emmanuel ciprian amoroso, 16 september 1901 - 30...

1901 - 30 October 1982Emmanuel Ciprian Amoroso, 16 September

R. V. Short, F. R. S.

, 2-30, published 1 November 1985311985 Biogr. Mems Fell. R. Soc.

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EM M ANUEL CIPRIAN AMOROSO

16 S eptem ber 1901 — 30 O ctober 1982

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EM M ANUEL CIPRIAN AMOROSO

16 Septem ber 1901 — 30 O ctober 1982

Elected F .R .S . 1957

B y R. V. S h o r t , F.R .S .

E m m a n u e l C i p r i a n A m o r o s o , or ‘A m o’ as he was affectionately known to his thousands of friends and acquaintances around the world, is the only T rin idad ian , indeed the only W est Indian, to have been elected into the Fellowship of the Royal Society— a signal achievement. T h a t he was able to overcome the prejudices of race and colour, so prevalent in British society particularly in the earlier decades of this century, is a tribute to his great personal charisma, his ability to speak in many tongues and so endear him self to people in nations and cultures very different from our own, and to the sheer force of his intellect. His studies on the formation, structure and function of the m am m alian placenta and its role in the evolution of viviparity form the basis of his international scientific reputation. He will be rem em bered more for his ability to absorb a mass of factual inform ation scattered throughout the w orld’s scientific literature over the last few centuries and to produce a concise, succinct sum m ary of the present state of the art, than as an experim entalist. His skills lay more in observation and interpretation, in the correlation of structure w ith function, than in the design and execution of planned experim ents. But it is as a speaker, indeed as an orator, and as a teacher that he will be best rem em bered throughout Europe, Australasia, Africa and the Americas. Listening to him deliver a lecture on a formal occasion, one had the distinct im pression that if he had not succeeded as a scientist he would have becorpe a great actor, or even an inspiring preacher; in a bygone age he would have been one of those great storytellers who could have held his audience enthralled. As Professor Courtenay Batholomew said of him , ‘He had a charisma and an aura about him whereby when he walked into a gathering one had to enquire: “who is this man?’’ ’

3



E a r l y L i f e

A m o or ‘M an n ie ’, as he was know n to h is fam ily, was b o rn in P o rt of Spain, T rin id ad , on 16 S ep tem b er 1901, the th ird o ldest o f a fam ily of twelve th a t included eight b ro th ers , one of w hom was adopted . I t was an exem plary C atholic fam ily and th is C atholic u p b rin g in g was to have a deep influence on A m o th ro u g h o u t the w hole o f his life; he a ttended M ass w ithou t fail every Sunday, and died in a state of grace. T h e ch ild ren w ere rem arkably ta len ted . O ne of his b ro th ers , Felix, tau g h t at S t M ary ’s College in T rin id ad ; Span ish was his m ajor subject, b u t he also tau g h t F rench and English . H e clearly shared A m o ’s delights in the beau ty of language. P rofessor B artholom ew recalled how one day in class Felix said to his pup ils ‘L e t us com pare the beau ty and the rh y th m of the Spanish language vis-a-vis E nglish . F o r exam ple in E nglish you say: “ V ery b eau tifu l” , in Spanish we say “ A bso lu tam en te b e lla” , em phasizing phonetically and dram atically the difference in the m usicality and rom ance of the languages’. O ne can ju s t hear A m o’s deep, rich resonan t voice saying those very w ords. A ndre was to becom e a B arris ter-a t-L aw ; he was called to the Bar at G ra y ’s In n , and at the tim e of his death was C om m issioner of In lan d R evenue of T rin id a d and T obago . M iguel was C om ptro ller of C ustom s and Excise in T rin id ad ; P edro , the adop ted son, now lives in Caracas; and the eldest b ro th e r, G ilbert, em igrated to the U n ited States. B ut it was Eric, the youngest, know n as H erm ann by the fam ily, w ho followed closest in A m o’s footsteps. E ric, like his elder b ro th e r before h im , qualified in M edicine in D ub lin , and for a tim e lectu red in A natom y at U n iversity College, D ub lin . H e th en w ent to E ngland and lectu red in A natom y at S t B artho lom ew ’s H osp ital before eventually becom ing a general p rac titioner in C lerkenw ell, w ith a private practice in W im pole S treet. H e died on 2 O ctober 1982, ju s t a few weeks before A m o. Val, the only surviving full b ro th er, rem ained in T rin id ad becom ing Secretary of the T r in id a d B uild ing and L oan A ssociation. R ita, one of his sisters, rem ained in T rin id ad , and Evelyn m arried and w ent to the U n ited States.

A m o’s father, T hom as A m oroso, was a w ell-respected W est Ind ian . As a young m an he had been em ployed as a book-keeper on one of the estates in T rin id ad , b u t he rose to becom e a p ro p rie to r, ow ning estates of his own. H e dealt m ainly in cocoa and at first his business p rospered , b u t it failed after a slum p on the m arket, w hich took its toll of m any dealers at th a t tim e. H e therefore reverted to his fo rm er occupation of bookkeeping. A m o’s m other, Ju liana C enteno, was of V enezuelan (Spanish) descent. She was ju s t un d er 5 feet tall, and was a sim ple person w ith trem endous charism a, reso lu tion and resourcefulness, to w hom Am o was deeply devoted. In the lean years th a t followed the slum p she did sewing on con tract at hom e, as well as m aking all the clothes w orn by her sons. A lthough always k ind and though tfu l, she was also a s tric t d iscip linarian in the hom e. She loved flowers and p lants, and always found the tim e to p o tte r in her little garden.

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Emmanuel Ciprian Amoroso 5

D uring his childhood Amo spent m any of his school holidays at the hom e of U ncle Em m anuel (his fa ther’s brother) and his wife Juliana. A unt Ju liana’s b ro ther Abeline, who was about A m o’s age and who was also to become a physician, became his constant com panion. T h e story is told of how these two boys were staying w ith Uncle Em m anuel and A unt Juliana at the tim e of the b irth of their second child. T hey noted that after all the hustle and bustle of the b irth was over, the midwife went down to the bottom of the garden where she was seen to bury a somewhat bloodstained parcel. T h a t evening, Amo and his reluctant co-conspirator, arm ed w ith a sharp knife and w ith light from a flambeau, dug up the placenta-—for so it proved to be— and proceeded to dissect it. T hey were so engrossed in this pursu it that they did not hear the approach of U ncle Em m anuel, who proceeded to give the two young boys a sound hiding— perhaps a baptism for A m o’s lifelong study of the placenta.

A m o’s school days were happy ones. He attended St T hom as’s Preparatory School until 1913 when he entered St M ary’s College, from which he took the Junior Cam bridge Certificate exam ination in 1917. He narrow ly missed obtaining one of the two coveted Island Scholarships that enabled the recipient to study Law or M edicine at an overseas university, although he was top of the applicants from St M ary s College. U nfortunately he was forced to leave school early because his sight failed him after a severe attack of typhoid fever. He subsequently recovered his vision in one eye, bu t could only dimly discern form with the other. Following this setback he m atriculated on his own, and subsequently returned to St M ary’s College for a brief period in order to teach.

Amo was always a lively and active person. Coming from such a large family there was always m uch activity in the home, with many guests and visitors. Cricket was a particularly popular pastime, with a box as the wicket and a handy piece of wood for a bat. He was also keen on football, playing back in his college team and later for one of the leading clubs, M aple. K ite flying from the rooftops, spinning tops and pitching marbles were other popular pastimes, as was the T rinidadian version of Pooh sticks’, known as ‘Jockey’. After rain, when there was water flowing in the street drains, pieces of charcoal or wood dipped in wax were dropped into the torrent, and all the boys from the neighbourhood would gather together to urge their ‘horses’ down the street. Amo also belonged to the D ram atic Society in St M ary’s College, playing the Earl in Little Lord Fauntleroy; he also took up weight lifting, and joined the Knights of St John, a religious society.

D uring the holidays the entire family would sometimes go to the country home of his m other’s aunt, known as T he D epot , near the present campus of the University of the W est Indies. It was here that the produce from the estates was stored to await transport to the city by train, and a favourite pastime was to search for sugar cane that had fallen off the wagons. ‘Sucking cane’ required strong jaws and sound teeth!



6 Biographical Memoirs

A c a d e m i c c a r e e r

A m o arrived in D u b lin at the age o f 21, w here he en ro lled in U n iv ers ity College of the N ational U n iv ers ity o f Ire land . H e m u st have been a q u ite ou ts tan d in g s tu d en t because he was aw arded first class ho n o u rs and cam e first in all the exam inations in science and m edicine. H e gained a n u m b e r o f College and U n iversity scho larsh ips, and prizes in bo tany , zoology, chem istry and physics (1923), anatom y and physio logy (1925), p a th o logy, pharm acology, m ateria m edica and th e rap eu tics (1927), and m ed icine, obstetrics, and the Jo h n M cA rd le M edal in su rgery (1929). T o m ake ends m eet A m o had to sell new spapers in th e s tree t ou tside the railw ay station . As was th e custom in those days, he also held a n u m b e r of p a rt-tim e dem o n stra to rsh ip s d u rin g the course o f h is m edical tra in in g . T h ese w ere in zoology (S ep tem b er 1923 to Ju n e 1924), anatom y and physiology (S ep tem b er 1925 to Ju n e 1926), patho logy (S ep tem b er 1927 to D ecem ber 1927), pharm acology (S ep tem b er 1927 to Ju n e 1928) and surgical anatom y (S ep tem b er 1927 to D ecem b er 1928). A ll th is ad d itional w ork obviously d id n o t d e trac t from his studies; ra th e r, it enabled h im to acqu ire the ancien t a rt o f lec tu ring— an expression of learn ing com bined w ith o ra to ry— the h a rd way, by teach ing his fellow studen ts. H e also found tim e for sp o rting activities, becom ing a boxer of som e renow n and an excellent sw im m er.

T h e 1920s w ere s tirrin g tim es in Ire land , th en in the g rip o f a civil w ar. T h is , perhaps coupled w ith the na tu ra l rom antic ism of its people and the p e tvad ing influence of C atholicism , developed w ith in A m o a deep and endearing love for the Irish ; w hen in his 81st year he was inv ited to address the Irish R egion of the A ssociation o f V eterinary T each ers and R esearch W orkers in D u b lin in A pril 1982, he accepted w ith alacrity . H e began his d iscourse by expressing his thanks in Irish and th en proceeded to tell a joke against h im self about a W est In d ian hosp ita l p o rte r, before em bark ing on a scientific d iscussion abou t the p lacenta.

A fter ob ta in ing a B .Sc. w ith honours in anatom y in 1926, A m o finally g radua ted M .B ., B .C h ., B .A .O ., cum , in Jan u ary 1929, and for the rem ainder o f th a t year he was a Surgical In te rn at the Jerv is S tree t H osp ital in D ub lin . In D ecem ber he was aw arded a T rav e llin g S tu d e n tship by the N ational U n iversity of Ire land for h is thesis on ‘M yelination of the cranial nerves of the p ig ’. T h is scholarsh ip was a key event in the shaping of his fu tu re scientific career, for it enabled h im to spend the years of 1930 to 1932 in G erm any , s tudy ing at the A lb ert-L u d w ig s U niversity in F re ib u rg , and th en at the K aiser W ilhelm In s titu t fu r Z ellforschung in B erlin , w here he learned physiology, h istological tech n iques and tissue cu ltu re m ethods u n d e r P rofessor R. T ren d e len b erg , P rofessor R. K rause and Professor R hoda E rd m an n . H e w orked on the factors contro lling the grow th , d ifferentia tion and de-d ifferen tia tion of epithelial cells in tissue cu ltu re , and in 1931 he pub lished , in G erm an , his first scientific paper on the em bryological developm ent o f the pancreas of




the chick (1)*. W hile in G erm any he was to learn m uch m ore than mere scientific techniques; his linguistic gifts soon enabled him to m aster the G erm an language, and this was to prove invaluable to him in later years as m uch of the im portan t early descriptive work on the placenta had been carried out by the great G erm an anatom ists of the 19th and early 20th centuries like Bischoff, G rosser, Reichert, Schauder and Strahl. T hus Amo becam e one of the few people in the English-speaking world who was fully acquainted w ith the details of their work.

In 1933 Amo was appointed to a D em onstratorship in Histology and Em bryology at U niversity College London, under Professor J. P. Hill, F .R .S ., and he em barked on a P h .D ., w hich was awarded in 1934 for his w ork ‘O bservations on the developm ent of the urogenital system of the rabbit, w ith special reference to the developm ent of the M ullerian ducts’ (2). I t m ust have been an exciting departm ent in which to work: J. P. Hill was by then one of the w orld ’s leading reproductive biologists, and his definitive accounts of the placentae of a wide variety of Australian m arsupials cannot have failed to influence the young Amoroso, who would not have been exposed to such a breadth of interests during the course of his orthodox m edical training.

W ith his thesis com pleted and no prospect for continued em ploym ent at U niversity College, Amo was in a dilem m a as to what to do next. He used to tell the story of how he w ent to Folkestone for the day, and gained some inspiration by looking at the statue of W illiam Harvey, erected to com m em orate his b irth there on 1 A pril 1578. W hile travelling on the train back to London that evening, he fell into conversation with a fellow passenger who turned out to be none other than D r H ew lett-Johnson, later to become famous as the ‘Red D ean’ of C anterbury Cathedral. He told Amo of a recently advertised vacancy for a Senior Assistant in charge of H istology and Em bryology at the Royal Veterinary College in Cam den Tow n, and encouraged him to apply for the post. Amo duly did, and was appointed w ith effect from O ctober 1934.

Those early days at the Royal Veterinary College m ust have been the m ost trying tim e in the whole of A m o’s life. As the first coloured m em ber of staff his presence was bitterly resented by many of his academic colleagues, who dismissed him as a ‘nigger’, not even the equal of that sprinkling of Asian students whom it was his duty to teach. Even his Professor, James M^cCunn, turned against him , and the students were encouraged to do likewise. W ere it not for those m em bers of the staff who rallied round him, like Jack Scorgie, ‘O scar’ Ottaway and Clifford Form ston, and the support of outsiders like Professor C. A. Lovatt Evans, F .R .S ., it is difficult to see how he could have survived those early years. N ot only was Amo resented because of his race and colour, but his medical degree, his intellectual superiority, his immense physical strength and his brilliance as a teacher all served to arouse strong feelings

* N um bers given in this form refer to entries in the bibliography at the end of the text.



of jealousy am ong his colleagues, m any of w hom were d istinguished by their m ediocrity.

As if this were not enough, in 1936 Am o em barked on a d isastrous m arriage to M iss ‘P e te r’ Pole. A t the w edding breakfast she openly announced to the guests tha t she w ould never bear him any children. She left him very soon thereafter, b u t because of his deeply-held Catholic beliefs he could not b ring him self to divorce her, and so she rem ained a financial m illstone round his neck for m ost of his days. T h is was undoubted ly the greatest tragedy of A m o’s life. H e seldom spoke about it, even to his closest friends, and he rem ained a very private and lonely person thereafter. H is only hobbies were reading thrillers, w atching football and cricket on the television, tend ing his roses and, of course, socializing. H e was so fond of ch ildren tha t he w ould have m ade an ideal family m an. As it was, he took refuge in work, and drove him self m ercilessly in his desire to succeed.

T o understand Amo it is necessary to understand those stresses and strains to w hich he was exposed in his form ative years. Sm all w onder tha t he was sensitive, quick to perceive an un in ten tioned slight and to take offence, and desperately anxious for recognition and approval by the Establishm ent. A nother problem tha t was to dog Am o th roughou t his entire life was his eyesight. H e had virtually lost the sight in his left eye as a result of an attack of typhoid fever and the righ t eye also requ ired a thick corrective lens, hardly an auspicious beginning for som ebody who was to make his nam e as a histologist, for by his own adm ission he was electrically and m echanically incom petent, and relied on his technician to cut his histological sections and take all his photom icrographs. H is skill lay in in terp re ting w hat he saw dow n the m icroscope, and in later years he was to be greatly troub led by failing eyesight and the fear of going com pletely blind.

W hen Amo joined the Royal V eterinary College in 1934, it was still a self-financing institu tion w ith no links to the U niversity of L ondon. A lthough founded as long ago as 1791 under the influence of John H un ter, F .R .S ., its activities were still centred on the diseases of the horse, w hich was the type anim al for anatom ical dissection. T h e College was slow to extend its interests to em brace the diseases of o ther farm livestock, let alone those of dom estic, laboratory or exotic anim als, w hich were regarded as very m uch th ird-class citizens. As the fortunes of agriculture declined betw een the wars during the econom ic depression, so too did the fortunes of the veterinary schools. T h u s w hen Amo jo ined the staff there w ould have been little in terest in or support for som ebody w ith no train ing in anim al husbandry or disease, who was concerned prim arily w ith developing his own research interests in the com parative anatom y of the placenta in a range of species tha t at the tim e were considered of absolutely no in terest to the veterinary profession w hatsoever. I f ever there was a classical exam ple of a m an being before his





tim e, and of adversity being the spur to success, it was A m oroso’s career at the Royal V eterinary College.

T h e next m ajor event was the outbreak of W orld W ar II in 1939, and the evacuation of the Royal V eterinary College to the U niversity of Reading, and of its F ield Station to nearby Streatley. Amo was given a N issen h u t at Reading U niversity for a laboratory, and m oved into ‘d igs’ w ith M rs Howes who was the housekeeper at the U niversity Halls. H er husband had died in 1934 leaving her w ith three children to bring up, and Amo becam e like a second father to them . One of the girls, M ay, was subsequently to type A m o’s famous chapter for Marshall’s Physiology of reproduction (36), and the o ther, Kay, who was a driver th roughout the war, was to becom e his ‘adopted’ daughter and lifelong friend and confident. H e telephoned her every evening of his life, no m atter w here in the w orld he happened to be, b u t such was his sense of decorum that he never deem ed it p roper to spend a n ight under her roof.

T h e w ar years m ust have been a difficult tim e for staff and students alike, b u t a great cam araderie was to develop in the Royal V eterinary College during tha t tim e, and vets still speak nostalgically about those early days. One has the feeling tha t Amo was becom ing increasingly accepted w ith in the College, and it was during those wartim e years, when he was already in his forties, that his research career first started to develop. H is close proxim ity to O xford enabled him to collaborate w ith A. E. Barclay, K. J. Franklin and M . L. P richard in their studies on the com parative anatom y of the fetal circulation, resulting in a num ber of jo in t publications, and he also worked w ith S. J. Folley, F .R .S ., at the N ational Institu te for Research in D airying, Reading. He also entered into collaboration w ith A. S. (later Sir Alan) Parkes, F .R .S ., at M ill Hill. H e was to becom e a lifelong friend of Parkes’s wife, D r R uth Deanesly.

A fter the war, the Royal V eterinary College returned once more to Cam den Tow n, and A m o’s research career really began to flourish. He collaborated w ith a num ber of physiologists at M ill Hill, such as O. G. Edholm , who was previously his professor at the Royal V eterinary College, and I. W . Rowlands, and began to build up a small group of his own w ith physiologists and anatom ists such as F. R. Bell, A. S. King, A. M . Law n and H . Rosenberg, some of whom had come under his spell as veterinary students. In later years, many of his form er students and colleagues were to become em inent teachers and research workers, holding chairs or directing active research groups in this country and abroad. One thinks, in particular, of people such as Professor J. M. Bedford, Professor F. R. Bell, Professor J. D. Biggers, Professor C. A. F inn, Professor P. A. Jewell, Professor A. S. King, Professor J. A. Laing, Professor G. E. Lam m ing and Professor D. G. Porter, and D r B. A. Cross, F .R .S ., D r L. W . Hall and D r Barbara W eaver, to name but a few. But his influence spread far beyond the Royal Veterinary College and he



encouraged m any o thers, m yself am ong them , to pu rsu e research careers in rep roductive biology.

T hose im m ediate postw ar years also saw the developm ent of A m o ’s grow ing in terest in com parative anatom y and physiology. In 1947 he becam e a Fellow of the Zoological Society of L ond o n , and kept in constant touch w ith the P rosec to rium w here the rep roductive trac ts of all the anim als th a t died w ere p reserved for h im , la ter to cram the shelves of his laboratory at the Royal V eterinary College. I t was en tire ly ap p rop ria te th a t in 1948 he should be appo in ted P rofessor o f Physiology in the Royal V eterinary College, and in 1950, w hen the College cam e u n d e r the w ing of the U niversity of L ondon , he becam e P rofessor of V eterinary P hysiology of th a t un iversity , a post he was to hold u n til his re tirem en t in 1968.

A m o’s com parative in terests w ere to serve h im in good stead w hen he visited the U n ited States for the first tim e in 1954; he soon estab lished a close personal friendsh ip w ith G . B. W islocki, the P rofessor o f A natom y at H arvard , w ho shared A m o’s in terests in the com parative m orphology of the p lacenta, and w ith E. W . D em psey, the P rofessor of A natom y at W ash ing ton U niversity , S t Louis, M issouri, w here he was to hold a V isiting P rofessorsh ip d u rin g the sum m er vacations of 1954, 1958 and 1961. I t was here th a t he first m et P rofessors C arl and A nn C ori, N obel L aureates, w ho were to becom e his lifelong friends. A m o used to tell a delightfu l story of how he, D em psey and W islocki w ere invited to dine one evening w ith a rich w idow in N ew York, w ho W islocki and D em psey were hoping w ould donate som e funds for th e ir research. T h e lady in question , w hom we shall call M rs W annam aker, was about to carve the turkey for th is d istingu ished gathering w hen, tu rn in g to A m o, she enqu ired w hat type of m eat he w ould like. A m o replied ‘T h e breast, p lease’, w hich clearly shocked M rs W annam aker. She th en politely inform ed A m o th a t in h e r coun try it was considered vulgar to use th a t w ord; one should refer instead to ‘w hite m eat’ o r ‘dark m ea t’. T h e following day, on the eve of his d ep artu re from Idlew ild (now K ennedy) A irport, A m o sent M rs W annam aker a beau tifu l o rch id w ith a note attached, thank ing her for the excellent d inner party , and expressing the hope th a t she w ould p in the o rchid to her ‘w hite m ea t’ in m em ory of the occasion.

In parallel w ith his developing scientific career, A m o began to assum e increasing adm inistrative responsibilities. H e was a founder m em ber of the Society for Endocrinology, inaugura ted in 1946, and he was elected to its com m ittee from 1950 to 1953, and in 1956 he becam e T reasu re r , a post he held un til he becam e C hairm an from 1961 to 1966. H is years as T reasu re r w ere m ost im p o rtan t ones for the Society. W ith the guidance of his friend and colleague Professor G ladw in B uttle, the P rofessor of Pharm acology at the School of Pharm acy in L ondon U niversity , and som ething a financial w izard, he was able to p u t the finances of the Journal o f Endocrinology and the Society for Endocrinology on a firm

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foundation, ensuring that the profits from the Journal company, w ith its charitable status, were used to finance and subsidize the activities of the Society— a situation that continues to this day. D uring his chairm anship the Society also had to play host to the Second International Congress of Endocrinology, which was held in London in A ugust 1964. At the opening session in the Royal A lbert Hall, Amo was able to welcome the guests in D anish, G erm an, French, Italian, Spanish and Portuguese. He also m ade a great im pression on Professor George Corner, For.M em .R .S ., the forem ost overseas guest, by furnishing him for the duration of his stay in London w ith a chauffeur-driven Rolls Royce. Such a gesture was typical of Amo: he had a great sense of occasion and always added a touch of class to everything he did. T h e bow tie, the small red rose in the buttonhole, the fine cigar, the charm ing letter of thanks— they were all part of his personal m ystique.

A m o’s success in helping to organize the International Endocrine Congress resulted in his election to the chairm anship of the Executive C om m ittee of the In ternational Society for Endocrinology from 1964 to 1967, in preparation for the next Congress, which was held in W ashington, D .C . Perhaps because of his heavy involvem ent w ith the Society for Endocrinology, Amo was slow to become involved w ith the affairs of the Society for the S tudy of Fertility, form ed in 1950 and for some time regarded as a rival splinter-group of the Society for Endocrinology. However, Amo was to become C hairm an of the Council, the Advisory Com m ittee and the Executive Com m ittee of the associated company, the Journals of Reproduction and Fertility L td , in the period from 1968 to 1976, when his experience with the Journal of Endocrinology was of great value in helping to make this new publishing enterprise a great financial success. He also helped the Society for the Study of Fertility by serving as Secretary to two highly successful International Symposia on Com parative Reproduction that it organized, one being held in Nairobi, Kenya, in 1968, and the other in Edinburgh in 1972.

Amo was to play an im portant role in the developm ent of the Zoological Society of London. In 1951 D r Leo H arrison M atthews, F .R .S ., was appointed Scientific D irector of the Zoological Society. He shared A m o’s interests in the reproductive biology of wild mammals and persuaded him to collaborate on a series of studies of the grey seals off the coast of Pem brokeshire, together with another anatom ist from the London Hospital, Professor R. J. H arrison, F .R .S ., and D r Idwal Rowlands. A m o’s academic interests in wild animals led him to give his full support to the Secretary of the Zoological Society, Lord Zuckerman, O .M ., F .R .S ., in his efforts to transform the Fellowship from a club for old gentlem en into an organization for the pursuit of scientific goals w ithin a zoo environm ent. Amo served on the Council of the Zoo from 1955 to 1958 and from 1962 to 1965, and was also a m em ber of the G arden and W hipsnade Committee, the Animal Welfare and Husbandry




Com m ittee, the Zoological Research Com m ittee, the Publications and L ibrary Com m ittee, and, notably, the Breeding Policy Com m ittee, on which he served from 1959 to 1968.

U ndoubtedly the crow ning achievem ent of A m o’s scientific career was the publication in 1952 of his m asterly review, ‘P lacentation’ (36), in the second volume of the th ird edition of Marshall's Physiology of reproduction, edited by Sir Alan Parkes. T h e fact tha t this volum e was reprin ted four tim es is a tribu te to A m oroso’s chapter, w hich is still to this day the definitive account of placental m orphology. T h e other contributions have become largely outdated w ith the passage of tim e, whereas A m oroso’s m eticulous account of his own m orphological o b servations, and his com prehensive review of the works of his p redecessors, have w ithstood the test of time. It was this publication, m ore than any other, that form ed the basis of his international scientific reputation and led to his election into the Fellowship of the Royal Society in 1957, soon to be followed by election to Fellow ships of the Royal College of Surgeons of England (1960), the Royal College of O bstetricians and Gynaecologists (1965), the Royal College of Physicians of London (1966) and the Royal College of Pathologists (1973), a rem arkable testim onial to his w idespread and growing popularity w ithin the medical profession. I t is sad to reflect that although he did so m uch m ore for the veterinary profession, the Royal College of V eterinary Surgeons could only see fit to make him an H onorary Associate (in 1959), presum ably denying him full Fellowship on the grounds that he did not possess a veterinary qualification.

Amo was also to become a Fellow of the Institu te of Biology (1957), of the Royal V eterinary College (1969), and of U niversity College London(1970) . He was aw arded an H onorary D octorate of Science by the National U niversity of Ireland in D ublin Castle on 11 July 1963, one of the other recipients tha t day being none other than President John Fitzgerald K ennedy of the U nited States. Amo also received H onorary D octorates of Science from the U niversity of Illinois (1967), the U niversity of N ottingham (1970), the U niversity of the W est Indies(1971) and the U niversity of G uelph, O ntario (1976), and an H onorary D octorate of V eterinary Science from the U niversity of Chile (1966). After his retirem ent from the chair of veterinary physiology he was also made an H onorary M em ber of, among others, the Society of E ndocrinology, the Society for the S tudy of Fertility, the Physiological Society and the Anatomical Society of G reat Britain and Ireland.

W hen Amo retired from the chair of veterinary physiology in 1968 to become Professor Em eritus, he took on a new lease of life. He moved from London to Cam bridge to live in a small house in D erw ent Close, Cherry H inton, and in 1969 he was appointed Visiting Scientist and given laboratory space at the A gricultural Research Council Institu te of Animal Physiology at Babraham , initially under the directorship of




D r R. D . K eynes, F .R .S ., and subsequently under D r B. A. Cross, F .R .S . T h is was the ideal env ironm ent for Amo; he becam e a valued colleague, collaborator and friend of the staff, and a constant source of stim ulus to the m any studen ts and visitors passing th rough the Institu te . H e w orked particu larly closely w ith D r J. S. Perry, w ith w hom he had collaborated in earlier years in the ir definitive study of the placenta of the A frican elephant, and also w ith D r R. B. H eap, whose w ork on the endocrine m echanism s responsible for the recognition of pregnancy in sheep and pigs was close to A m o’s heart.

P inned up on the no ticeboard of A m o’s office at B abraham was a quo tation , w ith no a ttribu tion , tha t he often copied and gave to research studen ts in the group. I t ran as follows:

‘ “ G en tlem en” , I have ven tu red to offer you these considerations upon the scholar’s place, and hope, because I thought tha t standing, as m any of you now do, on the th reshold of this College, g irt and ready to go and assum e tasks, public and private, in your country, you w ould no t be sorry to be adm onished of those prim ary duties of the intellect w hereof you will seldom hear from the lips of your new com panions. Y ou will hear every day the m axim s of low prudence. Y ou will hear tha t the first du ty is to get land and m oney, place and nam e. W hat is this T ru th you seek, w hat is this Beauty, m en will ask w ith derision? If nevertheless G od have called any of you to explore tru th and beauty , be bold, be firm , be true. W hen you shall say: “ As others do, so will I, I renounce, I am sorry for it, my early visions; I m ust eat the good of the land and let learning and rom antic expectations go un til a m ore convenient season;” then dies the m an in you; then once m ore perish the buds of art, and poetry, and science, as they have died already in a thousand thousand m en. T h e hour of th a t choice is the crisis of your history, and see that you hold yourself fast by the intellect. I t is this dom ineering tem per of the sensual w orld tha t creates the extrem e need of the priests of science. Be content w ith a little light, so be it your own. Explore and explore. Be neither chilled nor flattered out of your position of perpetual inquiry. N either dogm atize, nor accept ano ther’s dogm atism . W hy should you renounce your right to traverse the star-lit deserts of T ru th , for the p rem ature com forts of an acre, house, and barn? T ru th also has its roof, and bed, and board. M ake yourself necessary to the world, and m ankind will give you bread, and if not store of it, yet such as shall not take away your property in all m en’s affections, in art, in nature, and in hope.’

T h is quotation has an interesting history. I t is in fact part of Ralph W aldo E m erson’s O ration on L iterary Ethics, delivered before the L iterary Societies of D artm outh College, New H am pshire, on 24 July 1838. However, the perceptive will notice that the quotation contains a




num ber of m inor inaccuracies, and tw o sentences are m issing. T h e original, correct quotation had been sent to m e by m y father w hen I was a schoolboy in 1945, uncertain about the direction tha t m y own fu tu re was to take. I was so inspired by E m erson’s advice, w hich was to becom e m y credo, tha t I kept the quotation in m y wallet, and com m itted the w ords to m em ory. An unfortunate boating accident— a shipw reck at sea— resulted in salt-w ater dam age to the en tire contents of m y wallet, so I had to retype the quotation, largely from m em ory, and in so doing I inadvertently in troduced the errors.

In later years, as I came to know Am o personally and to value him as a friend, I though t tha t he too m ight appreciate the quotation , and so one day I sent him m y retyped version, unaw are of its inaccuracies. C u riously, Amo barely acknowledged m y letter, and never subsequently discussed the quotation w ith m e, although it m ust have im pressed him deeply; I have now discovered tha t his copy, w hich occupied such a prom inen t place on his notice board, corresponds exactly, w ord for w ord, m istake for m istake, w ith the version tha t I still carry in m y wallet to this day! Perhaps Amo w ould also have appreciated the advice in E m erson ’s two m issing sentences:

‘Bend to the persuasion w hich is flowing to you from every object in nature, to be its tongue to the heart of m an, and to show the besotted w orld how passing fair is w isdom . Forew arned tha t the vice of the tim es and the country is an excessive pretension, let us seek the shade, and find w isdom in neglect,’

As an Em eritus Professor w ith no teaching com m itm ents, Am o was free to travel, and during his tim e at Babraham he developed particularly close links w ith Professor G. E. Lam m ing and his students in the School of A griculture at the U niversity of N ottingham . Amo was appointed a Special Professor in the D epartm en t of Physiology and E nvironm ental Studies in 1973, and until his death he spent part of each year living and w orking at Sutton Bonington. H e gave a series of lectures, dem onstrations and tutorials each year to undergraduates on placentation and the im m unological aspects of pregnancy, and these were enorm ously popular. Amo obviously greatly appreciated and enjoyed this N o ttin g ham contact, as did the students who were fortunate enough to be taught by him .

D uring his retirem ent Amo also developed an in terest in the rep ro ductive physiology of the green tu rtle . T h is arose th rough an organization called M ariculture L td , which had set up a series of large salt-w ater enclosures for the com m ercial farm ing of tu rtles in the Caym an Islands, B ritish W est Indies. Because the green tu rtle is an endangered species tha t has been seriously over-exploited in the past, the idea was to establish a captive breeding colony on a self-sustaining basis tha t could supply the com m ercial m arket w ith eggs, soup, m eat and shells, thereby




taking the pressure off the rem aining wild population, w hich was being heavily poached. However, M ariculture soon encountered difficulties in getting the ir captive tu rtles to breed, and so in 1970 they approached Amo for help and advice. He was to become D eputy C hairm an of the Scientific and Research Division of M ariculture L td from 1970 to 1976, and rendered a great service to the organization. In particular, he helped to establish husbandry m ethods that ensured that m ating took place successfully in captivity, and he introduced routine procedures for exam ining the eggs during incubation to assess the extent of em bryonic m ortality.

A cting on A m o’s advice, M ariculture approached Sir Alan Parkes late in 1972 and invited him to collaborate w ith them in this enterprise. A lthough Amo and Parkes had been friends of long standing, a conflict arose betw een them over the financial m anagem ent of the turtle farm , and the possibility of establishing a Research Foundation for work on turtles, to be directed by Parkes; this deep disagreem ent was to term inate both the working relationship between these two m en and their friendship, and Amo resigned from the com pany in 1976.

Amo also developed an interest in manatees. T h is began in 1974 when the first m anatee workshop was held in Georgetown, British Guyana, at the instigation of D r Colin Bertram and his wife Kate, who had had a lifelong interest in these animals. Amo was an invited participant and his previous research interests in seals and in elephants made him ideally suited for work on these Sirenians, which are in evolutionary term s underw ater elephants. Amo was designated leader of a proposed new International C enter for Research on M anatees, but, alas, the venture was to come to nothing. Colin Bertram recounts how on one occasion he and Amo w ent to the K ennedy Space C enter at Cape Canaveral in Florida. As a result of the strict security surrounding the C enter, and the total exclusion of tourists and powerboats, the local manatee population had flourished and the animals had grown quite tame. Amo and Bertram were taken by a group of Am erican zoologists to the top of the obsolete Soyuz—Apollo launching gantry, at a height of over 100 metres, from w hich they obtained an excellent view of the manatees far below, one of which had a radio transm itter attached around its tail. T o his great delight, Amo found him self standing in an area fenced off w ith a chain and labelled ‘Only A stronauts beyond this po in t’.

Retirem ent brought w ith it num erous other opportunities for travel overseas. F rom N ovem ber 1968 to M arch 1969 Amo was the Royal Society Leverhulm e Visiting Professor in the D epartm ent of Veterinary Physiology in the U niversity of Chile at Santiago, and from Novem ber 1970 to M arch 1971 he was T . L. Pawlett Scholar in the D epartm ents of Veterinary Physiology and Animal Science at the University of Sydney, Australia. On his return to England in 1971 he was to have num erous health problem s. He had developed diabetes, which forced him to give up




his beloved spaghetti at B ertorelli’s, b u t m ore seriously he had developed a cataract in his right eye. T h is was operated on at St T h o m as’ H ospital, and he w ent to stay w ith the Principal of L ondon U niversity , Sir D ouglas Logan, to recover from the operation. All this, coupled w ith prostate problem s and the need for extensive dental treatm ent, m ade him quite depressed for a while.

In 1975, w ith his illnesses now behind him , he em barked once m ore on his w inter m igrations to w arm er climes, becom ing Royal Society Leverhulm e Visiting Professor in the D epartm en t of V eterinary Physiology at the U niversity of N airobi, Kenya, from N ovem ber 1975 to February 1976. H ere he was to have a great influence on the staff, and D r D om inic O dour-O kelo, D r C. S. Bam bra and Professor Sam son G om be speak w ith particular affection of the days he spent w ith them ; they were able to collaborate in some work on the placenta of the elephant shrew which was eventually published in 1980, and Professor G om be’s eldest son is christened Am oroso in his honour. Amo obviously enjoyed the visit because he was to re tu rn again the following year as Royal Society and In ter-U niversity Council V isiting Professor to the D epartm ent of Surgery, K enyatta H ospital, N airobi, from D ecem ber 1976 to F ebruary 1977. T he last bu t not least of his overseas visits were periods from F ebruary to June 1977, 1978 and 1982 spent as D istinguished V isiting Professor at the College of Biological Science, U niversity of G uelph, O ntario, Canada, where once again he became extrem ely popular w ith staff and students alike.

T h u s Amo can indeed claim to have been an am bassador of science to m any countries of the Com m onw ealth and beyond. Ironically, it was perhaps not until his retirem ent that his gifts as a teacher were m ost appreciated, as he came in contact w ith a wide variety of students from different backgrounds to w hom he had been denied access w ithin the narrow confines of a veterinary school.

Several national and scientific awards were also bestowed upon him . In 1969 H er M ajesty the Q ueen was graciously pleased to appoint him C om m ander of the O rder of the B ritish Em pire, and in 1977 he was awarded the T rin ity Cross of the Republic of T rin idad and Tobago for D istinguished Service. Amo used to recount how, when his plane had just landed at T rin idad airport, he was surprised to find a real red carpet laid out for him , and a cluster of reporters eagerly awaiting an interview. He announced that it was not his custom to give interviews to the press, bu t he could not escape the attention of one particularly persistent and attractive young lady, who begged to be able to ask him ju st one question. Relenting, Amo eventually consented to her request; she said ‘Professor Amoroso, do you re tu rn to T rin idad as a T rin idadian , or as a European?’ Seizing the m icrophone, he replied w ithout hesitation ‘Lady, I come here as an A fro-Saxon’. N ever has there been a m ore apposite sum m ary of Amo, the man.




In 1972 A m o was aw arded the M ary M arshall M edal of the Society for the S tudy of F ertility , its highest aw ard and a fitting tribu te to one who had m ade such a great con tribu tion to M arshall’s Physiology of reproduction. In 1977 he was also aw arded the L udw ig-S chunk Prize of the Ju stu s-L ieb ig U niversity , G iessen. An award th a t gave him particular delight was th a t of the D ale m edal of the Society for Endocrinology, m ade in his 80th year. T h e occasion was all the m ore poignant because Am o had been C hairm an of the Society w hen it had decided to strike a m edal to recognize the con tribu tions of Sir H enry Dale, O .M ., F .R .S ., to endocrinology, and Am o had gone to some lengths to m eet Sir H en ry ’s wishes as to how his likeness was to be graven; the die had to be modified to enable the bow tie to conceal Sir H enry ’s double chins m ore effectively.

T o celebrate his eightieth b irthday , A m o’s friends and colleagues from all a round the w orld organized a m eeting in his honour at the Royal Society in Septem ber, 1981, on ‘Placenta: struc tu re and function’, and it p rovided a fine opportun ity to review recent progress in all those fields in w hich A m oroso had had an in terest in the past. Sadly, he was to die only a few weeks before the full proceedings of this m eeting were published as supplem ent no. 31 of the Journal of Reproduction and Fertility in N ovem ber of the following year; the volum e w ould have given him such great delight. A fter the day’s scientific proceedings were concluded, Professor Peter Jewell, the M ary M arshall Professor of Reproductive Biology at the U niversity of C am bridge, organized a special dinner for Amo and all his friends and colleagues at B ertorelli’s restaurant in Soho, one of his favourite haunts in his younger days and a place in w hich he in his tim e had entertained m any of his students. It was a m em orable occasion, presided over by Sir Peter M edaw ar, F .R .S ., and Lady H uxley. I t was touching to see the endearm ent felt by the entire Bertorelli family and all the restaurant staff for their long-tim e friend and patron, a vivid testim ony of the fact that A m o’s personal charm extended far beyond the confines of the academic com m unity.

O n his eightieth b irthday, Amo was also to receive a birthday card from well-w ishers in U niversity College, D ublin, and his handw ritten letter of thanks to D r H ilda M acD onnell in the D epartm ent of A griculture, w ritten on 16 Septem ber 1981, tells us m uch about his thoughts and feelings at tha t tim e. I t reads as follows:

‘D ear H ilda,W hat a joy to receive your card and what nostalgic m em ories it brings back of the D ublin of yesterday.

I am m ost grateful to all those of my Alma M ater who have thought to rem em ber me w ith such affection.

I have never thought of my age num erically, bu t at 80 it has, for me, like Sum m er, the elem ent of sadness which colours all high




points in existence. Ju st as a rare m om ent of personal triu m p h is tinged by a daw ning sense of the barrenness of life suddenly deprived of a particu lar am bition , now fulfilled, so sum m er becom es too soon, alm ost at its very inception, a d rift in to au tum n, the passing of a prim e. A u tum n has its own beauties once it is accepted— in fact, it is often far lovelier than sum m er, b u t th is only once it is present, ju s t as an acceptance of old age can produce the m ost exquisite rew ards; b u t the first w hite hair sp rou ts in sum m er, and is given especial a tten tion because it is unseasonal. All I have tried to say, is thank you for your good wishes.

Sincerely,A m o.

But A m o’s activities d id no t cease at the age of 80. In 1982 he was to becom e the recipient of the Carl G . H artm an A w ard of the Society for the S tudy of R eproduction in the U nited States, its highest honour, in recognition of an outstand ing career of research and scholarly contribu tions in the field of reproductive biology, and in particu lar the role he played in stim ulating so m any individuals by his research, his publications and his teaching over m any active years. In A ugust 1982 he was elected an H onorary Fellow of the Royal A cadem y of M edicine in Ireland, and on his 81st b irthday , a few weeks before his death, he opened the F irst In ternational Sym posium on V eterinary A naesthesia, organized by the Association of V eterinary A naesthetists of G reat B ritain and Ireland, a society tha t owed m uch to the activities of two of his form er students, D r L. W . H all and D r B arbara W eaver.

H e died peacefully, at the hom e of his friends, Professor and M rs J. B. Lynch, in Leeds on Saturday, 30 O ctober 1982.

A m em orial service was held at St P e te r’s C hurch , Babraham , on 6 January 1983 for his close friends and colleagues, w ith tribu tes from Professor P. A. Jewell, the late Professor C. F. Cori, F o r.M em .R .S ., and D r B. A. Cross, F .R .S . Professor C ori, who was unable to be presen t in person, w rote the following w ords about his old friend:

‘Amo was involved in m any unusual events; he travelled widely and led a rich life. H ow ever, it was his gift for friendship , his w arm personality, his zest for life, his good com panionship and no t least his w isdom th a t endeared him to all. W hat he attained in life, seem ingly against insurm ountable odds, was entirely th rough his own effort. M any of his good deeds for o ther people are probably not known, since he was very reticent w hen it came to personal affairs. H e was a rare person who enriched our lives and con tribu ted im portan tly to science and helped o ther people, and w hom it is im possible to forget. T hose of us who were privileged to know him will always cherish him as one of the m ost rem arkable hum an beings.’




A m em orial m ass was held at the C hurch of S t A nselm and S t Cecilia, K ingsw ay, L ondon , on 28 F eb ruary 1983, w hen L ord Zuckerm an, O .M ., K .C .B ., F .R .S ., represen ting the D uke of E d inbu rgh as Patron of the Royal V eterinary College, delivered the M em orial A ddress.

S c i e n t i f i c c o n t r i b u t i o n s

A m o’s first publication (1), in 1931, dealt w ith the em bryonic developm ent of the pancreas in the chicken em bryo, and it was to be the only scientific com m unication th a t he published in G erm an. H is 1934 P h .D . thesis (2), on the developm ent of the M ullerian ducts in the rabbit, was never pub lished in a scientific journal; it was not un til 1947 tha t A lfred Jost, w orking Paris, discovered tha t the developm ent of the M ullerian ducts was under the control of an inhib itory horm one secreted locally by the foetal testis (Jost 1947 a,ban observation tha t depended on experim ental surgery, a technique in w hich Amo had no expertise.

In 1935 Am o w rote a b rief note to Nature (3) on the effects of colchicine on tu m o u r grow th. In later years he was to re tu rn once m ore to tu m o u r grow th; from 1962 to 1966, in collaboration w ith D r Stein- W erblow sky, he d id some w ork on the im m unological control of tum ours (78-80, 85, 86, 91, 96), and then in 1975, in collaboration w ith D r J. S. Perry (116), he w rote a m ajor speculative review of the m any ways in w hich the placenta m ight be able to defend itself from m aternal im m unological attack, e ither by the secretion of a protective m echanical b arrie r around the invading trophoblastic cells, or by the trophoblastic p roduction of a horm one such as chorionic gonadotrophin w hich itself m ight have im m unoprotective properties. T h is whole subject is still unresolved and rem ains a focus of cu rren t debate.

T h ree years were to pass before A m o’s next publication (4), a brief note to th e Journal of Anatomy in 1938 reporting the appearance of a recentlyovulated egg in the Fallopian tube of the goat. M ore extensive publications appeared in the following years (5 ,7), describing the cleavage stages of the fertilized goat’s egg as it traversed the Fallopian tube, and these really m ark the first papers tha t Amo was to produce in the field of reproduction. H e was clearly fascinated by early cleavage, and in collaboration w ith the Belfast anatom ist, W . J. H am ilton, he extended his observations to include the tubal ova of the horse and dog (6, 9). T h e next year was to be a productive one, and it also m arked Amo s first publication on the placenta (8), an account of the cotyledonary, syndesm o-chorial p lacentation of the blackbuck, an Indian species of antelope, in com parison w ith tha t of the m ore fam iliar cow, sheep and goat.

T h e move from London to Reading at the outbreak of war made it relatively easy for Amo to collaborate w ith the O xford group of foetal




physiologists A. E. Barclay, K. J. F ranklin and M . L . P ritchard , and from 1941 to 1943 he published in collaboration w ith them a nu m b er of descriptive papers (10—12, 15) on the lungs and cardiovascular system of a variety of m am m alian foetuses, including an A frican elephant. T h e elephant is of particu lar in terest in this regard because the heart is unusual in showing partial separation of the ventricles at the ir apices, and although developm ent of the pleural cavity is norm al in early foetal life, the parietal and visceral pleurae fuse before b irth so tha t the pleural cavity is com pletely ob literated for the rest of the an im al’s life (S hort 1962).

W hile at Reading Am o also developed a b rie f collaboration w ith D r S. J. Folley, F .R .S ., in experim ental lactational physiology (13), and perhaps this early in terest in lactation accounted for the in teresting paper he was later to publish w ith D r P. A. Jewell in 1963 (87) on the way in w hich prim itive peoples had exploited the m ilk-ejection reflex. In 1943 Amo published his only jo in t paper w ith A. S. Parkes, F .R .S . (16), who was then w orking at M ill H ill, on the effects of X -irrad iation on sperm atozoa, a field in w hich Parkes was particu larly active at tha t tim e.

F our years were to elapse before A m o’s next publication, in 1947, in collaboration w ith D r O. G. Edholm (17), describing the am azing array of valves in the jugu lar veins of those long-necked species, the giraffe, okapi, camel and ostrich, designed to p revent a sudden increase in hydrostatic pressure w hen the head is lowered for feeding or drinking. Clearly A m o’s developing connections w ith R egen t’s Park Zoo and the Prosectorial C om m ittee were responsible for this in teresting piece of work.

Perhaps.A m o’s first im portan t paper was th a t published w ith D r J. L. H ancock and D r I. W . Rowlands in Nature in 1948 (18), on ovarian activity in the pregnant m are. I t will be rem em bered tha t Am o was already 47 years old, at a tim e w hen m any people today are beginning to th ink tha t their active scientific careers are draw ing to a close as they dissipate their energies in ever-increasing adm inistrative duties. A m o’s late start, and great success, should therefore serve as a source of inspiration to us all.

As long ago as 1930 H . H . Cole had m ade the surprising discovery tha t at certain stages of gestation the serum of pregnan t m ares contains gonadotrophic activity (Cole & H art 1930), and this activity was later shown to be concentrated in, and presum ably produced by, ulcer-like structu res visible on the surface of the endom etrium , and known as endom etrial cups (Cole & Goss 1943). Amo was the first to show that du ring early gestation in the m are, w hen the gonadotrophic activity is greatest, the m are’s ovaries respond to this stim ulus in a unprecedented way, both by ovulation and by luteinization of u n ru p tu red follicles, in a m anner rem iniscent of the ovarian response of wom en following inadverten t hyperstim ulation w ith exogenous hum an gonadotrophins.




T h e endom etrial cups fascinated Amo for the rest of his life. T hey were first described by the G erm an anatom ist Schauder (1912); Amo went on to investigate their fine structure , and he described in detail the m orphological appearance of the unusual decidual cells, both in his 1952 chapter in M arshall (36), and in a British Medical Bulletin article in 1955 (46). He believed tha t these decidual cells, w hich we now know to be the source of equine chorionic gonadotrophin, were derived from plasm a cells:

‘W hen fully developed the cup cells are greatly enlarged and have rounded contours, granular cytoplasm and vesicular nuclei. Binucleate and m ultinucleate form s are frequently encountered. It is noticeable that transitional form s between small round cells, plasm a cells and typical decidual cells continue to appear even in the m ost m ature cups’ (46).

He was probably led into the m istaken belief that equine decidual cells were of m aternal origin by analogy with the decidual cells of rodents, w hich develop at the site of placental attachm ent through the hypertrophy of u terine strom al tissue. T h e elegant studies of D r W. R. Allen and his colleagues in Cam bridge on the ontogeny of endom etrial cup developm ent in a carefully tim ed series of pregnancies (Allen, H am ilton & M oor 1973) have established conclusively that the decidual cells of the equine endom etrial cup are of foetal origin. T hey are derived from the allantochorionic girdle of the developing conceptus during days 36-38 of gestation, as foreshadowed in the classical account by Professor J. Cossor Ew art, F .R .S ., in 1897 of the form ation of this girdle. Amo failed to recognize the ‘small round cells’ as invading m aternal lymphocytes, w hich penetrate the endom etrial cup in great num bers, presum ably in response to the expression of foreign antigens on the invading foetal cells. T h e subsequent destruction and rejection of the cup tissue bears all the hallm arks of a graft rejection reaction (Allen 1982), thus making it an outstanding and challenging exception to the general rule that the potential antigenic nature of the foetal placenta does not initiate a m aternal im m une response. It is both ironical and sad that Amo, with his great interest in this subject, and w ith the evidence before his eyes, should have failed to recognize the endom etrial cup for what it really was: a trophoblastic invasion of the endom etrium , destroyed by a maternal im m une response. H ad he been more of an experim entalist he would have been able to study the sequence of events leading up to the establishm ent of the endom etrial cups, and he would not have been led into the error of assum ing that lymphocytes and plasma cells were direct precursors of equine decidual cells.

M any other aspects of equine placentation continued to fascinate Amo throughout his career. In 1950 he wrote his first paper (23) on the secretion of ‘uterine m ilk’ in the mare. This was such a beguiling phrase; it was easy to imagine that ju st as the m am m ary gland secreted milk for




the neonate, so too m igh t the u terine glands secrete a m ilk for nourish ing the un im plan ted blastocyst. B ut A m o had initially used the te rm to describe the thick, viscous coagulum lying on the surface of the en d o m etrial cups as they begin to degenerate, a secretion th a t is only found in the Equidae, and has no co u n terp art in any o ther genus. B iochem ists searched in vain for som e un ifo rm chem ical constituen ts th a t m igh t characterize this ‘u terine m ilk ’ of m am m als, and a lthough in teresting proteins like ‘b lastok in in ’ or ‘u te rog lob in ’ w ere isolated, these were later shown to be species-specific. T h u s the te rm ‘u te rine m ilk ’ has gradually fallen into disuse for lack of an adequate definition of w hat is m eant by it.

H ow ever, there can be no doub t tha t the biochem ical com position of the u terine secretions holds the key to the spectacular arrest of em bryonic developm ent at the b lastocyst stage th a t is so characteristic of species like the roe deer, the seals, the bears, m any m ustelids, and m ost kangaroos and wallabies. I t was therefore en tirely appropria te th a t in 1963, Am o should be called upon to help organize and to sum m arize (84) the proceedings of the first in ternational conference on th is in trigu ing topic of delayed im plantation , held at Rice U niversity , H ouston , Texas.

A m o’s in terest in delayed im plantation , and his fascination w ith the historical aspects of reproductive biology, led h im astray on one point. In 1959 he w rote a review (69) of the h istory o f ovum im plan tation in collaboration w ith D r P. Eckstein and D r M . C. Shelesnyak, and he claim ed tha t W illiam H arvey had been the first to discover delayed im plantation in 1651 as a result of his studies of roe deer ( capreolus).

In W illiam H arvey’s classical w ork De generatione , p u b lished in L atin in 1651 and rep rin ted in English two years later, he had set ou t to determ ine how life began— w hether A risto tle was correct in his assertion tha t the em bryo developed from an egg form ed by coagulation of the ‘seed’ of the m ale, or w hether some o ther m echanism m ight be at work. H e therefore chose to dissect the deer being h un ted by his P atron , K ing C harles, on the reasonable assum ption th a t killing females at the tim e of the ru t w ould reveal the anatom ical na tu re of the early conceptus. H arvey makes it abundantly clear tha t the K ing was hun tin g principally red deer, ‘C ervus’ in the 1651 text ( Cerelaphus), and also some fallow deer or ‘D am a’ ( Damadama). Red deer ru t in Sep tem ber—O ctober, bu t H arvey could no t recognize any products of conception in the u te rus un til m id-N ovem ber, forcing h im to conclude, against his b e tte r judgem ent, tha t the act of copulation had no th ing w hatsoever to do w ith conception. W e now know that this was because red deer only m ate tow ards the end of the ru t in m id-O ctober, and the em bryo does no t becom e clearly recognizable to the naked eye un til about 2 weeks later, w hen it appears as a th in ribbon of tissue, m any centim etres in length. H arvey saw th is struc tu re , b u t it was so unlike the A ristotelean egg he expected to find tha t he dism issed it as m erely ‘p u ru len t m a tte r’, and it was no t un til m id-




N ovem ber th a t he could identify an em bryo w ith its pulsating heart ( punctum saliens) w ith in the am niotic vesicle (S hort 1977, 1978).

A m o and his colleagues, unaw are of these finer details of the rep ro ductive physiology of red deer, and using an inaccurate version of De generatione animalium pub lished in English by W illis in 1847 in w hich the ‘D am a’ of the original had been m is-translated as ‘ro e ’ instead of ‘doe’, erroneously concluded th a t H arvey m ust have failed to see the em bryo in O ctober and early N ovem ber because he was studying roe deer. In roe deer, ru ttin g and m ating take place in late July — early A ugust, followed by five m onths of delayed im plan tation so tha t the em bryo in the u terus only becom es visible to the naked eye at about C hristm as tim e. T h e phenom enon was, in fact, first described by the G erm an anatom ists Z iegler (1843) and Bischoff (1854).

P erhaps A m o’s m ost creative period was in the early 1950s. Between the years 1951 and 1954, he published a succession of papers in collaboration w ith F. R. Bell (24—28, 31, 43) on the central nervous contro l of vasom otor activity and respiration; these represent his only foray into the realm s of classical neurophysiology, and from th a t date his pub lications were concerned alm ost exclusively w ith reproduction . Also in 1951 he started w ork w ith D r L. H arrison M atthew s, F .R .S ., D r R. J. H arrison , F .R .S ., and D r I. W . Row lands (35) on rep roduction in the grey seal. A m oroso and Row lands, like m any before them , had been in trigued by the spectacular hypertrophy of the gonads of the foetal foal, a phenom enon that is still incom pletely understood, and they were therefore in trigued to find a degree of gonadal hypertrophy in the full- te rm seal foetus. In 1952 Am o was to publish im portan t papers (37-39) describ ing for the first tim e the placental m orphology of the seal, and docum enting the spectacular grow th rate of the new born seal pup as it ingests one of the m ost nu tritious m ilks found in the anim al kingdom .

T h is spate of research activity in the early 1950s coincided w ith the w riting of his fam ous 184-page review of p lacentation in Marshall’s Physiology of reproduction (36). N o t only did it have a bibliography of 438 references, b u t it is notew orthy that no t a single one of these was to A m o’s own grow ing list of publications— a rem arkable tribu te to his intellectual hum ility . Am o reviewed the whole history of the evolution of viviparity in elasm obranch and teleost fishes, am phibians and reptiles, before m oving on to discuss the various classifications of placental m orphology in m am m als, followed by detailed exam ples of the various placental types gleaned from a wide range of m onotrem e, m arsupial and Eutherian m am m als. H e also used the chapter as a m edium for publishing m any of his own original observations, and it is beautifully illustrated w ith his line draw ings, photom icrographs and gross dissections. I t was a m asterly review and synthesis of all tha t had gone before, and nothing has been w ritten since to supercede it. T h e chapter was also to provide a fram ew ork for all A m o’s fu ture research interests.




T h e reasons underly ing the spectacular diversity in placental s truc tu re betw een the species still escape our com prehension. F o r those who adhere to the belief tha t ontogeny repeats phylogeny, it m ust still come as som ething of a shock to realize tha t those so-called ‘p rim itive’ m am m als, the A ustralian m arsupial bandicoots, w hich hold the record for the shortest m am m alian gestation length, nevertheless have a ‘highly evolved’ chorioallantoic placenta, as first poin ted out by Professor J. P. H ill at the tu rn of the century. For those who like to believe tha t s truc tu re reflects function, the placenta presents a particu lar challenge because it seems im possible to relate the one to the other. T hese were the problem s w ith which Amo was to grapple for the rest of his scientific career; the placenta will continue to rem ain an enigm a to biologists until we can begin to understand the reasons for this high degree of placental diversity.

In later years Amo was to collaborate w ith a num ber of scientists including G. B. W islocki and J. S. Perry in provid ing definitive accounts of placentation in the wolverine, the hippopotam us, the spotted hyaena and the African elephant (52, 61, 90, 93). In 1958 he also collaborated w ith D r L orna Kellas (63) in an in triguing study of the ovaries of foetal and prepuberta l giraffes, showing that large G raafian follicles and num erous corpora lutea are present in the foetal ovaries at the later stages of gestation, presum ably in response to the placental production of a chorionic gonadotrophin. Amo also noted in passing in his 1955 British Medical Bulletin article (46) that the nilgai, the largest of the Indian antelopes, apparently accum ulates accessory corpora lutea in its ovaries during gestation, som ewhat rem iniscent of the situation in the pregnant mare.

Amo was also to develop a passing in terest in the environm ental control of seasonal breeding in b irds and m am m als, a phenom enon that is almost universal for species living in the tem perate and arctic regions of the world. In 1960, in collaboration w ith F. H . A. M arshall, he w rote a m ajor 93-page review of this subject for Marshall’s Physiology of reproduction (74). U nfortunately , this was ju s t before the key role of the pineal gland in m ediating the effects of changing photoperiod had been discovered, so their review was to have little im pact on a rapidly advancing field, and it is seldom referred to in cu rren t publications.

Amo was one of the few people to appreciate the intrinsic in terest of studying reproduction in the cat. I t is curious that the cat should have played such a central role in the developm ent of neurophysiology, and yet to this day we probably know less about reproduction in this species than in any other dom esticated animal. In collaboration w ith C. R. A ustin, then working at M ill H ill, Amo was able to show in 1957 (59) that the recently discovered sex chrom atin, first identified as an in tranuclear inclusion in the nerve cells of female cats by M urray Barr, could be used to determ ine the sex of early cat em bryos. T h is im portan t observation




was to pave the way for others who showed that it was possible to sex the hum an foetus inutero by looking for sex chrom atin in desquam ated epithelial cells obtained by amniocentesis.

In 1962 Amo was to write an im portant review in collaboration with D r C. A. F inn (77) on ovarian activity during gestation that was published in Solly Zuckerm an’s book, The ovary. Fifteen years later, Amo and J. S. Perry re-w rote and updated the article for the second edition of the book (120). These two reviews brought together all the inform ation, widely scattered throughout the scientific literature, on the effects of hypophysectom y of the m other during gestation. W hen this is pu t together with inform ation on the consequences of ovariectomy on the course of pregnancy it provides vital clues about the likely time of onset of progesterone production by the placenta, and w hether or not the placenta is capable of providing luteotrophic support for the corpus luteum. A lthough Amo him self did no research in this field, it was a subject that had always aroused his intellectual curiosity, and hence his reviews of the topic are both rewarding and thought-provoking to read. It was natural that Professor Geoffrey H arris, F .R .S ., and D r B. T . Donovan should tu rn to Amo and his form er student D r D. G. Porter to write a major review (99) of anterior pituitary function in pregnancy for their three- volume work The pituitary gland, published in 1966.

A m o’s retirem ent in 1968 did not call a halt to his scientific publications. In addition to writing an extensive biography (111) of H. M. Evans, For.M em .R .S ., published in 1972, he also wrote an account of the life of his old friend and colleague, G. B. Wislocki, for the Dictionary of American biography in 1980 (123). He was co-author of two im portant publications with his new colleagues at the Institute of Animal Physiology in Babraham; in 1972, D r F. A. Harrison, D r R. B. Heap, D r N. L. Poyser and Amo published a brief bu t none the less im portant note on the production of the luteolytic horm one prostaglandin F 2a by the uterus of the sheep (112). In the following year, w ith D r J. S. Perry and D r R. B. Heap, Amo reported in Nature the im portant finding that the unim planted pig blastocyst is capable of producing significant amounts of oestrogen (114). In 1976 Amo co-authored a major paper on the histology of the ovary of the green turtle (117), the result of all his efforts on the turtle farm in the Cayman Islands, and in 1978 he wrote a report summarizing the work that had been done on the manatee (121). But it was to his first love, the evolution of viviparity, that he was to return at the end of his days. He collaborated with D r R. B. Heap and D r M arilyn Renfree in their extensive review of the role of hormones in the evolution of viviparity (122), published in 1979, and almost his last paper (124), in 1981, was to be a review of viviparity in the preface to a book describing the exciting new discoveries that are now being made in the cellular and molecular biology of that ever-mysterious process, implantation.

So ended the scientific career of a truly remarkable man who was able




to tu rn w hat at one tim e had seem ed the insurm ountable obstacles of race and colour to his advantage; it was his determ ination to succeed, and the force of his intellect, tha t enabled him to becom e a m uch-loved in te rnational am bassador of science, an A fro-Saxon beyond com pare.

A c k n o w l e d g e m e n t s

I should like to express m y gratitude to the m any people who have helped me in the preparation of this m em oir, especially M rs Kay Jordan, Professor C. Form ston, Professor P. A. Jewell, D r C. B ertram , D r R. B. H eap and D r J. S. Perry.

T h e photograph, taken in 1981, is reproduced by courtesy of D r R. B. Heap of the Institu te of Anim al Physiology, Babraham .

(1) 1931

(2) 1934

(3) 1935(4) 1938(5) 1939(6)

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(9) 1940(10) 1941

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S e l e c t e d b i b l i o g r a p h y

Die Z uchtung von Epithel des em bryonalen H uhnerpankreas. Z. Zellforsch. mikrosk. A n a t. 12, 274-300.

O bservations on the developm ent of the urogenital system of the rabbit, w ith special reference to the developm ent of the M ullerian ducts. P h .D . thesis, U niversity of London.

Colchicine and tum our grow th. N ature, Lond . 135, 266-267.T h e first finding of the tubal ovum in the goat. J . A nat. 73, 359P.T ubal journey and rate of cleavage of the goat’s ovum , J . A n a t. 73, 672-674,(W ith W . F. B. G riffiths & W. J. H amilton) A living tubal ovum of the horse. Vet.

Rec. 51, 168.(W ith W. F. B. G riffiths & W. J. H amilton) Living tubal ova of the goat. Vet. Rec.

51, 1009-1010.(W ith W. F. B. G riffiths) T he placentation of the Blackbuck (Antelope cervicapra)

w ith rem arks on some features of the placentation of the cow, sheep and goat. J . A nat. 73, 675-676.

T ubal transit and rate of cleavage of the dog’s ovum. J . A na t. 74, 533.(W ith A. E. Barclay, K. J. Franklin & M .L , Prichard) O bservations on the

cardiovascular system and lungs of an African elephant foe tu s.^ . A nat. 76, 100-111.(W ith A. E. Barclay, K. J. F ranklin & M . L. Prichard) T h e valve of the foram en

ovale and its relations to pulm onary vein entries. Vet. J . 98, 29—41.(W ith A. E, Barclay, K. J. Franklin & M . L. Prichard) T h e bifurcation of the

posterior caval channel in the eutherian foetal heart. J . A nat. 76, 240-247.(W ith S. J. Folley & H. M . Scott-W atson) F u rth e r experim ents on lactation in

thyroidectom ized rats. T h e role of the parathyroids. J . Endocr. 3, 178-191.(W ith W. F. B. G riffiths & W. J. H amilton) T he early developm ent of the goat

{Capra hircus). y . A nat. 76, 377-406.(W ith A. E. Barclay, K. J. F ranklin & M . L. Pritchard) Incidence of bilateral

anterior vena cavae in a series of eutherian foetuses. Proc. zool. Soc . Lond. 113, 43-53.

(W ith A. S. Parkes) X -irradiation of rabbit sperm atozoa in vitro. Sem en collection, artificial insem ination and induction of super-ovulation. N ature , Lond. 153, 244-246.

(W ith O. G. Edholm & R. E. Rewell) Venous valves in the giraffe, okapi, camel and ostrich. Proc. zool. Soc. Lond. 117, 435-440.

(W ith J. L. H ancock & I. W. Rowlands) Ovarian activity in the pregnant mare. N ature, Lond. 161, 355-357.

(W ith F. R. Bell, A. S. K ing & H. Rosenberg) C arotid nerve reflexes and effects of adrenaline in the horse and mule, y. Physiol., Lond. 109, 29P.




(20) 1949

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(36) 1952

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(40)(41) 1953

(42) 1954(43)

(44) 1955

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(46)(47)

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(49)(50)

(51) 1956

(W ith C. B. M urray) Im plantation and the yolk-sac placenta in the cat. J . A nat. 84, 71-72.

T h e physiology of reproduction in the dog. Vet. Rec. 61, 777-779.Placental horm ones and viviparity. Proc. Linn. Soc. Lond. 161, 7—8.Source and com position of uterine milk in the mare. J . A nat. 84, 78.(W ith F. R. Bell & H. Rosenberg) T he localisation of respiratory regions in the ovine

rhom bencephalon.^* Physiol., Lond. 113, 2P.(W ith J. G . Bainbridge, F. R. Bell, A. S. K ing , A. M . Lawn & H. Rosenberg)

O bservations on circulation and respiration in the badger and lion. J . Physiol., Lond. 113, 2P.

(W ith J. G. Bainbridge, F. R. Bell, A. M. L awn & H. Rosenberg) Central respiratory spike potentials. N ature, Lond. 167, 603-604.

(W ith F. R. Bell & H. Rosenberg) T he localisation of respiratory regions in rhom bencephalon of the sheep. Proc. R . Soc. Lond. B 139, 128—140.

(W ith F. R. Bell, A. M . L awn & H. Rosenberg) Somatic m otor areas of the cerebral cortex of the goat. Vet. Rec. 63, 445.

T h e essential parenchym a of the p ig ’s placenta. J . A nat. 85, 409.(W ith A. G offin, G. H alley, D. J. M atthews & L. H. M atthews) Lactation in grey

seal. J . Physiol., Lond. 113, 4P.(W ith F. R. Bell, A. S. K ing & H. Rosenberg) T he aortic and sinus nerves of the lion

and badger. J . A nat. 85, 411.Interactions of the trophoblast and endom etrium in the ungulata at the time of

im plantation. Proc. zool. Soc. Lond. 121, 201.T he interaction of the trophoblast and endom etrium in the sheep. J . A nat. 85,

428-429.(W ith I. W. Rowlands) H orm onal effects in the pregnant mare and the foetal foal. J .

Endocr. 7, 1-liii.(W ith R. J. H arrison, L. H. M atthews & I. W. Rowlands) Reproductive organs of

near term and new born seals. Nature, Lond. 168, 771-775.Placentation. In M arshall's Physiology of reproduction 3rd edition (ed. A. S. Parkes),

vol. 2, pp. 127-309. London: Longm ans Green.(W ith L. H. M atthews) R eproduction and lactation in the seal. Proc. 2nd In t. Congr.

Anim . Reprod., Copenhagen 1, 193-203.(W ith L. H. M atthews) T he grow th of the G rey Seal (Halichoerus grypus F ) from

b irth to weaning. J . A nat. 85, 427-428.(W ith R. J. H arrison & D. J. M atthews) T he foetal placenta and m em branes of

Halichoerus grypus. J . A nat. 86 , 487—488.A llanto-chorionic differentiations in the Carnivora. J . A nat. 86 , 481-482.(W ith L. M . Kellas & L. H. M atthews) T he foetal m em branes of a waterbuck:

Kobus defassa. Proc. zool. Soc. Lond. 123, 477.U nusual features in the placenta of some carnivores. Proc. zool. Soc. Lond. 124, 916. (W ith F. R. Bell & H. Rosenberg) T he relationship of the vasomotor and respiratory

regions in the m edulla oblongata of the sheep. J . Physiol., Lond. 126, 86—95.T he comparative anatom y and histology of the placental barrier. In Gestation (ed.

Louis B. Flexner), pp. 119-224. Princeton: Josiah Macy Jr Foundation Publications.

(W ith L. H. M atthews) T he effect of external stimuli on the breeding cycle of birds and mammals. Br. med. Bull., 11, 87-92.

Endocrinology of pregnancy. Br. med. Bull. 11, 117-125.(W ith E. W. D empsey & G. W islocki) Electron microscopy of the p ig’s placenta, with

special reference to the cell m em branes of the endom etrium and chorion. Am . J . A nat. 96, 65-79.

(W ith G. S. D awes, J. C. M ott & B. R. Rennick) Occlusion of the ductus venosus in the m ature foetal lamb. J . Physiol., Lond. 129, 64P.

H orm one control of the oestrous cycle. Vet. Rec. 67, 1072-1084.De la signification du placenta dans revolution de la gestation chez les animaux

vivipares. Annls Endocr. 16, 435—447.T he endocrine environm ent of the foetus. Proc. 3rd Int. Congr. Anim . Reprod.,

Cambridge 1, 1-8.




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1957

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(61) 1958

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(65) 1959

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(70)(71)

1960

(72)(73)

(74)

(75)(76)

1961

(77) 1962

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(81) 1963

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(W ith G. B. W islocki) T h e placenta of the W olverine {Gulo gulo lusucs (Linnaeus) ) , Bull. M us. comp. Zool. H arv. 114, 91-101.

(W ith P. C. W illiams) C ortisone requirem ent of p regnant guinea pigs. Proc. 3rd In t. Congr. Anim . Reprod., Cambridge 2, 56—57.

R eproductive phenom ena in Gastrotheca marsupiata. R . Soc. Conversazione (23 May).R eproductive periodicity in the fem ale m am m al. Conspectus Endocr. 2(1), 19—23.T h e influence of environm ental factors on sexual periodicity . Conspectus Endocr. 2(2),

19-21.Effects of stim uli associated w ith courtship . Conspectus Endocr. 2(3), 19-28.N euro-hum oral factors in the control of the oestrous cycle. N E ast. M ink Fmr 50,

5-10.(W ith C. R. Austin) Sex chrom atin in early cat em bryos. E xp l Cell Res. 13, 419—421.(W ith R. M . L oosemore, C. R imington & B. E. T ooth) C ongenital porphyria in

bovines: first living cases in B ritain. N ature , Lond. 180, 230-231.(W ith N . A. H ancock & L. M . K ellas) T h e foetal m em branes and placenta of the

hippopotam us (Hippopotamus amphibius (L innaeus)). Proc. zool. Soc. Lond. 130, 437-445.

(W ith L. H. M atthews) O bservations on placental struc tu re in some ungulates. M am m alia 22, 175-185.

(W ith L. M . K ellas & E. W . van L ennep) O varies of some foetal and p repubertal giraffes (Giraffa Camelopardalis (L innaeus)). N a ture , Lond. 181, 487-488.

(W ith G. S. D awes & J. C. M ott) Patency of the ductus arteriosus of the new born calf and foal. Br. H eart J . 20, 92—96.

T h e biology of the placenta. In Gestation (ed. C. A. Villee), pp. 1-76. Princeton: Josiah M acy J r Foundation Publications.

C om parative anatom y of the placenta. A nn. N .Y . Acad. Sci. I S , 855-872.(W ith C. R. A ustin) T h e m am m alian egg. Endeavour 18, 130—143.(W ith J. D avies & E. W. D empsey). T he subplacenta of the guinea-pig. A na t. Rec.

133, 266.(W ith P. Eckstein & M . C. Shelesnyak) A survey of the physiology of ovum

im plantation in m am m als. M em . Soc. Endocr. 6, 3-12.Sex differentiation and developm ent. M em. Soc. Endocr. 7, 187-191.C om parative aspects of the horm onal functions of the placenta. In The placenta and

fe ta l membranes (ed. C. A. Villee), pp. 3-28. Baltim ore, M aryland: W illiam s & W ilkins.

V iviparity in fishes. Sym p. zool. Soc. Lond. 1, 153-181.(W ith J. S. E. D avid) Les causes de la m ortalite precoce des em bryons. In Les

fonctions de nidation uterine et leurs troubles, pp. 139-152. Paris: M asson et Cie.(W ith F. H. A. M arshall) External factors in sexual periodicity. In M arshall's

Physiology o f reproduction, 3rd edition (ed. A. S. Parkes), vol. 1, part 2, pp. 707-800. L ondon: Longm ans G reen.

Histology of the placenta. Br. med. Bull. 17, 81-90.(W ith J. D avies & E. W. D empsey) T he subplacenta of the guinea-pig: an electron

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Rep. Br. Emp. Cancer Campn , pp. 593-594.Irreparable damage of the reproductive organs. In Radiation effects in physics,

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1963 Sum m ary of conference. In Delayed implantation (ed. A. C. Enders), pp. 305-309.Chicago: U niversity of Chicago Press.

(W ith R. Stein-W erblowsky & R. W ynn) G row th of transplantable cancer in rats previously injected with placental tissue. N ature, Lond. 197, 606-607.

(W ith R. Stein-W erblowsky) A tum our-inhibitory fraction extracted from the post- coital fluid of the rat. In 41st Ann. Rep. B y . Emp. Cancer Campn , pp. 608-609.

(W ith P. A. Jewell) T he exploitation of the m ilk-ejection reflex by prim itive peoples. Occ. Pap. R. Anthrop. Inst. 18, 126-137.

1964 (W ith P. Scott & K. G. W illiams) T he pattern of external respiration in theunanaesthetized animal. Proc. R . Soc. Lond. B. 159, 325-347.

(W ith M . L. Blaxter, A. D. Chiquoine & R. H. N isbet) T he fine structure of neurons and other elem ents in the nervous system of the G iant African Land Snail (Archachatina m arginata). Proc. R . Soc. Lond. B 160, 167-180.

(W ith J. S. P erry) T he foetal m em branes and placenta of the African elephant (Loxodonta africana), Phil. Trans. R. Soc. Lond. B. 248, 1-34.

(W ith R. Stein-W erblowsky) A tum our-inhibitory fraction extracted from boar semen. In 42nd Ann. Rep. Br. Em. Cancer Campn, pp. 597-599.

(W ith A, N. Contopoulos, M . R yan & C. R. Contopoulos) Transplacental passage of thyroxine and iodine 131 in the sow, ewe, ferret, guinea-pig and rat. Proc. zool. Soc. Lond. 142, 699.

(W ith R. M , W ynn) Placentation in the spotted hyaena (Crocuta crocuta Erxleben), w ith particular reference to the circulation. A m . J . A nat. 115, 327-337.

1965 C orpus luteum in relation to pregnancy and evolution. In Proc. 2nd Int. Congr.Endocr.y In t. Congr. Series 83, pp. 665-668. Amsterdam: Excerpta Medica.

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1966 (W ith D. A, N ixon & A. St G. H uggett) Fructose as a com ponent of foetal blood andfoetal fluids of the bush baby (Galago senegalensis senegalensis). N ature, Lond. 209, 300-301.

(W ith D. G. Porter) A nterior pituitary function in pregnancy. In The pituitary gland (ed. G. W. H arris & B. T . Donovan), vol. 2, pp. 364-411. London: Butterworth.

(W ith R. Stein-W erblowsky) T um our immunology. In 44th Ann. Rep. Br. Emp. Cancer Campn , pp. 363-364.

(W ith R. Stein-W erblowsky) Immunological inhibition of induced rat tum our. In 46th Ann. Rep. Br. Emp. Cancer Campn, pp. 396-397.

1967 T he sym posium in perspective. In Reproduction in the female mammal (ed. G. E.Lam m ing & E. C. Amoroso), pp. 551-557. London: Butterworth.

(W ith R. A. M orton) Protein utilization by poultry. Edinburgh: Oliver & Boyd.1968 T he evolution of viviparity. Proc. R. Soc. Med. 61, 1187—1200.

Reproductive physiology: from a distinguished past to a prom ising future. J . Anim. Sc i, 27, Suppl, 1, 214-222.

1969 Physiological mechanisms in rep roduction .^ . Reprod. Fert. Suppl. 6 , 5-18.(W ith A. M. Lawn & A. D. Chiquoine) T he developm ent of the placenta in the sheep

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(W ith D. G. P orter) T he endocrine functions of the placenta. In Scientific foundations of obstetrics and gynaecology (ed. E. E. Phillip, J. Barnes & M. Newton), pp. 556-586. London: Heinemann.

1971 T he J. Y. Simpson Oration. Early theories of the placenta: from fancies to facts. In42nd Ann. Rep. R . Coll. Obstet. Gynaec., pp. 75-91.

1972 H erbert M cLean Evans (1882-1971) Biogr. Mem. Fell. R. Soc. 18, 83-186.(W ith F. A. H arrison, R. B. Heap & N. L. Poyser) T he production of prostaglandin

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(W ith J. S. Perry) T h e existence during gestation of an im m unological buffer zone at the interface betw een m aternal and foetal tissues. Phil. Trans. R . Soc. Lond. B 271, 343-361.

(W ith R. N. C. A itken & S. E. Solomon) O bservations on the histology of the ovary of the Costa Rican G reen T u rtle , Chelonia mydas L . J . exp. mar. Biol. Ecol. 24, 189-204.

(W ith D. H . Steven) A ndreas Vesalius and the zonary hum an placenta: egregious error or unfortunate circum stance? J . A na t. 122, 738.

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(W ith R. B. H eap & M . B. Renfree) H orm ones and the evolution of viviparity. In Hormones and evolution (ed. E. J. M . B arrington), vol. 2, pp. 925—989. L ondon: A cademic Press.

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