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Harold Munro Fox, 1889-1967

J. E. Smith

1968, 206-222, published 1 November141968 Biogr. Mems Fell. R. Soc.

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HAROLD MUNRO FOX

1889-1967

Elected F.R.S. 1937

H arold M unro Fox, Emeritus Professor in the University of London, died on the morning of 29 January 1967 at the age of seventy-seven of a coronary occlusion while enjoying his customary week-end ride on his horse. He was, at the time of his death and after thirteen years of retirement, daily engaged on his researches on the biology and systematics of ostracod Crustacea, busy in the editing of the Biological Reviews of the Cambridge Philosophical Society, and by his own wish active as librarian and unofficial tutor to students in the Department of Zoology of Queen Mary College, London. A great traveller and an accomplished linguist—it is said that he had a good working knowledge of six languages and was fluent in four of them—he achieved through the excellence of his researches and his ease of communication with scientists of many nationalities a truly international reputation as a zoologist which was matched by his own remarkable knowledge of contemporary developments in zoology.

Fox (at birth, Harold Munro Fuchs) was born in Clapham on 28 September 1889. His father, Georg Gotthilf Fuchs, had at one time been a Captain in the Prussian Army. His mother, Margaret Isabella Campbell Munro, was the daughter of Lt.-Colonel Andrew Munro who came of a Sutherland family and held a commission in the 19th (Yorkshire) Regiment. When Fox was only a few years old his parents separated, and though for a while they were re-united the marriage ended in a permanent separation. In consequence, Harold and his only (younger) sister Alison, now as Mrs Alison Settle well known as a distinguished fashion writer, a former editor of Vogue and Fellow of the Royal Society of Arts, were left in the care of their mother.

On reaching school age, Fox was sent to a preparatory school in Henley. During his holidays, spent mostly with his mother and sister in London, he would often make off on his bicycle to one or other of the main line stations to make notes on the many different kinds of engines and types of rolling- stock belonging to the then independent and distinctive regional railways. On one of these excursions, when he was ten or eleven years old he was thrown from his bicycle and run over by a horse-bus near Hyde Park. He suffered severe injuries to his legs and was for almost a year a patient

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208in St George’s Hospital. The accident left him with no permanent disability except for a slight shortening of one leg, which, however, was corrected by hospital treatment. About this time he was also operated on for the removal of tubercular glands. His doctors therefore advised that he should go to a day-school where he could convalesce under home care. So the family moved to his grandmother’s house in Brighton and, at the age of thirteen, Fox became a day-boy at Brighton College. At that time his main school interest was in mathematics in which he soon won a scholarship.He was a good shot and represented his school at Bisley; but he disliked most games and had a particular loathing for cricket which he managed to evade on account of his injured leg, though it is recorded that the disability did not prevent him from playing football, which he liked. Being a dayboy and freed from the chores of cricket he was able to get into the countryside at week-ends and on summer evenings to watch birds, collect wild flowers and to search the chalk cliffs for fossils; and with an increasing enthusiasm he began to seek out and to identify the plants and animals living on the rocks and in the tide pools of the sea-shore and in the fresh water ponds and streams on the outskirts of Brighton. Many carefully-selected specimens taken on these field excursions were brought back to be given a permanent home in the meticulously tended marine and fresh water aquaria which to the end of his life he counted among the most treasured of his household possessions. He seems also to have spent much of his time in his later years at school and, unbeknown to his teachers, in the Brighton library reading books on natural science, and in particular on biology and geology.His reading must have been of unusual width for a schoolboy, for when it was decided that he should try for a mathematics scholarship in Cambridge, and being without advice as to which College to apply, he chose Gonville and Caius because (as he says in his biographical notes) ‘of my interest in Dr Caius’. After sitting the papers in mathematics his mother was surprised to get a message saying that he was detained in Cambridge and would not be back for another two or three days. On returning home he explained that he had decided to stay on in order to sit the natural science papers. When the news came through that he had been awarded a scholarship in the natural sciences, in which he had no sort of formal training, his headmaster was convinced that a mistake had been made in the notification. When assured that there was indeed no error and that Fox was among the best of th e : candidates examined, the school authorities were so much put out by th e : : : unexpected turn of events that they were at first disinclined to award h im . . . the school scholarship of £60 which would have been attached to a mathe- • • • matics award. The scholarship was not in the end denied him, and in 1908 1 i i Fox entered Caius as a Senior Scholar to read zoology, botany, geology and . . . physiology for the Part I Tripos. In after years Fox thought that he had been i i till-advised to choose botany rather than chemistry for, as he says, ‘botany.......could have been got up as a hobby, and I have had the much more difficult; : : task of getting up chemistry from the small beginnings at school’. However,, , ,

Biographical Memoirs



he got a ‘first’ in Part I and went on to read zoology for Part II in which he took a ‘second’.

Fox graduated in 1911 and in the summer went to the Plymouth Laboratory to work for almost a year as junior partner with Gresswell Shearer and Walter de Morgan on the genetics of hybrid species of sea urchins reared from artificially fertilized eggs. Shearer, a Canadian by birth, had studied zoology at Johns Hopkins University and had later graduated in medicine at McGill where, at that time, E. W. MacBride was Professor of Zoology and in the full flow of his researches on the embryology of echinoderms. Shearer joined MacBride’s team and, with his encouragement, spent a good deal of time at Naples where Boveri, Driesch and Herbst—all of them then in their forties—were founding the science of experimental embryology with their experiments on the development of isolated blastomeres of urchin eggs and the effects of various ions on the course of a development. In 1910, J. Stanley Gardiner, newly appointed as Professor of Zoology in Cambridge, invited Shearer to give the first course of experimental embryology in the department. Fox attended these lectures and joined Shearer’s class at Plymouth in the summer vacation. After a year in Plymouth in 1911-1912 he spent ten months in Naples where ‘at first despondent for lack of guidance, I found my feet in work on fertilization’. In 1913 he was appointed by MacBride as lecturer in his department at Royal College of Science, London, and was there for a year until the outbreak of war when he enlisted in the Army Service Corps with an attachment to the London Mounted Brigade of Yeomanry. He served in the Balkans, Egypt and Salonica and during the last year of the war was attached to the French and Italian detachments in Palestine because of his fluency in their languages. When he joined the army he felt obliged to translate his name of Fuchs into Fox since a German name was then unacceptable, and although he intended this to be only for the duration of the war he found it so convenient that he later had it regularized by deed poll; and, ‘in order to distinguish myself from numerous other Foxes I began to use my second Christian name of Munro’.

When stationed in Egypt, Fox met Leonie Therese, the daughter of Henri Roger, a French official of the Suez Canal Company and they were married in 1917. In later years they became estranged and in 1931 he took as his second wife Natalia Lvovna, a daughter of Lev Yulevich Mertens of Saratov, Russia. ‘Natasha’ was his devoted companion to the end of his life, sharing with him his interest in natural history and supporting him in his busy social and professional life.

After the war, in 1919, Fox rejoined MacBride, but after six months in London he accepted an invitation from Edward Hindle, then Professor of Biology at the School of Medicine in Cairo, to join his staff as a lecturer. The post was especially attractive to him for he had suffered severely from hay-fever when in England and was free from it in Egypt; and Egypt was also his wife’s home. Prior to Hindle’s appointment as Professor at the Medical School there had never been any practical work within the courses

Harold Munro Fox 209



of biology. Fox was put in charge of the experimental side of the new courses and devised experiments, many of which were based on his own work, that were ideally suited to the limited facilities available. Together with Hindle he wrote a Laboratory notebook of elementary zoology and a Laboratory notebook of elementary botany as guides to the courses. These were published in 1922 by the Cairo Government Press and were used by successive generations of students within the University for many years. While in Cairo, Fox wrote up some work which he had begun at Plymouth in the summer of 1919, and later referred to in more detail, on the reactions of the flagellate protozoan Bodo to varying amounts of dissolved oxygen in water and he presented this work as a thesis for a Fellowship at Gonville and Caius. He was elected to a Fellowship but was unable to take it up until the following year when he returned to Cambridge on the termination of his Cairo appointment and his replacement, with the rise of Egyptian nationalism, by an Egyptian. He was no sooner back in Cambridge where, concurrent with his Fellowship he held the Balfour Studentship, before he began to organize, with the support of Stanley Gardiner, an expedition of four people to study the fauna of the Suez Canal. He had formed this project while in Egypt and was able to accomplish it largely through the financial support of the Canal Company. Fox returned to Cambridge to write up the results of the expedition and in 1927 was appointed to the Chair of Zoology in the University of Birmingham. He remained there for fourteen years before transferring to the Chair of Zoology at Bedford College in the University of London. The College had been evacuated to Cambridge at the beginning of the war and it gave Fox immense pleasure to be associated once again with Caius and to be teaching in the Department where he had earlier been a demonstrator and Balfour Student. When Bedford College returned to Regent’s Park at the end of the war the Department of Zoology was housed in Sussex Lodge. Fox had the difficult task of shaping a Regency residence to the requirements of a teaching department and, although the space allotted to him was small, he soon succeeded with the help of his small but able staff in making it into a vigorous and intimate teaching and research community. He retired with the title of Emeritus Professor in 1955 and at the invitation of the writer of this memoir made his home in the Department of Zoology of Queen Mary College. The College gave him the status of Research Associate and, in recognition of the distinction which he brought to the College through his scientific work and of his virtues as a colleague, conferred upon him a Fellowship of the College.

Fox’s researches spanned a period of more than fifty years and are recorded in some sixty-five papers and numerous shorter notes. His scientific work ranged widely over many aspects of zoology and included investigations in at least six different and largely unrelated fields of study. He began as an experimental embryologist and, after the interruption of the 1914-1919 war, turned to field studies of the breeding behaviour, ecology and distribution of the marine invertebrates of the eastern Mediterranean, Suez Canal and

210 Biographical Memoirs



Red Sea. In mid-life, as a university teacher and head of department, he undertook extended and important investigations into the nature and properties of the several kinds of respiratory pigments of marine and fresh water invertebrates, carrying along with this work in later years a contemporary study of the differing metabolic requirements and rates of heart beat and limb movement in species of marine invertebrates living in different latitudes and conditions of sea temperature. And finally, on his retirement, he changed course abruptly and concentrated his studies on the biology, anatomy and taxonomy of ostracod Crustacea.

Fox’s first published paper on the early larvae of two polychaete worms arose from a suggestion by Cresswell Shearer that, in preparation for the work on the hybridization of echinoids which they had planned, he should get some practice in making artificial fertilizations and in rearing larvae; and, having gained this experience, Fox, with Shearer and de Morgan, turned to the main problem. Some twenty years earlier Seeliger and other workers had successfully reared hybrid plutei from reciprocal male and female crosses of variously paired species of echinoids and had reported on the extent to which paternal and maternal characters were represented in the hybrids. Seeliger had noted a preponderance of paternal characters in the hybrid plutei; other investigators, on the other hand, had found maternal features to preponderate. There seemed to be two possible explanations of these discordant results. Either the characters which were being studied owed their variations of form to different conditions of culture or they were determined by the state of maturation of the egg at the time of its fertilization. In studies of Echinus esculentus—Echinocardium cordatum crosses Fox paid particular attention to the variations in pigmentation and skeleton construction of the aboral process of the larva and, although he found much variation in both these features, Fox concluded that inheritance showed a significant maternal dominance. However, on repeating these experiments in 1912 and extending them to other crosses, a high proportion of the larvae showed a dominance of paternal features. In further work, which included experiments with Mediterranean species, Shearer, Fox and de Morgan were able to show that the anomalies of the previous two seasons were not due to differing states of maturity of the eggs since successive fertilizations throughout the breeding period gave comparable types of plutei. Moreover, the larval features showed little variation under differing conditions of pH and temperature. It was tentatively concluded that the extent to which paternal or maternal features are represented in hybrid larvae may depend on the over-wintering conditions of gonad development. The outbreak of war put an end to further investigations and to work which Fox recognized as inconclusive and on the whole disappointing.

In 1913 when Fox was a lecturer at the Royal College of Science he brought to Professor MacBride’s notice the work of Paul Kammerer which purported to show from his experiments on salamanders, frogs and the




ascidian Ciona that characters acquired in response to the environmental conditions in which they were kept or as a result of certain kinds of operative treatment were transmitted to their offspring. MacBride’s stubborn advocacy of the inheritance of acquired characters is well remembered, and he was not disposed to think well of young men who strongly opposed his views on the subject. It says much for Fox’s courage therefore that, while still in MacBride’s department, he drew attention to the essential weakness of Kammerer’s experiments which had taken little account of the need to rear the offspring under conditions which eliminated the possibility of their acquiring their characteristics as environmental adaptations in the same way as had their parents. After the war Fox repeated Kammerer’s experiments on Ciona. This involved the amputation of the branchial siphons, after which (Kammerer had said) the siphons regenerate to a greater length, the animals thereafter transmitting this change to their offspring. Fox showed, however, that if the animals were kept in similar culture conditions before and after the amputation of the siphons there was no increase in length of the regenerated structures. However, an increase in the food supply induced the growth of long siphons and this happened both in siphon-amputated and in uninjured animals.

While carrying out his experiments on Ciona as Ray Lankester Investigator at the Plymouth Laboratory in 1919 Fox made some interesting observations on the behaviour of the flagellate Bodo under differing conditions of oxygen concentration. He noticed that when cultures of the flagellate, which he prepared from grass infusions, are examined under a coverslip the protozoans soon collect in a mass at the centre of the covered droplet of water. After a short time, however, they begin to move outwards towards the edge of the coverslip forming lines parallel to its edges. He further observed that Bodo aggregates on the surfaces of aquatic insects and other invertebrates in regions that are abstracting oxygen from the surrounding water, though they later move away with the progressive local deoxygenation of the water. Fox’s discovery that the flagellate avoids water of high and low oxygen concentration and finds its optimal conditions in water of intermediate oxygen concentration, and his exploitation of the behavioural response as a means of detecting the respiratory surfaces of animals formed the subject of the thesis which, as has already been noted, gained him a Fellowship in Cambridge.

On taking up his appointment as Edward Hindle’s assistant in Cairo he took the opportunity during the summer vacations of 1920 and 1921 to examine with Hindle the plankton and bottom fauna in the neighbourhood of Port Taufiq in the Gulf of Suez. It was here that he learned, in talking with the local fishermen, that they found it unprofitable to collect sea urchins at the period of the full moon as the gonads of the urchins—their only edible parts—are then empty. This observation led Fox to examine the breeding habits of a variety of marine invertebrates in order to discover the extent to which spawning may be determined by the moon’s phases and




Harold Munro Fox213the monthly rhythm of the tides. Although most of the animals he examined were found to be irregular in their spawning cycles or limited to a single emission of their gametes he was able to confirm that the urchin Centrechinus (Diadema) setosum undergoes a regular monthly cycle of gonad growth and discharge, spawning being confined to a few days around the period of the full moon. He was unable to relate the spawning cycle to variations of food supply, sea temperature, or salinity and he found no convincing evidence that spawning is conditioned by the changing regimen of the moon’s illumination. Fox was little given to speculation and the value of this work lies mainly in the accuracy and scope of his observations and in the comprehensive review presented in this paper of the phenomenon of lunar periodicity in animal and plant reproduction which has been used as a basis for many subsequent investigations.

After a brief period in Cambridge Fox returned in 1924 to Egypt as leader of an expedition in which Robert Gurney, V. C. Robinson and D. N. Twist also took part to survey and to make collections of the fauna of the Suez Canal and the adjacent areas of the Red Sea and eastern Mediterranean. The Red Sea and Mediterranean fauna are, in respect of a wide range of species, zoogeographically distinct and the main purpose of the expedition was to determine the extent to which the Suez Canal, opened in 1869, had been populated from the two seas and had enabled animals to pass from one region to another. The slightly higher level of the Red Sea causes slow and irregular currents to pass northwards through the canal, and it was found, in consequence, that the canal fauna is mainly composed of Red Sea species though a smaller number of Mediterranean species were also shown to have gained entry. The progress of the migrations was known, in some instances, from past records and in extension of these observations the expedition was able to trace the distribution of one species, the Red Sea swimming crab Neptunus pelagicus along the Mediterranean coast as far as Haifa. Fox enlisted the help of some of the leading taxonomists of the day to describe the material collected by the expedition and the published reports include contributions by Th. Mortensen, Hjalmar Broch, W. M. Tattersall, W. T. Caiman, E. T. Browne and Robert Gurney.

In the summer of 1923 while preparing for the Suez expedition Fox spent several weeks at the Roscoff laboratory in Brittany where, with the aid of a borrowed microspectroscope, he examined the absorption spectra of the dichroic (red-green) pigment in the blood of the sabellid worm graphis spallanzanii under varying conditions of oxidation and reduction ofthe pigment. In 1867 Ray Lankester had named the pigment chlorocruorin.

i He had shown it to occur in sabellid, serpulid and chlorhacmid worms and \ had found it to be similar in its chemical composition and properties to

haemoglobin. In a series of papers published intermittently over a period of thirty years Fox made an exhaustive study of the nature and physiological properties of the pigment. Having confirmed that chlorocruorin

' consists of an iron-containing haem combined with a porphyrin he went on



to show that derivative compounds prepared by denaturing the protein and by other physical and chemical reactions are essentially similar to the analagous derivatives of haemoglobin. Chlorocruorin was found to combine with oxygen in the proportion of one molecule of oxygen to each atom of iron in the pigment and (with J. Roche) he succeeded in obtaining oxychlorocruorin in crystalline form. Although the oxygen-carrying capacity of chlorocruorin was found to be significantly less than that of some haemoglobins Fox made the important discovery, which he later developed more fully in his investigations of haemoglobin, that the amount of pigment in the blood varies under different environmental conditions. Animals kept in partially deoxygenated waters synthesize the pigment, in well oxygenated waters on the other hand the concentration of chlorocruorin in the blood falls. In reporting on the occurrence of the pigment in four families of poly- chaetes Fox discovered that different species of serpulid worms may contain either haemoglobin or chlorocruorin, have both pigments, or have no pigment at all. As a broad generalization he concluded that chlorocruorin, a mutant form of haemoglobin, is, in spite of its lesser physiological efficiency, able to persist in animals with low oxygen requirements; and this conclusion was given especial point by the demonstration that when the pigment of worms such as Sabella is converted into carboxychlorocruorin the animal is able to meet its very limited oxygen needs by using the small amount of gas carried in solution in the blood.

Contemporary with his clarification of the nature and properties of chlorocruorin Fox carried out some collateral investigations on the oxygen uptake and rates of activity of related species of arthropods and annelids living in the contrasting environments of fresh water and sea water, of still and fast flowing fresh waters and of sea water in different latitudes and therefore of different temperature ranges. Consistently higher levels of oxygen uptake were recorded in freshwater amphipods and isopods than in related marine species; in fresh waters, ephemerid larvae living in flowing water were found to have a significantly higher rate of metabolism than those taken from the still waters of ponds. The high oxygen uptake of freshwater species was shown not wholly to reflect the metabolic needs of osmoregulation for the uptake was largely independent of the salt concentration of the blood. It was, however, found in some instances that high oxygen consumption is reflected in a rapid rate of heart beat.

Five papers reporting ‘The activity and metabolism of poikilothermal animals in different latitudes’ were based on work undertaken at Kristine- berg in Sweden, at Plymouth, and at Banyuls in the Mediterranean. Measurements were made of the oxygen consumption and the rate of the respiratory movements and heart beat of taxonomically related species from waters of different temperature ranges.

In this pioneer study of animals of different latitudes it was found that whereas animals of warmer waters have a higher oxygen consumption than related species from higher latitudes, the rate of their respiratory




Harold Munro Fox

movements and heart beat did not reflect a difference of metabolic rate. Fox found, however, that other activities such as ciliary beat and the rate of cleavage of echinoderm eggs are differently adapted to temperature variations, ciliary beat being faster in warmer and cleavage rate more rapid in colder waters. This work which was later taken up and in some respects brought into question by other workers was terminated by the Second World War and was judged by Fox himself to have been inconclusive.

‘After chlorocruorin’, says Fox in his personal record, ‘I turned my attention to the haemoglobin of Daphnia and later of other invertebrates. I showed that the haemoglobin content of the blood and muscles of Daphnia varies very greatly and is inversely proportional to the dissolved oxygen of the surrounding water. The utility of this to the animals was studied/ Fox is here referring to the first of a series of papers published between 1948 and 1955 in joint authorship with Barabara M. Gilchrist and Elizabeth A. Phear of Bedford College. The work began with the observation that the fresh water crustaceans Daphnia magna, D. obtura and D. pulex may be coloured red or pink. It was found that pink animals taken from oxygen-rich waters synthesize haemoglobin when transferred to partially deoxygenated water and that the change is reversible. Haemoglobin was shown to be present both in the blood and in the muscles as well as in parthenogenetically produced eggs. Fertilized eggs within the ephippia on the other hand were devoid of haemoglobin.

The causes and sequences of the fluctuations in haemoglobin content were investigated in further collaborative work. Haemoglobin synthesis was shown to be promoted by an increased food intake, by ferrous iron in solution, by vitamin Bia, by a rise in water temperature and, irrespective of these conditions, to vary from one instar to another and in particular with the approach of ovulation. Concurrent studies showed that an increase of haemoglobin and of oxygen carriage promoted a more rapid rate of feeding and swimming. Other animals such as conchostracan Crustacea, chironomid larvae and young water snails ( ) were later discoveredalso to gain or lose haemoglobin in response to environmental changes, and in the crustaceans studied there were concomitant fluctuations in the amount of cytochrome in the muscles. Annelids such as Arenicola, Scoloplos and Tubifex on the other hand showed no variation in their haemoglobin content in response to environmental changes, and an interesting outcome of this work was the discovery that animals adapted to habitats of low oxygen concentration are injured or killed in oxygen-saturated water.

Throughout the long period of his experimental studies Fox had published notes and papers (based on aquarium and field observations) on the natural history behaviour and habits of a number of aquatic invertebrates. They include observations on the swarming of Vorticella, on the remarkable longevity of a polychaete and of Aurelia scyphistomae, the appropriation by a hermit crab of its commensal anemone, and the rediscovery in Britain after a long interval of the crustacean Triops cancriformis.

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It is therefore not altogether surprising that on his retirement Fox found a deep satisfaction in an intensive study of the natural history, anatomy and’ taxonomy of fresh water and marine ostracods, a group which, though including some of the commonest of the smaller aquatic Crustacea, had in many aspects of their biology, distribution and relationships been little studied. Much of this work was done at the Istituto Italiano di Idrobiologia on Lake Pallanza in Northern Italy where Harold and Natasha Fox spent several summer vacations with their friends the late Professor Vittorio Tonolli (Director of the Institute) and Dr Livia Tonolli, but Fox’s accounts of ostracods include descriptions of animals collected from various parts of northern Italy, France and from many regularly visited localities in Britain. He became especially interested in ostracods distributed in underground waters and the writer’s last meeting with his friend was on a day spent in Cornwall searching for springs and netting them for specimens. Many were taken but none, alas, were hoped-for new species. The many beautifully prepared specimens on which Fox based his accounts in published papers are now in the collections of the British Museum (Natural History).

The remarkable range of Harold Fox’s zoological interests will be apparent from this far from complete review of his published work. He pioneered a number of original and productive lines of research which have subsequently been taken up and developed by other workers, and by his personal direction he guided the researches of many younger zoologists who learned from him the essential requirements of a scientific study and, by the example of his simple and direct style of writing, the value of an economical and lucid presentation. Dr Barbara M. Gilchrist, who was for many years, a close friend and professional colleague says of him that ‘he was a most stimulating and helpful supervisor of research. Not a week would go by but he would come in to see how things were going and invariably he would suggest a particular line to follow, and this would always prove most profitable. He was completely in touch with each step in the various lines of research going on in his department so that one could discuss the work at any stage without having to bring him up to date with what one was doing.’

Dr Gilchrist also speaks of Fox’s gift as a teacher. ‘He was a genius at explaining complicated things in simple terms—he would find an analogy with something quite common-place and so “light would dawn” for the student.’ The simplicity, directness and accurately informative style of Fox’s exposition are equally evident in the many popular books which he wrote and one of which, at least, The personality of animals has been translated into Italian and (what pleased Harold Fox most) into Japanese.

Fox’s forty years editorship of the Biological Reviews of the Cambridge Philosophical Society will perhaps remain the most abiding memorial of his international reputation and world-wide influence in contemporary zoology. The history of Biological Reviews is briefly outlined in the article he wrote in its pages in 1965. The jourrial under its present title and with its purpose




of presenting authoritative reviews on all aspects of biological thought and research developments was entirely of his creation. As its sole editor he was continuously involved in a search for new and suitable material. Almost every day he wrote letters to prospective authors and at regular intervals reminded them of their obligations. He always liked to have two or three numbers in hand and if the flow of articles seemed to be diminishing his colleagues would be taxed at tea-time conversations with a survey of other possible contributors. His standards were high and papers he thought inadequate or in need of revision were sent back to their authors copiously annotated with suggestions and criticisms which, from personal experience, the writer knows to have been uncompromising but invariably helpful. Those who have worked with Fox will know how great were the demands made upon him as editor of Biological Reviews. But for his immense powers of detachment and concentration which enabled him to revise manuscripts as readily and efficiently in a crowded underground train as in the quiet of his study he could not possibly have carried out his task concurrently with his many other activities.

Although Fox disliked committee work and thought most committees to be a waste of time he took a full share in the councils and consultative bodies of the universities in which he served. He also served on D.S.I.R. and other governmental committees. In the spring of 1946 he was invited by the Royal Society to travel around Europe to renew, on behalf of the Society, the contacts with European Academies which had been disrupted by the war. His ability to converse in several languages made him eminently suited for this renewal of understanding as a necessary preliminary to the reconstitution of the International Union of Biological Sciences. He was appointed a Vice-President of the I.U.B.S. and in 1950-1953 served as its President. He also travelled widely as a representative for natural sciences of U.N.E.S.G.O. giving lectures in Harvard and Yale as well as other American universities.

Fox was elected a Fellow of the Royal Society in 1937 and received many other honours. He was appointed Fullerian Professor of Physiology in the Royal Institution (1953-1956); elected as President d’Honneur of the Zoological Society of France in 1955. He received the degree of Honorary Doctor of Science of the University of Bordeaux in 1965, and was awarded a Darwin Medal of the Royal Society in 1966. He was also for many years, and up to the time of his death, Honorary President of the London Natural History Society.

Harold Fox was strikingly handsome both as a young man and in his later years. Upright of carriage and quick in movement he had an immense vitality which was reflected in a quickness of speech and a perpetual urgent questioning in conversation. He was a man of sharp prejudices; to him black was black and white was white and he was little given to compromise. Generous in his appreciation of the people and things he admired he could be devastatingly frank and outspoken on matters that irked him. Rudeness,




ugliness, bad or inefficient service, loud-voiced women, most contemporary art and any form of humbug were anathema to him. He delighted in vivid colours and when these were combined with other pleasant things such as auburn hair crowning a lovely face his admiration was unbounded. He was a man of simple tastes but enjoyed good things. He loved wine and good food, but drank and ate moderately, though he recalled with horror some formal occasions when he was expected to drink water. He was, in his younger days, a good swimmer and an accomplished ballroom dancer, and he remained to the end of his life an expert horseman. Music had few attractions for him and drama none, but he delighted in ballet. He was a great reader and enjoyed children’s books as much as, if not more than, sophisticated literature. Courtly and extremely modest he found salacious stories distasteful—and said so. Harold Fox loved the company of friends and dinner with Natasha and Harold as hosts was always a delightful occasion. He was for many years a member of the Savile Club and when on his retirement he felt that he could no longer afford the subscription he greatly missed the company of his club friends, though characteristically he came to say that Queen Mary College was the best club in London. To his friends he was generous and loyal. He never came to stay as a guest without bearing a gift and it was always necessary to pose for photographs on his departure. A man of fine integrity, of great gifts, and delightfully unusual in his idiosyncracies Harold Fox’s warm and vivid personality is gratefully remembered by his many friends.

I have been greatly helped in the preparation of this Memoir by a number of Harold Fox’s friends and former colleagues, including Dr Edward Hindle, F.R.S., Professor Sir James Gray, F.R.S., Dr Barbara M. Gilchrist, who also kindly provided a complete bibliography of Fox’s publications; and I am particularly indebted to Mrs Alison Settle on whose notes the account of Fox’s early life is largely based. Above all I am grateful for the talks I have had with Mrs Natasha Fox who also provided the photograph which illustrates the memoir. Harold Fox’s personal record has been used extensively in the preparation of the article.

J. E. Smith


BIBLIOGRAPHY

1911. Note on the early larvae of JVephthys and Glycera. biol. 9, 164.1911. Preliminary notice on the experimental hybridization of echinoids. J. mar. biol.

Ass. U.K. 9, 121.1912. The inheritance of the aboral process of the Echinocardium pluteus. Arch. EntwMech.

Org. 35, 558.1912. On paternal characters in'echinoid hybrids. Quart. J . micr. Sci. 58, 333.



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