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REFERENCES

BELLOT, H. H. (1929). University College London, 1826-1926. London: University of London Press. CRANEFIELD, P. F. (1974). The Way In and the Way Out. Frangois Magendie, Charles Bell, and the Roots of the Spinal Nerves. New York: Futura Pub. Co. FRENCH, R. D. (1975). Antivivisection and Medical Science in Victorian Society. Princeton University Press. HoLMEs, F. L. (1974). Claude Bernard and Animal Chemistry. Cambridge, Mass: Harvard University Press. SHARPEY-SCHAFER, E. (1927). History of the Physiological Society during its First Fifty Years, 1876-1926. Supplement to J. Physiol. London: Cambridge University Press.

The experimental notebooks of Claude Bernard BY J. T. FITZSIMONS The Physiological Laboratory, Cambridge Claude Bernard occupies a unique place in the history of British Physiology. Not only was he one of the greatest experimental physiologists ever, but his work and the opposition it aroused in some quarters were powerful influences in the foundation of the Physiological Society. He had a profound belief in the value and necessity of animal experimentation in physiological research and he was a superb practitioner in the art of vivisection. He was born on 12 July 1813 in the small village of Saint-Julien en Beaujolais, the son of the proprietor of a small vineyard. After receiving his schooling from the Jesuits at Villefranche he worked for a time as an assistant to a pharmacist in Vaise, a suburb of Lyon, where he wrote a vaudeville, La Rose du Rhone, which achieved a certain success. Encouraged by this he wrote a drama, Arthur de Bretagne, and deciding on a literary career he left Lyon for Paris in 1833. There an unusually percipient critic, Saint-Marc Girardin, Professor at the Sorbonne, to whom he showed his manuscript dissuaded him from becoming a professional writer and suggested that his experience in pharmacy fitted him for a career in medicine. Bernard followed this advice and became a medical student in November 1834. Arthur de Bretagne was eventually published, but posthumously, in 1887. One wonders what sort of writer Bernard might have become because, much later, the classical scholar Guillaume Patin, Secretary of the French Academy, in his speech of welcome to Bernard on the latter's reception by the Academy, said of the famous Introduction a l'etude de la medecine experimentale, 'You have created a style', so impressed was he by the elegance of the writing. As a student Bernard was hard working and thoughtful but not obviously outstanding. In 1839, in his fifth year of medical studies, he was

38P PROCEEDINGS OF THE placed 26th out of 29 successful candidates in the 'internat', but his skill at dissection had already attracted the attention of Magendie, who had used several of Bernard's anatomical preparations for his course at the College de France that same year. As an 'interne des hopitaux a Paris' Bernard worked under various chiefs, including Magendie at the HotelDieu, and he was also persuaded by Magendie to become his preparateur at the College de France in 1841. In May 1843 appeared Bernard's first paper, 'Recherches anatomiques et physiologiques sur la corde du tympan', and in December of that year he obtained the M.D. degree. Increasing personal strains between the authoritarian Magendie, an empirical experimentalist with an aversion for systematization of ideas, and Bernard, a gentler and more reflective person who looked for a rational ordering of the experimental facts, led to Bernard leaving Magendie's laboratory at the College de France in 1844 though later both men came to respect and understand one another. This event, his failure in the agregation of 1844 and increasing financial problems discouraged Bernard to such an extent that he was preparing to abandon physiology and return to Saint-Julien in order to practise medicine as a country doctor. Then followed one of the most fateful episodes in his life. Alarmed by his decision to leave Paris, Bernard's friends, among them the chemist Theophile Jules Pelouze (who gave Bernard curare for his experiments in the early summer of 1844), conceived the plan of his marrying a girl with a sufficient dowry to ensure his future material wellbeing. A bride was found, Marie-Frangoise Martin (known as Fanny), aged 26, the daughter of a medical practitioner, and the marriage took place in July 1845, Magendie's name appearing with those of Pelouze and the toxicologist Orfila on the marriage contract. The union was unhappy but it kept Bernard in Paris. Bernard now immersed himself in experimental work. In 1847 he was appointed Magendie's supplant or deputy at the College de France and in the same year he learned that the Academie des Sciences had awarded him the prize for experimental physiology for the year 1845. He later received the 1847 prize for his work on the digestive function of the pancreas. He was present at the meeting in May 1848 at which the Societe de Biologie was founded and he became one of its two first vice-presidents, and later its president. At about this time he abandoned any thought of practising medicine and decided to devote himself exclusively to laboratory work. This was the start of his marital problems. Madame Claude Bernard, her outlook limited by her bourgeois upbringing, wanted to enjoy the rewards of being married to a successful medical practitioner, and she was well aware that her husband had the ability to be eminently successful at this; whereas Bernard wished for nothing more than to pursue his

PHYSIOLOGICAL SOCIETY, JULY 1976 39P experiments in peace in the laboratory. Another source of friction was that she had a passion for dogs whereas he, she said, tortured them. She and their two daughters (the two sons died in infancy) became ardent and vociferous antivivisectionists. This unhappy menage finally broke up in 1869. Bernard's research activity was prodigious and far-reaching, and this was fully recognized and amply rewarded. In the dozen years between 1847 and 1859 he established a digestive role for the pancreas, discovered the glycogenic function of the liver and the placenta showing for the first time that animals as well as plants can synthesize complex substances, described piqufre diabete, discovered vasomotor mechanisms for the regulation of the blood supply to different parts of the body, found that CO displaced oxygen from the blood, and showed that curare affects neuromuscular transmission. Among the many visitors to Bernard's laboratory in these years were Henry Bowditch and Silas Weir Mitchell from the United States and John Scott Burdon-Sanderson from England. In 1854 Bernard was appointed to the Chair of General Physiology at the Sorbonne and the same year he was elected member of the Academie des Sciences. When Magendie died in 1855 he became Professor of Medicine at the College de France. It was about now that the concept of the milieu interieur was beginning to be formulated which was to occupy Bernard to the end of his days. He was elected to the Academie de Medecine in 1861. In 1868 he joined the ranks of the immortals when he was elected to the Academie Frangaise, to the seat previously occupied by Flourens, though he did not actually take his seat until 27 May 1869. In 1868 he also vacated his chair at the Sorbonne in favour of his preparateur Paul Bert and was appointed Professor of General Physiology at the Museum d'Histoire Naturelle. He was made a Senator by Imperial decree in 1869 but the war of 1870 and the fall of the empire deprived him of this office shortly afterwards. During this period of intense activity and increasing fame Bernard suffered from indifferent health. He had a form of chronic enteritis which almost killed him in 1865, and which obliged him to spend much of the time from 1862 onwards away from Paris at Saint-Julien. He had been accustomed to spend his vacations towards the end of each summer at Saint-Julien and he had acquired a permanent residence there in 1861 near the parental farm house. He loved the countryside and took an active interest in wine production on his estate. It was here in the country away from the laboratory that Bernard had time for meditation and reflexion and what emerged was one of the most influential works in physiology ever, Introduction a l'etude de la medecine expe'rimentale, published in 1865. In it the aims and methods of physiological inquiry were set down and

40P PROCEEDINGS OF THE clarified with examples drawn from the experimental work of Bernard and others. According to Bernard the only way in which the function of living organisms can be understood is through properly conducted experiments devised to test particular hypotheses. He later (1867) said, 'One cannot understand what one finds unless one knows what one is looking for'. Rationally conceived experimentation is superior to empirical observation and to the haphazard collection of facts such as practiced by Magendie, who himself described his activities as those of a 'chiffonnier' - Magendie was being less than fair to himself though there was an element of truth in the description. Nor is deduction of function from anatomy enough. Bernard exorcised the chimeras of vital force and the capriciousness of nature from physiology and replaced them by determinism - the theory that vital processes are determined by physicochemical conditions. In the Introduction and also in the Rapport sur les progress et la marche de la physiologic generate en France (1867) Bernard was at pains to assert the independence of physiology and its need for laboratories and resources. He saw the danger of physiological problems being investigated only from the point of view of the physicist, chemist or engineer. He also defined the relationship between physiology and clinical medicine: '. . physiology is the basis of scientific medicine, because it must give the explanation of morbid phenomena showing the relationships that they have with the normal state' (1865). Bernard's championship of physiology would seem to have particular relevance today. After the final rupture with his wife in 1869 the last nine years of Bernard's life was gladdened by his relationship with a strikingly beautiful and cultivated young woman, Marie Sarah Raffalovich, who became his friend and sometimes his collaborator and with whom he kept up a voluminous correspondence. Bernard's letters to Madame Raffalovich are the autobiography of a sensitive and perceptive man who had experienced suffering, physical as well as in his family life. His personality inspired Smile Zola to write Docteur Pascal. Claude Bernard gave his last lecture on 28 December 1877 at the College de France and he died from pyelonephritis on 10 February 1878 in his appartment, 40 rue des Pcoles, just opposite the main entrance of the College. His prestige had been such that the Chamber of Deputies voted him a state funeral, the first scientist to be so honoured. A number of Claude Bernard's notebooks, generously lent by the College de France, were exhibited including the manuscript which served as the basis for the Introduction a' le'tude de la medecine experimentale.

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REFERENCES

BERNARD, CLAUDE (1865). Introduction d l'ttude de la Medecine Experimentale. Paris: Bailliere. BERNARD, CLAUDE (1867). Rapport 8ur lea Progr"s et la Marche de la Phyaiologie Generale en France. Paris: L'Imprimerie imperiale. FOSTER, M. (1899). Claude Bernard. London: Fisher Unwin. GRMEK, M. D. (1967). Catalogue de8 Manuacrita de Claude Bernard. Paris: Masson. OLMSTED, J. M. D. & OLMSTED, E. H. (1952). Claude Bernard and the Experimental Method in Medicine. New York: Schuman. Philo8ophie et m6thodologie Wcientifiquem de Claude Bernard. Colloque international organism pour la celebration du Centenaire de la publication de l'Introduction 'a l'etude de la m6decine experimental de Claude Bernard (1965). Paris: Masson.

T. H. Huxley and the development of Physiology in Britain By A. F. HUXLEY. Department of Physiology, University College London, Cower Street, London T. H. Huxley (1825-95) is well known as a comparative anatomist and paleontologist, as an exponent of biological science, especially evolution, and as a major influence in the development of scientific education, but his contributions to Physiology are not widely appreciated. His interest in physiology began early, as is made clear in the brief Autobiography that he wrote in 1889 (de Beer, 1974, p. 103): As I grew older, my great desire was to be a mechanical engineer, but the Fates were against this; and, while very young, I commenced the study of Medicine under a medical brother-in-law. But, though the Institute of Mechanical Engineers would certainly not own me, I am not sure that I have not, all along, been a sort of mechanical engineer in partibul infidelium. I am now occasionally horrified to think how very little I ever knew or cared about Medicine as the art of healing. The only part of my professional course which really and deeply interested me was Physiology, which is the mechanical engineering of living machines; and, notwithstanding that natural science has been my proper business, I am afraid there is very little of the genuine naturalist in me. I never collected anything, and species work was always a burden to me; what I cared for was the architectural and engineering part of the business, the working out the wonderful unity of plan in the thousands and thousands of diverse living constructions, and the modifications of similar apparatuses to serve diverse ends.

He entered medicine chiefly because two of his brothers-in-law were doctors. He joined the Navy with no thought of a scientific career but was picked by Sir John Richardson, the former Arctic explorer who was then his chief at Haslar Hospital, for the post of assistant surgeon, with opportunities for scientific work, on the Rattlesnake, which was to make surveying voyages off the eastern and northern coasts of Australia in the years 1847-50. During these voyages he made numerous important discoveries in the comparative anatomy of marine organisms, for which he was elected

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to the Royal Society in 1851 and awarded a Royal Medal in 1852. He left the Navy in 1853 when he was allowed no further time for writing up his researches, and tried without success to obtain a position in either Physiology or Comparative Anatomy. In the end, he obtained a post at the Royal School of Mines, of which he writes in his Autobiography (p. 108): At last, in 1854, on the translation of my warm friend, Edward Forbes, to Edinburgh, Sir Henry De la Beche, the Director-General of the Geological Survey, offered me the post Forbes vacated of Paleontologist and Lecturer on Natural History. I refused the former point blank, and accepted the latter provisionally, telling Sir Henry that I did not care for fossils, and that I should give up Natural History as soon as I could get a physiological post. But I held the office for thirty-one years and a large part of my work has been paleontological.

Although Huxley never did move to a physiological post, he gave numerous courses of lectures in straightforward physiology. For example, in 1856-7 he lectured on Physiology and Comparative Anatomy at the Royal Institution, where he held the post of Fullerian Professor from 1855 to 1859 and from 1865 to 1869. Another course of lectures at the Royal School of Mines in 1863 formed the basis of his book Lessons in Elementary Physiology, published by Macmillan in 1866, in the 'School Class Books Series'. According to the Preface, it was 'primarily intended to serve the purpose of a text-book for teachers and learners in boys' and girls' schools'. It was reprinted on average about once a year for the rest of the century, later editions being revised by Michael Foster. A sixth edition considerably enlarged, was produced by Joseph Barcroft in 1915. Even the original edition contains much more than was in most biology 'A level' syllabuses until a few years ago. Huxley gave regular courses of lectures in Physiology for schoolteachers at South Kensington after the School of Mines had moved there from Jermyn Street in 1872 as the nucleus of what became the Royal College of Science and, finally, in 1907, the Imperial College of Science and Technology. His notes for some of these lectures are preserved with his other papers at Imperial College. It must not be supposed that because Huxley did no original research in Physiology, these lectures were based on second-hand knowledge: His regular lectures involved an immensity of labour, for he would never make a statement in them which he had not personally verified by experiment. In the Jermyn Street days he habitually made preparations to illustrate the points on which he was lecturing, for his students had no laboratory in which to work out the things for themselves. His lectures to working-meD also involved as much careful preparation as the more conspicuous discourses at the Royal Institution [L. Huxley,

1908].

Britain made almost no contribution to the tremendous progress in Physiology that took place between, say, 1840 and 1880. Apart from a few names like Claude Bernard and Brown-Sequard, almost all the leading

43P PHYSIOLOGICAL SOCIETY, JULY 1976 figures in this advance were German. It was clear to Huxley that the main reasons for the failure of Britain to participate in this advance were that full-time posts in physiology hardly existed and that most of the teaching of physiology in medical schools was performed as a side-line by clinicians. His address to the medical students of University College London, after he had acted as their examiner in 1870, contains the passage (T. H. Huxley, 1893, p. 315): I get every year those very elaborate reports of Henle and Meissner - volumes of, I suppose, 400 pages altogether - and they consist merely of abstracts of the memoir and works which have been written on Anatomy and Physiology - only abstracts of them! How is a man to keep up his acquaintance with all that is doing in the physiological world - in a world advancing with enormous strides every day and every hour - if he has to be distracted with the cares of practice? You know very well it must be impracticable to do so. Our men of ability join our medical schools with an eye to the future. They take the Chairs of Anatomy or of Physiology; and by and by they leave those Chairs for the more profitable pursuits into which they have drifted by professional success, and so they become clothed, and physiology is bare. The result is, that in those schools in which physiology is thus left to the benevolence, so to speak, of those who have no time to look to it, the effect of such teaching comes out obviously, and is made manifest in what I spoke of just now - the unreality, the bookishness of the knowledge of the taught.

Huxley was not content to diagnose and lament the situation, but played an important part in the first two effective steps that were taken to correct it. The first of these was the appointment by Trinity College, Cambridge, of a Praelector in Physiology, whose duties were to give lectures in Physiology to students not only of Trinity but of the other colleges, and to organize laboratory work (Sharpey-Schafer, 1927, p. 4). Huxley's share in the appointment is described thus by Bibby (1959, p. 183): In 1866 he was conferring with Sidgwick about the best person for a Praelectorship in Natural Science which it was hoped to establish at Trinity, but the proposal was defeated at the December College Meeting. Soon, however, with W. H. Thompson as Master, Trinity moved to the forefront of Cambridge reform, and in the spring of 1870 Huxley heard from a Fellow that there had been a more positive response to another such proposal: 'I read your letter to the Seniority yesterday. Your suggestion as to the Praelectorship of Physiology and the man to fill it, was most favourably received ... If Mr Foster will come and spend a day with me, I can explain everything to him ... If he is appointed, we will set to work about establishing the Physiological Laboratory.' His nominee once established in the new post. Huxley had a useful channel through which to influence science teaching at Cambridge, and for the next quarter of a century Michael Foster never ceased to consult him.

In 1877, a Chair of Physiology was established by the University, and Huxley was one of the electors. Michael Foster was the first occupant. The second major step in the establishment of Physiology in Britain was

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the endowment of a full-time Chair of Physiology at University College London by Mr T. J. P. Jodrell. Again, the influence of Huxley, who had known Jodrell for many years, was a major factor. To quote Bibby again

(p. 216): Three years after telling the medical faculty that its professors should be so remunerated as to abstain from private practice, he learned that T. J. P. Jodrell, a wealthy amateur of science, was prepared to provide an endowment of £7500. Following consultation with Huxley, Jodrell wrote to tell the President of University College that he was advised that the work of the two part-time professors in physiology might be given as efficiently by one full-time professor with the aid of a demonstrator, and he explained that the purpose of the proposed endowment was to induce men of eminent ability to forgo more lucrative sources of emolument and devote themselves so original research. At the first the College was reluctant to accept the rather stringent conditions which were attached to the offer, so Jodrell consulted with Huxley again, with an eminently satisfactory outcome: 'Since I saw you I have had another interview with the U.C. Deputation at which being fortified by yr. opinion I assumed a more confident tone & the result was that they acquiesced at once in my view.' The intention of the endowment was admirably fulfilled by the first Jodrell Professor of Physiology, Burdon-Sanderson; and, when a few years later the founder's wish to do something more for science took shape at Huxley's suggestion in the Jodrell Chair of Zoology and Comparative Anatomy, the first occupant was that well-loved prodigal protege, Ray Lankester.

Huxley was a member of the Royal Commission on animal experiments, 1875-6. He was the only member who was an active scientist, and we owe it to him that the Act of 1876 was not a great deal more restrictive than in fact it was. He was a member of the committee that started the Physiological Society and a founder-member of the Society itself. Finally, a quotation from his Rectorial Address at the University of Aberdeen in 1874 will show his appreciation of the educative value of a study of physiology (T. H. Huxley, 1893, p. 220): Moreover, I would urge, that a thorough study of Human Physiology is, in itself, an education broader and more comprehensive than much that passes under that name. There is no side of the intellect which it does not call into play, no region of human knowledge into which either its roots or its branches, do not extend; like the Atlantic between the Old and the New Worlds, its waves wash the shores of the two worlds of matter and of mind; its tributary streams flow from both; through its waters, as yet unfurrowed by the keel of any Columbus, lies the road, if such there be, from the one to the other; far away from that North-west Passage of mere speculation, in which so many brave souls have been hopelessly frozen up. REFERENCES

DE BEER, G. (1974). Charles Darwin, Thomas Henry Huxley: Autobiographies. Edited with an introduction by G. DE BEER. London: Oxford University Press. BIBBY, C. (1959). T. H. Huxley, Scientist, Humanist and Educator. London: Watts. HUXLEY, L. (1908). Life and Letters of Thomas Henry Huxley, 2nd ed., vol. 3, p. 385. London: Macmillan.

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HuxLEY, T. H. (1893). Science and Education: Collected Essays, vol. 3, London: Macmillan. SHARPEY-SCHAFER, E. A. (1927). History of the Physiological Society during it First Fifty Years, 1876-1926. London: Cambridge University Press.

George Henry Lewes, George Eliot and the Physiological Society BY D. NOBLE. University Laboratory of Physiology, Oxford The minutes of the first meetings of the Physiological Society in 1876 are signed by G. H. Lewes. He was a highly respected and much valued member of the Society. How did he come to occupy this position? It is well known that he lived with George Eliot from 1854 until his death in 1878. All of George Eliot's major novels were published during this period. Haight (1954) writes: 'In the transformation of Marian Evans into a great novelist Lewes had a major part. He helped her most by maintaining her self-confidence, which even at the height of her fame required constant bolstering. The ostracism they knew would follow their deliberate violation of convention was hard for one of her nature to bear.' Their 'violation of convention' was, of course, their living together. The manner of their coming together was wide open to scandalous interpretation. George Eliot was thought to be deeply involved with Herbert Spencer. Spencer certainly thought highly of her. To a friend he described her in 1852 as 'the most admirable woman, mentally, that I ever met'. However, their relationship was limited to a close friendship. George Eliot wrote: 'We have agreed that we are not in love with each other and that there is no reason why we should not have as much of each other's society as we like.' Her friendship with Spencer brought her into close contact with the main streams of liberal intellectual thought in Victorian England. Originally an evangelical Christian, she slowly made the painful transition away from her faith. She translated Strauss's Das Leben Jesu and, in so doing, produced a work (The Life of Jesus, 1846) that was important in the development of nineteenth-century rationalism. She was later to translate Spinoza's Ethics. In 1851 she became assistant editor of the Westminster Review. In 1852 Spencer used this periodical to publish his article on 'A theory of population 'in which he first linked the development of the species with the survival of the fittest. It was also at this time that G.H. Lewes was writing an influential series of articles (published in the Leader) on Comte's philosophy of the Sciences. The reaction to this series of articles was an important one for it drew G. H. Lewes into controversy with another founder of the Society: T. H. Huxley. I shall return to this controversy later.

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PHYSIOLOGICAL SOCIETY, JULY 1976 47P George Henry Lewes was married to Agnes Jervis in 1841. She was also a radical and helped Lewes in some of his French translations (notably of Balzac). Together with Thornton Hunt, they founded the Leader, a weekly periodical. Lewes had four sons by Agnes. In 1850 a fifth was born, but the father was Hunt. Lewes forgave the offence, but in 1851 another child was born, also fathered by Hunt. Lewes was trapped. Having condoned the first adultery his grounds for divorce were gone. His home was broken. It was about this time that he met George Eliot. The relationship developed slowly but, by 1853, Eliot wrote: 'Lewes has been quite a pleasant friend to me lately' (11 April 1853). Four days later she wrote to the same correspondent (Mrs Charles Bray) 'Mr Lewes especially is kind and attentive and has quite won my regard after having a good deal of my vituperation.' In 1854 they went to Germany together. Lewes was to collect material for one of his most important works, The Lyife and Works of Goethe, which was published in 1855. It was the first English biography of Goethe and was to remain the standard work for many years. On the 23 October 1854 George Eliot wrote from Weimar a letter to Charles Bray: It is possible that you have already heard a report prevalent in London that Mr Lewes has 'run away' from his wife and family. I wish you to be in possession of the facts which will enable you to contradict this report whenever it reaches you. Since we left England he has been in constant correspondence with his wife; she has had all the money due to him in London: and his children are his principal thought and anxiety. Circumstances, with which I am not concerned, and which have arisen since he left England, have led him to determine on a separation from Mrs Lewes, but he has never contemplated that separation as a total release from responsibility towards her. On the contrary he has been anxiously waiting restoration to health that he may once more work hard, not only to provide for his children, but to supply his wife's wants so far as that is not done by another. I have seen all the correspondence between them, and it has assured me that his conduct as a husband has been not only irreproachable, but generous and self-sacrificing to a degree far beyond any standard fixed by the world. This is the simple truth and no flattering picture drawn by my partiality. I have been long enough with Mr Lewes to judge of his character on adequate grounds, and there is therefore no absurdity in offering my opinion as evidence that he is worthy of high respect. He has written to Carlyle and Robert Chambers stating as much of the truth as he can without very severely inculpating the other persons concerned; Arthur Helps, who has been here since we came, already knew the whole truth, and I trust that these three [rational] friends will be able in time to free his character from the false amputations which malice and gossip have cast upon it. Of course many silly myths are already afloat about me, in addition to the truth, which of itself would be thought matter for scandal. I am quite unconcerned about them except as they may cause pain to my real friends. If you hear of anything that I have said, done, or written in relation to Mr Lewes beyond the simple fact that I am attached to him and that I am living with him, do me the justice to believe that it is false. Mr and Mrs Chapman are the only persons to whom I have ever spoken of his private position and of my relation to him, and the only influence I should ever

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dream of exerting over him as to his conduct towards his wife and children is that of stimulating his conscientious care for them, if it needed any stimulus. Pray pardon-this long letter on a painful subject; I felt it a duty to write it.

George Eliot does not exaggerate Lewes's generosity to his wife and children (see Haight, 1954). Let us return to 1853, when Lewes's relationship with George Eliot was still developing. On the 17 December 1853 George Eliot wrote a letter to the editor (John Chapman) of the Westminster Review, pleading with him that he should not print a review by T. H. Huxley of G. H. Lewes's articles on Comte. The letter is worth quoting in full for it reveals how much involved she was in Lewes's scientific work: PRIVATE Dear Friend May 1 beg that you will not send Mr Huxley's M.S. to the printer until you have seen me again? I have found out that he is in the wrong in his remark on the embryological doctrine at p. 33 of Mr Lewes'sbook, and also that the ridicule he throws on the remark about the gallionella ferruginea is not well founded. At all events I think you will wish for the sake of the Review as well as from your own sense of justice that such a purely contemptuous notice should not be admitted unless it be well warranted. The case is the more delicate as the criticism of Mr Lewes comes after the unmitigated praise of Miss Martineau. I hope to see you tomorrow afternoon. How came you to mention to Miss M. that you saw the proof of Mr Lewes's book 'in [my] Miss Evans's room?' I think you must admit that your mention of my name was quite gratuitous. So far you were naughty - but never mind.

It is also worth quoting Haight's (1954) comment on this letter: Huxley devoted more than 2 pages to an attack on the scientific errors in GHL's Comte's Philosophy of the Sciences. His suggestion (p. 225) that GHL was a mere book scientist 'without the discipline and knowledge which result from being a worker also' was particularly stinging. In his advocacy of the evolutionary hypothesis GHL was far in advance of Huxley at this time, his facility in languages having given him an acquaintance with the researches of Continental scientists of which Huxley knew nothing.* In an earlier skirmish GHL, reviewing an article by Huxley in the British and Foreign Medical Review questioned Huxley's position 'with the same freedom he has used towards Schleiden and Swann (Leader, 5 November 1853, pp. 1073-1075). Huxley's article was published unchanged. In a long signed letter to the Leader, 14 January 1854, p. 40, GHL replied that he is not a mere 'bookman', but has occupied himself with biology both practically and theoretically for more than eighteen years. Acknowledging one real error of fact in his book (sulphuric for sulphurous acid), he shows convincingly that the authorities Huxley relies on had often been superseded within the past two years by French scientists unknown to him. Huxley was later to come to admire Lewes. In 1859 Lewes presented * The implication that T. H. Huxley was not a good linguist is incorrect. He acquired fluent German as a boy and could read French without difficulty. His ignorance of the literature quoted by Lewes must have other causes.

49P PHYSIOLOGICAL SOCIETY, JULY 1976 three papers at the British Association: 'The necessity for a reform in nerve physiology', 'A demonstration of the muscular sense' and 'On the supposed distinction between sensory and motor nerves.' He was not present himself but he wrote in his diary. 'He [Beck] gave an amusing description of the 'row' they [the papers] excited and told me that Allen Thomson and Huxley both defended me. I expected the papers would create a disturbance, but hardly expected these eminent men to stand up for me.' Huxley in fact became a friend of the household. George Eliot wrote to a friend in 1866, 'It would be fortunate for us if you had nothing better to do than look in on us on Tuesday evening. Professor Huxley will be with us, and one or two others whom you know'. The 'one or two others' included Spencer. It appears that Huxley's view of Lewes changed when he read his Seaside Studies. On its publication, Huxley sent Lewes a complimentary letter and copies of his own papers. Huxley and others were to regret their initial view of Lewes as a 'book scientist'. In fact, he experimented a lot despite his prodigious efforts as a journalist and writer. In 1867 he was engaged on what he called 'long laborious researches into the Nervous System for my work on Psychology. I have had such illuminations on this subject, completely reshaping the whole scheme of nervous anatomy that I resolved to run over to Germany to inspect the preparations etc of the best men there, in the hope of finding bricks ready for my purposes.' A postscript to a letter written by George Eliot on the same day reads: 'Froggie continues to do better than even he expected without his head brain for months. He dies of starvation at last.' She must have watched a lot of his experiments and was fully aware of the issues when the vivisection controversy came to a head in the early 1870s. The reference to Psychology in Lewes's note is important. It is probable that, while reviewing Comte's work on the Philosophy of the Sciences, he introduced the science into a logical structure for the first time. In the last article published in the Leader on 14 August 1852 he wrote: 'I propose, therefore, to keep the Physical Sciences as Comte arranges them: and to introduce a new fundamental science - Psychology - as the basis of Sociology; that is to say, I begin the Science of Humanity with a preliminary science of Human Nature.' He was later to devote a considerable part of a major work, Problems of Life and Mind, to the study of Psychology, but this eventually involved Michael Foster (see below). Lewes's visit to Germany brought him into contact with virtually the whole of the very active German School. Haight (1956) lists the people he met. They included Boll, Pfluiger, Helmholtz and Kirchhoff. According to a letter of George Eliot he 'had a brilliant time, gained great instruction and has seen some admirable men who have received him warmly'. He

AOPPROCEEDINGS OF THE also found 'my Seaside Studies, and Physiology universally known to the men of science and the "Goethe " still continues to be read by everyone' (GHL's journal, 10 January 1868). Nevertheless, one must not overplay Lewes's experimental work. It remains true to say that his greatest contribution was to the intellectual development of Physiology and of Biology in general. In March 1868 he was asked to contribute a series of articles on Darwin to the Fortnightly Review. They appeared in April, June, July and November. Articles of this kind clearly won for him a deep respect amongst his scientific colleagues. The influence was also partly that of George Eliot herself. On 13 August 1868 he writes: 50P

Last week I spent at Oxford at the Medical Association and not the least of my pleasures was to hear the deeply reverential way in which men of eminence spoke of Mrs Lewes's genius and the evidence they gave that her work had profoundly entered into their souls. Paget, the great surgeon, and some lesser medical men spoke of her as the greatest genius - male or female - that we could boast of. The 'Gypsy' I find deeply affects all men except reviewers, and I pity the man it does not affect.

Actually, it is only fair to comment that The Spanish Gypsy, published in 1868, did not become one of George Eliot's major works. Much greater was to come, especially Middlemarch. George Eliot and G. H. Lewes were both avid readers of Darwin's work as it appeared. George Eliot's journal for 23 November 1859 reads: 'We began Darwin's work on The Origin of Species tonight. It seems not to be well written: though full of interesting matter, it is not impressive, from want of luminous and orderly presentation.' Faint damnation? One might expect her to be more concerned with the style but in fact she was also not all that moved by the content. In a letter (5 December 1859) she acknowledges that the work 'will have a great effect in the scientific world... but to me the development theory and all other explanations of processes by which things came to be, produce a feeble impression compared with the mystery that lies under the processes'. This quotation is important for it partly reveals that George Eliot and G. H. Lewes did not share similar opinions even on quite important matters. Haight (1954) qualifies his comments on the influence of Lewes on Eliot by writing But it is a mistake to believe that George Eliot's mind was deeply influenced by him. Every main bias had been taken before they met, and they respected each other too much to desire uniformity of opinion. Their attitude toward Positivism is a good example. Lewes, one of its earliest English advocates, had no sympathy whatever for the religion later formulated by Comte. George Eliot, on the other hand, who yearned always for the old faith she had lost, clung for years to the hope of finding a substitute for it in the Religion of Humanity. But 'I cannot submit my intellect or my soul to the guidance of Comte,' she told Sara Hennell. Though she never returned to orthodox Christianity, George Eliot was deeply religious.

PHYSIOLOGICAL SOCIETY, JULY 1976 51P Lewes had planned a review of Darwin's Origin of Species in 1859 but the publisher feared its controversial aspects and withdrew his offer to publish it. In the event, nothing appeared until 1865 when an article by Lewes in the Pall Mall Gazette became the book's 'first thoroughly appreciative and adequate welcome ... as Darwin always remembered' (Robertson-Scott, 1950). Following the articles in the Fortnightly Review in 1868, the Darwins became personally known to Lewes and Eliot. Thus by 1869, the Lewes household had not only lived out the scandal of its origins, it had become a social centre where the most eminent scientists and literary men of the period were to be found. Charles Eliot Norton gives us a revealing picture of the household in a letter written on 29 January 1869: The Leweses live in the St John's Wood district, not far from Regent's Park. Their house, called The Priory, is a little square two-storey dwelling standing in a half garden, surrounded with one of those high brick walls of which one grows so impatient in England. Lewes received us at the door with characteristic animation; he looks and moves like an old-fashioned French barber or dancing-master, very ugly, very vivacious, very entertaining. You expect to see him take up his fiddle and begin to play. His talk is much more French than English in its liveliness and in the grimace and gesture with which it is accompanied - all the action of his mind is rapid, and it is so full that it seems to be running over. 'Oh, if you like to hear stories', he said one day, 'I can tell you stories for twelve hours on end.' It is just the same if you like to hear science or philosophy. His acquirements are very wide, wider perhaps than deep, but the men who know most on special subjects speak with respect of his attainments. I have heard both Darwin and Sir Charles Lyell speak very highly of the thoroughness ofhis knowledge in their departments. In fact his talents seem equal to anything. But he is not a man who wins more than a moderate liking from you. He has the vanity of a Frenchman; his moral perceptions are not acute and he consequently often fails in social tact and taste. He has what it is hard to call a vulgar air, but at least there is something in his air which reminds you of vulgarity.

He also writes of George Eliot, The portrait of Mrs Lewes reminded me, not by its own merit, of Couture's drawing of George Sand - and there is a strong likeness to this drawing in her own face. The head and face are hardly as noble as George Sand's, but the lines are almost as strong and masculine; the cheeks are almost as heavy, and the hair is dressed in a similar style, but the eyes are not so deep, and there is less suggestion of possible beauty and possible sensuality in the general contour and in the expression. Indeed one rarely sees a plainer woman; dull complexion, dull eye, heavy features. For the greater part of two or three hours she and I talked together with little intermission. Her talk was by no means brilliant. She said not one memorable thing, but it was the talk of a person of strong mind who had thought much and who felt deeply, and consequently it was more than commonly interesting. Her manner was too intense, she leans over to you till her face is close to yours, and speaks in very low and eager tones; nor is her manner perfectly simple. It is a little that, or it suggests that, of a woman who feels herself to be of mark and is accustomed, as she is, to the adoring flattery of a coterie of not undistinguished admirers. In the course of the afternoon

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three or four men came in - the only one whom 1 knew was Professor Beesly. We came away just before sunset ... Every one who knows Mrs Lewes well seems attached to her, and those who know speak in the warmest terms of her relations to her husband and his family - of her good sense and her goodness.

On 15 December 1875 George Henry Lewes was examined before the Royal Commission on Vivisection. On 3 February 1876 George Eliot wrote in a letter to Mrs Bray: Mr Lewes gave evidence on the subject of Vivisection before the Committee of Inquiry, and has corrected the proofs for the Blue Book. He thinks it good that there should be a strong check from public opinion, but [thinks] that hardly any legislative measure on the subject can be taken without doing more harm than good. Your quotation from the 'Science Primer' shocks me very much. Mr Lewes knows Michael Foster, who is in many respects an admirable man - but men, like societies, have strange patches of barbarism in the midst of their 'civilization'. There are facts in almost everybody's habits for which they 'have eyes but see not'. Let us be thankful for any anointing that will make us see better. But one wishes that the advocates of good would be a little more scrupulous in their modes of advocacy.

The 'Science Primer' was Michael Foster's A Course of Elementary Practical Physiology. Following the Royal Commission, events moved very rapidly and very dangerously for the physiological world (see French, 1975). On 31 March 1876 there appears an entry in Lewes's diary that is so important that it should be quoted in full: Went to a conference of physiologists at Burdon Sanderson's: Huxley, Sharpey, Pavy, Marshall, Humphry, Ferrier, Lauder Brunton, etc. It was agreed to form a Physiological Society and I was asked to be one of the Council. Th& question was discussed of what steps should be taken with regard to the Vivisection Report, and it was agreed to let the Government quietly understand that we would not oppose a Bill framed in the spirit of that Report.

It was the minutes of this meeting that were signed by Lewes. Despite the seniority of Sharpey and the greater scientific reputation of Huxley, Lewes was clearly regarded as the eminence grise. Even Huxley and Darwin acknowledged his contributions to the intellectual growth of the subject. Lewes took the chair at two of the important early meetings and, at the inaugural dinner on 26 May 1876 he proposed the toast to 'Foreign Physiologists'. During the remaining two years of his life Lewes occupied himself with the work that he clearly thought would express the fullness of development of his own biological thinking. This was to be a five-volume work entitled Problems of Life and Mind. He was not able to finish it before he discovered in June 1878 that he was suffering from intestinal cancer. On 24 November 1878 George Eliot wrote: 'For the last week I have been in deep trouble. TMr Lewes has been alarmingly ill.' On the 25th she was 'absorbed in nursing him'. On 30 November he died. He had not been able to attend a meeting of the Society since January. At the meeting of 12 December,

PHYSIOLOGICAL SOCIETY, JULY 1976 53P 'It was unanimously resolved that a minute be entered to record the regret of the meeting at the death of one of the Society's original members, Mr G. H. Lewes.' The minute is signed by Francis Galton. Among the letters George Eliot received, there was one from Herbert Spencer. It is now kept in Balliol College Library: 5 December 1878 Athenaeum Club

Dear Friend I have hesitated these several days whether to write - feeling strongly how utterly vain are words. I can but dimly conceive what such a parting must be, even in an ordinary case. Still more dimly can I conceive what it must be in a case where two lives have so long bound together so closely, in such multitudinous ways. But I can conceive it with clearness enough to enable me to say, with more than conventional truth, that I grieve with you. ever yours sincerely, Herbert Spencer

This is a pained and formal letter but some of its significance becomes clear when one recalls Spencer's early relation with George Eliot, that he introduced her to Lewes and that he himself remained a bachelor. Lewes's death did not, however, end George Eliot's connexions with Physiology. First of all there were the manuscripts and proofs of Problems of Life and Mind to edit. She was helped in this task by Michael Foster. Volumes i-mn had already appeared. The volume that Foster helped with eventually appeared in 1879 under the title The Study of Psychology. Secondly, she decided to found a physiological studentship. She wrote on 20 February 1879 to Dr Clifford Allbutt. Can you help me with any suggestions as to founding (now while I live) some Lectureship or other efficient instrument of teaching Biology (including Psychology) in memory of my husband, and to be called by his name?

By 5 March the scheme had changed to that of a studentship and she was now being advised by Henry Sidgwick and Michael Foster. Foster visited George Eliot on 13 March and she writes: ... we arrived at a satisfactory clearness as to the conditions. He mentioned as men whom he had thought of as suitable trustees Huxley, Pye-Smith, Thistleton Dyer, F. Balfour and H. Sidgwick.

On 23 April she wrote to John Blackwood (her publisher) I am founding a studentship of Physiology to be called the George Henry Lewes Studentship. It will be placed in the first instance in Cambridge where there is the best physiological school in the Kingdom. But the trustees (with my consent during my life) will have the power of moving it where they judge best. This idea which I early conceived, has been a great stay to me.

By September the arrangements were all settled and the Studentship was advertised. The response was disappointing. In a postscript to a letter of 14 October 1879 she wrote

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There is a good student in view for the G.H.L. Studentship, but some of the applicants in answer to the advertisement were incredibly silly and presumptuous. But the real aspirants in physiology are so rare that every physiologist knows them all. It is the science least adequately studied in England.

On 18 October she nominated Dr C. S. Roy and he was duly accepted by the Trustees at their meeting on the 19 October. She wrote in December 1879 The Dr Roy who was elected to the Studentship is believed to be the most promising of our advanced physiologists in the generation that has its best days yet to come. This is a great satisfaction to me and the Studentship altogether is what I know my husband would have rejoiced in.

George Eliot did not live to choose another Student. She died a year later, on 22 December 1880. Portraits of Lewes and Eliot, Lewes's books and some of the correspondence referred to were on display at the Centenary meeting. REFERENCES FRENCH, R. D. (1975). Antivivisection and Medical Science in Victorian Society. Princeton, London: Princeton University Press. HAIGHT, G. S. (1954). The George Eliot Letters, volumes 1-7. Oxford University Press. ROBERTSON-SCOTT, J. W. (1950). The Story of the 'Pall Mall Gazette'.

E. Sharpey-Schafer and the jubilee of the Physiological Society BY A. V. HILL. 11a, Chaucer Road, Cambridge The history of the Physiological Society during its first fifty years is recorded in a supplement to the Journal of Physiology published in December 1927. According to Schafer (as he was then called), at a meeting on 31 March, 1876, with Burdon-Sanderson in the chair, a number of eminent physiologists and others met and decided to form a physiological society. Later that year 'the actual inaugural meeting was appropriately celebrated in the customary English fashion by a dinner'. This was held at the Criterion Restaurant in London on 26 May 1876. Twenty-two members were present, with Michael Foster in the chair. I have calculated, with the aid of Whitaker's Almanac, that the day was a Friday; the dinners of the Society are still usually held on a Friday. The jubilee celebration would naturally have been held in 1926, but in consequence of the time occupied in the negotiations for the acquisition of the Journal of Physiology [from Mrs Langley], and consequent changes in the Constitution of the Society, it was not found possible to make arrangements for the jubilee celebration during 1926. This was accordingly deferred until the following year. The arrangements which were suggested included: (1) A banquet to be held in London on Saturday May 13th 1927... This was duly beld

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(2) A commemorative medal to be struck. This suggestion was not acted upon. (3) A history of the Society during the first fifty years of its existence to be published in the Journal of Physiology. Now carried out.

I was in America between January and June 1927 and was unable to attend the banquet; however, the minutes show it to have been a great success. The 1927 minute book containing the records of the jubilee dinner were on display at the Centenary meeting. Schafer later adopted the title, Sharpey-Schafer, in honour of his teacher Sharpey, and left University College London (where he had worked from 1874 to 1899) to become Professor of Physiology at Edinburgh, where he died in 1935. After his death a Sharpey-Schafer Memorial Lecture was founded in the University of Edinburgh, the ninth of which I gave in 1951. Schafer lost two sons in the war; in spite of which, when the Congress of Physiologists was held in Edinburgh in 1920 (under his Presidency), only two years after the war was over, he insisted that all ex-enemy physiologists should be invited to come. This decision was approved at a meeting of the Society in Manchester and many of the ex-enemies turned up at the Congress. One son, who was killed in a Q-boat, had been named Sharpey-Schafer, to emphasize, as he said, his 'indebtedness to one who inspired his early work, a great scientist and a staunch friend'. This tribute of piety and affection, alike to teacher and son, and many private acts of kindness and hospitality, were signs of an inner sensibility little evident to those who knew him less. His 'superb intolerance of intellectual dishonesty', as it has been called, his accurate and logical mind, his power of controlling the largest and most boisterous class (even in this [Edinburgh] University !), his unremitting labours in physiology, his toughness, his implacable antipathy to false sentiment - all these gave an almost alarming impression to a new and hesitant acquaintance. For years I failed to realize the other side. I accepted the apparent roughness as a minor defect of his great qualities of scientific integrity and executive competence. Once, however, he seemed to me so rude that I was provoked to answer him back - and from then we became good friends. Maybe the provocation was planned deliberately to that end: probably he never respected a man who would not stand up for himself: anyhow from that moment I came to think of him with affection. Only once have I seen him discomfited, by a joint conspiracy of Henry Dale and myself, which we used to recall with satisfaction. A dinner was held in his honour on his 80th birthday (2 June 1930). On previous occasions he had expressed disapproval of the committee for recommending too many Honorary Members - and what was worse, we gave them free copies of the Journal of Physiology. On that evening Dale proposed that he 5

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56P A6PPROCEEDINGS OF THE should be elected an Honorary Member, and I followed this up quickly by insisting that he should receive free copies of the Journal. He accepted both proposals with a friendly growl and admitted defeat. A final little story tells the quality of the man. In Edinburgh, in his days as a Professor there, a gun was fired at noon. No other professor could possibly continue his lecture after the gun had gone off: but Schafer was permitted by his students to complete his lecture without disturbance.

Early research on electrophysiology BY R. D. KEYNES. Physiological Laboratory, Cambridge CB2 3EG Although the disconcerting effects of contact with an electric fish were familiar to classical physicians such as Scribonius (Compositions, chapters 11 and 162), investigations of their origin cannot be described as electrophysiology before some concept of the nature of electricity had arisen. Hence the credit for performing the first electrophysiological experiment appears to be due to Adanson (1757), for in September 1751 he showed that shocks produced by Malapterurus could be conducted along a metal rod, and likened them to the discharge of a Leyden jar, which had been described only a few years earlier. Not long afterwards, Allamand (1758) reported similar observations made by s'Gravesande in South America on Electrophorus. The first detailed study of the discharge of an electric fish was made by Walsh (1773) on Torpedo, and was published in the form of a letter to Benjamin Franklin in the Philosophical Transactions, accompanied by an account of the fish's anatomy by Hunter (1773). The specimens actually dissected by Hunter may still be seen in the Hunterian Museum of the Royal College of Surgeons (Dobson, 1970). The subject quickly became fashionable, achieving its first review in 1774, when the President of the Royal Society presented the Copley Medal to John Walsh (Pringle, 1775). The discharge of Electrophorus was described by Williamson in 1775, and Hunter (1775) again did some dissections. Cavendish (1776) carried out an extensive investigation on Torpedo, and built a model of the electric organ. The electric eel and Torpedo inspired scurrilous poems by an anonymous author (1777). A more familiar chapter in the history of electrophysiology began in 1791 with the publication of Galvani's De viribus electricitatis, quickly followed by the opening shots in the celebrated controversy with Volta (1792), an admirable account of which has been given by Cohen (1953). Although Volta made comparisons between his 'pile' and the electric organ, there was still doubt at this time whether the discharge of an electric fish could be regarded as 'ordinary' electricity. Faraday (1839) devoted considerable efforts to proving that electricity produced the same effects

57P PHYSIOLOGICAL SOCIETY, JULY 1976 whatever its source, and by 1840 Matteucci was equating the behaviour of the electric organ with that of nerve and muscle. During the latter part of the nineteenth century the electric organ continued to receive what modern electrophysiologists might regard as a disproportionate amount of attention, largely because it was relatively easy to study with the insensitive instruments of the day. In Biedermann's Electro-physiology (1898), the chapter on electrical fishes was nearly as long as that on nerve, and it was still assumed that studies of the electric organ would lead the way to a better understanding of the electrical properties of nerve and muscle. In the event, the boot proved to be on the other foot, for it was not until intracellular micro-electrodes had been applied by Nastuk & Hodgkin (1950) to the measurement of action potentials in muscle fibres that it became technically feasible to discover how the additive discharge of the electric organ is brought about (Keynes & Martins-Ferreira, 1953). Only now does it seem that the electric organ is coming into its own again, albeit in a manner not foreseen by the nineteenth-century electrophysiologists, as a source of acetylcholine receptor protein for biochemists! REFERENCES

ADANSON, M. (1757). Histoire naturelle du Sne'gal. Paris: Bauche. ALLAMAND, J. N. S. (1758). Kort verhaal van de Uitwerkzelen, welke een Amerikaanse Vis veroorzaakt op degeenen, die hem aanraken. Verhandelingen uitgegeven door de Hollandse Maatschappy der Wetenschappen te Haarlem 2, 372-379. ANONYMOUS (1777). The Electrical Eel: or, Gymnotus electricus. By Adam Strong, Naturalist. Bew, London. Also The Torpedo, a Poem to the Electrical Eel. London. BIEDERMANN, W. (1898). Electro-physiology, trans. WELBY, FRANCES A., vol. 2. London: Macmillan. CAVENDISH, H. (1776). An account of some attempts to imitate the effects of the Torpedo by electricity. Phil. Trans. R. Soc. 66, 196-225. COHEN, I. B. (1953). Luigi Galvani's 'Commentary on the effect of electricity on muscular motion', trans. FoLuY, MARGARET GLOVER. Norwalk, Connecticut: Burndy Library. DOBSON, JESSIE (1970). Descriptive Catalogue of the Physiological Series in the Hunterian Museum of the Royal College of Surgeons of England. London: Livingstone. FARADAY, M. (1839). Experimental researches in electricity. Fifteenth series. Notice of the character and direction of the electric force of the Gymnotus. Phil. Trans. R. Soc. pp. 1-12. GALvANI, L. (1791). De viribus electricitatis. Bologna. HUNTER, J. (1773). Anatomical observations on the Torpedo. Phil. Trans. R. Soc. 63, 481-489. HUNTER, J. (1775). An account of the Gymnotus electrics. Phil. Trans. R. Soc. 65, 395-407. KEYNEs, R. D. & MARTINs-FERREIRA, H. (1953). Membrane potentials in the electroplates of the electric eel. J. Physiol. 119, 315-351. MATTEUCCI, M. C. (1840). Essai sur les phenomrnens lectriques des Animaux. Paris: Carilian-Gouery and Dalmont. 5-2

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NASTUK, W. L. & HODGKIN, A. L. (1950). The electrical activity of single muscle fibres. J. cell. comp. Physiol. 35, 39-73. PRINGLE, J. (1775). A Discour8e on the Torpedo, delivered at the Anniver8ary Meeting of the Royal Society, November 30, 1774. London: Royal Society. VOLTA, A. (1792). Memoria prima sull' elettricita animale. BRUGNATELLI'S Grornale Fieico-Medico2, 146-187. WALsH, J. (1773). Of the electric property of the Torpedo. Phil. Tran8. R. Soc. 63, 461-480. WILLIAMSON, H. (1775). Experiments and observations on the Gymnotu8 electricu8 or electrical eel. Phil. Trans. R. Soc. 65, 94-101.

Some demonstrations on colour By C. HOOD and W. A. H. RUSHTON. Phyaioloqical Laboratory, Cambridge In 1666 Newton performed the celebrated prism experiment in Trinity College, and dispersed white light into all the colours of the rainbow. An 'original Newton prism' was demonstrated; also a spectrum. Individual spectral colours cannot be decomposed into new colours but all those rays added together form the original white. Does a mixture of yellow and blue make green or white? The spectrum will be Used to show how both answers may be true. Helmholtz first resolved the paradox. For thousands of years painters have mixed pigments to obtain new colours. By the eighteenth century many asserted that colour vision was compounded of three primaries; Lomonosov and later Brewster thought that light itself was a mixture of three kinds. Thomas Young reconciled the trichromacy of colour with the infinite gradation of Newton's spectral rays. Having proved by his interference bands that light was a vibration, he postulated three resonators in the eye, thrown into oscillation by these vibrations, and resonating more strongly, the stronger the light and the closer its frequency to the natural resonator frequency. If this is true all spectral colours can be matched by a suitable mixture of three primaries, e.g. red + green + blue. The mixture is physically quite different from the single monochromatic light matched, but it affects identically each resonator and so the eye cannot detect any difference they match. The trichromatic colour mixture that matches each light was measured by Clerk Maxwell and confirmed with great accuracy by W. D. Wright and W. S. Stiles. What are Young's resonators? They are electrons in or orbitals of the visual pigment molecules. Kuhne in Heidelberg studied the effect of light upon rhodopsin, the pigment of twilight vision in the rods. He showed that this rod pigment was bleached by light and regenerated in the dark if in good contact with the pigment epithelium (frog).

PHYSIOLOGICAL SOCIETY, JULY 1976 59P There are three kinds of cone in the human eye, each with a pigment that is not rhodopsin but is bleached by light and regenerates in darkness. Young's theory may thus be stated as follows: Any visible light will be absorbed by the cone pigments; the resulting colour sensation will depend upon the ratio of absorption in each of the three pigments. This does not mean that if we know that ratio we can predict the colour sensation experienced, for this depends also upon a multi-dimensional adaptation factor. It does mean that in two different light presentations when for each cone the quantum catch is the same from both presentations, the appearance will be the same, provided that the condition of the retina is otherwise identical. Grassmann's Laws, which define the properties of colour mixtures, follow at once from the quantum catch in each pigment. This means, as was demonstrated, that colour matches remain stable in all conditions where the visual pigments do not change their absorption spectra. But the colour appearance of these matching fields may change very greatly due to nerve adaptation, etc. Red-Green colour anomaly. John Dalton, the famous chemist, was the first to describe carefully his own colour vision in which he could not distinguish red from green, and saw red dimly. He thought his eyes contained a blue fluid that restricted the transmission of certain colours. Young (his contemporary) thought that Dalton lacked the red resonators. At post-mortem Dalton's eyes appeared of normal transparency. Clerk Maxwell studied the colour matches and confusions of his fatherin-law whose vision was similar to Dalton's. On the Maxwell colour triangle the colours confused all fell on straight lines running to the 'red corner' of the triangle. Maxwell saw that this meant that his subject lacked the red fundamental sensation - justifying Young's diagnosis of Dalton. Our retinal densitometer was able to confirm Young and Maxwell by showing objectively that these subjects (protanopes) lack erythrolabe, the pigment of the red-sensitive cones. The most common type of red-green anomaly is one not so severe as to confuse good reds and greens, but browns cannot be distinguished from olive greens. Lord Rayleigh classified these by their match of yellow with a mixture of red and green. About 5 % of males exhibit this anomaly and some members of the audience came forward and made the Rayleigh Match, showing that some need more red, some more green than average to match the yellow.

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W. H. Gaskell's demonstration of 'the electrical changes in the quiescent cardiac muscle which accompany stimulation of the vagus nerve' By JEAN BANISTER, HIARY BROWN, W. GILES, 0. F. HUTTER and SUSAN NOBLE. University Laboratory of Physiology, Oxford and Institute of Physiology, University of Glasgow W. H. Gaskell was one of the founders of the Physiological Society. As well as his contributions in other fields of Physiology, he made two major contributions to cardiac physiology. Both were of fundamental importance. In 1883, as the culmination of a long series of experiments, Gaskell reached the conclusion that 'the rhythm of the sinus and therefore of the whole heart depends upon the rhythmical properties of the muscle tissue of the sinus, and not upon any special rhythmical nervous apparatus'. Gaskell's second contribution concerned the mechanism of nervous inhibition of the heart. Having recognized that 'the muscular tissue of both the visceral and vascular systems is supplied by two sets of nerves, of which the one sets that tissue in activity and causes contraction, the other inhibits its action and causes relaxation' (Gaskell 1886a), he set out 'to compare the electrical change which takes place in a muscle when its inhibitory nerve is stimulated with those which are known to accompany the action of a motor nerve' (Gaskell, 1886b). He 'showed how a preparation could be made of the auricle of the tortoise in such a way that spontaneous beats cease for a time, owing to separation from the place where those beats arise in the sinus, while the supply of inhibitory fibres is left intact in a nerve (the coronary) which is separate from the tissues and can be left when they are cut' (Bayliss, 1915). He caused demarcation current to flow between an injured spot and intact auricle and found that 'stimulation of the vagus nerve in the neck causes some alteration in the non-beating muscle of the auricle which is manifested by an electrical change of an opposite sign to that which accompanies contraction of the muscle' (Gaskell, 1887) (see Fig. 1). Thus Gaskell was responsible for the first demonstration of the hyperpolarizing effect of vagus stimulation on heart muscle, and hence for the view that the seat of such inhibition is in muscular tissue rather than within the nervous system. On the heart of the frog, Gaskell (1887) found that 'stimulation of the sympathetic augmentorr) nerve causes an electrical variation of the same sign as that caused by a contraction', i.e. a depolarization. Gaskell chose to work on the quiescent auricle in order to test whether the inhibitory state differs from the resting state. His choice of preparation allowed him to use the slow but sensitive D'Arsonval galvanometer. In a

PHYSIOLOGICAL SOCIETY. JULY 1976

B

A.

26

R. Vag R.A. S

27

!

R. Vag R.A. 5

Fig. 1. A, Gaskell's preparation for recording electrical changes in the quiescent auricle of the tortoise on vagal stimulation. V, vagus nerve; C, coronary nerve; S, sinus and part of auricle in connexion with it; G, galvanometer; E, induction coil (Gaskell, 1887). B, Gaskell's record: galvanometer readings taken every 5 sec. The demarcation current caused by burning the apex of the auricular strip is gradually declining. Stimulation of the vagus (between the arrows) causes 'an electrical change of an opposite sign to that which accompanies contraction of the muscle'.

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62P 6PROCEEDINGS OF THE demonstration given to the Physiological Society in July 1887, Gotch used the faster capillary electrometer to examine the effect of vagus stimulation on the demarcation potential in the beating tortoise auricle. 'It was observed that during inhibition the top of the mercurial column remained steady at the level which is assumed in diastole.' A small difference between the demarcation occurring during inhibition and during diastole was, however, detected by Burdon Sanderson with a D'Arsonval galvanometer that was switched into the circuit only during diastole. 'In diastole and in the Gaskell preparation the heart is asleep, but not quite asleep as it is in inhibition' (Burdon Sanderson, 1887). A

0 5 mV [

B

i V, a term in V2 has been

dropped. The driving pressure resulting from the surface tension is opposed by the pressures resulting from the weight of the column, viscous drag and

PHYSIOLOGICAL SOCIET Y, JUL Y 1976

65P inertia. This is a second-order linear system. The undamped resonant frequency, W, is determined by the familiar pendulum equation:

W

1Ig 2 272 Hz,

where L is the length of tube which contains mercury, about 5 cm in Adrian's device.

Fig. 1. A capillary electrometer. The black represents mercury. The dotted area indicates dilute sulphuric acid. The two tungsten wires allow electrical connexion.

We have measured the frequency response of the capillary electrometer. The data are plotted on a log-log Bode plot in Fig. 2. Lines with slopes of 0, -1, and -2, were fitted by eye. They indicate an undamped resonant frequency of about 2 Hz, in agreement with the calculation. The complete transfer function represented by the lines is 1

(S+(2ir) 0-6)(S+(2r7) 3 0) Lucas (1912) made a machine to correct the records made from the capillary electrometer. The user inputs the abscissa, ordinate, and slope

66P PPROCEEDINGS OF THE of each point. The machine, using gears and levers, realizes the transferfunction (S + W0),

and outputs a new graph. By setting W0 = (27r) 0-6 and operating the machine, a new graph is created, related to the original by the transfer function (S (5+ + (27) (21T) 0-0.6) x (S+ (27T) 0.6)(S+ (2ff) 3.0) (S+ (27f) 3.0)'

1)1

While this is a fivefold increase (3.0/0.6) of the useable frequency range, it is surprising that Lucas makes no mention of using the machine a second O'

,l .

vigi

.9 |1411

I

1111"" 1 0-7 Hz

I

I

1

|

1l1

I

I

' ' "'i

W

=2-2 Hz

C~~~~~~~~~~~~~~ ,U~o .01

~ ~ ~

U

~

~ ~~~*

6 Hz

aE E E

°

0.001

0-01

0-1

1 Frequency (Hz)

10

100

Fig. 2. A log-to-log-Bode plot of the frequency response of Adrian's capillary electrometer. The straight lines represent the transfer function 0084 mm/mV

(S+(27T) 06) (S+(2ff) 3-0)

time, to completely correct the response. Of course, frequencies higher than 10 Hz will be so attenuated as to be lost before input into the machine anyway. A simpler way of improving the response would be to reduce the length of mercury which has to move; a threefold improvement in frequency response should be easily attainable. Both Adrian's capillary electrometer and Lucas's machine were demonstrated. Fig. 1 A and B of the following demonstration showed records made with a capillary electrometer by Waller (1887).

PH YSIOLOGICAL SOCIET Y, JUL Y 1976

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REFERENCES

LUCAS, K. (1912). J. Phy8iol. 44, 225-242. OVERBEEK, J. T. G. (1952). In Colloid Science, vol. i, ed. KRUYT, H. R., pp. 115122, 146-159. Amsterdam, Elsevier. WALLER, A. D. (1887). J. Physici. 8, 229-239.

The Einthoven string galvanometer and the interpretation of the T wave of the electrocardiogram BY R. H. ADRIAN, R. C. CnANNELL, I. COEEN and D. NOBLE. The Physiological Laboratory, Downing Street, Cambridge, and the University Laboratory of Physiology, South Parks Road, Oxford The flow of electric current during the heartbeat was first demonstrated in 1856 by Kolliker and Muller. They placed a frog nerve-muscle preparation in direct contact with the ventricle and were able to show that there are two distinct electrical changes during each contraction of the ventricle. The origin of the second of these changes (which, following Einthoven, is called the T wave) has remained controversial to the present day. The first recordings showing the magnitudes and directions of the components of the electrocardiogram were obtained using the capillary electrometer (see demonstration by Campbell & Pelli, 1976) and the first accurate analysis was performed by Burdon-Sanderson & Page (1880). Waller (1887) first used this method to record from the surface of the human body. The currents at the surface of the body were, however, too weak to allow good recordings to be obtained with this instrument (Fig. 1A). An example of the degree of resolution that could be obtained under more favourable conditions is shown in Fig. 1 B, which is a record obtained by Gotch (1910) from electrodes placed directly on the surface of the heart of a tortoise. The currents are then much stronger and the events occur more slowly in this case than in mammals (2.5 sec per cycle as opposed to less than 1 sec per cycle). There is clear resolution between the two waves corresponding to ventricular activity. This record is interesting in showing the second wave occurring in the opposite direction (negative) to the first (positive). This is the result expected when the excitation travels as a wave of constant duration from base to apex, and in this it corresponds to the simple diphasic wave obtained extracellularly from nerve or muscle (see Fig. 1 C). In 1892 Bayliss and Starling showed that in the mammalian heart the second wave of the record obtained from the surface of the body is in the same direction as the first (i.e. the T wave is positive). They explained the phenomenon by supposing that the excitation lasts longer at the base than at the apex.

PROCEEDINGS OF THE

68P

C

A

B

S

i

n~~~~~~

Fig. 1. A, electrocardiogram recorded from surface of human body using capillary electrometer (Waller, 1887). Time marker: sec. Note relatively poor quality of the record. B, electrocardiogram recorded from surface of tortoise heart (one electrode on sinus, second electrode on apex of ventricle) using capillary electrometer (Gotch, 1910). Time marker: * sec. The currents at the surface of the heart are much stronger and the capillary electrometer is then capable of accurately discriminating between the various components. In fact, this record is almost exactly similar to modern records from tortoise (see Noble, 1975, p. 11). The first positive wave and the following negative wave are associated with auricular excitation. The second positive wave and the following negative wave are associated with ventricular events. C, electromyogram (upper trace) recorded from frog gastrocnemius muscle using string galvanometer (Einthoven, 1913). The record consists of a positive wave followed immediately by a negative wave. Bottom trace is a signal marker and shows time of excitation of nerve. Time scale: 0-002 see; voltage scale, 7 mV. D, electrocardiogram (lower trace) recorded from surface of human body by Einthoven (1913) using string galvanometer. Note the high resolution obtained compared to capillary electrometer (4). The record also shows Einthoven's original labelling of the P, Q, R, S and T waves. The upper record shows the arterial pulse pressure. Time scale, 0 01 see; voltage scale, 100 V. E, electrocardiograms from surface of frog heart recorded with string galvanometer by Mines (1913). The first record shows the normal trace (frequency of beating). The second and third traces show records obtained during warming of the ventricular apex. The T wave becomes positive. The fourth record shows result obtained when warming ceased. The T wave then reverts to being negative.

69P PHYSIOLOGICAL SOCIETY, JULY 1976 Einthoven introduced the string galvanometer in 1901. This instrument is capable of better time resolution. A fine quartz fibre coated continuously with gold is stretched between the poles of a powerful electromagnet (Fig. 2). Currents flowing down the string then cause the string to be displaced laterally in the magnetic field. The movement of the string is recorded by means of a microscope and a photographic plate. The response time of the instrument could be decreased by increasing the tension on the

C

Fig. 2. The mechanism of the string galvanometer. The fine wire ('string'), CC, is stretched in the narrow gap between the poles N and S of a powerful electromagnet. When a current passes through the 'string' in the direction of the vertical arrows, the wire is deflected in the direction of the arrow a, that is, at right angles to the magnetic field NS. This small movement is observed, or projected on to a photographic plate, by means of a microscope, ED, the light of an arc lamp being condensed by the lens F on to the string. (From Bayliss, 1915.)

string. Einthoven's record of the human e.c.g., with his nomenclature, is shown in Fig. ID. Note, once again, that the T wave is positive. Mines (1913) provided strong evidence for Bayliss and Starling's hypothesis by showing that in an animal (the frog) with a negative T wave, the T wave could be made positive simply by warming the apex in order to produce a shorter period of excitation than at the base (Fig. 1 E). This had also been shown by Burdon-Sanderson & Page (1880, 1883) using the capillary electrometer. Finally, Samojloff (1910) made the important

70PPROCEEDINGS OF THE 70P observation that the direction of the T wave is independent of the direction of the initial spread of excitation. An extrasystole conducted from the apex shows the same T wave as a normal beat conducted from the base. This observation shows that the differences in duration of excitation are attributable to the intrinsic properties of different regions of the ventricle. They do not depend on the direction of propagation. The origin of these intrinsic differences was the subject of an illustrated communication at this meeting (Cohen, Giles & Noble, 1976). Thus, the major elements of the current explanation of the T wave of the electrocardiogram are based on the work of Burdon-Sanderson, Page, Bayliss and Starling nearly a century ago. Einthoven's major contribution was to provide an instrument that enabled the electrocardiogram to be used in routine clinical practice (see demonstration on the work of Sir Thomas Lewis (Hollman, 1976)) and which allowed more accurate recordings to be made from the body surface than was possible with the capillary electrometer. REFERENCES BAYLISS, W. M. (1915). Principles of General Physiology. London: Longmans. BAYLISS, W. M. & STARLING, E. H. (1892). Mon. Int. J. Anat. & Physiol. 9, 256-281. BURDON-SANDERSON, J. S. & PAGE, F. J. (1880). J. Physiol. 2, 384-435. BURDON-SANDERSON, J. S. & PAGE, F. J. (1883). J. Physiol. 4, 327-338. CAMPBELL, F. W. & PELLI, D. (1976). J. Physiol. 263, 64-67P. COHEN, I., GILES, W. & NOBLE, D. (1976). J. Physiol. 263. EINTHOVEN, W. (1901). Arch. Neerland, ser. ii, 6, 625-633. EINTIIOVEN, W. (1913). Pfiiger's Arch. 149, 65-86. GOTCH, F. (1910). Heart, 1, 235-261. HOLLMAN, A. (1976). J. Physiol. 263, 72 -74P. K6LLIJKER, H. & MULLER, H. (1856). Verh. phys.-med. Ges. Wiirz. 6, 528. MINES, G. R. (1913). J. Physiol. 46, 188-235. NOBLE, D. (1975). The Initiation of the Heartbeat. Oxford: Clarendon. SAMOJLOFF, A. (1910). Pflugers Arch. 135, 417. WALLER, A. D. (1887). J. Physiol. 8, 229-234.

Mackenzie's polygraph for simultaneous recordings of venous and arterial pulse By P. S. FREEDMAN.* The London Hospital Medical College, London, El 2AD James Mackenzie, born in 1853 near Scone, was a medical student at Edinburgh when the Physiological Society was founded. He was destined 'to bring reason and precision to the understanding of heart irregularities and by 1902 when he published "The Study of The Pulse" had undoubtedly become the world clinical authority on the heart' (McMichael, * Present address: Cambridge University School of Clinical Medicine, Addenbrooke's Hospital, Hills Road, Cambridge CB2 2QQ.

PH YSIOLOGICAL SOCIET Y, J UL Y 1976 71P) 1975). This position was attained while in general practice in Burnley where he made numerous observations of patients with heart disease and recorded the venous and arterial pulse simultaneously with an instrument of his own design. I began to consider whether some simple means of registering the venous pulse might not be devised whereby tracings of the heart or arteries might be taken at the same time, in order to determine the time of occurrence of events during a cardiac revolution [Mackenzie, 1893].

By linking a lead cone with rubber tubing to a Marey tambour and lever the pulsation of the vessel was recorded; two receivers and tambours allowed simultaneous recordings. The tracings were obtained on a small smoked drum or strips of smoked paper fed through a Dudgeon sphygmograph. It was a remarkable achievement to obtain satisfactory results given that the laboratory was generally the cramped room where his patients lay sick. Mackenzie was not the first to study the venous pulsations: in 1794 J. Hunter observed the dog, in 1865 Friedreich produced jugular venous curves from patients, and in 1867 P. C. E. Potain obtained simultaneous tracings of the jugular pulse, carotid and radial artery. C. S. Roy of Cambridge expressed reservations that the polygraph could produce decent curves unaffected by inertia vibration and was concerned about the use of air transmission (correspondence quoted, Mair, 1973). Mackenzie maintained that the delay in air transmission was very small compared to that between carotid and radial pulses and insisted he was rigorous in obtaining proper timings of the various pulses (Mackenzie, 1894). With the 'clinical polygraph' Mackenzie made significant observations on the origin of extrasystoles, on heart block and on the action of digitalis in auricular fibrillation. While stating that the normal periodic stimulus to contraction of the heart originated at the mouth of the great veins subsequently conducted through the auricles to the ventricles, he concluded that the stimulus to the extrasystole may arise in the auricle or ventricle itself; he appreciated that Gaskell had demonstrated reversal of the beats of the heart chambers in the tortoise. Mackenzie collected many hearts from autopsy and retained clinical histories and polygraph recordings of their owners in vivo. These hearts were sent from Burnley to The London Hospital for Keith to examine. One polygraph tracing showed rapid regular atrial systole with standstill of the ventricles lasting seventeen seconds: 'During the pause the patient lost consciousness which returned as soon as the ventricles began to beat.' A. Keith and M. W. Flack confirmed the existence of the atrio-ventricular bundle, acknowledged S. Tawara's earlier discovery, and noted that where there had been

72P PROCEEDINGS OF THE irregularity in auriculo-ventricular rhythm the connecting muscle bundle was replaced by fibrous tissue, 'thus interfering with the sole path by which the auricular wave of contraction passes to ... the ventricles'. The ink polygraph appeared in 1906 with help from a watch-maker, Sebastian Shaw: its use became widespread and contributed to Mackenzie's fame. The apparatus sold for the price of 10 guineas complete in plush lined box. A sphygmograph was connected by rubber tubing to a Marey tambour for recording the radial pulse and a lead cone similarly connected for the jugular pulse. A specially designed clockwork motor drove a roll of paper on which the lever movements were inscribed in ink. A governor was incorporated allowing for continuous variation in paper speed and a third pen attached to a vibrating fork marked time intervals of one-fifth of a second. The ink polygraph was portable and practical: it was superseded only by the solid-state electrocardiograph. The demonstration included (a) a Dudgeon sphygmograph similar to that incorporated in the 'clinical polygraph', (b) a working 'ink polygraph', (c) reproduction of polygraph tracings published by Mackenzie, and (d) copies of specific references. REFERENCES MAIR, A. (1973). Sir James Mackenzie, General Practitioner. Edinburgh: Churchill

Livingstone. MCMICHAEL, J. (1975). The Harveian Oration. Royal College of Physicians, London. MACKENZIE, J. (1893). J. Pacth. Bact. 1, 53-89. MACKENZIE, J. (1894). J. Path. Bact. 2, 84-154. MACKENZIE, J. (1902). The Study of the Pulse. Pentland: J. Young. MACKENZIE, J. (1925). Diseases of the Heart, 4th ed. Oxford Medical.

Sir Thomas Lewis (1881-1945) BY A. HOLLMAN. University College Hospital, Grafton Way, London WC1E 6DB The brilliant work done by James Mackenzie (later Sir James) whilst a single-handed general practitioner in Burnley using mechanical recordings of the jugular venous pulse, the arterial pulse, and the heart's apex beat was the first sustained scientific effort to unravel the mysteries of cardiac irregularities. His thoughtful and critical research led to substantial advances in this field and culminated in his monograph The Study of the Pulse, published in 1902. Soon after this Mackenzie moved to London and in 1908 he met Lewis who, having studied under E. H. Starling at University College, was working with Leonard Hill on the influence of respiration on the venous and arterial pulses. Mackenzie urged Lewis to study the nature of the puzzling 'irregular irregularity' of the heart beat which Mackenzie had ascribed to paralysis of the atria with nodal rhythm.

PHYSIOLOGICAL SOCIETY, JULY 1976

73P Lewis, a Beit Memorial Fellow, acquired the Einthoven string galvanometer as manufactured by the Cambridge Instrument Company and set it up in a basement room at University College Hospital Medical School. His stature as a medical scientist of outstanding intellect with an enormous capacity for unrelenting hard work then began to be revealed. He soon found that the irregularity which had intrigued Mackenzie was due to fibrillation of the atria and in 1909 felt able to entitle a paper in the British Medical Journal, 'Auricular fibrillation: a common clinical condition'. The nature of other forms of disordered heart action were soon uncovered by his work with the Einthoven electrocardiograph. Nodal rhythm was identified and was published in 1910 in the journal Heart, which he had founded the previous year. Animal work was undertaken when it could help in the analysis of clinical irregularities - for example, his research with A. G. Levy in 1911 on 'Heart irregularities resulting from the inhalation of low percentages of chloroform vapour, and their relationship to ventricular fibillation'. The year 1911 was eventful for Lewis. At a meeting of the clinical section of the Royal Society of Medicine, held at-University College Hospital Medical School, William Einthoven was the guest speaker and both he and Lewis read papers which described right axis deviation of the ventricular complex of the electrocardiogram (e.c.g.) in cases of congenital pulmonary stenosis. This was an early indication of the value of the e.c.g. in delineating abnormalities of the muscle of the ventricles as distinct from arrhythmias. Also in 1911, Lewis published his famous monograph 'The mechanism of the heart beat', which quickly became the bible of electrocardiographers the world over. Having worked previously with mechanical records of the arterial and venous pulses, e.g. 'The pulse in aortic disease: the relation of pulse curves to blood pressure' (Lewis, 1906), Lewis now began to record these events photographically and also produced excellent graphic records (phonocardiograms) of the heart sounds and murmurs. He then began his classical pioneer research on the spread of the electrical impulse in the heart and this was, and remains, electrophysiological work of the highest order. The first paper written with Paul D. White of Boston and Jonathan Meakins of Montreal was 'The excitatory process in the dog's heart; Part I, The auricles' (Lewis et al. 1914). Part 2 on the ventricles written with M. A. Rothschild came out a year later. Lewis was elected F.R.S. in 1918 and knighted in 1921. The former honour was conferred in respect of his electrocardiographic work. The latter was in recognition of his work during the war as a clinical cardiologist, when with Sir John Parkinson (born 10 February 1885, died June 1976), Sir William Osler and A. N. (now Sir Alan) Drury he did

74PPROCEEDINGS OF THE 74P much to help soldiers who erroneously were diagnosed as suffering from cardiac illness. In this respect his work published in 1918 on 'The soldier's heart and the effort syndrome' will be long remembered. The publication in 1925 of the third edition of The Mechanism and Graphic Registrhtion of the Heart Beat signalled his retirement from research work in electrocardiography. The second half of his long and fruitful career in medical research was concerned chiefly with The Blood Vessels of the Human Skin and their Responses and with Pain - these being the titles of his last two monographs. He wrote also on congenital heart disease and bacterial endocarditis, and his leisure activity found expression in the journal British Birds with a paper 'Notes on the breeding habits of the little tern'. Lewis's most important contribution to medicine may have been his concept of Clinical Science; his insistence that progress in medicine would come chiefly from scientific studies on living men in health and disease rather than from the basic science laboratories and animal experimentation. It is difficult for us now to re-live the time when Lewis, almost alone, fought for the creation of established career posts in medical research now numbered in their thousands. Of his numerous national and foreign distinctions none was more significant than the Copley Medal of the Royal Society (1941), which had previously been awarded to only one clinician, Lord Lister. Lewis's first scientific paper, written whilst a pre-clinical student, with Sale Vincent his teacher in physiology in Cardiff, was published in the Journal of Physiology in 1901. He contributed a further twenty-nine papers to the Journal. REFERENCES

LEWIS, T. (1902). The Study of the Pulse. Edinburgh and London: Young Pentland. LEWIS, T. (1906). Lancet ii, 714. LEWIS, T. (1909). Br. med. J. ii, 1528. LEWIS, T. (1910). Heart 2, 127. Lewis, T. (1911). Proc. R. Soc. led. 4, Med. Sec. 81. LEWIS, T. (1911). The M1echanism of the Heart Beat. London: Shaw and Sons Ltd. LEWIS, T. (1918). The Scldier's Heart and the Fffort Syndrome. London: Shaw and Sons Ltd. LEWIS, T. (1920). British Birds 14, 74. LEWIS. T. (1925). The Mechanism and Graphic Registraticn of the Heart Beat (3rd edition). London: Shaw and Sons Ltd. LEWIS, T. (1927). The Blocd Vessels of the Human Skin and their R&sponses. London: Shaw and Sons Ltd. LEWIS, T. (1942). Pain. New York: Macmillan Co. LEWIS, T. & LEVY, A. G. (1911). Heart, 3, 99. LEWIS, T., MEAKINS, J. & WHITE, P. D. (1914). Phil. Trans. R. Soc. B 205, 375. LEWIS, T. & ROTHSCHILD, M. A. (1915). Phil. Trans. R. Soc. B 206, 181. LEWIS, T. & VINCENT, S. (1901). J. Physiol. 26. xixP.