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  and by Max Blythe in 1993 . Much of the information and views quoted here are from records of discussions I have had with him over many years.
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 , 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  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 , with poor mathematics and next to no science but eight years of Latin and four of Greek. None of his teachers inspired him.
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 . Luckily, at this stage Otto attended a lecture on plant genetics by Professor Erwin Baur, a charismatic personality and lecturer , 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' .
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.
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.
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 .
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.
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' .
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' .
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 , 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.
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.
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 . 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'.
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  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 , 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'.
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.
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' .
For Otto, as for François Jacob, 'genetics became a bastion of reason. To do genetics was to say no to intolerance and fanaticism' . 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  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.
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 .
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  – 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 , 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' . 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).
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 . 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  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 .
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.
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' .
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' .
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' , 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.
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.
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.
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