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Herbert Graham Cannon, 1897-1963

J. E. Smith

, 55-68, published 1 November 196391963 Biogr. Mems Fell. R. Soc.

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HERBERT GRAHAM CANNON

1897-1963

H erbert G raham Cannon , Beyer Professor in the University of Manchester, died in a London hospital on 6 January 1963, at the age of sixty-five. He had been for more than forty years a leading figure among British zoologists. An inspiring teacher, a vigorous and controversial writer, he had contributed greatly to the advancement of zoology by his researches on arthro- podan animals and, in particular, by his interpretations of the embryology of the Crustacea and of the nature of their feeding mechanisms.

Graham Cannon was born on 14 April 1897, in Wimbledon, the third child of a family of four. His father David William Cannon was a compositor in the firm of Eyre and Spottiswoode and was for several years engaged in the preparation of maps and papers for the India Office; his mother was the daughter of a Charles Graham who owned and drove one of the first horse- buses to run on a regular service in south London. David Cannon’s weekly wage was little enough for the needs of a largish family and when Graham was about five years old the parents moved from Wimbledon to a house in Brixton near to which there were some good free schools. Cannon was sent for a time to the local council school and, on winning a scholarship, he moved on to Wilson’s Grammar School in Camberwell where he followed the normal grammar school curriculum specializing, in the higher forms, in science subjects in preparation for entrance to a University. He took little part in field games, but he was interested in chess and rifle shooting. He also enjoyed making things, and early developed a facility for drawing and sketching. As evidence of this I have been told that one of his schoolmasters, the father of a present Professor of Zoology, thought sufficiently well of the young Cannon’s drawings to keep them to the day of his retirement as examples of the work of his more talented pupils. But Graham’s main love was music. He had a good singing voice and was for several years a choirboy at the local church where, on special occasions such as the Christmas and Easter services, he and his brother would almost invariably be called upon to take the solo parts.

Cannon’s ability as an artist and a singer were to stand him in good stead in his later career. But there is little doubt that, in its later consequences, the most significant event of his schoolboy days was the accident he suffered at the age of sixteen. He had been to the theatre to see a performance of ‘The Arcadians’ where, in one of the scenes, a man jumps from an aeroplane as it descends from the wings to the floor of the stage. Cannon



^6 Biographical Memoirs

remembered that later that night he had dreamed about this incident; and, in the vividness of the dream, he had walked in his sleep and jumped out of the window. As a result of the fall he sustained compound fractures of both wrists and damage to the pelvis and lumbar vertebrae. His injuries left him with permanent physical disabilities, and for the rest of his life he was subject to attacks of acute and often crippling pain. These he bore with great fortitude, but he always found it difficult to sit for long periods at a bench and towards the end of his life he found it impossible to work with a microscope and to continue his anatomical studies.

Cannon acknowledged with characteristic honesty one immediate advantage he gained from his accident, for, being unfit for war service, he was able to start his University courses immediately on leaving school. He went up to Cambridge in the Michaelmas term of 1916 as a Choral Scholar of Christ s College, where F. H. A. (‘Tibby’) Marshall, for whom Cannon had a great affection, was his tutor, and began to read for the Part I Natural Sciences Tripos. His subjects in addition to zoology, were chemistry and physiology; and in the final examination of 1918 he was awarded first class honours. Instead, however, of choosing to spend the third year on a Part II course he obtained leave of absence to take up a temporary post as Naturalist with the Board of Fisheries. He worked during the summer of 1918 at the fisheries laboratory at Conway, and it seems that by the autumn of that year he had moved to London, for he used to recall how he had viewed from his office window the scenes in Whitehall on the morning of 11 November 1918, when news came through of the signing of the armistice. The war having ended, and having been excused the one term, he was free to return to Cambridge to complete the three years of residence required as a qualification for the B.A. degree. During the war the teaching of zoology in Cambridge had been mainly in the hands of Professor Stanley Gardiner, Hans Gadow, L. A. Borradaile, Leonard Doncaster, H. H. Brindley and Cecil Warburton, some of whom, however, had been away from Cambridge on war service for longer or shorter periods. By January 1919, James (later Professor Sir James) Gray, F. A. Potts, J. T. Saunders and other members of the staff of the Zoology Department had returned from war service, but they needed to pick up the threads of their long interrupted researches and had not yet fully taken up their teaching duties. Leonard Doncaster, who was somewhat older than the others, was a Quaker and had served in France for a year as a member of the Friends’ Ambulance Unit. He was a man universally admired for his friendliness, high principles and sterling character, and his studies in cytogenetics, which included work on sex determination in sawffies and gallflies, phenotypic variation in sea-urchin hybrids and the demonstration of instances of sex-linkage, had earned him election into the Fellowship of the Royal Society at an early age. Doncaster took the young Cannon under his wing and they worked together on a study of the spermatogenesis of the louse Pediculus. The study had been prompted by an observation made by Captain (later Professor) Edward Hindle that the offspring of some matings



Herbert Graham Cannon

of Pediculus were all, or almost all, of one sex. Doncaster and Cannon, while not fully confirming Hindle’s results, found that there was indeed a considerable disparity in the sex ratio of the Fx generation which they related to peculiarities of the meiotic division of the spermatogonial nuclei. They described their findings in a paper published early in 1920 for which Cannon drew most of the figures. Doncaster left Cambridge in October 1919, on his election to the Chair of Zoology in Liverpool, and died a few months later.

Cannon had now to decide on the line of research he was to follow. One who knew him well at this time has said that he thought seriously of taking up an experimental study. Experimental zoology was in its early and exciting days of discovery and the young men of the Cambridge school, led by James Gray, were being attracted in increasing numbers by the new approaches to the understanding of animal form and function. Cannon was well equipped for experimental work. He had a competent knowledge of chemistry and mathematics, was handy in the making of gadgets and apparatus, and he had a capacity for clear thinking which would have ensured a thorough understanding of the purposes and limitations of an experiment. He was, however, a little ahead in time of the main stream of the experimental work in Cambridge, and after considering the opportunities open to him—for there was no prospect of his remaining in Cambridge—he decided to continue for a while with the cytological studies begun with Doncaster.

In 1920 Cannon was appointed as a demonstrator in the Department of Zoology of Imperial College under Professor E. W. MacBride. The department housed in the Royal College of Science building was not large but was an active centre of research. MacBride’s ‘tea-parties’ provided regular meeting places for zoologists in the various London colleges and in the Natural History Museum, and the discussions of research topics, particularly when they touched upon phylogeny and evolution, were invariably lively and provocative. It was not possible to be in MacBride’s company for long and not be goaded into an argument, and Cannon recalls in his memoir of W. T. Caiman, written in 1953, that ‘when the Lamarckian atmosphere got too exciting, I could always run across the road to cool down in the old brick building at the back of the Museum’. It was here that he sought out Caiman to talk with him for a while and to enjoy a clandestine smoke.

Cannon’s friendship with Caiman, who had an immense knowledge of the Crustacea, marked a turning point in his career and a change in the direction of his research. For in 1922 he completed the last of his five papers on the cytology of germ cells and began the long series of informative and penetrating investigations of the embryology, feeding mechanisms, and general anatomy of the Crustacea which were to establish his reputation as a zoologist of high distinction. The reports of the Council of the Marine Biological Association of the early 1920’s provide in the lists of workers at the Plymouth Laboratory some indication of the way in which the re-alinement of his

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^8 Biographical Memoirs

researches came about. In 1919, while still at Cambridge, he had spent a short time at Plymouth getting to know marine animals and he is noted in the report as studying ‘general zoology’. On his return in 1920, no doubt having discussed with MacBride, who was an embryologist, possible research problems, he put himself down as working on the embryology of nudibranch molluscs. Nothing seems to have come of this but the interest in embryology remained and, when in 1923 Caiman gave him some dried mud containing eggs of the conchostracan Estheria which had been sent to him by an army officer in Baghdad, and from which he had hatched out nauplii and reared them to the adult stage, Cannon reared his own cultures and used them for preparing sections of the successive stages and for describing their organogeny.

Cannon’s early papers on the embryology of the Crustacea are notable for two important discoveries. He showed in his studies of the embryology of Estheria (1924, 1925) that certain of the muscles of this animal are of ectodermal origin, and in this and a later investigation of the post-embryonic development of the fairy-shrimp Chirocephalus (1926) he demonstrated the occurrence of coelomic cavities within a number of the mesodermal somites and was able to trace the fate of these cavities and the further development of their mesodermal walls. The idea that muscles can be developed from ectoderm was at variance with the then generally accepted view that they are always mesodermal or mesenchymal in origin, but Cannon’s interpretation has since been fully substantiated, and it is now known that homologous systems of muscles may in some crustaceans arise from mesoderm and in others from ectoderm. His discovery of coelomic cavities in the embryonic mesoderm of crustaceans is perhaps of even greater interest, for it not only rationalizes the position of the Arthropoda within the protostome line of invertebrate phylogeny but leads to an understanding of the homologies of the systems of organs and structures with which the coelom is associated.

Cannon’s papers on the embryology of the Crustacea which included, in addition to the papers mentioned, an important study of the genesis of the segmental excretory organs of ostracods (1925), earned him in 1926 the D.Sc. degree of the University of London, and in 1927 the high quality of his researches was recognized by his election as a Fellow of the Royal Society of Edinburgh and the award of the Crisp Medal of the Linnean Society. Meanwhile he had been elected, at the early age of 29, to the Chair ol Zoology in Sheffield as the successor of L. C. Miall. Shortly after settling in Sheffield, he married. His bride, Miss Annie Helen Fyfe, was a zoologist and a graduate of the University of Edinburgh. Their first meeting had been at the Plymouth Laboratory where Graham Cannon was occupying a research table and Nannie Fyfe was a member of one o fj. H. Orton’s celebrated Easter Classes for university students. It was the beginning of a singularly happy partnership and of a family life enriched by the mutual devotion of the parents and of the four children born of their marriage. Those of us who were privileged to receive the frequent invitations to their homes at




Chinley and at Chapel-en-le-Frith in the beautiful Derbyshire countryside remember with an abiding pleasure the evenings spent with Graham, Nannie and the children, the fun of their firework parties, the enjoyment of listening to music and of talking by their fireside and the relaxed sense of well-being and friendship which grew naturally in their company and in the serene companionship of their family life.

Cannon remained in Sheffield for five years and though he had a full teaching programme he published during this period four important papers on the feeding mechanisms of the Crustacea. Two papers (‘On the feeding mechanism of a mysid crustacean, Hemimysis ’ (1927) ; ‘On the feeding- mechanism of the syncarid Crustacea’ (1929)) were on work done in collaboration with Dr Sidnie M. Manton; of two (‘On the feeding mechanism of Nebalia bipes’ (1927); ‘On the feeding mechanisms of the copepods Calanus Jinmarchicus and Diaptomus gracilis’ (1928)) he was the sole author. It had long been known that many crustaceans are able to feed by filtering small particles and organisms held in suspension in the surrounding water. The particles are drawn towards the body in currents generated by the movements of the limbs, trapped in the meshes of the limb setae from whence, by various mechanisms, they are combed off and conveyed to the mouth. And crustaceans which use this method often do so as an alternative to or in conjunction with the seizing and manipulation of larger food masses by the mandibles and the adjacent mouth parts. The mechanisms of current production, filtration, and transport had been examined in a few instances, notably in the cladoceran ‘water-fleas’. Cannon & Manton had, however, been impressed by the fact that explanations of the mechanisms were often faulty or inadequate, largely because of the failure of previous observers to relate the methods of food capture to the shapes, postures and relative motions of the limbs as they were to be seen in the living animal. Their methods of study were simple and elegant. The directions of the feeding currents on their approach to the animal and within the interlimb spaces were recorded by observing the movements of particles and of dyes when the animal was free to exercise its limbs without restraint. The movements of the limbs were observed and their rhythms analyzed by stroboscope viewing or by other appropriate methods. The sites of filtration, the methods of combing and the sequences of particle transport were then deduced by observing the distribution of particles and the form and positions of the limbs in animals fixed and embedded by methods ensuring a minimum of distortion, and subsequently stained and cut into thick slices so as to allow whole series of limbs to be viewed in their natural relationships.

These studies not only furnished satisfactory explanations of the feeding mechanisms of a wide variety of crustaceans but gave new meaning to the relation of the form of the limbs to their functions, for the analysis required examination both of the general shape of the appendages and of the detailed structure of their several parts, for example the character of their lobes and the varied development and differing character of their setae. Moreover, by

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comparing the feeding mechanisms in the different groups of Crustacea it became evident, as Cannon showed in a later paper ‘On the feeding mechanisms of the Branchiopoda’ (1933) that similar mechanical principles of current production and particle collection could involve, in different animals, different component parts and different limb series. It was further shown that the mechanisms employed could, in some groups of the Crustacea, be fitted into an evolutionary series with the more primitive forms often using a predominantly filtratory method of feeding, the more specialized animals relying wholly or mainly on a raptatory method of food capture. And, in describing the feeding mechanism of Cannon demonstratedits essentially malacostracan character, thus anticipating Manton’s embryo- logical studies on this group which were to show that the supposed similarities of the Leptostraca to the ‘entomostracan’ Crustacea were due to the convergent development of somewhat similar types of limbs, the Leptostraca showing in their body segmentation and in other features of their anatomy undoubted affinities with the Malacostraca.

In 1931 Cannon left Sheffield to become the Beyer Professor of Zoology in the University of Manchester. The Manchester school had acquired under Professor S. J. Hickson a high reputation for sound teaching and good research, and provided through its numerous research students and assistant staff greater opportunities than had been available in the smaller Sheffield department for the direction and encouragement of original work. Cannon numbered among his staff in these early years in Manchester four lecturers, James Brough, B. J. Marples, Ralph Dennell and the present writer who were later to be appointed to University chairs. Norman Millott and P. M. Butler, who were also destined for chairs, joined his staff a little later. All of them, and many more, readily acknowledge the debt they owe to their old chief for his invaluable advice and for his sympathetic encouragement of their early researches.

After publishing, in 1933, one further paper ‘On the feeding mechanism of certain marine ostracods’ Cannon brought this aspect of his work on the Crustacea to a close with the magnificently comprehensive review ‘On the feeding mechanism of the Branchiopoda’ (1933). Two years later he was elected into the Fellowship of the Royal Society.

He now turned to anatomical studies of a more general kind. The collections in the British Museum of Natural History included a great deal of undescribed expedition material and it was natural that Cannon should be invited to report on such Crustacea as might be of interest to him. He selected for examination the rock-boring barnacle Lithotrya a giantcherry-sized ostracod Gigantocypris mulleri, and a southern ocean representative of the Leptostraca, Nebaliopsis typica. The reports he subsequently made on these animals, published in the Discovery Reports and the Reports of the Great Barrier Reef Expedition, are models of anatomical description and illustration and, as always in Cannon’s papers, the functions of organ systems are convincingly interpreted in relation to their structural design.

60 Biographical Memoirs



Herbert Graham Cannon 61The last of these papers appeared in 1947, by which time Cannon had

begun to acquire a heavy load of administrative work. He had always taken a full part in the work of university committees, had been Dean of the Faculty of Science and, since 1943, Chairman of the Manchester Museum Committee. In 1946 he was appointed Pro Vice-Chancellor of the University for a period of four years and, on relinquishing his office, he was appointed one of the two Manchester Academic Sponsors of the newly-founded University College of North Staffordshire at Keele. These additional duties, to which he devoted a great deal of time and attention, and the frequent indispositions which were the legacy of his boyhood injuries left him little time for research. He continued, however, to compile a monograph of the Leptostraca and had the satisfaction of seeing it published in 1960 as a volume of Bronn’s Klassen und Ordnung des Tierreichs.

Early in 1962 Cannon was operated upon for acute appendicitis. The operation was successful but recovery slow. In the summer of that year the University of Manchester gave him a generous grant towards the expenses of a recuperative holiday. In company with Mrs Cannon he embarked on a leisurely voyage to West Africa where, in Freetown, he had the pleasure of seeing his eldest son David and his wife and family. He appeared to be making a good recovery, but on reaching Las Palmas on the return journey he again became ill and was admitted to hospital. He was found to be suffering from ulceration of the liver and it was advised that he should be flown home. He died in London shortly after admission to St Bartholomew’s Hospital.

The distinguishing qualities of the researches which have been referred to in this brief review of Cannon’s career are accuracy of observation, clarity of presentation, and an insistence on the need to relate the design of animals and the structure of their parts to their habits and machinery of function. And there is little doubt that Cannon’s approach to his researches and his methods of study go far to explain the attitudes he took and the opinions he expressed on two matters which lay outside the narrower range of his observational studies and to which he devoted an almost passionate attention, namely the teaching of biology and its role in education, and the interpretation of the Lamarckian view of animal evolution.

It was Cannon’s custom in carrying out his anatomical investigations to make use of rather few preparations, for anything that he regarded as imperfect he rejected. But having got the preparation he wanted he would set it up under his microscope and return to it again and again in order to make sure that no part of it had escaped his attention or any detail of its structure overlooked. One such preparation, immersed in alcohol and pinned to a piece of black velvet in order to reduce background reflexions and which he had been viewing and sketching over a period of some weeks, led to his discovery of the usefulness as a biological stain of the dye Chlorazol Black. Some of the dye had leached from the velvet and had penetrated the preparation to outline features of it that had not previously been clear. Cannon



obtained samples of the dye, tested its capabilities on material fixed in various ways, and reported on its merits as a whole mount and as a nuclear stain in two papers published in 1937 and 1941.

His meticulous attention to detail and regard for accuracy of observation is evident in the beautifully drawn figures which illustrate his papers. He was not content merely to give an impression of what he saw; his figures faithfully portray the actual appearance of the features he intends to describe and show them, as photographs would not, unencumbered by irrelevant surrounding structures. In his little book A method of illustration for scientific papers (1936), written at the request of the Association of British Zoologists, he describes how he prepared his illustrations and gives some invaluable advice on the practice of line drawing, shading and labelling of pen and ink drawings suitable for reproduction by block printing. Finally, a word must be said about Cannon’s style of writing. His descriptions of animals are framed in simple direct language and are phrased in short sentences free from a clutter of dependent clauses. When he uses technical terms he invariably examines their meaning, and if (as when, for example, he uses the term ‘gnathobase’) he considers them to have an inappropriate connotation he does not hesitate to redefine them or to indicate their limitations. His texts are, without exception, easy to read and perfectly suited to the purposes of anatomical description.

As might be expected of a man who delighted in the forms of animals and in the infinite variety of their structural designs and functional adaptations, and who wrote so lucidly in the explanation of them, Cannon was an admirable teacher. He loved an audience and was quick to sense its reactions. He prepared his lectures to the last detail and sometimes wrote them out in full. But he never gave the impression of being tied to his notes and, when explaining the anatomy of an animal, he would build up its structure in beautifully executed blackboard drawings, pausing at times to fashion by the movements of his hands the form and action of its movable parts.

Cannon’s university teaching was mainly confined to the courses for first year students. He held strongly to the view that it was the duty of a professor to lay the foundations of his subject, and the foundations of zoology he regarded as consisting primarily of a knowledge of the nature and variety of animal design and an understanding of their systems of relationship and evolutionary development. His first year lectures were therefore directed towards a survey of the major phyla of animals during the course of which he explained the character of their distinctive architectural styles and the range of their functional adaptations by examining in detail a few representative animals of each phylum. And, in following the ‘type system’, he dealt with other organismal attributes, such as their physiological mechanisms, life histories, fossil ancestry, behaviour and distribution, in conjunction with the groups which he thought most appropriate for their illustration. Cannon’s conviction that animals should be viewed, first and foremost, in the wholeness of their interrelated attributes of structure, function, habits, history and





relationships led him to become increasingly impatient, in the framing of school examination syllabuses and in the teaching of biology in schools, of the neglect of these approaches. In letters to The , the (Manchester)Guardian and to Nature he protested vigorously against the introduction into sixth form teaching of topics based on the latest and most fashionable lines of research, on the emphasis laid on Mendelian genetics which he regarded as too often treated merely as an exercise in arithmetic, and on the inclusion of other specialized aspects of zoology of necessity treated superficially and as snippets of information. There is no doubt that in taking this extreme view of the dangers of early specialization Cannon was overstating his case. Young people not only need to know what zoology is about, they ought also to know where it is leading; and as a subject of education its particular forms of analysis are no less important than are its synoptic viewpoints and broader generalizations. The problem, which with the extension of knowledge is becoming increasingly difficult of solution, is how to preserve the balance. But Cannon was certainly getting the priorities right in insisting that it is necessary to know what animals are, how they live, and what they do, if the detailed analysis of their functional processes is to have a purpose and a meaning.

In his later years Cannon began to give increasing thought to the problem of evolution and, in particular, to the origin and nature of the heritable variations implicit in evolutionary change. In a series of articles which include ‘Is the problem of evolution solved?’ (1954) and ‘What Lamarck really said’ (1957), and in two books The evolution of living things (1958) and Lamarck and modern genetics (1959) he criticized the neo-Mendelian explanations of the nature of evolutionary processes and argued that the Lamarckian theory, when correctly interpreted, offers a more satisfactory explanation of them. Cannon’s main reasons for rejecting the neo-Mendelian view of evolutionary mechanisms were (1) that the phenotypic variations consequent on gene mutation and chromosome recombination are, almost without exception, restricted to trivial and localized modifications of structure and of function which have no obvious adaptive value or evolutionary significance, and (2) that in primitive organisms such as viruses, which lack an organized nucleus, inheritance must be vested in the cytoplasm; if, therefore, cytoplasmic inheritance is demonstrable in the lower organisms it cannot be ruled out in plants and animals generally. Cannon therefore ascribed the ability of organisms to adapt to environmental change to adjustments made within the complex systems of the cytoplasm, allowing that these adjustments can be modified, but not wholly directed, by the activities of the chromosomes. This concept of the cytoplasmic mediation of adaptive change led Cannon to re-examine Lamarck’s writings which, though they preceded by half a century Mendel’s demonstration of factorial inheritance and by almost a century the identification of factors with genes, seemed to him to embody ideas which presupposed cytoplasmic inheritance. Cannon showed that Lamarck’s statement of his theory of evolution in the

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Zoologie Philosophique had subsequently been grossly misrepresented and distorted, mainly as the result of the much publicized and abominably petty eulogy which Cuvier (who had no love for Lamarck) read before the French Academy shortly after Lamarck’s death. Cuvier had given the impression that Lamarck believed that animals could acquire body adaptations by striving for them or (as Cuvier said) for the wishing of them. The critical word in Lamarck’s French is ‘besom’ not ‘ ’ and, as Cannon pointed out,what Lamarck really said was that if new organs were needed they would, in the course of evolution arise, and this seems to be self-evident when we view the variety and environmental fittedness of animal adaptations. And, as Cannon further goes on to show, Lamarck’s own theory of the inheritance of acquired characters is thereby rendered nugatory, for if an animal acquires a character in response to a particular environmental need and the environment subsequently changes so, by Lamarck’s own requirements, will the adaptation change.

Cannon’s views on evolution must be respected both for their critical re-appraisal of Lamarck’s original statements and for his indication of the failure of classical genetics to provide an explanation of the origin of the kinds of phenotypic variation that are of evolutionary significance. What he perhaps fully failed to realize or to take account of were the remarkable advance^ which the geneticists themselves were making in the fields of molecular biology and microbial genetics, and which were already beginning to provide the explanations which he thought to be lacking. The following passage is taken from the most recent Leeuwenhoek Lecture (Pontecorvo 1962). ‘Thus mutation . . . is a non-adaptive process, or, in other words, the structure of the genetic material is not subject to regulatory change. It is even clearer, however, that the expression of the genetic material, both in micro-organisms and in higher animals and plants, is subject to regulation—qualitative and quantitative—at all levels of organization; the chromosome, the nucleus, the cell, the tissue, and even the whole organism in relation to its environment. A most important consequence of regulation of gene expression is that cells of identical genotype . . . can have different properties as the result of different previous experience. This difference in properties may even be maintained for an indefinite number of cell generations, though the difference in experience which caused it has been transitory.’ These, though the sentiments of Lamarckism, are the facts of genetics.

Forthright and uncompromising in the defence of his beliefs, quick to react to criticism and not giving to pulling his punches, Graham Cannon was rarely free from the dust and combat of scientific controversy. Yet he never harboured resentments; he was by nature warm-hearted and generous, a good companion and a loyal friend. He loved things that are beautiful and satisfying and he made his main hobby the collection of the elegantly worked Japanese swords and furnishings on which he spoke and wrote with authority. But his deepest contentment was found in the enjoyment of nature and of the simple things of everyday experience. Among his miscellaneous papers there





were found some notes of a talk he gave at a girls’ school at St Helens. He concludes his talk with this self-revelation. ‘You are blessed (he says) with certain senses, you can feel, you can smell, you can see, you can hear and you can taste, and in the exercise of all these senses you can experience beauty. The feeling of a little child’s hand grasping yours, the smell of the first narcissus of spring, the sight of the sun breaking through dark clouds on a stormy day, the sound of shingle pounded by the waves on a beach, the taste of a ripe gooseberry eaten straight off the bush—all these are good.’ And he found them so.

I wish gratefully to acknowledge the valuable assistance which I have received in the preparation of this Memoir from Mrs Cannon and from the many friends from whom I have sought advice. I am particularly indebted to Professor Ralph Dennell, Mr Gordon Blower and Dr Sidnie M. Manton, F.R.S., each of whom furnished me with some of Professor Cannon’s notes and papers. Mr Blower also very kindly prepared the Bibliography.

J. E. Smith

65

BIBLIOGRAPHYPublications on cytology

1920. (With D oncaster, L.) On the spermatogenesis of the louse corporis andP. capitis) with some observations on the maturation of the egg. Quart. micr. Sci. 64, 303-328, pi. 15.

1921. The early development of the summer egg of the cladoceran ( vetulus).Quart. J . micr. Sci. 65, 627-642, pi. 25.

1922. A further account of the spermatogenesis of lice. Quart. J . micr. Sci. 66, 657-667.1923. On the nature of the centrosomal force. J. Genet. Cambridge, 13, 47-78.1923. Spermatogenesis of the Lepidoptera. Nature, Lond. I l l , 670-71.

Publications on the Crustacea

1922. On the labral glands of a cladoceran {Simocephalus vetulus) with a description of itsmode of feeding. Quart. J. micr. Sci. 66, 213-234, pis. ix, x.

1923. A note on the zoea of a land-crab Cardisoma armatum. Proc. £ool. Soc. Lond. 94, 11-14.1924. On the development of an estherid crustacean. Phil. Trans. 212, 395-430, pis. 18-24.1925. Ectodermal muscles in a crustacean. Nature, Lond. 115, 458-459.1925. On the segmental excretory organs of certain fresh-water ostracods. Phil. Trans. B,

214, 1-27, pis. 1 and 2.1926. On the feeding mechanism of a freshwater ostracod, Pionocypris vidua (O. F. Muller).

J.Linn. Soc. (£oo/.), 36, 325-335, pi. 11.1926. On the post-embryonic development of the fairy shrimp ( diaphanus).

J. Linn. Soc. {Zool.), 36, 401-416, pis. 22, 23. (Awarded the Crisp Medal by the Linn. Soc. London, j'. Linn. Soc. Z°°t- 36.)

1927. (With M anton, S. M.) On the feeding mechanism of a mysid crustacean Hemimysislamornae. Trans. Roy. Soc. Edinb. 55, pt. 1, 219-253, pis. I-IV.

1927. On the feeding mechanism of Nebalia bipes. Trans. Roy. Soc. Edinb. 55, pt. II, 355-370.



1927. (With Manton, S. M.) Notes on the segmental excretory organs of Crustacea. Pts.I-IV. J.Linn. Soc. (Z°ol-)> 36,439-456.

1928. On the feeding mechanism of the fairy shrimp, diaphanus Prevost.Roy. Soc. Edinb. 55, 807-822.

1928. On the feeding mechanisms of the copepods and Diaptomus gracilis.Brit.J. Exp. Biol. 6, 131-144.

1929. (With M anton, S. M.) On the feeding mechanism of the syncarid Crustacea. Trans.Roy. Soc. Edinb. 56, Pt. 1, 175-189.

1931. On the anatomy of a marine ostracod Cypridina ( ) levis (Skogsberg). DiscoveryReports, 2, 435-482, pis. VI and VII.

1931. On the blood system of Parabathynella malaya G. O. Sars. Ann. Mag. Nat. Hist. (10), 8, 109-114.

1931. Nebaliacea. Discovery Reports, 3, 199-222, pi. XXXII.1933. On the feeding mechanism of certain marine ostracods. Trans. Roy. Soc. Edinb. 57.

Pt. I ll, 739-764.1934. Feeding mechanism of the fairy shrimp. Nature, Lond. 133, 329.1933. (With Leak, F. M. C.) On the feeding mechanism of the Branchiopoda. (By H.G.C.

with an appendix on the mouth parts of the Branchiopoda by H.G.C. and F.L.) Phil.Trans. B, 222, 267-352.

1935. On the rock-boring barnacle Lithotrya valentiana. Sci. Rep. Gr. Barrier Reef Exped. 1928-1929, London (Trustees Brit. Mus.), 5, no. 1, 1-17, 2 pis.

1935. Function of the labral glands in Chirocephalus. Nature, Lond. 136 758.1935. A further account of the feeding mechanism of Chirocephalus diaphanus. Roy.

Soc. B, 117, 455-470, pi. 24.1938. On the term ‘Gnathobase’ (Lankester), a correction. Proc. <oo/. Soc. Lond. B. 107.

539-541.1940. Ostracoda. Sci. Rep. \John Murray’ Expedn. 1933-34, 6, no. 8, 319-325.1940. On the anatomy of Gigantocypris mulleri. Discovery Reports, 19, 185-244, pis. XXXIX-

XLII.1946. Nebaliopsis typica. Discovery Reports, 23, 213-222, pi. XV.1947. On the anatomy of the pedunculate barnacle Lithotrya. Phil. Trans. B, 233, 89-136.1960. Leptostraca. In Bronn's Klassen und Ordnung deTierreichs. Bd. 5, Abt. 1. 4 Buch. 1 Teil.

pp. 1-81. Leipzig.


Publications on university teaching

1948. Undergraduate zoology. Advancement of Science. 5, 194-204.1948. The place of biology in a curriculum. Nature, Lond. 162, 1006. (Part of an address at a

joint session of Sections D and L of the B.A. meeting at Brighton on 14 September 1948.)

1949. The teaching of biology in schools. Nature, Lond. 163, 577.1949. University entrance requirements and the new examinations. Journal of Education.

April.1949. The teaching of biology in schools. Nature, Lond. 163, 846.1949. University awards. Journal of Education. May.1949. University education in natural science. Nature, Lond. 163, 882.

Publications on genetics and evolution

1954. Is the problem of evolution solved? School Sci. Rev. 35, 232-236.1956. An essay on evolution and modern genetics. J . Soc. (Zool.), 43, 1-17.1957. What Lamarck really said. Proc. Linn. Soc. Lond. 168, 70-85.1958. Opening address. Joint Discussion with the Systematics Association on ‘The inheri

tance of acquired characters’. Proc. Linn. Soc. Lond. 169, 41-45.1958. The evolution of living things. Manchester Univ. Press.



Herbert Graham Gannon 671959. Lamarck and modern genetics. Manchester Univ. Press, 1959. (Supplement, Chapter

VIII "The DNA racket’, 1963.)1959. Darwin and Lamarck. Their roles in the history of science as seen in the centenary

year 1959. A lain Currents in Modern Thought, pp. 106-110.1960. The myth of the inheritance of acquired characters. The New Scientist, 7, 798-800. 1960. Lamarck. In Collier's Encyclopedia, New York.

Other publications1924. (With Baylis, H. A.) A new trematode from the grass snake. Ann. Mag. Nat. Hist. (9),

13, 194-196, pi. VI.1924. (With Baylis, H. A.) Further note on a new trematode from the grass-snake. Ann.

Mag. Nat. Hist. (9), 13, 558-559.1927. (With Grove, A. J.) An aerating and circulating apparatus for aquaria and general

use. J. Roy. Micr. Soc. 47, 319-322.1936. A method of illustration for zoological papers, pp. x + 36. Norwich: The Association of

British Zoologists.1937. A new biological stain for general purposes. Nature, Lond. 139, 549.1941. A note on fine needles for dissection. J. Roy. Micr. Soc. 61, 58-59.1941. On chlorazol Black E and some other new stains. J. Roy. Micr. Soc. 61, 88-94.1950. The technique of biological staining. Endeavour, 9, 188-195.1953. William Thomas Caiman. 1871-1952. Obit. Not. Roy. Soc. 8, 355-372.

Letters to the press (mainly on education)1947. More graduates. Manchester Guardian. 16 May 1947.1948. New certificate examination. University entrance requirements. The Times. 16 Oct.

1948.1948. University entrance requirements. The Times. 11 Dec. 1948.1951. University standards. The Times. 20 Nov. 1951.1953. Disadvantages of specialization. The Times. 7 Jan. 1953.1954. The perfect athlete. Manchester Guardian. 26 May 1954.1955. The teaching of history. The Times. 7 Jan. 1955.1955. Illiterates. Manchester Guardian. 10 Oct. 1955.1955. Scientific language. Manchester Guardian. 21 Nov. 1955.1956. The shortage of science masters. Manchester Guardian. 10 Feb. 1956.1956. Recovery from poliomyelitis. Courage alone not the answer. The Times. 25 May, 1956.1956. Cambridge science tripos. Alanchester Guardian. 30 June, 1956.1956. The science tripos. Manchester Guardian. 9 July, 1956.1957. Specialization at school. Manchester Guardian. 8 Jan. 1957.1957. Specialization in the sixth form. The Times. 25 Jan. 1957.1957. Specialization at school. Manchester Guardian. 25 Jan. 1957.1957. Britains two cultures. Sunday Times. 31 March, 1957.1957. A most specialized creature. The Times. 10 April, 1957.1958. The misuse of penicillin. Manchester Guardian. 27 Jan. 1958.1958. Use of penicillin. Alanchester Guardian. 4 Feb. 1958.1958. Origin of species. The Times. 21 May, 1958.1958. The idea of evolution. Manchester Guardian. 16 June, 1958.1958. The idea of evolution. Darwin and Lamarck. Manchester Guardian. 26 June, 1958.1958. How the giraffe got a long neck. Alanchester Guardian. 10 July, 1958.1959. Language requirements. The Times, 8 June, 1959.1959. Oxford and Cambridge scholarships. Aianchester Guardian. 23 June, 1959.1959. Quiz programmes and education. Manchester Guardian. 27 Oct. 1959.1959. Balance sheet. The Times. 21 Nov. 1959.1959. Basis of life. Manchester Guardian. 10 Dec. 1959.1960. The future of man. The Listener. 21 Jan. 1960.



I960. The professor’s job. The Guardian. 4 Feb. 1960.1960. Specialization in the sixth. The Guardian. 22 Feb. 1960.1960. (Wider view of learning.) The Times. 27 May, 1960.1960. Mass graduation. The Times. 28 Dec. 1960.1961. Genetics at the universities. The Times. 1 Aug. 1961.1961. The universities’function. The Guardian. 14 Dec. 1961.1961. Animals on Tristan da Gunha. The Times. 19 Oct. 1961.1961. Sir Bertram Jones. An adventurous life. The Times. 20 Oct. 1961.1962. Education in depth. The Times. 7 Sept. 1962.




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