Foster, Michael

(b. Huntingdon, England, 8 March 1836; d. London, England, 28 January 1907)

physiology.

Foster was descended from a well-known family of religious Nonconformists who had farmed for many generations in Hertfordshire and Bedfordshire. His father, also named Michael, broke the yeoman tradition and became a medical practitioner. From 1831 to 1833 the elder Michael Foster studied medicine at the University of London (later University College), where he won many prizes. He practiced at Huntingdon from 1833 and was created fellow of the Royal College of Surgeons in 1852. A fervent Baptist, he was a prominent religious and civic leader in Huntingdon.

The younger Michael Foster, the eldest of ten children, was educated at the local grammar school in Huntingdon until 1849, when he was sent to University College School in London. In 1852 he entered University College, from which he graduated B.A. in 1854, placing first on the honors list in classics and receiving the college scholarship in that faculty. Foster might have pursued a career in classics at the University of Cambridge, but religious tests prevented his competing for a fellowship there. Instead, he immediately entered the medical school at University College, where in 1856 he won gold medals in anatomy and physiology and in chemistry. He graduated M.B. in 1858 and M.D. in 1859, and spent part of the next two years studying clinical medicine in the Paris hospital schools.

In the autumn of 1860, after impairment of his health led to fears of consumption, Foster signed on as ship’s surgeon of H.M.S. Union, which was bound for the Red Sea to take part in the building of a lighthouse near Mt. Sinai. He signed on partly for the sake of his health and partly in the hope of studying the natural history of the area. After his attempts to make such studies were repeatedly discouraged or prevented, he returned to Huntingdon in March 1861. His health remained uncertain for many years and contributed to his later decision to leave London for Cambridge. From 1861 to 1866 he practiced medicine with his father in Huntingdon, but during all this time he longed for a career in science.

In January 1867, at the invitation of William Sharpey, Foster became instructor in practical physiology and histology at University College. In 1869 he was promoted to a professorship in the same subject. Meanwhile he had established a course of laboratory instruction, including elementary experimental physiology, that was the first of its kind in England. Also in 1869 he was appointed Fullerian professor of physiology at the Royal Institution.

In May 1870, Foster was appointed to a newly established prelectorship in physiology at Trinity College, Cambridge. In choosing both physiology as the subject of the prelectorship and Foster as the man to fill it, the Trinity seniority followed the recommendation of Thomas Henry Huxley. Foster remained prelector at Trinity College until 1883, when he was chosen to occupy the first chair of physiology in the university. Upon his resignation in 1903, the chair went to his former student John Newport Langley.

When Foster arrived at Cambridge in 1870, the biological sciences and the medical school were largely moribund. Neither Charles Babington, professor of botany, nor Alfred Newton, professor of zoology, was particularly receptive to the movement toward laboratory training in biology, and both were indifferent teachers. From the first Foster was determined to change this situation and to build a great school of biology and physiology at Cambridge, even though his original position made him responsible only to Trinity College. Before teaching his first class, Foster obtained the consent of the Trinity seniority to open his course to all students in the university. The university responded by giving him the use of one small room, which was furnished with the basic necessities by Trinity College. This accommodation very soon proved inadequate as Foster attracted ever larger numbers of students to his courses in physiology and elementary biology. New buildings were completed in 1879 and 1891, but even these were becoming overcrowded by the time Foster retired. From 1870 to 1883, the number of students attending his courses in physiology grew from about twenty to 130, while the number attending his course in elementary biology grew from about forty-five at his initial class in 1873 to more than eighty in 1883. In that year Foster turned the latter course over to two of his former students, and enrollment continued to grow dramatically. He then concentrated on the courses in physiology, which were drawing about 300 students when he retired in 1903.

Like other great teachers, Foster is probably best remembered for his students, many of whom remained at Cambridge to develop the principles and programs he had inaugurated. On leaving London for Cambridge in 1870, Foster had invited his two favorite students at University College to join him. Edward Sharpey-Schafer declined, on his father’s advice, and remained at University College as assistant and later successor to John Burdon-Sanderson in the Jodrell chair of physiology. H. Newell Martin joined Foster at Cambridge and served as his righthand man until 1876, when he was called to the United States as first occupant of the chair in biology at the newly established Johns Hopkins University. Martin thus carried to America the methods of teaching he had learned from Foster and Huxley.

While at Cambridge, Foster attracted a group of students as remarkable for the breadth of their interests as for their later eminence. Apart from physiologists, they include the embryologist Francis Maitland Balfour; the biologist G. J. Romanes; the anthropologist A. C. Haddon; the psychologist C. S. Myers; the neurologist Henry Head; the pathologist J. G. Adami; the botanists S. H. Vines, F. O. Bower, and H. Marshall Ward; and the morphologists A. Milnes Marshall, Adam Sedgwick, D’Arcy Wentworth Thompson, and A. E. Shipley. Of these, Balfour, Vines, Haddon, Ward, Sedgwick, and Shipley became leading members of the Cambridge faculty; and through them Foster remained a living influence on Cambridge biology long after his own direct role had come to an end.

But Foster was above all else the founder of the Cambridge School of Physiology, and the eminent physiologists trained while he was there are his chief contributions to science. Three of his earliest students were John Langley, Walter Holbrook Gaskell, and Arthur Sheridan Lea. Except for brief periods of study in Germany, all three remained at Cambridge throughout their careers, and all three were elevated to university lectureships in physiology when Foster was appointed to the professorship in 1883. Especially through the work of Langley on glandular secretion, of Gaskell on heart action, and of both on the involuntary nervous system, Cambridge soon became recognized as one of the world’s leading centers for physiological research.

The list of physiologists trained at Cambridge later in Foster’s career is one of almost staggering eminence. It includes Charles Scott Sherrington (matriculated at Cambridge in 1879, major work on reflexes and the integrative action of the nervous system); W. B. Hardy (1884, colloid chemistry); Walter Morley Fletcher (1891, muscle metabolism); Joseph Barcroft (1893, blood gases, respiration, and homeostasis); Henry H. Dale (1894, chemical transmission of nerve impulses); T. R. Elliott (1896, sympathomimetic drugs); and Keith Lucas (1898, conduction of nerve impulses). Sherrington and Dale went on to win the Nobel Prize in physiology or medicine in 1932 and 1936, respectively. Hardy, Fletcher, Barcroft, and Lucas joined the staff at Cambridge, with Barcroft succeeding Langley as professor of physiology in 1925.

Because Foster epitomizes the concept of a great teacher, it is interesting to consider his approach and the reasons for his success. He owed much to William Sharpey, who taught him at University College and first aroused his interest in physiology, and to Thomas Huxley, who very early perceived the remarkable kinship of mind and spirit between Foster and himself. Both taught Foster that physiology should be viewed broadly, as one of the biological sciences, and both encouraged his appreciation of the experimental approach, although neither made much use of that approach in his own work. Of the two, Huxley’s influence was the more lasting and profound.

Foster first met Huxley in 1856, when the latter examined him in anatomy and physiology at University College. From then on, Huxley was his main guide and chief agent. The two rarely, if ever, disagreed on any issue of substance, and their educational philosophies are virtually indistinguishable. Both Huxley and Foster insisted that science must take equal place with mathematics and classics in the English educational system, and both urged students to undertake original laboratory research at an early stage. There are even two striking parallels in the course of their careers: Foster succeeded Huxley as Fullerian professor of physiology in 1869 and as biological secretary of the Royal Society in 1881.

In the summer of 1871, when Huxley introduced a laboratory course in elementary biology to a group of schoolmasters at South Kensington, he selected Foster as the first of his demonstrators. After a second summer as Huxley’s demonstrator, Foster established his own one-term course in elementary biology at Cambridge in 1873. Both courses were taught on evolutionary principles, with a very few organisms being dissected and studied as representative “types.” Huxley’s course, which was the model, is considered the origin of the modern method of teaching introductory biology. Foster’s very similar course seems to have been the first such course taught in a true university setting. It illustrates his broadly biological approach to physiology and helps to explain the breadth of his influence on Cambridge biology and physiology, for Foster designed this course in such a way that it became the standard means by which students were introduced to all of the biological sciences at Cambridge. In this way he was able to exert an influence on students who became botanists or morphologists as well as on those who became physiologists.

Another important factor in Foster’s success was that the University of Cambridge was in a state of transition when he arrived. After decades of defending the virtual monopoly enjoyed by mathematics and classics at Oxford and Cambridge, many Cambridge dons were ready at last to give a more sympathetic hearing to the advocates of science and other “modern” studies. It was an excellent opportunity for Foster, who had a clear vision of how much could be done and how to go about doing it. With the support of several influential allies, especially G. M. Humphry, professor of anatomy, and Coutts Trotter, a fellow of Trinity College who was a leading force in university administration, he pressed the claims of physiology upon the university with remarkable success for a mere collegiate lecturer. That this success depended in part on a new attitude in the university at large is suggested by the comparable success of the famous Cambridge School of Physics, whose development paralleled almost exactly that of the School of Physiology.

Foster’s achievement at Cambridge depended also on his own great clarity of aim and charm of manner. Even more important was his capacity for inspiring others to undertake original research. Many of his students have testified that they chose a career in research only because Foster promoted the enterprise with so much enthusiasm and with such a sense of adventure. Because Foster’s own contributions to original research were few and largely ignored, he has acquired a reputation as a discoverer of men rather than of facts, as a great teacher of research who did not himself practice what he so effectively taught. But it is crucial to recognize that Foster did engage in original research. Throughout his brief career in research, the problem of the heartbeat held a special fascination for him.

The issue to be settled about the heartbeat was whether it depended ultimately on nerve discharges or rather on an inherent rhythmicity in active cardiac muscle—in other words, whether the heartbeat was neurogenic or myogenic. When Foster began his work, the issue had apparently been decided in favor of the neurogenic theory, especially because a number of German investigators had found that in the vertebrate heart, nerve ganglia were concentrated precisely in those regions of the cardiac tissue where excision or ligature disturbed the spontaneous rhythm of the heartbeat.

In 1859 Foster found that even very tiny pieces cut from the beating heart of a snail continued to beat rhythmically for some time. But no ganglia had been found anywhere in the snail’s heart; and even if they were there, it seemed impossible to Foster that they could be so widely diffused as to appear in each and every part of the heart. He therefore concluded that, in the snail at least, the rhythmic beat must depend not on nerve ganglia but on the inherent properties of the general cardiac tissue.

In 1869 Foster published his first challenge to the neurogenic theory in vertebrate hearts. He showed that the lower two-thirds of a frog’s ventricle, where no ganglia were known to exist, could nonetheless be induced to rhythmic pulsations by direct application of an interrupted current of appropriate strength. He therefore argued that the frog’s cardiac musculature, like the snail’s, must possess an inherent tendency to rhythmic pulsation.

Returning in 1871 to the snail’s heart, Foster showed that when weak currents were applied directly to its muscular tissue, the result was an inhibition exactly like that produced in the vertebrate heart by stimulating the vagus nerve. Since in the ganglionfree snail’s heart this inhibition could not be the result of any nervous mechanisms, Foster attributed it instead to the direct effects of weak electrical currents on the fundamental properties of active contractile tissue.

Between 1875 and 1877 Foster published four papers on the heartbeat, three of them in collaboration with his student A. G. Dew-Smith. Two of these were elaborate extensions of Foster’s own earlier work on the snail’s heart and on the frog’s. The third, in 1877, was written in reply to a German scientist’s claim that he had detected ganglia in the snail’s heart. Foster and Dew-Smith argued that what the German had taken for ganglia were in fact pyriform connective-tissue cells.

The fourth paper, by Foster alone, concerned the effects of the poison upas antiar on the frog’s heart. He emphasized the remarkable effect that vagus stimulation produced in a heart thus poisoned. The eventual result was not the usual inhibition but an opposite accelerating effect, which increased as the influence of the poison increased. Again Foster rejected the idea that nerve ganglia were somehow responsible. Antiar was known to be a muscular poison, and if it were admitted that vagus inhibition also resulted from direct action on the cardiac musculature, then the marked acceleration eventually observable under their combined influence could be explained simply as an exaggerated reaction by the muscle tissue to a previously exaggerated inhibition.

In all of this, the general trend of Foster’s conclusions is clear. The causes both of inhibition and of the rhythmic heartbeat itself were to be sought not in nerve ganglia but in the basic properties of contractile tissue. Less obvious, but very definitely present in two of the papers with Dew-Smith, is the evolutionary basis for Foster’s conviction. Impressed by the rhythmic capacity of undifferentiated protoplasm—in amoebas, in ciliates, and in the simple snail’s heart—he saw no reason why differentiated nerve ganglia should be necessary for the same function in higher organisms. Apart from some ambiguity about whether the general cardiac tissue was purely muscular or rather neuromuscular in character, the chief difficulty in Foster’s scheme was his concession that in the frog (and presumably other vertebrates) the ganglia might serve to coordinate the sequence of beats. In the undifferentiated and ganglion-free snail’s heart, Foster attributed coordination to a muscular sense inherent in the general cardiac tissue, so by his own evolutionary criteria, differentiated ganglia should not have been required to perform the same function in higher organisms.

By 1900 the myogenic theory had largely replaced the neurogenic, but Foster’s work was for a long time generally ignored and was not itself greatly influential in changing the direction of the debate. It is nonetheless important for at least three reasons. First, it reveals Foster as a competent research physiologist and suggests that this experience may have been a crucial factor in his success as a teacher. Without some such experience, it seems unlikely that he could have promoted research so convincingly or that he could have developed the critical acumen so essential to a director of research. Second, the problem of the heartbeat was the starting point or focus for much of the research carried out by the Cambridge School of Physiology during its crucial early years. Under Foster’s direction Lea, Langley, Romanes, Francis Darwin, and Gaskell were all attracted to the problem during the 1870’s. Like Foster himself, Romanes and Gaskell saw the problem of the heartbeat as part of the more general problem of rhythmic motion, and they pursued the issues with significant success. Although the others rather quickly found different research interests, their work on the heart was crucial in establishing a research tradition in physiology at Cambridge. Third, Foster’s work on the heart provides a concrete illustration of his broadly biological and evolutionary approach to physiology and suggests again that this approach may have contributed greatly to his success and to that of the embryonic Cambridge School of Physiology. This seems even more likely because Gaskell, who did more than anyone else to resolve the problem in favor of the myogenic theory, depended very heavily on the evolutionary approach he had learned from Foster.

Foster’s impact on physiology and science extended far beyond his contributions as original investigator and as founder and director of the Cambridge School of Physiology. His name was attached to several important textbooks, including the pioneering Handbook of the Physiological Laboratory, edited by John Burdon-Sanderson (1873), for which he wrote the section on nerve and muscle; The Elements of Embryology, which he published with his student F. M. Balfour in 1874; and A Course of Elementary Practical Physiology and Histology, which he published with Langley’s assistance in 1876. The first edition of his famous Text-Book of Physiology appeared in 1877. Distinguished for its literary style, balanced judgement, and evolutionary perspective, this work was translated into Russian, Italian, and German. It went through six complete editions and part of a seventh.

Foster was also the leading figure in the professionalization of physiology in Victorian England. A conspicuous opponent of popular antivivisection sentiment, he was largely responsible for the founding in 1876 of the British Physiological Society. In 1878 he founded the Journal of Physiology, which he edited until 1894.

As he became increasingly occupied with these and other organizational activities, Foster abandoned original research in physiology and delegated to former students most of the responsibility for teaching the advanced classes at Cambridge and for directing the day-to-day activities of the research laboratory. By about 1880 the shift was complete. Foster concentrated thereafter on elementary teaching and on exercising his talent for organization. At Cambridge he led the fight for the establishment of a school of scientific agriculture in the 1890’s. In 1898 he induced Frederick Gowland Hopkins to join his staff at Cambridge as teacher of biological chemistry. Hopkins went on to become founder of the Cambridge School of Biochemistry and to share the Nobel Prize in physiology or medicine in 1929.

Foster became a leader as well in the national and international organization of science. At the Royal Society, of which he was elected fellow in 1872, he served as biological secretary from 1881 to 1903. In this influential and burdensome office he supported a wide range of scientific expeditions and was a vigorously successful advocate of a closer partnership between the Society and the government. He was vice-president of the Society in 1903–1904. He was also active in the affairs of the Royal Horticultural Society and of the British Association for the Advancement of Science, of which he was president in 1899. In the founding of the International Physiological Congresses, the first of which was held in 1889, he played a major role, as he did in the establishment of the International Association of Academies and in the preliminary arrangements for the International Catalogue of Scientific Literature.

Foster served on national commissions dealing with vaccination, tropical disease, disposal of sewage, and the reorganization of the University of London. In 1901 he was designated chairman of the Royal Commission on Tuberculosis. From 1900 to 1906 he was Member of Parliament for the University of London. Because he opposed the Liberal bill for Irish home rule, he stood originally as a Conservative; later he joined the Liberal opposition because of his stand on the education bill. On seeking reelection in 1906, he lost by the narrow margin of twenty-four votes.

Despite all these administrative duties, Foster coedited The Scientific Memoirs of T. H. Huxley (1898- 1902) and several times revised Huxley’s Lessons in Elementary Physiology. In 1899 he published a biography of Claude Bernard and in 1901 Lectures on the History of Physiology During the Sixteenth, Seventeenth and Eighteenth Centuries. Even in gardening, which was his chief source of relaxation, Foster exhibited leadership. He hybridized several new varieties of iris and was for a long time the internationally acknowledged expert on the genus. In fact, he published more original papers in horticulture than in physiology.

A member or fellow of a vast number of scientific societies, both British and foreign, Foster received honorary doctorates from the universities of Glasgow, St. Andrews, McGill, and Dublin. At Cambridge he was made honorary M.A. in 1871, and the full degree was conferred upon him in 1884. He was created K.C.B. in 1899.

Foster was twice married to women from Huntingdon: in 1863 to Georgina Edmonds, who died in 1869, and in 1872 to Margaret Rust, who survived him. By his first wife he had a daughter, Mercy, and a son, Michael George.

BIBLIOGRAPHY

I. Original Works. Foster’s papers on the heartbeat are the following: “On the Beat of the Snail’s Heart,” in Report of the Twenty-ninth Meeting of the British Association for the Advancement of Science (London, 1860), transactions of the sections, p. 160; “Note on the Action of the Interrupted Current on the Ventricle of the Frog’s Heart,” in Journal of Anatomy and Physiology, 3 (1869), 400–401; “Ueber einen besonderen Fall von Hemmungswirkung,” in Pflüger’s Archiv für die gesamte Physiologie des Menschen und der Tiere, 5 (1872), 191–195; “On the Behaviour of the Hearts of Mollusks Under the Influence of Electric Currents,” in Proceedings of the Royal Society, 23 (1875), 586–594, written with A. G. Dew-Smith; “The Effects of the Constant Current on the Heart,” in Journal of Anatomy and Physiology, 10 (1876), 735–771, written with A. G. Dew-Smith; “Some Effects of Upas Antiar on the Frog’s Heart,” ibid., 586–594; and “Die Muskeln und Nerven des Herzens bei einigen Mollusken,” in Archiv für mikroskopische Anatomie und Entwicklungsmechanik, 14 (1877), 317–321, written with A. G. Dew-Smith.

The most complete bibliography may be found in the article by Henry Dale (see below). A slightly less complete bibliography can be obtained by combining the citations in the Royal Society’s Catalogue of Scientific Papers, II , 674; VII , 692; IX , 906; XV , 69: and in the British Museum General Catalogue of Printed Books, LXXVI , cols. 241–243. Several publications are omitted from all of these sources. A series of popular articles on science (some unsigned), including one on the snail’s heart, appeared in the Christian Spectator in 1863 and 1864. Foster expresses his views on education in the unsigned article “Science in the Schools,” in London Quarterly Review, 123 (1867), 244–258; in “Vivisection,” in Macmillan’s Magazine, 29 (1874), 367–376, he defends physiologists;a gainst the charges of the antivivisectionists; and in “Reminiscences of a Physiologist,” in Colorado Medical Journal, 6 (1900), 419–429, he gives a rambling account of his education and early career.

There is no central repository for Foster’s private papers and correspondence. Of the Foster-Huxley correspondence, more than 200 letters are preserved in the Huxley Papers at Imperial College, London, and nearly as many, from Huxley to Foster only, at the Royal College of Physicians. Many letters from both collections are quoted in Leonard Huxley, Life and Letters of Thomas Henry Huxley, 2 vols. (London, 1900). Sir Robert Mordant Foster, grandson of Sir Michael, possesses more than twenty letters sent home by Foster during his brief career as ship’s surgeon. The rest of Foster’s extant letters are scattered among various collections in England, including the J. D. Hooker and W. T. Thistleton-Dyer letters at the Royal Botanical Gardens in Kew, and the E. A. Sharpey-Schafer Papers at the Wellcome Institute for the History of Medicine.

At the library of the Cambridge Physiological Laboratory is a bound MS volume in Foster’s hand with the heading “Three Lectures on the ‘Involuntary Movements of Animals’ Delivered Before the Royal Institution of Great Britain, February 1869.” This MS reveals the central thrust—as well as the difficulties and ambiguities—of Foster’s later ideas on the rhythmic heartbeat.

II. Secondary Literature. This article is based on Gerald L. Geison, “Sir Michael Foster and the Rise of the Cambridge School of Physiology, 1870–1900,” unpublished Ph.D. thesis (Yale, 1970). Of the many available sketches of Foster’s work and career, the most valuable are J. N. Langley, in Journal of Physiology (London), 35 (1907), 233–246; and in Dictionary of National Biography, suppl. I (1901–1911), 44–46; W. H. Gaskell, in Proceedings of the Royal Society, 80B (1908), lxxi–lxxxi; and Henry Dale, in Notes and Records. Royal Society of London, 19 (1964), 10–32.

For general background, see E. A. Sharpey-Schafer, History of the Physiological Society During Its First Fifty Years, 1876–1926 (London, 1927); G. L. Geison, “The Stagnancy of English Physiology, 1850–1870,” in Bulletin of the History of Medicine (in press); and Richard D. French, “Some Problems and Sources in the Modern Foundations of British Physiology,” in History of Science (in press). French has also drawn attention to the evolutionary context of the cardiological research of Foster and Gaskell. See “Darwin and the Physiologists, or the Medusa and Modern Cardiology,” in Journal of the History of Biology, 3 (1970), 253–274.

Gerald L. Geison