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Sydney John Folley, 1906-1970

Alan S. Parkes

1972, 240-265, published 1 November181972 Biogr. Mems Fell. R. Soc.

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SYDNEY JOHN FOLLEY

1906-1970

Elected F.R.S. 1951

Sydney J ohn F olley was born on 14 January 1906 into a working-class family in Swindon, Wilts. He died on 29 June 1970. His father, Thomas John Folley of 75 Graham Street, Swindon, who died in 1950 at the age of 83 years, was for more than 40 years an engine litter in the Great Western Railway Running Shed at Swindon. His mother, Katie Folley (n£e Baggs), died in 1938 at the age of 67 years. John Folley, as he was always known, had an elder sister, but was an only son. He was twice married, first, in 1935, to Madeline Kerr of Altrincham, Cheshire, and, then, when this marriage was dissolved in 1947, to Mary Lee Muntz (n£e Harnett) of Reading. There were no children of either marriage.

Folley was proud of the rail way men in his family and he was fascinated by trains. His maternal grandfather and his paternal great-grandfather both worked in one capacity or another for the Great Western and his paternal grandfather was a well known passenger driver on that railway. He recalls that his childhood circumstances were those usual in households of the skilled artisan class, but that his parents had a great respect for and were fully aware of the value of education. They were anxious to make it possible for both their children to take full advantage of every educational opportunity and were prepared to make considerable sacrifices to this end. ‘Perhaps the number of books in my home, when I was a child, was unusual for a working class home at that time. Probably, many of them were school books of my sister who was 8 years my senior and had received a secondary education after winning a scholarship. From an early age I had a passion for reading and took full advantage of these books’.

Scholastic achievements

His parents must have felt well rewarded for their sacrifices—Folley’s scholastic career was brilliant: From 1912 to 1918 he attended Sanford Street Boys’ Elementary School, Swindon, and gained a secondary school scholarship. Then for 6 years he went to the Swindon and North Wilts. Secondary School and Technical Institution. In 1922 he passed the London University General School Examination, obtaining exemption from the matriculation examination and gaining an Advanced Course Scholarship.

241



242Thereafter, he specialized in science (chemistry, physics and mathematics) and passed the Bristol University Higher School Certificate in 1924 with distinction in chemistry; he was awarded a Wilts. County Scholarship on the results of this examination. At this time he also sat for an open scholarship examination at Manchester University and though not gaining a University scholarship he was awarded an Exhibition at Hulme Hall, Manchester University.

Folley had a great respect for his school and especially for its Headmaster. He recalls:

‘In my time, the Swindon and N. Wilts. Secondary School was an excellent school in the best sense and was particularly strong on the science side, especially in chemistry. This may well have been due to the Headmaster, M r G. H. Burkhardt, M.A., M.Sc., who was an outstanding educational personality (he was given an Honorary M.A. by Bristol University for his services to education) and had carried out research in the laboratory of W alther Nernst.Pupils taking the two-year advanced course in science, in preparation for proceeding to a university, were given a thorough grounding in classical physics and chemistry and received special lectures from the head himself in scientific method. The quality of the practical instruction in organic chemistry may be gauged from the fact that at school I carried out reactions and made preparations which I was later required to repeat in the third year of my honours course in chemistry at Manchester University (one of the most famous schools of chemistry in the world).’

Folley had not taken Latin at school and was therefore unable to try for an Oxford or Cambridge Scholarship and since he wished to read chemistry, his headmaster advised him to enter the Honours School of Chemistry at Manchester. Folley records that he never regretted this ‘since in my day the intellectual level of the teaching staff of a number of Faculties in the University of Manchester was probably equal to that of any University in the world’. Besides taking the prescribed courses leading to an Honours Degree in Chemistry, which involved attending lectures by A. Lap worth and R. Robinson among others, he attended lectures on physics by W. L. Bragg, botany by F. E. Weiss and biochemistry by H. S. Raper. In 1927, he obtained a First Class Honours Degree in Chemistry, coming out at the head of the list, and accordingly was awarded the Mercer Scholarship in Chemistry as well as a Department of Industrial and Scientific Research Maintenance Grant, to enable him to do research for two years.

Early research

Before graduating, he had attended a special course in colloid chemistry given by D. C. Henry, then in charge of the Graham Colloid Laboratories at Manchester, and was so interested that after graduation in 1927 he began

Biographical Memoirs



research in colloids under Henry’s direction. At the end of the year, having obtained the degree of M.Sc. for a thesis on the properties of organosols o f nickel, he decided to change to biochemistry, and joined the biochemical laboratory of H. S. Raper, Brackenbury Professor of Physiology at M anchester. Here he worked for three years on plastein formation, for the latter two as R aper’s personal research assistant. In 1931, he was awarded the degree of Ph.D. (Manchester) for a thesis on plastein and was then appointed Assistant Lecturer in Biochemistry in the University of Liverpool. There he continued the plastein work for a few months until forced to resign his post owing to a slight attack of tuberculosis.

Sydney John Folley 243

National Institute for Research in Dairying

On recovery in 1932, on R aper’s recommendation, he was appointed Research Assistant in the Physiology Department at the National Institute for Research in Dairying at Shinfield, Reading. I had visited the Institute several times in the middle 1920s, originally with Jack Drummond, but my first indication of Folley’s existence was early in 1934 (see p. 245). Two years later a letter from S. K. Kon, Head of the Nutrition Laboratory, dated 4 February 1936, asking whether he and his colleague Folley could come to see me about pregnancy tests for cows showed that something new was stirring at Shinfield. On the day following the arrival of R on’s letter I received one from Folley himself and the three of us met on 11 February 1936. This started a close connexion which continued for many years, though we collaborated in the writing of only one paper. During the rest of 1936 and in 1937 there was a continuous correspondence between us and frequent meetings, mainly about the effects of oestrogens, androgens and prolactin on lactation and also in connexion with attempts to raise antiprolactin sera in a pig, for which Folley had arranged facilities.

During 1936, I invited him to contribute the chapter on lactation to the third edition of F. H. A. Marshall’s Physiology of reproduction which I was to edit. Later he attended a meeting of prospective contributors and records:

‘This led to my meeting and coming under the influence of F. H. A. Marshall, J . Hammond and S. Zuckerman, with the latter of whom I collaborated in a study of the monkey mammary gland. Contact with all of these eminent workers in the field of reproductive physiology had a profound influence on my scientific development. About this time I also met F. G. Young and began a series of joint studies with him, lasting many years, on the galactopoietic effect of anterior pituitary extracts.’

By this time, therefore, he was well entrenched in the lactation field and three main lines emerged, (a) metabolic aspects of milk formation, (b) endocrinological aspects of mammary growth and milk secretion, (c) neurophysiology of milk secretion.



244Much of the later work was greatly facilitated by Folley’s far-sighted

development of the goat as an experimental animal. The early work of the physiology department was necessarily confined to small laboratory animals, mainly rats and guinea-pigs, and to such cows as could be made available by other departments. About 1937 Folley realized the value of goats as prototype ruminants for lactational research. At first, a few goats were purchased but he soon realized the importance of using pedigreed animals with high milk yield potential and during 1938 and 1939, with advice from the British Goat Society, he started to build up a herd of British Saanen goats. The herd grew gradually during the war and the available accommodation became too small. Soon after the war plans were drawn up for a new goat house which was completed in 1952. This was one of the most modern in the country and, at times, housed about 140 goats, the nucleus being a breeding herd of some 30 animals. In this way, a magnificent herd of pedigreed animals was developed, of which Folley was justly very proud and which played a major part in the department’s work on the hormonal control of mammary growth, on the hormonal requirements for lactation and in neuro-endocrine studies.

Metabolic aspects of milk formation

The work on the metabolic aspects of lactation started immediately after his arrival at Shinfield, where he worked with the sole graduate member of the staff, G. L. Peskett, another biochemist, on the relation between blood electrolytes and the lipid constituents of milk. Then, with H. D. Kay, newly arrived from Toronto as Director, he worked on the alkaline phosphatase of the mammary gland. His attention then switched, as already indicated, to endocrinological studies and the outbreak of war in 1939 kept applied research of this kind in the forefront for many years. In the late 1940s, however, in collaboration with T. H. French, Folley studied the respiratory metabolism of slices of mammary gland from ruminants and non-ruminants. The results indicated a curious difference between ruminants and nonruminants in that, in the former, but not the latter, it seemed that acetate must be an important substrate for the synthesis of milk fatty acids, particularly those of shorter chain-length. This work, recorded at a meeting of the Biochemical Society, attracted the attention of G. J . Popjak, who suggested collaboration on lipogenesis in the mammary gland, using r e labelled acetate, at that time a scarce reagent for which official sanction was necessary. Happily, this was forthcoming and a goat was selected as the experimental animal. I now quote Folley’s own account of the experiment and its results, as recorded in the last paper he wrote, a personal perspective in the 1970 Biennial Review from the National Institute for Research in Dairying:

‘The first major experiment by the use of 14C in Britain was carried out on this goat. After being injected with 5 mCi of [1-14C] acetate its head was placed into an improvised respiration chamber, constructed by M r




245

S. C. Watson from the astrodome of a bomber aircraft; the air supply to the chamber was provided by a milking machine and the expired 14COa was collected over 6 hours. The animal did not object to its head being in the chamber. . . .This splendid animal provided evidence (a) that the short-chain fatty acids in milk triglycerides were synthesized in the mammary gland from “acetate” and were not derived by the breakdown of longer-chain acids transported from the blood; (b) that in the synthesis of fatty acids chain-elongation occurred by the addition of “acetate” units to the carboxyl end of a shorter acid; (c) that the galactose moiety of lactose arose by an inversion of glucose; (d) that hexose was the direct source of glycerol in triglyceride synthesis; and (e) that glycerol was treated in enzymic reactions in an asymmetric manner.’

Sydney John Folley

Endocrinology of lactation

Folley records, very generously, that his interest in endocrinology was aroused by reading my book on the Internal secretions of the , which directed his attention to the endocrine control of lactation as a promising field of research. Our first recorded correspondence, early in 1934, related to this book. In Folley’s own words (1970) ‘Parkes and his colleagues were among the first to show, by indirect means, that oestrogens inhibited lactation in laboratory rodents and I thought it would be interesting to find out if the same was true of cattle, in which the milk yield could be measured directly, and see what changes in milk composition accompanied the inhibition.’ This was no doubt the background to his letter to me of 5 February 1936, which ran:

‘I would like to study the effect of oestrone on lactation in the cow. I am writing to ask you whether you are able to give me some idea of the twelve hourly dosage necessary for the production of physiological effects in an animal weighing about 600 kilos.’

When we met on 11 February, as already recorded, I suggested multiplying the mouse dose, which was known, by the weight factor of 24 000. Folley must then have become very active, because early in July he wrote to me the letter reproduced in facsimile in figure 1.

The De Fremery referred to was on the staff of N. V. Organon, Oss, Holland, but worked in close touch with E. Laqueur. Folley subsequently described the results of his first experiment with oestrogens as follows: ‘I managed to acquire sufficient of the then extremely expensive and scarce purified natural oestrogenic hormones to experiment on three cows and found that the inhibition was only temporary but was accompanied by very prolonged increases in the concentration of milk solids, a phenomenon which I called the “enrichment” effect.’ The full account was published later in 1936 in the Biochemical Journal, to be followed soon afterwards by a



246 Biographical Memoirs University of Reading.

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second one of the effects of thyroxine on lactation, published in the Proc. R. Soc. Lond.

The preparation by Dodds and his colleagues of di-ethylstilboestrol and related artificial oestrogens, soon to be available in large quantities, enabled observations on oestrogenized cows to be carried out on a much larger scale, and during the early years of the war strenuous efforts were made to improve milk yield and to induce lactation in dry cows and barren heifers by feeding or implanting massive doses of hexoestrol. The work on dry cows and heifers was based on the observation that oestrogen would induce lactation in virgin goats, which in turn derived from the well-known fact that virgin guinea-pigs could be made to lactate by treatment with oestrogen. The results on cows were complicated by the fact that massive dosage with oestrogen caused slackening of the pubic ligaments, causing ‘dropped hip’, a dangerous condition likely to lead to fracture. This work was under the nominal direction of an A.R.C. Committee on Endocrinology, chaired by Joseph Barcroft, of which Folley was a prominent member. I well remember one spirited meeting, when discussing dosage, Folley said we should probably get pelvic complications anyway, so we decided to give really massive dosage to ensure lactational effects. And massive they were, 5 X 0.5 g tablets of compressed dry hexoestrol. However, partly because the milk yield never went above 2 gal a day, a relatively insignificant amount, the work on dry cows and barren heifers did not get beyond the pilot experiment stage. The use of compressed tablets for this work turned Folley’s attention to this method of administering steroid and similar substances, and he and his colleagues made the curious observation that in time, the interstices of the compressed tablets became infiltrated with protein material so that, on dissolving away the residual substance, a ‘ghost’ of the tablet remained.

Another war-time project arose from Folley’s work on the effects of thyroxine in increasing milk production. His results seemed to be sufficiently promising to test on a field scale, and the inadequacy of thyroxine supplies led to a considerable project for preparing artificial products with thyroidlike activity by the iodination of proteins, especially casein. Large amounts of such preparations, with thyroidal activity up to twice that of dried thyroid tissue, were made under the supervision of a committee chaired by C. R. Harington and including Folley. These preparations were tested on large groups of lactating cows with varying degrees of the expected effect, but, as with the oestrogen work, no worthwhile results in terms of overall milk supply were obtained.

Folley’s interest in the hypophysial lactogenic hormone prolactin arose from his meeting with F. G. Young, who was working on the effects of hypophysial hormones on carbohydrate metabolism and was fractionating extracts of dried pituitary tissue from several species. This meeting resulted in close collaboration for a number of years, the first fruit of which was the idea that lactogenesis involved a complex of anterior hypophysial hormones, of which prolactin and adrenocorticotrophin were the most important. This




248idea became widely accepted, but required some qualification in the light of later work at Shinfield. Folley and Young also suggested that a galactopoietic complex of anterior pituitary hormones was necessary for the maintenance of lactation, and that hypophysial growth hormone was an essential factor. The involvement of growth hormone was afterwards confirmed by the discovery of the galactopoietic action of purified somatotrophin. Later work at Shinfield, based on hypophysectomized and stalk- sectioned goats gave further information about requirements for the maintenance of lactation and added the interesting fact that in the goat, as in the rat, the anterior pituitary gland deprived of its vascular connexions with the hypothalamus, continued to secrete prolactin, but not the other trophic hormones.

Studies on hypophysectomized rats and rabbits revealed species differences in the hormonal requirements for maintaining lactation; rats needed prolactin+AGTH whereas prolactin alone was sufficient in the rabbit. The story regarding other species was further complicated by the fact that prolactin could not be isolated from primate pituitaries. Human growth hormone, however, was found by Folley and his colleagues, working on pseudopregnant rabbits, to be as lactogenic as sheep prolactin—ruminant growth hormone showing no such activity. The belief grew that primate prolactin probably did not exist and that primate growth hormone with its inherent lactogenic activity played a dual role. However, in Folley’s laboratory discrepancies came to be observed between the results of bioassays for lactogenic activity in the plasma of lactating women and the results of radioimmunoassays for human growth hormone in the same plasmas, indicating the presence of a lactogenic hormone other than G H ; similar observations were reported about the same time from the U.S.A. Two years later, at a Giba Foundation symposium, the successful isolation of human and monkey prolactin was reported by American workers.

Neuroendocrine effects in lactation

Much of the work at Shinfield in the 1960s dealt with neuroendocrine mechanisms, which are concerned both with the stimulation of prolactin secretion and the milk-ejection reflex involving oxytocin. The work on oxytocin at Shinfield was made possible by the development of a method for the determination of relatively small amounts of the substance and its secretion was found to be stimulated by suckling or other stimulation of the udder and, in conditioned animals, by auditory and visual signals associated with milking time.

Extensive studies were carried out on the ascending pathways, within the brain, of the milk-ejection reflex and of the reflex responsible for prolactin release. As far as the mid-brain these reflexes appeared to share a common path, thereafter there was some divergence. The pathway for prolactin release was found to have connexions with the medial pre-optic area and the




249

prefrontal cortex. This connexion with the cerebral cortex opened new fields of study on the routes by which environmental stimuli may influence lactation.

Sydney John Folley

Personal appreciation

In his undergraduate days Folley was something of an athlete, representing Manchester University at cross-country running and being secretary of the Swimming Club. I t is a sad irony, therefore, that for much of his life he had to struggle with physical disabilities. In 1931, soon after taking it up, he had to resign his appointment in Liverpool because of an attack of tuberculosis. On recovery in 1932 he went to Shinfield and it is odd to think that had he not become ill, he might well have stayed in Liverpool working on colloids, so that Shinfield and with it lactational physiology in this country would have been the poorer. In 1938, a second attack forced him to come back prematurely from the U.S.A., where he had planned to spend a year with Edgar Allen at Yale and collaborate with W. U. Gardner and G. van Wagenen. This visit, which started when he was greatly worried about the Munich episode, got off to a bad start and ended in near disaster. Then, in 1946, he suffered a severe deterioration in vision, which was very worrying, but which at that time became no more than a severe handicap to his scientific work. In 1958 when he returned from a third visit to the U.S.A. he underwent an eye operation which it was hoped would improve his sight. Unhappily, the operation was a calamitous failure and to his bitter disappointment by April 1959, at the age of 53, he became almost totally blind and had to rely entirely on the services of readers and the use of tape recording equipment. This stunning blow would have incapacitated many people permanently, but his dogged resolution and fortitude, and, above all, the devoted assistance of his wife Mary, who earned the admiration and affection of all who knew them and for whom no tribute can be too great, enabled him to carry on to a surprising extent. Up to the time of his death he attended meetings, read papers and sat effectively on committees, aided by a memory now highly trained for the job. I remember at a Scientific Group on Lactation convened by W.H.O. in Geneva in 1964, Folley listened attentively to a long memorandum read by one of the other participants and then commented on various parts of it, quoting accurately the various passages with which he disagreed. His Dale Lecture to the Society for Endocrinology was delivered entirely from memory of a carefully prepared script, a very different thing from extempore speaking.

Folley had wide interests outside science. Politically, he was towards the left, and in his early days took part in left-wing activities. But outside the laboratory his main interest was in the arts. In his own words:

‘For many years I have been interested in the arts, particularly music, painting, ballet, cinema, architecture, etc. and I have formed collections of drawings and paintings by contemporary French and English



250painters and of gramophone records for the reproduction of which I have assembled a fine electrical reproducer.’

This love of music was a great consolation to him after his sight failed, and his collection of gramophone records was famous. Personally, I never experienced one of his musical evenings, but according to the appreciation published in the Journal o f Endocrinology ‘He was particularly fond of eighteenth-century orchestral and chamber music, in particular the works of Handel, J . S. Bach and Mozart, and he loved to listen to these with the volume control set at a level appropriate for a fair-sized auditorium. He would listen, beaming with obvious delight, while his visitors sat amid the tumult and speculated uneasily on the resilience of their own auditory apparatus.’ He had, however, little patience with the human voice in song, and opera had no appeal.

There can be no doubting Folley’s intellectual capacity and the value of his contributions to science. When he joined the Institute in 1932, the newly established Department of Physiology at Shinfield consisted of a biochemist and a technician, housed in a converted back-bedroom of the old manor house. When he died, after having been Head for 25 years, the Department had extensive accommodation and animal quarters—as many as 50 people worked in it at one time—and the Department occupied a prominent place in world research on the physiology of lactation; it was a Mecca for overseas workers. Some of this development was no doubt due to the general upsurge of scientific research after the war, but mainly it must be attributed to Folley’s insistent pressure for his subject outside the Department and his galvanic influence within it.

As a scientist he was fertile-minded, clear-headed, resourceful and critical, a powerful combination. Although his background was essentially biochemical at the time he went to Shinfield, he very soon acquired a profound knowledge of the mammary gland. Early in the war, when we were working on the induction of lactation in dry cows and barren heifers, by administration of oestrogen, I suggested (having in mind the case of the male guinea-pig) that, while we were on the job, we should bring a few steers into milk. Folley immediately explained that the rudimentary mammary tissue of the billy-goat showed little response to oestrogen and it was not likely that steers would react differently. Shortly afterwards, as Cowie recalls, a local farmer made the same suggestion about steers and Folley put him off by replying that he did not have any steers available. Next day a steer arrived in a truck with the farmer’s compliments and the message that the steer would be collected again when it was in milk. Malpress and Cowie were instructed to do something with it. Oestrogen was implanted and its teats grew, but there was very little glandular growth and the maximum yield obtained was about 20 ml of a milky fluid!

Folley was undoubtedly a first-rate scientist. I t is less easy to assess the human qualities of anyone so full of contrasts and apparent contradictions.




251In the last thing he wrote before his death, he acknowledged most generously, and not for the first time, his debt to those who had helped and encouraged him in his early days. On his side, he was always ready to provide material or facilities for those who, like myself, had no direct access to farm animals or for other reasons needed his help. My first abortive experiments on freezing bull semen were carried out in his laboratory and the later highly successful ones at the N .I.M .R. were made possible by a supply of bull semen from Shinfield organized by him.

His outstanding quality was his utter devotion to his department and his colleagues in it, a devotion which was whole-heartedly reciprocated. In his own words, published a few days before he died ‘I have had the help, friendship and loyal collaboration of numerous colleagues with whom it has been a great pleasure and privilege to work’. Among those should be mentioned especially F. H. Malpress, A. T. Cowie and S. G. Watson. This intense dedication to his department naturally led to other consequences. For instance, he tended to be resentful of anything which he regarded as obstruction or interference from within the Institute, or of encroachment or lack of understanding from outside. Moreover he was intensely critical and intolerant of anything he considered to be shoddy work or a bogus personality. Yet he was in no sense insular or introverted. He travelled widely and, on his side, welcomed overseas visitors at Shinfield. He had close ties with the Laboratoire de Physiologie de la Lactation at Jouy-en-Josas in France and organized with it a series of symposia held at two-year intervals at Shinfield or at Jouy.

Even before his sight failed, he had little capacity to see the funny side of things in his work or to handle situations with a light touch—possibly a legacy from his efforts, which must have been intense at school and university, to make a way for himself. Contretemps of which most people would take little notice aroused his serious ire, and badinage (as when I said to him some years ago ‘Happily, I am no longer an endocrinologist!’) could be dangerous. Yet he was never vindictive and he loathed injustice, cruelty and violence, and Mary Folley notes that above all, he delighted in the little private jokes which are the leaven of domesticity. Unfortunately, too few people saw this side of him and for the most part his fellow scientists outside Shinfield, regarded him with respect and admiration, but perhaps with less overt affection than should have been the case. Yet under the surface it was there, as witness the symposium on Lactogenic Hormones planned by Ciba Foundation to honour him on his retirement but perforce dedicated to his memory. Even this heavy cloud over the meeting had a silver lining, for no more fitting a memorial than one epitomizing his life’s work could have been found for Sydney John Folley.

I am much indebted to Dr A. T. Cowie for assistance in the preparation of this memoir. The photograph is by Peter Tring.

Sydney John Folley

A lan S. Parkes.



252 Biographical Memoirs

1927.

1929- 1931.1930- 1931.1931- 1932.

1932- 1945.

1938-1939.

1943-1945.

1945-1970.1946.1951.1953.1953.1964.1964.

1931-1937-1941-1946-

1960-1963-

1945-1947.1940-1942.1948.1952-1957.1954.1952-1960-1963.

1963.

Professional Record

B.Sc., Manchester. Awarded the Mercer Scholarship in Chemistry for two years.

Research Assistant to Professor Raper in Manchester.Resident Tutor, Hulme Hall, Manchester.Assistant Lecturer in Biochemistry and Resident Tutor, Rankin Hall,

University of Liverpool.Research Assistant, Physiology Department, National Institute for Research

in Dairying, University of Reading.Rockefeller Research Fellow, Department of Anatomy, Yale University,

U.S.A.In charge of the teaching of Biochemistry at the Royal Veterinary College,

which was evacuated to the University of Reading during the war.Head of Physiology Department, National Institute for Research in Dairying. Cantor Lecturer, Royal Society of Arts, London.Elected Fellow of the Royal Society.Promoted D.C.S.O.Professeur a titre Etranger, College de France.Research Professor of the University of Reading.Honorary Doctorate, University of Ghent.

Membership o f British SocietiesBiochemical Society (Committee 1952-1957).Physiological Society.Royal Society of Medicine (Council of Section of Endocrinology, 1948). Society for Endocrinology (Secretary 1946-1951, Chairman 1951-1956, Dale

Medallist 1969).Society for the Study of Fertility.Blair Bell Research Society.

Membership o f Advisory Bodies, etc.Committee on Human Fertility, Medical Research Council.Committee on Endocrinology, Agricultural Research Council.Editorial Board of Journal of Endocrinology (Chairman 1960- ).Chairman, Technical Committee on Endocrinology, A.R.C.Committee on Anterior Pituitary Hormone Standards, M.R.C.Advisory Panel of Ciba Foundation.Member of the International Jury for the award of the Prix Francqui (President, 1963).

Scientific Group on Lactation, World Health Organization.

Visits abroadFolley was well known in Western Europe which he visited many times, and he made

many visits to N. America, but he never got as far as S. America, Africa or the East. In Europe, between 1937 and 1953 he visited laboratories in Stockholm, Uppsala, Helsinki, Basel, Paris, Brussels, Ghent, Leiden, Amsterdam, Oss, Groningen, Copenhagen and Strasbourg. He was a guest speaker at several International Congresses and Symposia.



253abroad, including the Laurentian Hormone Conference in 1951, and before several foreign Societies, including the Dutch-Belgian Endocrinological Societies in 1949 and 1953. In 1961 he visited the U.S.S.R. under the official scheme for the exchange of agricultural research workers, visiting universities and research institutes, and delivering a lecture in the Ministry of Agriculture in Moscow.

PublicationsThe three lectures given in Paris, in French, were afterwards published in book form

under the title Recherches rScentes sur la physiologie et la biochemie de la sScretion lactSe.He contributed chapters to Marshall’s Physiology of reproduction, 3rd Edition, Volume 2,

1952 entitled ‘Lactation’ and to various other collaborative works listed below.

Sydney John Folley

BIBLIOGRAPHY

A

Scientific papers

1928- (With D. C. H enry.) The preparation of colloidal solutions of nickel in dry1929. acetone and some preliminary observations on their stability. Mem. Manchr

lit. phil. Soc. 73, 99-110.1930. An improved design of the Van Slyke apparatus for the estimation of amino-

nitrogen. Biochem. J . 24, 961-964.1932. The nature of plastein. Biochem. J . 26, 99-105.1933. Note on the preparation and fractionation of the a-naphthyl-wo-cyanate

compound of plastein. Biochem. J . 27, 151-152.1933. (With G. L. Peskett.) Some observations on cow’s milk poor in non-fatty

solids. J . Dairy Res. 4, 279-284.1933. Standardized collodion membrane in low pressure ultrafiltration. Biochem. J .

27, 1775-1778.1934. (With G. L. Peskett.) Experiments on variations in blood composition. J .

Physiol. 82, 486-495.1934. (With G. L. Peskett.) The effect of salts on cell permeability as shown by

studies of milk secretion. Proc. R. Soc. Lond. B, 116, 396-402.1935. (With H. D. Kay.) The alkaline phosphomonoesterase of the mammary

gland. Biochem. J . 29, 1837-1850.1936. (With P. White.) The effect of thyroxine on milk secretion and on the phos

phatase of the blood and milk of the lactating cow. Proc. R. Soc. Lond. B, 120, 346-365.

1936. (With H. D. Kay.) Variations in the phosphomonesterase content of the milk of the cow in relation to the progress of lactation. Enzymologia, 1, 48-54.

1936. The effect of oestrogenic hormones on lactation and on the phosphatase of the blood and milk of the lactating cow. Biochem. J . 30, 2262-2272.

1938. (With S. K. K on.) The effect of sex hormones on lactation in the rat. Proc. R. Soc. Bond. B, 124, 476-492.

1938. Experiments on the relation between the thyroid gland and lactation in the rat. J . Physiol. 93, 401-412.



1938. (With F. G. Young.) The effect of anterior pituitary extracts on established lactation in the cow. Proc. R. S Land. B, 126, 45-76.

1938. (With A. G. Bottomley.) The effect of androgenic substances on the growth of the teat and mammary gland in the immature male guinea-pig. Proc. R. Soc. Lond. B, 126, 224-240.

1938. (With A. G. Bottomley.) The effect of high doses of androgenic substances on the weights of the testes, accessory reproductive organs and endocrine glands of young male guinea-pigs. J . Physiol. 94, 26-39.

1938. (With E. W. Ikin, S. K. K on and H. M. Scott Watson.) Observations onspecific nutritional factors in lactation. Biochem. 32, 1988-1999.

1939. (With A. N. Guthkelch and S. Zuckerman.) The mammary gland of theRhesus monkey under normal and experimental conditions. Proc. R. Soc. Lond. B, 126, 469-491.

1939. (With F. G. Young.) The effect of continued treatment with anterior pituitary extracts on milk volume and milk-fat production in the lactating cow. j Biochem.J. 33, 192-197.

1939. (With P. Bacsich.) The effect of oestradiol monobenzoate on the gonads, endocrine glands and mammae of lactating rats. J . Anat., Lond. 73, 432-440.

1939. Interactions of estrone and prolactin with special reference to the effect of estrone on the pigeon crop-gland response. , 24, 814-822.

1939. (With J . Van Heuverswyn and W. U. Gardner.) Mammary growth in male mice receiving androgens, estrogens and desoxycorticosterone acetate. Proc. Soc. exp. Biol. Med. 41, 389-392.

1939. (With A. C. Bottomley, F. H. A. Walker and H. M. Scott Watson.)Effect of subcutaneous implantation of adrenalin tablets on blood-sugar and milk composition in lactating ruminants. J . Endocr. 1, 287-299.

1939. (With G. Van Wagenen.) The effect of androgens on the mammary glandof the female Rhesus monkey. J . Endocr. 1, 367-372.

1940. (With A. C. Bottomley and H. M. Scott Watson.) Experiments on the useof chorionic gonadotrophin (pregnancy urine extract) for the treatment of sterility in dairy cattle. J . agric Sci., Camb. 30, 235-243.

1940. (With F. J . Dyer and K. H. Coward.) The assay of prolactin by means of the pigeon crop-gland response. J . Endocr. 3, 179-193.

1940. (With F. G. Young.) Further experiments on the continued treatment oflactating cows with anterior pituitary extracts. J . Endocr. 2, 226-236.

1941. (With H. M. Scott Watson and A. G. Bottomley.) Experiments on thechemical enrichment of cows’ milk by the administration of diethylstilboestrol and its dipropionate. J . Dairy Res. 12, 1-17.

1941. (With W. M. Allcroft.) Observations on the serum phosphatase of cattle and sheep. Biochem. J . 35, 254-266.

1941. (With G. W. Scott Blair, F. H. Malpress and F. M. V. Coppen.) Variations in certain properties of bovine cervical mucus during the oestrous cycle. Biochem. J . 35, 1039-1049.

1941. (With H. M. Scott Watson and A. C. Bottomley.) Studies on experimentalteat and mammary development and lactation in the goat. J . Dairy Res. 12, 241-264.

1942. (With H. M. Scott Watson and E. G. Amoroso.) Further experiments onlactation in thyroidectomized rats: the role of parathyroids. J . Endocr. 3, 178-191.




1944.

1944.

1944.

1944.

1944.

1944.

1944.

1945.

1945.

1945.

1945.

1946.

1946.

1946.

1947.

1947.

1947.

1947.

Studies on the absorption of subcutaneously implanted tablets of hexoestrol. Proc. R. Soc. Lond. B, 132, 142-163.

(With F. H. Malpress.) The response of the bovine ovary to pregnant mares* serum and horse pituitary extract. Proc. R. Soc. Lond. B, 132, 164-188.

(With F. H. Malpress.) The artificial induction of lactation in the bovine by the subcutaneous implantation of synthetic oestrogen tablets. J . Endocr. 4, 1-18.

(With F. H. Malpress.) Artificial induction of lactation in bovines by oral administration of synthetic oestrogens. J . Endocr. 4, 23-36.

(With F. H. Malpress.) The chemical composition of bovine mammary secretions induced by the subcutaneous implantation or oral administration of synthetic oestrogens. J . Endocr. 4, 37-42.

(With D. L. Stewart and F. G. Young.) Experiments on the use of tablets containing 50% hexoestrol for the artificial induction of lactation in the bovine. J . Endocr. 4, 43-52.

(With A. T. Cowie.) Adrenalectomy and replacement therapy in lactating rats. Tale J . Biol. Med. 17, 67-74.

(With F. H. Malpress and F. G. Young.) Induction of lactation in goats and cows with synthetic oestrogens and anterior-pituitary extracts. Endocr. 4, 181-193.

(With F. G. Young.) The galactopoietic action of pituitary extracts in lactating cows. 1. Dose-response relations and total yields during declining lactation. J . Endocr. 4, 194-204.

(With H. T. Fawns and F. G. Young.) The galactopoietic action of pituitary extracts in lactating cows. 2. The response during the peak of lactation. J . Endocr. 4, 205-211.

(With F. H. Malpress and F. G. Young.) The galactopoietic action of pituitary extracts in lactating cows. 3. Comparison of extracts of pituitary glands from different species. J . Endocr. 4, 212-218.

(With A. L. Greenbaum.) Effects of adrenalectomy and of treatment with adrenal cortex hormones on the arginase and phosphatase levels of lactating rats. Biochem. J . 40, 46-51.

(With A. T. Cowie.) Some factors affecting the absorption rate of subcutaneously implanted hormone tablets. JEndocr. 4, 375-385.

(With R. Deanesly and A. S. Parkes.) Further observations on the formation of ‘ghosts’ in subcutaneously implanted tablets. J . Endocr. 4, 422-425.

(With A. T. Cowie.) The measurement of lactational performance in the rat in studies of the endocrine control of lactation. J . Endocr. 5, 9-13.

(With A. T. Cowie.) Adrenalectomy and replacement therapy in lactating rats. 2. Effects of deoxycorticosterone acetate on lactation in adrenalectomized rats. J.Endocr. 5, 14-23.

(With A. T. Cowie.) Adrenalectomy and replacement therapy in lactating rats. 3. Effects of deoxycorticosterone acetate and 11-oxygenated cortical steroids on lactation in adrenalectomized rats maintained on stock or high-protein diets. J . Endocr. 5, 24-31.

(With A. T. Cowie.) The role of the adrenal cortex in mammary development and its relation to the mammogenic action of the anterior pituitary. Endocrinology, 40, 274-285.




256

1947. (With A. L. Greenbaum.) Changes in the arginase and alkaline phosphatase contents of the mammary gland and liver of the rat during pregnancy, lactation and mammary involution. Biochem. J . 41, 261-269.

1947. (With K. M. Henry and S. K. K on.) Reproduction and lactation in the raton highly purified diets. Br. J . Nutr. 1, 39-53.

1948. (With A. L. Greenbaum.) Adrenalectomy and replacement therapy inlactating rats. 4. Effect of deoxycorticosterone acetate on the water content of mammary tissue after adrenalectomy. J . Endocr. 5, 236-242.

1948. (With A. T. Cowie.) Adrenalectomy and replacement therapy in lactating rats. 5. The effect of adrenalectomy on lactation studied in pair-fed rats. J . Endocr. 5, 282-289.

1948. (With A. L. Greenbaum.) Determination of the arginase activities of homogenates of liver and mammary gland: effects of pH and substrate concentration and especially of activation by divalent metal ions. Biochem. J . 43, 537-549.

1948. (With A. L. Greenbaum.) Effect of adrenalectomy on the arginase levels ofliver, mammary gland and kidney in lactating rats studied by the paired feeding technique. Biochem. J . 43, 581-584.

1949. (With T. H. French.) The intermediary metabolism of the mammary gland.1. Respiration of lactating mammary gland slices in presence of carbohydrates. Biochem. J . 45, 117-125.

1949. (With T. H. French.) The intermediary metabolism of the mammary gland.2. Respiration and acid production of mammary tissue during pregnancy, lactation and involution in the rat. Biochem. J . 45, 270-275.

1949. (With A. L. Greenbaum and A. R oy.) The response of the ovary of the an-oestrous goat to pregnant mares’ serum gonadotrophin.^. Endocr. 6, 121-131.

1950. (With T. H. French.) The intermediary metabolism of the mammary gland.3. Acetate metabolism of lactating mammary gland slices with special reference to milk fat synthesis. Biochem. J . 46, 465-472.

1951. (With G. PopjAk and T. H. French.) Utilization of acetate for milk-fatsynthesis in the lactating goat. Biochem. J . 48, 411-416.

1951. (With P. M. F. Bishop.) Absorption of hormone implants in man. Lancet, ii, 229-232.

1951. (With J udith H. Balmain.) Further observation on the in vitro stimulation by insulin of fat synthesis by lactating mammary gland slices. Biochem. J . 49, 663-670.

1951. (With A. T. Cowie, W. G. Duncombe, T. H. French, R. F. Glascock, L.Massart, G. J . Peeters and G. PopjAk.) Synthesis of milk fat from acetic acid (CH314COOH) by the perfused isolated bovine udder. Biochem. J . 49, 610-615.

1951. (With S. C. Watson.) Comparative effects of cortisone and 11-deoxycorticosterone on tissue arginase levels of adrenalectomized lactating rats. Proc. Soc. exp. Biol. Med. 78, 473-476.

1952. (With A. T. Cowie, F. H. Malpress and K. C. R ichardson.) Studies onthe hormonal induction of mammary growth and lactation in the goat. J . Endocr. 8, 64-88.

1952. (With Judith H. Balmain and R. F. Glascock.) Inhibition of fatty acid synthesis in rat mammary gland slices by cortisone in vitro and its antagonism by insulin. Nature, Lond. 169, 447-449.




1952. (With J udith H. Balmain.) In vitro effects of prolactin and cortisone on the metabolism of rat mammary tissue. Archs. Biochem. Biophys. 39, 188-194.

1952. (With J udith H. Balmain and R. F. Glascock.) Effects of insulin and of glycerol in vitro on the incorporation of C] acetate into the fattyacids of lactating mammary gland slices with special reference to species differences. Biochem. J . 52, 301-306.

1952. (With G. PopjAk and R. F. Glascock.) Incorporation of C] acetateinto lactose and glycerol by the lactating goat udder. Biochem. J . 52, 472-475.

1954. (With J udith H. Balmain and R. F. Glascock.) Relative utilization of glucose and acetate carbon for lipogenesis by mammary gland slices, studied with tritium 18C and 14C. Biochem. J . 56, 234-239.

1954. (With S. Bartlett, A. W. A. Burt and S. J . R owland.) Relative galactopoietic effects of 3 :5 rS'-triiodo-L-thyronine and L-thyroxine in lactating cows . J. Endocr. 10, 193-201.

1954. (With D. S. Flux and S. J . R owland.) The effect of adrenocorticotrophic hormone on the yield and composition of the milk of the cow. J . Endocr. 10, 333-339.

1954. (With K. C. R ichardson.) Synthetic activity of a single epithelial cell of the lactating goat udder. Nature, Land. 174, 828.

1954. (With J . H. Balmain, C. P. Cox and M. L. M cNaught.) The bioassay ofinsulin in vitro by manometric measurements on slices of mammary glands.

J. Endocr. 11, 269-276.1955. (With M. L. M cNaught, R. F. Glascock and J. H. Balmain.) Effects of

adrenal corticoids on fatty acid synthesis in mammary gland slices in vitro. Biochem. J . 60, 102-108.

1955. (With G. K. Benson, A. T. Cowie, C. P. Cox and D. S. Flux.) Studies onthe hormonal induction of mammary growth and lactation in the goat. II. Functional and morphological studies of hormonally developed udders with special reference to the effect of ‘triggering’ doses of oestrogen. J . Endocr. 13, 46-58.

1956. (With G. K. Benson.) Oxytocin as stimulator for the release of prolactinfrom the anterior pituitary.) Nature, Lond. 177, 700.

1956. (With A. J . Marshall.) The origin of nest-cement in edible-nest swiftlets( Collocaliaspp.) Proc. zool. Soc. Lond. 126, 383-389.

1957. (With G. K. Benson.) The effect of oxytocin on mammary gland involutionin the rat. J . Endocr. 16, 189-201.

1958. (With E. C. Dodds, R. F. Glascock and W. Lawson.) The excretion ofmicrogram doses of hexoestrol by rabbits and rats. Biochem. J . 68, 161-167.

1960. (With G. K. Benson and J . S. Tindal.) Effects of synthetic oxytocin andvalyl oxytocin on mammary involution in the rat. J . Endocr. 20, 106-111.

1961. (With A. Chadwick and G. A. Gemzell.) Lactogenic activity of humanpituitary growth hormone. Lancet, ii, 241-243.

1965. (With I. A. Forsyth and A. Chadwick.) Lactogenic and pigeon crop- stimulating activities of human pituitary growth hormone preparations. J . Endocr. 31, 115-126.

1965. (With G. K. Benson, A. T. Cowie, C. P. Cox and Z. D. H osking.) Relative efficiency of hexoestrol and progesterone as oily solutions and as crystalline suspensions in inducing mammary growth and lactation in early and late ovariectomized goats. J . Endocr. 31, 157-164.




258

1965. (With A. T. Cowie, G. P. Cox, Z. D. H osking and J . S. T indal.) Relative- efficiency of crystalline suspensions of hexoestrol and of oestradiol monobenzoate in inducing mammary development and lactation in the goat; and effects of relaxin on mammogenesis and lactation. J . Endocr. 31, 165-172.

1965. (With G. S. Knaggs.) Oxytocin levels in the blood of ruminants with special- reference to the milking stimulus. In Advances in oxytocin research (ed. J . H. M. Pinkerton), pp. 37-49. Oxford: Pergamon Press.

1965. (With A. T. Cowie, C. P. Cox, Z. D. H osking, M. Naito and J . S. T indal.)The effects of the duration of treatments with oestrogen and progesterone on the hormonal induction of mammary growth and lactation in the goat.

J . Endocr. 32, 129-139.1965. (With G. S. Knaggs.) Levels of oxytocin in the jugular vein blood of goats

during parturition. J . Endocr. 33, 301-315.1966. (With G. S. K naggs.) Milk-ejection activity (oxytocin) in the external

jugular vein blood of the cow, goat and sow, in relation to the stimulus of milking or suckling. J . Endocr. 34, 197-214.

1967. (With E. M. Rivera, I. A. Forsyth.) Lactogenic activity of mammaliangrowth hormone in vitro. Proc. Soc. exp. Biol. Med. 124, 859-865.

1970. (With J . D. Cleverley.) The blood levels of oxytocin during machine milking in cows with some observations on its half-life in the circulation. Endocr. 46, 347-361.


B

Scientific reviews

1936. (With H. D. Kay.) The phosphatases. Ergebn. 5, 159-212.1937. (With H. D. Kay.) The phosphatases. Tabul. biol., Berl. 6, 268-279.1938. The role of the anterior pituitary in lactation. A review of recent work.

Lancet, 234 (2), 389-390.1940. Lactation. Biol. Rev. 15, 421-458.1941. Effects of oestrogens on lactation. Lancet, 240 (1), 40-41.1941. (With F. G. Young.) Prolactin as a specific lactogenic hormone. Lancet, 240

(1), 380-381.1941. Algunos aspectos del control enddcrino de la lactancia. Prensa Argent.

28, 1397-1404.1944. Recent advances in lactational endocrinology as applied to farm animals.

Vet. Rec. 56, 9-10.1944. Limitations and uses of the comparative method in medicine. Proc. R. Soc. Med.

37, 443-446.1945. The hormonal control of lactation. J . R. Soc. Arts, 93, 114-121.1947. Recent researches on the physiology of mammary development and function

Proc. R. Soc. Med. 40, 903-906.1947. Endocrine control of the mammary gland. 1. Mammary development. Br. med.

Bull. 5, 130-134.1947. Endocrine control of the mammary gland. II. Lactation. Br. med. Bull. 5, 135-

142.1947. The nervous system and lactation. Br. med. Bull. 5, 142-148.



2591948. (With F. H. Malpress.) Hormonal control of mammary growth. In The

hormones (ed. G. Pincus and K. V. Thimann), Vol. 1. Chap. 16, pp. 695-743. New York: Academic Press.

1948. (With F. H. Malpress.) Hormonal control of lactation. In The hormones (ed.G. Pincus and K. V. Thimann), Vol. 1. Chap. 16, pp. 745-805. New York: Academic Press.

1949. Biochemical aspects of mammary gland function. Biol. Rev. 24, 316-354.1949. Nutrition and female fertility. Br. J . Nutr. 3, 91-96.1949. Neuro-hormonal mechanisms in lactation. J . Endocr. 6, xxii-xxv.1950. The physiology of prolactin with special reference to the relation between

the anterior pituitary and the mammary gland. Ned. Tijdschr. Geneesk. 94, 1090-1091.

1950. Lactational physiology. In Modejrn trends in obstetrics and gynaecology, (ed. K. Bowes), Chap. 30, pp. 441-453. London: Butterworth.

1950. Hormones of the anterior-pituitary gland. Ann. Rep. chem. Soc. 47, 360-372. 1952. Aspects of pituitary-mammary gland relationships. Recent. Progr. Horm. Res. 7,

107-137.1952. Enzymic and metabolic aspects of hormonal stimulation of the mammary

gland. 2nd. Congr. Int. Biochim. July 1952, (4), 5-19.1952. Aspects of fat metabolism in the ruminant with special reference to the

biosynthesis of milk fat. Biochem. Soc. Symp. No. 9, pp. 52-65.1952. (With F. G. Young.) The galactopoietic action of anterior-pituitary extracts-

in milking cows. Xlth Int. Congr. Chem. 4, 77-79.1951. Biochemical aspects of the hormonal control of lactation. Colloques int. Cent.

natn. Rech. Scient, X X X II1950, 81-95.1952. Lactation in Marshall’s Physiology of reproduction, 3rd edn. (ed. A. S. Parkes),

Vol. 2, pp. 525-647. London: Longmans, Green.1953. The adrenal cortex and the mammary gland. In The suprarenal cortex (ed.

J . M. Yoffey), pp. 85—90. London: Butterworth Scientific Publications.1953. The use of hormones in nutrition: some practical possibilities.

Suppl. 1, pp. 214-227.1954. Invloed van thyroied-actieve stoffen op lacterende runders. Vlaams dier~

geneesk. Tijdschr. 23, 1-4.1954. (With A. T. Cowie.) Terminology in lactational physiology. Lancet, ii, 601-602. 1954. Some applications of hormones in animal husbandry. J . Soc. Dairy Tech. 7,

100- 101.

1954. Endocrine influences in lipogenesis. Int. Conf. Biochem. Problems of Lipids, Brussels, 1953, 275-280.

1954. M6canisme hormonal du d^clenchement et de l’entretien de la s£cr6tion lact£e. C.r. Seanc.Soc. Biol., Paris, 148, 5-14.

1954. Etudes sur le d^clenchement hormonal de la s£cr6tion lact^e sp&dalement chez la vache et la chevre. Annls Endocr. 15, 220-223.

1954. Recherches rScentes sur la physiologie et la biochimie de la secretion lactie. Paris:Masson; Li&ge: Desoer.

1955. (With A. T. Cowie.) Physiology of the gonadotrophins and the lactogenichormone. In The hormones, (ed. G. Pincus and K. V. Thimann) Vol. 3. Chap. 8, pp. 309-387. New York: Academic Press.

Sydney John Folley




1955. Effects of somatotrophin and other pituitary hormones on the lactating mammary glands. In The hypophyseal growth hormone, nature and actions (ed. R. W. Smith, O. H. Gaebler and G. N. H. Long). Chap. 27. Philadelphia and Toronto: Blakiston.

1955. Hormones in mammary growth and function. Br. med. Bull. 11, 145-150.1957. (With A. T. Cowie.) Neurohypophysial hormones and the mammary gland.

In The neurohypophysis (ed. H. Heller), pp. 183-197. London: Butterworths Scientific Publications.

1958. (With Mary L. M cNaught.) Hormonal effects in vitro on lipogenesis inmammary tissue. Br. med. Bull. 14, 207-211.

1958. Discussion on physiology and biochemistry of lactation: introductory remarksand concluding remarks. Proc. R. Soc. Land. B, 149, 310-303, 424.

1959. (With G. K. Benson, A. T. Cowie and J. S. T indal.) Recent developmentsin endocrine studies on mammary growth and lactation. In Recent progress in the endocrinology of reproduction (ed. C. W. Lloyd), pp. 457-490. New York and London: Academic Press.

1960. Fiziologiya i biokhimiya laktatsii. Moskva: IzdatePstvo Inostrannoi Literatury. 1960. Disorders of mammary development and lactation. In Clinical Endocrinology,

Vol. 1, pp. 518-525. New York: Grune and Stratton.1960. (With A. L. Greenbaum.) Insulin and metabolism of fatty acids. Br. med.

Bull. 16, 228-232.1960. (With Mary L. M gNaught.) Biosynthesis of milk fat. In M ilk: The mammary

gland and its secretion (ed. S. K. Kon and A. T. Cowie), Vol. 1, pp. 441-482. London and New York: Academic Press.

1961. ‘Lactation5. In The Encyclopedia of the biological sciences (ed. by P. Gray),pp. 541-542. New York: Reinhold.

1961. Recent advances in the physiology and biochemistry of lactation. Dairy Sci. Abstrs. 23, 511-528.

1961. (With A. T. Cowie.) The mammary gland and lactation. In Sex and internal secretions, 3rd edn. (ed. by W. C. Young), Vol. 1, Chap. 10, pp. 590-642. Baltimore: Williams and Wilkins.

Fiziologiya i biokhimiya laktatsii: 2-e ispravlennoe i dopolnennoe izdanie.Moskva: Izdatel5stvo Inostrannoi Literatury.

Some observations on prolactin secretion. In Advances in neuroendocrinology (ed. by A. V. Nalbandov), pp. 277-282. Urbana: University of Illinois Press.

(With E. A. J ones.) Enzymic and metabolic aspects of hormonally dependent changes in mammary gland function. Archs Anat. microsc. Morph, exp. 56, 584-595.

1969. The milk-ejection reflex: a neuroendocrine theme in biology, myth and art.J . Endocr. 44, x-xx.

1970. Lactation. In The Encyclopedia of the biological sciences, 2nd edn. (ed. byPeter Gray), pp. 478-479. New York, Cincinnati, Toronto, London, Melbourne: Van Nostrand Reinhold Company.

1970. (With G. S. Knaggs.) Physiological and pharmacological effects: mammary action. In International encyclopedia of pharmacology and therapeutics, Section 41, Vol. 1; Pharmacology of the endocrine system and related drugs: the neurohypophysis (ed. H. Heller and B. T. Pickering), Chap. 8c, pp. 295-320. Oxford and New York: Pergamon Press.

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1970. (With I. A. Forsyth.) Prolactin and growth hormone in man and other mammals. In Ovo-implantation, human gonadotropins and prolactin (ed. P. O. Hubinont, F. Leroy, G. Robyn and P. Leleux), pp. 266-278. Basel: Karger.

1970. (With A. T. Cowie.) The mammary gland and lactation. In Scientific foundations of obstetrics and gynaecology (ed. E. E. Philipp, J . Barnes and M. Newton), pp. 423-432. London: William Heinemann Medical Books Ltd.

1970. Some aspects of mammary gland research: a personal perspective. Rep. natn. Inst. Res. Dairy, pp. 15-20.

1970. The milk-ejection reflex: a neuroendocrine theme in biology, myth and art. Perspectives in Biology and Medicine, 13, 476-490.

GShort communications: Scientific journals

1931. (With E. Davis.) Effect of activated charcoal on frog heart perfusate. J . Physiol. 73, 9-10.

1933. (With G. L. Peskett.) Automatic pressure control. J . Physiol. 78, 13-14.1933. (With A. T. R. Mattick.) A stainless steel high-pressure ultrafilter. Biochem.

J . 27, 1113-1115.1934. (With G. L. Peskett.) Blood composition in relation to milk secretion. Nature,

Lond. 133, 142.1934. An electric furnace for micro-Kjeldahl digestions and similar purposes.

Biochem. J . 28, 890-891.1935. The phosphatase of the mammary gland. J . soc. chem. Ind., Lond. 54, 86.1935. A modification of the King and Armstrong method for estimation of phos

phatase. Mikrochemie, 17, 263.1935. A shaking gear for use with water-baths. J . sci. Instrum. 12, 257.1935. (With E. A. Rowsell.) An improved screw plunger for use with mercury

piston micro burettes. Mikrochemie, 18, 303-304.1935. (With P. L. Temple.) An improved thermoregulator and circuit for D.C.

supply. J . sci. Instrum. 12, 392-393.1936. Effect of oestrogenic hormones on lactation in the cow. Nature, Lond. 137,

741-742.1936. The effect of oestrogenic hormones on lactation in the cow. J . Soc. chem. Ind.,

Lond. 55, 509.1937. (With F. G. Young.) Stimulation of milk production by prolactin in the

cow. J . Soc. chem. Ind.9 Lond. 56, 96-97.1937. (With S. K. K on.) Effect of progesterone on lactation in the rat. Nature,

Lond. 139, 1107.1937. (With P. White.) Response of the pigeon crop gland to prolactin: inhibition

by oestradiol monobenzoate. Nature, Lond. 140, 505.1937. (With A. C. Bottomley.) Stimulation of nipple growth in the guinea-pig by

preparations of p-anol and its benzoate. J . Soc. chem. Ind., Lond. 56, 1054-1055.

1938. (With A. C. Bottomley.) The effect of androgenic substances on the testesand accessory reproductive organs of the immature male guinea-pig. J. Physiol. 92, 15-16.

1938. (With A. C. Bottomley.) The effect of androgenic substances on the rate of teat growth in the young male guinea-pig. J. Physiol. 92, 33-34.





1938. (With H. M. Scott Watson.) Some biological properties of diethylstilboestrol. Lancet, 253 (2), 423-424.

1938. (With S. Zuckerman.) Development of the mammary gland in the Rhesusmonkey. J . Anat. Land. 72, 613.

1939. (With A. G. Bottomley and F. H. A. Walker.) Induction of prolongedhyperglycaemia in the lactating ruminant by subcutaneous implantation of adrenalin tablets. J . Soc. chem. Ind., Lond. 58, 470.

1939. (With H. M. Scott Watson.) Induction of abortion in the cow by inunction with stilboestrol dipropionate. Lancet, 237 (2), 788-789.

1939. Sex difference in the response of the pigeon crop-gland to prolactin.Lond. 144, 834.

1939. (With A. C. Bottomley.) Inactivation of prolactin by treatment with phenylisocyanate. Nature, Lond. 145, 304.

1940. (With H. M. Scott Watson and A. C. Bottomley.) Induction of lactationin goats with diethylstilboestrol dipropionate. J . Physiol. 98, 15-16 P.

1941. (With G. W. Scott Blair, F. M. V. Coppen and F. H. Malpress.) Rheological properties of bovine cervical secretions during the oestrous cycle.

N a t u r e , Lond. 147, 453-454.1941. (With A. C. Bottomley.) Observations on teat growth in immature female

goats. J . Physiol. 99, 5-6.1941. Parathyroid glands and lactation in the rat. Nature, Lond. 147, 744.1941. (With A. G. Bottomley.) Relative growth in the individual. Nature, Lond.

148, 169.1941. (With H. M. Scott Watson and A. G. Bottomley.) Initiation of lactation

in nulliparous heifers by diethylstilboestrol. J . Physiol. 100, 7-8 P.1941. (With F. G. Young.) Artificial induction of lactation in virgin animals.

Nature, Lond. 148, 563-564.1942. Non-effect of massive doses of progesterone and desoxycorticosterone on

lactation. Nature, Lond. 150, 266.1942. (With K. M. H enry and S. K. K on.) Lactation and reproduction on highly

purified diets. Nature, Lond. 150, 318.1942. ‘Ghost* formation in subcutaneously implanted tablets of synthetic oestrogens.

Nature, Lond. 150, 403-404.1942. Retarding effect of ghost formation on absorption from subcutaneously

implanted tablets of hexoestrol. Nature, Lond. 150, 735-736.1942. (With G. W. Scott Blair and A. T. Cowie.) Rheological properties of

bovine cervical mucus. J . Physiol. 101, 11-12.1943. Tablet absorption and ghost formation. Nature, Lond. 152, 134-135.1944. (With P. M. F. Bishop.) Implantation of testosterone in cast pellets. Lancet,

246 (1), 434-435.1944. (With A. T. Cowie.) Effect of adrenalectomy and anterior pituitary injections

on mammary development. Nature, Lond. 154, 302.1945. (With P. M. F. Bishop.) Absorption of subcutaneously implanted hormone

pellets. Endocrinology, 36, 283.1945. (With A. T. Cowie.) Parathyroidectomy and lactation in the rat. Nature,

Lond. 156, 719-720.1947. (With A. L. Greenbaum.) Decrease in the arginase of the liver and mammary

gland in adrenalectomized lactating rats as compared with pair-fed controls. Nature, Lond. 160, 364.



1947.

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1948.

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1951.

(With A. T. Cowie, T. H. French and A. L. Greenbaum.) Effect of sodium intake on adrenalectomized rats receiving cortical steroids. J . Endocr. 5, xxxiii.

(With F. H. Malpress.) Some observations on oestrogen-anterior pituitary relationships. Abstr. Commun., XVIIth physiol. Congr. pp. 340-341.

(With A. T. Cowie.) The effect of adrenalectomy on lactation in the rat studied by the paired-feeding technique. J . Endocr. 5, lxxxviii.

(With S. G. Watson.) A high-speed tissue homogenizer. Biochem. J . 42, 204-206.

(With T. H. French.) Respiratory quotient of the mammary gland. Nature, Land. 161, 933-934.

(With T. H. French.) The respiratory quotient of the mammary gland. Biochem. J . 42, xlvii-xlviii.

(With S. Bartlett, S. J . Rowland, D. H. Curnow and S. A. Simpson.) Oestrogens in grass and their possible effects on milk secretion. Nature, Land. 162, 845.

(With T. H. French.) Utilization of acetate by tissues of the ruminant. Biochem. J . 43, lv.

(With T. H. French.) Acetate as a possible precursor of ruminant milk fat, particularly the short-chain fatty acids. Nature, Land. 163, 174.

(With G. L. Bailey and S. Bartlett.) Use of L-thyroxine by mouth for stimulating milk secretion in lactating cows. Nature, Land. 163, 800.

(With A. T. Cowie, T. H. French and A. L. Greenbaum.) Further observations on the effects of adrenalectomy on lactating rats studied by the paired-feeding technique. J . Endocr. 6, ii-iii.

(With A. T. Cowie.) Relative growth of the mammary gland in the normal, gonadectomized and adrenalectomized rat. J . Endocr. 6, vi-vii.

(With T. H. French.) Effect of glucose on the utilization of acetate by lactating mammary gland slices. Biochem. J . 44, xlv.

(With G. PopjAk and T. H. French.) Synthesis of the short-chain fatty acids of milk fat from acetate. Archs. Biochem. 23, 508-510.

(With P. M. Cotes, J . A. Crichton and F. G. Young.) Galactopoietic activity of purified anterior pituitary growth hormone. Nature, Land. 164,992.

(With A. L. Greenbaum.) Effect of experimental diabetes on tissue arginase levels. J . Endocr. 6, x-xi.

(With A. L. Greenbaum.) Studies on tissue arginase levels in mammals. Abstr. Commun., 1st int. Congr. Biochem. pp. 409-410.

(With T. H. French.) Studies on the intermediary metabolism of lactating mammary gland slices. Abstr. Commun., 1st int. Congr. Biochem.pp. 606-608.

(With J . H. Balmain and T. H. French.) Stimulation by insulin of in vitro fat synthesis by lactating mammary gland slices. Nature, Land. 165, 807-808.

(With G. PopjAk and T. H. French.) Utilization of acetate for milk-fat synthesis in the lactating goat. Biochem. J . 46, xxviii-xxxix.

(With J . H. Balmain.) Further observations on the stimulation by insulin of fat synthesis by lactating mammary gland slices. Biochem. J . 48, i.

(With A. T. Cowie, W. G. Duncombe, T. H. French, R. F. Glascock, L. Massart, G. Peeters and G. PopjAk.) Studies of the synthesis of milk fat by the perfused, isolated bovine udder. Biochem. J . 48, xxxix.




2641951. (With A. T. Cowie, F. H. Malpress and K. G. R ichardson.) Studies on the

induction of mammary growth and lactation in virgin goats with oestrogen, thyroxine and progesterone. J . Endocr. 7, xxi-xxii.

1951. (With A. T. Cowie, B. A. Cross, G. W. H arris, D. Jacobsohn and K. C.R ichardson.) Terminology for use in lactational physiology. Nature, Lond. 168, 421.

1951. (With J . H. Balmain and R. F. Glascock.) Stimulation by insulin of in vitrofat synthesis by mammary tissues studied with carbon-14 and tritium. Nature, Lond. 168, 1083-1084.

1952. (With J . H. Balmain and R. F. Glascock.) Isotope and respiration studieson in vitro fatty acid synthesis by mammary tissue: species differences in effects of insulin and glycerol. Biochem. J . 50, xxix.

1953. (With J . H. Balmain and R. F. Glascock.) Relative utilization of glucoseand acetate carbon for lipogenesis by mammary gland slices, studied with tritium, 13C and 14C. Biochem. J . 53, xxvi.

1954. (With J . H. Balmain, R. F. Glascock and M. L. M cNaught.) Effects ofadrenal corticoids on lipogenesis in mammary tissue in vitro. Biochem. J . 56, vi.

1954. (With T. R. Bradley, F. W. Landgrebe and G. M. Mitchell.) Effect of melanophore-dispersing hormone (‘B’) on lactating mammary gland slices in vitro. Biochim. biophys. Acta, 13, 449-450.

1957. (With G. K. Benson.) Retardation of mammary involution in the rat by oxytocin. J . Endocr. 14, xl-xli.

1960. (With G. K. Benson.) Effects of oxytocin on the functional capacity of pituitary grafts with particular reference to prolactin secretion. Acta endocr., Copenh. 35 (supp. 51), 1147.

1962. (With A. Chadwick.) Lactogenesis in pseudopregnant rabbits treated withACTH. J . Endocr. 24, xi-xii.

1963. (With A. Chadwick.) Lactogenic and pigeon-crop-stimulating effects ofhuman pituitary growth hormone. J . Endocr. 26, xiii-xiv.

1963. (With P. Andrews.) Molecular weights of bovine, ovine and porcine pituitarygrowth hormones estimated by gel filtration on Sephadex. Biochem. J . 87, 3 P.

1964. (With G. S. Knaggs.) Observations on-oxytocin release in ruminants. J .Reprod. Fert. 8, 265-266.

1970. (With A. S. McNeilly.) Blood levels of milk-ejection activity (oxytocin) in the female goat during mating. J . Endocr. 48, ix-x.

D

Other Publications

1936. The hormonal control of lactation. Agric. Progr. 13, 120-125.1938. Ductless glands and milk secretion. Tearb. Brit. Goat Soc. pp. 87-89.1938. Extending animal productivity. Fmr & Stk-Breed. 52, 1622.1941. (With F. H. Malpress.) Bovine sterility: the use of oestrogens for the induction

of oestrus in anoestrous cattle. Vet. Rec. 53, 551.(With G. W. Scott Blair, F. H. Malpress and F. M. V. Coppen.) The

oestroscope: an instrument for detecting oestrus in cows. Vet. Rec. 53, 693-695.


1941.



2651942. An aid to back-end calvings. Fmr & Stk-Breed. 56, 18.1942. Artificial lactation in the goat. Mon. J . Brit. Goat. Soc. 35, 109-111. 1944. Modem trends in zootechnology, Mon. J . Brit. Goat Soc. 37, 30-32.1944. The problem of hermaphrodism. Tearb. Brit. Goat Soc. pp. 11-14.1950. Physiological mechanism of milk secretion. Land. 166, 853.1954. Comparative endocrinology of vertebrates. Nature, Land. 174, 543.1955. Henry Stanley Raper—Obituary Notice. J . Chem. Soc. pp. 2987-2988.1962. The brain’s influence on milk production. New Scientist, 14, 654-656.1963. Concluding remarks: Biochemical Society Symposium No. 24 (Oxford: 19

July 1963). In The control o f lipid metabolism (ed. by J . K. Grant),pp. 181-183. London and New York: Academic Press.

1964. Opening remarks. In Small animal anaesthesia (ed. O. Graham-Jones), pp. 1-3.Oxford: Pergamon Press.

1965. The hypothalamo-neurohypophysial system and the milk-ejection reflex.Nature, Lond. 207, 1267-1268.

1967. Research on the physiology of lactation. Anim. Prod. 9, 572.1969. Chairman’s introduction: Symposium on lactogenesis. In Lactogenesis: the

initiation of milk secretion at parturition (ed. by M. Reynolds and S. J . Folley), pp. i-3. Philadelphia: University of Pennsylvania Press.

1968. Lactation symposium : chairman’s introductory reflexions. Proceedings of theInternational Union of Physiological Sciences, Vol. VI, pp. 206-207. XXIV International Congress, Washington, D.C.

Sydney John Folley



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