Bayliss, William Maddock

(b. Wednesbury, Staffordshire, England, 2 May 1860; d. London, England, 27 August 1924)

general physiology.

W. M. Bayliss was the only son of Moses Bayliss and Jan Maddock to survive infancy. His father was a manufacturer of galvanized goods and founder of the Wolverhampton firm of Bayliss, Jones and Bayliss.

William received his early education at a private school in Wolverhampton, Staffordshire, and was then apprenticed to a local general practitioner.

When, in 1880, the elder Bayliss moved to London, William continued his studies at University College, London, where in 1881 he gained a medical entrance exhibition and a science exhibition at the preliminary scientific examination. In 1882 he took the B.Sc. degree with a scholarship in zoology and physics, and began to study physiology and anatomy, but at the second M.B. examination he failed in anatomy. He then relinquished medical studies to concentrate on physiology.

Two men who influenced him at this stage were John Burdon-Sanderson, Jodrell professor of physiology at University College, and Ray Lankester, professor of zoology. Physiology was at that time emerging as a subject separate from anatomy, and the Jodrell chair, founded in 1874, was the first separate chair of physiology in Great Britain. When the Waynflete chair of physiology was established at Oxford in 1882, Burdon-Sanderson was elected to it, and in 1885 Bayliss followed him to Oxford as an undergraduate of Wadham College. He took a first-class degree in physiology in 1888 and then returned to University College, where, in the meantime, E. A. Schäfer (afterward Sir Edward Sharpey-Schafer) had succeeded to the Jodrell chair. Bayliss worked for the remainder of his life at University College, first as assistant, then in 1903 as assistant professor, and from 1912 onward as professor of general physiology.

He attended an Anglo-Catholic church, but was largely tolerant in religious matters. Politically, he was somewhat to the left of liberal, with a sympathy for reasonable minority movements, but had little interest in politics as such.

In 1893 he married Gertrude Starling, the sister of E. H. Starling; and, after his father’s death in 1895, he returned to live in the family home in Hampstead, London. He had three sons and one daughter; the youngest son, Leonard (1900–1964), was also a physiologist. Bayliss was well-off financially, his married life was happy, and until shortly before his death from a blood dyscrasia his health was excellent. In their large house, set in a four-acre garden with two tennis courts and a private laboratory, the Baylisses entertained often and liberally, mainly colleagues and visiting physiologists.

Bayliss was elected to membership in the Physiological Society in 1890 and served as secretary from 1900 to 1922, and treasurer from 1922 to 1924. He was editor of Physiological Abstracts from 1923 to 1924, and joint editor of the Biochemical Journal from 1913 to 1924.

In 1903 he was elected a fellow of the Royal Society, serving on its council from 1913 to 1915. He was Croonian lecturer (jointly with Starling) in 1904 and was awarded a Royal Medal in 1911 and the Copley Medal in 1919. He also received honorary degrees from the universities of Oxford, St. Andrews, and Aberdeen and was elected to the Royal Danish Academy of Science, the Royal Academy of Belgium, and the Sociétéde Biologie of Paris. In 1917 he was awarded the Baly Medal of the Royal College of Physicians and in 1918 gave the Oliver-Sharpey lectures, and the Sylvanus Thompson lectures in 1919. He was created a knight bachelor in 1922.

During World War I he served on the Food (War) Committee of the Royal Society, and on the Wound Shock Committee of the Medical Research Committee; he visited the front in France in 1917. From 1917 to 1924 he served on the Medical Research Council’s committee to study the biological action of light.

In 1922 he visited the United States to deliver the Herter lectures at Baltimore, a Harvey lecture at New York, and to talk to the Research Club at Harvard.

The placid tenor of his life was interrupted in 1903 by the urgent necessity to take out an action for libel against Stephen Coleridge, secretary of the National Antivivisection Society. The trial, before the lord chief justice, occupied four days and Bayliss won the day, with £2,000 damages. He presented the money to University College for furtherance of research in physiology. The interest on the capital sum is still used for that purpose.

Bayliss devoted his life almost solely to his work in physiology. He was fond of music and played the violin. He was of a merry and gentle disposition, good with his hands, and would accept little help from others, although he gladly gave his own. Photography was more than a hobby to him; the illustrations for his books and papers were his own work.

Bayliss’ generation entered the scene at the beginning of a period of exponential growth of science, and his outlook can be seen as a projection into the twentieth century of that of Claude Bernard (1814–1878), Ludwig (1816–1895), and Helmholtz (1821–1894). To him all science was one, although fundamentally he was a biologist who was deeply interested in the emergence of physical chemistry and biochemistry and in their relations to the problems of general physiology. The foundations of Hoppe-Seyler’s Zeitschrift für physiologische Chemie (1877) and Ostwald’s Zeitschrift für physikalische Chemie (1887) were harbingers of what was to follow, and Bayliss availed himself of the tools offered by these new subjects.

The frequent coupling together of the names of Bayliss and Starling in important publications might lead to the inference that their partnership was an essential condition for either or both of them, but nothing could be further from the truth. They were both great physiologists and in their collaboration were largely complementary. Bayliss was the more fundamental and erudite, but of a retiring disposition; Starling was a pragmatic and forceful extrovert, with an essentially medical outlook. Bayliss preferred to work alone, without even technical help, but collaboration was imposed upon them for a time; until a new Institute of Physiology was built there was inadequate space and equipment in the department. After 1909 they never collaborated, although they frequently conferred at the daily tea meetings.

Bayliss’ researches can be divided into roughly the following six phases:

Electrophysiology. When Bayliss started at University College, electrophysiology was the central feature of the research work in the department. Bayliss worked with Dr. (afterward Sir John) Rose Bradford in studying, with the aid of a Thomson galvanometer, the changes in electric potential involved in the act of secretion, first in the salivary glands of a frog and then in the skin. In his second paper, “The Electromotive Phenomena of the Mammalian Heart,” he collaborated with Starling, partly at Oxford; they were able to use the relatively rapidly responding capillary electrometer, which Lippmann had introduced in 1873. The hearts of mammals, frog, tortoise, and man (the two authors) were studied, the electrical changes shown to be triphasic, and the time relations of the cardiac cycle observed.

Vascular System. This work, which was begun in 1892 with Starling, resulted in two papers, one on a new form of optically registered blood pressure apparatus and one “On Some Points in the Innervation of the Mammalian Heart.” He also worked with Bradford on the innervation of the vessels of the limbs. At about this same time, he began his own investigations, starting with the physiology of the depressor nerve, which led on to an examination of vasomotor reactions in general and of their central coordination. The nature of the antidromic reaction, which he showed to be an axon reflex, was another important outcome of this work. By 1894 Bayliss and Starling were working on circulatory problems, including venous and capillary pressures and intraventricular and aortic pressure curves.

Intestinal Movements. The partners now turned their attention to the study of the movements of the small and large intestines, and of their innervation. The most important result of this work was the elucidation of the peristaltic wave.

Pancreatic Secretion. Bayliss’ interest in secretion led him, with Starling, to study the anomalous secretion of pancreatic juice which occurred when acid was introduced into the duodenum. At that time the only known means by which glands were excited to secrete was by the mediation of secretory nerves, as exemplified by the pioneer work of Ludwig (1851) on the submaxillary gland. Pavlov had shown that secretion of pancreatic juice was evoked on stimulation of the vagus nerves, but it could also be produced by the introduction of acid into the upper intestine, even after section of both vagi; this acid-provoked secretion, he concluded, could only be brought about by a local reflex from the intestinal mucosa. Bayliss and Starling showed, however, that introduction of acid into the duodenum still induced secretion of pancreatic juice after division of both vagi and of all the nerves to the upper intestine, so that the only means of communication of the pancreas with the intestine was via the bloodstream. The excitant must, therefore, be not a nervous but a chemical stimulus, a substance derived from the intestine and carried in the bloodstream to the pancreas, which it excited to secrete. This they proved by showing that the intravenous injection of an acid extract of intestinal mucosa resulted in the secretion of pancreatic juice. They called the excitant substance Secrelin, and thus recognized the existence of a new class of chemical messengers, the hormones (from the Greek horman, “to set in motion”), a name first promulgated by Starling in his Croonian lecture of 1905, in which he foreshadowed other hormone actions.

Enzyme Action. Bayliss and Starling next studied the activation of trypsin in the pancreatic juice. As secreted, the juice has no proteolytic action, but it becomes activated by contact with enterokinase in the intestinal juice. Pavlov regarded this activation to be the result of an enzyme action by which the precursor, trypsinogen, was converted into the active form, trypsin, by the separation from it of part of its molecule. Opponents had thought of the action as a combination between inactive enzyme and activator. Bayliss and Starling gave support to Pavlov’s view by showing that the reaction was catalytic and not stoichiometric.

Bayliss now turned to study enzyme action, in the light of physical chemistry. He used trypsin as the enzyme, caseinogen or gelatin as substrate, and followed the course of the digestion by measurement of the electrical conductivity, correlated with other measurements such as viscosity, refractive index, optical rotation, osmotic pressure, etc. These studies led on to further work on the velocity of enzyme action, final equilibria, reversibility, and the effect of temperature, in the course of which he showed that enzyme action is largely influenced by adsorption effects in which there is combination between enzyme and substrate, a viewpoint developed by Michaelis and Menten in 1913.

Colloids. Bayliss’ study of the osmotic pressure of Congo red, showing that this colloid exerted the same osmotic pressure that it would if it were in true solution, led on to the study of the electrical equilibrium set up when Congo red and sodium chloride solutions are separated by a semipermeable membrane. This work gave support to the studies of Donnan on the theory of membrane equilibria, and its relation to Nernst’s equation for a concentration cell.

War Problems (1914–1918). Bayliss’ most valuable contribution to the war effort was his work on wound shock, which led to the practice of replacing lost blood with a gum saline solution. He had shown the importance of maintaining an adequate colloidal osmotic pressure in the circulating blood.

Books. Bayliss’ great book, on which he spent many years, was his Principles of General Physiology, which first appeared in 1914. It can be regarded as an extension into the twentieth century of Claude Bernard’s Phénomènes de la vie commune aux animaux et aux végétaux (1878–1879). The Principles stands as a landmark in the history of biological literature. When Bayliss visited the United States in 1922, he was pleased, but embarrassed, to find that at some universities there were Bayliss clubs, formed to discuss the contents of the book.

BIBLIOGRAPHY

I. Original Works. Articles. “On the Physiology of the Depressor Nerve,” in Journal of Physiology. 14 (1893), 303–325; “On the Origin From the Spinal Cord of the Vaso-Dilator Fibres of the Hind Limb and on the Nature of These Fibres,” ibid., 26 (1901), 173–209; “Further Researches on Antidromic Nerve Impulses,” ibid., 28 (1902), 276–299; “The Kinetics of Tryptic Action,” in Archives des sciences biologiques11 (1904), 261–296, supp., Pavlov Jubilee Volume; “On Some Aspects of Adsorption Phenonmena, with Especial Reference to the Action of Electrolytes, and to the Ash Constituents of Proteins,” in Biochemical Journal, 1 (1906), 175–232; “On Reciprocal Invervation in Vaso-Motor Reflecxes, and on the Action of Strychnine and of Chloroform Thereon.” in Proceedings of the Royal Society, 80B (1908), 339–375; “The Excitation of Vaso-Dilator Fibres in Depressor Reflexes,” “in Journal of Physiology, 37 (1908), 264–277; “The Osmotic Pressure of Congo Red, and of Some Other Dyes,” in Proceedings of the Royal Society, 81B (1909), 269–286; “On Adsorption as Preliminary to Chemical Reaction,” ibid., 84B (1911), 81–98; “The Osmotic Pressure of Electrolytically Dissociated Colloids,” ibid., pp. 229–254; “Researches on the Nature of Enzyme Action. II. The Synthetic Properties of Anti-Emulsin,” in Journal of Physiology, 43 (1912), 455–466; “Researches on the Nature of Enzyme Action. III. The Synthetic Action of Enzymes,” ibid., 46 (1913), 236–266; “The Action of Insoluble Enzymes,” ibid., 50 (1915), 85–94; Methods of Raising a Low Arterial Pressure,” in Proceedings of the Royal Society, 89B (1916), 380–393, and in British Medical Journal (1917); “The Action of Gum Acacia on the Circulation,” in Journal of Pharmacology and Experimental Therapeutics, 15 (1920). 29–74; “Reversible Gelation in Living Protoplasm,” in Proceedings of the Royal Society, 91B (1920), 196–201.

Collaborations. J. R. Bradford: “The Electrical Phenomena Accompanying the Process of Secretion in the Salivary Glands of the Dog and Cat,” in Internationale Monatsschrift für Anatomic und Physiologic, 4 (1885), 109–136; “Proc. Physiol. Soc.,” in Journal of Physiology, 6 (1889), xiii–xvi; and “The Innervation of the Vessels of the Limbs,” ibid., 16 (1894), 10–22.

W. B. Cannon: “Note on Muscle Injury in Relation to Shock,” in Special Report of the Medical Research Committee no. 26 (1919), 19–23.

Leonard Hill: “On the Intra-Cranial Pressure and the Cerebral Circulations.” in Journal of Physiology, 18 (1895), 334–362.

E. H. Starling: “On the Electromotive Phenomena of the Mammalian Heart,” in Internationale Monatsschrift für Anatomie und Physiologie, 9 (1892), 256–281; “On Some Points in the Innervation of the Mammalian Heart,” in Journal of Physiology, 13 (1892), 407–418; “On the Form of the Intraventricular and Aortic Pressure Curves Obtained by a New Method,” in Internationale Monatsschrift für Anatomie und Physiologie, 11 (1894), Heft 9; “The Movements and Innervation of the Small Intestine,” in Journal of Physiology, 24 (1899), 99–143; “The Mechanism of Pancreatic Secretion,” ibid., 28 (1902), 325–353: “The Proteolvtic Activities of the Pancreatic Juice,” ibid., 30 (1903), 61–83; and “The Chemical Regulation of the Secretory Process,” Croonian Lecture pub. in Proceedings of the Royal Society, 73B (1904), 310–322.

Books. The Nature of Enzyme Action, 5 eds. (London, 1908–1925); An Introduction to General Physiology (London, 1919); Principles of General Physiology 4 eds. (London, 1919–1924); The Physiology of Food and Economy in Diet (London, 1917); Intravenous Injections in Wound Shock, Oliver-Sharpey Lectures (London, 1918); The Vasomotor System (London, 1923); The Colloidal State in Its Medical and Physiological Aspects (Oxford, 1923); Interfacial Forces and Phenomena in Physiology, Herter Lectures (London, 1923).

II. Secondary Literature. L. E. Bayliss, “William Maddock Bayliss (1860–1924),” in Perspectives in Biology and Medicine4 (1961), 460–479; C. Lovatt Evans, First Bayliss-Starling Memorial Lecture, Reminiscences of Bayliss and Starling (Cambridge, 1964), p. 17; E. H. Starling, obit, of W. M. Bayliss in The Times (28 Aug. 1924).

Charles L. Evans