his standards were high and his knowledge of procedures profound.

Raymond's attitude towards science, and its responsibilities, as well as its benefits, for mankind generally, reflected his caring personality and I learnt much in discussion of such questions with him.

His company, and that of Cathie, were always enjoyable. We looked forward to the occasions when we could meet. He had a true sense of humour, without malice or rancour. He never boasted or sought the limelight, but his stature as a scientist is clear from the fact that he was elected to the Royal Society of London as an Australian chemist.

The memory of Raymond warms my heart, for, to paraphrase a quotation used by Chadwick of Rutherford, 'he was a good man, who did good things'.

Personal

At the age of sixty-three, Le Fèvre suffered a heart attack but was able to resume his duties after a period of convalescence. His love for research was undiminished and it was with great sadness that in 1970, having reached the compulsory retiring age of sixty-five, he was obliged to vacate his Sydney laboratories. Le Fèvre was made Emeritus Professor in 1971 and thereafter he moved to Macquarie University at North Ryde where he continued his research as an Honorary Professorial Fellow. His contributions to the University of Sydney were honoured in 1985, the year of the Science Centenary, by the award of a Doctorate of Science honoris causa.

An extremely modest man, Le Fèvre seemed almost oblivious of honours or esteem accorded him. An exception, perhaps, was his stated appreciation of the unusual honour bestowed on him with his inclusion on the 'wall of fame' at the Technion in Haifa, Israel.

He made generous donations in recent years to, inter alia, Queen Mary College, the University of Sydney, and Macquarie University. In each of these recipient institutions, the decision was made that such funds be used to establish awards to younger chemists.

Raymond Le Fèvre's record of achievement, though great, tells only part of the story. Some other aspects of his life deserve special mention. He was blessed with an extraordinarily happy marriage with Catherine, his wife of fifty-five years. It was a trusting, loving relationship, enhanced by their two children, Nicolette and Ian. They were totally supportive of each other in matters personal, in regard to their family, in times of happiness and of sadness. The tragic loss of their son Ian in November 1977 brought them, if anything, even closer together. It was, to all who knew them, the complete partnership. Catherine's research contributions led to the award to her of a DSc by the University of London (1960). In the 1960s she became very much involved with research issues of social importance such as the drug problem in New South Wales and, later on, with aspects of forensic science.

To generations of Sydney science graduates, Raymond Le Fèvre was an inspiration. He propagated to so many that which he himself felt, the excitement of exploration in science. Many of his past students now occupy senior posts in academia and elsewhere – testimony, in large measure, to his example and encouragement. He will be remembered as a warm, gregarious man with a subtle and gently wicked humour. Above all, he was compassionate and feeling – a man who evoked loyalty and deep affection.

It is fitting that Cathie, in whose arms he died, should have shared his odyssey.

About this memoir

This memoir was originally published in Historical Records of Australian Science, vol.7, no.3, 1988. It was written by:

• M.J. Aroney, School of Chemistry, University of Sydney.
• A.D. Buckingham, University Chemical Laboratory, Cambridge, U.K.

Acknowledgements

The authors are very grateful to Dr Catherine G. Le Fèvre for making available papers dealing with Professor Le Fèvre's early life and wartime experiences, for providing the photograph, and for her helpful comments. We are indebted also to Dr R.S. Armstrong, Professor R. Bonnett, Professor D.P. Craig, F.R.S., Professor H.J.A. Dartnall, Professor M. Davies, Dr J.M. Eckert, Professor P.H. Gore, Mr A.J. Harle, Dr H.G. Holland, Professor L.E. Lyons, Professor Sir Mark Oliphant, F.R.S., Professor B.J. Orr, Dr B. Purnachandra Rao, Professor G.L.D. Ritchie, Professor I.G. Ross, Dr P.J. Stiles, and Professor W.C. Taylor. Most of all, we are indebted to the late Professor Le Fèvre for his beautiful autobiographical material from which we have quoted at length.

Notes

• (1) M.M. Harris, Chemistry and Industry, 1966, 1953; C.K. Ingold, Biographical Memoirs of Fellows of the Royal Society, 14 (1968), 449.
• (2) Contribution to the 1983 Australian Department of Defence account, Mustard Gas Field Trials during World War II, by R.G. Gillis. Australian Government – released for publication, June 1988.
• (3) C.K. Ingold, Structure and Mechanism in Organic Chemistry, Cornell University Press, 1953.

Written by D. J. Adams and P. H. Barry.

• Introduction
• Family background, education and marriage
• PhD at the Australian National University, 1963–1965
• Postdoctoral research at Duke University, North Carolina, 1965–1968
• Academic positions at the University of New South Wales, 1968–1984
• Australian National University, 1984–2005
• Scientific achievements in biophysics and neuroscience
• Service to Australian science
• Awards and affiliations
• Some personal comments
• About this memoir
  • Reference Material and Acknowledgments
  • Bibliography

Introduction

Peter William Gage (1937–2005) was recognised nationally and internationally as one of Australia’s leaders in membrane physiology, biophysics and neuroscience. His research on neurotransmission, muscle and the structure–function of ion channels was extraordinarily productive, with over 7, 000 citations. A gifted speaker with a great enthusiasm for research and for the introduction of cutting-edge technology, Peter Gage influenced and encouraged a great many research students, postdoctoral fellows and senior colleagues in their scientific careers.

Peter Gage died peacefully of myeloid leukemia in Canberra Hospital on 13 August 2005 with his partner, Angela Dulhunty, his sister, Janice Ryan, and his adult children around him. Australia had lost one of its foremost investigators in membrane biophysics and neuroscience. As Bertil Hille commented from the USA: ‘For almost 40 years Peter was a leading practitioner and advocate of membrane biophysics in Australia. He had many students. He was imaginative and brave in his range of work. ’ Peter was awarded a DSc from the University of New South Wales (UNSW) in 1976, elected a Fellow of the Australian Academy of Science (AAS) in 1977, awarded an Australian Research Council (ARC) Centre of Excellence in 1982, awarded the Bob Robertson Medal of the Australian Society for Biophysics in 2004 and elected an Honorary Member of the Australian Physiological Society in 2005. His research on neurotransmission, muscle and the structure–function of ion channels was extraordinarily productive, resulting in over 200 publications and more than 7, 000 citations, with nineteen of his papers receiving more than 100 citations each. He had a great enthusiasm for research and for the introduction of cutting-edge technology to Australia. He was a gifted and dynamic lecturer who received innumerable invitations to speak at national and international conferences. All this, together with the quality of his research, attracted many PhD students, postdoctoral fellows and other senior colleagues to his laboratory. He also contributed greatly to the Australian research community by organizing International Union of Physiological Sciences (IUPS) satellite conferences, patch-clamp workshops, numerous Curtin Conferences over many years, and a GABA 2000 International Symposium in Cairns. In addition, he was a warm and engaging person with a keen sense of humour whose presence will be greatly missed by his many former students, postdoctoral researchers and colleagues, family members and friends in Australia and around the world.

The authors of this memoir, currently holding professorial positions in their respective Australian universities, first met Peter at different stages in their careers. Peter H. Barry (PHB) as a postdoctoral fellow at UCLA first met Peter in the late 1960s, when Peter encouraged him to contact him when PHB wanted to return to Australia. PHB did this and with Peter’s support came from Cambridge University to work with him at UNSW as a Queen Elizabeth II (QEII) fellow in 1972, continuing to work closely with him there until Peter left to go to the Australian National University (ANU) in 1984. David J. Adams (DJA) first met Peter in 1972, as a science student in Peter’s third-year membrane physiology course, becoming his BSc (Hons) student and subsequently his PhD student until he left to go to the USA as a postdoctoral fellow in 1978.

Peter had a special ability for attracting around him students, colleagues and postdoctoral researchers from a range of different disciplines, and for forming successful collaborations with other scientists both within and outside Australia. This was aided by the fact that 1. Peter was invariably involved in cutting-edge research in areas of membrane physiology; 2. he knew the leading international researchers in the field; and 3. he had an outstanding reputation for his own research in these areas. His attributes included considerable practical expertise in electrophysiological experiments and an enthusiasm for advances in instrumentation—for example, the use of operational amplifiers and computers in electrical recording and data analysis, and techniques such as noise analysis and, later, single channel recording, as the latter was being developed. Peter had a solid background in membrane biophysics, together with a good understanding of the physics and mathematics involved in the modelling of processes like neurotransmission, circuit theory and cable analysis, and that was also required for understanding signal analysis and its relevance to ion channel properties. This gave him the ability either to solve problems himself or to encourage, guide and mentor students and postdoctoral researchers to do this. Where more specialist input was required, Peter had the ability to collaborate effectively with specialized biophysicists, engineers and mathematicians. Furthermore, Peter had a great enthusiasm for basic science and research that he communicated to those in his research group. A high proportion of Peter’s research students and postdoctoral fellows have been successful in their careers, greatly helped by his mentoring and training and the quality of his research group. Peter’s international contacts and research reputation were also of considerable value when recommending his students to other leading international laboratories. In addition, research in the field of biophysical and biomedical research of necessity requires the collaboration with research students, postdoctoral researchers and other research colleagues that Peter was able to provide.

Family background, education and marriage

Peter William Gage was born in Auckland, New Zealand, on 21 October 1937 to John Gage, an accountant, and Kathleen Mary Gage (née Burke). He was the third child with two brothers (John and Michael) and one sister (Janice).

Peter was educated at the Sacred Heart College in Auckland and then studied medicine at the University of Otago, being awarded his MB ChB from the University of New Zealand (the only degree-awarding university in New Zealand from 1870 to1961) in 1960. In the same year, he married Jillian (Jill) Christine Shewan, the daughter of James and Carla Shewan, whom he had met while she was studying physiotherapy and he medicine in Dunedin. They had two daughters, Michelle and Jennifer, and two sons, Peter and David, and have eleven grandchildren. Peter did his internship as a house surgeon at Auckland Hospital in 1961 and was a Research Fellow at Green Lane Hospital, Auckland, in 1962.

PhD at the Australian National University, 1963–1965

In early 1963, he and Jill moved to Canberra with their two daughters, for him to undertake a PhD with Professor John Hub-bard in Sir John Eccles’ department in the John Curtin School of Medical Research (JCSMR) at the ANU in Canberra. This was an exciting time to be at the JCSMR, not only because Eccles was awarded the Nobel Prize that year, but also because it was a very dynamic department. Peter’s PhD research on post-tetanic potentiation, post-tetanic hyperpolarization and neurotransmission was extraordinarily productive and was highly cited. From this period, he produced six papers in Nature (3–7, 9), one in the Journal of Pharmacology and Experimental Therapeutics (8) and three in the Journal of Physiology (11–13), with the whole set of papers being cited more than 560 times in other publications. In addition, during this time he had input to two other papers (one in Nature and the other in Vision Research) with Ken Brown.

Postdoctoral research at Duke University, North Carolina, 1965–1968

The Gages, now also with a young son Peter, went to the USA in 1965. Peter (senior) commenced work in Paul Horowicz’s department at Duke University on a prestigious National Institutes of Health International Postdoctoral Fellowship in 1965–7. This was followed by an appointment as Assistant Professor in the same department, 1967–8. During this period he worked with a number of leading electrophysiologists including James R. Bloedel, Clay Arm-strong, Robert (Bob) Eisenberg, Rudolpho Llinas, John Moore, David Quastel and Paul Horowicz himself. In addition, Peter worked on neurotransmission at mammalian and amphibian endplates and, with his colleagues, he also investigated the properties of neurotransmitter release and the ionic nature of the underlying postsynaptic current in voltage-clamp experiments on the squid giant synapse. At least seven papers were published from this research, including two in Nature (15, 22) and one in Science (27), that have obtained more than 270 citations. Peter’s keenness to introduce new techniques is well illustrated by his second Nature paper (22), with Clay Armstrong, in which they recorded synaptic currents at the end plate, and ‘used, then, novel operational amplifiers to build the voltage clamp’, an innovation that paved the way for further developments by Charles F. Stevens and others in their work on understanding synaptic transmission at the endplate [2]. Peter was the first to use these voltage-clamp techniques (22). In addition, Peter published papers in connection with his previous work with David Quastel and Ken Brown.

The impact of all this productive and important research, significant as it was, was, however, somewhat dwarfed by a series of innovative and comprehensive classical experiments. In collaboration with Bob Eisenberg, Peter investigated the role of the transverse tubular system (TTS) in muscle fibres that was then believed (but not yet proven) to be ‘an essential link between the action potential and the activation of the contractile apparatus’ in the fibre (19). In two brief papers in Science in December 1967, Gage and Eisenberg (18, 19) documented their development of a technique, using a glycerol treatment, to electrically isolate the TTS from the surface membrane of muscle fibres. In their microelectrode experiments, they showed that the glycerol treatment, while it did not affect the production of action potentials in the surface membrane of a muscle fibre, had blocked contraction of the muscle and changed the electrical properties of the fibres, radically reducing their electrical capacitance and eliminating a resistance ‘creep’ phenomenon, as would be predicted if the TTS was disrupted (and hence isolated from the surface membrane of the fibres) (18, 19). They followed this up with three detailed and comprehensive papers in the Journal of General Physiology (24–26), that were published in early 1969, to quantify clearly contributions of the TTS and surface membranes to the conductances and capacitance of whole muscle fibres and to confirm the role of the TTS in excitation–contraction coupling in muscle. This was also a breakthrough in muscle electrophysiology, making it possible to dissect out the individual electrical parameters of underlying muscle fibre components and enabling detailed simulation of currents and ionic diffusion within the muscle fibre and its TTS. The importance of this TTS work is well demonstrated by the fact that these five papers alone have been cited over 770 times, underscoring the fact that Peter Gage’s period at Duke University was highly productive and had a significant impact.

Academic positions at the University of New South Wales, 1968–1984

In addition to research, Peter was a committed educator. He had organized and taught a unit on cellular neurophysiology at Duke University, and also held a teaching position during the latter part of his time there. Then, in 1968, he returned to Australia to take up a Senior Lectureship in the School of Physiology and Pharmacology in the Faculty of Medicine at UNSW. He enthusiastically undertook the heavier teaching loads typical of Australian universities compared with those of major US universities. Throughout Peter’s time at UNSW, Walter (Darty) Glover was Head of School (HOS; 1969–85), except when Peter arrived in 1968 when Robert Holland was Acting HOS. During his time in the School, Peter taught both medical and science students. He was very concerned that medical students should be given a strong grounding in basic science and he taught them the biophysics of excitable cells, on topics such as membrane potentials, action potentials (Hodgkin & Huxley equations), muscle and neurotransmission. He also set up a unit in Biophysics for Physiology II for the third-year science students, which provided an excellent background for students wanting to do a BSc (Hons) year in membrane biophysics. On the home front, about two years after the Gage family returned to Australia, Peter’s youngest son, David, was born.

Soon after he arrived at UNSW, Peter was able to establish an active research group and obtain support from the Australian Research Grants Committee (ARGC; the forerunner of the ARC) and the National Health and Medical Research Council (NHMRC). The early members of Peter’s research group included PhD students Angela Dulhunty, Ron Balnave and Robert (Bob) McBurney, a chemical engineer— Dirk Van Helden—and an MSc student, Susan Andrews. Research over the next few years included: 1. measuring the electrical properties of single isolated skeletal muscle fibres and investigating the mechanism of glycerol treatment for disrupting the TTS in those fibres (with Dulhunty: 38, 41, 42); 2. investigating the properties of transmitter release at the toad neuromuscular junction, the temperature dependence of facilitation on release and the inhibitory effect of manganese on release (both with Balnave: 29, 35, 44) and the stimulation of acetylcholine (ACh) release by lithium (with a staff member, John Carmody: 34); and 3. exploring the mechanism of action of blue-ringed octopus toxin to block neuromuscular transmission by blocking presynaptic action potentials (with Dulhunty: 30) and that of miniature endplate currents and potentials, including their simulation, generated by quanta of acetylcholine in toad muscle fibres without transverse tubules (with McBurney: 33, 37, 45).

It should be noted that, early on, Peter had attracted funding to obtain a ‘large’ 4K (word; 8K byte) Digital Equipment Corporation (DEC) PDP8I computer, which was used for data acquisition, theoretical calculations and simulation studies. It was the first computer in the School. An 8K PDP8e was added shortly after for the same cost as an extra 4K of memory for the PDP8I.

Peter was promoted to Associate Professor in 1971. In 1972, Peter Barry (PHB; see Introduction) a QEII Fellow, also arrived to join the group, remaining a collaborator with Peter at UNSW for the rest of Peter’s time there.

In keeping with Peter’s enthusiasm for new technologies and new research projects, he wanted to develop a marine laboratory, similar to the laboratories at Woods Hole in the USA and Plymouth in the UK, in order to work on the giant axon of the squid. To work on squid required setting up a laboratory close to where the squid were caught, so in the early 1970s Peter obtained research funding and set up a small preliminary laboratory in a caravan on a farming property near Shell Harbour, south of Wollongong. The laboratory was established with Dirk Van Helden but, while the idea was excellent, there proved to be logistical problems in doing research so far from Sydney and unfortunately it never quite took off.

As mentioned earlier, Peter’s PhD students generally gained prestigious fellowships and positions in leading international laboratories, in no small measure a reflection of Peter’s international reputation, his international links and the excellent research training imparted to his students. Dulhunty, with her PhD completed in 1973, was awarded a Muscular Dystrophy Association Fellowship at the University of Rochester (USA), to work with Paul Horowicz and Clara Franzini-Armstrong; McBurney, with his PhD also completed in 1973, received a Florey Fellowship to work with Andrew Crawford at the University of Cambridge; Van Helden, with his PhD completed in 1975, was awarded a Ramaciotti Fellowship at UNSW in 1976–7, followed by a Nuffield Travelling Fellowship to work with Bernhard Frankenhaeuser at the Karolinska Institute in Stockholm and with Richard Keynes at the University of Cambridge.

David Adams (DJA; see Introduction) joined Peter’s research group as a BSc (Hons) student in 1973, before starting a PhD in 1974. He worked with Peter on the electrophysiology of Aplysia neurons, investigating various monovalent and divalent ionic currents, conductance changes, gating currents, and the effects of depressant drugs, obtaining his PhD in 1978 (52, 54, 57, 59, 63, 72–74, 81, 91; including two papers in Nature, one in Science and four in the Journal of Physiology). He received a Muscular Dystrophy Association Fellowship to work with Bertil Hille at the University of Washington in Seattle and a Beit Memorial Fellowship at University College, London. Dirk Van Helden, under Peter, developed computer programs to use the technique of noise analysis to determine indirectly the conductance and lifetime of single ionic channels in biological membranes. At this time, Owen Hamill had arrived in Sydney with a BSc from Monash University to do a PhD with Peter. He worked on synaptic transmission investigating the effects of general anaesthetics and ethanol (e. g. 49, 54, 59). Van Helden, with some early input from Hamill, used the noise analysis technique to investigate the effect of permeant cations and some anaesthetics on the lifetime and conductances of ACh endplate channels in muscle fibres (e.g. 60, 64, 67, 69–71, 75, 76, 79). In 1978, Van Helden left for the Karolinska Institute, returning in 1980 on a QEII Fellowship. Hamill completed his PhD in 1977 and after working as a Postdoctoral Fellow with Peter at UNSW (1978–9), obtained a Von Humboldt Fellowship to work at the Max-Planck-Institut für biophysikalische Chemie in Göttingen, Germany, with Erwin Neher and Bert Sakmann on the patch-clamp technique (for which Neher and Sakmann were awarded the Nobel Prize in 1991).

In 1976, following an invitation to write a major review on ‘Generation of end-plate potentials’ for Physiological Reviews (55; to date cited 164 times), Peter was awarded a DSc from the UNSW and was promoted to a personal chair in the School of Physiology and Pharmacology. The following year, 1977, he was elected a Fellow of the AAS.

In the late 1970s, Peter stimulated the interest of the physical chemist Ray Golding in physiological problems, and they co-supervised a physical chemistry student, Tatsanee Mallanoo, in a PhD project on a physico-chemical analysis of synaptic transmission (1978). Peter also supervised the anaesthetist Thomas Torda’s MD project to investigate the effects of some anaesthetic drugs on neurotransmission at the mammalian neuromuscular junction (1978) (53, 62, 78). In the early 1980s, a PhD student from Canada, Ken Takeda, came to work with Peter and subsequently, together with PHB, investigated the effects of nitrate ions, divalent ions and external sodium concentration on the properties of endplate ACh channels (80, 90, 93). Takeda obtained a postdoctoral position in the Centre National de la Recherche Scientifique (CNRS) at Gif-sur-Yvette, France, and subsequently a permanent academic position with the CNRS at Strasbourg. Around this time, Peter also supervised the PhD project of an electrical engineer, Nick Datyner, on the control of acetylcholine secretion at mammalian motor nerve terminals (1981) (83, 94). Over this period, Peter had also been particularly interested in setting up the new technique of patch-clamping that was being developed in Germany by Neher and Sakmann. This directly measured the current in pA (10^-12 A) passing through a single protein channel, and was thus able to determine the conductance and open duration of these channels directly, rather than inferring them indirectly by noise analysis. Through his international connections Peter was able to obtain the electronic circuits, generously supplied by Erwin Neher, to build the appropriate amplifier. A new PhD student, Nino Quartararo, jointly supervised by PHB and Peter Gage in PHB’s laboratory, then successfully built a patch-clamp amplifier and electrophysiology set-up, thus initiating patch-clamp measurements in Australia.

By this time, Peter had built up a large research facility that also included PHB’s laboratory. It comprised a PDP 11/34 computer for data analysis and theoretical studies, eight fully equipped electrophysiological laboratories with three PDP 11/03 computers for online data acquisition and analysis, and the older PDP8e and PDP8I computers. In this period Peter also attracted a number of international visitors who spent their study leave in his laboratory, such as Nancy Lane-Perham (1972) (40) from the University of Cambridge, James McLarnon (1978), Peter Vaughan (1979) from the University of British Columbia, John Mac-Donald (1980) from the University of Auckland, Alan Harvey (1979–80) (85) from the University of Strathclyde and Paul Adams (1981–2) from the State University of New York, Stony Brook. There were also two postdoctoral fellows, Dirk Van Helden who had returned as a QEII Fellow, and a Muscular Dystrophy Association Fellow, Ruth Wachtel from Duke University, as well as four PhD students, an MD student, an MSc student and a BSc (Hons) student, an electrical engineer as an NHMRC Research Associate, a senior Technical Officer, Rodney Malbon, an ARGC Student Assistant and some other support staff.

In 1982, Peter was awarded one of the first ARC Centres of Excellence, the ‘Nerve Muscle Research Centre’, to investigate the normal and abnormal function of nerve and muscle and the signal transmission between them. The staff included three full-time permanent academic research staff (Peter as Director, PHB as Assistant Director and Angela Dulhunty, seconded from the University of Sydney; Fig. 1). New postdoctoral fellows over the next couple of years included Graham Collingridge (on leave from the University of Bristol) and Graham Lamb from the University of Melbourne; new PhD students included Brian Robertson from the UK, W. Roland Taylor, David McKinnon, Pankaj Sah, Chris French and Gavin Schneider, together with BSc Hons and BMedSci students and a number of other support staff including a computer programmer, two electrical engineers and an administrative assistant. The new facilities included a faster and more powerful PDP 11/44 computer to enable development of programs and data analysis to cope with twelve active research projects.

Some of Peter’s collaborative research highlights at the Centre included the following:

1. Single channel currents were successfully recorded and analysed by Nino Quartararo, and later by David McKinnon, in intact denervated single muscle fibres (e.g. see fig. 7 in 104, 106, 121), for what was arguably the first time in Australia. The laboratories subsequently helped to train researchers in other Australian universities in this technique.
2. Collingridge and Robertson began using acute brain slices for electrophysiology, a technique that was then practised in only a few laboratories worldwide, and successfully recorded spontaneous inhibitory postsynaptic currents in neurons in hippocampal slices using a single electrode voltage-clamp system (98, 101) and demonstrated that some general anaesthetics prolong the inhibitory currents (107). Peter’s group was the first to record such currents and use them to examine the effects of general anaesthetics on GABAergic synaptic transmission.
3. Schneider started to record some very large chloride currents in cultured pulmonary alveolar cells, which would later lead to the observation of equally spaced current sublevels and a demonstration of ‘co-channel’ behaviour (116, 163).
4. Chris French observed an interesting slow inward current in rat hippocampal neurons, which was blocked by tetrodotoxin. It was subsequently shown to be a persistent sodium current (108, 132) that plays a role in hypoxia (166).
5. Peter and colleagues set up a three-microelectrode end-of-fibre voltage-clamp system and were the first to record asymmetrical charge movement in mammalian muscle fibres. There were some highly cited papers by Dulhunty and Peter Gage (e.g. 96, 105) on measurements showing a close link between asymmetrical charge movement across muscle membranes and activation of contraction in muscle in normal and paraplegic rats, and an interesting ultrastructural observation by them of internal cysternae of muscle fibres in normal and paraplegic rats (97, 102) that were later identified with crucial elements in the link between depolarization of the muscle tubular system and release of calcium from the sarcoplasmic reticulum that directly enables muscle contraction.

In 1984, after his productive tenure at UNSW, Peter was appointed as Professor and Head of Physiology in the JCSMR at the ANU, with an arrangement worked through with the Director of the JCSMR, Professor Bob Porter. Under the arrangement, Peter brought with him a support staff member and a number of his students, and the ANU provided some postdoctoral fellowships and a senior academic position. Peter was also able to take a significant amount of his equipment, including the PDP 11/44 computer and many of the electrophysiological set-ups, to the ANU.

Peter took with him the laboratory administrator, Rodney Malbon, six PhD students—Robertson, Sah, Taylor, McKinnon, French and Schneider—and a postdoctoral fellow, Graham Lamb (funded initially by UNSW and then on an ANU fellowship). Peter was joined in Canberra by other new ANU postdoctoral fellows—Alasdair Gibb from the UK, Mauri Krouse from the USA and Kevin Buckett from the UK. In addition, Dulhunty was the successful academic applicant for the position at the JCSMR. PHB’s laboratory remained at UNSW with the PDP 11/34 computer and some additional equipment. Nino Quartararo completed his PhD on ion channel measurements and continued for a few years with postdoctoral research in that laboratory as it subsequently expanded independently. It is clear that while Peter’s move from UNSW meant that the Nerve-Muscle Research Centre now ceased to exist as a funded unit, the core of its personnel had moved to the JCSMR and the research that had been stimulated and expanded during its lifetime continued not only with the large group at the ANU but also in PHB’s laboratory at UNSW, with good collaboration between the two groups.

On the personal side, Peter and Jill separated in the early 1980s, some years before his move to the ANU (the marriage being officially dissolved in 1991), and from that time on until the end of his life, his partner was Angela Dulhunty, his research colleague.

Australian National University, 1984–2005

Peter’s period at the ANU was marked by major academic achievements. He had been appointed to the John Eccles Chair of Physiology and Head of the Department of Physiology. The Physiology Department was dissolved in 1988 and replaced by the Division of Neuroscience. Peter remained in that Division for several years and then transferred to the Division of Biochemistry and Molecular Biology in 1999 and was a part of the Membrane Biology Program at the ANU. Peter’s science continued to flourish at the ANU and he made a major contribution to Australian physiology and biophysics, holding two very successful Patch Clamp Workshops at the JCSMR between 1986 and 1989 and then a series of Curtin Conferences at Canberra Grammar School from 1995 to 2003. These conferences were re-established in 2009 by the Ion Channels and Transporters community in his honour.

Peter’s work between 1984 and 1990 was in part a continuation of a number of projects begun at UNSW, with PhD student and postdoctoral fellows who transferred to the ANU with him. This work included an extensive study of post synaptic currents in the hippocampus, with important work on anaesthetic effects on these currents, with C. French, Robertson, Sah, Gibb and Frances Edwards (107, 108, 110, 123, 125, 127, 128). Fran Edwards (123) started as a PhD student with Peter for a couple of years before she transferred her ANU scholarship to Germany for personal reasons, to complete her PhD with the subsequent Nobel prize winner, Bert Sakmann.

Peter continued the first single-channel work to be performed in Australia, which started at UNSW, with an analysis of anion channels in pulmonary epithelia with Schneider and Krouse in a strong collaboration with John Young and David Cook from the University of Sydney. This research included some ground-breaking observations and analysis of multiple conductance levels in the channels (111) and led to a publication in Nature (113). The novel and productive studies of gating charge movement and excitation–contraction coupling in mammalian muscle continued with Dulhunty, Lamb, Bruce Wakefield and a collaboration with Ian Neering and Martin Fryer at UNSW (115, 118, 122, 126, 129, 130). The long-term collaboration with PHB and Quartararo on ion permeation also continued into this period (121). A stream of visitors to the ANU during this time included Florian Dreyer (Germany), David Leaver (Australia) (124), John Macdonald (NZ), Bob Martin (USA), Sue Pockett (NZ), Bob French (Canada), Joe McArdle (USA) (133) and Pei-Hong Zhu (China), together with a collaboration with David Hirst (151) from the University of Melbourne.

Peter began an extensive collaboration with the physical chemist/theoretical biophysicist Shin-Ho Chung and the mathematician John Moore at the ANU, which led to the development of the Hidden Markov Model (HMM) for high-powered analysis of single-channel currents and definition of the probability of small subconductance levels that were difficult to distinguish from background noise (136). Other collaborators in this work included Louis Premkumar and Derek Laver (134, 136, 141, 146, 163, 169). HMM was also used to illustrate a novel phenomenon of coupled gating between channels in hippocampal neurons (137). This important, innovative model has been used by numerous investigators in Australia and overseas and was developed commercially. The model was applied to choride channels in the sarcoplasmic reticulum of skeletal muscle in a collaboration with Dulhunty and Laver (153).

Another discovery by Peter’s laboratory during the 1980s was the persistent sodium current that underlies the massive increase in intracellular calcium during ischaemia in cardiac myocytes and neurons. This collaborative work included an extensive characterization of the current and its sensitivity to the redox environment started with Lamb and Wakefield (130). It remained a mainstream interest in the laboratory over the following fifteen years with collaborative contributions from Buckett, French and Sah (132), David Saint and Yue-Kun Ju (135, 140, 147, 151, 156, 158) and Anna Hammarström (166, 173, 179, 184, 185, 192).

The late 1980s to early 1990s also resulted in the beginning of a long-term project with GABA-induced currents and GABA_A receptors. This work included Peter’s collaborations with research assistant John Curmi, and PhD students and postdoctoral fellows including L. Premkumar, Chung, Saint, Bryndis Birnir, Louise Tierney, Julie Dalziel, Andrea Everitt, Michelle Lim and Mansoureh Eghbali (134, 135, 143, 144, 148, 149, 161, 162, 175, 176, 178, 182, 187, 188, 201). To pursue these studies in greater detail and to take advantage of the rapidly evolving application of molecular biology techniques to ion channels in the 1990s, Peter began a rewarding and fruitful collaboration with Graeme Cox, a biochemist and molecular biologist at the JCSMR. They expressed GABA_A receptors, investigated the role of various subunits in determining the characteristics of GABA_A receptor channels and undertook extensive mutation analysis to understand the conductance and gating of these channels and the molecular determinants of their drug responses, including anaesthetic actions. This work is continuing at the JCSMR under the direction of Louise Tierney. The first publication in this extensive work was in 1992 (142) and the most recent publication was in 2008 (202), with numerous communications in between, including a paper in Nature. The work has been in collaboration with numerous PhD students and postdoctoral fellows, including Tierney, Birnir, Lim, Eghbali, Susan Howitt, Brett Cromer and Tien (Cindy) Luu (152, 159, 162, 164, 167, 168, 171, 174, 177, 190, 196, 198). This novel research led to a paradigm shift in understanding ion channel conductance by showing that single channel conductance could vary with ligand and drug application. This resulted in Peter developing the concept that channels could associate and synchronously gate together to produce variable conductance units, and predicting that the increased conductance of such resultant ‘channels’ under certain circumstances may be due to the direct interactions between the individual channels. This hypothesis has subsequently been supported by experiments indicating that proteins like GABARAP, which facilitate channel clustering, also lead to high conductance state ‘channels’ (198).

The early 1990s produced the unprecedented discovery of viral ion channels, which also formed a major research area for Peter’s group over the following fifteen years and led to two patents. This research was also with Graeme Cox, together with Gary Ewart, Anita Premkumar, Sabine Piller, David Jans, Patricia Jans, Julian Melton, Lauren Wilson, Philip Board and Eric Gowans, a virologist from the Burnet Institute in Melbourne (154, 155, 157, 165, 170, 172, 183, 185, 186, 188, 191, 194, 197, 199). The viral ion channel research formed the impetus for setting up the biotechnology company Biotron. Drugs developed by the company and based on the ANU research have been shown to block viral ion channels and prevent viral replication. These drugs are currently in phase Ib/IIa clinical trials for the hepatitis C virus.

Finally, another area of research for Peter opened up in the early 2000s with the remarkable discovery that glutathione transferases and CLIC proteins are novel modulators of ryanodine receptor ion channels and form one of the few endogenous inhibitors of the cardiac ryanodine receptor. This work, which began in collaboration with Dulhunty and Board, and with Pierre Pouliquin and Yasser Abdelatiff (181, 193, 195, 200), is continuing at the ANU. Other research collaborators with Peter not mentioned elsewhere over this recent period at the ANU included postdoctoral fellows Nesrin Oszarac, N. P. Pillai (152, 159, 160) and Weiping Wu, PhD students Steven Weiss, Rolla Khorie and Victoria Seymour, and research assistant Terry Sutherland (157).

During the period 1998–2005, Peter was instrumental in developing the biotechnology company Biotron. Figure 2 shows Peter’s elation when Biotron started to trade on the Australian Stock Exchange. He was passionate in his belief that research should be developed commercially. The company was originally developed in collaboration with a business partner, Peter Scott, and included JCSMR researchers Chris Parish, Board, Dulhunty and Cox. The company was eventually listed on the Australian Stock Exchange and Peter remained a Board member until his death in 2005.

Scientific achievements in biophysics and neuroscience

Throughout his research career, Peter’s work on neurotransmission, skeletal muscle and the structure and function of ion channels was highly productive. Some particularly noteworthy areas of his collaborative research include:

• with Bob Eisenberg (24–26), the use of glycerol treatment to isolate and investigate the electrical properties and role of the TTS in muscle, continued with Angela Dulhunty (41, 42), and extended to the generation of end-plate potentials in muscle (33, 37, 55)
• showing that general anaesthetics prolong inhibitory postsynaptic currents in hippocampal neurons (107)
• with Dulhunty, being the first to measure asymmetric charge movement in mammalian muscle, which showed a fibre-type dependence (96), and elucidating the role of the dihydropyridine receptor in skeletal muscle excitation–contraction coupling (126)
• suggesting that activation of ion channels in pulmonary alveolar cells was coupled (163)
• with Peter Barry describing cation permeability through acetylcholine receptors (68, 71, 76, 80, 90, 93, 104)
• investigating asymmetrical charge movements in neurons with David Adams (52, 63, 74)
• with Chris French characterizing the persistent sodium channel (108, 132), later showing it to be increased in hypoxia, and reversed by reducing agents
• showing that a series of virus proteins also formed ion channels in lipid bilayers (154, 155, 157) and that some of them could also form channels in neurons, depolarizing and killing them (165, 170)
• uncovering spontaneously opening GABA_A channels in neurons that could be modulated by drugs (e.g. 158, 159, 175, 176, 178, 182)
• showing that members of the glutathione transferase structural family modulate ryanodine receptor Ca²⁺ channels (181)
• showing, in a collaborative project with plant biologists, that a wheat protein that has anti-bacterial and anti-fungal activity, forms ion channels (180).

Overall, Peter’s research resulted in more than 200 publications, the majority in high-impact journals (including sixteen in Nature and three in Science), that included a large number of invited reviews and invited book chapters. His work was cited well over 7,000 times, with nineteen publications receiving over 100 citations each. He also held numerous research grants, mainly from the NHMRC and ARC.

Service to Australian science

Peter introduced many cutting-edge electrophysiological techniques to Australia. He established his laboratory as a central resource for training other researchers, and his organization of annual Curtin Conferences in Canberra on ion channels from 1995, various Patch Clamp Workshops and the GABA 2000 International Symposium reinforced this role. The new techniques included: voltage-clamping with operational amplifiers; the three-electrode voltage clamp for measuring asymmetric charge movement in muscle (with Dulhunty); the patch-clamp technique for directly studying ion channels (with PHB and Nino Quartararo); and (with members of his group at the ANU) the hippocampal slice technique to study synaptic currents.

Peter had more than thirty successful PhD students, many of whom went on to establish international reputations, and at least twenty-two postdoctoral fellows. He also attracted numerous distinguished senior scientists from around the world who visited and collaborated with his group at both UNSW and the JCSMR. Peter was an excellent speaker with innumerable invitations to speak at national and international conferences. His founding role in Biotron, and on its Board, together with two patents of his own, will be of continuing value in encouraging the commercialization of basic research discoveries in Australia.

Awards and affiliations

• In 1977, Peter was elected a Fellow of the Australian Academy of Science. He served on the Council (1983–6) and was Vice-President in 1985–6.
• In 1976, he was awarded a DSc from the UNSW.
• In 1982, he was awarded one of the first ARC Research Centres of Excellence, the Nerve–Muscle Research Centre at UNSW.
• In 2004, he was awarded the Bob Robertson Medal of the Australian Society for Biophysics, named in honour of Sir Rutherford (Bob) Robertson, in recognition of Peter’s outstanding contributions to the field of biophysics in Australia, his contributions to the Society— of which he had been a member for many years—and to Australian science in general. Because he was too ill at the last moment to have it presented at the Society’s meeting, it was awarded a few months later by the Society’s then President, Peter Barry, at a special presentation in the JCSMR with Judith Whitworth, the Director (Fig. 3).
• In 2005, Peter was elected an Honorary Member of the Australian Physiological Society (AuPS), formerly the Australian Physiological and Pharmacological Society (APPS), of which he had been a member since 1964, Treasurer in 1973–5 and President in 2000–4.
• He was also a member of the International Brain Research Organisation and the Australian Society for Biochemistry and Molecular Biology.

Some personal comments

Peter was a very engaging person with a keen sense of humour, who was always very supportive of research colleagues and staff. He was especially proud of his children— Michelle, a general practitioner, Jennifer, a lawyer, Peter, an aerospace engineer and David, a successful business man—and of his grandchildren. He was also keenly interested in music, in native vegetation regeneration on his hobby farm, and in activities such as tennis, skiing, horse riding and camping. He is survived by his former wife, Jill, their four children and eleven grandchildren, and his subsequent partner, Angela.

About this memoir

This memoir was originally published in Historical Records of Australian Science, vol.20, no.2, 2009. It was written by:

• D. J. Adams. Health Innovations Research Institute, RMIT University, PO Box 71, Bundoora, Vic. 3083, Australia
• P. H. Barry. Department of Physiology, School of Medical Sciences, University of New South Wales, Sydney, NSW 2052, Australia. Corresponding author. Email: p.barry@unsw.edu.au

Reference material and acknowledgments

Reference material includes: Who’s Who in Australia, 2000 [1]; P. W. Gage Curriculum Vitae (2004) [2] and other material accompanying his nomination for the Bob Robertson Award [3]; JCSMR Web-site information (27 September 2004) [4]; information from Gage reprints; information from Nerve–Muscle Research Centre documents; personal information of DJA and PHB and from other colleagues; information from and discussions with Jill Gage; and considerable input from Professor Angela Dulhunty, especially on Peter’s research and activities at the ANU. The portrait photograph is of Peter Gage in 2001, courtesy of Multimedia, JCSMR, ANU.

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101. Collingridge, G. L., Gage, P. W., and Robertson, B. (1984). Inhibitory postsynaptic currents in rat hippocampal CAI neurones. J. Physiol. 356, 551–564.
102. Dulhunty, A. F., Gage, P. W., and Valois, A. A. (1984). Indentations in the terminal cisternae of denervated rat EDL and soleus muscle fibers. J. Ultrastr. Res. 88, 30–43.
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105. Dulhunty, A. F., and Gage, P. W. (1985). Excitation-contraction coupling and charge movement in denervated rat extensor digitorumlongus and soleus muscles. J. Physiol. 358, 75–89.
106. Gage, P. W., and McKinnon, D. (1985). Effects of pentobarbitone on acetylcholine activated channels in mammalian muscle. Br. J. Pharmacol. 85, 229–235.
107. Gage, P. W., and Robertson, B. (1985). Prolongationof inhibitory postsynaptic currents by pentobarbitone, halothane and ketamine in CAl pyramidal cells in rat hippocampus. Br. J. Pharmacol. 85, 675–681.
108. French, C. R., and Gage, P. W. (1985). A threshold sodium current in pyramidal cells in rat hippocampus. Neurosci. Lett. 56, 289–293.
109. Gage, P. W. (1985). Ion channels and signal transmission in animal cells. (Invited lecture. )Proc. Aust. Physiol. Pharmacol. Soc. 16, 61–77.
110. Sah, P., French, C. R., and Gage, P. W. (1985). Effects of noradrenaline on some potassium currents in CA1 neurones in rat hippocampal slices. Neurosci Lett. 60, 295–300.
111. Schneider, G. T., Cook, D. I., Gage, P. W., and Young, J. A. (1985). Voltage-sensitive, high-conductance chloride channels in theluminal membrane of cultured pulmonary alveolar (type II) cells. Pflügers Arch. 404, 354–357.
112. Gage, P. W., McKinnon, D., and Robertson, B. (1985). The influence of anaestheticson postsynaptic ion channels. In: Molecular Mechanisms of Anesthesia. (Eds S. H. Rothand K. W. Miller. ) (Plenum Press: New York. )
113. Krouse, M. E., Schneider, G. T., and Gage, P. W. (1986). A large anion-selective channel has seven conductance levels. Nature 319, 58–60.
114. Finkel, A. S., and Gage, P. W. (1985). Conventional voltage clamping with two Peter William Gage 1937–2005 251intracellular microelectrodes. In: Voltage and Patch Clamping with Microelectrodes. (EdsT. G. J. Smith, H. Lecar, S. J. Redman and P. W. Gage. ) (American Physiological Society: Bethesda, Maryland. )
115. Dulhunty, A. F., Gage, P. W., and Lamb, G. D. (1986). Differential effects of thyroid hormone on T-tubules and terminal cisternae inrat muscles: an electrophysiological and morphometric analysis. J. Mus. Res. Cell Motil. 7, 225–236.
116. Krouse, M. E., Schneider, G. T., and Gage, P. W. (1986). A large anion-selective channel has seven conductance levels. Nature 319, 58–60.
117. Gage, P. W. (1987). Channels – an introduction. Proc. Aust. Physiol. Pharmacol. Soc. 18, 15–17.
118. Dulhunty, A. F., Gage, P. W., and Lamb, G. D. (1987). Potassium contractures and asymmetric carge movement in extensor digitorumlongus and soleus muscles from thyrotoxic rats. J. Mus. Res. Cell Motil. 8, 289–296.
119. Barry, P. H., and Gage, P. W. (1987). Artificialnerve and muscle. Proc. Aust. Physiol. Pharmacol. Soc. 18, 167–171.
120. Gage, P. W. (1987). Ion channels in nerve andmuscle. Proceedings of the First Congress of the Asian and Oceanian Physiological Societies, Thailand: 99–110.
121. Quartararo, N., Barry, P. H., and Gage, P. W. (1987). Ion permeation through single channels activated by acetylcholine in denervated toad sartorius muscle fibres: effects of alkalications. J. Membr. Biol. 97, 137–159.
122. Fryer, M. W., Gage, P. W., Neering, I. R., Dulhunty, A. F., and Lamb, G. D. (1988). Paralysis of skeletal muscle by butane dionemonoxime, a chemical phosphatase. PflügersArch. 411, 76–79.
123. Edwards, F. A., and Gage, P. W. (1988). Seasonal changes in inhibitory currents in rat hippocampus. Neurosci. Lett. 84, 266–270.
124. Leaver, D. D., Schneider, K. M., Rand, M. J., Anderson, R. M., Gage, P. W., and Malbon, R. (1988). The neurotoxicity of tunicamycin. Toxicology 49, 179–187.
125. Sah, P., Gibb, A. J., and Gage, P. W. (1988). The sodium current underlying action potentials in guinea pig hippocampal CAI neurons. J. Gen. Physiol. 91, 373–398.
126. Dulhunty, A. F., and Gage, P. W. (1988). Effects of extracellular calcium concentration and dihydropyridines on contraction in mammalian skeletal muscle. J . Physiol. 399, 63–80.
127. Sah, P., Gibb, A. J., and Gage, P. W. (1988). Potassium current activated by depolarization of dissociated neurons from adult guinea pig hippocampus. J. Gen. Physiol. 92, 263–278.
128. Gage, P. W. (1988). Ion channels and postsynaptic potentials. Biophys. Chem. 29, 95–101.
129. Dulhunty, A. F., and Gage, P. W. (1989). Effects of cobalt, magnesium and cadmiumions on contraction in rat skeletal muscle fibres. Biophys. J. 56, 1–14.
130. Gage, P. W., Lamb, G. D., and Wakefield, B. T. (1989). Transient and persistent sodium currents in normal and denervated mammalianskeletal muscle. J. Physiol. 418, 427–439.
131. Gage, P. W., Dulhunty, A. F., Lamb, G. D., and Wakefield, B. T. (1989). Effects of denervation on excitation-contraction coupling in mammalian muscle. In: Neuromuscular Junction. (Eds L. C. Sellin, R. Libelius and S. Thesleff. ) pp. 405–412. (Elsevier: Amsterdam, NewYork. )
132. French, C. R., Sah, P., Buckett, K. J., and Gage, P. W. (1990). A voltage-dependent persistent sodium current in mammalian hippocampal neurons. J. Gen. Physiol. 95, 1139–1157.
133. Gage, P. W., McArdle, J. J., and Saint, D. A. (1990). Effects of butanedione monoxime on neuromuscular transmission. Br. J. Pharmacol. 100, 467–470.
134. Premkumar, L. S., Chung, S., and Gage, P. W. (1990). GABA-induced potassium channels in cultured neurons. Proc. R. Soc. Lond. B241, 153–158.
135. Saint, D. A., Thomas, T., and Gage, P. W. (1990). GABA_B agonists modulate a transient potassium current in cultured mammalian hippocampal neurons. Neurosci. Lett. 118, 9–13.
136. Chung, S., Moore, J. B., Xia, L., Premkumar, L. S., and Gage, P. W. (1990). Characterization of single channel currents using digital signal processing techniques based on Hidden Markov Models. Phil. Trans. R. Soc. Lond. B329, 265–285.
137. Premkumar, L. S., Gage, P. W., and Chung, S. (1990). Coupled potassium channels induced by arachidonic acid in cultured neurons. Proc. R. Soc. Lond. B 242, 17–22.
138. Gage, P. W., Chung, S., Moore, J. B., and Premkumar, L. S. (1990). Detection and interpretation of multiple conductance levels in ion channels in membranes. In:Exocrine Secretion II. (Eds P. Y. D. Wong252 Historical Records of Australian Science, Volume 20 Number 2and J. A. Young. ) pp. 53–56. (United League Graphic and Printing Co. Ltd: Hong Kong. )
139. Saint, D. A., Ju, Y., and Gage, P. W. (1992). A persistent sodium current in rat ventricular myocytes. J. Physiol. 453, 219–231.
140. Ju, Y., Saint, D. A., and Gage, P. W. (1992). Effects of lignocaine and quinidine on the persistent sodium current in rat ventricularmyocytes. Br. J. Pharmacol. 107, 311–316.
141. Vasudevan, S., Premkumar, L., Stowe, S., Gage, P. W., Reilander, H., and Chung, S. (1992). Muscarinic acetylcholine receptor produced in recombinant baculovirus infected Sf9 insect cells couples with endogenous G-proteins to activate ion channels. FEBS 311, 7–11.
142. Birnir, B., Tierney, M. L., Howitt, S. M., Cox, G. B., and Gage, P. W. (1992). A combination of human α₁ and β₁ subunits is requiredfor formation of detectable GABA-activatedchloride channels in Sf9 cells. Proc. R. Soc. Lond. B 250, 307–312.
143. Gage, P. W. (1992). Activation and modulation of neuronal K channels by GABA. TINS15, 46–51.
144. Curmi, J. P., Premkumar, L. S., Birnir, B., and Gage, P. W. (1993). The influence of membrane potential on chloride channels activated by GABA in rat cultured hippocampal neurons. J. Membr. Biol. 136, 273–280.
145. Gage, P. W., Premkumar, L. S., and Chung, S. (1993). Influence of GABA on potassium channels in hippocampal neurons. In: Molecular and Cellular Biology of Pharmacological Targets. (Eds H. Glossmann and J. Striessnig. )pp. 165–188. (Plenum Press: New York. )
146. Gage, P. W., and Chung, S. (1994). Influence of membrane potential on conductance sublevels of chloride channels activated by GABA. Proc. R. Soc. Lond. B 255, 167–172.
147. Ju, Y., Saint, D. A., and Gage, P. W. (1994). Inactivation-resistant channels underlying the persistent sodium current in rat ventricular myocytes. Proc. R. Soc. Lond. B 256, 163–168.
148. Premkumar, L., and Gage, P. W. (1994). Potassium channels activated by GABA_B agonists and serotonin in cultured hippocampal neurons. J. Neurophysiol. 71, 2570–2575.
149. Birnir, B., Everitt, A. B., and Gage, P. W. (1994). Characteristics of GABA_A channels in rat dentate gyrus. J. Membr. Biol. 142, 93–102.
150. Gage, P. W., Sunstrom, N. A., Premkumar, L. S., Laver, G. W., and Cox, G. B. (1994). Structure and function of a novel ion channel. In: Studies in Honour of Karl Julius Ullrich: an Australian Symposium. (Eds P. Poronnik, D. I. Cook and J. A. Young. ) pp. 11–14. (Fast Books, Wild &Woolley Pty Ltd: Sydney. )
151. Ju, Y., Saint, D. A., Hirst, G. D. S., and Gage, P. W. (1995). Sodium currents in toad cardiac pacemaker cells. J. Membr. Biol. 145, 119–128.
152. Birnir, B., Tierney, M. L., Pillai, N. P., Cox, G. B., and Gage, P. W. (1995). Rapid desensitization of α₁β₁ GABA_A receptors expressedin Sf9 cells under optimized conditions. J. Membr. Biol. 148, 193–202.
153. Kourie, J. I., Laver, D. R., Junankar, P. R., Gage, P. W., and Dulhunty, A. F. (1996). Characteristics of two types of chloride channel in sarcoplasmic reticulum vesicles from rabbit skeletal muscle. Biophys. J. 70, 202–221.
154. Piller, S. C., Ewart, G. D., Premkumar, A., Cox, G. B., and Gage, P. W. (1996). Vprof human immunodeficiency virus type 1forms cation-selective channels in planarlipid bilayers. Proc. Natl. Acad. Sci. USA 93, 111–115.
155. Sunstrom, N. A., Premkumar, L. S., Premkumar, A., Ewart, G., Cox, G. B., and Gage, P. W. (1996). Ion channels formed by NB, an influenza B virus protein. J. Membr. Biol. 150, 127–132.
156. Ju, Y., Saint, D. A., and Gage, P. W. (1996). Tetrodotoxin-sensitive inactivation-resistant sodium channels in pacemaker cells influence heart rate. PflügersArch. 431, 868–875.
157. Ewart, G. D., Sutherland, T., Gage, P. W., andCox, G. B. (1996). The Vpu protein of human immunodeficiency virus type 1 forms cation selective channels. J. Virol. 70, 7108–7115.
158. Ju, Y., Saint, D. A., and Gage, P. W. (1996). Hypoxia increases persistent sodium currentin rat ventricular myocytes. J. Physiol. 497, 337–347.
159. Tierney, M. L., Birnir, B., Pillai, N. P., Clements, J. D., Howitt, S. M., Cox, G. B., and Gage, P. W. (1996). Effects of mutatingleucine to threonine in the M2 segment of α₁and β₁ subunits of GABA_Aα₁β₁ receptors. J. Membr. Biol. 154, 11–21.
160. Gage, P. W., Birnir, B., Tierney, M. L., Dalziel, J. E., Cromer, B., Pillai, N. P., Howitt, S. M., and Cox, G. B. (1996). Effects on functionof mutations in α₁β₁ GABA_A receptors. In: Studies in honour of John Atherton Young. (Eds A. Dinudom and P. Komwatana. ) pp. 109–114. (University of Sydney Printing Service: Sydney. )
161. Birnir, B., Tierney, M. L., Dalziel, J. E., Cox, G. B., and Gage, P. W. (1997). A structural Peter William Gage 1937–2005 253determinant of desensitization and allosteric regulation by pentobarbitone of the GABA_A receptor. J. Membr. Biol. 155, 157–166.
162. Eghbali, M., Curmi, J. P., Birnir, B., and Gage, P. W. (1997). Hippocampal GABA_A channel conductance increased by diazepam. Nature388, 71–75.
163. Laver, D. R., and Gage, P. W. (1997). Interpretation of substates in ion channels: uniporesor multipores? Prog. Biophys. Mol. Biol. 67, 99–140.
164. Birnir, B., Tierney, M. L., Lim, M., Cox, G. B., and Gage, P. W. (1997). The nature of the 5_residue in the M2 domain affects function of the human α₁β₁ GABA_A receptor. Synapse26, 324–327.
165. Piller, S. C., Jans, P., Gage, P. W., and Jans, D. A. (1998). Extracellular HIV-1 Vpr causes a large inward current and cell death in cultured hippocampal neurons: implications for AIDS pathology. Proc. Natl. Acad. Sci. USA95, 4595–4600.
166. Hammarström, A. K. M., and Gage, P. W. (1998). Inhibition of oxidative metabolism increases persistent sodium current in rat CAI hippocampal neurons. J. Physiol. 510, 735–741.
167. Gage, P. W. (1998). Signal transmission inligand-gated receptors. Immunology and CellBiology 76, 436–440.
168. Tierney, M. L., Birnir, B., Cromer, B., Howitt, S. M., Gage, P. W., and Cox, G. B. (1998). Two threonine residues in the M2 segment ofthe α₁β₁ GABA_A receptor are critical for ion channel function. Receptors and Channels 5, 113–124.
169. Chung, S. -H., and Gage, P. W. (1998). Signal processing techniques for channel current analysis based on Hidden Markov Models. Methods. Enzymol. 293, 420–437.
170. Piller, S. C., Ewart, G. D., Jans, D. A., Gage, P. W., and Cox, G. B. (1999). The amino terminal of VPR from HIV-1 forms ion channels and kills neurons. Journal of Virology 73, 4230–4238.
171. Dalziel, J. E., Birnir, B., Everitt, A. B., Tierney, L. M., Cox, G. B., and Gage, P. W. (1999). A threonine residue in the M2 regionof the β₁ subunit is needed for expression of functional α₁β₁ GABA_A receptors. European Journal of Pharmacology 370, 345–348.
172. Ewart, G. D., Greber, D., Cox, G. B., and Gage, P. W. (1999). Ion channels formed by Vpu, an HIV-1-encoded protein (a potential target for AIDS therapeutic drugs?). Australian Biochemist 30, 11–13.
173. Hammarström, A. K. M., and Gage, P. W. (1999). Nitric oxide increases persistent sodium current in rat hippocampal neurons. J. Physiol. 520, 451–461.
174. Dalziel, J. D., Cox, G. B., Gage, P. W., and Birnir, B. (1999). Mutant human α₁β₁(T262Q) GABA_A receptors are directly activated but not modulated by pentobarbital. Eur. J. Pharmacol. 385, 283–286.
175. Birnir, B., Everitt, A. B., Lim, M. S., and Gage, P. W. (2000). Spontaneously opening ABA(A) channels in CAl pyramidal neuronesof rat hippocampus. J. Membr. Biol. 174, 21–29.
176. Birnir, B., Eghbali, M., Everitt, A. B., and Gage, P. W. (2000). Bicuculline, pentobarbital and diazepam modulate spontaneous GABA(A) channels in rat hippocampal neurons. Br. J. Pharmacol. 131, 695–704.
177. Dalziel, J. E., Cox, G. B., Gage, P. W., and Birnir, B. (2000). Mutating the highly conserved second membrane-spanning region 9_leucine residue in the alpha(1) or beta(1) subunit produces subunit-specific changes in the function of human alpha(1)beta(1) gamma-aminobutyric acid(A) receptors. Mol. Pharmacol. 57, 875–882.
178. Eghbali, M., Gage, P. W., and Birnir, B. (2000). Pentobarbital modulates gamma-aminobutyric acid-activated single-channel conductance in rat cultured hippocampal neurons. Mol. Pharmacol. 58, 463–469.
179. Hammarström, A. K., and Gage, P. W. (2000). Oxygen-sensing persistent sodium channels in rat hippocampus. J. Physiol. 529, 107–118.
180. Hughes, P., Dennis, E., Whitecross, M., Llewellyn, D., and Gage, P. (2000). The cytotoxic plant protein, beta-purothionin, forms ion channels in lipid membranes. J. Biol. Chem. 275, 823–827.
181. Dulhunty, A., Gage, P., Curtis, S., Chelvanayagam, G., and Board, P. (2001). The glutathione transferase structural family includes a nuclear chloride channel and aryanodine receptor calcium release channel modulator. J. Biol. Chem. 276, 3319–3323.
182. Birnir, B., Eghbali, M., Cox, G. B., and Gage, P. W. (2001). GABA concentration sets the conductance of delayed GABA_A channels in outside-out patches from rat hippocampal neurons. J. Membr. Biol. 181, 171–183.
183. Ewart, G. D., Mills, K., Cox, G. B., and GageP. W. (2002). Amiloride derivatives block ion channel activity and enhancement of virus like particle budding caused by HIV-1 protein Vpu. Eur. Biophys. J. with Biophys. Lett. 31, 26–35.
184. Hammarström, A. K. M., and Gage, P. W. (2002). Hypoxia and persistent sodium current. Eur. Biophys. J. with Biophys. Lett. 31, 323–330.
185. Nielsen, K. J., Watson, M., Adams, D. J., Hammarström, A. K., Gage, P. W., Hill, J. M., Craik, D. J., Thomas, L., Adams, D., Alewood, P. F., and Lewis, R. J. (2002). Solution structure of mu-conotoxin PIIIA, a preferential inhibitor of persistent tetrodotoxin-sensitive sodium channels. J. Biol. Chem. 277, 27247–27255.
186. Melton, J. V., Ewart, G. D., Weir, R. C., Board, P. G., Lee, E., and Gage, P. W. (2002). Alphavirus 6K proteins form ion channels. J. Biol. Chem. 277, 46923–46931.
187. Eghbali, M., Gage, P. W., and Birnir, B. (2003). Propofol increases GABA_A channel conductance. Eur. J. Pharmacol. 468, 75–82.
188. Eghbali, M., Birnir, B., and Gage, P. W. (2003). Effects of propofol on GABA(A)channel conductance in rat-cultured hippocampal neurons. J. Physiol. 552, 13–22.
189. Premkumar, A., Wilson, L., Ewart, G. D., and Gage, P. W. (2004). Cation-selective ion channels formed by p7 of hepatitis C virus areblocked by hexamethylene amiloride. FEBSLett. 557, 99–103.
190. Everitt, A. B., Luu, T., Cromer, B., Tierney, M. L., Birnir, B., Olsen, R. W., and Gage, P. W. (2004). Conductance of recombinant GABA_A channels is increased in cellsco-expressing GABARAP. J. Biol. Chem. 279, 21701–21706.
191. Premkumar, A., Ewart, G. D., Cox, G. B., and Gage, P. W. (2004). An amino-acid substitutionin the influenza-BNBprotein affects ion channel gating. J. Membr. Biol. 197, 135–143.
192. Hammarström, A. K., and Gage, P. W. (2004). Methods to study oxygen sensing sodium channels. Methods. Enzymol. 381, 275–290.
193. Board, P. G., Coggan, M., Watson, S., Gage, P. W., and Dulhunty, A. F. (2004). CLIC-2modulates cardiac ryanodine receptor Ca2+release channels. Int. J. Biochem. Cell. Biol. 36, 1599–1612.
194. Ewart, G. D., Nasr, N., Naif, H., Cox, G. B., Cunningham, A. L., and Gage, P. W. (2004). Potential new anti-human immunodeficiency virus type 1 compounds depress virus replication in cultured human macrophages. Antimicrob. Agents Chemother. 48, 2325–2330.
195. Dulhunty, A. F., Pouliquin, P., Coggan, M., Gage, P. W., and Board, P. G. (2005). A recently identified member of the glutathione transferase structural family http://www. publish. csiro. au/journals/hras modifies cardiac RyR2 substate activity, coupled gating and activation by Ca2+ andATP. Biochem. J. 390, 333–343.
196. Luu, T., Cromer, B., Gage, P. W., and Tierney, M. L. (2005). A role for the 2_ residue in the second transmembrane helix of the GABA_A receptor gamma2S subunit in channel conductance and gating. J. Membr. Biol. 205, 17–28.
197. Premkumar, A., Horan, C. R., and Gage, P. W. (2005). Dengue virus M protein C-terminal peptide (DVM-C) forms ion channels. J. Membr. Biol. 204, 33–38.
198. Luu, T., Gage, P. W., and Tierney, M. L. (2006). GABA increases both the conductance and mean open time of recombinant GABA_A channels co-expressed with GABARAP. J. Biol. Chem. 281, 35699–35708.
199. Premkumar, A., Dong, X., Haqshenas, G., Gage, P. W., and Gowans, E. J. (2006). Amantadine inhibits the function of an ion channel encoded by GB virus B, but fails to inhibit virus replication. Antivir Ther. 11, 289–295.
200. Abdellatif, Y., Liu, D., Gallant, E. M., Gage, P. W., Board, P. G., and Dulhunty, A. F. (2007). The Mu class glutathione transferase is abundant in striated muscle and is an isoform-specific regulator of ryanodine receptor calcium channels. Cell Calcium. 41, 429–440.
201. Gaul, S., Ozsarac, N., Liu, L., Fink, R. H., and Gage, P. W. (2007). The neuroactive steroids alphaxalone and pregnanolone increase the conductance of single GABA_A channels in newborn rat hippocampal neurons. J. SteroidBiochem. Mol. Biol. 104, 35–44.
202. Tierney, M. L., Luu, T., and Gage, P. W. (2008). Functional asymmetry of the conserved cystine loops in alphabetagammaGABA_A receptors revealed by the response to GABA activation and drug potentiation. Int. J. Biochem. Cell Biol. 40, 968–979.

Patents

1. Method for determining ion channel activity of a substance. Provisional patentPO2581/96, 27 September 1996; international patent application PCT/AU97/00638, 26 September 1997.
2. A method of modulating ion channel functional activity. Provisional patent PP6464/9812 October 1998; international patent application PCT/AU99/00872, 12 October 1999.

Peter Hall, in the 40 years of his research career, produced work in both probability and statistics, whose breadth and depth must be regarded as phenomenal. He displayed extraordinary technical skills together with remarkable intuition in developing and applying multifaceted mathematical approaches in the whole of his work. The impact of this wide-ranging use of powerful mathematical methods has had a profound effect on much of modern mathematical statistics. 

After completing his DPhil at Oxford, he remained in Australia for almost all his career although he was renowned as one of the major international figures in probability and statistics. 

Peter was a mentor to a large group of postgraduate students and postdoctoral colleagues giving encouragement and guidance and he attracted many research visitors contributing greatly to the whole of Australian statistical research. 

Remarkably, given his immense research output, he took a significant role in both editorial duties in major international journals and in advocacy for mathematics and statistics in Australia. 

Peter was a man of great charm whose modest demeanour belied his staggering abilities. His loss to mathematics and statistics is great, but is matched by the personal loss to us and to his many friends.

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Corrigendum

The original Supplementary Material 1 and 2 for this paper, available online, contained some errors. Supplementary Material 1 and 2 have now been corrected and replaced online.

About this memoir

This memoir was originally published in Historical Records of Australian Science, vol. 28(2), 2017. It was written by John Robinson and Alan H. Welsh.

Written by F.J. Morgan.

Pehr Victor Edman was born in Stockholm, Sweden, in April 1916 and died in Munich, FRG, in March 1977. He was born into a lawyer's family and received his schooling in Stockholm. In 1935 he began medical studies at the Karolinska Institute and graduated with his primary medical qualifications in 1938. He became interested in research and, following graduation, continued to work at the Karolinska Institute, largely in the laboratory of Professor Eric Jorpes. He appears to have systematically taught himself organic chemistry at this time by extensive reading. During the war years his research was interrupted by a long period of service in the medical corps of the Swedish Army. He was awarded the degree of Doctor of Medicine in 1946. The subject of his thesis was the purification and analysis of angiotensin from bovine blood. His earlier published studies concerned heparin and secretin, which were interests of his mentor Jorpes.

At this juncture Edman began to take the independent research direction which he followed almost uninterruptedly for the rest of his career. He accepted a grant to work for a year in the Northrop-Kunitz laboratory at the Princeton branch of the Rockefeller Institute for Medical Research. Swedish medical research had been isolated during the war and he was anxious to learn of the progress made in the United States. Moreover, his work on angiotensin had made him realize that simple compositional analysis would not be helpful in providing a basis for understanding the biological function of peptides or proteins. The realization that proteins were not colloids but that each had a definite molecular weight and a specific structure was beginning to emerge, especially as a result of the work of the Uppsala group. Edman knew that the order of aminoacids linked by peptide bonds was an essential part of the unique makeup of any given protein. At Princeton he began experiments to try to find a way to chemically decode the aminoacid sequence of proteins.

In the early years of Edman's attempts in this area two general procedures were being used to attack the sequence problem. Various reagents had been found useful in labelling the aminoterminal (or first) aminoacid through its reactive amino group and allowing identification as a derivative. One of these reagents, fluorodinitrobenzene (FDNB), which gave the dinitrophenol (DNP) derivative of the aminoterminus, was used by Sanger in his epochal work on the structure of insulin. By using the FDNB reaction with sets of overlapping peptides derived from partial cleavage of insulin, Sanger, by 1956, was able to deduce a unique structure for the insulin molecule. This was the first primary structure of a protein to be decoded, but despite the undoubted importance of the feat it was clear that the method was too cumbersome to have wide application.

Another reagent used for aminoterminal determination was phenylisocyanate (PIC), introduced for this purpose by Abderhalden and Brockmann in 1930. As with FDNB the hydrolysis to release the aminoterminal aminoacid derivative destroyed many of the other peptide bonds, leaving the remaining protein useless for analysis. In Princeton, Edman realised that if phenylisothiocyanate (PITC) were used the nucleophilic sulphur would weaken the adjacent peptide bond, raising the possibility of finding conditions for its hydrolysis that did not cleave the remainder of the molecule. This remaining peptide could then be subjected to a second reaction with PITC and the second aminoacid determined, and so on theoretically to the carboxyterminal end of the molecule. Whether Edman thought of this solution completely independently, whether some unrelated paper uncovered in his wide reading drew PITC to his attention, or whether some colleague in Princeton or Stockholm suggested its use is not known. In view of the success which the reaction ultimately achieved, the latter seems unlikely in the absence of any claims or reminiscences to this effect. In his review of aminoacid sequencing methods written in 1969 Edman is at pains to stress that the PITC reaction was not at all derived from the earlier PIC reaction, as they had different mechanisms of action. However, in view of the superficial similarities between the reagents and the similar uses to which they were put in protein chemistry, this seems a little strained. By the time Edman returned to Sweden in 1947 he had performed enough experiments to know that the idea was practicable and could form the basis of a protein sequencing technique.

Edman took up an associate professorship at Lund and continued to work almost exclusively on protein degradation. The derivative resulting from the coupling of PITC with an aminoacid, the 3-phenyl-2-thiohydantoin (PTH) aminoacid, proved to be a stable compound in almost all cases. Edman synthesised the PTH derivatives of all the amino acids found in proteins and developed chromatographic systems to identify and quantify them; conditions for the coupling of PITC to the aminoterminus and the cleavage of the PTH derivative which worked smoothly for all peptide bonds were found. After two years work Edman was able to publish the chemical details of a method capable, in theory, of solving the problem of primary structure of proteins and of providing the essential information about innumerable proteins essential for further advances in protein biochemistry. The characteristic ultraviolet absorption spectra of the PTH-aminoacids made them particularly suited to quantitative studies. It permitted useful measurements of the subunit structure and molecular weight of proteins and, in conjunction with column chromatography, it was an alternative to the ninhydrin reaction for aminoacid analysis. However, these possibilities could not be fully exploited until the recent developments in high performance liquid chromatography. The method became widely known and was given the eponym 'Edman degradation' by Kai Linderstrom-Lang of the Carlsberg Laboratories.

In the early 1950s Mr 'Jack' Holt, a well-known Victorian racehorse trainer, died and his will provided that the income from his estate, to be held in trust, was to be used for medical research at St. Vincent's Hospital in Melbourne. By 1956 the hospital authorities had decided to establish a separate research institution in the hospital rather than disburse the funds as research grants to existing hospital units. The decision had also been taken to develop a non-clinical basic science area, preferably biochemistry. The School of Medical Research, as it was originally known, had its own governing board and for practical purposes functioned independently of the hospital administration.

Edman applied for the position of Director of Research. He was clearly the outstanding candidate and moreover his interests coincided with the preference for biochemistry as the focus of the School. In 1957 Edman accepted the offer of the position of first Director of Research at St Vincent's School of Medical Research in Melbourne. His reasons for this move, which were said to have been a mixture of general dissatisfaction with scientific resources in Lund and the impending breakdown of his marriage, must have been strong and not primarily directed to career improvement. He had just accomplished an outstanding piece of individual biochemical research which would have made him welcome in many leading centres in the northern hemisphere. In Australia, in Melbourne, he would be largely isolated from these centres; the School was a new institution, with no traditions, no established workers, nor any support staff; and although situated in a teaching hospital of the University of Melbourne, it had itself no academic or university affiliation. Moreover, Australia at that time was not known for generous government research funding. Despite this formidable array of disincentives Edman decided to move alone to Melbourne to continue his work, initially without trained help and without close colleagues.

In Australia Edman completed a few small projects on other aspects of protein structure that he had begun in Sweden, but otherwise he worked almost entirely on the phenylisothiocyanate (PITC) degradation. This work fell into three phases: improvements in the conditions for the degradation, largely focussed on the elimination of side reactions; 'automation' of the reaction sequence; and application of the degradation to various sequence problems. The latter phase overlapped the other two and usually involved the interests of visiting scientists who had come to Edman's laboratory to learn or use his technique.

By 1960-61 the three-stage degradation reaction had been essentially perfected. Its universal application and repetitive nature suggested to G.S. Begg, Edman's Australian technical assistant, that it would be suitable for automation. Edman realized that the number of existing proteins (about ten million) made manual sequencing an impossible task and was quickly converted to the idea of automation. The close control of reaction conditions possible with automation also gave promise of higher and more constant repetitive yield than were possible manually. High repetitive yields are crucial to repetitive processes, whether synthetic or degradative. Geoffrey Begg had been one of the early technical staff employed by Edman after his arrival in Melbourne. He had no formal qualifications but from a combination of courses at technical college and self-instruction had achieved a remarkable expertise in practical chemistry and glassblowing, mechanical engineering, and electronics. The sequenator project provided a perfect opportunity to use these multiple talents which complemented Edman's academic and theoretical knowledge. Edman and Begg worked as a team on the sequence automation project, with no sustained input from other workers. It was typical of Edman's thorough approach to all tasks that he became a sufficiently adept toolmaker in this period to do much of the fitting and turning himself.

The basis of what was to become the protein sequenator was developed to a prototype stage in a period of a few weeks in the autumn of 1961 – the glass cup spinning on its cylindrical axis, addition of reagents via a catheter, reactions in a thin liquid film on the wall of the spinning cup, and extractions by solvent moving upwards over the film into a groove. Within two years Edman and Begg had built, in their own workshop, a machine capable of reliably carrying out the reactions of the degradation. They had found new conditions and reagents suitable for the physical conditions of the spinning cup; for example, the open cup in general required less volatile chemicals and the narrow delivery and effluent tubes demanded special attention to the surface tension and rheological properties of the solvents. Edman's wide knowledge of classical organic chemistry enabled quick progress in converting the manual reaction to its automated form. In 1964 Edman reported his preliminary findings to a meeting in Scotland. In 1967 in the first issue of the  European Journal of Biochemistry  with Begg as co-author he published his definitive paper demonstrating an unbroken automated determination of the aminoterminal sixty aminoacids of humpback whale myoglobin at the rate of one residue per hour. The extent of this advance can be gauged from the knowledge that at that time the most extensive manual degradation encompassed about fifteen residues at a rate of one per day. Many laboratories could not establish the manual degradation at all, owing to a failure to appreciate the importance of pure reagents in eliminating side reactions. During the next few years Edman's aim was to improve the repetitive yield obtained from the machine; an increase from the 98% of the 1967 paper to 99% was calculated to double the length of determinable sequence. The protein sequenator in Melbourne remained unique until late in 1969 when the Beckman Instrument Company in the United States put on the market a commercial version based on Edman's design. Edman played no part in the commercialisation of his machine. The Board of the School discussed the possibilities of patenting the sequenator but soon accepted Edman's strong view that he should publish fully without patent protection. Edman was elected a Fellow of the Australian Academy of Science in 1968 and a Fellow of the Royal Society of London in 1974. Edman became an Australian citizen in the mid sixties.

In 1972 Edman resigned from St Vincent's School of Medical Research and became Director of Protein Chemistry I at the Max Planck Institute for Biochemistry at Martinsreid near Munich. In 1968 he had remarried; his second wife Agnes Henschen had come from Stockholm and this had given him a reason to think of a return to Europe. In the ten years since his arrival in 1957 the School had remained small, and attempts to raise support for expansion on the basis of the success of the sequenator project proved not very successful. Now, as many years before in Lund, he believed that the importance of his work was not properly recognised and that he would continue to have inadequate resources in Melbourne. A move to the new laboratories of the Max Planck Institute seemed to provide an answer to his needs. Edman set up his laboratory in Munich along the lines of that in Melbourne and with the same aim of increasing the efficiency of the degradation. In addition, with his aid, Agnes Henschen began to make substantial progress in her studies of fibrinogen structure. Sadly, Edman developed a cerebral tumour and died after a short period of coma in 1977.

Edman played little if any role in broader scientific administration or politics in Australia. Although his School had no formal academic affiliation, there is no evidence that he would not have been accepted in these arenas. Some efforts to arrange a personal appointment at the University of Melbourne came to nothing. Thus he remained something of an enigma in the scientific community. He was slow to publish, with approximately one and a half papers per year during his Australian period, which made difficulties for those wishing to implement the method. If his impact on the Australian scene was limited, it was paradoxically the result of his single-minded pursuit of the sequence degradation. Such work, despite Edman's reputation, was not very attractive to students and he never built up a tradition of a flow of graduate students. Once the initial work on the manual or especially the automated reaction was complete, the details would easily have been completed by others in his or other laboratories. One cannot help thinking that his impact would have been so much greater had he seen himself able to move strongly into new areas of protein structure and function. Biological research often requires the appreciation of the importance of an approximate result for advancement.

Nevertheless the Fellowships of the Australian Academy of Science and the Royal Society of London indicate in how much esteem his work was held internationally, and this judgement has been supported by later events. Technical advances in related fields, especially in liquid chromatography and sensitive ultra violet detectors, have led to the development of low-level or microsequencing techniques which still employ the reactions described by Edman forty years ago and which play an indispensible role in gene isolation and molecular cloning.

Edman's reputation as a reclusive person, often difficult to deal with, did not arise from those who worked with him in the laboratory. He was reserved by Australian standards, but courteous and always helpful, often humorous, and took pleasure in organising social occasions both at his home and outdoors in the country. To those who came to know him in the laboratory two aspects probably had a lasting influence. In those days he was a rare example in the hospitals and the world of Australian medical research of someone who devoted himself full-time to nonclinical research. This served as an example, to those who came across him, of the possibility of such a career. At a time when biochemistry in Australia was largely concerned with the intricacies of metabolic pathways, an area where the great discoveries had already been made, Edman understood and stressed the importance of the information-containing macromolecules. The double helical structure of DNA had been proposed by Watson and Crick only a few years before Edman's arrival in Australia. The possibility of obtaining a corresponding understanding of the more complex structures of proteins which Edman's work opened up inspired his colleagues with the belief that they were in a position to participate directly in a new era of biological investigation. fic basis for consciousness. Cognitive Studies 5, 95-109.

About this memoir

This memoir was originally published in Historical Records of Australian Science, vol.8, no.2, 1990. It was written by F.J. Morgan, Department of Biochemistry, La Trobe University.

Written by L.T. Evans.

Introduction

Sir Otto Frankel – whom I shall refer to as Otto because that is how we all addressed him – was a geneticist by training, plant breeder by occupation, cytologist by inclination and genetic conservationist by acclaim. Apart from his personal research, Otto was a highly effective builder and leader of research groups, Socratic gadfly to the scientific establishment, and high prophet of the genetic resources conservation movement. His career in science was unusual in that his most widely acclaimed work was done after his official retirement. A man of inexhaustible variety of opinions, Otto had a complex personality that could be rough or kindly, bored or engaged, impossible or altogether charming by turns, and he did not wish this memoir paint him otherwise. He refused to write an autobiographical sketch and did not keep many records. However, two interviews with him about his career were recorded, by Gavan McCarthy in 1985 [1] and by Max Blythe in 1993 [2]. Much of the information and views quoted here are from records of discussions I have had with him over many years.

Family background and early life

Otto Herzberg-Frankel was born in Vienna on 4 November 1900, the third of four sons of a prominent and wealthy lawyer. Frankel was a relatively uncommon name in Vienna at that time, but to differentiate himself from a more traditionally Jewish branch of the family, Otto's paternal grandfather, a well-known author, added Herzberg from his mother's name to become Herzberg-Frankel. After his father's death, Otto chose to drop the hyphen and revert to Frankel.

Ludwig Herzberg-Frankel, Otto's father, was a highly successful barrister in Vienna, effective both in the courtroom and as a public orator. Otto's enjoyment of, and effectiveness in, public debate and persuasion were clearly shared with his father. What he called his 'peasant instincts', so important in his motivation to make his career in agriculture, came from his mother's family. Thérèse Sommerstein was the youngest child of a family with several rural estates in Galicia. Her older sister Ann was an able and progressive farmer, and Otto's early rural experiences were associated with visits to his aunt's estate [3], where he was also impressed by the abilities and aristocratic manners of her husband, Joseph Bernstein vel Niemirovski. Their son, the British historian who later changed his name to Lewis Namier, also played a significant role in Otto's career.

Fin-de-siècle Vienna was a culturally sophisticated and affluent city which attracted the adventurous and talented in many fields to become a hot-bed of cultural change in art, architecture, music, theatre and philosophy. This was the background to Otto's life and a formative effect on his youth, which had many elements in common with that described by Victor Weisskopf [4] and others. But he also witnessed the collapse of the old Empire, along with the relative impoverishment of his family, at the end of the First World War. At its outset his father had volunteered for military service along with his motor car and his chauffeur. By its end there was no car but, as Otto's brother Paul put it, the brothers had acquired the incentive to work.

Max, Otto's oldest brother (1895-1983), qualified in law but after joining Otto in New Zealand in 1938 he became an accountant. Theo (1897-1986), who had to flee Vienna hurriedly in 1938, became a progressive paper manufacturer in Great Britain, establishing the Scottish Pulp and Paper Mills enterprise in the Highlands. Paul (1903-1992) also moved to Britain, from Poland in 1937. An economist by training, he founded Petroleum Economics Ltd. in 1955 and became a distinguished international authority on the oil industry. Their family home was not a happy one, according to Otto. Paul told me that Otto's youth was quite tempestuous and unhappy, more because of his own problems than of his father's domination. One of these was Otto's sense of 'homelessness' which he ascribed to his father's 'distance' and to not having a country to call 'home': they were Viennese Jews but not Jewish in any religious sense, nor Austrian, nor Polish.

Not only did Otto subsequently claim that he had no country, he also insisted that he had 'no education'. In Otto's early years, his father employed a tutor for his sons as well as a French governess. From 1910 to 1918 Otto attended the Piaristen Staatsgymnasiums Wien VIII, as did Karl Popper. Otto's claim to no education was based on this being a classical rather than a modern school [5], with poor mathematics and next to no science but eight years of Latin and four of Greek. None of his teachers inspired him.

University education

The end of school coincided with the end of the war, when there was little chance of a young man without military service being admitted to the University in Vienna. However, under Otto's leadership, a group of young people took over a disused military laboratory, obtained a copy of the practical course work from the Chemical Institute of the University, worked through it together without any lectures and were subsequently given credit for the course.

Besides chemistry, the young Otto had also become involved in communism, and was arrested on one occasion for addressing a street crowd. At about this time there had been a communist putsch in Munich and Otto went there to be interviewed by the celebrated professor of chemistry, Richard Willstätter. As a result he was admitted to Munich University (1919-1920) to study chemistry, botany and physics. However, after three semesters he had lost some of his enthusiasm for chemistry. The young idealist, concerned about the fight against hunger, wanted to do something more practical like agriculture.

The Agricultural Institute of the University of Giessen was recommended to Otto's father, and he studied there under Professor Paul Gisevius for two semesters in 1920/21. Otto did not find his professor congenial and packed his bags again. He was still determined to learn about farming, however, and in 1922 returned to his aunt Ann's estate in Galicia but soon decided he was not really interested in farm management. His aunt then persuaded him to go back to university, with her support.

In the autumn of 1922 he began his studies at the Agricultural University of Berlin, having been given credit for his earlier studies in Vienna, Munich and Giessen, as well as for his practical farm work [6]. Luckily, at this stage Otto attended a lecture on plant genetics by Professor Erwin Baur, a charismatic personality and lecturer [7], which opened a new and wholly fascinating world. He was challenged by Baur's claim to be able to work with genes and the genetic combinations of plants exactly like the chemist with his molecules and his formulae. Otto asked Baur in 1923 if he could begin research under him before his diploma was completed, and Baur agreed in view of the educational disruptions following the war. However, this arrangement entailed a lot of commuting between the city of Berlin, where he was completing his diploma studies, and the laboratories at Dahlem where Otto grew the large populations of Antirrhinum needed for his research, and this added to his sense of isolation from his fellow students.

In his book on the German genetics community, 1900-1933, Harwood divides them between the 'comprehensives' and the more practically-orientated 'pragmatics'. The former were drawn from the upper classes, had a classical education, and maintained broad biological and cultural interests, while the latter were of more lowly social origins, often with a modern education and more specialized interests, and wished to use their expertise to solve agricultural, industrial and social problems. By social and cultural background Otto could have been expected to join the 'comprehensives' like Kuhn, Goldschmidt and von Wettstein, but his wish to do something of practical value led him instead to Baur, the exemplar of the pragmatics but also with an active interest in 'racial hygiene' [8].

The research problem allocated to him by Baur was one of the earliest studies of genetic linkage in plants. Baur suggested that he clarify the linkage relations between one specific mutant (A, fuchsin red) and another nine mutants in Antirrhinum majus. In this Otto was rather unlucky because, after an extensive crossing and back-crossing programme, he found that all but one of the mutations segregated independently of A, and to a large extent of one another. However, the introduction to his thesis was a comprehensive review of linkage in plants that brought high praise from Baur, led to his first published paper (1) and earned his doctorate in agriculture from the University of Berlin in 1925.

When comparing his university experience with that of his later colleagues Otto concluded that while he had learned the value of self-reliance in research, he had missed out on many important elements of university education. His first degree courses were neither coherent nor comprehensive and he was left to himself in his research. Neither Baur nor Elizabeth Schiemann (his nominal supervisor) gave him significant attention, he was isolated by his work (and his genes) from the other students, and he was not taught cytological techniques. He did present a seminar on linkage in plants, and another on speltoid wheats, a subject he later revisited. And he spent his aunt's supporting grant on a psychoanalysis.

Itinerant post-graduate

Through a client of his father, Otto was then employed for two years (1925-1927) as a plant breeder on a large private estate at Dioseg, near Bratislava, after marrying his first wife, Mathilde Donsbach (1899-1989). Although sugar beet crops and their processing were the major activities on the estate, Otto began wheat and barley breeding programmes, which helped his subsequent appointment in New Zealand.

At this point his cousin Lewis Namier re-entered the scene. He had become an ardent Zionist and advisor to Chaim Weizmann. A British group had arranged for a small team of scientists to be sent to Palestine to establish a plant and animal breeding programme there and to act as a bridge between the Zionist Organization and the Empire Marketing Board under the direction of John Boyd Orr, then Director of the Rowett Research Institute, Aberdeen.

Namier had suggested his cousin as a potential recruit with experience in genetics and plant breeding and Otto was brought to London for interview and briefing. In Palestine Otto found that he was not fully occupied because the main emphasis of the project was on animal improvement. However, he made friends with J.D. Oppenheim who had a microscope on which Otto began his cytological career by counting the chromosomes of the Jaffa orange, which led to his third paper. The highly political nature of the Palestine project resulted in its being visited in 1928 by an influential group led by Walter Elliot, Under-Secretary of State for Scotland. In the course of this visit Otto apparently made it clear that he did not wish to remain in Palestine and temporary support for him in England was arranged until a permanent position became available.

Rowland Biffen, Director of the Plant Breeding Institute in Cambridge, arranged for Otto to work on fatuoid oats. In the course of his work at the Institute, Otto became friendly with A.E. Watkins, an excellent cytologist and evolutionist who introduced him to the cytological complexities of wheat. He also took Otto's still-imperfect English in hand, recommending that he should read all of Jane Austen, which Otto did with pleasure and profit. This brief period in Cambridge was seminal in developing Otto's interest in cytology and evolution, as well as his understanding of the English way of life.

During his stay in Cambridge Otto was invited to accompany F.B. Smith on a secret trip in August/September 1928 to Brazil and Argentina to advise the banking group Lazard Bros. on the establishment of a wheat industry in the southern state of Parana, a mission he enjoyed in many ways as 'my first real overseas trip'.

However, a more permanent position and home was what Otto sought, and it came about quite unexpectedly. His patron, Boyd Orr, was travelling across Canada by train on which he met the newly-appointed Secretary of the New Zealand Department of Scientific and Industrial Research (DSIR), Ernest Marsden. In the course of their discussions Marsden said he was looking for a plant breeder and geneticist for the newly-established Wheat Research Institute in New Zealand. Boyd Orr said he knew of one, and a cable was sent to Professor Biffen for Otto: 'Will you accept post New Zealand beginning £400 per annum plus passage?' There was no indication of what kind of work was involved, but Otto trusted Orr and accepted. Watkins encouraged him to take to New Zealand a subsample of his comprehensive collection of wheat varieties, many of which had been obtained by British government officers from overseas markets. Otto and Tilli arrived in New Zealand, after a month at sea, in March 1929.

Wheat breeding in New Zealand

In view of how significant his 22 years in New Zealand were for his scientific career, it is surprising that Otto says virtually nothing about them in his 1985 interview with McCarthy. He had come to New Zealand hoping to find a 'home'. Indeed, he did feel at home there with the countryside, and especially in the mountains, but 'I was never accepted in New Zealand, I always felt a foreigner and was made to feel that. Only in the ski huts was I accepted.'

At Lincoln College, near Christchurch, where he was to work until 1951, Otto was briefed on the role of the Wheat Research Institute (WRI) by Professor F.W. Hilgendorf, the director, with whom Otto was to have excellent relations. Hilgendorf had conceived of the Institute to improve New Zealand wheat, especially its baking quality, with support from levies on wheat growers, millers and bakers to be matched by government funding. After much proselytizing by Hilgendorf, all parties had agreed to its establishment in 1928, within the DSIR [9].

Hilgendorf had previously crossed the Canadian variety White Fife with Tuscan, and the progeny of the seventh cross were handed over to the newly established WRI. Otto began his breeding programme with these, introducing quantitative assessments of grain yield and of milling and baking quality, which eventually led to the release of the widely grown variety 'Cross 7' in 1934. Being more resistant to lodging and better suited to direct heading, this variety aided the mechanization of the wheat industry as well as improving the quality of New Zealand bread. Although the improvement of milling and baking qualities was the prime objective of his breeding programme, Otto also began an analysis of the yield components in wheat crops, reflecting the influence of F.L. Engledow's work at Cambridge (12, 17, 19).

However, further studies of the effects of selection for yield (26) showed that the efficiency of selection was not enhanced by using yield components rather than yield per se. Otto's overall experience in breeding for higher yield was summarized in an influential review (27) which, in the light of the striking advances in yield potential since that time, now seems far too conservative, indeed pessimistic. Like Farrer in Australia, he was much more confident of being able to improve the baking quality of New Zealand wheat, and put a considerable effort into optimizing the role of quality-testing in the selection process (8, 20). That this was highly effective was shown by the outstanding baking qualities of his cv. Hilgendorf.

Otto had hoped to use Watkins' varietal collection, which he enlarged in New Zealand, particularly for increasing grain size, but in the event this did not prove desirable because of associated loss of baking quality. However, the expansion and maintenance of the wheat collection put him in touch with the great Russian geneticist, N.I. Vavilov, and other plant breeders around the world, and led on to his later interest in genetic resources.

Early cytological investigations

Otto soon felt on top of the demands of his wheat breeding programme and sought the permission of his Director to undertake cytological research on native plants in the off season. Hilgendorf encouraged him and consulted New Zealand's leading ecologist, Leonard Cockayne, who suggested Hebe as a large genus with some interspecific hybrids. The Hebe-Veronica complex required considerable travel, often in the mountains, which was attractive to Otto and he began cytological research on it in 1929 although he was aware, as he indicated in a letter to the DSIR head office, that such 'fundamental' work 'is not the type of work I am here for'.

When Cockayne recommended Hebe for Otto's cytological investigation, he did so as an ecologist who considered it a diverse and interesting genus, but ecological and evolutionary significance do not bestow cytological suitability. In the event Hebe proved to be rather unsatisfactory from a cytological point of view, with its many small chromosomes. However, his cytological evidence suggested several taxonomic revisions, including the grouping of the New Zealand Veronica species with Hebe (14). Further cytological studies (21, 23) supported the establishment of Hebe as a separate genus, and suggested that the New Zealand species of Euveronica and the genus Pygmaea should also be included within Hebe, which subsequent taxonomic work has confirmed.

In wheat, Otto observed two abnormalities among the progeny of an F4 plant from a Tuscan x White Fife cross he made in 1928/29, which provided him with the material for his finest cytological research. The first involved two inverted duplications, one long and one short, that Otto interpreted as having arisen by breakage of dicentric chromatid bridges. The broken ends presumably then 'healed', becoming a functional telomere, thereby preventing sister reunion of chromatids (32). Homozygotes and heterozygotes for these duplications, as well as combined long/short heterozygotes, were analysed for their meiotic behaviour (33) in what M.J.D. White calls 'a piece of classic cytogenetics' [10].

In the same family that displayed the inverted duplications there also occurred a chlorophyll defect that Otto called striatovirescens, the leaves having alternate white and green sectors (35). The defect was shown to be due to three recessive mutations, presumably on homeologous chromosomes of the A, B and D genomes, but how these occurred is uncertain. The independent mutation of three genes, or the mutation of one which was then transferred to the homeologous chromosomes, was considered highly improbable. The association of the defect with the occurrence of the inverted duplication was presumably not coincidental, and the duplication may therefore have acted as a destabiliser of heredity, comparable with those found in maize, which Otto once referred to in a 1954 letter as 'the curious case of Barbara McClintock' [11].

Overseas visit to Darlington and Vavilov, 1935

Otto first canvassed the possibility of an overseas trip between wheat harvests early in 1934. He wanted to extend his wheat collection, have discussions with wheat breeders in Europe, and get some guidance and criticism in 'modern Karyology' which had 'grown up since I came here'. He hoped to finish off his work on Hebe during his stay of several months at the John Innes Horticultural Institution at Bayfordbury. In one sense Otto's work on Hebe was finished off there. C.D. Darlington took one look at his slides and said 'If you want to work on small chromosomes, go to Karpechenko. He likes small chromosomes.' Darlington then introduced him to the chromosomes of Fritillaria, 'on which you can do real cytology'. In any case, as Otto put it, 'the Hebe work was an evolutionary study and I'd got an inferiority complex about the taxonomy'. So although the work on Hebe continued for many years, this first overseas visit changed the direction of his cytological research. He made a systematic search for evidence of inversions among almost 30 species of Fritillaria, using Darlington's collection of slides. None were found in most species, but the results of crossing-over in the four species with them were analysed (13) and the nucleolar cycle examined (15).

Otto's overseas visit in 1935 introduced him to two of the scientists who most influenced his life. When Otto first met Cyril Darlington, he had written his highly influential Recent Advances in Cytology and was in the throes of discussing and revising it. Otto was also stimulated by the wide-ranging lunch time discussions at John Innes between J.B.S. Haldane, Darlington and others.

The other major impact of Otto's overseas trip in 1935 was his meeting with Nicolai Vavilov who warmly welcomed his visit to the USSR, arranged his visa and itinerary [12], and spent much of his time with Otto during the week he was in Leningrad (129). Otto was impressed by Vavilov's passionate drive to identify general principles, by his style of leadership and by his stamina, if not by the poor state of his experimental plots. At the time of Otto's visit Vavilov was preparing the second and third volumes of his Theoretical Bases of Plant Breeding, yet shared his time and ideas generously with Otto. Many years later Otto and Erna Bennett dedicated Genetic Resources in Plants (72) to Vavilov, of whom Otto always had a photograph in his office.

After Leningrad, Otto visited several of Vavilov's research stations, before continuing his visits to European plant breeding institutes on his way back to England. From Kiev Otto sent some 'friendly criticism and suggestions' that Vavilov took seriously in his reply. After his return to New Zealand Otto again wrote urging Vavilov to adopt Brabender's cereal chemistry procedures, criticising the Lysenkoist papers presented at the International Botanical Congress in Amsterdam, and letting him know of the discussions at the succeeding Imperial Botanical Conference in London.

Proselytizing for plant breeding

Otto's return to New Zealand opened a more than usually tempestuous phase in his life. He was divorced from Tilli in 1937, resumed his vitriolic exchanges with E. Bruce Levy (Director of the DSIR Plant Research Station in Palmerston North) over the question of who should oversee pasture plant breeding, and got into hot water for some off-the-cuff comments to a reporter about the Department of Agriculture. Like other extempore remarks by Otto, these found their way into The Press, to the delight of some and the chagrin of the DSIR headquarters.

From early in 1934, Otto had pressed for his wider involvement in DSIR plant breeding activities, initially in relation to pasture plants such as perennial ryegrass. After his return from overseas Otto put forward a more ambitious proposal for a Plant Breeding Section of DSIR, with five Divisions spread between Lincoln and Palmerston North. For a time, Otto held high hopes of its establishment. He had also formulated plans for a Genetics Bureau to undertake fundamental genetic research with both plants and animals and to collaborate with the various groups of plant and animal breeders in New Zealand. However, strong opposition from Levy and others on the plant side, and lack of interest by Dr Dry of Massey College on the animal side, led to these proposals being dropped.

Another activity in which Otto was engaged at that time (1937/9) was in trying to assist the immigration to New Zealand of Jewish refugees from Europe following the Anschluss. Otto was the secretary of a committee on which Karl Popper, who had come to Christchurch early in 1937, was also a member, one who tended to favour intellectuals whereas Otto had to deal with the Minister for Immigration who thought there were already too many of them in the country.

In 1939 Otto again went overseas by ship, on a second pilgrimage to the John Innes Institution, his research there resulting in a paper on chiasma formation in Fritillaria (22). This visit was less stimulating and significant than his first one. Darlington was preoccupied with his new duties as Director of the Institution. Haldane had left and laboratory discussions were less lively and wide-ranging. Otto was impressed by how quickly the research atmosphere of an institution could change.

Remarriage and recognition

Otto returned to New Zealand by ship in the early stages of the Second World War, and married Margaret Anderson (1902-1997), an artist and art teacher from a well-known Christchurch family, within a few hours of getting home on 8 December 1939. They had met many years before and theirs was a wonderfully enduring and happy marriage of two strong personalities. Unlike Tilli, Margaret was always welcomed by Otto's family and he, after they recognized his gardening skills, by hers. The Andersons lived in an historic house called Risingholme, set in large grounds, and Margaret's father gave them some of the land on which to build the first of the three houses and gardens from which they gained so much delight. When he died, Risingholme was given to the City of Christchurch, and Otto and Margaret suggested that it be turned into a community centre, in which they were active for many years. Their first house, completed in 1940, was designed by the great Viennese-born contemporary of Otto, Ernest Plischke, and is illustrated in two books on his architecture.

When Hilgendorf died in 1942, Otto was appointed Chief Executive Officer of the Wheat Research Institute and his powers of leadership, his capacity for planning and his vision were at last given some scope. He had already proved himself to be an able plant breeder whose varieties Cross 7 (released in 1934), Taiaroa and Tainui (1939) and Fife-Tuscan (1941), and subsequently WRI-Yielder (1947), had raised wheat yields, while Hilgendorf (1948) had quite outstanding baking quality [13]. Although the work of the WRI was appreciated by wheat growers, millers and bakers, it was not widely known until 1947 when the seed harvested from the experimental plots of a very promising line was stolen. Otto's appeal for its return received headline treatment, bringing the Institute and its work into welcome prominence. He was reminded of the opera diva who became more famous for her stolen pearls than for her singing.

Nevertheless, these years were among Otto's most scientifically productive. In 1947 he published a paper on plant collections (25), another on selection for yield in wheat (26), and an influential review on the theory of plant breeding for yield (27). Over the next three years his papers included accounts of two newly-released wheat varieties (28, 29) and of what he regarded as his best cytological work, on an inverted duplication in wheat (32, 33), as well as the first paper in his long series on base sterile mutants in speltoid wheat, which he had first observed in 1929 (31).

In 1948 he visited the John Innes for a third time, to find Darlington preoccupied with the controversy over Lysenko's presidential address to the Soviet Academy of Agricultural Science on 31 July. Darlington, S.C. Harland and R.A. Fisher had expressed deep concern over this development in a discussion on the BBC's Third Program and Otto entered the fray with a letter to The Listener of 9 December 1948 questioning J.B.S. Haldane's public defence of Lysenkoism. When they met a little later there was, as Otto put it, 'an exchange of grunts'.

In 1949 the wheat breeding section of the WRI was merged with the DSIR Agronomy Division, also based at Lincoln. For the first year, Otto was associate director of the combined group, becoming Director in March 1950. As he says in his interview with McCarthy: 'for the first time I was able to look for scientific staff...and I could think of the scientific content of a job rather than purely breeding barley or some root crop or other, and I could look for quality...' However, he encountered bureaucratic resistance to his attempts to build a stronger research group at Lincoln, and was soon tempted to leave New Zealand.

Although administrative frustrations precipitated his departure from New Zealand, his sense of intellectual isolation there would probably have led him to leave eventually, as well as the lack of 'old stones and modern art'.

Chief of the CSIRO Division of Plant Industry, 1951-1962

Although Otto felt less at home with the landscape in Australia than in New Zealand, he was 'never made to feel a foreigner', a welcome difference. Two other components led to his remembering his early years in Australia as the most rewarding of his career. Funding for research in the Commonwealth Scientific and Industrial Research Organization (CSIRO) burgeoned in the fifties, giving opportunities for recruitment and building that Otto seized. And in Sir Ian Clunies Ross he had an idealistic, cultured and brilliant but enigmatic Chairman with whom his relations remained warm. Otto's memoir of him [14] radiates appreciation, and a photograph of Clunies Ross was prominent in all the offices Otto later occupied.

Yet when Otto first visited the Division of Plant Industry in Canberra with Carl Forster (an associate of the then CSIRO Executive) before being interviewed in Melbourne, he was reluctant to be considered. It was much larger and more diverse than his Crop Research Division but the buildings were run down and the staff demoralized, and he thought the environment ugly. Forster must have conveyed Otto's impressions to Clunies Ross because when Otto went, still wavering, to see him, the Chairman cut off his retreat with 'Frankel, Frankel. I'm exceedingly sorry you won't take it – and that job is so terribly difficult'.

Clunies Ross had made it quite clear that Otto's major role was to strengthen the Division's research, especially the more fundamental research, particularly in genetics, all of which appealed to him. In fact, when Clunies Ross first joined the CSIRO Executive, one of his main goals had been the building of a national programme in genetics education and research [15], and in a paper to the Executive Committee in 1947 he had argued the need to strengthen genetic research in the Division of Plant Industry. Otto's efforts along similar lines in New Zealand must have encouraged Clunies Ross in his subsequent negotiations with him.

Prior to Otto's interview in mid-1951, there had been a review of the Division that had also recommended a strengthening of basic research, so his brief was clear and he set about it vigorously on a series of recruiting trips overseas. He was sceptical of the effectiveness of advertising positions: 'People who apply are rarely the best, they are footloose, they are available. You want to get people who are not available'. Much of the agrostological and associated research moved to Brisbane, where it eventually became a separate Division. Distinguished older scientists and promising young ones were recruited to give new life to old activities. In other areas, such as plant introduction, Otto simply challenged them to do better. The small Divisional unit in plant physiology already established at the Waite Research Institute in Adelaide was moved to Canberra and enlarged. Plant biochemistry also required extensive recruiting. Some projects were tapered off and the younger research staff given new opportunities, as in the case of the phytochemical survey staff in Queensland who were refocused on rainforest ecology.

The most profound changes, however, were in the area of genetics and cytology, Otto's own, where he felt most confident in his recruiting and where he had the strong support of Clunies Ross. Although the fields of quantitative genetics and cytology received initial emphasis, a powerful and wide-ranging group in evolutionary genetics was quickly established, and became highly influential both in Australia and internationally. Otto's personal commitment to his research on the genetics of floral development in wheat, the high standards he demanded, his interest in broader genetic issues and his provocation of their discussion made the genetics seminar a lively centre of Divisional activities.

As he sought to rejuvenate the Division and re-orientate it to 'science for the second half of the 20th century' there was, naturally, some resentment among some of the older staff. However, his efforts soon transformed the scientific life and standing of the Division by building up a range of strong discipline-based groups, such as that in agricultural physics under the leadership of J.R. Philip. The recruiting of additional staff in new fields greatly enlarged the Divisional requirements for additional laboratory space and new equipment, both of which proved hard to obtain. Looking back, Otto thought these were his greatest difficulties. When he became Chief, the Division was still largely housed in its original 1930s building and in spite of great efforts by Otto there was no major addition until the 'Genetics' building was completed in 1958. Even that required a personal visit by the Prime Minister, R.G. Menzies, to the Division. After that, the new buildings came more readily, the Genetics extension being completed in 1960, the Biochemistry building in 1961 and the phytotron in 1962.

The capstone to Otto's reconstruction of the Division was his campaign to build an Australian phytotron to serve as a national facility for research on the responses of plants to climatic factors. On his first visit to the phytotron at the California Institute of Technology in 1953, he was greatly impressed by its potential value for agricultural research in a country like Australia with such a wide range of climatic conditions. This would be an expensive facility, especially for the biological sciences at that time, and the 'big science' element was a challenge. But Otto also wanted the Australian phytotron to be novel and distinctive in both engineering design and architecture. Clunies Ross was supportive, and Otto soon enlisted the enthusiastic cooperation of R.N. Morse, officer-in-charge of the CSIRO Engineering Section, and his staff.

A novel design was developed, in which the major components were thoroughly tested before financial commitments were sought, and in 1958, the federal government decided to provide the requisite funds. The phytotron was officially opened in August 1962 by the Prime Minister, Mr Menzies, by which time Otto had joined the CSIRO Executive. Nevertheless, its scale, originality and style symbolized, for many, Otto's leadership of the Division of Plant Industry.

Other research areas in the Division were also stimulated and enlarged right up to the end of Otto's term as Chief. Perhaps his greatest disappointment as Chief was that Clunies Ross, not long before he died in June 1959, began to express concern that Otto had neglected, even impoverished, applied research by the Division. Yet Otto continually challenged his research staff to pursue their work within a framework of potential relevance to agriculture.

What particularly saddened him was that, as he makes plain in his interview, his initial attraction to the CSIRO culture was the enthusiastic and idealistic encouragement given by Clunies Ross and his Executive colleagues to basic research related to applied problems, and he sensed that such enthusiasm was faltering. Despite this, he told McCarthy that the biggest thing in his life, which he felt 'everyday in spite of all my wars and my arguments', was his admiration and respect for CSIRO. On one occasion when Otto expressed this rather forcibly in a comparison with the Australian National University to Lord Florey, its Chancellor, the latter was moved to comment: 'Frankel, I'm not here to award marks'.

On the CSIRO Executive, 1962-1966

Otto was persuaded by R.N. Robertson to succeed him in 1962 as a member of the Executive of CSIRO. He would rather have remained in the Division, preferring always to fight for something than against it, distrusting the 'management' of science, and enjoying the irreverence of young colleagues. But he sensed that a need to fight for basic research within CSIRO was emerging. He also sensed a need to protect the Division of Plant Industry, by then the largest in CSIRO, from being split, which would reduce the interactions and cross-fertilization between disciplines that he had tried to foster. He may also have had expectations of succession to the Chairmanship of CSIRO.

However, he found little satisfaction in his work, missed the contact with active researchers and missed his home and garden through having to spend most of his time in Melbourne. Otto's record as Chief made it clear that he was excellent at choosing staff given his subtle, merciless, multidimensional judgement of people, and he had expected to deploy those skills in the choice of new Chiefs of Divisions, but even there he was thwarted.

It was fortunate, therefore, that just at this juncture a subject in which he had long been interested, especially since visiting Vavilov in Leningrad, namely the conservation of genetic resources, began to emerge on agendas for international action. Otto soon became a key figure in the movement and remained so for thirty years after his official retirement from CSIRO in 1966.

Research after retirement

On retirement Otto returned to the Division of Plant Industry as an Honorary Research Fellow, which allowed him to resume active research in genetics and to play a more active role in the International Genetics Federation.

Throughout his life, after hearing that lecture by Baur when he was in his early twenties, Otto saw himself primarily as a geneticist. Altogether he attended ten International Genetics Congresses, beginning with the 6th at Edinburgh in 1939. He was a Vice-President and Treasurer of the International Genetics Federation from 1968 to 1973 and, according to M.J.D. White, 'played a large part in bringing together pure and applied geneticists in order to confront these most critical problems of the earth's biota in an intelligent, informed and humane manner' [10].

For Otto, as for François Jacob, 'genetics became a bastion of reason. To do genetics was to say no to intolerance and fanaticism' [16]. Many of Otto's closest friends were geneticists, and he admired the way Darlington, Dobzhansky, Haldane and others viewed humankind and its problems from a genetic perspective.

Otto's post-retirement research was focused on the base sterile mutants of speltoid wheats. In the evolution of wheat, the appearance of the free-threshing 'naked' grain character, was a highly significant step in domestication, associated with the presence of the Q-factor on the long arm of chromosome 5A. Speltoid mutants, with a brittle rachis, long internodes and tight glumes, arise by a cytologically-observable deletion of the Q segment.

Otto had found several speltoids in a crop of Yeoman wheat in 1929/30, among which one plant was sterile in most of the basal (first) florets. This sterility was found to be caused by the recessive allele of a single gene, subsequently called Bs and found to be located on chromosome 5D, and homeologous with the Q factor (31, 85). Even in a single dose, Bs prevents basal sterility in the second and higher florets, whereas sterility can extend up to them in the double recessive, to an extent depending on the polygenic background (68, 92). Eventually a series of speltoid lines was developed that ranged from full fertility to sterility of the first three florets, or even of the first eight or nine (65). At the other extreme of this scale of fertility were the 'compactoids', in which grains are found in the axils of the normally sterile basal glumes, associated with the addition of a second Q factor.

On the question of the relation between the genomes in floral morphogenesis of vulgare (bread) wheats, it is striking that the dominant Q factor is not present in any of the ancestral A, B or D genomes (50), which nevertheless had genetic systems ensuring the fertility of their first florets. According to Kuckuck [17] the Q factor may have arisen from unequal crossing over in chromosome 5A, and Frankel and Roskams (92) proposed that a gene homeologous to Bs was included in the multiple repeat. They also examined the effect of a period in short days at high temperatures at the beginning of floral initiation on the pattern of floret sterility in several normal and base sterile genotypes. These shock treatments increased basal sterility in the speltoid lines but not in the normal wheats. Further work identified the most sensitive period for each floret (94), and the stage at which cell division failed (113). Even the initiation of sterile floret primordia required prior lemma initiation (110). Although no further work has been done on them, the speltoid lines developed by Otto still have much to offer those interested in the molecular control of floret differentiation and fertility in wheat and other cereals and grasses.

FAO, IBP and genetic resources

As long ago as 1923, N.I. Vavilov had warned of the need to conserve, as well as to use, the range of genetic variation within crop plants in the face of agricultural change. With the initiation of the International Biological Program (IBP) in 1963, concern for such 'genetic erosion' was heightened. Otto had not been keen to be involved in the IBP, but at the urging of R.N. Robertson, of Ledyard Stebbins (who had introduced 'plant gene pools' as a major theme of IBP) and of C.H. Waddington (Vice-President of IBP and an old friend from his early days in Cambridge), he took part in the 1st General Assembly at Paris in 1964. The entry of the IBP, and of Otto, into the field resulted in a transformation in public awareness of the problem and plans for action. The programme, drafted by Otto in 1965, led to a clearer definition of the various kinds of genetic resources, a strategy for their conservation with priority on the land races, and an emphasis on information and availability. The realisation that IBP could not achieve this on its own led Otto to meet R.B. Sen, the Director-General of FAO, in 1965, to explore the prospects for joint efforts. Both the IBP and FAO welcomed the proposed collaboration, and Otto was invited by Sen to act as a consultant in 1966, to review the activities and responsibilities of FAO, and to prepare plans for a meeting in 1967. This integration of effort by IBP and FAO continued until the end of the IBP in 1974 [18].

During his consultancy at the FAO in 1966, Otto was visited by a fellow student from Baur's group, Professor Hermann Kuckuck, who painted an alarming picture of the accelerating loss of land races and wild relatives in Turkey and Ethiopia. His account brought home to Otto the need for urgent and comprehensive action, which he stressed in his report to FAO.

The 1967 FAO/IBP Conference on 'The Exploration, Utilization and Conservation of Plant Genetic Resources' was a landmark for the genetic resources movement. In both its planning and the reworking of its proceedings, Otto was joined, at his personal request, by Erna Bennett, and together they coined such phrases as genetic resources and genetic erosion. The conference itself led to a programme for FAO-initiated international action, while the book (72) had a substantial impact on the scientific community. In particular, the book emphasised the importance of what Otto liked to refer to as the 'generalist strategy' of Vavilov as against 'mission-oriented' collecting (122).

After the 1967 conference there followed a frustrating period of bureaucratic inaction by FAO. The Panel of Experts was reconstituted with a membership representing both IBP and FAO under Otto's chairmanship, and many issues were considered at their four meetings. The high priority for collection of endangered land races was retained. Consideration was given to the problems of evaluating accessions more broadly, to computerizing the information, to long-term seed storage, to the establishment of a global network of genetic resource centres, and to the respective roles of the various kinds of collection. Throughout the late sixties and early seventies it was the Panel of Experts under Otto's activist chairmanship, and Erna Bennett within FAO, who kept the genetic resources issues alive. As Otto wrote later: 'Conservationists who became so concerned in the eighties when the battle was essentially over were notably uninterested when their publicity might have been invaluable' (127).

Otto published several papers on genetic resources issues through this period, many of them aimed at increasing public awareness of the problems. One of the finest of these – in the estimation of Soulé and Mills [19] – was his Macleay Memorial Lecture entitled 'Variation – the essence of life' (77), in which he argues that the scale of human impact on genetic variation within both domesticated and natural communities is now such that we can no longer claim evolutionary innocence: 'We have acquired evolutionary responsibility' and must develop an 'evolutionary ethic'.

A conference of experts was convened at Beltsville, USA in 1972, to consider the proposal of the IBP/FAO Panel of Experts for the establishment of a network of regional genetic resource centres plus a coordinating centre to recommend priorities and organize training and other activities of the network, which would be associated with FAO. Otto was invited to present the report of the Beltsville meeting to the Technical Advisory Committee of the Consultative Group on International Agricultural Research (CGIAR) in April 1972.

Two months later Otto unexpectedly found himself given an opportunity to address the United Nations Conference for the Human Environment, in Stockholm, on genetic resources. He had been asked by FAO to prepare a background paper on this subject for the conference, with recommendations. Several delegates moved the adoption of these recommendations, and another requested that Otto be allowed to address the conference. He relished the opportunity, his recommendations were adopted in Articles 39-45 [20], and the world's news media carried his message. He became a cult figure at Stockholm and genetic resources became an international issue, requiring consideration by national governments and inviting the concern of public interest groups. The genetic conservation wave began to roll, fourteen years before the term 'biodiversity' was coined.

In 1973 the CGIAR established an International Board for Plant Genetic Resources (IBPGR). Otto, who was widely expected to be the first chairman of the Board, was not even a member. Moreover, FAO, which had sponsored the cause of genetic conservation when no other organization did so, abandoned its own Panel of Experts once IBPGR was established. Although the panel members had expected their accumulated experience in the area to be retained, the Panel was barred from contact by the Board, and disappeared (128). A round-up technical conference was held in Rome in 1973, the proceedings being edited by Otto and Professor J.G. Hawkes (88).

Otto's most widely admired and influential paper, 'Genetic conservation: our evolutionary responsibility' (84), presented at the 13th International Congress of Genetics in Berkeley, had already been published. Regarded by M.J.D. White 'as a landmark in the cultural evolution of the human species', and by Soulé and Mills as 'prophetic...(presenting) the conceptual and moral agenda for the discipline of conservation genetics', this paper signalled the end of Otto's most active, creative and influential role in the genetic resources movement. Otto then collaborated with Michael Soulé in the writing of Conservation and Evolution (109). Published in 1981, this was a pioneering book, particularly in placing the genetic resources movement within the wider context of the conservation of biological diversity and of the opportunity for continuing evolution.

Otto was now freer to speak out on genetic resources issues as he continued to think and write about them. He urged greater activity by the national gene banks (128) and more comprehensive evaluation and documentation of accessions. He proposed the use of representative 'core collections' as being more accessible for plant breeders (118). Nevertheless, Otto had always regarded the global network of base collections as the backbone of the genetic conservation strategy, and was appalled when one of his erstwhile colleagues suggested a shift of emphasis to the national collections. At the age of 90 he still responded vigorously (140). He had earlier engaged in public debate with P.R. Mooney on the subject of 'farmers' rights', and in 1988 locked horns with J.R. Kloppenberg and others on the 'sovereignty of seeds' and our 'genetic debts' to developing countries (131). He expressed his views on the FAO International Undertaking, on the Commission on Plant Genetic Resources, and on the Keystone International Dialogue (137). He became less and less sanguine about the role of botanic gardens in genetic conservation, and more and more convinced of the benefits of in situ conservation of wild species, while remaining unsure to the very end on where to draw the line between the impossible goal of conserving everything and the utilitarian approach of conserving only species of likely usefulness.

Otto's views on several issues in genetic conservation evolved, but his commitment to the effective conservation and use of genetic resources never wavered. However, there were times when the populist excesses and errors of Mooney, the genetic debts movement and the FAO commission made him want to dissociate himself from the issue: 'I sometimes wonder whether the ideas of the early days – which became the Genetic resources dogma – did harm through overstatement and over-acceptance' [21]. When political fashion and rhetoric displaced reasoned debate, or when bureaucratization and management issues predominated, he often wished to quit but continued to debate them to the very end. He was a worthy successor to Vavilov, and took great pleasure in the decision of the erstwhile IBPGR to establish a Vavilov-Frankel Fellowship Program in genetic resources, with the first awards in 1993. At 95 years of age Otto, with two younger colleagues A.H.D. Brown and J. Burdon, published The Conservation of Plant Biodiversity (142).

Other activities related to science

In New Zealand throughout the 1930s and 1940s, Otto was an outspoken advocate of high-quality basic research. He was elected to fellowship of the Royal Society of New Zealand in 1948, of the Royal Society of London in 1953 and of the Australian Academy of Science in 1954, becoming vice-president of the latter in 1959/60. He once observed, somewhat ruefully: 'I tend to be held at the Vice-presidential level, being too unpredictable to be made President'.

In 1956, when designs for an Academy building were first being discussed, a sketch in the classical style, decorated with columns, was produced by a member of Council. Otto found it appalling and suggested that a design committee should be appointed [22]. Otto, as a member of the committee under the chairmanship of the President, Sir Mark Oliphant, sought advice on architects to be approached and procedures to be followed and chaired the meeting at which the design by Grounds, Romberg and Boyd was recommended to Council. Otto was then appointed to the Building Committee where, as he later put it: 'In everything I was concerned with style and Oliphant with quality and I think we made a very good team'.

The bold, modern, symbolic design of the Academy dome delighted him. In his tribute to Roy Grounds [23] he wrote: 'The Academy building helped to generate a corporate consciousness and, thanks to its architectural distinction, it enhanced a growing pride in the Academy. For the public it became a symbol of Australian science.' Otto had enjoyed working with Roy Grounds. He became a close friend and Otto and Margaret asked him to design their third house, built in 1971.

In the early 1960s when the International Biological Program was first being discussed, Otto had been an outspoken critic of the proposed projects. However, he was persuaded by R.N. Robertson to convene the Academy's ad hoc committee to report on the advisability of Australian participation in the IBP. Given the strength of local research in many areas of the IBP, Otto's committee recommended that Australia should take an active part, and Otto was made chairman of the National Committee for IBP. Despite his early reservations, he provided sustained and dynamic leadership of the Australian efforts, and was a member of ICSU's Special Committee for IBP.

Through his service on Council and on several committees, Otto had opportunities to shape the practices as well as the architecture of the Academy. In his 1972 Falk memorial lecture (79), Otto had suggested the establishment of a 'Science and Society Forum' by the Academy, to concern itself with scientific issues of likely or emerging public impact and to provide a forum where 'dissent and constructive criticism are given the orderly freedoms of institutionalization'. The first such Forum, however, was not under his chairmanship and proved to be disastrously disorderly and the experiment was abandoned [24].

In 1979, Otto was appointed chairman of a committee of review to report to the 25th anniversary meeting of the Academy on the appropriateness of its activities and structure for the next decades. Here the canvas was much wider and the committee made a lot of recommendations for change, many of which were acted upon. 'Whatever the Academy is now, it ought to be something better', was the attitude Otto took to the review. He wished the Academy to become more outward-looking and responsive in its second quarter-century and, looking back later, he felt it had.

The man

Complex, mercurial, charismatic, acerbic, persuasive, polarizing, practical, ironical, elegant, concerned: these are some of the adjectives that spring to mind when colleagues and friends recall Otto. His complex mixture of practical peasant and intellectual aristocrat flowered when he played host in the elegant houses and gardens he and Margaret had created. Clunies Ross had once suggested that the corner of the CSIRO land where Otto hoped to build his second house was not regarded as an appropriate environment for a Chief, to which Otto replied: 'We don't mind. We make our own environment' – which they did.

Whatever opinion you expressed, whatever side of an issue you took, he would challenge it to sharpen both your thinking and his, a practice he learned from his uncle Joseph Niemirovski. Where some focus on points of agreement, Otto homed in on points of disagreement. It could be tiresome, but also illuminating, because he was quick and resourceful in debate and liked to test all facets of any idea. Arguments with Otto were 'energetic', as Boswell found those with Dr Johnson. Such a man does not have heroes. He would always want to argue with them. He told me he had none, but for many years there were photographs of Clunies Ross, Darlington, Dobzhansky and Vavilov displayed in his offices. They were friends, and they had influenced him at various stages, 'played God in his life' as he had with others.

His friends were mostly men and mostly scientific colleagues. Science was important to him, but so also were music and the arts, architecture and skiing, in all of which he had cherished friends. He particularly relished the company of young colleagues who would argue with him irreverently yet seriously about their research and his, and about science in general. Otto cherished these generations of intellectual offspring, not having had children of his own.

He also enjoyed meeting the great and famous but once wrote to a social scientist: 'I am no longer top brass, which in my eyes I never was, nor do I feel the need to listen to it. It might be a problem for your consideration why inescapably we turn to the Establishment when we organize a serious discussion of social problems...I hope I am right in thinking that in the natural sciences on the whole we have slightly better ways of discerning and using talent at a less exalted level' [25].

Otto was always quick to remind protégés of the dangers of the 'slippery slope' away from the real world of science, whenever they were tempted to take on organizational responsibilities or international commitments. Life was theatre for him, and like Picasso he would play games with people, especially those he cared about. Picasso's reply to Max Jacob as to why he did this could have come from Otto: 'Since I cared very much about my friends, it seemed to me I should put our friendship to the test every once in a while just to make sure it was as strong as it needed to be' [26].

Whenever Otto called himself a peasant, it was partly out of pride in his practicality. He enjoyed the manual skills needed in cytological work, 'doing his root tips' as his wife referred to it, and he often noted the absence of such skills in his colleagues with surprise. Dobzhansky was 'clumsy', while Vavilov was 'not terribly good on techniques, and plant breeding with which I was familiar was not second nature to him' [27], a comment as revealing of Otto as of Vavilov. One look at Otto's hands, or at his gardening tools, sufficed to indicate that he was a practical man, expert in the arts of pruning (both plants and people). At the entrance to the Canberra phytotron he had inscribed 'Cherish the earth for man will live by it forever', which truly reflected his values and concern for the world's resources. He cherished his garden and the fruits of it, even the imperfect ones. He was a true husbandman.

Music was also important to him and even in his nineties he enjoyed hearing new compositions. Trout fishing was, for a time, a consuming hobby, in which he enjoyed playing the fish, much as he enjoyed playing his colleagues at times. Besides gardening, skiing was his most abiding joy, which he practised whenever he could in Europe, New Zealand, Australia and the USA. Near the Blue Cow Club, which he helped to found at Guthega, there is a slope now called 'Sir Otto's run', on which he was to be seen each year until he was 90.

Otto's practicality was also expressed in his attitude to research. He was an unwavering proponent of the need for basic research, provided it was 'first class', as the key to enlarging our understanding of the world about us. His loyalty to CSIRO and to Clunies Ross derived from their support of that approach in the CSIRO culture of the 1950s, when he joined. Nevertheless, he was also an agriculturist at heart, deeply concerned with the world's food and population problems, and he encouraged long-term research with a bearing on those problems. The long term was his 'time scale of concern', and the internationality of science his delight.

Humane was an important word in Otto's vocabulary. Not having grown up with English as his native language, he savoured his subsequent mastery of it and was resourceful in his use of words. His early exposure to Jane Austen left its mark on his unremitting search for elegance of expression, fine manners, and love of irony, the salt of life to him. Always fluent and persuasive in speech, especially extempore speeches such as his address to the Stockholm conference in 1972, he nevertheless laboured hard – with pencil, eraser and scotch tape – on his written drafts. To the end of his career, he envied those scientists who wrote with an elegant style, such as E.O. Wilson and his colleague J.R. Philip. Elegance of expression was as significant for him as perpetual challenge, and he sought both to the very end.

In old age he remained determinedly active, alert, involved and irreverent, consciously exercising both his body and his mind. He would 'not go gentle into that good night', as Dylan Thomas had urged his own father. He died on 21 November 1998.

Degrees and honours

• D. Agr. (Berlin, 1925)
• D. Sc. (New Zealand, 1951)
• FRS NZ (1948)
• FRS (1953)
• FAA (1954)
• Knight Bachelor (1966)
• Correspondant Štranger, French Academy of Agricultural Science (1969)
• Honorary Life Fellow, Pacific Science Association (1979)
• Distinguished Economic Botanist (1983)
• Honorary Member, The Japan Academy (1983)
• Foreign Associate, US National Academy of Science (1988)

About this memoir

This memoir was originally published in Historical Records of Australian Science, vol.12, no.4, 1999. A shorter version of this memoir is also published in Biographical Memoirs of Fellows of the Royal Society of London, 1999. It was written by L.T. Evans, Honorary Research Fellow, CSIRO Division of Plant Industry, Canberra, ACT.

Acknowledgements

Reminiscences and comments from many colleagues and friends of Otto and Margaret have shaped this memoir. For detailed comments on earlier drafts I am grateful to Dr R.D. Brock, Dr A.H.D. Brown, Professor R.W. Home, Dr Elizabeth Whitcombe and Professor S.G. Wildman. The photographic portrait of Otto was taken in his seventies by Colin Totterdell; that of him participating in the protest was published in Search, 21(2) (1990), 47.

Notes and references

1. McCarthy, G. (1985) Transcript in Frankel papers, to be deposited in the National Library of Australia, Canberra.
2. Blythe, M. Videotaped interview, 15 September 1993, deposited in the Australian Academy of Science.
3. Otto's aunt Ann and the Niemirovski estate at Koszylowce where he worked are described by Julia Namier in Lewis Namier, A Biography (London, 1971).
4. Weisskopf, V. The Joy of Insight – Passions of a Physicist, (New York, 1991).
5. In his book Styles of Scientific Thought: The German Genetics Community 1900-1933 (Chicago, 1993), J. Harwood describes the difference between the 'classical' and 'modern' schools of Germany and their relation to the later careers of German geneticists.
6. Such frequent shifts from one university to another, and between subjects, were not uncommon in Germany at that time, cf. R.B. Goldschmidt, Portraits from Memory (Seattle, 1956).
7. Schiemann, E., 'Erwin Baur', Berichte der Deutschen Botanischen Gesellschaft, 52 (1934), 51-114.
8. Baur had come to genetics from the practice of psychiatry and believed that the new knowledge of genetics should also be applied to human populations and to the question of racial hygiene, as discussed by R.D. Harvey in 'Pioneers of genetics: A comparison of the attitudes of William Bateson and Erwin Baur to eugenics', Notes and Records of the Royal Society of London, 49 (1995), 105-117.
9. The background to the establishment of the WRI and the work which led to it becoming one of the early successes of DSIR are described more fully in DSIR: Making Science Work for New Zealand, by Ross Galbreath (Wellington, 1998).
10. White, M.J.D., 'Otto Frankel – contributions to wheat genetics', pp. 279-283 in Wheat Science – Today and Tomorrow, ed. L.T. Evans and W.J. Peacock (Cambridge, 1981).
11. Frankel to S. Smith-White, 15 June 1954.
12. Vavilov to E. Marsden, 29 September 1934. A notebook records the details of Otto's visits to the USSR and European wheat breeders.
13. Copp, L.G.L., 'Wheat breeding in New Zealand', New Zealand Wheat Review, No. 10 (1967), 78-86.
14. Frankel, O.H., 'The very human touch', pp. 131-135 in Ian Clunies Ross, Memoirs and Papers, with some Fragments of Autobiography (Melbourne, 1961).
15. McCann, D.A. and Batterham, P., 'Australian genetics: a brief history', Genetica, 90 (1993), 81-114.
16. Jacob, F., The Statue Within (Transl. F. Philip) (New York, 1988).
17. Kuckuck, H., 'Neuere Arbeiten zur Entstehung der hexaploiden Kulturweizen', Zeitschrift fur Pflanzenzüchtung, 41 (1959), 205-226.
18. Worthington, E.B., The Evolution of IBP (Cambridge, 1975).
19. Soulé, M.F. and Mills, L.S., 'Conservation genetics and conservation biology: a troubled marriage', pp. 55-68 in Conservation of Biodiversity for Sustainable Development, ed. O.T. Sandlund, K. Hindar and A.H.D. Brown (Oslo, 1992).
20. Pistorius, R., Scientists, Plants and Politics (Rome, 1997).
21. Frankel to M.F. Day, 19 March 1985.
22. Fenner, F., ed., The Australian Academy of Science: The First Forty Years (Canberra, 1995).
23. Frankel, O.H., 'Sir Roy Grounds, 1905-1981', Historical Records of Australian Science, 5 (1982), 89-91.
24. Johnston, R., 'Social responsibility of science: the social mirror of science', pp. 308-325 in MacLeod, R. (ed.) The Commonwealth of Science (Melbourne, 1988).
25. Frankel to S. Encel, 20 November 1977.
26. Gilot, F. and Lake, C., Life with Picasso (New York, 1964).
27. Frankel to B.M. Cohen, 31 August 1977.

Bibliography

1. 1925 Frankel, O.H. Faktorenkoppelung bei Pflanzen. Zeitschrift für induktive Abstammungs-und Verebungslehre 38: 324-348
2. 1929 Baur, E., Herzberg-Frankel, O., Husfeld, B., Saulescu, N. and Schumann, E. Koppelungserscheinungen bei Antirrhinum majus. Zeitschrift für induktive Abstammungs-und Verebungslehre 50: 314-343
3. 1929 Oppenheim, J.D., Frankel, O.H. Investigations into the fertilization of the 'Jaffa Orange'. I. Genetica 11: 369-374
4. 1929 Frankel, O.H. Pflanzenzüchtung in Neuseeland. Der Züchter 1:9
5. 1930 Genetics and plant-breeding. New Zealand Journal of Science & Technology 11: 401-408
6. 1930 Frankel, O.H. Analytical yield investigations of New Zealand wheat. 1. Wheat Research Institute Annual Report pp. 42-59
7. 1932 Frankel, O.H. Analytische Ertragsstudien an Gertreide. Der Züchter 4: 98-109
8. 1933 Frankel, O.H. and Donald, H.P. Some critical observations on quality testing in wheat breeding. Proceedings of the World Grain Exhibition and Conference 2: 400-408
9. 1933 Frankel, O.H. A case of mass-occurrence of non-inherited chlorophyll defects in wheat. Transactions of the Royal New Zealand Institute 63: 141-143
10. 1934 Frankel, O.H. 'Cross 7' wheat. A new combination of high yield and baking-quality. Bulletin New Zealand Department Scientific Industrial Research 46
11. 1935 Frankel, O.H. The differentiation of grain samples of closely related varieties of wheat by means of a simple mechanical test for grain quality. Journal of Agricultural Science 25: 461-465
12. 1935 Frankel, O.H. Analytical yield investigations on New Zealand wheat. 2. Five years' analytical variety trials. Journal of Agricultural Science 25: 466-509
13. 1937 Frankel, O.H. Inversions in Fritillaria. Journal of Genetics 34: 447-462
14. 1937 Frankel, O.H. and Hair, J.B. Studies on the cytology, genetics and taxonomy of N.Z. Hebe and Veronica (Part 1). New Zealand Journal of Science Technology 18: 669-687
15. 1937 Frankel, O.H. The nucleolar cycle in some species of Fritillaria. Cytologia 8: 37-47
16. 1938 Frankel, O.H. The evolution of cultivated plants. Journal of the New Zealand Institute of Horticulture 8: 27-34
17. 1938 Frankel, O.H. and Hair, J.B. Analytical yield investigations on N.Z. wheat. III. Nine years' observations on two varieties. New Zealand Journal of Science Technology 20A: 224-259
18. 1939 Frankel, O.H. Tainui, a new spring wheat variety. New Zealand Journal of Science Technology 20A: 319-323
19. 1939 Frankel, O.H. Analytical yield investigations on N.Z. wheat. IV. Blending varieties of wheat. Journal of Agricultural Science 29: 249-261
20. 1940 Frankel, O.H. A critical survey of breeding wheat for baking quality. Journal of Agricultural Science 30: 98-112
21. 1940 Frankel, O.H. Studies in Hebe. II. The significance of male sterility in the genetic system. Journal of Genetics 40: 171-184
22. 1940 Frankel, O.H. The causal sequence of meiosis. I. Chiasma formation and the order of pairing in Fritillaria. Journal of Genetics 41: 9-34
23. 1941 Frankel, O.H. Cytology and taxonomy of Hebe, Veronica and Pygmaea. Nature (London) 147: 117-118
24. 1941 Frankel, O.H. 'Fife-Tuscan' wheat: a new variety for 'Tuscan land'. New Zealand Journal of Science and Technology 22A: 303-308
25. 1947 Frankel, O.H. Plant collections. Journal of the Australian Institute of Agricultural Science 13: 122-124
26. 1947 Boyce, S.W., Copp, L.G.L. and Frankel, O.H. The effects of selection for yield in wheat. Heredity 1: 223-233
27. 1947 Frankel, O.H. The theory of plant breeding for yield. Heredity 1: 109-120
28. 1948 Frankel, O.H. Hilgendorf wheat of outstanding baking quality. New Zealand Journal of Agriculture 76: 117-119
29. 1948 Frankel, O.H. A new high-yielding wheat variety – WRI-yielder. New Zealand Journal of Agriculture 76: 221-222
30. 1948 Frankel, O.H. Wheat varieties in New Zealand. Canterbury Chamber of Commerce Agricultural Bulletin No. 22
31. 1948 Frankel, O.H. and Fraser, A.S. Basal sterile mutants in speltoid wheat. Heredity 2: 291-397
32. 1949 Frankel, O.H. A self-propagating structural change in Triticum. I. Duplication and crossing-over. Heredity 3: 163-194
33. 1949 Frankel, O.H. A self-propagating structural change in Triticum. II. The reproductive cycle. Heredity 3: 293-317
34. 1950 Frankel, O.H. The development and maintenance of superior genetic stocks. Heredity 4: 89-102
35. 1950 Frankel, O.H. A polymeric multiple gene change in hexaploid wheat. Heredity 4: 103-116
36. 1951 Frankel, O.H. The multiple mutation in wheat. Heredity 5: 349
37. 1954 Frankel, O.H. Genetic adaptation of cultivated plants in Australia. Proceedings Pan Indian Ocean Science Congress pp. 71-74
38. 1954 Frankel, O.H. Invasion and evolution of plants in Australia and New Zealand. Caryologia Supplement 6: 600-609.
39. 1957 Frankel, O.H. The biological system of plant introduction. Journal of the Australian Institute of Agricultural Science 23: 302-307
40. 1958 Frankel, O.H. The biological system of plant introduction. Indian Journal of Genetics and Plant Breeding 17: 336-342
41. 1958 Frankel, O.H. The dynamics of plant breeding. Journal of the Australian Institute of Agricultural Science 24: 112-123
42. 1958 Frankel, O.H. and Williams, J.D. A record of natural crossing in subterranean clover. Journal of the Australian Institute of Agricultural Science 24: 162-163
43. 1958 Frankel, O.H., Gani, R. and Munday, A. Two independent gene systems for floral induction in wheat. Proceedings 10th International Congress of Genetics 2: 86
44. 1959 Frankel, O.H. Variation under domestication. Australian Journal of Science 22: 27-32
45. 1959 Morley, F.H.W. and Frankel, O.H. An ecogenetic research program with introduced plants. Monographiae Biologicae 8: 577-586
46. 1960 Brock, R.D. and Frankel, O.H. Plant improvement. Journal of the Australian Institute of Agricultural Science 26: 170-182
47. 1960 Frankel, O.H. and Munday, A. The genetics of floral development in wheat. Journal of the Wheat Information Service, Biology Laboratory, Kyoto University 11: 1-3
48. 1961 Frankel, O.H. The F.A.O. Freedom from Hunger Campaign. Journal of the Australian Institute of Agricultural Science 27: 79-84
49. 1962 Frankel, O.H. Agricultural science and productivity in the next decade – plant science. Journal of the Australian Institute of Agricultural Science 28: 84-91
50. 1962 Frankel, O.H. and Munday, A. The evolution of wheat. In: The Evolution of Living Organisms (Royal Society of Victoria, Melbourne) pp. 173-180
51. 1963 Frankel, O.H. Agricultural scientists among scientists. Journal of the Australian Institute of Agricultural Science 29: 95-103
52. 1963 Frankel, O.H. Concluding remarks – the next decade. In: Environmental Control of Plant Growth (L.T. Evans ed., New York, Academic Press) pp. 439-441
53. 1963 Frankel, O.H. The social responsibility of agricultural science. Australian Journal of Science 25: 301-307
54. 1963 Frankel, O.H. and Munday, A. Canalization of flower morphogenesis in wheat. Proceedings XIth International Congress Genetics 1: 10.31
55. 1963 Frankel, O.H. The role of long range research in agricultural development. Proceedings of the World Food Congress, 4-18 June, 1963. FAO, Rome.
56. 1964 Barnard, C. and Frankel, O.H. Grass, grazing animals, and man in historic perspective. In: Grasses and Grasslands (C. Barnard ed.) (London: Macmillan) pp. 1-12
57. 1965 Frankel, O.H. Agricultural education for research workers. Agricultural Education. (Australian Institute of Agricultural Science, Melbourne) pp. 35-41
58. 1966 Frankel, O.H. Adaptability of crops. New Scientist 31: 144-145
59. 1966 Frankel, O.H. Internationalism in agricultural science. Australian Journal of Science 28: 314-320
60. 1966 Frankel, O.H. The international biological program. Australian Journal of Science 28: 324-325
61. 1967 Frankel, O.H. Guarding the plant-breeder's treasury. New Scientist 35: 538-540
62. 1968 Frankel, O.H. Human welfare and international cooperation. Proceedings of the National Academy of Science of the United States of America 60: 33-41
63. 1968 Frankel, O.H. International collaboration in plant exploration and conservation. Journal of the Australian Institute of Agricultural Science 34: 22-27
64. 1968 Frankel, O.H. Man in the biosphere – an international study. In: Biology in the Modern World (Canberra: Australian Academy of Science) pp. 4-12
65. 1968 Frankel, O.H. and Shineberg, B. The genetic system of basal fertility in wheat. Proceedings 3rd International Wheat Genetics Symposium (Canberra: Australian Academy of Science) pp. 279-281
66. 1969 Frankel, O.H. Pacific centres of genetic diversity. Malayan Forester 32: 356-360
67. 1969 Frankel, O.H. The dynamics of plant breeding. In: Proceedings XII International Congress Genetics (C. Oshima ed.) 3: 309-325
68. 1969 Frankel, O.H., Shineberg, B. and Munday, A. The genetic basis of an invariant character in wheat. Heredity 24: 571-591
69. 1970 Frankel, O.H. Genetic conservation of plants useful to man. Biological Conservation 2: 162-169
70. 1970 Frankel, O.H. Genetic dangers in the green revolution. World Agriculture 19: 9-13
71. 1970 Frankel, O.H. Save the genetic treasuries in the Sabrao region. Sabrao Newsletter 2: 1-6
72. 1970 Frankel, O.H. and Bennett, E. (eds.) Genetic resources in plants – their exploration and conservation. In: Genetic Resources in Plants – Their Exploration and Conservation (Blackwell: Oxford and Edinburgh) IBP Handbook 11, 554 pp.
73. 1970 Frankel, O.H. Preface. In: Genetic Resources in Plants – Their Exploration and Conservation (O.H. Frankel and E. Bennett, eds.) (Blackwell: Oxford and Edinburgh) IBP Handbook 11, pp. 1-4
74. 1970 Frankel, O.H. and Bennett, E. Genetic resources. In: Genetic Resources in Plants – Their Exploration and Conservation (O.H. Frankel and E. Bennett, eds.) (Blackwell: Oxford and Edinburgh) IBP Handbook 11, pp. 7-17
75. 1970 Frankel, O.H. Evaluation and utilization – introductory remarks. In: Genetic Resources in Plants – Their Exploration and Conservation (O.H. Frankel and E. Bennett, eds.) (Blackwell: Oxford and Edinburgh) IBP Handbook 11, pp. 395-401
76. 1970 Frankel, O.H. Genetic conservation in perspective. In: Genetic Resources in Plants – Their Exploration and Conservation (O.H. Frankel and E. Bennett, eds.) (Blackwell: Oxford and Edinburgh) IBP Handbook 11, pp. 469-489
77. 1970 Frankel, O.H. Variation – the essence of life. Proceedings of the Linnean Society of New South Wales 95: 158-169
78. 77a. 1971 Frankel, O.H. The International Biological Program. Frontier Research on our Biosphere. The Australian Science Teachers Journal 17: 13-19
79. 1972 Frankel, O.H. Australia and the international biological program. Search (Sydney) 3: 105-108
80. 1972 Frankel, O.H. Genetic conservation – a parable of the scientist's social responsibility. Search (Sydney) 3: 193-201
81. 1972 Frankel, O.H. Only one earth: the United Nations conference on the human environment, Stockholm, 5-16 June 1972. Search (Sydney) 3: 406-408
82. 1973 Frankel, O.H. Introduction. In: Survey of Crop Genetic Resources in Their Centres of Diversity (O.H. Frankel, ed.) (Rome: FAO/IBP) pp. ix-xiv
83. 1973 Frankel, O.H. (ed.) Survey of Crop Genetic Resources in Their Centres of Diversity (Rome: FAO/IBP) 164 pp.
84. 1973 Frankel, O.H. The citizen in the knowledgeable society. Public Administration 32(2): 128-130
85. 1974 Frankel, O.H. Genetic conservation – our evolutionary responsibility. Proceedings XIII International Congress Genetics 78: 53-65
86. 1975 Frankel, O.H. Base-sterile speltoids – the location of the Bs gene of Triticum aestivum. Proceedings of the Royal Society of London B Biological Sciences 188: 163-166
87. 1975 Frankel, O.H. Conservation in perpetuity: ecological and biosphere reserves. In: A National System of Ecological Reserves in Australia (F. Fenner, ed.) (Report 19) (Canberra: Australian Academy of Science) pp. 7-10
88. 1975 Frankel, O.H. Genetic conservation – why and how. In: South East Asian Plant Genetic Resources (J.T. Williams, C.H. Lamoureux and N. Wulijarni-Soetjipto, eds.), Proceedings Symposium Bogor, 1975 (Bogor: International Board for Plant Genetic Resources and others) pp. 16-32
89. 1975 Frankel, O.H. and Hawkes, J.G. (eds.) Crop Genetic Resources for Today and Tomorrow (International Biological Programme 2) (London: Cambridge University Press) 492 pp.
90. 1975 Frankel, O.H. and Hawkes, J.G. Genetic resources – the past ten years and the next. In: Crop Genetic Resources for Today and Tomorrow (O.H. Frankel and J.G. Hawkes, eds.) (International Biological Programme 2) (London: Cambridge University Press) pp. 1-11
91. 1975 Frankel, O.H. Genetic resources survey as a basis for exploration. In: Crop Genetic Resources for Today and Tomorrow (O.H. Frankel and J.G. Hawkes, eds.) (International Biological Programme 2) (London: Cambridge University Press) pp. 99-109
92. 1975 Frankel, O.H. Genetic resources centres – a cooperative global network. In: Crop Genetic Resources for Today and Tomorrow (O.H. Frankel and J.G. Hawkes, eds.) (International Biological Programme 2) (London: Cambridge University Press) pp. 473-481
93. 1975 Frankel, O.H. and Roskams, M.A. Stability of floral differentiation in Triticum. Proceedings of the Royal Society of London B Biological Sciences 188: 139-162
94. 1976 Frankel, O.H. Biological structure of the landscape. In: Man and Landscape in Australia – Towards an Ecological Vision (G. Seddon and M. Davis, eds.), Symposium Canberra 1974, Australian UNESCO Committee for Man and the Biosphere, Publication 2 (Canberra: Australian Government Publishing Service) pp. 49-62
95. 1976 Frankel, O.H. Floral initiation in wheat. Proceedings of the Royal Society of London B Biological Sciences 192: 273-298
96. 1976 Frankel, O.H. IRRI phytotron – science in the service of human welfare. In: Proceedings of the Symposium on Climate and Rice (Los Baños, Philippines: International Rice Research Institute) pp. 3-9
97. 1976 Frankel, O.H. The time scale of concern. In: Conservation of Threatened Plants (J.B. Simmoons, R.I. Beyer, P.E. Brandham, G.L. Lucas and V.T.H. Parry, eds.) (New York: Plenum Press) pp. 245-248
98. 1977 Frankel, O.H. Genetic resources. Annals of the New York Academy of Science 287: 332-344
99. 1977 Frankel, O.H. Genetic resources as the backbone of plant protection. In: Induced Mutations Against Plant Diseases, Proceedings of a symposium held in Vienna 1977 (Vienna: International Atomic Energy Agency) pp. 3-12
100. 1977 Frankel, O.H. Natural variation and its conservation. In: Genetic Diversity in Plants (A. Muhammed, R. Askel and R.C. von Borstel, eds.) (New York: Plenum Press) pp. 21-44
101. 1978 Frankel, O.H. Value of wilderness to science. Proceedings National Wilderness Conference, Canberra. ed. G. Mosley, Australian Conservation Foundation. pp. 101-105
102. 1978 Frankel, O.H. Germplasm 'preservation'. Plant Genetic Resources Newsletter No. 34: 18-19
103. 1978 Frankel, O.H. Natural resources and technology – evaluation, use and conservation of biological resources. Proceedings 3rd Inter-Congress Pacific Science Association (Bali, Indonesia: July 1977) pp. 303-323
104. 1978 Frankel, O.H. Philosophy and strategy of genetic conservation in plants. Forest Tree Breeding. In: Third World Consultation of Forest Tree Breeding, Canberra 1977, Documents (Canberra: CSIRO) 1: pp. 2-11
105. 1978 Frankel, O.H. Biosphere reserves: the philosophy of conservation. In: Conservation and Agriculture (J.G. Hawkes, ed.) Duckworth, London. pp. 101-106
106. 1978 Frankel, O.H. Conservation of crop genetic resources and their wild relatives: an overview. In: Conservation and Agriculture (J.G. Hawkes, ed.) Duckworth, London. pp. 123-149
107. 1980 Frankel, O. Our evolutionary responsibility. UNESCO Courier, May 1980 pp. 25-27
108. 1980 Frankel, O.H. Trees in the Australian rural landscape. In: Focus on Farm Trees. The Decline of Trees in the Rural Landscape (N.M. Oates, P.J. Greig, D.G. Hill, P.A. Langley and A.J. Reid, eds), Proceedings of a National Conference, Melbourne 1980 (Melbourne:Organising Committee) pp. 14-18
109. 1981 Frankel, O.H. Evolution in jeopardy: the role of nature reserves. In: Evolution and Speciation: Essays in Honor of M.J.D. White (Atchley, W.R. and D.S. Woodruff, eds.) (Cambridge University Press: New York.) pp. 417-24
110. 1981 Frankel, O.H. and Soule, M.E. (eds.) Conservation and Evolution. (Cambridge: Cambridge University Press) 327 pp.
111. 1981 Frankel, O.H., Knox, R.B. and Considine, J.A. Development of the wheat flower: genetics and physiology. In: Wheat Science – Today and Tomorrow (L.T. Evans and W.J. Peacock, eds.) (Cambridge: Cambridge University Press) pp. 167-190
112. 1981 Frankel, O.H. Definition of gene pools. In: Evolution Today. Proceedings of the 2nd International Congress of Systematic and Evolutionary Biology. (G.G.E. Scudder and J.L. Reveal, eds.) Hunt Institute of Botanical Documentation, Pittsburgh. pp. 385-386
113. 1981 Frankel, O.H. Maintenance of Gene Pools: Sense and Nonsense. In: Evolution Today (G.G.E. Scudder and J.L. Reveal, eds.) pp. 387-392
114. 1982 Considine, J.A., Knox, R.B. and Frankel, O.H. Stereological analysis of floral development and quantitative histochemistry of nucleic acids in fertile and base-sterile varieties of wheat. Annals of Botany (London) 50: 647-663
115. 1982 Frankel, O.H. Role of conservation genetics in the conservation of rare species. In: Species at Risk: Research in Australia (R.H. Groves and W.D.L. Ride, eds.) pp. 159-162
116. 1982 Frankel, O.H. Can genetic diversity survive? Advances in Cytogenetics and Crop Improvement (R.B. Singh, R.M. Singh and B.D. Singh, ed.) Kalyani, New Delhi.
117. 1983 Frankel, O.H. The place of management in conservation. In: Genetics and Conservation (C.M. Schonewald-Cox, S.M. Chambers, B. MacBryde and W.L. Thomas, eds.) pp. 1-14
118. 1984 Frankel, O.H. Genetic diversity, ecosystem conservation and evolutionary responsibility. In: Ecology in Practice. Part l: Ecosystem Management (F. Di Castri, F.W.G. Baker and M. Hadley, eds.) (Dublin/UNESCO Paris: Tycooly International Publishing Limited) pp. 414-427
119. 1984 Frankel, O.H. Genetic perspectives of germplasm conservation. In: Genetic Manipulation: Impact on Man and Society (W. Arber, K. Illemensee, W.J. Peacock and P. Starlinger, eds.) (Cambridge: Cambridge University Press) pp. 161-170
120. 1984 Frankel, O.H. Genetic principles of in situ preservation of plant resources. In: Conservation of Tropical Plant Resources, Proceedings Regional Workshop on Conservation of Tropical Plant Resources in South-East Asia, New Delhi, 1982 pp. 55-65
121. 1984 Frankel, O.H. and Brown, A.H.D. Current plant genetic resources a critical appraisal. In: Genetics : New Frontiers, Volume IV Applied Genetics, Proceedings XV International Congress of Genetics, New Delhi, 1983, V.L. Chopra, B.C. Joshi, R.P. Sharma and H.C. Bansal, eds.) (New Delhi: Oxford and IBH Publishing Co) pp. 3-13)
122. 1984 Frankel, O.H., Brown, A.D.H. Plant Genetic resources today: a critical appraisal. In: Crop Genetic Resources: Conservation and Evolution (J.H.W. Holden and J.T. Williams, eds.) (Allen & Unwin, London).
123. 1985 Frankel, O.H. Genetic resources: the founding years [Part One]. Diversity 7: 26-29
124. 1985 Frankel, O.H. Into the second decade: genetic resources and the plant breeder. Proceedings International Symposium South East Asian Plant Genetic Resources, Jakarta, 1985 (K.L. Mehra and S. Sastrapradja, eds.) pp. 26-31
125. 1986 Frankel, O.H. Conservation – science, ethics and society. N.W. Briton Oration, Queensland Agricultural College, 1986, 8 pp.
126. 1986 Frankel, O.H. Genetic resources – museum or utility. In: Plant Breeding Symposium DSIR 1986, Agronomy Society NZ, Special Publication 5: 3-8
127. 1986 Frankel, O.H. Genetic resources: the founding years. Part Two: the movement's constituent assembly. Diversity 8: 30-32
128. 1986 Frankel, O.H. Genetic resources: the founding years. Part Three: the long road to the International Board. Diversity 9: 30-33
129. 1987 Frankel, O.H. Genetic resources: the founding years Part Four: after twenty years. Diversity 11: 25-27
130. 1987 Frankel, O.H. Nikolai Ivanovich Vavilov 1887-1943: a memoir. Plant Genetic Resources Newsletter 72: 1-2
131. 1987 Frankel, O.H., Gerlach, W.L. and Peacock, W.J. The ribosomal RNA genes in synthetic tetraploids of wheat. Theoretical and Applied Genetics 75: 138-143
132. 1988 Frankel, O.H. Genetic resources: evolutionary and social responsibilities. In: Seeds and Sovereignty – The Use and Control of Genetic Resources (J.R. Kloppenburg (Jr), ed.) pp. 19-46
133. 1988 Frankel, O.H. Nikolai Ivanovich Vavilov 1887-1943 a memoir. FAO/IBPGR Plant Genetic Research Newsletter 72: 1-2
134. 1989 Appels, R., Reddy, P., McIntyre, C.L., Moran, L.B., Frankel, O.H. and Clarke, B.C. The molecular – cytogenetic analysis of grasses and its application to studying relationships among species of the Triticeae. Genome 31: 122-133
135. 1989 Brown, A.H.D., Frankel, O.H., Marshall, D.R. and Williams, J.T. (eds.) The Use of Plant Genetic Resources (Cambridge: Cambridge University Press) 382 pp.
136. 1989 Frankel, O.H. Principles and strategies of evaluation. In: The Use of Plant Genetic Resources (A.H.D. Brown, O.H. Frankel, D.R. Marshall and J.T. Williams, eds.) (Cambridge: Cambridge University Press) pp. 245-260
137. 1989 Frankel, O.H. Point of view: perspectives on genetic resources. In: CIMMYT 1988 Annual Report: Delivering Diversity (International Maize and Wheat Improvement Center: Mexico) pp. 10-17
138. 1989 Frankel, O.H. The Keystone international dialogue on plant genetic resources: A scientist's evaluation. Diversity 5: 59-60
139. 1990 Frankel, O.H. Cultivation and conservation. Trends in Ecology and Evolution 5: 129-130
140. 1990 Frankel, O.H. Germplasm conservation and utilization in horticulture. In: Horticultural Biotechnology (Wiley-Liss, Inc.) pp. 5-17
141. 1990 Frankel, O.H. The future of the global genetic resources network: Activation or dissolution? Diversity 6: 59-60
142. 1995 Frankel, O.H., Burdon, J.J. and Peacock, W.J. Landraces in Transit – The threat perceived. Diversity 11(3): 14-15
143. 1995 Frankel, O.H. Brown, A.H.D. and J.J. Burdon. The Conservation of Plant Biodiversity. Cambridge University Press 500 pp.

Alan Walker (he never used ‘Norman') developed pioneering electrophysiological methods to make major contributions to our understanding of mechanisms and energetics of transport of solutes across plant cell membranes. He used giant cells of characean algae to measure membrane electrical potential differences and conductances that were combined with measurements of fluxes of inorganic ions and organic nutrients obtained with radioactive tracers. 

Alan built up a strong research group at the University of Sydney and had many external collaborations that considerably widened the scope of his research in relation to membrane biology of characean cells and well beyond. His considerable skills in mathematical modelling contributed significantly to research led by others on plant-fungal (mycorrhizal) symbiosis, movement of solutes through the cytoplasmic connections between plant cells (plasmodesmata), the membrane transport mechanisms that regulate the release of nutrients from coats of developing legume seeds, and even nutrient cycling in Arctic ecosystems. 

Alan had strong political views and definite aesthetic tastes that included aspects of visual arts (ceramics and impressionist paintings) and music. He was very interested in good food and wine, and other interests included vertebrate palaeontology and bird-watching. He had many friends with similar tastes, who share fond memories of him.

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About this memoir

This memoir was originally published in Historical Records of Australian Science, vol. 26(2), 2015. It was written by F. Andrew Smith and Mary J. Beilby.

Mollie Holman was a biophysicist whose work on the autonomic nervous system and the innervation of smooth muscle was seminal in advancing knowledge of its behaviour at a cellular level. 

She was particularly known for her technical expertise in microelectrode recording of membrane potential from single smooth muscle cells, and the interpretation of their electrical activity, both spontaneous and in response to transmitters released from their autonomic nerves.

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About this memoir

This memoir was originally published in Historical Records of Australian Science, vol. 24(2), 2013. It was written by Elspeth M. McLachlan and G. David S. Hirst.

Written by W.J. Peacock and D. McCann.

• Place in science
• Early life and education
• Early scientific career
• The United States—science and politics
• Australia—an intellectual and cultural home
• Research on morabine grasshoppers
• Stasipatric speciation
• Epilogue
• Awards and positions
• About this memoir
  • Acknowledgements

Place in science

Michael White put chromosomes into evolutionary thinking and made a primary contribution to the emerging neo-Darwinian evolutionary synthesis. He emphasised cytogenetic systems and argued that genic and chromosomal evolution were of seminal importance in the process of speciation and evolution. His major scientific contribution was Animal Cytology and Evolution (1945), a book that summarised, analysed and synthesised current information on animal chromosomes. White held a somewhat parallel place in cytogenetics to C.D. Darlington, whose book Recent Advances in Cytology (1932, 1937) had earlier synthesized observations on plant chromosomes. For many years from the late 1930s there was a lively competition between these two industrious, innovative and self-assertive figures. Undoubtedly White found satisfaction greater than that of science alone when (contra Darlington) he described an achiasmate meiosis in a mantid during his first period of research in the United States in the late 1930s.

White had an enormous personal capacity for research that was matched by a voracious appetite for the literature of his subject. He had an extraordinary ability to absorb and retain essential technical information, and this contributed towards the integrated approach he achieved in his books. White's contributions to Australian science were largely the result of his own research efforts. He was one of the most consistent and effective participants in meetings of the Genetics Society of Australia and of the Australian Entomological Society. Apart from the impressions that he made on students and other research workers in presenting his results and ideas, he contributed more generally to Australian science by taking a leading role in the Flora and Fauna Committee of the Australian Academy of Science in preparing policies for the development of the biological sciences in Australia. His concerns for taxonomic studies on the Australian fauna eventually led to the fauna work initially carried out by the Australian Biological Resources Study. White worked with colleagues such as Fenner, Waterhouse and Ride on these causes, and later was involved in the early plans for the development of a Museum of Australia. He was a strong advocate of the formation of a Research School of Biological Sciences in the Institute of Advanced Studies in the Australian National University.

Particularly in his later years, Michael White appeared a rather formidable person on first meeting. His unique voice and manner of speaking and somewhat craggy appearance belied what was underneath, an emotional, shy and endearing man for whom many people had a real affection. White was a major figure in the fields of cytology, genetics and evolution. In Australia, where he spent the major part of his scientific life, he was one of the principals in biology, one of a small group of individuals who brought new standards of rigour into Australian cytology and genetics. In presentations to professional societies, he set high standards both as to content and in his mode of delivery. He was a fine example for students and young scientists to emulate, although his manner of presentation was so unique that one was always tempted to listen to the way in which he said things rather than to what he was saying. He liked to be the centre of attention, to create a presence, but he had a wonderful sense of humour and his persona was always softened by mischievous comments and wry smiles. White was an erudite, literate man whose writings even on complex scientific subjects were a pleasure to read. Papers, grant proposals and books were always written longhand in the first draft, and more often than not his first draft was a close approximation to the last.

White's career involved work in many countries, but probably he identified principally with Australia and the country of his boyhood, Italy. In his last years Michael showed particular enthusiasm for Italy. He had been honoured by election to the Accademia Nazionale dei Lincei, but in addition he had active research programmes that took him back to Italy several times. He organised a meeting for the Accademia Nazionale dei Lincei and was obviously delighted to be part of the senior scientific community of that Academy during the meeting, which involved visiting scientists from other parts of the world. His principal leisure activity, one of few, was to read the classics of Italian literature, in the language in which they were first published. It was a unique experience to travel in Italy with White and to benefit from the manifest pleasure he derived in imparting some of his knowledge and observations on the history and culture of the country.

Early life and education

Michael White was born in London on 20 August 1910 to James Kemp White and Una Chase White. James White made a modest living tutoring students who were about to enter Oxford and Cambridge universities or the civil Service, in mathematics, Greek and Latin. In 1915, when Michael was five years of age, the family migrated to the Tuscany region of Italy. His father, who held fairly bohemian values, did not favour a formal education for Michael and nurtured his education in the home environment. Although, as described in Michael's own writings, his father could not be regarded as having been a successful man in the formal sense, he apparently was a capable teacher and certainly inculcated a desire for the acquisition of knowledge in his son. Michael learned a great deal of natural history in his surroundings in Tuscany and even at the age of seven made some incisive observations on local insects and their life histories, and so gained an early interest in entomology. He was also a keen observer of the native flora and made collections in a systematic manner. His father fostered his developing interest in natural history by providing him with books on appropriate subjects. He admired his father greatly and was distressed when his father died when Michael was only 14 years of age. Before this the family had moved to southern France close to the Italian border, after five years in Italy. They lived a total of seven years in France, although during this period the major cultural influence remained that of Italy. As well as cultivating an interest in natural history, Michael had by this time formed an emotional attachment to Italy and the Italian people. After his father's death, an uncle supported Michael's interest in natural history and provided him with books on botany. His mother, loyal to her late husband's wishes, arranged that his schooling continue through a correspondence course with the University Correspondence College in Cambridge.

In 1927, after almost three years of secondary school studies by correspondence, Michael returned to London to study for a degree at University College (part of the University of London). He was initially disposed towards a botanical career, but under the influence of the Professor of Zoology, D.M.S. Watson, he became enthralled with the possibilities presented in this area of science. In particular, Michael was influenced by Watson's discussions on evolutionary biology and his interest was strengthened by a young lecturer of the College, Richard Palmer, who added a genetic aspect to Michael's thinking about evolution. In his third year of university life White made a special effort in entomological subjects, perhaps because of his childhood interests. He supplemented his University College studies with courses in entomology at the Royal College of Science, where one lecturer, O.W. Richards, had a particular influence. Richards lectured on evolution and discussed problems of the nature of species and speciation. Michael was awarded a prize for excelling in his third year and in choosing his gift made a decision that was to influence the direction of his whole career. Instead of receiving the Gold Medal to which he was entitled as a top student he chose instead to receive a copy of E.B. Wilson's classic book The Cell in Development and Heredity. Wilson's book influenced his choice of his first research topic. In his book Wilson mentioned that there were seeming contradictions between the genetic and cytological data concerning sex-linked inheritance in birds, so White decided to conduct an investigation into the chromosomes of the domestic chicken. Although he encountered technical difficulties in this study, it did confirm his interest in chromosomal and genetic questions. During these and subsequent studies at University College, he was able to take advantage of the University of London system and attend lectures in other colleges in various aspects of biology and evolution. In this way he was exposed to a variety of evolutionary ideas ranging from Lamarckian to Darwinian.

Early scientific career

White was awarded a Master of Science degree in 1932 for his cytogenetic studies in chickens. Probably the most important outcome of his Master's degree was that he became convinced that he had special interests and abilities in analyzing genetic and evolutionary matters from a chromosomal observational starting point. Given this realisation it was natural that he might turn to the Orthoptera (an insect group including grasshoppers, locusts, crickets, coackroaches etc.) for his subsequent studies because they presented excellent cytological advantages; in particular, large chromosomes.

His first studies also showed that his mind was attuned to the genetical implications of cytological phenomena. He studied the effect of external environmental factors on the frequency of recombination in grasshoppers, first by looking at the influence of temperature on chiasmata frequencies and later at the consequences of x-irradiation. These studies were important because they were experimental in character. This emphasis on experimentation was an underlying theme in all of White's subsequent research. However, what attracted him most were the puzzles presented by cytological observations on natural populations. He believed that seemingly anomalous observations were likely to provide a key to understanding the normal and later often referred to his 'treasured exceptions'. The first unusual situation that attracted him was the strict centromeric localization of chiasmata in Mecostethus, a grasshopper found in some of the sphagnum bog areas within striking distance of London. These investigations brought him into contact with a figure who was to have a major influence on him, namely J.B.S. Haldane.

Haldane had arrived at University College in 1932, already a charismatic figure in genetics and evolutionary biology. Although White later denied that Haldane had a strong influence on his work, there is no doubt that he was greatly impressed by him, perhaps as much by his behaviour as by his ideas. White was to develop into a colourful figure himself. Many of White's acquaintances had the pleasure of listening to Michael in some of his memorable feats of story telling, and many of those centred around incidents involving J.B.S. Haldane. Haldane was unquestionably impressed by White and in 1938 invited him to move from the Department of Zoology to the Department of Genetics for his research, but White chose to stay in the zoological milieu, perhaps indicative of his conviction that one always had to study genetic systems in a biological framework. Also, Haldane's department emphasized population and analytical genetics with a mathematical slant which was not one of White's major interests or strong points. However, White maintained close contact with Haldane, who accompanied him on some of his collecting trips. Rigour in analysis of experimental results was certainly one conviction that White absorbed from Haldane's approach to science.

In these first years of study, White cemented the major directions of his research and intellectual interests for the next several decades. It was clear to him that he wanted to work with chromosomes and that the Orthoptera was an ideal group to work with because they were cytologically amenable. Nevertheless, his interest in chromosomes was not in them as cytological objects per se but because they provided a key for some incisive thinking about genetic systems and the role of genetic processes in speciation and evolution. His analytical and genetical approach to cytology differed substantially from the more descriptive outlook of most other cytologists of the day. In this early period, too, he worked with grasshoppers, one of the three groups of insects that would be his dominant experimental material throughout his research career. In later research, as a consequence of his first visit to the United States, he would add mantids and gall midges.

1932 was an eventful year in White's life. He married Margaret Thomas, a fellow student with similar scientific and political interests to his own. This was also the year that Darlington's Recent Advances in Cytology was published, as too was Haldane's book, The Causes of Evolution. Darlington's synthesis of cytological studies in the plant kingdom was of major importance in cytology and cytogenetics. There is no doubt that it had a strong influence on White, probably triggering a desire to make a comparable mark with the chromosomes of animals, particularly insects. Just as White was impressed by Haldane's behaviour he was also impressed by the demeanour of Darlington. Above all, Darlington's book strengthened White's conviction that chromosomal observations were of value in developing an understanding of genetics and evolution. However, Darlington's approach to cytology, genetics and evolution differed markedly from White's, and several times during their careers they clashed with some relatively intense disputes emerging in the scientific literature.

As a young academic, White briefly became involved politically, and although he was dedicated primarily to his research it is not surprising that, as an intellectually active person, he responded to political concerns of the day. The radical physicist J.D. Bernal had a particular influence in White joining the Communist Party in 1932. Although White was politically concerned, it is clear that he was not a great enthusiast. He was more a supporter than an activist, and was criticised by his political peers for his apparently indifferent attitude. A couple of years later he resigned and joined the British Labour Party.

White went through a troubled period when his marriage effectively ended in 1933 and it was not until 1936 that he fully regained his drive in research and life. At that time he met Sally (Isobel Mary Lunn), whom he would later marry and who for the remainder of his life was an influential and supportive colleague and partner. In 1935, too, he was made a lecturer at University College. In his research he began demonstrating one of his characteristic traits, that of thinking beyond his own immediate experimental studies and placing them in a broader picture, looking for any appropriate generalizations that they indicated. Darlington was of the same conviction but went far beyond the experimental data with many of his generalizations. White was more conservative in this regard and additionally saw the value of experimental cytology in assisting in general understanding. A very good example was work he published on the nature of distributed centromeric activity in the chromosomes of Ascaris, in which he settled contemporary differences of opinion with an unambiguous experimental analysis.

White first synthesised his thoughts about the nature and importance of chromosomes in a small monograph, The Chromosomes, first published in 1937. Although this book did not contain the theoretical insights and major tracks of conceptual thinking that were prevalent in his subsequent book Animal Cytology and Evolution, it filled a niche in the field at the time and went to seven editions. It was published in several languages and for many years was used extensively in universities throughout the world. In this small monograph White signalled his appreciation of the genetic consequences of meiotic events. The book included what was essentially his first public discussion of the importance of chromosomal systems in evolution. This was quite an achievement for a young scientist of 27 years. The publication of his book undoubtedly contributed to his gaining a Rockefeller Fellowship in 1937 to travel to the United States for a period of research and study.

White went to work at Columbia University with Franz Schrader and Sally Hughes-Schrader, two insect cytologists. He took with him some cytological material and slides from his London studies and carried out an analysis of this material that established that the direction of chromosome coiling was basically random. This contravened generalizations made by Darlington. After the completion of this work, White saw no way to take it any further and began searching for new problems. Through contact with one of the Schraders' students, Kenneth Cooper, he became interested in one of the central cytogenetic problems of the day, the question as to whether meiosis could proceed regularly if there were no chiasmata between homologous chromosomes at metaphase 1. Cooper's views differed markedly from those of Darlington, who insisted that chiasmata were essential to the normal and orderly progress of meiosis. White studied meiosis in a male mantid, Callimantis antillarum, and discovered that there was a complete absence of chiasmata in meiosis in the male. White was excited by this discovery and promptly published his findings in the Proceedings of the Royal Society of London, shocking the Schraders with what they viewed as a precipitous approach to science. When reminiscing about working with the Schraders, White always compared their thorough Germanic descriptive approach with his own, which was more dynamic and always genetically oriented.

At Columbia, White made another vital contact. Theodosius Dobzhansky was visiting Columbia from the California Institute of Technology where he worked in association with T.H. Morgan and colleagues in Drosophila genetics. Dobzhansky had just published his seminal book Genetics and the Origin of Species, and already it was obvious to White that he was one of the world's most dynamic evolutionary geneticists. Dobzhansky and White quickly established a rapport and, discovering that they both planned to travel to Mexico on collecting trips, arranged to rendezvous in Mexico City. This they did and participated in a lecture series at the Instituto Politcnico Nacional in Mexico City. White had a vivid memory of one of his lectures in which he discussed sex chromosome systems involving either two or three X chromosomes. He was surprised at the enthusiastic applause that punctuated his lecture, not realising at the time that 2 X's and 3 X's signified brand names of popular Mexican beers.

While in the United States, White met a large number of cytologists and geneticists and was clearly influenced by the high level and freedom of academic exchange of ideas. This stimulating period probably strongly influenced his subsequent decision to write what was to become a major book, Animal Cytology and Evolution. His somewhat idyllic academic experience in the United States as a Visiting Fellow also influenced his later decision to return to work in that country. However, the Rockefeller Fellowship required him to go back to England. He returned accompanied by Sally, who had come over to the United States to spend some time with him towards the end of his stay, while he worked at the Woods Hole Laboratory.

Not long after their return to England war broke out. Although for a while his research work continued at University College, the Zoology Department then closed down. After a brief time at an entomological laboratory in Slough, White was placed as a statistician in the Ministry of Food, where he served for the duration of the war. Michael and Sally had married only a few months before war broke out. An amusing episode in an otherwise sombre year of political crises occurred at this time. Michael found himself standing in a queue in order to acquire the marriage license, and only after some considerable time discovered that he was in fact in the wrong queue. He had unknowingly joined the one for petrol coupons!

So far as his scientific work was concerned, White was frustrated by the inconveniences caused by war-time restrictions. His cytogenetic research languished. He was not intellectually stimulated by his war duties and generally resented the interruption to his career. Nevertheless, this wartime hiatus paradoxically led to one of his most important contributions to science. In the evenings and on weekends he worked on his book, Animal Cytology and Evolution, and he had a manuscript ready by the end of the war. It was published in 1945.

A chapter he wrote for the book Cytology and Cell Physiology (1942) signalled many of the concepts that he further developed in Animal Cytology and Evolution. White recognised that chromosomes were complicated organelles and that ultimately a molecular understanding of them was going to be fundamental to both physiological and evolutionary biology. In the 1942 review he struggled with the fragmentary state of nuclear chemistry and was not able to attach any conceptual meaning to what was then known of nucleic acids and proteins in the chromosome. In this he was not alone. However, this section of his paper was in marked contrast to the masterly sections dealing with structural behaviour of the chromosome. The latter was his field and it is what he understood best. It would not be until the last couple of years of his life that White would fully return again to the recognition that the control of gene expression was central to an adequate understanding of development and evolution. The review was of interest too because, although White had considerable empathy with the wonderful work of the Drosophila geneticists, he was able to question their notion of equating heteropyenotic chromosomes with genetic inertness. White's analysis came from a cytological viewpoint and it was interesting that he did not feel overwhelmed by the genetical studies. His paper contained other important ideas. For example, he concluded that polytene chromosomes of insects' salivary glands and other tissues represented a particular form of endopolyploidy and suggested that there were probably different levels of replication of the chromosome thread in different regions of some polytene chromosomes. Again this was a suggestion stemming from his cytological knowledge of chromosomes, extending far beyond the salivary chromosomes of Drosophila. In this review paper he contributed a penetrating consideration of induced chromosome structural rearrangements in terms of their consequences as mutations. He also emphasised that not all was known in genetics and he pointed to some significant holes in the knowledge fabric.

Animal Cytology and Evolution gathered into a coherent whole a mass of descriptive cytology and conflicting theories of cytogenetics and evolution. In this respect it paralleled Darlington's earlier plant-oriented book, Recent Advances in Cytology. White's Animal Cytology and Evolution was the first critical survey of cytology since E.B. Wilson's The Cell in Development and Heredity published twenty years earlier, the book that initially inspired White to enter the field. It integrated the cytological approaches of Belar, Dobzhansky and Darlington. White's book must be regarded as the foundation of modern animal cytogenetics and it established him as one of the major conceptual contributors to the neo-Darwinian evolutionary synthesis. White examined cytological and evolutionary observations of diverse sources from a genetical viewpoint, and the resulting synthesis probably provided his single most important contribution to the development of modern evolutionary theory.

Animal Cytology and Evolution ranks with Dobzhansky's Genetics and the Origin of Species as one of the seminal treatises in animal evolutionary biology. White's book emphasized that the principles of evolution applied to individual chromosomes and the chromosome complement just as they did to more classical morphological characters. He also stressed that the chromosome complement, principally through its meiotic properties, could influence the course and rate of evolution of any taxon. White had a great understanding of the mechanism of meiosis and its significance, and an almost intuitive grasp of the complexities and consequences of chromosomal rearrangements, especially in regard to speciation. The various chapters of the book testify to what were his major areas of creative study. One of the dominant themes in the book concerned the evolution of sex chromosomes, sex-determining mechanisms, and the phenomenon of thelytoky, the subject on which he concentrated for many of his later research years. Before the war had interrupted White's research, he had published his first papers on sex chromosome mechanisms in both mantids and grasshoppers. He retained an experimental and theoretical interest in sex chromosome systems throughout his career and made many contributions in this area. This topic and his stimulating experiences in the United States were major factors in convincing him to write Animal Cytology and Evolution.

In various editions of the book, as in his research career generally, White's focus was always the chromosome and the chromosome complement. This was true whether he was concerned with a particular aspect of an insect's genetic system, whether he was probing the causes of speciation, or thinking even more widely about mechanisms of consequence for evolution. White personally attached a great deal of importance to the book and put in an enormous effort in later years into producing a second and third edition of comparable quality. The third edition published in 1973 demonstrated that he was attempting to keep abreast of the vast changes that were occurring in genetics with the advent of molecular biology. It revealed his strong interest in molecular analyses of genetic events and showed his determination to embrace these findings within an evolutionary perspective. All three editions of the book bear witness to White's encyclopedic knowledge and familiarity with the published work of animal cytogenetics. White wrote in the preface to his third edition: 'If the present book helps to re-establish chromosomal mechanisms in the centre of the evolutionary stage, the labour of writing it will not have been in vain.'

The United States—science and politics

In 1946 White was elevated to a readership in University College, but he remained dissatisfied with the scientific community in England. He felt deeply the interruption the war had brought to his research career. While writing Animal Cytology and Evolution his ideas were developing rapidly, but he felt he was extremely isolated in the English academic environment. He was becoming increasingly annoyed at what he perceived to be personality cults surrounding a few central figures who dominated the scene in British biology such as Darlington, Mather, Ford, Fisher and to some extent Waddington. As soon as an opportunity presented itself, he travelled again to the United States, looking for a suitable position there. His favourable memories of the stimulating academic environment in the pre-war United States were confirmed by his return visit to the Genetics Department of the Carnegie Institute of Washington at Cold Spring Harbour. It was thus not surprising that he accepted an offer of a job at the University of Texas at Austin, one of the strongest centres of genetics research in the United States. In 1947 White spent six months as Visiting Fellow at the Cold Spring Harbor laboratories, then, accompanied by Sally, moved to Texas to begin a new phase of his career.

For most of the time at the University of Texas White was engaged in productive research. He appreciated the opportunity of working with colleagues such as Patterson, Stone, Wheeler, Griffiths and Wagner. It was during this time that he studied the peculiar meiotic systems of the gall midges, Cecidomyidae, a group to which he had been introduced by the Schraders while at Columbia University. White saw that study of the bizarre meiosis of these organisms could further advance the understanding of regular meiotic systems. He also continued work on the chromosomes of grasshoppers which confirmed for him the pleasure to be derived from working with natural populations of insects, particularly those living in arid environments. While at Texas White commenced regular annual summer collecting trips in the deserts of the south-west. These regular and extensive collecting pilgrimages to outback areas were also to be a feature of his later research work in Australia. In Australia, even in the years of his 'retirement', White mounted major collecting trips into the arid Nullarbor Plain in the heat of summer. His wife, Sally, proved to be his only durable companion on these arduous safaris.

Some of the south-west grasshopper taxa had chromosome rearrangements that were polymorphic in various populations. His study of these, particularly in Trimerotropis, kindled his interest in population cytogenetics. During a prolific period, he published not only his own experimental results but also commentaries on a range of other cytological and evolutionary matters.

But White's time at Texas was to be troubled, and this phase of his career was brought to an end by political issues. The McCarthy era of political witch-hunts had begun. Michael was investigated by the US Immigration Authority because of his connections with the Communist Party during his student days in England. His problems were acute largely because a state law had been passed that required employees of public universities and other institutions to sign an oath indicating that they had never had any Communist affiliations. Of course White was unable to do so and this ultimately resulted in his resignation. In the end it was a matter of White either resigning or being deported. During this period he found himself in an untenable situation at Texas. Once again White found safe haven in the intellectual cocoon of the Cold Spring Harbor laboratories. He took sabbatical leave, without pay, and for a year was personally and materially supported and provisioned at Cold Spring Harbor by Miloslav Demerec, Barbara McClintock and others. This gave the White family some respite, allowing them to regain a measure of equanimity before Michael returned to teach again at the University of Texas. However, shortly afterwards this excruciating and deplorable political saga came to a conclusion when in 1953 the White family voluntarily left the United States for Australia.

Australia—an intellectual and cultural home

Through the efforts of Dobzhansky and other colleagues, in 1953 Michael White was offered an appointment in the Genetics Section of the CSIRO Division of Plant Industry in Canberra. Dobzhansky made contact with Otto Frankel, who had recently been appointed Chief of the Division, requesting his help. Frankel, himself a prominent geneticist, was happy to provide a position for White because he was attempting to build up the research strength of the Genetics Section of his Division. Frankel made the full facts of White's predicament known to CSIRO's senior executive officers, Frederick White and Ian Clunies Ross, who had no hesitation in supporting Frankel in his efforts to secure a position for White. There is no doubt that White would have warranted the appointment through his scientific reputation alone, but they were also sympathetic to the unfortunate political intrusions into his career. It was a courageous decision by Frankel and colleagues because anti-Communist sentiment was also raging in Australia. At Canberra, White was put under no pressure to work on plants. Frankel felt that White's presence and active research would be of general benefit to the programme at CSIRO and it certainly proved to be so, with White providing an international perspective and the setting of new standards for other geneticists. He played an important role in raising the standing of the Division's genetics group as a whole.

White spent three productive years in the Canberra laboratories. In 1956, the final year of his appointment, he began working on the morabine grasshoppers. This proved to be a turning-point in his career. This large group of endemic, wingless grasshoppers would be his central experimental organisms in subsequent years. One piece of work he carried out in Canberra was an extension of his observations on pericentric inversions in the trimerotropine grasshoppers of the south-west United States. He collaborated with another CSIRO geneticist, Fred Morley, in exploring the genetic consequences of polymorphism for pericentric inversions in populations of a species of local grasshoppers. White was proud of this paper which demonstrated his ability to collaborate with other workers who had complementary skills - in this case, Morley's conceptual mathematical thinking. White also unearthed some other remarkable chromosomal rearrangements in the morabine grasshoppers and this resulted in the first of a series of papers published over many years.

Although his work was stimulating and he was well pleased with his colleagues and the environment, White was not completely satisfied at CSIRO. The CSIRO was a purely research institution and he found himself missing the stimulation and pleasures of teaching. During his time at Canberra he received entreaties to return to the United States where his genetical colleagues fully appreciated his capabilities and his stature as a scientist. No doubt they were anxious to redress the wrongs of the McCarthy period.

With McCarthy's decline in late 1954 the United States returned to a saner political environment and White decided to return to a university position where he hoped to experience the stimulation of both research and teaching. He accepted a position as Professor of Zoology at the University of Missouri, which was one of the strong centres of genetic research in the United States, having an impressive history of both Drosophila and maize genetics. But unfortunately it was not a good choice for White. His particular brand of cytogenetics was not well represented there. Both he and Sally found the general culture and religious environment of Missouri not to their liking. White realised that he and the family had been much more at home in the social and scientific environment of Australia. With the intervention of Frankel and Clunies Ross, an invitation came to White to return to Australia as Professor of Zoology at the University of Melbourne. So, after just eighteen months in Missouri, in 1958 White once again found himself a major figure in the Australian genetics community.

White built up a strong genetics section within the Department of Zoology at the University of Melbourne. In 1964 he became the foundation professor of genetics at the University, moving in 1973 to a separate building of which he was, perhaps uncharacteristically, very proud. He put a considerable effort into raising money for the building and insisted that it made a statement about the importance of genetics as a discipline. White built up a first class genetics department, something much needed in Australia at that time. His personal reputation allowed the department to maintain a strong international flavour, attracting a succession of visiting scientists and scholars. Nevertheless, from the perspective of a dynamic research scientist not all was ideal for him at the University of Melbourne. Particularly onerous were the administrative duties associated with the heavy bureaucracy of the University. White was one of the outstanding scholars in the faculty and this was probably under-recognised by the University. During his professorship he maintained an active research effort and found himself in touch with a wealth of cytogenetic opportunity, particularly with the morabine grasshoppers. He also 'travelled and collected' the Orthoptera of other countries, for example in South Africa and Madagascar.

White had a succession of graduate students during this period, some of whom including Ross Crozier, Jon Martin, John Thomson and Graham Webb, went on to academic and research positions in Australia. However, he personally supervised relatively few postgraduate students. This was perhaps due to his single-minded concern with his own research programme and his lifelong tendency to operate as a lone-worker. He required complete freedom in his own work and thinking and granted the same to others. He was not orientated towards building a strong 'school' as such, although he certainly hoped that this would happen naturally, as a by-product of his own inspiration and devotion to the cause. He was driven on by a desire to uncover yet another piece of the genetic puzzle and fit it into the larger picture. Nevertheless, he made a huge contribution to Australian genetics. Apart from his own students, he also influenced others elsewhere in Australia, primarily through his presentations at the annual meetings of the Genetics Society of Australia. By his later years he had developed into a wonderful example of an idiosyncratic but gifted academic. His addresses at the Genetics Society meetings were inevitably of a high standard with new findings from his experimental programme presented every year. Above all they were colourful and entertaining and fired the imagination of his audience. Along with Otto Frankel, Jimmy Rendel and Spinny Smith-White, Michael White stimulated many of us to aspire to excel in our quests for understanding of the mechanisms and consequences of genome dynamics in plants and animals.

Following his retirement from Melbourne in 1975, White accepted a visiting fellowship in the Research School of Biological Sciences at the Australian National University (ANU) in Canberra. Along with David Shaw he was a major figure in an active cytogenetic team headed by Professor Bernard John, an ex-Darlington student. Even though nominally retired, White attracted, on merit, research grants from both Australia and the United States. He published a substantial part of his total research papers during a very active period at the ANU. During this time he also renewed associations with the CSIRO laboratory at Plant Industry in which he had worked in earlier years. The CSIRO laboratory and the ANU collaborated in seminars and the organization of journal clubs. Jim Peacock and his colleague, Elizabeth Dennis, molecular biologists in CSIRO, were able to provide him with another set of complementary skills to probe the evolutionary puzzles of the morabines.

In this period in Canberra, his last major period of research, he also renewed his ties with Italy. In 1978 he was elected a Fellow of the Accademia Nazionale dei Lincei, an honour he prized above almost any other, and he took every available opportunity to lecture and conduct research in the country that along with Australia he had identified as homeland territory. It was in Italy, in what proved to be his last visit, that he was stricken with a cancer that not long afterwards ended his life. From his last visit he returned elated because he had been able to visit Corsica and Sardinia, islands he had long wanted to see.

Research on morabine grasshoppers

Michael White had an extremely wide knowledge of genetics and evolution. He was a walking encyclopaedia of research in cytology and cytogenetics of the animal kingdom; but to many scientists he was 'the grasshopper man'. Many of his major contributions to science and particularly to an understanding of the modes of speciation and evolutionary change came from his research in the Orthoptera. His particular interest was in the short-horned grasshoppers and particularly the Australian group of Morabine species. He began his work on the Morabines in Canberra, interrupted it when he returned to the USA for the period in Missouri, and then re-established it when he returned to Australia as Professor of Zoology at the University of Melbourne. White's research intensified when he returned to Canberra on retiring from the University of Melbourne.

Population Cytology of Moraba scurra

In his first paper on the morabine grasshopper Moraba (later Keyacris) scurra, White analysed hybrids of chromosome races in an attempt to explain their distribution in the field, carrying out both laboratory and field experiments. Here we have once again evidence of White's experimental approach to cytogenetics. This modus operandi delineated him as a major figure in the field. White analysed the heterotic effect of the polymorphisms he had identified and related his work to the extensive studies made in various Drosophila groups. He was intrigued by the positive heterosis that was associated with chromosome rearrangements in Moraba scurra and postulated negative heterosis for hybrids between races with different chromosome numbers, arguing that this could provide a basis for raciation even in small geographic areas. Incidentally, White did often work with the morabines in small geographic areas. He frequently studied these wingless grasshoppers in cemeteries of country towns that provided 'islands' of natural vegetation. More than one citizen in Australia was startled by the figure crawling around on hands and knees in the local cemetery. White summarised his findings on chromosomal polymorphisms and their effects in a major paper in the Cold Spring Harbour Symposium of 1958. This paper marked a milestone in his contributions to primary concepts of population dynamics.

In his prolonged and productive interaction with the morabine grasshoppers, White was fortunate in having a series of colleagues with complementary sets of expertise. Ken Key, an orthopteran taxonomist in the CSIRO Division of Entomology was associated with White for many years. They first published together in 1957 on the grasshopper genus Austroicetes, a genus of some economic importance in Australia, but the Key-White axis really matured during the long-term collaboration on the subfamily Morabinae that provided a series of papers on the systematics, genetics and evolutionary biology of these remarkable species. In his first period in Canberra, as well as working with Ken Key on the taxonomic side, White formed an association with Fred Morley, a stimulating geneticist in the CSIRO Division of Plant Industry. Morley contributed a quantitative and analytical dimension to White's study of the effect of pericentric inversion polymorphisms in natural populations of Moraba scurry. The White-Morley work was published in 1955.

When White returned to Australia in 1958 to take up the post as Professor of Zoology at the University of Melbourne he continued his analyses of the biological effects of various chromosomal arrangements in Moraba scurra, searching for evidence for heterosis and its basis. In collaboration with Richard Lewontin, he constructed a series of adaptive topographies for various inversion frequencies. They published their results in a strong theoretical paper in 1960. Lewontin, one of the brightest of Dobzhansky's students, provided a link back to the Dobzhansky-White interaction of earlier years. Indeed, Lewontin strongly influenced a number of Australian geneticists during his sabbatical period in the late 1950s in Australia. At a personal level White and Lewontin got on exceptionally well. Before White's work on Moraba scurra it was generally thought that chromosomal polymorphisms in natural populations were maintained in equilibrium by simple heterozygote superiority. His research indicated that the situation was far more complex. The research on Moraba scurra, and especially the work carried out in collaboration with Lewontin on adaptive topographies, was extensively cited and discussed in the literature. The Moraba scurra project was one of the most detailed studies of chromosomal polymorphisms in natural populations beyond the standard Drosophila work.

William Atchley, another visitor to Melbourne, provided biometrical analyses in some of the Moraba work, and another visiting scientist, Robert Blackith, collaborated with White in the mid-1960s, applying multivariate analyses to some of the population studies. Of White's students, probably Graham Webb made the major direct contribution to White's own research area, in particular on the parthenogenetic grasshopper Warramaba virgo. Later, in Canberra, Webb was able to bring molecular-biological analyses to bear on the research through his associations with Liz Dennis and Jim Peacock.

These were fruitful collaborations, and in many cases were crucial in the development of White's scientific work, but throughout there was no doubt in the minds of any of the collaborators that White was in charge! Although his own research work always had priority, he did appreciate the contributions of others and collaborated only with scientists whom he respected and trusted.

Parthenogenesis in Warramaba virgo

White's tireless fieldwork on the morabines further paid off when he happened on the first example of a parthenogenetic species of grasshopper. On a field trip in western New South Wales in January 1961, White discovered a population of a Moraba (later Warramaba) species and was surprised that he could locate only females. After a fruitless additional search for males by his observant son Nicholas, White became convinced that this was possibly a parthenogenetic species. No mind could have been better prepared to come to such a conclusion. In an earlier study in Austin on the mantid Brunneria borealis White had described an exclusive parthenogenetic reproduction system and had pondered on the genetic consequences of parthenogenesis for a number of years. He sent off a short note to the Australian Journal of Science about his discovery, which was published in August 1962. White enthusiastically took Ken Key, his taxonomist colleague, to look at the all-female population. Key was initially sceptical that this would prove to be a valid species. However, he was soon convinced that no males were present and provided a suitable taxonomic place for the species, with a joint publication in the Australian Journal of Zoology in 1963.

White's first public announcement of his finding of the parthenogenetic grasshopper was at the Genetics Society of Australia meeting in Sydney in 1962. Many Australian geneticists have fond memories of Michael describing what was then called Moraba virgo, which he had discovered in two populations by that time, in New South Wales and north-western Victoria. The scientific excitement was accentuated by his presentation, and in particular his enunciation of the word 'femalllllllllle', this word sometimes using many seconds of his valuable presentation time! White was clearly fascinated by thelytoky, as he persisted in calling it in the scientific meetings of that time (perhaps because of the resonance he could give to that particular word rather than because of its precise biological meaning). White studied Warramaba virgo not just for its own sake, however, but because he felt that a detailed study of this unusual species would lead him to an understanding of processes involved in regular speciation events.

Ironically, his initial conclusions about the origin of Warramaba proved to be incorrect. Along with his graduate student, Graham Webb, White studied the patterns of chromosomal replication and concluded that the difficulties they had in recognising homologous pads of chromosomes in the genome were due to differential heterochromatisation that had occurred since the origin of the parthenogenetic species. It was a prevalent concept in the cytological literature of the day. Another of White's students, David Porter, suggested that perhaps the puzzling chromosome complement was a result of a hybrid origin of the parthenogenetic species. But it was not until 1975 that Godfrey Hewitt, a visiting scientist at the Australian National University, made the convincing suggestion that Warramaba virgo originated as a hybrid between two particular Western Australian sexual species of the grasshopper. Initially White's work on Warramaba virgo had been only in eastern Australia but he subsequently found that it also occurred on the other side of the arid centre of the continent in Western Australia, where there was a complex of related sexual species. His results were recorded first in 1973 in a paper in Chromosoma. Hewitt, of course, was able to propose a hybrid origin only because of White's extensive data on the sexual species P169 and P196 from the Western Australian location. White, who had earlier dismissed a hybrid origin, gradually came to accept this as a real possibility. His experimental hybridisation studies using bisexual relatives led him in 1977 to publicly admit that the hybrid origin was a distinct possibility, and indeed, a high probability.

White had also surmised that the parthenogenetic species undoubtedly had a single origin and he developed elaborate hypotheses as to the probable point of origin and the rate and directions of subsequent migrations to extant localities. But work with Graham Webb on chromosome banding patterns showed two clones of Warramaba virgo which were clearly different, and White was forced to consider the possibility that there had been two separate hybrid origins of Warramaba virgo. This was confirmed in Canberra, following his retirement as Professor of Genetics in Melbourne. He established a collaboration with Jim Peacock and Elizabeth Dennis at the nearby laboratories of CSIRO's Division of Plant Industry. Here, analyses on repeated DNA sequences established beyond any doubt that there had been more than a single origin of Warramaba virgo and that there were probably many.

Michael White's place in the Australian scientific landscape was paralleled by his seamless fit into the Australian physical landscape. Collecting with him in the harsh sunlight, of the Mulga and Cassia country near Broken Hill left an indelible imprint of a dedicated, excited scientist perfectly at home in that demanding, xerophytic ecosystem.

Stasipatric speciation

In addition to his experimental studies on Moraba scurra and Warramaba virgo, White conducted a third major analysis with morabine species. This was based on the Vandiemenella (formerly Moraba) viatica species group of morabines in the Eyre Peninsula and the surrounding region of central southern Australia. This research sharpened his ideas on the mechanisms by which speciation occurs and provided the stimulus for his last book Modes of Speciation.

White's work on Moraba scurra and his other cytogenetic studies had strengthened his conviction that the chromosomal and genetic system of a taxon was of considerable importance to its future. In thinking about how new species developed, White adhered to the basic genetico-biological view that a species was a collection of individual organisms that could be considered to have an interchanging gene pool, so that a species perimeter was drawn by the limits of freedom of exchange of genetic information. Conceptually, he accepted that the gene pool of a taxon could differentiate into two or more subsequent distinct gene pools, that is new species, with genetic mechanisms playing a primary role as isolating mechanisms. He gradually modified his early acceptance of the generally accepted proposition that geographical isolation was a prerequisite to speciation. In several of his publications, White commented on the complexity of biological mechanisms involved in speciation. He recognised that geographic, behavioural, and genetic and cytogenetic mechanisms could all play a role, and in different incipient speciation complexes these factors could have different weightings.

During his studies on the morabines, with their low vagility, he found that he was dealing with taxa associated with small geographic areas, often intimately interdigitated. In his earliest writings on Moraba scurra it is possible to see that he was moving towards the ideas that genetic barriers could be the major isolation mechanism needed for the development of two subsequently independent gene pools. In the viatica species group in the Eyre Peninsula in South Australia, he was confronted with a mosaic of karyotypic chromosomal systems with very little geographical separation. Not only could he find situations where it seemed that new taxa, as defined by the cytogenetic system, arose from an apparent peripheral population isolate of an existing group, but he found other situations where individuals with a new chromosomal system seemed to have arisen within the distribution of another taxon. He coined the term stasipatric (stationary place) speciation to describe the latter situation as apparently demonstrated in the viatica species group.

Vandiemenella viatica was a regular single species over much of its distribution range, but in the coastal region of central South Australia White found a multitude of chromosomal forms. These forms or races were often contiguous (or parapatric in White's terminology). Because the hybrids could be identified cytologically he was able to determine that in many situations the hybrid zones could be extremely narrow, a matter of one or a few metres. This emphasized to him that chromosomal rearrangements could function as strong primary genetic isolating mechanisms. White saw that chromosomal variants with the appropriate properties did not always occur on the geographic peripheries of the species distribution. Rather, White found situations which he interpreted as meaning that the origin had been within the distribution with a subsequent expansion on one or more fronts. This convinced him that geographic isolation was not a mandatory requirement in the speciation process. He developed his stasipatric speciation concept in a lead article in Science in 1968, realising that in doing so he was throwing up a challenge to what he considered to be an overly narrow concept of geographically based speciation promulgated by orthodox neo-Darwinian contributors to this field such as Ernst Mayr. White developed his ideas further in his 1978 book, Modes of Speciation, where he went to some lengths to explain why we might envisage many different paths of speciation, dependent on chromosomal, genetic, behavioural and other biological factors as well as geographic considerations. In developing his concept of stasipatric speciation, White emphasised his lifelong view of the importance and complexity of cytogenetic processes in population dynamics and hence in evolution.

White, an admirer and colleague of Mayr, felt that Mayr had underestimated the importance of genetics related processes in the mechanisms of speciation and evolution. The viatica group of the morabines gave him the opportunity to illustrate his view of the many factors involved in speciation. Characteristically, he drew on data derived from his own field collections and experimentation. Although White may not have succeeded in achieving a general acceptance of his views on speciation processes and may not have convinced the broad range of taxonomists and evolutionists that stasipatric speciation may occur, he certainly re-established the importance of genetic mechanisms in the isolation processes involved in the generation of species. Mayr, in a thorough review of Modes of Speciation in Systematic Zoology in 1978, paid tribute to White in this regard.

Epilogue

Michael White was one of the most distinguished scientists of his generation to work in Australia. Throughout his career he made important contributions to many aspects of cytology and cytogenetics and to evolutionary biology, including speciation theory and systematics. He had an awesome capacity for unremitting hard work and continued his research activities up until a few days before his death. He died from cancer on the 16 December 1983 at age 73, still at the height of his career. At the time of his death he was acknowledged as the world's leading cytogeneticist. His importance to science is indicated by his membership of many of the world's most prestigious academic societies and in the variety of international honours bestowed on him (see below). White was honoured by a Festschrift on his seventieth birthday (Evolution and Speciation, 1981; edited by W.R. Atchley and D.S. Woodruff).

Michael is survived by his wife Isobel (Sally), an anthropologist who specialises in research on the Australian aborigines. In addition to her own academic work, Sally had an extensive involvement in Michael's field work. Michael is also survived by his three children: son Nicholas, a virologist, son Jonathan, a university lecturer in humanities, and daughter Charlotte, a medical practitioner. Michael White's passing was a major loss to Australian and international science and to family and friends. His legacy, however, is immense.

Awards and positions

Degrees

• BSc in Zoology and Human Physiology (First Class Honours), University of London, 1931
• MSc, University of London, 1932
• DSc, University of London, 1940
• MSc, University of Melbourne, 1959
• Dottore in Science Biologiche honoris causa, University of Sienna, 1979.

Fellowships of Learned Societies

• Fellow, Australian Academy of Science, 1955
• Member of Council, AAS, 1960-1962
• Fellow, Royal Society of London, 1961
• Honorary Foreign Member, American Academy of Arts and Sciences, 1963
• Fellow of University College, London, 1962
• Foreign Member, American Philosophical Society, 1978
• Socio Straniero, Accademia Nazionale dei Lincei, 1978
• Fellow of the Linnean Society of London honoris causa, 1979
• Foreign Associate, U.S. National Academy of Sciences, 1981.

Medals

• Mueller Medal, Australian & New Zealand Association for the Advancement of Science, 1966
• Silver Medal for Research, Royal Society of Victoria, 1979
• Linnean Medal for Zoology, Linnean Society of London, 1983
• Minerva Medal of the University of Rome, 1983.

Academic appointments

• 1976 Visiting Fellow, Department of Population Biology, Australian National University
• Jan.-Mar. 1968 Visiting Agassiz Professor, Harvard University
• Aug. 1964-1975 Professor of Genetics, University of Melbourne
• July 1958-July 1964 Professor of Zoology, University of Melbourne
• 1963 Visiting Fellow, Witwatersrand University, South Africa
• Jan. 1957-June 1958 Professor of Zoology, University of Missouri
• July 1953-Dec.1956 Senior Research Fellow, CSIRO, Canberra
• Sept. 1947-June 1953 Professor of Zoology, University of Texas
• Mar.-Sept. 1947 Guest Investigator, Department of Genetics, Carnegie Institution of Washington, Cold Spring Harbor, N.Y.
• Jan.-Mar. 1947 Reader in Zoology, University of London
• 1940-1945 Wartime positions as Statistician and Entomologist in British Ministry of Food
• 1937-1938 Rockefeller Research Fellow, Columbia University
• 1935-1946 Lecturer in Zoology, University College, London
• 1932-1935 Assistant Lecturer in Zoology, University College, London.

About this memoir

This memoir was originally published in Historical Records of Australian Science, Vol.10, No.2, 1994, and also in Biographical Memoirs of Fellows of the Royal Society of London, 1994. It was written by:

• W.J. Peacock, who works in the CSIRO Division of Plant Industry; and
• D. McCann, who works in the Department of History and Philosophy of Science, University of Melbourne.

Acknowledgements

The authors are indebted to Philip Batterham, Linden Gillbank, Rod Home and Sally White for advice and critical comments on this manuscript.

Written by J. Barrett and R.N. Robertson.

Introduction

Max Rudolf Lemberg, who died on 10 April 1975, was one of the Foundation Fellows of the Australian Academy of Science. Known to a wide circle of friends around the world as Rudi, he was affectionately referred to by his younger Australian colleagues as Lemmy. Born and educated in Germany, he made Australia his home for the latter half of his life and adapted himself well to the ways of his country of adoption. Other accounts of Lemberg's life and influence have appeared elsewhere: his autobiographical chapter written at the invitation of the Annual Review of Biochemistry (1965); Encounter with Rudi Lemberg published privately by his friends (1975); the Royal Society (London) Memoir (1976) compiled by C. Rimington FRS and C.H. Gray.

Early life

Lemberg was born in Breslau on 19 October 1896, into a cultured, educated family. His father was a leading lawyer specializing in civil law and law was in the family tradition, particularly on the maternal side. His younger brother became a lawyer. However, many relatives and close family friends were noted scientists such as Albert Neisser, bacteriologist, Martin Freund, organic chemist, Minkowski and 'Augen' Cohn, medical scientists, and Cohn's son who, under the name of Emil Ludwig, became an internationally known writer. Thus as a small boy he was exposed not only to an intellectual heritage but also to contact with many professional and academic adults. He was educated at the same liberal, humanist gymnasium in Breslau as Bonhoffer, who became an eminent protestant theologian and was later killed by the Nazis. This gymnasium gave excellent instruction in mathematics, Greek, and Latin – much of which Lemberg retained in his later years despite his modest disclaimers. As might have been expected he was more attracted to the intellectual glories of the Greek civilization than to the majestic splendour of the Roman heritage.

Lemberg had a keen love of music mostly of the middle Germanic tradition, music by composers such as Telemann, whose music he greatly enjoyed, Bach, Mozart, and Beethoven. He was not drawn much to the creations of the French and English schools, though he loved Purcell and had some attachment to the works of Britten. He has recorded that there was a great deal of music in his home. His beautiful mother, who was later to die in a concentration camp, had a fine alto voice and his younger brother who also migrated to Australia played the viola and continued to do so until his death two years before that of Lemberg. Though Rudi Lemberg was never a practising musician, he had great ability for analysing music immediately on hearing it.

Aesthetic appreciation was an important component in his mental disposition. Though a somewhat overprotected and frail child, he escaped to experience the joys of the open countryside and the woods beside the River Oder. Thus from childhood and throughout his student days he was keenly aware of the beauty of natural surroundings. In his autobiographical chapter (1965) there are several references to the beauty of the countryside he experienced in all countries he visited. He also enjoyed the colour and rhythm of verse, and frequently read Goethe, Morgenstern, and Rilke. He has recorded that he dreamt in colour and certainly he enjoyed the splendid reds of his porphyrin compounds.

His religious background was Jewish, although not strict or orthodox. His mother, who had a great influence on his cultural life, had a protestant education. He had many contacts with the Lutheran Church and was converted to the Christian faith during World War I. Thus he experienced the flexibility of the Jewish spirit combined with protestant application and uprightness which probably accounted for his strong conscience throughout his life.

War and universities

His high school education veered to the classics but he was impressed by his mathematics master and responded to his teaching. On leaving school he studied chemistry, physics, mineralogy, and geology at Breslau University, but the war came and, believing in the justice of the German cause, he tried to enlist. Twice he was refused on medical grounds and so continued his studies at Breslau, Munich, and Heidelberg until 1917. He was attracted to chemistry but recalled the teaching of this subject in both Munich and Heidelberg at that time as being dull. He enjoyed his period in Munich because of opportunities for skiing and walking in the Alps. In 1917 he was finally able to enlist in the German army. He served in the trenches as a telephonist. Like so many of his time, he gave loyal support to the Fatherland but the brutal experiences in the German army left him a confirmed pacifist with a keen contempt for the military establishment and the mentality of its officers. He was repelled by the brutalization of men by the war, not only by the crudity of the trenches, but also by the senseless cruelty of the officers and sub-officers. He was no coward, being able to face up to the hardships and death of the front line, and was awarded the Iron Cross (2nd class) for a daring attempt to repair a telephone line during the Somme battle of March 1918, being wounded in the attempt.

In 1919 he was able to resume his university work at Breslau University, where he studied methyl-substituted uric acid derivatives with Heinrich Biltz, working towards a PhD. Lemberg remarks that he had not been fitted to become a biochemist. He had had no lectures on biological subjects and, except for a few hours teaching in microbiology and botanical class work, his extensive knowledge of biology was self-taught. Following the award of PhD (summa cum laude) he went to Mannheim in 1923 to work with Bayer, the pharmaceutical manufacturer. Biltz had told him that he was not suited to an academic life and should go into industry. On 21 December 1924 he married Hanna Claussen who was to share the rest of his life in Germany, in England, and in Australia. In 1926 following retrenchment in the prevailing economic crisis, he went to Heidelberg, obtaining a grant from the Notgemeinschaft der Deutschen Wissenschaft which was supplemented by a three-year severance allowance from Bayer. Encouraged by Freudenberg, whose courage and kindness he greatly admired, he began research for his 'venia legendi', a qualification which would permit him to give lectures. Freudenberg recalled recently that he liked Lemberg very much because of his modesty. In Freudenberg's laboratory in the old Institute in the Märzgasse, the organic chemistry was directed to the solution of biochemical as well as physiochemical problems, and there Lemberg had his first contact with the borderland between chemistry and biology which became his future working field. At that time there was a great interest in metallo-organic compounds. Karl Ziegler, later to become a Nobel laureate for his researches on organo-metal complexes, provided many ideas for Lemberg's fertile mind. Hieber was then working on metal carbonyl compounds and Werner Kuhn had begun his studies on the optical rotatory dispersion of macromolecules. Kautsky had recently come from Warburg's laboratory to carry out fluorescence studies which became of importance in the understanding of photosynthesis.

With Freudenberg, whose work on the stereochemistry of organic compounds he admired, he worked on catechins for about six months, but then began independent work on the chromoproteins of red algae, stimulated by Czapek's accounts of them in his Biochemie der Pflanzen. Freudenberg recalls that they ordered a sample of seaweed from Japan, a most precious plant material in Heidelberg, which arrived in two trunks, one of which contained a crab. The algal chromoproteins largely occupied Lemberg's research efforts until 1934 but he retained a lasting interest in these pigments and was particularly pleased when one of his later colleagues (Barrett) took up the study of the interaction of the chromophores with the protein moiety in phycocyanin. Intuitively he recognized that the phycobiliprotein pigments were pyrrole-derivatives and was forced to immerse himself in the work of Hans Fischer whose school at Munich was prodigious in its output of tetrapyrrole literature, sometimes unfortunately premature and incorrect. Lemberg has recorded that there was some disagreement – or polemics – between Fischer and himself over the question of the structure of certain bile pigments. Lemberg and Fischer were antithetical and, though he recognized the immensity of Fischer's achievement, he was more drawn to the imaginative explorations of the Cambridge school of biochemistry, although inclined there to be critical of what he regarded as a lack of chemical thinking by Hopkins and Barcroft and certain of their associates. His Habilitation as Privatdozent at Heidelberg was awarded in 1930 for his demonstration that the prosthetic groups of the algal chromoproteins, phycoerythrin and phycocyanin, were bile pigments. The zinc complexes of the split products were like those of urobilin and 'mesobiliviolin', giving him the first clue that the prosthetic groups of these crystalline chromoproteins were bile pigments. This study also familiarized him with work on proteins.

On Freudenberg's recommendation he applied for a Rockefeller Foundation fellowship to go to the biochemistry department under Gowland Hopkins at Cambridge. At that time Cambridge had outstanding scientists such as Barcroft, Robin Hill, Hopkins, Szent-Györgi, and the Needhams. Lemberg had been greatly impressed by the work of Keilin, Barcroft, and Robin Hill on haem compounds and cytochromes, and elected (1930-31) to continue his studies on the bile pigments rather than to participate directly in the work of the Hopkins school. This decision he later somewhat regretted. However there was interchange of ideas particularly between the departments of physiology and biochemistry and the Molteno Institute. One day Keilin told him that Barcroft had a green pigment in the placenta of the dog. It turned out to be 'uteroverdin', which was identical with oocyan, the green pigment that Lemberg had isolated from gull's egg shells. The 'uteroverdin' was more readily purified than oocyan and was analysed as tetrapyrrolic dehydrobilirubin. In Cambridge Lemberg worked in the same laboratory as Robin Hill, who at that time was doing his brilliant pioneering work on photosynthesis.

Retreat from Germany

Lemberg was a Christian socialist who had been a member of the democratic socialist party. Freudenberg recalls that he displayed no political divergences and no student agitated against him. However the end of his scientific career in Germany came in 1933 when he was compelled by the encroaching Nazi antisemitic oppression to flee from Heidelberg. His English friends were aware of the danger he was in and had sent a message by Szent-Györgi personally for him to leave immediately for Cambridge. Freudenberg had obtained orders from the authorities in Karlsruhe to dismiss him but, though he felt he had to pass on the information to Lemberg, he did not feel he had to serve him notice. However, though Lemberg was a qualified lecturer, he was working only as an assistant and, from that position, Freudenberg was obliged to give him notice. To the Lembergs' everlasting gratitude, Freudenberg gave them shelter in his own home during their last few days in Heidelberg. The Freudenbergs arranged a small farewell party attended by Dr and Mrs Ziegler, Dr and Mrs Kautsky and Werner Kuhn, who was a bachelor. Before his departure they all took a short walk through Heidelberg and accompanied him to the railway station. It had been a pleasant evening but Freudenberg recalls that though the departure was quite unsentimental, they were all overwhelmed and in a very reflective mood.

An associate of David Keilin during his second stay at Cambridge, Lemberg was increasingly aware of the excitement associated with the rediscovery of the cytochromes, first observed by McMunn in the nineteenth century, and of the beginnings of the unravelling of the complex pathways of biological oxidation.

The Australian opportunity for Lemberg to obtain some measure of financial and political security and to be established as a completely independent worker came through the foresight of Dr Wilson Ingram of the Royal North Shore Hospital, Sydney. Dr Ingram, a Scot, of great pioneering spirit, travelled as surgeon to Antarctica with Mawson, and then in the mid-1920s founded the biochemical laboratories that were to grow into the Institute of Medical Research, containing the Kolling Laboratories. There in the crises of 1935 Lemberg found a haven and a base for his future growth as a scientist. The Academic Assistance Council of the UK sought all over the world at that time to resettle academic refugees in academic positions and Ingram had responded to their enquiries. Lemberg, who had been recommended by Sir Frederick Gowland Hopkins as an outstanding scientist with a good command of English, accepted an offer to become a director of the Research Biochemical Laboratories, a position which had been advertised both locally and overseas. Ingram's decision to appoint Lemberg was both courageous and farsighted at a time when xenophobia and a lingering dislike of German nationals, even though many were victims of Hitlerism, still existed in some Australians. The appointment of Lemberg was subject to questions and protests by the Australian Medical Association and chauvinistic scientists; questions were also asked in Federal Parliament. Fortunately, the Australian immigration authorities were more humane and Ingram himself persisted with the appointment and protected Lemberg from those who criticized him for lack of a medical degree. Lemberg and Ingram both originated in the Northern Hemisphere, but were different in personality. They were nevertheless complementary so that their combined organizational and scientific strengths contributed greatly for 40 years to the output of research from the Institute. A bond between them was their love of exploration and both men had suffered the miseries of the western front of World War I.

The experience of travel to the far distant Sydney was a shock for the Lembergs and, during the long ship voyage, provided many doubts on the wisdom of isolating themselves from the centre of western learning and research. However the Australia of the thirties and forties provided a fertile, if unfamiliar, soil for the growth of Lemberg's ideas in biochemistry and gave him an opportunity to provide intellectual leadership in his branch of science. Cast out of his homeland he sought, perhaps, to play a fuller role as an exemplar of liberal humanistic thought and to contribute to his adopted country some of the intellectual heritage of European society. Lemberg was quick to find support for his research and, though funds were modest, he considered himself fortunate to have obtained such support so soon after his arrival, when some others, who were less fortunate than he, had proceeded westwards after their ejection from Germany by the Hitler regime, and had languished alone. In their turn, he and his wife, Hanna, working particularly with the Hon. Camilla Wedgewood, assisted refugees who had come from Europe to Sydney. This involvement with displaced people of that period and from the later upheavals and political disturbances of Europe, with the consequential loss of personal liberty of many scientists, gave the Lembergs much opportunity for challenging humanitarian work. This also caused him to think deeply about the wider issues of human liberty and in his later years the increasing violence and barbarities of the post-war scene. He sought to avoid allying himself to any movement politically inclined to the right or to the left but rather applied himself to the furtherance of humane missions through the Society of Friends.

Early years in Australia

The first few years were difficult, as he had inherited little equipment in his laboratory. The staff consisted of one graduate doing hospital analyses, one research graduate, R.A. Wyndham, and one technician, M. Norris, later to become a distinguished industrial chemist. He had to set about acquiring staff and was fortunate in obtaining the help of two very able younger colleagues in J.W. Legge and W.H. Lockwood who joined him in 1937 and 1938 respectively. At this time his eminence in the area of biological pigments had received international recognition, and he was invited to be author of the first review on animal pigments for the Annual Review of Biochemistry of 1937. The first X-ray crystallographic study on an azaporphyrin – a tetrapyrrole closely related to biological porphyrins – had just been completed by J.M. Robertson. The recognition of its planar structure led Lemberg to speculate on the binding of the iron-complexes of porphyrins to protein as in the oxygen carriers, haemoglobin and myoglobin, and in the haematin enzymes. Here he showed his brilliant insight and the beginning of his later preoccupation with the intimate relationship of the haems with their associated proteins, leading him to emphasize the importance of the conformational changes of the protein moiety of haemoproteins in determining and controlling the reactivity of the central iron of haematin enzymes, particularly cytochrome oxidase.

Those were the days of the threat of Nazism. Legge the Marxist and Lemberg the social democrat argued through the historical antagonism of ideologies to form a united front of two. Lemberg's pleasure at finding someone who spoke the same language and Legge's respect for his intellect developed into an understanding and affection that was life-long.

The advent of war in 1939 and the consequent increased isolation of Australia hindered the development of his laboratory researches. The exigencies of the war, particularly Australia's isolation from her normal sources of pharmaceuticals and other chemicals, provided a stimulus to find effective local means to overcome deficiencies of supply. To that end many notable scientists including Lemberg were coopted to give advice. He and his colleagues carried out a number of ad hoc investigations which arose from wartime needs. It was during this period that he isolated an orange pigment from a fungus gathered on his frequent walks in the Sydney bush. Named by him polystictin (now cinnabarin), it was the only nitrogen-containing fungal pigment then discovered. Lemberg studied its decarboxylation and later a colleague, Dr Peter Clezy (now associate professor of organic chemistry in the University of NSW) established cinnabarin to be 2-amino-phenoxazin-3-one and thus related to the antibiotic actinomycins.

During this period Lemberg intensified his survey of the accumulated literature of tetrapyrrole chemistry and biochemistry for his writing of the text on the haematin compounds and bile pigments – a book which was firmly to establish his authority in the field of tetrapyrrole biochemistry. Within this book he sought to bring into relation the many and varied biochemical manifestations of the porphyrin molecule as well as the linear tetrapyrroles, the animal bile pigments and the algal phycobilins, which had first drawn him into this diverse realm of chemical biochemistry. The book, Hematin Compounds and Bile Pigments, was published in 1949 with J.W. Legge as co-author and some collaboration from J.P. Callaghan. This book was a high-water mark in Lemberg's scientific development and thinking. It led to the shaping of his future major lines of personal research, which included the elucidation of the structure of porphyrin a and his investigations into the complexities of the interaction between molecular oxygen and the two haem a components of cytochrome oxidase (cytochromes a + a₃). Quite early, Lemberg realized and put forward the view that the haem of the haemoproteins must lie in a crevice formed by the polypeptide chain of the protein.

Post-war years

The immediate post-war period brought to the research programmes of Lemberg some more very able men when Ernest Foulkes and John Falk joined his research group. Foulkes later became professor of physiology at Cincinatti, USA. Falk, who made important contributions to our knowledge of the biosynthesis of porphyrins and was the author of a classical book on porphyrins, later became Chief of Division of Plant Industry, CSIRO.

Lemberg committed himself and his colleagues for the next 14 years to the study of the structure of the prosthetic groups of cytochrome oxidase (cytochromes a + a₃), cytochrome a₂ (the terminal oxidase of many bacteria), lactoperoxidase and myeloperoxidase, and the interaction of these haems with their protein moieties. Significant contributions were made to the determination of the structure of these porphyrin prosthetic groups, some of which were first identified and isolated by Lemberg's school. The achievement was all the more remarkable considering the relatively simple apparatus and paucity of technical assistance. Great use was made of the hand spectroscope and the Hartridge reversion spectroscope, supplemented in 1951 by the first manual electronic spectrophotometer. Previously the quantitative spectral analysis – then a vital method in the determination of porphyrin structure – had been carried out using an optical spectrophotometer.

The haem moiety of cytochromes a + a₃, the Atmungsferment of Warburg, proved hard to isolate and purify. Warburg chose to attempt to purify the haem, but Lemberg with his greater command of tetrapyrrole chemistry elected to purify the porphyrin. The task was difficult because of the extreme lipophilic nature of the molecule and the presence of complex lipid impurities from the heart tissue, and even obtaining enough fresh hearts was not easy! However, by the late forties Lemberg in Sydney, C. Rimington (with John Falk) at University College, London, W.A. Rawlinson of Melbourne University (in collaboration with Hale of St Mary's Medical School, London) had achieved the isolation of porphyrin a from heart muscle and bacteria, and had demonstrated that it had formyl and vinyl functions. Appropriately a joint communication was presented, in the presence of the pioneer, David Keilin, at the first International Congress of Biochemistry at Cambridge. The complete purification and crystallization of porphyrin a (as the dimethyl ester) and the definitive determination of the complete structure and subsequent synthesis of the porphyrin at the Cambridge Chemical Laboratories, under Alan Battersby, in collaboration with one of us (J.B., a former member of Lemberg's research group) and at the University of NSW chemistry school under Clezy, required 25 more years during which major contributions to the elucidation of the structure were made by the Lemberg school.

In 1949 Lemberg went overseas for the first time since his arrival in Australia – a span of 14 years. The international biochemical scene had changed greatly since his departure from Cambridge. With the stimulus provided by the post-war revival of biochemical research, there were advances in instrumentation, such as those of Britton Chance in spectroscopy, which were to have a profound influence on the development of dynamic studies in bioenergetics. New vistas were opening like that of Perutz at Cambridge, who had begun his X-ray crystallographic study of haemoglobin, of immense interest to Lemberg. By 1948 also, Shemin and Rittenberg in the USA, and Altman, had experimentally confirmed Lemberg's prediction that glycine and succinate were precursors of porphyrins and had opened up the whole problem of the complex biosynthesis of the natural tetrapyrroles.

Following the first International Biochemistry Congress at Cambridge, where he met several of the English and Continental porphyrin biochemists and investigators into bioenergetics of the cell, Lemberg proceeded to the USA. He was there also to meet several of the leaders of research into porphyrin biosynthesis and into bile pigment metabolism. As a former Rockefeller scholar of the 1930s, he visited the Rockefeller University. He then worked for two months in Chicago with David Shemin whose recent work with Rittenberg had shown by labelling with ¹⁴C that glycine and succinic acid were the precursors of the tetrapyrroles. Shemin(1) has recounted that they sought to identify the C₁ compound that escaped on the splitting of the ring when haem was degraded to biliverdin. He recalls that they mistakenly looked for CHO rather than correctly for CO. Lemberg also encountered Watson and Schmidt of Minnesota who have spoken of their immense admiration for Lemberg's scientific insight, wide-ranging intelligence, and personal charm.

Lemberg journeyed to the west coast visiting Berkeley and the laboratory of Calvin, who was later to receive a Nobel award for his discovery of the C₃ photosynthesis cycle. Although not engaged in photosynthesis chemistry, apart from his earlier discoveries of the nature of the chromophores of the algal phycobiliproteins, Lemberg thought of and contributed to the discussion of the specific features of the chlorophylls which were especially pertinent to the primary act of photosynthesis.

On his return to Australia Lemberg was enabled by a grant from the Rockefeller Foundation to purchase a modern high speed centrifuge and other equipment for a joint investigation with his next generation of colleagues into the bioenergetics of animal and microbial cells. The same year brought to the laboratory the first manual electronic spectrophotometer of high optical resolution. This instrument, and its successors, were to play an important role in the elucidation of the structure of porphyrin. Though Lemberg gradually acquired, and used with great skill, the new equipment that more sophisticated technology made available, he was a master of simple improvization which he maintained could often solve difficult technical problems. He would recount, with some glee, how his analytical chemistry teacher, 'old' Jannash at Munich, had removed the starched shirt cuffs from his wrists to demonstrate their use as shields when grinding refractory silicates in a mortar. He had a delicate, artistic approach to bench work which accounted for much of his success, exemplified by the great skill that enabled him to crystallize the dimethyl ester of biliverdin which had defied purification since its discovery in the 1850s.

Lemberg was catholic in his acceptance of people with innate scientific ability. Not overly impressed by formal degrees, he would give opportunities to those who had not experienced the particular scientific disciplines encompassed by his school. Because of his attitude, two of his colleagues, Frank Moss, a medical bacteriologist (later associate professor of biochemistry, University of NSW) and one of us (J.B.), formerly a microbiologist, were enabled to make significant contributions to the work of the Lemberg school. Lemberg's relationship with his co-workers was that of a gentle aristocrat and savoured of the enlightened court of a nineteenth century German principality, echoing another era. 'Let a thousand flowers flourish' could well have described Lemberg's laboratory in its most creative period.

His undoubtedly high reputation for his work on tetrapyrrole chemistry and biochemistry was recognized by the Royal Society of London which elected him to its fellowship in 1951.

Later years

Because of his dominant interest with the organic moieties of the many fascinating tetrapyrrole-proteins investigated at Royal North Shore Hospital, his laboratory was essentially a natural-product style organic chemistry laboratory until the advent of David B. Morell, a student of David Keilin. Later, the grants of large NIH funds enabled support of research, particularly in the protein field, so that work in the sixties and seventies was directed towards topics in haemoprotein and phycobiliprotein biochemistry rather than porphyrin chemistry per se. Perhaps the turning point can be seen to be marked by the Haematin Enzyme Symposium organized under the auspices of the Australian Academy of Science and held in Canberra in 1959. Lemberg was the president and principal editor of the proceedings; R.K. Morton was the convener, and Falk the local organizer. The proceedings were published under the title of Haematin Enzymes. This international symposium was something of a watershed in the development of haematin biochemistry, bringing together for the first time on this subject, workers from mathematics, physics, and biology. It was important for another reason also, for it was the first time that a party of Japanese scientists had visited Australia since before World War II and marked the beginning of the collaboration between Australian and Japanese biochemists working in this field. The Japanese delegation was headed by Professor K. Kaziro who had been at Cambridge University. Others who attended included Y. Ogura and F. Egami, and the younger, brilliant T. Horio. By then most of the purely organic-type chemistry on the porphyrins and bile pigments in Lemberg's laboratory had been carried out and the later sustained interest in the interaction of their metal complexes – or the free tetrapyrroles in the case of the phycobiliproteins – was developing. Lemberg himself had just returned from a six months visit overseas where, during 1958, he had worked on the biosynthesis of porphyrin a for two months in Rimington's laboratory at University College Hospital, London. He had also visited university departments in Europe, including those of Lynen and Kiese in Munich. He had also lectured at the Academia Anatomica-Chirugica di Perugia (and in 1959 took much pleasure when membership of the ancient Academy was conferred on him, so reinforcing his 'European identity').

That invigorating confrontation in 1959 with many eminent workers on the area of cytochromes and haematin enzymes stimulated his research into the complexities of cytochrome oxidase, for him perhaps the most important of haematin enzymes – or of any – because of its vital role in respiration. This study of a highly sophisticated haem-copper protein complex absorbed Lemberg's attention to the end of his working life. From that time on he was also increasingly involved in international collaboration. In 1966 he was guest professor with Britton Chance at the Johnson Foundation, University of Pennsylvania, where he continued the collaboration with Dr Marion Gilmour who had been a visiting fellow in his laboratory in Sydney. He contributed vigorous discussions on cytochrome oxidase during his USA visit, particularly at the Heme and Hemoproteins Colloquium in Philadelphia (a colloquium dedicated to him) and at the first Gordon Conference on porphyrins which was also attended by four of his collaborators – Falk, Barrett, Sinclair, and Gilmour.

In 1967, accompanied by two of his associates, he went to Japan as chairman of the Symposium on Structure and Function of Cytochromes honouring Okunuki, the leader of haematin-enzyme research in Japan. He prepared himself (aged 70) by learning some Japanese. His preeminence in tetrapyrrole biochemistry was generally recognized and he was sought after by many Japanese scientists both at Kobe and at the Congress of Biochemistry in Tokyo. He was accorded the status of Dai Sensei (distinguished teacher). On his part, Lemberg admired the Japanese scientists for their experimental skills and keen powers of observation. As always, he embraced new experiences with appreciative vigour and attended the classical theatres, One of us (J.B.) remembers both the vigour with which he scaled the hills above Lake Hakone to arrive at the sulphurous plateau of Ohwakidani and to examine enthusiastically the bubbling sulphur vents, and his enjoyment of the solitude at a wayside eating place. In his final year of professional life (1972) he revisited the centres of haemoprotein work in Japan, en route to the symposium on porphyrin chemistry convened by the New York Academy of Sciences.

In his last decade he was particularly engrossed with the problem of the interaction of the two molecules of haem a (not rigidly proven to be identical in every structural detail) and the two copper atoms of the complex mammalian cytochrome oxidase (cytochrome a,a₃). This phase of his life work culminated in a definitive and encyclopaedic review in 1969 in Physiological Reviews, which brought some thousand requests for reprints from many countries. A final and more synoptic account of his views was given in his second book, Cytochromes, with J. Barrett, published in 1973. There he expressed his standpoint that, though the evidence did not support the view that cytochrome oxidase consisted of two separate haem a proteins, there was strong evidence for two different types of binding of the two haem a groups to protein in cytochrome-oxidase. In this view he differed from the eminent and admired Okunuki school. In his later experimental work Lemberg intensively studied, with Professor Ron Williams of Canada and Dr Marion Gilmour of the USA, the ferric state and the oxygenated state (discovered by Okunuki) of solubilized cytochrome oxidase. He concluded that mechanistically the oxygenated state was important as being indicative of a highly reactive, transient Fe³+, or ferryl (Fe IV) state of membraneous cytochrome oxidase in the mitochondrion.

Influence on Australian science

Lemberg had experienced, in the 1930s, the value of the frequent meetings of the British Biochemical Society as a forum of interchange of ideas for spreading the burgeoning knowledge of biochemistry. In Sydney he actively participated in the late 1940s and early 1950s with his colleagues in the (now defunct) Society for Experimental Biology of NSW. From the time of World War II he attended regularly the weekly forum at the Botany School, Sydney University, inaugurated and led by one of us (R.N.R.). The appreciation of this reading group (affectionately termed the bible class!) for the critical discussion and dissemination of current international research, particularly in what is now termed bioenergetics, is attested to by the assiduous weekly attendance of the 'Lemberg group', considering the time and hazards of travel across Sydney. Lemberg's contributions to the education of many younger scientists at that reading group over about 15 years were of great value.

The Australian Academy of Science was founded in 1954 by a group of scientists most of whom were fellows of the Royal Society of London. Lemberg, who had become an FRS in 1952, was much concerned with the discussions leading up to the foundation of the Academy and regarded it as a very important development for Australian science. He subsequently took part in various activities, being a member of Council (1956-58) and vice-president (1957-58). When the Academy began it took over the National Committee for Biochemistry which had been under the auspices of the Australian National Research Council and on which he had been serving. Lemberg was elected to the National Committee for Biochemistry of the Academy and served until 1966 and took part in the activities of the Sectional Committees. He was particularly interested in the Academy building, both before and after it was completed and welcomed the addition of its imaginative architecture among the more conventional Canberra buildings; he referred to it as the 'mushroom'. He regularly attended the dinners of the Sydney fellows' dining club. Lemberg was also active in the affairs of the Royal Society of NSW and was its president in 1956.

The growth of biochemistry in Australia, largely in the capital cities, was brought about by increased funding by the Federal Government through the National Health and Medical Research Council and, later, the Australian Research Grants Committee. It was assisted also by the formation of several special disease-oriented medical funds and by the expansion of that fount of biological research in Australia, the CSIRO. The growing interest in biochemistry and the increasing number of biochemists resulted in a search for a more general venue for an exchange of current biochemical research within Australia. At the national level this had previously been effected to some extent at ANZAAS. Lemberg was a strong supporter of that organization and in 1954 he was president of the section comprising biochemistry and physiology at the ANZAAS Congress. By this time there was a strong feeling that biochemists needed a specialist association to advance their science. Extensive consultation led to the formation in 1955 of the Australian Biochemical Society, of which Rudi Lemberg became the first president, and subsequently its first honorary life member. Recently the principal annual lecture of the Society and its associated gold medal were named after him.

From his viewpoint as director of the Biochemical Laboratories of the Royal North Shore Hospital, Lemberg realized the need to upgrade the standard of clinical biochemical analysis. He saw particularly the need to achieve complete biochemical values for normal subjects to assist the clinician in his diagnosis of pathological states. Following consultation with D. Roman of Adelaide and other clinical biochemists, he encouraged his deputy in the clinical area, F. Radcliff, to collaborate in this task with K.M. Mattocks (Sydney), J. Owen (Melbourne), and D.H. Curnow (Perth). Having provided a stimulus to action, Lemberg left this task to his younger confreres, while continuing to play the role of an elder statesman. He was gratified to see shortly the formation of the Association of Clinical Biochemists – and to become its patron.

Lemberg served on the Advisory Committee of the NH&MR Council for ten years: he was always concerned about the financial plight of the young and promising research worker at the threshold of an independent career. For some years Lemberg was also on the Advisory Committee of the NSW State Cancer Council, where he sought to apply strict scientific principles to the assessment of applications for grants.

It is appropriate to record that, throughout his scientific career in Australia, he was generously supported – in the context of the limited funds available – by grants, first from the National Health and Medical Research Council, and later from the Australian Research Grants Committee. Between 1961 and 1969 the National Institutes of Health (USA), Heart Division, gave considerable amounts of money for major equipment, salaries, and substantial alteration and refitting of part of one floor at the Kolling Institute. The successful development and redeployment of the research within the scientific ambit of Lemberg and his colleagues during that period was dependent on this foreign aid which, thereby, made an important contribution to advancing biochemistry in this country.

It is a significant comment on the university scene in Australia, and in Sydney in particular, that Lemberg received no official recognition from any university until the University of Sydney conferred on him an honorary DSc in 1970, largely due to the representations made by L.C. Birch and R.J.W. Le Fèvre. Though he contributed in many ways to university life, his potential as a teacher and leader of thought in the biological sphere was never recognized by any formal academic affiliation or by a personal chair, such as would have happened in other countries. His scientific eminence was recognized by his peers in this country and especially overseas. In 1956 his old university, Heidelberg, conferred on him the status of Professor Emeritus, and he was a foreign member of the Heidelberg Academy of Science. In 1965, he was awarded the James Cook Medal of the Royal Society of NSW and, in 1971, the Walter Burfitt Prize and Medal of the same Society.

His philosophy

Lemberg thought deeply about life, the significance of man's existence in the cosmic scheme, and his personal role therein. He was a deist and his philosophical thinking was influenced and permeated with the light of his particular understanding of man's existence. Although a protestant and a member of the Society of Friends, shaped by his education and his long association with his gifted and devoted wife Hanna, he had within him elements of the Hebraic faith and would often emotionally identify with the Jewish cause in controversial issues.

As a scientist he thought critically about the physical origin of life, devoting a chapter in each of his books to discussion of the evolution of tetrapyrroles and haematin enzymes. Lemberg accepted the geobiological evolutionary theories of I.A. Oparin, who has expressed his great respect for Lemberg and his exploratory discussions of this topic.

He was preoccupied with the truths of human life. He was not only analytical in his approach but also thought creatively. He was emancipated, humane, and compassionate in his approach to social questions, but elements of the Jewish concept of a just, albeit strict and righteous, God would obtrude into the more generally tolerant vein of his attitudes. His stature as a theological thinker was recognized by the invitation of the Society of Friends of Australia to give the annual Backhouse Lecture in 1966. This lecture 'Seeking in an age of Imbalance' has been widely acclaimed.

His desire for open discussion of the philosophical and sociological, on a real-world plane, led him for many years to participate actively in and often lead the wider discussions provided by the Friday evening forum of the Society of Friends, to which not only senior members of the community came, but also many students. For some young people these discussions, especially in the 1960s, left an indelible impression. Men such as Dr H.C. Coombs, chairman of the Reserve Bank and later chairman of the Australian Council of Aboriginal Affairs, Thomas Keneally, novelist, Charles Birch, biologist and theologian, Peter Mason, physicist, women such as Faith Bandler, aboriginal leader and spokeswoman, and Dorothy Butler, the mountaineer, presented their views on major contemporary issues at these forums, the venue of which was a Meeting House given by Lemberg to the Society of Friends, and set in the beautiful native bush garden of his home at Wahroonga. This Meeting House had been built from the money of his Britannica (Aust.) Prize for Science, presented in 1965.

As his life advanced, he became increasingly concerned with the spiritual crises of western technological man and with the problems increasingly encountered by the Asian countries confronted with the impact of this technology. He raised his voice against the abuse of scientific knowledge, against its misuse by military-economic juntas, and against political power-blocks' ability to lay waste man's environment, and his artistic and social heritage. He was troubled by the use of violence in political and social disputes, and saw in the unbridled uses of violence and torture as political tools the possible ultimate destruction of humane rationalistic western society. As a pacifist, his conscience led him to protest against the dangers to the peace of the world caused by the development of the H-bomb. He deplored the idiocies and cruel suffering of the Vietnamese war. His convictions were such that when he was in his seventies he endured a silent vigil throughout the night outside the Sydney Town Hall.

His character

In his early years, Rudi Lemberg had a private teacher who was a naturalist; his mother also knew the names of nearly all the local species of wildflowers and, in his university years, he received some instruction in botany. Freudenberg recalls he was a gardening enthusiast in Heidelberg where he rented a small garden to grow flowers and vegetables and a few grape vines. His early interests matured into a great love of the flora of the native bush of New South Wales and of the alpine slopes of the Snowy Mountains. With loving meticulousness, he plotted the distribution, and catalogued the identity and appearance of the many wildflowers, bushes and trees that filled the one-acre native bush garden of his home in the hills of Sydney. His special love was for the wild orchids; these beautiful, sometimes solitary plants, he photographed systematically.

Lemberg was a very warm person despite his sometimes austere attitudes, and was sincerely interested in the continuing fortunes of younger scientists with whom he had had association either as supervisor of their research in his own laboratories, as an adviser to a PhD student, or later as examiner. He took pride in their achievements even if they were not his own pupils. On one occasion he was heard to refer to his own scientific entourage as his 'science children' and the students of John Falk, who was then heading a flourishing porphyrin and chlorophyll biochemistry research group in Canberra, as his 'science grandchildren'. If there was a slight possessiveness in this remark, it was far out-weighed by his sense of continuing responsibility to those he had guided in their careers.

He took great interest in imparting his knowledge to his associates, to students, and to children. Though never having formal teaching responsibilities in Australia, he gave many authoritative lectures in the scientific field, characterized by insight and breadth of scholarship and which were delightful and endearing in their delivery.

Lemberg had tremendous powers of concentration, and the ability to exclude all extraneous distractions when working at the bench or writing, to the extent that it was sometimes impossible to break into his consciousness if he was approached without prior arrangement. During the writing of the book The Cytochromes, in his seventies, after a day in his laboratory going through his card index and journals he would work every evening until late at night, collating and writing the text of the chapter currently in hand.

When in deep consideration of some of his favourite scientific principles, Lemberg was oblivious of the mechanical and organizational machinery of society. Colleagues remember, with fond amusement, his presidential address to the Royal Society of NSW. Having filled the blackboard, he sought only briefly for the duster before, to the sudden consternation of the secretary of the Society, he firmly grasped the scarlet side curtains with the very evident intent of putting them to practical use as dusters! Likewise, he was not interested in, or good at, the mundane administrative tasks which he rarely delegated but simply by-passed, believing usually correctly, that someone would take over.

Similarly, Lemberg did not organize the members of his research groups into a coordinated multi-faceted attack on a problem. Unless frustrated by his personal inability to cope with the necessary scale of operations, he worked frequently as a 'one man band' thus allowing his colleagues to work with him or on (sometimes rather distantly) related topics, as they chose. Thus, it was difficult to imagine Lemberg as the motivating head of a university department with all its organizational as well as human needs. Nevertheless, he was exceptional as one who would motivate his colleagues through his ad hoc 'think sessions' in the research laboratory or the discussion group where his leadership stemmed simply from the high quality of his probing and postulating mind.

Never having a large staff, at the most four senior colleagues with an equal number of supporting assistants and the occasional visitor, he was an assiduous bench worker and only in his later years delegated his experiments to others.

Trained as a chemist, though by nature drawn to the enormous variety of form and range of colour of the biological realm, he delighted in the coloured solutions of porphyrins and bile pigments and the formal beauty of their crystallized state. Until his death he retained, and displayed with great pleasure, the first crystals of phycocyanin prepared in 1932 at Cambridge. To his great joy, his wife Hanna, through her sensitive skill, was able to capture the colour and form of the Australian bush in her remarkable tapestries.

Lemberg gave a distinctive and intellectual stature to the science of biochemistry in Australia equalled by only a few others in his time. He brought to his adopted country a rich heritage from the associations of his youth and early manhood. In his life he bore himself with a dignity and was an exemplar of the value of reason and scholarship. He conducted himself with no interest in personal gain or in the acquisition of power. He brought with him the impress of his years at Cambridge where he had grown to love the gentleness and warmth of his many Cambridge friends, some of whom were of the Society of Friends. He invested any office he held with a sense of dignity and purposefulness to deal with the tasks ahead, committing himself with zeal to the furtherance of its cause.

Though Lemberg was sure of his position as a scientist and aware of the esteem with which his work was held by those familiar with the field of haematin enzymes he, nevertheless, bore some evidence of the insecurity of his younger years. This in part had been generated by the economic collapse of the new German Republic and by the destruction of his career in Germany due to the emergence of the Nazi regime. As his wife has noted, the shock of having to fight for his very existence, because he had been publicly stamped as a Jew, contributed greatly to this sense of insecurity.

His insecurity appeared also in later life because he seemed to feel that he had not been accepted by the establishment at Cambridge, despite his two periods in that mecca of biochemists, and he never revisited Cambridge after the first Biochemical Congress in 1949. At some stage in his earlier life, he had prepared a paper for the Proceedings of the Royal Society and it had been rejected and returned with comments which he regarded as unscientific.

Perhaps compensating for this uncertainty made him appear arrogant at times. Though somewhat autocratic in approach and somewhat impatient with specious arguments, which he would contemptuously dismiss, he was not arrogant but rather was committed to maintaining standards of excellence and intellectual probity, sometimes in situations where these standards had been obscured. He was a somewhat reticent person, though accurately described as an intellectual elitist. Although quick to speak his case in any discussion, private or public, he did not appear to be fitted to participate in the hurly-burly of academic politics and certainly would never engage in 'horse-trading' to secure personal or professional advancement.

With the passage of time perhaps his greatest contribution to science (and above all to its quality) in this country may be seen not only in his intellectual scientific achievements, great as they were, but also in the leadership he gave and in the maintenance of the highest standards of conduct and judgement, often before conflicting claims. This he did with firmness but with an essential humility only by which a man can advance the causes of enlightenment of humanity and of even partial comprehension of the universe.

Sir John Eccles, Nobel laureate, has written to one of us:

Rudi Lemberg was one of the most gracious and gentle men I have ever known. In a wonderful way he adopted his new homeland with a deep love and understanding of nature. He became an expert in Australian wildflowers, particularly in the great national parks of the Sydney area. I remember vividly several long walks with him in the enchanting Kuringai Chase, in August, in wildflower time. Out of his love grew his acre of beautifully planted wildflowers and trees in his paradisal 'Sanctuary' at Wahroonga.

He was deeply religious and extremely sensitive to the wonder and mystery of existence. After much effort he was able to develop a philosophy in which science and religion had a complementary relationship at a mystical level. It was a synthesis of vital interest to this present age of disillusionment, where it has been assumed that science had destroyed religion and yet had not replaced it by a system of beliefs whereby human individuals could live in harmony and dedication and face death with serenity.

I had long ago urged him to write his message to mankind, and at last he started, but unfortunately too late. It was to be a great enterprise and was entitled Complementarity of Religion and Science. Alas, he died after completing only 8 of the projected 50 chapters. Here is a brief extract:

'We are creatures of the earth and part of nature, and also made in God's image in a sense deeper than that nature is also God's creation. We are in a special way God's helpmates to whom some creativity has been delegated. We remain as part of nature and can as such enjoy its beauty. The knowledge of the really great scientists has not diminished but enhanced their sense of wonder and mystery. Far from being a hindrance to the freedom of our souls, matter is in fact the complement, providing the handholds and footholds on the mountain of our spiritual climb.'

I believe that this fragment of eight chapters gives a unique message by a great scientist. I hope for a publisher who will link these eight chapters with some six earlier publications by him on religion and science.

About this memoir

This memoir was originally published in Records of the Australian Academy of Science, vol. 4(1), 1978. It was written by:

• Jack Barrett MSc, who was a member of the Biochemical Research Group of the Kolling Institute, Royal North Shore Hospital, Sydney, 1953–73, and is at the time of publication a Visiting Fellow with the CSIRO Division of Plant Industry, Canberra.
• Sir Rutherford Robertson CMG DSc FRS, who was director of the Research School of Biological Sciences, The Australian National University, and is at the time of publication Emeritus Professor of Botany in the University of Adelaide. He was elected to the Academy in 1958, was secretary (Biological Sciences) in 1958, a member of Council 1961-64, and president of the Academy 1970–74.

Acknowledgements

We are grateful to a number of people who helped us with material and comments when we were writing this biographical memoir. Special thanks are due to Mrs Lemberg for her substantial and thoughtful help at all times and to Mrs Katherine Carson, secretary of the Institute for Medical Research at Royal North Shore Hospital, who played such an important role over many years in the research group.

Former research colleagues of Lemberg who have helped are P. Clezy, J.W. Legge, W.H. Lockwood, D.B. Morell and Norma Scott (Newton). Helpful comments also came from C. Appleby and the letter quoted came from Sir John Eccles. We are indebted to Dr G.F. Kolar for obtaining and translating into English the reminiscences of Professor Karl Freudenberg.

Notes

(1) Personal communication.