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Department of Zoology


Carl Pantin, the fifth Professor of Zoology, was born in the 19th century; just, in 1899. Pantin’s time as Professor was next to the shortest; he served for seven years. (The truncated reign of Adam Sedgwick was the shortest at just two years).

The photographic portrait in Pantin’s entry in Royal Society Biographical Memoirs gives the impression of a scholarly man, peering benignly at what must be one of his own, very beautiful, watercolours of a transverse section of a sea anemone. The impression of enduring calm in the photograph is, however, rather misleading, because it is quite clear from those who knew him that Carl Pantin was a huge enthusiast: an enthusiast for all aspects of the natural world, and indeed for life in general, serially possessed of energetic and deep interests in those topics that engaged him. He communicated this enthusiasm through his teaching, and he was a prolific writer over an eclectic range of subjects: my count of his publications – papers, articles, and reviews  – comes to around 130. The majority of these publications has Pantin as single author, but some of the co-authors whose names may be recognized by those reading this biographical sketch include Lancelot Hogben, Frank S.J. Hollick, J.F. Danielli, A.V. (‘Bill’) Grimstone, Elizabeth (‘Betty’) Batham, and Elaine Robson. And Carl Pantin was hardly unworldly: he became a director, and then Chairman, of the family firm, specialising in timber and machinery, in which he took a keen interest.

It is fitting that Pantin’s entry in Biographical Memoirs was written by F.S. Russell, one of the great names in British marine biology, because the scientific lives of the two developed much in parallel. Sir Frederick Russell was educated at Gonville and Caius and spent his career, from 1924, at the Marine Biological Association (MBA) in Plymouth, becoming its Director in 1945. Russell and Pantin were contemporaries as Cambridge undergraduates but did not really meet (Pantin was at Christ’s) until both were on the staff of the MBA. Russell was elected to the Royal Society in 1938, just one year after Carl Pantin, and the two shared research interests, particularly in coelenterates (Russell’s The Medusae of the British Isles is a classic of marine biology).

Carl Pantin’s parents were Herbert and Emilie. Emilie Abel had distinguished German ancestry. Indeed, Carl received his germanic-sounding first name from an antecedent, the composer Karl Friedrich Abel, a pupil of J.S. Bach. Several of the Abels were notable scientists. As a young boy, Carl was reading H.G. Wells and Arthur Mee’s Children's Encyclopaedia, with its many articles on science and industry. He was educated at Tonbridge School from 1913 to 1917, where he was a model pupil, which positioned him ideally for joining the Royal Engineers as an Officer Cadet in late 1917. He was sent to France in early 1918, in time to see service towards the end of the First World War. On Carl’s return from the War he came to Cambridge in 1919. F.S. Russell notes that Carl’s original intention was to study physics, probably as a result of his interest in the family firm and his training and experience in the Royal Engineers. However, he was attracted to zoology, perhaps as a result of his early enthusiasm for natural history (his father introduced him to angling, which became a permanent passion). The following quotation from Russell gives a flavour of the teaching then on offer in the Department.

‘J. Stanley Gardiner was then Professor and the lecturers included Hans Gadow, for vertebrates, L. A. Borradaile and F. A. Potts for invertebrates, and F. Balfour-Browne and G. P. Bidder on insects and sponges respectively. Experimental zoology and ecology were then creeping into the syllabus with lectures given by James Gray on experimental cytology and J. T. Saunders on freshwater ecology. Johan Hjort gave a special series of lectures on marine science.’

So, zoology it was.

Most of the names in Russell’s account will already be known to at least some of those reading this sketch, but probably the names of J.T. Saunders and Johan Hjort will be rather less familiar. So I include here brief digressions on both personalities to give a better idea of the teaching environment in which the undergraduate Carl Pantin would have received his initial inspiration.

Mr J.T. Saunders was a Fellow of Christ’s College (Pantin’s own College), was appointed Demonstrator in Animal Morphology, and became Lecturer in Zoology; he is the ‘Saunders’ of the textbook BEPS (Borradaile, Eastham, Potts and Saunders, The Invertebrata: A Manual for the Use of Students, Cambridge University Press) familiar to generations of students, and the editor, with the then Professor of Physiology, Joseph Barcroft, of the series Cambridge Comparative Physiology, also published by Cambridge University Press. Saunders was an enthusiast for experimental research at the crucial time when, although such work was still a minority interest, younger workers (among them the young James Gray, later to become the fourth Professor of Zoology, and hence Pantin’s immediate predecessor in that post) were beginning to move in the experimental direction. However, the lectures Pantin will have received from Saunders were in freshwater ecology, another area in which Saunders is conspicuous as a pioneer. (Although the Cambridge-trained ecologist G. Evelyn Huthchinson, later to achieve legendary status as ‘the father of modern ecology’ at Yale, who attended Saunders’s lectures, was singularly unimpressed, according to his memoirs).

The Norwegian Johan Hjort’s name will certainly be familiar to those interested in the history of marine biology, but otherwise, for most zoologists, it seems that he has somewhat sunk beneath the waves. In 1919 Hjort would have been 50 years old, and his time in Cambridge brought expertise in both pure and applied marine biology. Hjort was described as ‘ … one of the great leaders in oceanography whose names will live in the annals of that science … ’ by no less an authority than Sir Alister Hardy. In 1921 Hjort left Cambridge for a professorship in Oslo; he received an honorary degree from Cambridge.

The combination of experimental and ecological teaching (the latter particularly in marine biology) from Gray, Saunders, and Hjort appear to have gone a long way to setting Carl Pantin’s lifelong research interests. It was, of course, also the case that a thorough training in morphology could be taken for granted, and, perhaps more surprisingly, Pantin maintained a strong fascination for palaeontology.

In 1922, after his graduation, Pantin moved to the Marine Biological Association in Plymouth, to the physiology division. His initial work at Plymouth involved a variety of aspects of amoeba physiology, using the so-called limax form of amoeba (limax denotes a body form and movement-type of amoebae spanning several families). He went on to investigate osmotic regulation in the triclad platyhelminthe Procerodes ulvae, notable for its chosen residency in freshwater run-off onto a marine foreshore, thereby exposing it to both salt and fresh water over each tidal cycle. (Incidentally, a search for this species on the current WORMS, the World Register of Marine Species, entertainingly produces the message, ‘Oops! This taxon is out of scope! The taxon you have searched for is non-marine’.) The secret seemed to be, as so often, in the mucus, which persists more assiduously in the presence of higher concentrations of calcium ions.

From 1925 for five years, Pantin ran experimental courses at Plymouth; and in 1927/8, while still at Plymouth, he lectured on physiology at University College, London. In 1929, however, Pantin returned to Cambridge, to a Fellowship at Trinity College, and membership of the Department of Zoology. He stayed for the rest of his life, but not without excursions; to the Naples Zoological Station in 1933, for example. This period of study at Naples saw him initiate the work on sea anemones (he started with Calliactis) for which he has generally been best known. His work led him to the conclusion that, ‘The whole scheme is thus 100 to 1000 times slower than that which characterises the nervous organisation of the vertebrates.’ To anyone who has watched anemones for long, this is not, perhaps, too surprising, but the graded nature of responses demonstrated by Pantin helps to elucidate the natural history observation that the anemone’s life attached to a hermit crab must expose it to a continual series of small, challenging shocks: instant and total responses would be inappropriate. Also in the early 1930s, Pantin extended his neurophysiological studies to the leg musculature of the shore crab, Carcinus maenas, concluding, ‘There is a close analogy between the neuromuscular mechanism disclosed here and the neuromuscular mechanism of Coelenterata. In both there is a tendency for an entire effector to behave as a single system in which the response is governed by the number and frequency of impulses received by the muscle. This system is distinguished sharply from that of vertebrate skeletal muscle in which gradation of response is brought about through the multiplicity of motor units.’ Work on anemones continued, as Metridium and Anemonia were added to the suite of experimental subjects, and Pantin gave attention to pigment properties and the control of stinging-cell discharge. In some of this work he was assisted by Betty Batham (for details of her eventful life, see the Dictionary of New Zealand Biography: and Elaine Robson (who subsequently carried out her own studies on the swimming behaviour of the anemone Stomphia). In 1933 Pantin contracted tuberculosis, which removed him from research for a year.

Under James Gray, who became Professor of Zoology in 1937, Carl Pantin was asked to take on the invertebrate teaching in Part II. In 1938, Pantin recruited J. Eric Smith from Sheffield to a  Demonstratorship in Zoology. The two taught the invertebrates courses together and became close friends. Eric Smith left Cambridge in 1950 to take up the chair at Queen Mary College, London, leaving QMC to become Director of the Plymouth Marine Laboratory and Secretary of the Marine Biological Association in 1965 until his retirement in 1974. In these posts he succeeded F.S. Russell, whose Biographical Memoir of Carl Pantin has provided much of the material on which this current sketch is based. In turn, Pantin, himself, served as President of the Marine Biological Association from 1960 to 1966.

Nemertime worms

In the early 1940s, Carl Pantin began a new series of studies on terrestrial nemertine worms. He became characteristically absorbed in their natural history, morphology, distribution, and evolution, although his publications on nemertines are far fewer than those on sea anemones. This work was continued after Pantin’s death by Janet Moore, who was able to use the specimens that he had collected and, in many cases, prepared as serial sections.

Apart from Carl’s field trips, both zoological and geological, in Britain, the Pantins visited Uganda in 1962, Carl in his capacity as Chairman of a Nuffield Trust supporting tropical animal ecology. (Carl married Edith Smith, known as ‘Amy’, a botanist, his contemporary as a Cambridge undergraduate, and later one of his assistants in the anemone work.) There were other visits to America, Kenya, Sudan, New Zealand, and Brazil. In 1965, when he visited America, he was already ill with the leukaemia that was to kill him in 1967, a mere year after his retirement.

Russell’s Biographical Memoir of Pantin gives exhaustive details of his many presidencies, chairmanships, medals and honorary degrees. He was both a dedicated Cambridge academic and a person who gave freely of himself in wider public service: a scientist given to highly detailed work, but also one who could see the bigger picture. In his obituary of Carl Pantin, written for Trinity College, Mark Pryor, himself both a member of the Department of Zoology and a Fellow of Trinity, writes, ‘He was an enthusiast, he shook hands from the shoulder, his eyes shone “Splendid, splendid”. I suppose he did it on purpose to encourage the young, but it came straight from the heart all the same.’

Hole in the ground

F.S. Russell quotes J.A. (Arthur) Ramsay (another subject of an earlier one of these biographical sketches) who wrote to Russell, concerning Pantin, ‘What I think he set himself to do, as head of department, was first to secure the position of various members of the staff who were not yet confirmed in their appointments, and second to hand over a good going concern to his successor. In the first he succeeded gloriously. In the second he was defeated by circumstances outside his control. He should have been able to hand over a bright new building; instead, he handed over a hole in the ground.’ The handover was to Torkel Weis-Fogh, the sixth Professor of Zoology, and that ‘hole in the ground’ saw the completion, after several years, of what is now the ‘David Attenborough Building’.

Written by Dr Adrian Friday