eric john underwood, 7 september 1905 - 19 august...

19 August 1980Eric John Underwood, 7 September 1905 -

Sir Kenneth Blaxter, F. R. S.

1981, 578-602, published 1 November271981 Biogr. Mems Fell. R. Soc.

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E R IC JO H N U N D E R W O O D

7 September 1905 — 19 August 1980

Elected F.R.S. 1970

By S ir Kenneth Blaxter, F.R.S.

E ric J o h n U n d e r w o o d was born in London on 7 September 1905, the youngest of the three children of James Underwood and his wife, Elizabeth ( nee Lowe) who was the daughter of a Baptist minister. Eric’s great grandfather on his father’s side was a farm labourer, and his great grandmother a domestic servant while his paternal grandfather was a gardener on an estate in Middlesex. Eric’s father, James, was apprenticed to the trade of saddlery and practised as a master saddler in London. In 1906, when Eric was but a year old, his m other died and six months later his father left all three children in the care of his brothers and sisters in London and emigrated to Australia. His intention was to become a farmer and create a home for his children so that they could join him at the first opportunity. James’s first step was to take over a small saddler’s shop and apple orchard at M ount Barker, some 220 miles south of Perth. A long correspondence then ensued with an old friend, Katie Louise Taseman, about the possibility of her joining him. Eventually Katie emigrated to Australia in 1913 bringing with her James’s three children, Gilbert aged 11, Marjorie aged 9 and Eric aged 8. When Katie arrived in Fremantle she and James were married immediately, the ceremony taking place on the lawn in the front of the registry office.

The reunited family then set off by horse and cart for M ount Barker to the wooden shack which was to be their home. An aboriginal corroboree took place that very night outside causing considerable fright to the children, a matter Eric never forgot. Eric started school on his arrival at M ount Barker Primary School. W hen W orld W ar I broke out James closed the M ount Barker business and moved to Perth, taking employment first with a saddle-maker and then with the harness department of a larger company. At this time the family lived in a suburb of North Perth and the children then transferred to the North Perth State School. From this school Eric won an open scholarship to enable him to enter Perth M odern School were he was a student from 1919 to 1923. There he was

579



580 Biographical Memoirs

very happy and developed a particular interest in chemistry which was well taught by his masters in that subject, M r Henry Pearson and M r T . W. Ellison. Eric lodged in Perth at this time and though he liked school he found the succession of landladies unconvivial and even mean. He was lonely and often hungry. M rs Chandler, who was later to become his mother-in-law, once described him as heart-breakingly deprived at this time. On Eric’s outstanding performance in the University Entrance Examinations at the end of his school career he was awarded a cadetship by the W estern Australian State D epartm ent of Agriculture to study at the University of W estern Australia.

C o o ro w

As the motor car slowly replaced the horse in the city of Perth so work for saddlers declined. James gave up his job and became an itinerant worker travelling by horse and trap to surrounding country properties where his skills were still required. Learning of a new land development project arising from the construction of the M idland Railway between Perth and Geraldton, some 310 miles to the north, James’s first ambition came to fruition. For a minimal sum he purchased at Coorow, a property of 2000 acres some 200 miles north of Perth. A small house was built and 120 acres were cleared of scrub before the family moved from Perth in 1920. James, helped by his two sons during their school holidays, then cleared the remaining land, fenced the property, planted cereals and stocked the property with sheep. Eric’s brother, Gilbert, eventually took over the farm, to sell it in 1950 when he retired to live in Perth.

It is pertinent to record the hardship and vicissitudes of life for James and his family in those early days at Coorow. Eric referred to them rarely, perhaps because they were commonplace to his generation, but occasional remarks in his papers relating to the nutrition of man indicate the difficulties he encountered during those times. Erica Underwood has written ‘While the farm was being established life was tough and precarious as there was no capital to call on and the family lived from harvest to harvest and from one wool sale to the next. They could buy little more than staple foods; otherwise living off what they could grow or catch . . . kangaroo, rabbit and scrub turkey.’ The cadetship of the Departm ent was absolutely necessary to enable Eric to have a career at all for the grant paid his university and living expenses and provided vacation employment. Such early difficulties and the comparative isolation of the family must surely have created a deep sense of collective responsibility. Certainly his sister Marjorie, who became a primary school teacher, helped Eric financially later in his career when he went to Cambridge and Eric’s own attitude and concern for his own family may well have had roots in those early experiences.



U n d e r g r a d u a t e st u d ie s at t h e U n iv e r s it y of W ester n A u st r a l ia

At the University Eric first took a degree in agriculture under the tutelage of T . W. Paterson, the first Hackett Professor of Agriculture. Paterson, trained in Edinburgh, modelled his course on that in the Scottish Universities. Underwood, in the Hector and Andrew Stewart Memorial Lecture a half century later, recalled that Paterson was ‘a dour but not unfriendly figure but not as progressive or forward-looking as one could reasonably expect*. In the margin of the typescript of this speech in the University archives is a handwritten note by Underwood ‘A primrose by the river’s brim , A yellow primrose was to him, And it was nothing more*. Nevertheless, Underwood graduated B.Sc.(Agric.) with five distinctions and received the Norman Albert Prize given to the best student in the final year. He also won the Amy Saw Scholarship of the University which enabled him to undertake the honours course. This entailed making a study of Australian pastures, the results being embodied in a thesis which was to become the first scientific publication of the University of W estern Australia. Underwood travelled from north of Geraldton almost to Albany, a distance of about 600 miles, collecting samples of pasture grass. These he analysed botanically and chemically and related these attributes to assessments made by graziers of the quality of the pasture in terms of their stock-carrying capacity. He concluded that differences in pasture quality were related to the protein and phosphorus content of the dry herbage and that the latter both responded to application of superphosphate. This work, remarkable at the time for its appreciation of Australia’s pastoral problems, resulted in his graduation in 1928 with first class honours.

Eric John Underwood 581

C a m b r id g e

Immediately after graduation Underwood was awarded the Hackett Research Studentship and took this up at Gonville and Caius College and at the Animal Research Institute of the School of Agriculture at Cambridge, specifically to continue his studies on pasture. It was arranged that he should work with T . B. Wood, D raper’s Professor in the School, and on his arrival in January 1929 started studies under W ood’s guidance on milk production in ewes. Wood, however, died in November of that year and Underwood’s supervision was transferred to H. E. Woodman. The work he then undertook was part of W oodman’s marathon study of the nutritive value of pasture and entailed cutting 13 pasture plots at four weekly intervals for two successive years and determining chemically 12 constituents in each harvest. This labour was obviously a continuation of studies he had begun in Australia; no doubt it was regarded as good training in application and perseverance! He also attended many of the lectures in biochemistry and physiology for Part II. After being awarded




the degree of Ph.D . he was appointed as temporary assistant in the Institute for a period of eight months undertaking studies on the iron content of feedingstuffs and devising a method for determining sodium. He left to return to Australia in September 1931.

During the following ten years Underwood continued some work on the chemical composition of pastures, much of which was along similar lines to those he had employed at Cambridge. Soon, however, he encountered far more profound problems related to the use of pastures where livestock died or became infertile or cachectic although subsisting on pastures that seemed chemically quite normal.

M arriage a n d f a m il y

Underwood met his future wife, Erica Reid Chandler, daughter of a W estern Australian schoolmaster, when both were scholarship pupils at Perth Modern School. They were close friends at school and later at the University of W estern Australia where Erica graduated B.A. with a teacher’s qualification and a special interest in psychology. After graduation, she spent four years teaching in a country school during which time Eric Underwood was taking his Ph.D . at Cambridge. M rs Underwood recalls that their courting days, after he returned in 1931, were disrupted by Eric’s constant trips in connection with his work on Denmark wasting disease—one and a half days by train each way, and two days in the field. They were married in 1934 and, to use M rs Underwood’s words, ‘For the next forty-six years, we shared all aspects of our lives.’ Their first child, Elizabeth, was born in 1936, their second daughter Jennifer in 1939, and two sons were born, Roger in 1941 and Peter in 1943. In spite of this growing family, M rs Underwood, warmly encouraged by her husband, continued her involvement with education and community work. In the late 1940s, she began a series of broadcasts for the Australian Broadcasting Commission W omen’s Session—she was later to take over the programme and to become widely known to women throughout the State. In 1949, she became one of the first women in W estern Australia to be appointed to the Children’s Court Bench to assist the magistrate. But when Underwood was sent on assignments to Nigeria, to the Philippines and to the West Indies, he was always accompanied by his wife, who helped him with paperwork and with setting up interviews.

In the course of his career, Eric Underwood paid frequent and public tribute to the value of his close and happy family life and in particular to his wife, in whose activities and achievements he took great pride. He shared the domestic work of the family so that she also could develop her career. She became the first woman on the Churchill Fellowship Award Committee and the first woman government nominee on the council of the W estern Australia Institute of Technology, of which she became deputy chairman in 1977. At the time of his death she was deputy




chairman of the W estern Australia Arts Council and held many other appointm ents on committees of importance to the community. In the Q ueen’s New Year’s Honours List of 1978, Erica Underwood was awarded an M .B.E. in recognition of her community work, and in 1981, the W estern Australia Institute of Technology conferred on her an honorary doctorate. The four Underwood children chose careers that repeated the parental pattern of service to their community. Elizabeth (Jill) became a school psychologist, Jennifer a school health sister, Roger a Superintendent of Forestry in W estern Australia and Peter a doctor. All are married and there are fifteen grandchildren.

D e n m a r k w a s t in g disease a n d c o ba lt

In 1921 the attention of the W estern Australian Departm ent of Agriculture was drawn to a malady of cattle occurring in the Denmark area close to the port of Albany. As land was opened up under the group Settlement Scheme it became evident that ‘Denmark wasting disease’ or ‘Enzootic marasm us’ had such a high incidence that the whole resettlem ent programme on at least 5000 acres (2000 ha) was in jeopardy. The disease was characterized by progressive emaciation of the stock with a concurrent anaemia. In May 1928 work on the disease began by A. F. Flood and J. T . Armstrong and in October of that year J. F. Filmer, Veterinary Officer of the Departm ent, took over direction of the work. Underwood joined Filmer in October 1931 on his return from Cambridge.

A very similar disease, ‘Coast disease’, had been noted as early as 1880 on the calcareous sandy dunes of the coastal regions of South Australia. Investigation of this disease was the responsibility of the Division of Animal N utrition of the Council for Scientific and Industrial Research (C .S.I.R .). Studies of Coast disease began under Brailsford Robertson in 1929 who was succeeded on his death in 1931 by Sir Charles M artin, F.R .S. M artin set up a ‘Coast Disease Committee’ consisting of H. R. M arston, F.R .S., L. B. Bull and J. A. Prescott and it was M arston who took major responsibility for the work. There were thus two separate investigations in progress in the early 1930s but no official links between them. This was despite the fact that Underwood’s requests to the State Departm ent for expensive equipment were referred to C .S.I.R .

It was soon realized that Denmark wasting disease was also similar to a num ber of other wasting diseases in other countries, notably ‘Vinquish’ or ‘Pine’ in Scotland, ‘Nakuruitis’ in Kenya, ‘Salt sickness’ in Florida and ‘Bush sickness’ in New Zealand. B. C. Aston in 1928 in New Zealand had recommended the dosing of animals affected with Bush sickness with iron ammonium citrate and some of the Denmark settlers reported improvement in their stock but not lasting cures when they adopted similar measures. Filmer had shown that moving stock from affected land to




normal pasture resulted in cure, that the disease occurred in sheep as well as cattle and had provided a good pathological description of the disease.

W hen Underwood joined Filmer, a farm was leased by the Departm ent on unsound land at Denmark and experimental work began. By 1933 it was established that the disease could indeed be cured by massive doses of iron compounds. These, however, varied in potency. A crude ferrous carbonate imported from New Zealand and found effective in Bush sickness was not entirely satisfactory in curing Denmark wasting disease; limonite, an hydrated ferric oxide from New South Wales, however, was fully effective and was immediately recommended as a practical solution to the settlers’ problems. In this respect, throughout the investigations there was constant political and public pressure on the team to produce cures and on several occasions they were summoned by the State Premier, James Mitchell, to explain why a remedy was not forthcoming.

There were reasons besides the varying efficacy of different iron sources to doubt whether the disease was due to iron deficiency. First, the amounts of iron provided each day were really massive—25 g for a heifer. Secondly, Underwood showed that the iron content of the organs of affected sheep was higher than that of normal animals and that there were no differences between normal and affected farms in the iron content of the pastures they grazed. Additionally, both raw and dried liver, despite containing very little iron, were effective in prevention. Liver was used by Underwood since he was aware that pernicious anaemia in man would respond to the extrinsic factor present in liver; the use of gastric extracts, that is of the intrinsic factor implicated in pernicious anaemia was found to be without effect in sheep with Denmark wasting disease.

Studies on the fractionation of the standard dose of 50 g limonite then began and continued through the winter of 1933. An extraction with dilute acid gave a solution which provided only 35 mg iron and this prevented the disease. The residue from the extraction was ineffective. Extraction of the acid extract with ether resulted in an ‘iron free’ residue containing only 0.2 mg iron and this cured the disease while the iron fraction extracted with ether did not. This work eliminating iron as the preventative factor was followed by fractionation of the limonite extracts by classical methods of group analysis, namely precipitation of acid soluble and insoluble sulphides to separate a ‘copper group’ and a ‘zinc group’ and precipitation of the hydroxides of iron, chromium and aluminium. The filtrate from such sequential precipitations constituted the ‘sodium group’. All fractions were tested using marasmic sheep beginning in April 1934. The ‘copper group’ and ‘sodium group’ were ineffective in curing the sheep, and both the other groups cured the disease. The ‘hydroxide group’ was obviously contaminated with an element from the ‘zinc group’. A series of tests was then made with the latter group, separating in particular and not too successfully a nickel fraction by precipitation with dimethyl glyoxime. Simultaneously nickel




as nickel oxide was tested and found partially effective. This test was unfortunate. Because of the equivocal results with the glyoxime separation it focused attention on nickel. Further work started in June 1934 showed that neither nickel alone, nor zinc alone, nor zinc and nickel together cured the disease. The effectiveness of the nickel oxide must therefore have been due to a contaminant. In December 1934 the contaminants of the nickel oxide preparation, manganese, cobalt and zinc and also nickel, were tested in pairs. The results showed that nickel and cobalt were completely effective and experiments in January 1935 provided complete confirmation of the essentiality of cobalt.

Simultaneously, M arston in South Australia was taking another approach. It was at first suspected that coast disease was due to phosphorus deficiency but M arston and his group also investigated limonite to find it not wholly effective. Following a suggestion of R. G. Thom as, a geochemist attached to the Division of Animal N utrition in Adelaide, that the concentrations of transition elements would be low in these calcareous soils, they showed that these and other elements were present in limonites from various sources. M arston then reported early in 1935 that ‘we have gathered very suggestive evidence that cobalt—in m inute doses—exerts markedly beneficial effects in sheep suffering from the extreme symptoms of coast disease’. E. W. Lines, M arston’s colleague, published a preliminary note giving the protocols of two experiments in which ‘coasty’ sheep were given cobalt (1935). From internal evidence from Lines’s paper, these experiments commenced in October 1934.

W hen Underwood and Filmer published their work also in 1935 they acknowledged the few weeks of priority established by this publication by Lines, but Underwood wrote many years later ‘Despite this acknowledgement M arston was very unhappy and made many unfair allegations which led to some acrimony lasting for many years.’ After a further lapse of time, there is no reason to dissent from the conclusion which R. L. M. Synge, F.R.S., reached when he prepared the biographical memoir on Hedley M arston in 1967. Referring to the Adelaide and Western Australian work he wrote ‘There were during these stages of the work, personal exchanges of information about progress by the two groups which were rapidly moving towards the same conclusion. Simultaneous publication should have been concerted.’

The demonstration of the dietary essentiality of cobalt for ruminant livestock was soon confirmed throughout the world and led to control of these characteristic wasting diseases in many countries. Underwood continued with studies involving nickel showing that if dietary concentrations of cobalt were very low then a synergism with nickel could be demonstrated. He showed unequivocally the difference between affected and normal pastures in the Denmark area in terms of cobalt concentrations in herbage and animal tissues, ruled out the possibility that




concomitant copper deficiency contributed to the enzootic marasmus syndrome and with remarkable prescience concluded that the effectiveness of liver in preventing the disease must be due to a factor for whose formation cobalt is necessary. This was based on his observation that liver ash was not effective in providing a cure and that the concentrations of cobalt in an effective dose of dried liver provided less than 0.02 mg cobalt when his estimates of daily requirement showed the requirement of the element to be in excess of 0.1 mg. It was not until 1948 that it was demonstrated that vitamin B12 was a cobalt containing compound; and it was even later that cobalt deficiency in ruminants was shown to be a vitamin B12 deficiency induced by failure of its microbial synthesis in the rumen through lack of cobalt.

There were, of course, many difficulties for Underwood to overcome in tackling the Denmark wasting disease problem. He found his knowledge of chemistry somewhat lacking and during the early period he consulted D r Eric Watson of the Perth Technical College where there was a considerable interest in separatory techniques associated with the mining industries. More of a problem was the provision of analytical equipment. He urgently required a microcolorimeter costing £30, for he had to borrow one from Perth Hospital who in turn had to borrow it from a private practitioner, D r Breidahl. The W estern Australian Departm ent could not afford to buy the instrum ent and wrote in July 1932 to the C .S.I.R . State Committee asking for help. A massive correspondence then ensued over the principle of C .S.I.R . giving funds to a State Departm ent and eventually Sir Charles M artin was asked first whether it was really necessary for Underwood to have a colorimeter and secondly if Adelaide could lend him one. Sir Charles said ‘yes’ and ‘no’ to the questions and eventually, several months later, C .S.I.R . bought the instrum ent and lent it to Underwood. The loan was not forgotten; Underwood, while in the U.S.A. in November 1936, was asked to return it to the Trustees.

T he U n iv e r sit y of W is c o n s in

In December 1935, Underwood applied for a Commonwealth Fund Service Fellowship (now the Harkness Fellowships). The reason was obvious. G. L. Sutton, then Director of Agriculture for Western Australia, wrote at the time ‘Unfortunately this Department cannot offer facilities for further pursuing the investigation into the relation of cobalt to metabolism.’ Sutton also stated that the work on cobalt had been stopped and that ‘Underwood has been detailed, as Senior Investigator, to a large scale investigation into the nutritional aspects of Toxic Paralysis (Botulism) of sheep and other farm animals in W estern Australia . . .’ Although it would clearly have been inconvenient to replace him, Sutton nevertheless agreed to support Underwood’s application. He was also




supported by Sir Arnold Theiler who considered U nderwood’s work ‘to be of immense importance to a more thorough understanding not only of the so-called “ iron deficiency diseases” of animals generally but also possibly those of m an.’

Underwood was successful in his application and he and his wife and their infant daughter, Elizabeth, spent two years ending in June 1938 at the University of Wisconsin where he worked with Professors E. B. H art and C. A. Elvehjem. There he attempted to produce cobalt deficiency in rats and had considerable difficulty in producing diets low in the element. He concluded, in an unpublished report, that rats were in no way as susceptible as were ruminants to deficiency of the element.

T he Underwoods spent some time touring in the U.S.A. and were driven by W. M. Neal of the Florida Agricultural Experimental Station at Gainesville virtually all over the Florida peninsula inspecting the work being done on ‘salt sick’ cattle, work which stemmed from the Australian discovery of the essentiality of cobalt.

Underwood noted the different approach used in graduate training in American universities and at Cambridge where he had studied and concluded ‘In our Australian universities graduate work on any scale is only just developing. Such influence as I have will definitely be thrown towards some blend of the two systems rather than too slavish an imitation of English methods towards which there is a definite tendency at present.’ It may be remarked that Underwood had very few Ph.D. students when he achieved that influence; his blend was possibly more English in flavour than American.

Attempts were made by E. B. H art to encourage Underwood to stay in the U.S.A. At the same time it is evident from correspondence that Sir David Rivett, Chief Executive Officer of C .S.I.R . was considering Underwood for a post in the Animal Health Division of the C .S.I.R . at the M cM aster Laboratory, Sydney. Difficulties of financing such a post arose and Underwood returned to W estern Australia.

He did not, however, return directly but, supported by a grant from the Science and Industry Endowment Fund, visited Onderstepoort Veterinary Research Station, Pretoria, S.A. He spent two weeks there and two weeks travelling to see something of the livestock problems of the South African veldt.

Toxic p a r a l y sis or b o t u l ism in sh eep

At the time that Underwood was investigating enzootic marasmus he was also conducting investigations with F. L. Shier and H. W. Bennetts on another disease which was seriously limiting sheep production in W estern Australia. A fatal botulism occurred during the dry summers to disappear with the onset of rain in the autumn and the growth of green herbage. Bennetts had shown the involvement of Clostridium botulinum in




the disease and also that it arose because sheep ate rabbit carcases infected with the organism. The carcases were present because of compulsory poisoning of rabbits, usually by fencing off water dams and providing cyanided water for the rabbits. The reasons for the pica were unknown; by analogy with the pica associated with the aphosphorosis (lamsiekte) in South Africa, phosphorus deficiency was suspected and phosphorus supplements were advocated. In a series of studies over several years Underwood showed that phosphorus deficiency was not the primary cause, that the phosphorus requirements of sheep were much lower than was hitherto thought and that the depraved appetite could be prevented by giving adequate supplementary feed during the critical summer months. No specific nutrient deficiency was involved; the pica was the result of semi-starvation. He pointed out, incidentally, that the sheep only consumed the flesh of the dead rabbits not their bones. In aphosphorosis, oesteophagia is the characteristic behavioural sign.

Supplementary feeding of the sheep to prevent this depraved appetite was, however, too expensive a process for farmers to adopt. In collaborative work with L. B. Bull, Chief of the Division of Animal Health of C .S.I.R .O ., preventative vaccination was tried. Bull produced the toxoid vaccine in Melbourne and Filmer and Underwood tested it in 1935-36 in field trials with success. It was widely used in the 1930s to 1940s, indeed until subterranean clover was used extensively to improve the quality of the grazings and thus avoid the depraved appetite, consequent upon undernutrition of the sheep.

T he pr a c tic a l n u t r it io n of sh eep

During the war years Underwood was closely involved in studies designed to increase sheep production under W estern Australian conditions, and with T . J. Robinson and Shier he began a series of investigations designed to increase the fertility of ewes. Among factors studied were those of increasing feed provision before mating to influence ovulation rate, increased feeding in late pregnancy to affect the subsequent size and growth of the lamb and the extent of breed differences in the duration of anoestrus and in fertility. Additionally, a num ber of studies were made of the vitamin A reserves of sheep and the vitamin A content of their milk in relation to the season and to the |3 carotene content of the species of pasture plant on which they grazed or which was conserved as hay for them to eat. This work was largely concerned with the application and extension of established work to the grazing and animal husbandry conditions of W estern Australia. It was essential to farming progress; it did not, however, lead to major new discovery but rather to a closer appreciation of the determinants of increased productivity.




S u b t e r r a n e a n clover

D uring the early 1930s pasture improvement in the S.W. Division of W estern Australia relied heavily on the use of an early flowering strain of subterranean clover, Trifolium subterraneum L. var. Dwalganup. The use of the clover doubled animal production per hectare. Suddenly, in 1941— and the suddenness was surprising—breeding problems appeared in ewes grazing these clover-dominant swards. H. W. Bennetts of the Agricultural Departm ent, W estern Australia, described the field condition. It consisted of dystocia, infertility in the ewe, lactation in virgin females and castrated males, and uterine prolapse in older ewes. Underwood was directly involved in the experimental work to elucidate the causal factors involved and eventually the problem became of such magnitude that the D epartm ent, the C .S.I.R . and the Institute of Agriculture pooled their resources and established a Technical Committee with Underwood as Chairman to further the work. W hen it was realized that the problem of infertility was not restricted to the clover belt of West Australia but was occurring in South Australia, Victoria and New South Wales, the Commonwealth Government provided funds for the purchase of a farm on suitable land where work could be done.

Underwood, with Bennetts and F. L. Shier who was a close friend throughout his life, provided a complete clinical and pathological description of the disease concluding that concordance of observations and particularly the presence of a cystic endometrium in affected ewes suggested excessive oestrogenic activity. The unequivocal proof that this activity was present in subterranean clover was first obtained with guinea pigs and later with sheep. D. H. Curnow and T . J. Robinson then joined the team and they showed that a crude ethyl ether extract which was oestrogenic could be obtained from the clover. The syndrome could be induced by administration of diethyl stilboestrol and the effects of both stilboestrol and the clover oestrogen could be modified by simultaneous administration of androgen. In further biological work, Underwood and his collaborators showed that in most respects the clover oestrogen when ingested functioned like a natural oestrogen but, unlike oestradiol, it did not interfere with ovulation or lead to follicular atresia: the infertility in the sheep was not due to a failure of oestrus. They showed that drying reduced the oestrogenic activity of the clover and defined the minimal dose levels in terms of stilboestrol equivalents necessary to induce the syndrome. Considerable effort was exerted to isolate the oestrogen. Curnow visited the Courtauld Institute to work with E. C. Dodds, F.R .S., to characterize the oestrogens in the ether extracts and showed that they were not steroids. Eventually, it was shown that activity was associated with presence of genistein (5,7,4'-trihydroxyisoflavone). Biological assay of different varieties of subterranean clover, however, did not always give results in accord with chemical determination of geni-




stein, and it was then shown by R. B. Bradbury, D. E. White of the Organic Chemistry Department of the University of W estern Australia and by A. B. Beck that other isoflavones were present, notably formononetin (7-hydroxy-4'-methoxyisoflavone) and biochanin A (5-7 dihydroxy-4/-methoxyisoflavone). Concentrations of the isoflavones differed from variety to variety.

The discovery of the oestrogenic potency of the naturally occurring isoflavones was an achievement of considerable importance to the farming of the dry lands of Australia. Considerable work then took place in Underwood’s department to find ways of using subterranean clover effectively, including studies of genetic improvement and cultural practices. Underwood, while obviously associated with all this work, was particularly involved with studies related to the use of subterranean clover in rotations. He showed that wheat grown on land which had been under subterranean clover for 8-11 years was higher in N and thiamin content than that from land frequently cropped and that the higher gluten content of wheat grown following subterranean clover led to improvement in the bread making properties of W estern Australian wheat and hence its market acceptability.

Most of the work, however, was concerned with finding or producing isoflavone-free subterranean clovers. Underwood set up a joint programme of the University and the W estern Australian Departm ent of Agriculture leading to the establishment of subterranean clover seed collections in both centres and eventually varieties low in formononetin content emerged from the breeding work.

It may be remarked that Underwood’s Institute, realizing the importance of legumes in W estern Australian agriculture, studied alternative species including those of Medic ago, Trifolium ornithopus, Vicia and Lupinus. These legumes were tested at the Institute of Agriculture not simply for yield and agronomic characteristics but also for oestrogenic activity to show that another isoflavone, coumestrol, was present in the medics and this too had mild but discernible oestrogenic activity. A photograph of 1951 shows Underwood among the cultivars of subterranean clover at the Institute and he formulated the overall programme of investigation of ways in which pasture legumes could contribute to W estern Australia’s wheat and animal industry.

T he U n iv e r sit y a n d th e In s t it u t e of A g r ic u ltu r e

While working for the State Department, Underwood gave many specialist lectures in the University, indeed the University Department, through shortages of staff, relied heavily on help of this nature from several sources. In 1946, he was appointed from a large field of applicants to the Hackett Chair of Agriculture and to the Directorate of the Institute of Agriculture and he held these twin posts for 25 years until his




retirem ent late in 1970. The Institute had been founded in 1938 by a release of funds from the Hackett T rust to provide a permanent home for the University Departm ent of Agriculture and it was in the same year that the Departm ent was granted Faculty status. Underwood was thus also Dean of Agriculture.

His first task was to accommodate a massive increase in undergraduate num bers swelled by returning ex-service men. Numbers rose from 16 before the war to 104 in 1948 and thereafter more slowly. The degree course was altered to abolish the obligatory year spent in farm work and became a bachelor’s degree of four years’ duration which Underwood insisted should be broadly based, and a Ph.D . degree was instituted in 1949. Eventually, four departments were created within the Faculty and some specialization introduced at undergraduate level to accommodate growth of new knowledge, but the premise he adopted was that those trained in agriculture should be acquainted with all its parts. He himself had a considerable lecturing commitment. Many of his lectures are preserved in the University archives and Professor Quirk, appointed to the chair of Soil Science in the Faculty in 1963, has written ‘The only complaint I have heard about his lectures was from a student who said he lectured too well. The student had been so intrigued that he forgot to take appropriate notes.’

Obviously, as Dean, Underwood was heavily involved in university administration at this time; he was chairman of the Professorial Board, President of the University Staff Association and representative of the staff on the Senate. Additionally, he was deeply committed to wider aspects of agricultural education. In 1949, he had been chairman of a Government Committee on agricultural education and extension which produced plans for its reorganization and these plans were adopted in W estern Australia. Later, in 1952, he was asked by the University of Sydney to advise on the development of teaching and research in animal husbandry and again these suggestions were adopted. On his sabbatical visits abroad he rarely failed to enquire about aspects of University organization and teaching and their relevance to his own University. In 1957, for example, he spent 5^ months in the U.S.A. visiting 27 universities and research establishments in 33 states, laconically remarking in his report ‘This was a fairly full programme.’ Aware of the load of committee work in the University of W estern Australia, he commented on the power wielded by his American counterparts, ‘In the U.S.A. there is less democracy and more efficiency.’ Nevertheless, he advised caution before accepting the American system as a way of ‘keeping our professors doing what they are appointed for, namely teaching and research’!

T he f u n d in g of u n iv e r sit y research

As important to Underwood as the establishment of sound undergraduate training was the provision of adequate funds for research within



the University. Indeed, it was central to his whole view of the role of the universities that, as he often put it ‘The universities have not only to teach they have also to learn’. W hen he succeeded to the chair he regarded it as essential to provide funds to further research. He was instrumental in negotiating a number of research grants to enable Research Fellows to be appointed and additionally a num ber of bequests were made, the terms of which reflected the high regard in which the University was held. By 1953 the University of W estern Australia’s Institute of Agriculture had the largest graduate research group of any university department in Australia.

T he S o il F e r t il it y R esearch F u n d

In his first report as Director, in 1948, Underwood wrote of ‘the importance of the wheat industry to Australia, the vital part played by wheat and its mill products in the nutrition of human and stock populations, and the necessity for thorough scientific investigation of the agricultural and technological factors involved in the wheat industry’. He described, too, the studies which he and his colleagues had begun at the Institute to further this research for the benefit of the industry in W estern Australia.

The effect of this work showed when in 1951 the Trustees of the W heat Pool of Western Australia donated £10000 to the Institute to assist research into soil fertility and in 1953, the W estern Australia Millowners Association agreed to donate the initial sum of not less than £2000 each year for three years to support the same broad programme. But in July 1953, Underwood devised a scheme which was later to be followed by the country as a whole in the whole area of research support. In a newspaper article in the Western M ail in April 1952 Underwood suggested that to provide ‘the invigorated programme of research and extension the wheat industry needs’ a levy of one penny a bushel on all cereals marketed should be paid into a trust fund to provide finance for research. The response from farmers and farmers’ unions was considerable and favourable but a penny a bushel seemed too much for some, others thought farmers without a wheat acreage might benefit without paying, still others wondered whether the levy was to be taxed. Underwood accommodated by reducing his suggested penny to three farthings and then to between a farthing and a half-penny. Eventually the Wheat Section of the Farm ers’ Union of Western Australia agreed to impose a voluntary levy of a farthing a bushel on all wheat sold in Western Australia and in 1954, the Soil Fertility Research Act was passed by the State Parliament on the basis of these negotiations. Before the Act was passed there were numerous difficulties of a legal nature to be overcome and the inevitable law’s delay relating to whether it was constitutional to impose a levy under the existing State Wheat Marketing Act, who should collect it and who should pay legal costs if farmers refused to pay.





Underwood was deeply involved in all these negotiations as judged by a voluminous correspondence both privately and through the press. In the first three years, almost 90% of wheat growers contributed voluntarily to the scheme. The money from the levy was placed under the control of four trustees, one of them being the Director of the Institute of Agriculture, to be used for the promotion of soil fertility research at the Institute. This Soil Fertility Research Fund, Underwood estimated, should provide not less than £30000 annually to assist research and remove the insecurity which had handicapped the financing of research projects at the Institute.

Three years later, in 1957, after the passage of the W heat Industry Research Acts through the Federal Parliament, a countrywide levy of a farthing per bushel was made on all wheat to promote research and advisory work. The Commonwealth Government contributed an equal amount and the funds were distributed by the M inister for Primary Industry, on the recommendation of a central W heat Industry Research Council, to Research Committees in each state. It was only proper that Underwood was appointed representative of the universities on the W heat Industries Research Council when it was formed in 1957 and he served on it for ten years. The same pattern of levy support of research was subsequently developed for other commodities.

T he u n iv e r s it y a n d C.S.I.R .O .

In 1965, research funds were made available to universities throughout Australia by the Australian Research Grants Committee and Underwood was appointed to be a member of the committee. He regarded the work of this committee as invaluable to the universities for the advancement of scholarship and scientific research and when in 1966 he had necessarily to reduce his commitments to enable him to serve on a part-time basis for six years on the Executive of the Commonwealth Scientific and Industrial Research Organization (C .S.I.R.O.), he stated to Sir Stanley Prescott, his vice-chancellor, that membership of the Research Grants Committee was one appointment he would think it unwise to relinquish. There ensued what may be called Underwood’s championship of the university case for more active participation in national research. Real problems had to be faced. There were constitutional ones related to the use of Commonwealth funds for the support of State universities and the Commonwealth institution, C .S.I.R .O ., was thought by some to be better able to study national problems than purely State institutions. Underwood clearly resented the fact that C .S.I.R .O . had been able to obtain funds to develop first class research institutions in both the biological and physical sciences while the universities had been lamentably unsuccessful, and said so at the Canberra conference on Science in Australia in 1951. He continued this approach in correspondence with




C.S.I.R .O ., indeed one letter in reply by Otto Frankel refers to his ‘inevitably pugnacious’ approach to educational and research problems. Perhaps Underwood’s view of the organization of Australian science is best expressed in the John Falk Memorial Lecture given in 1973, three years after he had retired but while still a part-tim e member of the C .S.I.R .O . Executive. He commented on

‘the slowness with which the Australian community has come to recognize the essential functions of a university. . . . It has a duty to enrich as well as to transmit and conserve knowledge. This slowness is perhaps a reflection of a persistent anti-intellectualism in this country, linked with the peculiarly Australian predilection for support of government departments and agencies with the opportunities they provide for bureaucratic control. . . W hat can be stated with certainty is that the universities should have sufficient research resources to ensure that the staff are able to pursue their specialities at the boundaries of knowledge with reasonable financial security. In this way they can remain stimulated and informed and the training of graduate students in research can be conducted in an ambience of real and active research vital to its success.’

Underwood’s championing of the university cause did not, however, prevent him taking a full part in the activities of C .S.I.R .O . As Director of the Institute he frequently requested funds for research for his staff from C.S.I.R .O . and these were invariably granted. From 1949 to 1956 Underwood was chairman of the W estern Australian State Committee for C .S.I.R .O . and in 1966 he was appointed, initially for a period of three years, to be a member of the Executive of C .S.I.R .O . This appointment was regularly renewed until 1975. On his retirement from the Chair at Perth, he was asked by D r J. R. Price (later Sir Robert Price), chairman of the C .S.I.R .O ., to review the whole of their research programme in agricultural and biological fields. This was an enormous task. It involved review of a wide spread of projects and disciplines involving some 400 research scientists located in more than 40 laboratories and field stations throughout Australia. The review took almost 14 months to complete, and Underwood’s report to the C .S.I.R .O . Executive concluded with some 33 recommendations, the majority of which were accepted and implemented. Sir Robert Price refers to this report and its implementation as marking a turning point in the development and documentation of C .S .I.R .O .’s agricultural and biological research activities and says that it undoubtedly contributed to the continuing viability and relevance to national needs of the Divisions concerned.

R esearch at th e in s t it u t e

Administration, the search for funds, his national activities in agriculture and his lecturing commitments did not prevent Underwood from continuing to undertake investigational work. Besides the studies on




subterranean clover, Underwood instigated and took an active part in other researches in a num ber of fields. W ith R. J. M oir he embarked on studies, continued by M oir in collaboration with others, related to the role of the symbiotic ruminal flora, particularly as a source of protein for the sheep. The biological value of the microbial protein was determined and its limiting amino acids identified as methionine and isoleucine. A ttention was drawn to the high sulphur content of ruminal bacteria, the sulphur largely consisting of that in cystine and methionine. A large requirem ent of sulphur by the rum inant could thus be inferred. The group also showed relations between dietary carbohydrate components and the mass of bacteria in the forestomachs of the sheep. This work was contemporaneous with a considerable international effort to elucidate the role of the microflora in rum inant metabolism, and demands for funds for such work by Australian research workers were considerable and competitive. Underwood’s application in 1947 to the Wool Research T rust Fund on behalf of M oir for funds to investigate seasonal effects on the ruminal flora met with the reply from Clunies Ross who was the Executive Officer of C .S.I.R .O . that Hedley M arston’s Division of Biochemistry and General Nutrition was already investigating the matter. Underwood’s reply was that it would be quite useless to write to M arston so to avoid duplication since ‘the reply would be unsatisfactory in the sense that it would claim that everything was already “sewn up” or about to be . . . This may sound a little hard, but it is a statement based on experience and not on prejudice.’ Underwood got the grant!

During the late 1940s and early 1950s Underwood undertook a series of studies already referred to and mostly with J. W. Lee to provide information about the nutritive value of wheat and its milling products extending these to include effects of previous cropping policies on the nutritive value of the grain. In the 1950s and in the 1960s his interest in the trace elements again became evident in his work.

Arising from his earlier studies on phytate in cereals, Underwood realized that zinc could become limiting in poultry diets high in phytate, and he surveyed Australian feedingstuffs to ascertain their adequacy as zinc sources. Then, in association with M. Somers, he began a series of studies to extend to sheep observations made with other species that indicated that testicular function was impaired in zinc deficiency. There was an additional practical problem in that Underwood thought that part of the loss of wool in sheep in Australia during drought might be due to zinc deficiency. Zinc certainly limited crop growth on millions of acres of land, notably the Esperance Plain area of W estern Australia. Somers and Underwood found that the zinc required for testicular growth and spermatogenesis was greater than that needed for growth. Further studies showed changes in RNA and DNA contents of testicular tissue and it was suggested that a primary role of zinc was to modulate ribonuclease activity.



I n t e r n a t io n a l a c t iv it ie s for F.A.O.

In 1953, Underwood was invited to lead the Australian delegation at the F.A.O. Conference on Agriculture in Asia and the Far East, held in Bangalore, India. He was asked to act as Nutrition Expert to the F.A.O. mission to Nigeria in 1963, and in 1965 accepted a two-month assignment as Food and Nutrition Policy Consultant to the West Indies. His task there was to assess, and suggest improvements for, the use of the resources in the Caribbean and to study and suggest solutions to the problems of nutrition encountered there. At the same time, he was requested to prepare plans for setting up a nutrition training and research centre. It was, as Underwood wrote in a letter to the Vice-Chancellor of his University ‘A challenging and formidable assignment’. His success in it was made evident when he was invited to undertake a similar task in the Philippines in 1971. Here, as F.A.O. Food and Nutrition Policy Consultant, he prepared a coordinated plan for the Government of the Philippines designed to achieve satisfactory food production and consumption levels by 1974. Additionally, Underwood served as a member of the W .H .O ./F.A .O . Expert Panel on Human Nutrition and on the F.A.O. Expert Panel on Animal N utrition from 1964.


T race elem en ts in h u m a n a n d a n im a l n u t r it io n

A review of trace element metabolism which Underwood published in 1940 entitled “ Trace elements in nutrition’’ and a chapter with the same title published in Biochemistry and physiology of nutrition in 1953 were both well received and may well have prompted Underwood to write his textbook Trace elements in human and animal nutrition. This was first published in 1956. It was completely rewritten for publication in 1962 and the two subsequent editions entailed considerable extension and revision. Underwood maintained a wide correspondence with many workers in the fields concerned so to ensure that each edition was fully up-to-date and that any omissions from the published work on which the account was based were made good in the account of it he gave. His command of the field and his considerable facility in summarizing and integrating work from many sources necessarily meant that in his later years he received many invitations to lecture on the nutritional aspect of the trace elements. He was in the process of writing the introductory paper for a Royal Society Symposium on the metabolic and physiological consequences of trace element deficiency in animals and man im mediately before his death. The holograph manuscript of this last paper is preserved in the University of W estern Australia archives, and at the Symposium many of the lecturers paid tribute to his contributions to




knowledge. Shortly before his death he was also engaged in the organization of the 4th International Symposium on Trace Element M etabolism in Animals which was held in Perth, W estern Australia, in May 1981. At that meeting the high regard in which his work was held was once more evident in remarks that were made.

Even so, the tributes on those occasions were more than an expression of appreciation of Eric Underwood’s scientific accomplishment and organizing ability. They reflected also the high regard in which he was held as a man. He had a complete and uncompromising integrity, he was fair in all his dealings with others and, though not gregarious by nature, had a facility to establish relationships with people from very different walks of life. His energy, his directness in conversation and in his writing and the efficient way in which he worked equally reflect his character. He was an early riser, awake long before his family at 6.30 in the morning, to spend a full day at the University. Each evening he retired to his study for at least two hours, but never to work on administrative matters but to read and to write. The latter part of the day was always spent in conversation with his wife for the relationship between Eric and Erica Underwood was an uncommon one. It was rich in understanding and to both stimulating and rewarding.

Both his father and stepmother were Christian Scientists and Eric had been reared in that faith. This faith he abandoned when in his ’teens and in his later years avowed that he had escaped from things of religion to those of reason and a more direct concern for others. Something of this early upbringing may well have remained, for though not adhering to the Christian Scientists’ view of illness and disease, he regarded these as somewhat akin to weakness, and his reaction to his own illness was almost to deny its existence. He suffered two serious ischaemic seizures in the 1950s which resulted in angina of effort, the pain of which he controlled with nitroglycerin. Few knew of this and he made no concessions to it by changing his pattern of life. M uch later the pain could not be ignored and he was admitted to hospital where surgical restoration of the coronary circulation was successfully performed. A major cerebral haemorrhage occurred, however, and he died on 19 August 1980.

Peter Underwood, Eric’s youngest son, has written to me of his father ‘If from my account a picture has emerged of a dry individual obsessed with work and duty it would be a very incomplete picture. While not a bon viveur he was ebullient in manner and character. He was an excellent raconteur, with a fund of witty and wise stories and one who, especially as he got older, enjoyed convivial company especially amongst his large and devoted family. He was proud in different ways of each of his children and their families. Above all he loved to laugh and, after his death, we have all noticed with surprise and profound sadness the gap left by the loss of that explosive and infectious laugh, somehow as big, as strong and as uncompromising as the man himself.’




H o n o u r s a n d a w a r d s

Underwood was elected to Fellowship of the Australian Academy of Science in 1954 and following receipt of the Medal of the Australian Institute of Agricultural Science elected to Fellowship of that body in 1958. He was elected to Fellowship of the Royal Society in 1970.

He was made a C.B.E. in Her Majesty The Queen’s Birthday Honours List in 1963 ‘for services to agriculture, education and administration’ and in 1976 was made an Officer of the Order of Australia (A.O.) ‘for a contribution of exceptional merit to agricultural teaching, research and administration ’.

Eric Underwood’s contributions were equally acknowledged by the award of honorary degrees by the University of New England (1967), his own University of W estern Australia (1969), the University of Melbourne (1973) and the University of Wisconsin (1980). Additionally, he was Kelvin Medallist of the Royal Society of W estern Australia, Burnett Medallist of the Australian Academy of Science, and he received the A.N.Z.A.A.S. Medal in 1975 for his ‘contribution to the advancement of science in Australia’. He was awarded the Klaus Schwartz Commemorative Medal posthumously in 1981.

I am grateful to the C .S.I.R .O ., the Commonwealth Fund and the University of W estern Australia all of which made available information from their archives relating to Professor Underwood. Among individuals to whom I am indebted for helpful material and information are: M r P. F. Butler; Professor D. H. Curnow; D r D. V. Frost; D r A. MacPherson; Professor R. J. Moir; D r E. C. Owen; D r A. E. Pierce; Sir Robert Price; M r W. J. Ridgman; Professor J. P. Quirk, F.A.A.; Ms Christine Shervington; D r W. R. Stern; Professor R. Street, F.A.A.; Professor J. W. Suttie; Professor R. L. M. Synge, F.R.S. and Ms Sheila Widra. M rs J. Boyne gave much assistance in collating material and preparing the reference list.

In particular, I would wish to acknowledge the indispensable help given by D r Erica Underwood and her family.

The photograph is by courtesy of C .S.I.R .O . Australia.

Bibliography

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1932 (With H. E. Woodman) Nutritive value of pasture. VIII. The influence of intensive fertilising on the yield and composition of good permanent pasture (seasons 1 and 2). J. agric. Sci., Camb. 22, 26-71.

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1941 (With F. L. Shier) Studies in sheep husbandry in Western Australia. I. The effect ofnutritional flushing on fertility of sheep. J. Dep. Agric. West. Aust. 18, 13-21.

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1942 (With F. L. Shier) Studies in sheep husbandry in Western Australia. II. The influence offeeding during late pregnancy on birthweight, growth rate and quality of the lamb. J. Dep. Agric. West. Aust. 19, 37—47.

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(With D. H. Curnow & F. L. Shier) Further biochemical studies of pregnancy toxaemia in sheep. Aust. vet.J. 19, 164—173.

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(With R. J. Moir) Studies in cereal hay production in Western Australia. II. The influence of time of cutting upon the chemical composition and digestibility of wheaten and oaten hay. J. Dep. Agric. West. Aust. 21, 41—51.

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(With H. W. Bennetts & F. L. Shier) A breeding problem of sheep in the South-West division of Western Australia. J . Dep. Agric. West. Aust. 23, 1-12.

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(With T. J. Robinson & D. H. Curnow) The manganese content of Western Australian cereal grains and their by-products and of other poultry feeds. J. Dep. Agric. West. Aust. 24 259-270.

(With A. J. M illington) Further studies with cereal hays in WA. Dep. Agric. West. Aust 24, 250-259.

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(With C. L. Bangs) A comparison of methods of estimating sperm concentration in ram’s semen. Aust. vet. J. 24, 89-92.

(With R. G. Coleman & D. H. Curnow) Carotene losses in curing and storing cereal hay. J . Aust. Inst, agric. Sci. 14, 175-179.

(With J. W. Lee) The total phosphorus, phytate phosphorus and inorganic phosphorus content of wheat, and its mill products. Aust. J. exp. Biol. med. Sci. 26, 413-418.

(With L. E. A. Clare) The thiamin and riboflavin contents of wheat and its mill products. Aust. y. exp. Biol. med. Sci. 26, 413—418.

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(With J. W. Lee) The influence of variety on the thiamin and nitrogen contents of wheat. Aust. y. exp. Biol. med. Sci. 28, 543-548.

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1951 (With H. W. Bennetts) The oestrogenic effects of subterranean clover—uterine maintenancein the ovariectomised ewe on clover grazing. Aust. J. exp. Biol. med. Sci. 29, 249-253.

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(With R. J. Moir) Aspects of protein nutrition in animals. J. Aust. Inst, agric. Sci. 19, 214-221.

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Trace elements. In: Biochemistry and physiology of nutrition vol. II (ed. G. H. Bourne & G. W. Kidder), pp. 427-504. New York: Academic Press.

1956 (With R. J. Moir) Protein utilisation in the ruminant. In: Proc. 7th International GrasslandCongress, New Zealand, 1956, pp. 241-250.

Trace elements in human and animal nutrition. New York: Academic Press. Second edition, completey rewritten 1962. Third edition, revised and enlarged 1971. Fourth edition, revised 1977.

1957 Plant oestrogens in animal breeding and production9 Schiff Foundation Lecture, CornellUniversity, U.S.A. October.

Trace elements in animals. In: Trace elements. Proc. Wooster, Ohio, Conference, pp. 241-250. New York: Academic Press.

Trace elements in ruminant nutrition. Aust vet. Jr. 33, 283-286.1958 (With D. R. Grant-F rost) Zinc toxicity in the rat and its interrelation with copper. Aust. y.

exp. Biol. med. Sci. 36, 339-346.1959 (With A. J. Anderson) Trace element deserts. Scient. Am. 200, 97-106.

(With F. L. Shier, N. D avenport & H. W. Bennetts) Further studies of the effects of prolonged injections of stilboestrol on the ewe. Aust. vet. y. 33, 84-90.

Mineral metabolism. A. Rev. Biochem. 28, 499-526.1960 Animal production, y. Aust. Inst, agric. Sci. 26, 153-169.

Minerals in pastures: soil-plant-animal interrelations. 7th Int. Congr. Soil Science, Madison, Wisconsin, U.S.A.

1961 Recognition and correction of deficiency states. Fedn Proc. Fedn Am. Socs exp. Biol. 20,284-290.

1962 Agricultural organization for tomorrow: research and extension.^. Aust. Inst, agric. Sci. 28,267-273.

A preliminary statement on the sources of zinc for poultry in Australia. In: Proc. 12th Wld’s Poult. Congr. Sydney, vol. I, pp. 216-218.

Trace elements and the nutrition of sheep. In: The simple fleece (ed. A. Barnard), pp. 133-144. Melbourne University Press.

1963 Clinical and physiological aspects of the trace elements. In: Proc. 6th Int. Congr. of Nutrition,pp. 289-312. Edinburgh: E. & S. Livingstone.

(With E. H. Morgan) Iron in ruminant nutrition. I. Aust.y. exp. Biol. med. Sci. 41, 247-253. 1966 The mineral nutrition of livestock. Aberdeen: Central Press for FAO and CAB. Second edition,

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1967 The role of science in the development of the animal industries of Australia. Inaugural A. S. Niven Memorial Address: The University of New England.

Hedley Ralph Marston. Rec. Aust. Acad. Sci. 1, 73—78.Man, land and food. Aust. J. Sci. 29, 395-403.

1969 (With M. Somers) Ribonuclease activity and nucleic acid and protein metabolism in the testesof zinc-deficient rats. Aust.J. biol. Sci. 22, 1277—1282.

(With M. Somers) Studies of zinc nutrition in sheep. I. Aust. J. agric. Res. 20, 889-897. (With M. Somers) Studies of zinc nutrition in sheep. II. Aust. J. agric. Res. 20, 899-903.

1970 Geographical and geochemical relationships of trace elements to health and disease. In: 4thAnnual Conf. on Trace Substances in Environmental Health, Columbia, Missouri (ed. D. D. Hamphill), pp. 3-17.

The cobalt story. Rec. Aust. Acad. Sci. 2, 23-28.Progress and perspectives in the study of trace element metabolism in animals. In: Proc. 1st

Int. Symp. on Trace Element Metabolism in Animals (ed. C. F. Mills), pp. 5-21. Edinburgh:C, & F. Livingstone.

1971 The history and philosophy of trace element research. In: The newer trace elements in nutrition(ed. W. Mertz & E. Cornatzer), pp. 1-18. New York: Marcel Dekker.

1973 Trace elements. In: Toxicants occurring naturally in foods (2nd ed.), pp. 43-87. WashingtonD. C.: National Academy of Sciences.

Agricultural research in Australia: a critical appraisal. Search 4, 155—160.Science development and the environment. Presidential Address, 1973 ANZAAS Congress.

Search 4, 413-417.1974 Human nutrition in our changing environment. Fd Nutr. Notes Rev. 31, 1-11.

The organization of agricultural research in Australia. Agric. Admin. 1, 73-81.Trace elements in human health and nutrition. Rec. Aust. Acad. Sci. 2, 48-65.

1975 Nutrition in our changing environment. The Ministry, February pp. 29-32.Cobalt. Nutr. Rev. 33, 65-68.Trace elements and their physiological roles in the animal. In: Trace elements in

soil-plant-animal systems (ed. J. D. Nicholas & A. R. Egan), pp. 227-241. Academic Press. The rich harvest—science in the development of Western Australia. Second Walter Murdoch

Lecture: Murdoch University, Western Australia.1976 Mineral imbalances in farm animals and their study and diagnosis with isotopic tracers. Atom.

Energy Rev. 14, 591-598.Methodology of trace element research. Trace elements in human health and disease, vol. II (ed.

A. S. Prasad), pp. 269-279. New York: Academic Press.Molybdenum in animal nutrition. In: Molybdenum in the environment (ed. W. R. Chappell &

K. K. Petersen), pp. 9-31. New York: Marcel Dekker.1977 Interactions of trace elements. In: Toxicity of heavy metals in the environment, (ed. F. W.

Oehme), pt II, ch. 28. New York: Marcel Dekker.1978 Trace element imbalances of interest to the dietitian. J. Am. diet. Ass. 72, 177-179.1979 (With D. I. Paynter & R. J. Moir) Changes in activity of the Cu-Zn super-oxide dismutase

enzyme in tissues of the rat with changes in dietary copper. J. Nutr. 109, 1570-1576. Trace metals in human and animal health. Jl R. Soc. Arts December, pp. 45-55.Trace elements and health: an overview. Phil. Trans. R. Soc. Lond. 288, 5-14.Concluding remarks. Phil. Trans. R. Soc. Lond. 288, 215-216.Environmental sources of heavy metals and their toxicity to man and animals. Prog. Wat.

Technol. 2, 33-45.The effect of sulphur on 75Se absorption and retention in sheep. J. Nutr. 109, 1448-1455. The detection and correction of trace mineral deficiencies and toxicities. In: Man's un

conquerable mind. Proc. Florida Nutrition Conf. pp. 202-233. Florida: I FAS.1981 The incidence of trace element deficiency diseases. Phil. Trans. R. Soc. Lond. B294, 3-8.



eric john underwood, 7 september 1905 - 19 august...

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