VII The Nervous System
As with the circulatory system, so with the nervous system, the work of Galen was centuries ahead of its time. Before him, Aristotle, who twice refers to experiments on animals, had observed the brain during life: for he says, “In no animal has the blood any feeling when it is touched, any more than the excretions; nor has the brain, or the marrow, any feeling, when it is touched”: but there is reason for believing that he neither recognised the purpose of the brain, nor understood the distribution of the nerves. Galen, by the help of the experimental method, founded the physiology of the nervous system:—
“Galen’s method of procedure was totally different to that of an anatomist alone. He first reviewed the anatomical position, and by dissection showed the continuity of the nervous system, both central and peripheral, and also that some bundles of nerve fibres were distributed to the skin, others to the muscles. Later, by process of the physiological experiment of dividing such bundles of fibres, he showed that the former were sensory fibres and the latter motor fibres. He further traced the nerves to their origins in the spinal cord, and their terminations as aforesaid. From these observations and experiments he was able to deduce the all-important fact that different nerve-roots supplied different groups of muscles and different areas of the skin.... An excellent illustration of his method, and of the fact that we ought not to treat symptoms, but the causes of symptoms, is shown very clearly in one of the cases which Galen records as having come under his care. He tells us that he was consulted by a certain sophist called Pausanias, who had a severe degree of anæsthesia of the little and ring fingers. For this loss of sensation, etc., the medical men who attended him applied ointments of various kinds to the affected fingers; but Galen, considering that that was a wrong principle, inquired into the history, and found that while the patient was driving in his chariot he had accidentally fallen out and struck his spine at the junction of the cervical and dorsal regions. Galen recognised that he had to do with a traumatism affecting the eighth cervical and first dorsal nerve; therefore, he says, he ordered that the ointments should be taken off the hand and placed over the spinal column, so as to treat the really affected part, and not apply remedies to merely the referred seat of pain.”5
Galen, by this sort of work, laid the foundations of physiology; but the men who came after him let his facts be overwhelmed by fantastic doctrines: all through the ages, from Galen to the Renaissance, no great advance was made toward the interpretation of the nervous system. Long after the Renaissance, his authority still held good; his ghost was not laid even by Paracelsus and Vesalius, it haunted the medical profession so late as the middle of the seventeenth century; but the men who worshipped his name missed the whole meaning of his work. This long neglect of the experimental method left such a gap in the history of physiology, that Sir Charles Bell seems to take up the experimental study of the nervous system at the point where Galen had stopped short; we go from the time of Commodus to the time of George the Third, and there is Bell, as it were, putting the finishing touch to Galen’s facts. It is true that experiments had been made on the nervous system by many men; but a dead weight of theories kept down the whole subject. For a good instance, how imagination hindered science, there is the following list, made by Dr. Risien Russell, of theories about the cerebellum:—
“Galen was of opinion that the cerebellum must be the originator of a large amount of vital force. After him, and up to the time of Willis, the prevalent idea seems to have been that it was the seat of memory; while Bourillon considered it the seat of instinct and intelligence. Willis supposed that it presided over involuntary movements and organic functions; and this view, though refuted by Haller, continued in the ascendency for some time. Some believed strongly in its influence on the functions of organic life; and according to some, diseases of the cerebellum appeared to tell on the movements of the heart.... Haller believed it to be the seat of sensations, as well as the source of voluntary power; and there were many supporters of the theory that the cerebellum was the seat of the sensory centres. Renzi considered this organ the nervous centre by which we perceive the reality of the external world, and direct and fix our senses on the things round us. Gall, and later Broussais, and others, held that this organ presided over the instinct of reproduction, or the propensity to love; while Carus regarded it as the seat of the will also. Rolando looked on it as the source of origin of all movements. Jessen adduced arguments in favour of its being the central organ of feeling, or of the soul, and the principal seat of the sensations.”
It is plain, from this list, that physiology had become obscured by fanciful notions of no practical value. If a better understanding of the nervous system could have been got without experiments on animals, why had men to wait so long for it? The Italian anatomists had long ago given them all the anatomy that was needed to make a beginning; the hospitals, and practice, had given them many hundred years of clinical facts; nervous diseases and head injuries were common enough in the Middle Ages; and by the time of Ambroise Paré, if not before, post-mortem examinations were allowed. The one thing wanted was the experimental method; and, for want of it, the science of the nervous system stood still. Experiments had been made; but the steady, general, unbiassed use of this method had been lost sight of, and men were more occupied with logic and with philosophy.
Then, in 1811, came Sir Charles Bell’s work. If any one would see how great was the need of experiments on animals for the interpretation of the nervous system, let him contrast the physiology of the eighteenth century with that one experiment by Bell which enabled him to say, “I now saw the meaning of the double connection of the nerves with the spinal marrow.” It is true that this method is but a part of the science of medicine; that experiment and experience ought to go together like the convexity and the concavity of a curve. But it is true also that men owe their deliverance from ignorance about the nervous system more to experiments on animals than to any other method of observing facts.
1. Sir Charles Bell (1778-1842)
The great authority of Sir Charles Bell has been quoted a thousand times against all experiments on animals:—
“Experiments have never been the means of discovery; and a survey of what has been attempted of late years in physiology, will prove that the opening of living animals has done more to perpetuate error than to confirm the just views taken from the study of anatomy and natural motions.”
He wrote, of course, in the days before bacteriology, before anæsthetics; he had in his mind neither inoculations, nor any observations made under chloroform or ether, but just “the opening of living animals.” He had also in his mind, and always in it, a great dislike against the school of Magendie. Let all that pass; our only concern here is to know whether these words are true of his own work.
They occur in a paper, On the Motions of the Eye, in Illustration of the Uses of the Muscles and Nerves of the Orbit; communicated by Sir Humphry Davy to the Royal Society, and read March 20, 1823.6 This essay was one of a series of papers on the nervous system, presented to the Royal Society during the years 1821-1829. In 1830, having already published four of these papers under the title, The Exposition of the Nervous System, Bell published all six of them, under the title, The Nervous System of the Human Body.
In his Preface to this book (1830) he quotes the earliest of all his printed writings on the nervous system, a pamphlet, printed in 1811, under the title, An Idea of a New Anatomy of the Brain, Submitted for the Observation of the Authors Friends. We have therefore two statements of his work, one in 1811, the other in 1823 and 1830. The first of them was written when his work was still new before his eyes.
Those who say that experiments did not help Bell in his great discovery—the difference between the anterior and the posterior nerve-roots—appeal to certain passages in the 1830 volume:—
“In a foreign review of my former papers, the results have been considered as a further proof in favour of experiments. They are, on the contrary, deductions from anatomy; and I have had recourse to experiments, not to form my own opinions, but to impress them upon others. It must be my apology that my utmost efforts of persuasion were lost, while I urged my statements on the grounds of anatomy alone. I have made few experiments; they have been simple and easily performed, and I hope are decisive....
“My conceptions of this matter arose by inference from the anatomical structure; so that the few experiments which have been made were directed only to the verification of the fundamental principles on which the system is established.”
If it were not for the 1811 pamphlet, the opponents of all experiments on animals might claim Sir Charles Bell on their side. But while his work was still a new thing, he spoke in another way of it:—
“I found that injury done to the anterior portion of the spinal marrow convulsed the animal more certainly than injury to the posterior portion; but I found it difficult to make the experiment without injuring both portions.
“Next, considering that the spinal nerves have a double root, and being of opinion that the properties of the nerves are derived from their connections with the parts of the brain, I thought that I had an opportunity of putting my opinion to the test of experiment, and of proving at the same time that nerves of different endowments were in the same cord (nerve-trunk) and held together by the same sheath.
“On laying bare the roots of the spinal nerves, I found that I could cut across the posterior fasciculus of nerves, which took its origin from the posterior portion of the spinal marrow, without convulsing the muscles of the back; but that on touching the anterior fasciculus with the point of the knife, the muscles of the back were immediately convulsed.
“Such were my reasons for concluding that the cerebrum and cerebellum were parts distinct in function, and that every nerve possessing a double function obtained that by having a double root. I now saw the meaning of the double connection of the nerves with the spinal marrow; and also the cause of that seeming intricacy in the connections of nerves throughout their course, which were not double at their origins.”
It is impossible to reconcile the 1830 sentences with this vivid personal account of himself; I had an opportunity of putting my opinion to the test of experiment... an opportunity of proving... Such were my reasons for concluding... I now saw.... It is just what all men of science say of their experiments: the very phrase of Archimedes, and Asellius, and de Graaf. If Sir Charles Bell had been working at the facts of chemistry or of botany, who would have doubted the meaning of these words?
This same inconsistency of sentences occurs elsewhere in his Nervous System of the Human Body. In one place he says that he has made few experiments: They have been simple, and easily performed, and I hope are decisive. In another he says: “After making several experiments on the cerebrum and cerebellum, I laid the question of their functions entirely aside, and confined myself to the investigation of the spinal marrow and the nerves; a subject which I found more within my power, and which forms the substance of the present volume.”
Next, take his account of the cranial nerves:—
“It was necessary to know, in the first place, whether the phenomena exhibited on injuring the separate roots corresponded with what was suggested by their anatomy....
“Here a difficulty arose. An opinion prevailed that ganglions were intended to cut off sensation; and every one of these nerves, which I supposed were the instruments of sensation, have ganglions on their roots. Some very decided experiment was necessary to overturn this dogma. (Account of the experiment.) By pursuing the inquiry, it was found that a ganglionic nerve is the sole organ of sensation in the head and face: ganglions were therefore no hindrance to sensation; and thus my opinion was confirmed.... It now became obvious why the third, sixth, and ninth nerves of the encephalon were single nerves in their roots....
“Observing that there was a portion of the fifth nerve which did not enter the ganglion of that nerve, and being assured of the fact by the concurring testimony of anatomists, I conceived that the fifth nerve was in fact the uppermost nerve of the spine.... This opinion was confirmed by experiment.... (Account of an experiment on the dead body.) On dividing the root of the nerve in a living animal, the jaw fell relaxed. Thus its functions are no longer matter of doubt: it is at once a muscular nerve and a nerve of sensibility. And thus the opinion is confirmed, that the fifth nerve is to the head what the spinal nerves are to the other parts of the body, in respect to sensation and volition.”
The value of the experimental method could hardly be stated in more emphatic words. He supposed something, conceived it, had an opinion about it. Anatomy had suggested something to him. He put his opinion to the test of phenomena, that is to say, to the test of visible facts; and then his opinion was confirmed. As with the spinal nerve-roots, so with the fifth cranial nerve—his work was successful, because he followed the way of experiment.
He was by nature of a most complex and sensitive temperament, full of contrary forces—one man in 1811, another in 1830. In 1811 he wrote, I now saw the meaning of the double connection of the nerves; in 1830 he had come to hate the stupid sterile materialism of the French school: he beheld anatomy falling behind physiology, and his Windmill Street school perishing to make way for the Hospital schools and for the University of London. He was before everything else a great anatomist: he stood up for the honour of anatomy against the new physiology, and for the honour of the Monroes and the Hunters against Magendie: he hated the notion that any man should proceed to experiments on function till the very last secrets had been got out of structure. He died a few years afterward. The 1830 writings are his last stand for the defence of his country, his school, and his beloved anatomy, against the methods of Magendie; who said of himself, “I am a mere street scavenger, chiffonier, of science. With my hook in my hand and my basket on my back, I go about the streets of science, collecting what I find.”
This open conflict between Bell’s first and last thoughts is a part of his character: he was brilliant, impulsive, changeable, inconsistent; and, what is more important, his honour kept him from trying to evade this trumpery charge of inconsistency; and he reprinted the 1811 Preface in the book that he published in 1830. Doubtless he would have picked his words more carefully if he had foreseen that one of the 1830 sentences would be wrested out of its place in his life’s work, and used as false evidence against the very method that he followed.
His observations on the cranial nerves brought about an immediate change in the practice of surgery:—
“Up to the time that Sir Charles Bell made his experiments on the nerves of the face, it was the common custom of surgeons to divide the facial nerve for the relief of neuralgia, tic douleureux; whereas it exercises, and was proved by Sir Charles Bell to exercise, no influence over sensation, and its division consequently for the relief of pain was a useless operation.” (Sir J. Erichsen.)
The relation of Magendie’s work on the nerve-roots to Bell’s work need not be considered here. The exact dates of Bell’s observations are given by one of his pupils in the Preface to the 1830 volume. Magendie finally proved the sensory nature of the posterior nerve-roots: “The exact and full proof which he brought forward of the truth which Charles Bell had divined rather than demonstrated, that the anterior and posterior roots of spinal nerves have essentially different functions—a truth which is the very foundation of the physiology of the nervous system—is enough by itself to mark him as a great physiologist.” (Sir M. Foster, loc. cit.)
2. Marshall Hall (1790-1857)
Reflex action had been studied long before the time of Marshall Hall. The Hon. Robert Boyle (1663) had observed the movements and actions of decapitated vipers, flies, silkworms, and butterflies. Similar observations were made on frogs, eels, and other lower animals, by Redi, Woodward, Stuart, Le Gallois, and Sir Gilbert Blane. According to Richet, it was Willis who first gave the name reflex to these movements.
It cannot be said that these first studies of reflex action did much for physiology. But the following translation from Prochaska (1800) shows how they cleared the way for Marshall Hall’s work, by the proof that they gave of the liberation of nervous energy in the spinal cord:—
“These movements of animals after decapitation must needs be by consent and commerce betwixt the spinal nerves. For a decapitated frog, if it be pricked, not only draws away the part that is pricked, but also creeps and jumps; which cannot happen but by consent betwixt the sensory nerves and the motor nerves. The seat of which consent must needs be in the spinal cord, the only remaining portion of the sensorium. And this reflexion of sensory impressions into motor impressions is not accomplished in obedience to physical laws alone—wherein the angle of reflexion is equal to the angle of incidence, and reaction to action—but it follows special laws as it were written by Nature on the spinal cord, which we can know only by their effects, but cannot fathom with the understanding. But the general law, whereby the sensorium reflects sensory impressions into motor impressions, is the preservation of ourselves.”
It was not possible, in 1800, to go further, or to put the facts of reflex action more clearly: but this fine sentence gives no hint of the truth that guided Marshall Hall—that the “consent and commerce” of reflex action are to be found at definite points or levels in the spinal cord; that the cord no more “works as a whole” than the brain. The greatness of Marshall Hall’s work lies in his recognition of the divisional action of the cord: he proved the existence of definite centres in it, he discovered the facts of spinal localisation, and thus foreshadowed the discovery of cerebral localisation. In his earlier writings (1823-33) he showed how the movements of the trunk and of the limbs are only one sort of reflex action; how the larynx, the pharynx, and the sphincter muscles, all act by the “consent and commerce” of the spinal cord. Later, in 1837, he demonstrated the course of nerve-impulses along the cord from one level to another, the results of direct stimulation of the cord, and other facts of spinal localisation. He noted the different effects of opium and of strychnine on reflex action; and he extended the doctrines of reflex action beyond physiology to the convulsive movements of the body in certain diseases.
3. Flourens (1794-1867)
Beside his work on the nervous system, Flourens studied the periosteal growth of bone, and the action of chloroform;7 but he is best known by his experiments on the respiratory centre and the cerebellum. The men who interpreted the nervous system followed the anatomical course of that system: first the nerve-roots, then the cord, then the medulla oblongata and the cerebellum, and last the cerebral hemispheres; a steady upward advance, from the observation of decapitated insects to the localisation of centres in the human brain. Flourens, by his work on the medulla oblongata, localised the respiratory centre, the nerve-cells for the reflex movements of respiration:—
“M. Flourens a circonscrit ce centre avec une scrupuleuse précision, et lui a donné le nom de nœud vital” (Cl. Bernard.)
Afterward came the discovery of cardiac and other centres in the same portion of the nervous system. Flourens also showed that the cerebellum is concerned with the equilibration of the body, and with the coordination of muscular movements; that an animal, a few days old, deprived of sensation and consciousness by removal of the cerebral hemispheres, was yet able to stand and move forward, but, when the cerebellum was removed, its muscles lost all co-ordinate action. (Recherches Expérimentales, Paris, 1842.) And from his work, and the work of those who followed him, on the semicircular canals of the internal ear, came the evidence that these minute structures are the terminal organs of equilibration: that as the special senses have their terminal apparatus and their central apparatus, so the semicircular canals and the cerebellum are the terminal apparatus and the central apparatus of the sense of equilibrium.
4. Claude Bernard (1813-1878)
The discovery of the vaso-motor nerves, and of the control of the nervous system over the calibre of the arteries, was made by Claude Bernard at the outset of his work on the influence of the nervous system on the temperature.8 The evidence of Professor Sharpey before the Royal Commission of 1875 shows how things had been misjudged, before Bernard’s time, in the light of “views taken from the Study of Anatomy and Natural Motions”:—
“I remember that Sir Charles Bell gave the increased size of the vessels in blushing, and their fulness of blood, as an example of the increased action of the arteries in driving on the blood. It turns out to be just the reverse, inasmuch as it is owing to a paralysis of the nerves governing the muscular coats of the arteries.”
Claude Bernard’s first account of his work was communicated to the Société de Biologie in December 1851. The following description is taken from his Leçons de Physiologie Opératoire:—
“I will remind you how I was led to the discovery of the vaso-motor nerves. Starting from the clinical observation, made long ago, that in paralysed limbs you find at one time an increase of cold, and at another an increase of heat, I thought this contradiction might be explained by supposing that, side by side with the general action of the nervous system, the sympathetic nerve might have the function of presiding over the production of heat; that is to say, that in the case where the paralysed limb was chilled, I supposed the sympathetic nerve to be paralysed, as well as the motor nerves; while in the paralysed limbs that were not chilled, the sympathetic nerve had retained its function, the systemic nerves alone having been attacked.
“This was a theory, that is to say, an idea leading me to make experiments; and for these experiments I must find a sympathetic nerve-trunk of sufficient size, going to some organ that was easy to observe, and must divide this trunk to see what would happen to the heat-supply of the organ. You know that the rabbit’s ear, and the cervical sympathetic nerve of this animal, offered us the required conditions. So I divided the nerve; and immediately my experiment gave the lie direct to my theory—Je coupai donc ce filet et aussitôt l’expérience donna à mon hypothèse le plus éclatant démenti. I had thought that the section of the nerve would suppress the function of nutrition, of calorification, over which the sympathetic system had been supposed to preside, and would cause the hollow of the ear to become chilled; and here was just the opposite, a very warm ear, with great dilatation of its vessels.
“I need not remind you how I made haste to abandon my first theory, and gave myself to the study of this new state of things. And you know that here was the starting-point of all my researches into the vaso-motor and thermic system; and the study of this subject is become one of the richest fields of experimental physiology.”
Waller, in 1853, studied the vaso-motor centre in the spinal cord; and Schiff, in 1856, found evidence of the existence of two kinds of vaso-motor nerves—those that constrict the vessels, and those that dilate them. This view was finally established in 1858 by Claude Bernard’s experiments on the chorda tympani and the submaxillary gland.
The Leçons de Physiologie Opératoire were published in 1879. Twenty years later, Sir Michael Foster says of Bernard’s work:—
“It is almost impossible to exaggerate the importance of these labours of Bernard on the vaso-motor nerves, since it is almost impossible to exaggerate the influence which our knowledge of the vaso-motor system, springing as it does from Bernard’s researches as from its fount and origin, has exerted, is exerting, and in widening measure will continue to exert, on all our physiological and pathological conceptions, on medical practice, and on the conduct of human life. There is hardly a physiological discussion of any width in which we do not sooner or later come on vaso-motor questions. Whatever part of physiology we touch, be it the work done by a muscle, be it the various kinds of secretive labour, be it the insurance of the brain’s well-being in the midst of the hydrostatic vicissitudes to which the changes of daily life subject it, be it that maintenance of bodily temperature which is a condition of the body’s activity; in all these, as in many other things, we find vaso-motor factors intervening. And if, passing the insecure and wavering line which parts health from illness, we find ourselves dealing with inflammation, or with fever, or with any of the disordered physiological processes which constitute disease, we shall find, whatever be the tissue specially affected by the morbid conditions, that vaso-motor influences have to be taken into account. The idea of vaso-motor action is woven as a dominant thread into all the physiological and pathological doctrines of to-day; attempt to draw out that thread, and all that would be left would appear as a tangled heap.”
5. Cerebral Localisation
Finally, moving upward along the anatomy of the nervous system, physiology came to study the motor-centres and special sense-centres of the cerebral hemispheres. The year 1861 may fairly be said to mark the beginning of the discovery of these centres, when Broca, at a meeting of the Anthropological Society of Paris, heard Aubertin’s paper on the connection between the frontal convolutions and the faculty of speech. But, of course, some sort of belief in cerebral localisation had been in the air long before Broca’s time. Willis (1621-1675), who was contemporary with Sir Isaac Newton, had written of the brain as though its convolutions, or “cranklings” as he called them, showed that its work was departmental:—
“As the animal spirits for the various acts of imagination and memory ought to be moved within certain and distinct limits, or bounded places, and these motions to be often iterated or repeated through the same tracts or paths, for that reason these manifold convolutions and infoldings of the brain are required for these divers manners of ordinations of the animal spirits—to wit, that in these cells or storehouses, severally placed, might be kept the species of sensitive things, and as occasion serves, may be taken from thence.”9
And Gall, a century after Willis, had collected and published, in support of his system of phrenology, many cases and post-mortem examinations showing the differentiation of the work of the brain. Gall is a warning for all time against the dangers of deduction; he had but one idea, and he drove it to death; but the clinical and pathological facts which he amassed, in the hope of establishing a set of doctrines out of all relation to facts, are as true now as ever; and, if he had been content to go the way of induction, and to set himself to the accumulation of facts, he might have become a great physiologist. In his knowledge of the anatomy of the brain, and in the dissection of the brain, he was far ahead of the men of his time; but he followed his own imaginings, and left nothing that could last, except those cases and pathological instances that are buried in the ruins of his system. But there they are, and are still of value. For example, Gall’s case of loss of speech, after an injury involving the speech-centres, ought to have commanded the attention of all physiologists: but it came to nothing, because he used it to support his doctrine of organs and bumps, and it shared the fate of that doctrine. Phrenology is gone past recall; it died of that congenital disease, the deductive fallacy; but there was a time when it might have been turned to the service of science.
The excitement that Gall aroused by the spread of his ideas shows that some belief in cerebral centres was waiting for development. All men are by nature phrenologists; the commonplace excuses that are offered for lapses of memory, venial offences, and inherited weaknesses, all appeal to the comfortable notion that the offender is not wholly perverted, and that some very small and strictly localised group of cells is at fault. And it is probable that the physiology of the central nervous system, with its present strong tendency toward psychology, will some day be back, at a far higher level, above the point where phrenology went wrong. As Mme. de Staël said, L’esprit humain fait progrès toujours, mais c’est progrès en spirale. But the question, whether the general desire for a rational system of psychology will ever commend itself to physiology, belongs to the future. All that is of present concern is the steady, continuous, and successful advance, by the way of induction, and by the help of experiments on animals, toward a clear and accurate statement of the departmental work of the brain.
It is one of many instances how science and practice work together, that the modern study of these centres began not in experiment but in experience. The first centres that were thus studied were the speech-centres; and the observation of them arose out of the cases recorded by Bouillard in 1825, and Dax in 1836. Clinical observation, and post-mortem examination, found the speech-centres; physiological experiments had nothing to do with it; and phrenology had, as it were, found them, and then lost them. But at once, so soon as practice gave the word to science, physiology set to work. These clinical facts had been there all the time; loss of speech had gone with disease or injury of “Broca’s convolution” ever since man had been on the earth, and nobody had seen the significance of this sequence. Then, after 1861, everything was changed; and in a few years physiology had mapped out a large part of the surface of the brain, and had charted the motor-centres.
The story of Broca’s convolution is told in Hamilton’s Text-Book of Pathology:—
“In 1825, Bouillard collected a series of cases to show that the faculty of speech resided in the frontal lobes. In the year 1836 M. Dax, in a paper read to the Medical Congress of Montpellier, stated as a result of his researches that, where speech was lost from cerebral causes, he believed the lesion was invariably found in the left cerebral hemisphere, and that the accompanying paralysis of the right side of the body is consequent upon this. This paper for long lay buried in the annals of medical literature, but was unearthed years afterwards by his son, and presented to the French Academy. Bouillard’s views were also disinterred by Aubertin, and in the year 1861 were brought by him before the notice of the Anthropological Society of Paris. Broca, who was present at the meeting, had a patient under his care at the time who had been aphasic (without power of speech) for twenty-one years, and who was in an almost moribund state. The autopsy proved of great interest, as it was found that the lesion was confined to the left side of the brain, and to what we now call the third frontal convolution. Broca was struck with the coincidence; and when a similar case came under his care afterwards, unaware of what had been done by Dax, he postulated the conclusion that the integrity of the third frontal convolution, and perhaps also part of the second, is essential to speech. In a subsequent series of fifteen typical cases examined, it was found that the lesion had destroyed, among other parts, the posterior part of the third frontal in fourteen. In the fifteenth case the destruction had taken place in the island of Reil and the temporal lobe.”
After 1861, physiology took the lead, and kept it. But, through all the work, science and practice have been held together; the facts of experimental physiology have been and are tested, every inch of the way, by the facts of medicine, surgery, and pathology. The infinite minuteness and complexity of the investigation, and its innumerable side-issues, are past all telling. They who are doing the work, in science and in practice, have always had in their thoughts the fear of fallacies in the interpretation of these highest forms of life. Sir William Gowers, fourteen years ago, wrote as follows of the earlier workers:—
“Doubt was formerly entertained as to the existence of differentiation of function in different parts of the cortex, but recent researches have established the existence of a differentiation which has almost revolutionised cerebral physiology, and has vastly extended the range of cerebral diagnosis. The first step of the new discovery was constituted by the clinical and pathological observations of Hughlings Jackson, which suggested the existence, on each side of the fissure of Rolando, of special centres for the movements of the leg, arm, and face. These observations led to the experiments of Ferner, which resulted in the demonstration of the existence in the cortex of the lower animals of well-defined regions, stimulation of which caused separate movements, or evidence of special sense excitation, while the destruction of the same parts caused indications of a loss of the corresponding function. Hence he came to the conclusion that these regions constitute actual motor and sensory centres. Ferrier had, however, been anticipated in many of these results by two German experimenters, Fritsch and Hitzig, whose results, differing a little in detail, correspond closely in their general significance. Many other investigations of the same character have since been made, of which those of Munk are especially important. The original observations of Hughlings Jackson left little doubt that the general facts, learned from experiments on animals, are true of man; and this conclusion has been to a large extent confirmed by pathological and clinical observations directed to the verification on man of the pathological results. To this verification the labours of Charcot and his coadjutors have largely contributed. But the verification has already made it probable that some differences exist between the brain of man and that of higher animals (even of monkeys), and that the conclusions from the latter cannot be simply transferred to the former.”
Many and great difficulties, beyond this danger of the fallacy of “simple transference,” beset every step of the work: it required the right use of the most delicate and susceptible instruments and tests, and the right understanding of anatomy, microscopic anatomy, comparative anatomy, organic chemistry, electricity, and physics: every moment of advance must be guarded, every word must be weighed. Among the earlier difficulties, was the failure of almost all the physiologists, before Hitzig, to produce muscular action by excitation of the cerebral cortex. Longet, Magendie, Flourens, Matteuci, Van Deen, Weber, Budge, and Schiff, had all failed. Hitzig (Untersuchungen über das Gehirn, Berlin, 1874) had observed, in man, that it was easy to produce movements of the eyes by the passage of the constant current through the occipital region.10 Taking this fact for a starting-point, he used a very low current, and thereby succeeded in producing certain definite muscular movements by stimulation of the cortex in animals. Of Hitzig’s work, Sir Victor Horsley says:—
“It was not till 1870 that the next absolute proof (after Bell’s work in 1813) was obtained of the localisation of function, so far as the highest centres of the nervous system were concerned. In that year Fritsch and Hitzig discovered that electrical excitation, with minimal stimuli, of various points of the cortex, caused those storehouses, of which Willis spoke, to discharge, and to reveal their function by the precise limitation of the groups of muscles which they were able to throw into action. These researches were abundantly confirmed and greatly extended by Professor Ferrier, and thus has been constructed in the history of this subject the most recent great platform or stage of permanent advance.”11
The thirty years since Hitzig’s work cannot be put here, for they would take a volume to themselves. There have been differences of interpretation of this or that fact, diversities of results, and problems too hard to solve, and other difficulties, such as befall all the natural sciences; but these imperfections amount to very little, when the whole result comes to be reckoned. The marvel is that the work is so nearly perfect, seeing its immeasurable complexity.
Let any man, who has but touched the study of physiology, consider what is involved in even the most superficial observation of the simplest facts of the nervous system: for instance, the ordinary nerve-muscle preparation that is taught to every medical student, or the microscopic structure of the spinal cord, or the Wallerian method. Or let him consider how the physiology of the nervous system has been founded on the lower forms of life: the work of Romanes and others on the Medusa and the Echinodermata, and Huxley’s work in biology, and the endless chain of forces that are alike in man and in jelly-fishes. Then let him try to estimate the output of hard thinking, for the advance from lower to higher structures, and up to man; the vigilant criticism of all theories and foregone conclusions, the incessant self-judgment and wearisome doubts and disputes all the way, elusiveness of facts, and vagueness of words. And the results thus wrung out of science had still to be stated in terms of practice, and tested by the facts of medicine, surgery, and pathology, and used in every hospital in the civilised world, not only for the saving of life, but also for the diagnosis and medical or surgical treatment of innumerable varieties of disease or injury of the brain, the cord, or the nerves. Sir Michael Foster, in a short summary of the problems of physiology, puts clearly these consummate difficulties of the physiology of the nervous system:—
“In the first place there are what may be called general problems, such as, How the food, after its preparation and elaboration into blood, is built up into the living substance of the several tissues? How the living substance breaks down into the dead waste? How the building up and breaking down differ in the different tissues in such a way that energy is set free in different modes, the muscular tissue contracting, the nervous tissue thrilling with a nervous impulse, the secreting tissue doing chemical work, and the like? To these general questions the answers which we can at present give can hardly be called answers at all.
“In the second place there are what may be called special problems, such as, What are the various steps by which the blood is kept replenished with food and oxygen, and kept free from an accumulation of water; and how is the activity of the digestive, respiratory, and excretory organs, which effect this, regulated and adapted to the stress of circumstances? What are the details of the vascular mechanism by which each and every tissue is for ever bathed with fresh blood, and how is that working delicately adapted to all the varied changes of the body? And, compared with which all other problems are insignificant and preparatory only, how do nervous impulses so flit to and fro within the nervous system as to issue in the movements which make up what we sometimes call the life of man?”
The physiology of the nervous system is wrought to finer issues now than in the time of Bell and Magendie; and this generation of students may live to see the present facts and methods of cerebral localisation as the mere rudiments or elements of science. Happily for mankind, science has already so far elucidated them that they have done good service for the diagnosis and treatment of disease, and for the saving of lives.
Some examples have been given, in the foregoing chapters, of the value of physiological experiments on animals. It would be easy to lengthen the list, for there is no general subject in all physiology that does not owe something to this method: as Mr. Darwin said, in his evidence before the Royal Commission of 1875, “I am fully convinced that physiology can progress only by the aid of experiments on living animals. I cannot think of any one step which has been made in physiology without that aid.” Many examples have been left out altogether—the work of Boyle, Hunter, Lavoisier, Haldane, Despretz, and Regnault, on animal heat and on respiration; of Petit, Dupuy, Breschet, and Reid, on the sympathetic system; of Galvani, Volta, Haller, du Bois-Reymond, and Pflüger, on muscular contractility: nothing has been said of the work lately done on the suprarenal glands and “adrenalin,” and on the blood-pressure in its relation to secretion. For the most part, only those examples have been taken that occur far back in the history of physiology: more has been said about the past than about the present. First, because it was necessary to put an end to the false statements that are made, by those who are opposed to all experiments on animals, about the work done in the past. Next, because the abstruse details of physiology, in the present, are not intelligible for general reading. Next, because it is impossible now to isolate physiology, or to say what belongs to physiology alone, to have back the simpler problems of the past, to discover the circulation of the blood twice. But the experimental method, alike in the past and in the present, has been the chief way of advance. And if a forecast may be made without offence, it is certain that the work of physiology, as in the past and the present, so in the near future, will exercise a profound influence for good on medical and surgical treatment. Among the subjects that especially occupy physiologists now are, the more exact localisation and interpretation of the special sense-centres, and the better knowledge of the internal secretions and chemical influences of the glands and tissues of the body. It would be hard to find two fields of work more sure to favour the growth of the arbor vitæ side by side with the arbor scientiæ.
But the last word here must be said by a physiologist of the very highest authority, Professor Starling. He has kindly given me, for this edition, the following note:—
“Among the researches of the last thirty years, those bearing on the Circulation of the Blood must take an important place, both for their physiological interest and for the weighty influence they have exerted on our knowledge and treatment of disorders of the vascular system, such as heart disease. We have learned to measure accurately the work done by the great heart-pump; and by studying the manner in which this work is affected by different conditions, we are enabled to increase or diminish it, according to the needs of the organ. Experiments in what is often regarded as the most transcendental department of physiology—i.e. that which treats of muscle and nerve—have thrown light on the wonderful process of ‘compensation,’ by which a diseased heart is able to keep up a normal circulation.
“Vaso-motor System.—Largely by the labours of British physiologists, the exquisite control exercised by the nervous system over every blood-vessel in the body has been brought to light, the paths tracked out, and the mechanisms elucidated, by means of which the circulation through each part of the body is subordinated to the needs of the whole. Since the chief vaso-motor nerves take their course through the sympathetic system, the researches on their distribution have led to the mapping out of the whole of this system, and to an accurate knowledge of its functions. We are now acquainted with the course, to all parts of the body, of the nerves which not only determine the changes in the calibre of the blood-vessels, but affect also the secretion of sweat and the erection of the hairs. Incidentally, the mapping out of these nerves, in the hands of Mackenzie, Head, and others, has led to more power of localising the seat of visceral disease.
“Digestion.—Our knowledge of the processes of digestion has of late years received a great accession by the work of Professor Pawlow, of St. Petersburg. His success is largely due to his recognition of the importance of keeping his experimental animals under the most normal conditions possible, and of studying the different parts of the alimentary tract in animals which were not anæsthetised, but which were free from any pain or even discomfort, either of which conditions materially interferes with the activity of the digestive glands. He therefore established in dogs fistulæ in chosen portions of the alimentary canal, analogous to the fistula which accident rendered so valuable in the case of Alexis St. Martin. Not only has the knowledge thus gained enabled the physician to understand the sequel of events in disordered digestion, but the success of the operative measures undertaken by physiologists for the elucidation of their science has emboldened surgeons to attack disease in the most various parts of the alimentary canal.
“Renewed study of the secretion of pancreatic juice evoked by the passage of the acid digestive products from the stomach into the small intestine, which had been described by Pawlow, has resulted in the discovery of a new class of chemical agents, which act as special messengers from one part of the body to another, and exercise an important function in determining the action of all parts to one common end.
“Respiration.—The investigation of the chemical properties of the colouring matter of blood, and of its compound with carbon monoxide, has resulted, in the hands of Dr. Haldane, in the laying down of measures for the prevention of accidents from choke-damp or after-damp in mines. The same investigation has resulted in the discovery of a method of determining the total amount of blood circulating in the body of a living man. The application of this method has already added largely to our knowledge of the pathology of different forms of anæmia, as well as of the conditions obtaining in heart disease. Experiments by Hill and others on the physiological effects of compressed air have shown the precautions which should be observed in all diving operations. A proper appreciation of these results by diving-engineers would not only entirely obviate the cases of ‘caisson disease,’ but would enable diving to be carried on safely to a greater depth than has hitherto been attempted.
“It is impossible, however, to enumerate all the physiological gains of the last twenty or thirty years, or to point out their manifold applications in the cure and prevention of disease. The full control of the processes of disease, which is the goal of the physician and the surgeon, can only be attained through an accurate knowledge of the conditions governing the functions of the healthy body. The foundation of medicine and surgery is physiology: and it is only on living animals that the processes of life can be investigated.”