sir michael foster, k.c.b., f.r.s. a secretary of the royal society

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Sir Michael Foster, K.C.B., F.R.S. A Secretary of the Royal Society Author(s): Henry Dale Source: Notes and Records of the Royal Society of London, Vol. 19, No. 1 (Jun., 1964), pp. 10- 32 Published by: The Royal Society Stable URL: http://www.jstor.org/stable/3519858 . Accessed: 16/06/2014 11:19 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. . The Royal Society is collaborating with JSTOR to digitize, preserve and extend access to Notes and Records of the Royal Society of London. http://www.jstor.org This content downloaded from 91.229.229.203 on Mon, 16 Jun 2014 11:19:05 AM All use subject to JSTOR Terms and Conditions

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Sir Michael Foster, K.C.B., F.R.S. A Secretary of the Royal SocietyAuthor(s): Henry DaleSource: Notes and Records of the Royal Society of London, Vol. 19, No. 1 (Jun., 1964), pp. 10-32Published by: The Royal SocietyStable URL: http://www.jstor.org/stable/3519858 .

Accessed: 16/06/2014 11:19

Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at .http://www.jstor.org/page/info/about/policies/terms.jsp

.JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range ofcontent in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new formsof scholarship. For more information about JSTOR, please contact [email protected].

.

The Royal Society is collaborating with JSTOR to digitize, preserve and extend access to Notes and Records ofthe Royal Society of London.

http://www.jstor.org

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SIR MICHAEL FOSTER, K.C.B., F.R.S. A SECRETARY OF THE ROYAL SOCIETY

By SIR HENRY DALE, O.M., G.B.E., F.R.S.

[Plate I]

AMONG other reasons for including some account of Sir Michael Foster's life and achievements in these Notes and Records, we naturally

recall his special services to the Royal Society, as one of its Secretaries for the unusually long period of 22 years, from 1881 to 1903. A glance at the Society's Record will show, indeed, that he was the last to hold that Office, before the present convention of a ten years' limit to its tenure came to be adopted. And it may further be noted, in the same connexion, that the only Secretaryship as long as Sir Michael Foster's, in the whole history of the Society, was that of Sir George Stokes, whose period of service overlapped with his, and was even substantially longer. Sir George's Secretaryship lasted indeed for 31 years, from 1854 onwards; and it might even be thought that it required his election as President, in 1885, to bring it to a natural end.

Both Stokes and Foster, in any case, did very distinguished service to the Society; and in Foster's case, with which we are here directly concerned, we shall see that he was able to use the opportunities which came to him in that connexion, in such ways as to strengthen the influence and advocate the claims of the Royal Society with the Government of the day, to enhance its reputation with a wider public, and in many directionsto enlarge the exercise of its historic functions, of promoting the advancement of scientific know- ledge, and its uses for the general benefit. Foster's influence, indeed, on the development and the teaching of the natural sciences, especially in functional biology covering a far wider range than that of his own special department of animal and human physiology, had become noteworthy for more than a decade before his official connexion with the Royal Society's activities began, and even for some years before his election to its Fellowship.

Michael Foster was born in Huntingdon, on 8 March I836, as the eldest son of Michael Foster, F.R.C.S., well known in that neighbourhood as an able practitioner of medicine and surgery, and esteemed as a man of strongly marked character, staunchly nonconformist and radical in his religious and political convictions. We can think, accordingly, of Michael Foster, the son, as passing his boyhood and development in such an atmosphere of opinions and loyalties, as might be expected to retain some of its influence on those which were to be his own.

He attended the Huntingdon Grammar School until he was 13 years old,

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Plate 1

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SIR MICHAEL FOSTER, K.C.B., F.R.S.

Photograph in the Library of Trinity College, Cambridge. [Facing payge 1 o

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and his education was then continued, from 1849 to 1852, at the University College School, in London. There he matriculated at the University of London; and at what, by modern standards, would seem to be the early age of 16, he entered University College as a student. An even greater precocity might be suggested by his graduation as B.A. only two years later, in I854, at the age of 18. It should be remembered, however, that at that time, and indeed till the end of the Igth century, the University of London was only an examining and degree-giving organism. And in the same connexion I might recall a more recent case of such early graduation at London. In 1892 Joseph Barcroft, then a schoolfellow of mine, graduated B.Sc. as an 'external student' of London University, while still at school; and it was he, the later Sir Joseph Barcroft, who was in 1925 to become a successor at one remove, in the Chair of Physiology at Cambridge, to Sir Michael Foster, who had been its first incumbent in 1883.

Untilhe graduated, Foster's studies at school and university had evidently been largely on the literary side, and his record in the B.A. examination had appeared to give further promise of natural gifts in that special direction; for he was placed first in the Honours list in Classics, and was awarded the Scholarship in that Faculty. When the choice of a career, however, called then for an immediate decision, the possibility ofjoining his father in medical prac- tice at Huntingdon seemed to offer the most obvious opening; nor is there any reason to suppose that this prospect was other than entirely congenial to the young Foster's own ideas. He began, accordingly, to attend the preliminary scientific lectures of the medical course at University College, and at the same time to obtain what clinical experience was available at that stage, by attendance at the University College Hospital. He won a Gold Medal for his examination success in anatomy and physiology, and another in chemistry; and he then brought his regular clinical studies to the stage which enabled him to qualify in medicine, by obtaining the London M.B. in 1858, and the M.D. in the following year, at the early age of 23. He must obviously have been a diligent as well as a highly intelligent student; and we should hardly have expected, perhaps, to find that, at this early stage, he could have given much time to activities outside the range of his formal studies. It is on record, however, that he played cricket at University College with such success as to become captain of its eleven; and it is evident that cricket retained a special interest for him. For even when he had become a professor at Cambridge, he contrived for some years to organize, from his colleagues and assistants, a sufficient team to play an annual match; though it can hardly be supposed that this was up to the standard conventional at a university centre. In any

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case, cricket was the only kind of outdoor game in which Foster appears ever to have become interested.

Although he had finished his formal medical studies and graduation by 1859, it was evidently regarded as desirable that he should obtain further clinical experience, and some broadening of the scientific basis of his know- ledge, before beginning country practice with his father. Accordingly he spent most of 1859 and 1860 at University College and the associated hospital, and in Paris, which he visited in each of these years. It was there, we may suppose, that he first began to acquire his special interest in the fundamental contributions to physiology and experimental medicine, being made even then by the great Claude Bernard. In the dedication to the study, which he published some 40 years later, of the life and work of Bernard (I), Foster, with regretful reference to himself as one of those who 'never saw his face', claims that Bernard had been to him, nevertheless, 'a father in our common science'. And this short, but very attractive biography of his great scientific exemplar is full of his enthusiasm, not only for Bernard's great contribution to the foundations of modemrn physiology, but also for his attitude to the philosophy of experimental science in general.

It is evident that during those same two years, 1859-1860, Foster was also beginning his more direct experience of the methods and principles ofphysio- logical research, under the stimulating and distinguished guidance of William Sharpey, then the Professor of Anatomy and Physiology at University College. Sharpey, in fact, must have had a remarkable gift of inspiration as a teacher. Foster, as we shall see, after some years of medical practice with his father at Huntingdon, was to return to Sharpey's Department in University College, as Teacher of Practical Physiology. In his earlier years there, I859- 1860, immediately following his medical graduation, he had already begun experiments on the origin of the rhythmical activity of the heart, using the iso- lated heart of a snail-an experimental subject to which he was later to return -and finding that any separated part of it would still beat rhythmically, so that the beat must be 'the peculiar property of the general cardiac tissue'. And he made a communication on these observations to the British Association at its Aberdeen Meeting in 1859. In the following year he studied the resistance to freezing shown by isolated muscles from frogs, finding that they could regain their normal contractile response to a stimulus after being frozen for 5 to Io minutes, but passed into rigor mortis on being thawed after a longer freezing; and these results were communicated to the Royal Society and published in its Proceedings (2). We might indeed, think of Foster as thus receiving his first official recognition as an experimental biologist, of evident

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promise. I think that it ought to be said at once, however, that it was not as a personal experimenter in any direction, that he was to win his great position of influence and authority in science in general, or in physiology in particular. At more than one interval, when an opportunity presented itself among other calls on his time and interest, he resumed for a time such direct activities in experimental research; but all his experiments were of the same relatively simple, observational kind as those already mentioned, involving neither novelty of method, nor anything special in the way of instruments or appara- tus. I think, indeed, that it may safely be concluded, that Foster's eventually outstanding contribution to the rise and rapid development in this country, and in the other English-speaking countries of the world, of experimental physiology and, indeed, of the application of the experimental method to a wider range of the biological disciplines, owed relatively little to his own direct enterprises as an experimenter. Such experiments as he devised for himself, and carried out with his own hands, were sound enough; but their results were such as to leave but a modest mark on scientific history. It is unlikely that, by themselves, they would have called for any special record here of Foster and his achievements.

After the two years following graduation, spent in Sharpey's laboratory and in educational visits to Paris, the year 1861 made an unfortunate, though not a wholly unfruitful gap, in the course of Foster's career. As happened all too often to the young men and women of those days, an otherwise un- explained impairment of his health had aroused a suspicion of pulmonary consumption. It may be recalled that more than two further decades had to pass, before Koch's discovery of the tubercle bacillus, in I882, brought a new conception of the nature of that disease. And it is further of interest for us, perhaps, to note in advance that, forty years after this temporary threat to his own health and the resulting delay of his plans, Foster himself, in 9IgoI, was to be appointed Chairman of a Royal Commission on Tuberculosis. The particular task of the Commission was 'to enquire into the Relations of Human and Animal Tuberculosis'. And quite recently, in connexion with the celebration in November I963 of the Jubilee of the Medical Research Council, which came into being (as the Medical Research Committee) in 1913, attention has been drawn to the small-scale but important precedent created by Foster's Royal Commission, twelve years earlier. For this body had taken, doubtless under Foster's stimulus, 'the remarkable step of deciding not to hear evidence-that is opinions-but to conduct experimental investi- gations of its own, to determine facts; and for that purpose it was provided with funds enabling it to employ a scientific staff and meet other costs. In a

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sense, within the limits of its subject, the Commission was a true precursor (3)-i.e. of the Medical Research Council. The recommendations of this Commission were indeed to play an important part in the inauguration of a new period, in which policies based upon active and continuous researches, and securing a prompt application of their verified results, have now reduced the formerly serious problems of tuberculous infection, in man and in farm animals, to relatively small proportions, in all scientifically organized com- munities. Foster was fortunate, perhaps, in that the early threat of 'consump- tion' in his own case proved to be mild enough to be dispelled by a sea voyage, then regarded as the appropriate expedient in such a condition. He obtained appointment as ship's surgeon on a vessel bound for the Red Sea, in connexion with a lighthouse-building project; and he was able to use the opportunity which this voyage presented, to make scientific observations on marine fauna, as well as to get rid of the suspicion concerning his health. It may be, perhaps, that Foster found encouragement, to use this opportunity for trying his hand as a marine biologist, from the knowledge that T. H. Huxley, eleven years his senior, but later to become his close friend and adviser, had begun his own distinguished career in natural science as a surgeon in the Royal Navy, and had been able as such, during a four-year voyage to Far Eastern waters in H.M.S. Rattlesnake, to make observations and collections of such outstanding importance and interest, as rapidly to establish his position as a coming leader, among biological investigators. Foster and Ray Lankester many years later, from 1898 onwards, were to act asjoint editors of a collected edition ofHuxley's scientific communications.

Some years, in any case, had yet to pass before Foster made personal contact with Huxley; and meanwhile, with health restored, he was able in 186I to resume the plan which had been interrupted, returning to Huntingdon to join his father in medical practice, and, being thus established, to marry Miss Georgina Edmunds in 1863. He appears to have maintained his interest in more general problems of physiology, in his use of such leisure as was left to him by the demands of a busy practice. He wrote an account of the develop- ment of observation and theory concerning the process of the clotting of the blood, in which he reflected, philosophically, that knowledge and theory move in spirals rather than in circles, so that-'Our faces may be again looking towards the same point of the compass, but we are on a higher level, and with a wider horizon.' The contacts of a country practice enabled him also to follow the lead of a curiosity about the physiology of the round worms living in the intestine of the pig, among other higher animal species. Following a clue, no doubt, from the work of Claude Bernard on the livers and muscles

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I5 of vertebrates, and finding that these worms, and their muscular body walls in particular, had a remarkably high content of glycogen, he was puzzled to find an explanation for its function there. For there was no apparent need for the heat of its oxidation to keep such a creature warm in the intestine of its mammalian host, or to provide energy for more than the apparently small muscular activity required in its pampered life. He wondered whether it could meet some energy-requirement of the process of reproduction, which seemed to be the main functional activity of an adult in such parasitic species. Such examples may provide evidence, perhaps, not only of Foster's continued interest in functional biology; they appear to me to illustrate also a char- acteristic feature of his attitude to experimental research, in that, while his contributions of new knowledge to the general stock, from his own observa- tions and experiments, may seem relatively few and simple, as coming from a man of his scientific eminence and authority, he seemed always eager to extract from them, as also from the well-authenticated evidence presented by other investigators, any stimulating suggestion which they could properly be made to yield to a well controlled speculation, and a sound scientific philosophy. It was largely, I imagine, this lively philosophical attitude which was to make him so inspiring a teacher, and to make the school which he was to establish at Cambridge, especially in his earlier years there, so richly pro- ductive of distinguished pupils, many of whom were to carry something of his special stimulus to new schools of experimental biology elsewhere in Britain, and beyond it.

Foster stayed in the Huntingdon practice for only six years. In 1867 he moved back again to University College London, accepting an opportunity to devote most of his time and interest to experimental physiology, then offered by his old teacher Professor William Sharpey, with whom he had evidently been a favourite pupil. The position offered was that of Teacher of Experimental Physiology, and he was to act also as Curator of the Museum of Pathology. Soon afterwards the holder of the Professorship of Experimen- tal Physiology, Dr George Harley, fell ill; and Foster, after lecturing for a year or two as his deputy, succeeded to the Chair when Harley retired, in 1869. Among Foster's pupils, during his short tenure of this Chair, was Edward A. Schiifer, who was eventually to succeed Sharpey at University College London, and later, as Sir Edward Sharpey-Schfifer, to be Professor of Physio- logy at Edinburgh. He has recorded a conviction that it was Foster's influence which decided him to choose a career in physiology instead of in medical practice, and that he had almost been persuaded to accompany him to Cam- bridge (4). About this time also Foster must have made his first personal

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contact with T. H. Huxley, and their acquaintance soon ripened into a close friendship. Huxley soon recognized Foster's special ability; and they evi- dently had similar views concerning the proper claims of the natural sciences as factors in education, and as contributors to the general progress of civiliza- tion. In 1869, when Huxley retired from the Fullerian Professorship of Physio- logy at the Royal Institution, Foster was appointed to succeed him; and in the following year, I870, when Huxley gave his first course of practical biology at South Kensington, he engaged Foster, Ray Lankester and Ruther- ford as his demonstrators. Foster was to retain, for many years, his association with Huxley in the South Kensington Science and Art Examinations; and, as we shall see later, he was eventually to succeed him in the Secretaryship of the Royal Society.

We must note, however, that this opening of Foster's scientific prospect, towards the end of the I860's, had come at a time of tragic change in his personal life; for in I869 his first wife died, when they had been married for only six years, leaving him a son and a daughter. In the summer of 1870, however, he was able to join his official chief Professor Sharpey in a tour, in the course of which they visited several of the German Universities, where the medical schools had already established physiology in the teaching cur- riculum, and recognized it as a subject of active research. The planning of this tour, and the nature of the scientific contacts which it enabled Foster to make, had an obvious relevance to a most important new appointment which had been offered to him earlier in that year, and which it must be assumed that he had already accepted. And this was, indeed, to bring for him a new start in his career and to provide him with his life's main opportunity as a teacher, and as an inspiring promoter of research, in what had already become his chosen field of activity.

This offer had come from the Master and Fellows of Trinity College, Cambridge, who had decided in May 1870 to appoint Foster to a Praelector- ship in Physiology at that College. The story of how such an appointment came to be made is rather too complicated to be followed here in full detail. By the time of my own student years at Trinity (I894-I898) it had even become enlivened, by what recent consultation on the history of the College has shown me to have been an essentially misleading legend. In the circle of my undergraduate contemporaries, with whom I then enjoyed the discussion of this among many other such matters, it had somehow come to be accepted as well-known fact, that the University would have created a Professorship of Physiology in 1870, with Foster as its first holder, if his conscience had not forbidden him to declare his belief in the 'Thirty-nine Articles', then still

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required by the Test Act as a condition of such a university appointment; and that accordingly, at the particular instance of an influential Fellow of Trinity, the Reverend Mr Coutts Trotter, our College had then nobly intervened, by creating a Trinity Praelectorship in Physiology especially for Foster, and thus meeting the need of the College for teaching in that subject, and, by extending the privilege to students from other colleges, that of the University as well.

Even now I must confess to some feeling of regret, at having to relinquish belief in what proves to have been an imaginary instance-it would, of course, have been easy later to find genuine ones-of Trinity's r8le as an enlightened champion of intellectual freedom, as well as of what were then the nascent claims of the experimental sciences. It was at least not implausible to imagine that, if Foster had actually been faced in 1870 by this requirement of the Test Act-not finally repealed till I87I--he might indeed have found himself unable to comply with it, on account of his nonconformist inheritance and his own mature convictions, so far as these could be inferred from his intimacy with such famous heretics as Huxley. Coutts Trotter, again, was certainly an early and staunch advocate of the claims of the natural sciences, and of physiology in particular, in Trinity and in the University as a whole. And it is further clear from a memoir which Foster published in Nature (5), on Coutts Trotter's all too early death in 1887, that they had actually met and formed a friendship, even before the question of a Cambridge appoint- ment for Foster had taken shape; and that after he arrived there, during his 13 years' tenure of the Praelectorship, and the 4 years for which he had then held the University Professorship before Trotter died, Foster had been deeply and gratefully dependent upon his advice, support and steady comradeship. Even I, eleven years later again, had reason to think gratefully of Coutts Trotter's devotion to the cause of the natural sciences. For it was my great privilege, as a callow apprentice to research in physiology, to share from 1898 to 19oo in a studentship for research in physics or physiology, founded on a bequest which Coutts Trotter made to Trinity for the purpose. Fortunately for me the tenure of this had become vacant just in time for the completion of my tripos, through the departure of the previous holder, Ernest Ruther- ford, for Montreal.

With regard, however, to the appointment which actually brought Foster to Cambridge in 1870, it is clear, in the first place, that the hindrance to any consideration by the University of an appropriate Professorship at that date, and even for 13 further years, had nothing to do with the Test Act, but was purely financial. The University had no funds of its own, or any

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prospect then in view of an endowment, which it could have applied to such a purpose. Trinity, on the other hand did not, as we students had imagined, create a Praelectorship specially to bring Foster to Cambridge. The creation of more than one such Praelectorship, primarily for the purpose of extending the resources of the College for teaching, and especially in the direction of the experimental sciences, had in fact been under discussion for a good many years-as a consequence, indeed, of a recommendation by a Statutory Com- mission appointed to give detailed effect to that of a Royal Commission, which had reported as early as I857. After years of delay, attributable, partly no doubt, to the difficulty of arriving at ready agreement in such a constit- uency, distinguished Fellows of Trinity, such as the philosopher Henry Sidgwick, had begun to exercise effective pressure towards the end of the I860's, for the claims of physics and physiology in particular. As a result, the offer of a Praelectorship in Experimental Physics had actually been made to William Thomson of Glasgow-the later Lord Kelvin; but he had found himself, for family reasons, unable to accept it. Attention appears then to have turned to the possibility of finding a suitable chemist or physiologist; and enquiries, made through personal contacts in London, had led to a recom- mendation of Michael Foster. It can hardly be doubted that Huxley's support of this suggestion would have been largely effective in its adoption by Trinity, or that Coutts Trotter's advocacy would have had an important influence on the decision of the College to offer the appointment, and also on Foster's willingness to accept it. In his history of The Cambridge Medical School (1932), the late Sir Humphry Rolleston, without quoting the source of his informa- tion, mentions also George Eliot and her special friend, George Henry Lewes, as having pressed the claims of Foster for consideration by Trinity. The Trinity records offer no evidence of this. On the other hand, Foster's circle of friends in London would have included Lewes and George Eliot; Lewes, indeed, was later to be associated with him in the group of enthusiasts who founded the Physiological Society in 1875. And we may find some contingent support for Rolleston's suggestion, in the fact that George Eliot, with her intimate knowledge of Lewes's plans and interests, commemorated him by endowing a George Henry Lewes Studentship for Research in Physiology, to be awarded by Cambridge University.

Whatever may have been the personal influences which were thus effective in the offer of the Trinity Praelectorship, and in its acceptance by Foster, the late Professor Langley was certainly justified in his conclusion that 'No election could have had more fortunate results for the University' (6). There would be no proper ground, however, for any suggestion that its sponsors

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at Trinity had in view such a development as it was in fact to produce. Their immediate object was to increase the range and efficiency of teaching in the College, and to make this, on suitable terms, available also to students from other colleges. The Praelector's stipend was provided partly from the cor- porate revenue of the College, and partly from the tuition fund, derived from the fees paid by undergraduates. Foster's programme of lectures had to be approved by the College authorities, to whom he had also to apply for grants, to meet his additional requirements in the way of equipment or staff; and the record of these makes a remarkably modest impression, when they are com- pared with those which could now be regarded as normal for the teaching apparatus alone, of a well equipped modern department; to say nothing of the much more elaborate and costly equipment which it might be expected now to need for its programme of researches. It is clear, however, that, at the time and under the conditions of Foster's migration to Cambridge, no kind of special equipment for research had been envisaged by those respon- sible for his appointment; and it can hardly be supposed that he himself had started with any specific plans for conducting or encouraging it.

In any case, he began his lectures in the Michaelmas term of I870, and the practical classes which he regarded as an even more important aspect of his teaching, with accommodation and equipment so primitive and meagre that a modern teacher or student would find it difficult to believe that any- thing serious could be attempted with it. Cambridge University, unable then to give any financial support to the enterprise, did what it could by lending the use of a room. According to Langley (6), who was a member of Foster's first class in it, this room had later become incorporated into the Philosophical Library; and this library, as I remember it, was then still housed in a single room of no great dimensions. Foster's activities then, and those of his classes in those early years from 1870, had to make do with the part of this simple structure which already existed; and this, having served as lecture room and as laboratory for the practical classes, had eventually to serve also for such experimental researches as Foster himself, and most of the members of those early classes, somehow contrived to take in hand. Trinity was responsible for equipping this room, and for any assistant salary which was needed. The initial outfit included tables for use as laboratory benches, three of which could, as required, be joined, to define the limit of the lecturer's 'platform'; together with some bottles, chemical reagents, etc., a few microscopes and items of simple apparatus. The College accounts for 1870 show that a sum of 4oo00 was granted for such initial provision, with ?80 a year for the wage of an assistant. In 1874 Foster received ?30o for the purchase of some resistance

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coils, and an equal sum for 'a recording apparatus'. Was the latter, perhaps, a kymograph? And was somebody embarking already on a simple research in electrophysiology? It may appear significant, in any case, that permission had already been given to Foster by 1873, to have the Arms of Trinity College printed on publications, describing results which had been obtained by researches in this primitive laboratory-cum-lecture-room.We may remember, however, that Claude Bernard's fundamental discoveries, as Foster was to describe them, had been made under conditions at least as primitive.

Foster showed, from the first, a determination to encourage research, and also to provide in Cambridge the scientific background required for the effective education of the students of medicine, by a course based upon direct observation and experiment. Physiology, in the examinations for the Natural Sciences Tripos, or for a medical degree, had figured till then merely as a subsidiary item, in the papers on zoology and on human anatomy respec- tively. Foster gave the Michaelmas and Lent Terms to lectures and practical work on different aspects of physiology, so widely conceived as to include embryology, for example; and for the Summer Term he offered a general course of elementary biology. At first the classes were small; but that would have given him, in the intimate personal contacts with his pupils which it made easy, an optimal opportunity for the exercise of his most brilliant faculty as a teacher-that for eliciting from his pupils an enthusiasm, to match his own, for the interest and the challenge of a scientific problem. Langley, himself a member of this class, comments that 'The majority of the small band who attended his lectures during the first three years adopted a scientific career'. And he mentions five of them in particular-Gaskell, Balfour, Dew- Smith, Sheridan Lea and himself-who, in his opinion, would probably not have done so, without Foster's inspiring influence. 'The desire to undertake research', he continues, 'was imbibed from him in lecture and in familiar talk. It was the air we breathed.' And these enterprises in research, for which Foster made his pupils so eager, were by no means restricted to such as the conventions of those days, or, indeed, those which are now generally accep- ted, would assign to the particular scope and domain of'physiology'. Foster's generous conception of the proper field of his activities evidently extended, indeed, to practically any biological problem which required the experimen- tal method for its study; whereas, before his arrival, the provision made by Cambridge for the biological disciplines had been limited, in effect, to their systematic aspects.

This breadth ofFoster's scientific interests and influence was made evident by the wide range of the subjects in which his early pupils at Cambridge

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became active and eminent. There were some, of course, who followed him in physiology, at Cambridge and elsewhere. Newall Martin, who had followed him from London to Cambridge, was to become the first Professor of Physiology in the Johns Hopkins Medical School in Baltimore. Gaskell and Langley remained in Cambridge; and, as other demands began to divert Foster's activities, they became, for a time, the effective leaders of physio- logical researches there. In succession, they had attacked the complicated anatomical and physiological problems presented by the 'involuntary' or 'autonomic' nervous system. It was Langley, however, who was to devote the remainder of his active career, with H. K. Anderson in particular among his distinguished collaborators, to the unravelling of the details of its pattern and its functions. Gaskell, on the other hand, a man of private fortune and always more of a 'free lance', had become diverted, by my time as a student, to a morphological adventure, which was to lead him, in fact, to no con- vincing discovery. Langley was in due course to succeed Foster in the Cam- bridge Chair of Physiology; but, to us who had the privilege of attending Gaskell's special lectures, all of them dealing with the historical backgrounds and the detailed courses of his own earlier researches, it seemed clear that he was the more effective transmitter of the Foster tradition, with its special genius for awakening the spirit of research, and the eagerness to undertake it.

Sheridan Lea had early shown a special interest in the chemical aspect of physiology, as presented in Foster's lectures. He was later to contribute a chemical section to an edition of Foster's famous text-book, before he became the Professor of Pathology at Cambridge; but his tenure of that Chair was to be cut short early, by fatal illness. On the other hand there were members of Foster's early class who were to win distinction and occupy university chairs in subjects further beyond the accepted range of physiology-Liver- sidge, for example, in chemistry, and Milnes Marshall in zoology. And there were others, again, who had been able first to relieve Foster of some of the teaching obligations which he had created for himself; thus Vines and Adam Sedgwick had jointly taken over the course of elementary biology, before Vines became Professor of Botany at Oxford, while Sedgwick was to become Reader and eventually Professor of Zoology in Cambridge. What had given promise of becoming one of the most famous, as well as one of the earliest, of these off-shoots from Foster's original physiology class, was embryology, the responsibility for which he had been able soon to transfer to the charge of his brilliant pupil F. M. Balfour-less known to a general public than were his brothers, Arthur and Gerald Balfour. Francis Balfour had quickly become recognized as a British leader in what was then, in many respects, a new

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direction of scientific enterprise. In due course he had been appointed to a new University Chair of Animal Morphology, at Cambridge; but then, to the deep distress of Foster, among many devoted friends and admirers, Balfour had been killed in an Alpine accident.

Foster, meanwhile, in spite of the growing calls of other kinds on his time and his interest, had not yet abandoned the attempt to continue some re- searches of his own; though these hardly amounted now to more than exten- sions of those already mentioned. In conjunction with Dew-Smith-an interested amateur, who had been one of his early pupils and was to remain a close personal friend-Foster carried somewhat further his observations on the origin of the heart-beat, and had concluded, from experiments with isolated parts of the frog's heart, that 'The well-known, easily recognized ganglia of the heart play a subordinate part in the production of the heart's spontaneous rhythmic pulsations'. At which point, we may suppose, the problem passed for a time to Gaskell. Foster himself had quite early become involved in writings and publications, which must surely have claimed most of the time which he had left, at least during term-time, after fulfilling the requirements of personal teaching as he himself had conceived them. In 1873 he contributed the section on 'Functions of Muscle and Nerve', to The Hand- book for the Physiological Laboratory, edited by John (later Sir John) Burdon- Sanderson, who had preceded Foster in Sharpey's laboratory and had suc- ceeded Sharpey in the Chair at University College London, before he became Professor of Physiology at Oxford, during Foster's tenure at Cambridge. In 1874 Foster published, with Balfour, The Elements of Embryology, dealing in a practical manner with embryonic development in the bird's egg; and, but for Balfour's early death, this was to have served as the introduction to a more systematic survey. Then, in 1876, came A Course ofElementary Practical Physiology, by Foster and Langley; and, in 1877, the first edition of Foster's great Text Book of Physiology. This made something of a new tradition for a students' text-book, with its easy and attractive literary style, and its careful presentation and philosophical appraisement of the experimental evidence. Properly used and enjoyed, it offered the student a sound lesson in scientific method, as well as a reasoned survey of the known facts and conclusions. It was widely welcomed and used in America, as well as in the country of its origin; and it was to be translated into Italian, German and Russian. Even Langley, who had by then succeeded Foster in the Cambridge Chair, writes of it in 190o7, after seeing it in several editions, that 'it will always remain a work to which the jaded physiologist can turn with interest'. In later years, apart from a number of occasional lectures and addresses, dealing with a

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range of scientific and medical themes, Foster was to publish two scholarly books of scientific and historical interest-that on Claude Bernard, already mentioned, and A History of Physiology during the 16th, 17th and 18th centuries, based upon a course of lectures which he delivered in 1900 at the Cooper College, San Francisco.

By the middle of the I870's Foster's influence, in promoting the develop- ment of physiology, was spreading beyond Cambridge. In 1875 he had joined with Burdon-Sanderson and others, including Huxley, in organizing the formation of the Physiological Society. This bore from its beginning some impress of Foster's easy and generous personality, and his methods and ideals as a teacher, in the informal friendliness of its proceedings, the priority which it accorded to practical demonstrations, and the good-humoured frankness of its discussions. These characters it has essentially retained, in spite of the recently prodigious growth of the numbers attending its meetings; and it has transmitted those traditions to the daughter societies, which have naturally budded in succession from its parent stock-Biochemical, Pharmacological, Endocrinological Societies, and others, not improbably, yet to come.

Some plan for an International Congress of Physiologists seems to have been privately discussed, on the occasion of a meeting of the British Medical Association in 1881. Foster's contribution to this discussion, as reported, would seem to have been characteristic enough; namely, to urge that such a Congress should be essentially devoted to the demonstration of experiments and methods of research, and that it should not publish any formal record of its proceedings. Nothing, however, came of this, till Foster again took the initiative, with the Physiological Society, in 1888; with the result that an International Committee was then appointed, and a first Congress was successfully held in Basel in I889, with Professor Kronecker as its President. Foster was naturally the President of one held in Cambridge, in I898; and by the time the next Congress was held, in 190IgoI, at Turin, his influence and initiative in this connexion had become so generally acknowledged, that, at the instance of its immediate President, Professor Angelo Mosso, Foster was there made Honorary Perpetual President of the International Congress of Physiologists, and presented with a congratulatory tablet.

Another direction in which Foster did an inestimable service to physiology and to experimental science in general, was in the foundation in 1878 of The Journal of Physiology, with the financial support of his friend and pupil Mr Dew-Smith, and with the scientific support of the academic leaders in physiology of those days, in the United States of America as well as in Great Britain. The first number was issued from the Cambridge University Press

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in March 1878, with Foster as its Editor. He held that responsible position alone till I894, when Langley joined him and relieved him of most of the editorial routine; and, after passing to Langley's full control when Foster died, the Journal eventually became, and remains, the property of the Physio- logical Society, with an Editorial Committee which the Society appoints. Its output, like that of all such standard scientific journals, has naturally undergone a great and progressive enlargement, in bulk and in the range and complexity of the researches which it presents. I do not think, however, that it is altogether fanciful, to feel that it still retains something of the special character which it so long ago acquired, from its founder and first editor.

It is evident that, after his arrival in Cambridge in 1870, Foster had become quickly and easily acclimatized to its special academic and social atmosphere. The University made him an Honorary M.A. in 1871, and Trinity elected him in the same year to a Fellowship, in addition to his Praelectorship. Then, in 1872, he made a second marriage, with Miss Margaret Rust of Huntingdon, who became his constant companion in their home and on his travels abroad, and shared with him in the welcome extended to his colleagues, friends and visitors from other countries, in the house which they built some miles out of Cambridge, near Great Shelford, on a hillside of the Gog-Magog range. There Foster planted shelter trees and made a beautiful garden, to the care and development of which he devoted a large part of such leisure as he could contrive to keep from the growing claims, academic and public, on his scientific experience, judgement and initiative. His success as a gardener brought him, indeed, another kind of reputation, extending far beyond the local and academic ranges; for he became widely known in horticultural circles, as a producer of new and attractive hybrids, of Irises especially. I am indebted to Sir George Taylor, Director of the Royal Botanic Gardens, Kew, for the following estimate of Foster's achievement in scientific and practical horticulture:

'Sir Michael Foster's interest in horticulture was real and practical. Not only was he prominent in the affairs of the Royal Horticultural Society and by his efforts and advice largely responsible for restoring its fortunes at a time when the future of the Society was insecure, he was assuredly a very knowledgeable and devoted gardener. For many years he was the accepted leader in the cultivation of irises and the acknowledged international authority on the genus. In his letters to Sir Joseph Hooker and Sir William Thiselton- Dyer preserved at Kew there are many references to his Iris investigations and also to the Royal Horticultural Society. He grew many Iris species in his garden at Shelford, a considerable number being new introductions which he

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acquired through his personal contacts with botanists and plant collectors overseas. Of the new species and varieties which he described several retain their rank and he also was responsible for raising a number of interspecific hybrids. Among his many contributions to Iris literature, his most substantial work was an account of the bulbous irises based on a lecture given to the Royal Horticultural Society and published as a separate brochure in 1892. Foster's studies in Iris could reasonably have led to a monograph of the genus but, according to W. R. Dykes, he was reluctant to prepare such a com- prehensive work. It was left to Dykes to do so and in The Genus Iris (1913) the debt to Foster, who encouraged the author to investigate the group and placed his plants and knowledge unstintedly at his disposal, it generously recognized.'

My own six University years at Cambridge, as undergraduate and research student, came too late for any enjoyment of such close personal contacts with Foster, as those of which we heard such glowing reminiscences from the teachers and demonstrators of our time, who had studied and begun their researches when he still had freedom and opportunity to take interest in their individual doings-men of the generation of W. B. Hardy and H. K. Ander- son. Charles Sherrington, the greatest of all Foster's physiological pupils, had left for London some ten years before my time at Cambridge, so that I did not know him until later; but he too was to recall that Foster 'took a genuine interest in younger men and their work, and he rejoiced in their scientific achievements with an almost paternal delight' (4). 'I love the Professor', I remember Hugh Anderson saying to me; 'there is still nobody like him as a critic of one's researches.' By the time I entered Trinity, however, in I894, the classes for the ordinary course in physiology, for Tripos Part I, had already become so large, that contacts by Foster with individual students, if any could still have been attempted, would have had to be highly selective. The outside claims on his time and activities, moreover, had already reached the stage at which it seemed to us normal, as we surged out of the theatre after a lecture, to see the Professor climbing into a waiting hansom cab, to catch the train which would take him to duties in London for the Royal Society and, in increasing measure, for the Government of the day. Under these various conditions, it was hardly to be expected that his lectures would have retained all their legendary sparkle of interest and inspiration. In fact, to a diligent and retentive student of the physiology so well presented in Foster's Text-Book, lectures which had come to consist mostly, and I suppose almost inevitably, of passages from that familiar work, rather monotonously recited by its author, with a steady rhythm of gesture for emphasis, had few of the attrac-

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tive features of earlier days. They disappointed my expectation; and even now, in long retrospect, I cannot rid myself of a feeling that, by my time as a student, it would have been better for Foster to have handed over the elementary course of physiology to others, and to have devoted his own teaching activity to more advanced lectures, on chosen physiological themes, in which he surely would still have been able to transmit some of the inspira- tion, which his earlier pupils remembered with such enthusiasm.

The Royal Society had elected Foster to its Fellowship in 1872, when he was 36 years old, two years after his migration to Cambridge. Nine years later, in 1881, he had become one of its Secretaries, in succession to his friend, T. H. Huxley; so that he had already been serving in that capacity for thirteen years, at the time which I have been recalling, when I became a Cambridge freshman. It was not, of course, to be expected of a man with such a width of interests as Foster's, that he would be content to limit his activities, as an Officer of the Royal Society, to those required by its more immediate routine, or even to matters concerned with his own expert knowledge and main scientific interests, in experimental biology and its medical applications. We find records, accordingly, of his activities, doubtless in co-operation with his opposite numbers in the Secretaryship, in the organization of the Meteoro- logical Office, and the establishment of The National Physical Laboratory; of which we may regard the latter as an important landmark, representing an early recognition, by a British Government, of a national obligation to give support to fundamental researches in science, even beyond the range of any visible need, or opportunity, for practical applications. Foster was also active in starting the International Congress of Geodesy-one of a number of enterprises, indeed, on behalf of the international organization of scientific activities and information, with which he became specially identified. The Royal Society, through its Council, had become the recognized adviser of the British Government, and the normal channel of application to it, con- cerning grants-in-aid of scientific expeditions, dealing with a variety of natural objects and phenomena-coral reefs, eclipses, earthquakes, countries then still unexplored, etc. And it evidently became recognized that, in dealing with the Treasury and with other departments appropriate to particular applica- tions, Foster had the instincts and the experience of an acceptable negotiator, and, more generally, of an effective representative to the Government of the scientific authority of the Royal Society. In due course he had been made a member of several Royal Commissions, especially of those dealing with problems of medical importance-Vaccination, Disposal of Sewage, Tuber- culosis; of the last of these, as already mentioned, he was the Chairman. And

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he was made Chairman also of what was to be a very important departmental committee, appointed by the Colonial Secretary to investigate, and to make recommendations on, 'the prevention of malaria and other tropical diseases'. This field of research, in tropical medicine, was one in which British investi- gators-several of them having, and using to great purpose, their special opportunities as officers in the medical services of the British armed forces and of the Indian Government-were able for a time to give an important lead to the rest of the scientific world. Foster's great services, in these various capacities, were recognized in 1899 by the award ofa K.C.B.

With all these claims on his time, I had no reason to wonder why, even in my two last Cambridge years, as a research student from 1898 to 1900, I was able to make hardly any direct contact with Foster, who, indeed, was becoming so loaded with other duties than those belonging to his Professor- ship, that he applied in 19oo for the appointment of Langley as his deputy, until he retired from the Chair three years later, in 1903, after completing a 20 years' tenure of it. In the same year he retired also from his Royal Society Secretaryship, having held it for 22 years. Meanwhile, however, he had done a last, and certainly one of the greatest of all his many services to natural sciences at the University. This was his invitation to Frederick Gowland Hopkins to come to Cambridge, in order to make good a serious deficiency which had developed, in the chemical aspect of teaching and research in physiology. The characteristic manner in which Foster made this proposal to Hopkins, and the major part which its acceptance was eventually to play in the emergence of biochemistry as an independent, but now so widely per- vasive, scientific discipline, has recently been the subject of a brilliant contri- bution to these Notes and Records by one of Hopkins' early Cambridge pupils, Joseph Needham (7). Here I may, perhaps, allow myself to emphasize the extent to which a decisive event in Foster's career was to be nearly repeated in that of Hopkins; in that Trinity College stepped into a breach, actually threatened in this case by a breakdown in Hopkins' health. Trinity then appointed him also to a Praelectorship, of course in Biochemistry, and elected him likewise to its Fellowship.

The Secretaryship of the Royal Society, was, of course, by far the longest in duration, of the services which Foster rendered to the more general interests of natural science; and, together with others which arose from it more or less directly, it was obviously the one which was eventually to entail for him the greatest diversion of energies and interests from his Cambridge Professorship and from physiology. It was by no means the only one, however. As early as 1859 he had joined the British Association, when, as already mentioned, he

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made the first of all his scientific communications to its annual meeting, at Aberdeen. He was to become one of its General Secretaries from 1872 to 1876; President of its Section of Physiology in 1897; and President of the Associa- tion in 1899, when he was still holding his Royal Society Secretaryship. One can further understand his need to apply in the following year for a Deputy in his Cambridge Chair, when it is realized that he had then, in 19oo, incurred yet another major distraction from its duties, by accepting election as the Member of Parliament for the University of London. The House of Com- mons, however, with its discipline of Party Whips, was evidently unsuited to his scientific independence. He had entered it as a Liberal Unionist; but when the Government claiming his allegiance introduced its Education Bill, in connexion with which Foster might have been expected to find his con- structive opportunity, his principles obliged him to vote against the clauses dealing with religious instruction. And then, on the Government's Tariff Reform proposals, he crossed the floor of the House, to sit and vote with the Opposition. It was hardly a matter for surprise that, though he stood again, he was not re-elected.

In rather striking contrast to the humdrum tendency of his later routine lectures to students, Foster's numerous occasional addresses, and his lectures on special subjects, had won him a well justified and growing reputation for the easy grace of their style, as well as for the intellectual distinction of their contents. This faculty served him well, of course, in connexion with the variety of calls and opportunities which came to him, many of them in connexion with his long tenure of the Royal Society Secretaryship. In lighter vein also, he won a wide and, doubtless, a well justified reputation, as an after-dinner speaker. According to Langley's estimate of his success in this graceful and ingenious art, he 'at his best, was in the very first rank'. W. B. Hardy, as I recall, was another who savoured his memories of Foster, as incomparably happy and successful on such occasions. In a different connexion, his friend Dr Pye Smith wrote of him that 'Few public speakers knew better how to express sympathy or sorrow in dignified and gracious terms.

We have seen how an early threat of illness had disappeared, and it seems to have left no mark; for the equability of Michael Foster's health, till a few years from the end of his life, must have been an essential factor in the great extent and the high distinction of his achievement. In 190o4, however, he suffered such pain, attributed to dyspepsia, that work and enjoyment were almost suspended. He seems then to have found sufficient relief to become normally active again; and it was not till January 1907 that, having spoken to the British Science Guild in the afternoon, he became acutely ill in the

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evening through the bursting of an oesophageal ulcer. Being in London, he was attended by his friend Sir John Rose Bradford-who was to become a Secretary of the Royal Society in the following year. The condition was beyond remedy, and the end came quickly.

The death of Sir Michael Foster brought a deep sense of bereavement to his devoted pupils, and to a much wider circle of many friends and admirers, attracted by the spontaneous warmth of his generous personality, and by a recognition of what his life of devotion had done for physiology, for experi- mental biology, and for the claims of the natural sciences as a major com- ponent of a reasonable culture, and as a central factor in the progress of civilization. 'In Cambridge', as Professor Langley wrote, 'it will not be forgotten that the modern developments of Biological Science in the Uni- versity are primarily due to Sir Michael Foster.' And the Royal Society will surely remember him, as one of the most enterprising and successfully active and, indeed, in every sense, one of the greatest of its Secretaries.

My own direct contacts with Sir Michael Foster having been but few, and late in his career, I have obviously depended for much of this account of him on memories of talks with my seniors and teachers, and on several good biographical memoirs-those by the late Professor Langley in the Journal qf Physiology and the Dictionary of National Biography, and those which appeared in The Lancet, I907 i, p. 349, and the British Medical Journal, I9o7 i, p. 397. In addition, I have been greatly indebted to Lord Adrian, and especi- ally so to Dr R. Robson, Fellow of Trinity College, Cambridge, and learned in the records of its history, for what was to me so illuminating an account of the academic background, and of the personalities involved in the offer by the College of the Praelectorship to Foster, and of the manner of its full support of his physiological enterprise in Cambridge, during its first thirteen years. I hope that I have made a reasonable use of the information so gener- ously given.

The charming, informal portrait of Sir Michael Foster used in illustration of this record, was recently discovered in the Bowditch Library at the Har- vard Medical School by Dr J. R. Pappenheimer, and by him sent to Lord Adrian, who was kind enough to lend it to me for reproduction, before adding it to the collection of Foster material in the Library of Trinity College, Cambridge. According to Dr Pappenheimer it was probably taken at the Putnam Camp, of which Bowditch was one of the founders, in the Adiron- dack Mountains, in connexion with Foster's visit to the U.S.A. to deliver his Lane Lectures on the history of physiology, at the Cooper College, San Francisco. That was in I900oo, and the portrait certainly represents Foster's

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appearance at about that date, three years before his retirement from the Cambridge Chair, and from his Royal Society Secretaryship. In his letter to Lord Adrian, Dr Pappenheimer writes that:

'It probably took Bowditch and Foster two or three days to reach Put- nam Camp from Boston-a day in the train, a day's drive with horses and supplies, and almost another day's walk into the camp. Having gone to this trouble, it would hardly be worth while to spend less than two weeks at the camp. It seems to me, from looking at this photograph, that Foster must have come to know his host pretty well; he probably learned more of American physiology in this delightful and leisurely fashion than his modern counterpart could learn from a dozen hurried trips by jet.'

Apart from the note on Sir Michael Foster as a gardener, which Sir George Taylor was kind enough to let me have for inclusion in this Memoir, he has been so good as to review the list of papers on Horticulture in the Biblio- graphy. And I have further been greatly indebted to Mr N. H. Robinson, the Assistant Editor of these Notes and Records, for much help in the preparation and completion of the Bibliography as a whole.

NOTES

(i) Claude Bernard, by Michael Foster, M.A., M.D., D.C.L., etc., T. Fisher Unwin, 1899. (In the Series-Masters of Medicine.)

(2) Proc. Roy. Soc., Io, 523 (I859-60). (3) Sir Landsborough Thomson-Original Development of the Medical Research Council.

Brit. Med.]., 1963, ii, p. 1290. (4) Obituary Notice of Foster. Lancet, 1907, i, p. 351. (5) Nature, Lond., 37, 153 (1887). (6) In memoriam-Sir Michael Foster.J. Physiol., 35, 233-246 (1907). (7) Notes and Records Roy. Soc. Lond., 17, 117(1962).

OBITUARY NOTICES OF FOSTER

Proc. Roy. Soc., B, 8o, lxxi-lxxxi (1908). Proc. Linn. Soc. Lond. [by A. E. Shipley] 1906-7, pp. 42-5. Kew Bull. 1907, pp. 65-6. 'Michael Foster-a recollection' by W. T. Thiselton-Dyer. Camb. Rev., 1907, pp. 439-4o. Gdnrs' Chron., ser. 3, 41, 78 (1907). Brit. Med. J., 9 Feb., 1907. Lancet, 1907, i, p. 351. 'In memoriam-Sir Michael Foster' [byJ. N. Langley]. J. Physiol., 35, 233-46 (1907).

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BIBLIOGRAPHY

Books

Report on modern microscopes, and recent improvements in microscopical apparatus. London, I1867. 'Function of muscle and nerve.' Handbook for the Physiological Laboratory. Edited by J. Burdon-

Sanderson. London, 1873. The elements ofembryology. (With F. M. Balfour.) London, 1874. The elements ofembryology. (With F. M. Balfour.) Second edition. London, 1883. A course ofelementary practical physiology. (WithJ. N. Langley.) London, 1876. A text book ofphysiology. London, I1877. A text book ofphysiology. Sixth edition. London, 1893-19oo. Science primer ofphysiology. London, 1890. Bulbous Irises. London [1893]. Physiology for beginners. (With L. E. Shore.) London, 1894. A course ofelementary practicalphysiology and histology. (WithJ. N. Langley.) London, 1896 A course of elementary practical physiology and histology. (With J. N. Langley.) Seventh edition.

London, 1902. Life of Claude Bernard. London, 1899. Lectures on the history of physiology during the sixteenth seventeenth and eighteenth centuries. (Lane

Lectures, Cooper College, San Francisco.) Cambridge, 19ol.

Papers on Physiology and General Science 1859-60. On the effects produced by freezing on the physiological properties of muscles.

Proc. Roy. Soc., xo, 523.

1860. On the beat of the snail's heart [1859]. Abstract p. I6o. Rep. Brit. Ass. 1864. The coagulation of blood (Review). Nat. Hist. Rev. IV, 157. 1865. On the existence of glycogen in the tissues of certain Entozoa. Proc. Roy. Soc., 14,

543. 1867. Notes on amylolytic ferments.]. Anat. Lond., I, 107. 1869. On some points of the epithelium of the frog's throat.J. Anat. Lond., 3, 394- 1869. Note on the action of the interrupted current on the ventricle of the frog's heart.

J. Anat. Lond., 3, 400oo. 1870. On imbedding substances for microscopic section. Quart.J. Micr. Sci., xo, 124. 1871. On the physiological action of the codeia derivatives. Proc. Roy. Soc., 19, 510o. 1872. Ueber einen besonderen Fall von Hemmungswirkung [1871]. Arch. Physiol., 5,

191.

1874. On the effects of a gradual rise of temperature on reflex actions in the frog. Camb. Stud. Physiol. Lab., I, 36.

1875. (With A. G. Dew-Smith.) On the behaviour of the hearts of mollusks under the influence of electric currents. Proc. Roy. Soc., 23, 318.

1876. (With A. G. Dew-Smith.) The effects of the constant current of the heart.J. Anat. Physiol., Io, 735.

1876. On some conditions of reflex action [1873]. Proc. Camb. Phil. Soc., 2, 309. 1876. On the use of the term 'endothelium' [I874]. Proc. Camb. Phil. Soc., 2, 340. 1876. On the effects ofupas-antiar on the heart [1875]. Proc. Camb. Phil. Soc., 2, 398. 1876. Some effects ofupas-antiar on the frog's heart.]. Anat. Physiol., Io, 586.

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1877. (With A. G. Dew-Smith.) Die Muskeln und Nerven des Herzens bei einigen Mollusken. Arch. Mikr. Anat., 14, 317.

1893. Weariness. Nature, Lond., 48, 178. 1894. The organisation of science. Int. Congr. Med., Vol. I, p. 246. 1894-95. On the teaching of physiology in schools [1895]. Nature, Lond., 51, 487. 1896. The Huxley lecture. Recent advances in science and their bearing on medicine and

surgery. Nature, Lond., 54, 580. 1897. [Presidential address to the Physiol. Sect.] Physiology between 1884 and 1897.

Rep. Brit. Ass., p. 798. 1898. On the physical basis of psychical events. Mem. Manch. Lit. Phil. Soc., No. 12, 46

PP. 1899. [Presidential address to the British Association.] Rep. Brit. Ass., p. 3.

Papers on Horticulture

1883. Iris bartoni [n. sp.]. Gdnrs' Chron., n.s. 19, 275. 1883. Notes on irises. Gdnrs' Chron., n.s. 20, 231. 1884. Some new varieties of Iris. Gdnrs' Chron., n.s. 22, 524. 1885. Iris stylosa (unguicularis), var. alba. Gdnrs' Chron., n.s. 23, 340. 1885. New garden plants. Gdnrs' Chron., n.s. 23, 438. 1885. The reticulate group of irises. Gdnrs' Chron., n.s. 23, 567. 1886. Iris cengialti. An attempt towards garden nomenclature. Gdnrs' Chron., n.s. 25, 554. 1887. Some new irises. Gdnrs' Chron., ser. 3, I, 611. 1888. Hybridisation. Gdnrs' Chron., ser. 3, 3, 151. 1888. Freesias. Gdnrs' Chron., ser. 3, 3, 588. 1888. Iris korolkowi. Gdnrs' Chron., ser. 3, 4, 36. 1888. (WithJ. G. Baker.) Irises. Gdnrs' Chron., ser. 3, 4, 182. 1889. Iris caucasica and I. orchioides. Gdnrs' Chron., ser. 3, 5, 588. 1889. On irises.J. Hort. Soc., II, 131. 1892. Iris loreti (Barbey). Gdnrs' Chron., ser. 3, 12, 152. 1893. New or noteworthy plants. Gdnrs' Chron., ser. 3, 13, 711. 1895. New or noteworthy plants. Gdnrs' Chron., ser. 3, 17, 612. 1899. New or noteworthy plants. Gdnrs' Chron., ser. 3, 25, 225. [899. New or noteworthy plants. Gdnrs' Chron., ser. 3, 26, 389. 1901oI. Iris ewbankiana. Gdnrs' Chron., ser. 3, 29, 397. 1902. Iris aschersoni. Garden, Lond., 61, 288. 1902. New Irises. Gdnrs' Chron., ser. 3, 31, 385. 1902. Iris leichtlini. Gdnrs' Chron., ser. 3, 32, 242. 1903. The identity of Iris hookeri and the Asian L setosa. Rhodora, 5, I57. 1905. A remarkable hybrid Narcissus. Gdnrs' Chron., ser. 3, 37, 82. 190o6. Iris (Xiphion) taittii. Gdnrs' Chron., ser. 3, 4o, I45.

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