Bell's discovery was epochal; but its full significance was not appreciated for a decade, nor, indeed, was its validity at first admitted. In Paris, in particular, then the court of final appeal in all matters scientific, the alleged discovery was looked at askance, or quite ignored. But in 1823 the subject was taken up by the recognized leader of French physiology - Francois Magendie - in the course of his comprehensive experimental studies of the nervous system, and Bell's conclusions were subjected to the most rigid experimental tests and found altogether valid. Bell himself, meanwhile, had turned his attention to the cranial nerves, and had proved that these also are divisible into two sets - sensory and motor. Sometimes, indeed, the two sets of filaments are combined into one nerve cord, but if traced to their origin these are found to arise from different brain centres. Thus it was clear that a hitherto unrecognized duality of function pertains to the entire extra-cranial nervous system. Any impulse sent from the periphery to the brain must be conveyed along a perfectly definite channel; the response from the brain, sent out to the peripheral muscles, must traverse an equally definite and altogether different course. If either channel is interrupted - as by the section of its particular nerve tract - the corresponding message is denied transmission as effectually as an electric current is stopped by the section of the transmitting wire.

Experimenters everywhere soon confirmed the observations of Bell and Magendie, and, as always happens after a great discovery, a fresh impulse was given to investigations in allied fields. Nevertheless, a full decade elapsed before another discovery of comparable importance was made. Then Marshall Hall, the most famous of English physicians of his day, made his classical observations on the phenomena that henceforth were to be known as reflex action. In 1832, while experimenting one day with a decapitated newt, he observed that the headless creature's limbs would contract in direct response to certain stimuli. Such a response could no longer be secured if the spinal nerves supplying a part were severed. Hence it was clear that responsive centres exist in the spinal cord capable of receiving a sensory message and of transmitting a motor impulse in reply - a function hitherto supposed to be reserved for the brain. Further studies went to show that such phenomena of reflex action on the part of centres lying outside the range of consciousness, both in the spinal cord and in the brain itself, are extremely common; that, in short, they enter constantly into the activities of every living organism and have a most important share in the sum total of vital movements. Hence, Hall's discovery must always stand as one of the great mile-stones of the advance of neurological science.

Hall gave an admirably clear and interesting account of his experiments and conclusions in a paper before the Royal Society, "On the Reflex Functions of the Medulla Oblongata and the Medulla Spinalis," from which, as published in the Transactions of the society for 1833, we may quote at some length:

"In the entire animal, sensation and voluntary motion, functions of the cerebrum, combine with the functions of the medulla oblongata and medulla spinalis, and may therefore render it difficult or impossible to determine those which are peculiar to each; if, in an animal deprived of the brain, the spinal marrow or the nerves supplying the muscles be stimulated, those muscles, whether voluntary or respiratory, are equally thrown into contraction, and, it may be added, equally in the complete and in the mutilated animal; and, in the case of the nerves, equally in limbs connected with and detached from the spinal marrow.

"The operation of all these various causes may be designated centric, as taking place AT, or at least in a direction FROM, central parts of the nervous system. But there is another function the phenomena of which are of a totally different order and obey totally different laws, being excited by causes in a situation which is EXCENTRIC in the nervous system - that is, distant from the nervous centres. This mode of action has not, I think, been hitherto distinctly understood by physiologists.

"Many of the phenomena of this principle of action, as they occur in the limbs, have certainly been observed. But, in the first place, this function is by no means confined to the limbs; for, while it imparts to each muscle its appropriate tone, and to each system of muscles its appropriate equilibrium or balance, it performs the still more important office of presiding over the orifices and terminations of each of the internal canals in the animal economy, giving them their due form and action; and, in the second place, in the instances in which the phenomena of this function have been noticed, they have been confounded, as I have stated, with those of sensation and volition; or, if they have been distinguished from these, they have been too indefinitely denominated instinctive, or automatic. I have been compelled, therefore, to adopt some new designation for them, and I shall now give the reasons for my choice of that which is given in the title of this paper - 'Reflex Functions.'

"This property is characterized by being EXCITED in its action and REFLEX in its course: in every instance in which it is exerted an impression made upon the extremities of certain nerves is conveyed to the medulla oblongata or the medulla spinalis, and is reflected along the nerves to parts adjacent to, or remote from, that which has received the impression.

"It is by this reflex character that the function to which I have alluded is to be distinguished from every other. There are, in the animal economy, four modes of muscular action, of muscular contraction. The first is that designated VOLUNTARY: volition, originated in the cerebrum and spontaneous in its acts, extends its influence along the spinal marrow and the motor nerves in a DIRECT LINE to the voluntary muscles. The SECOND is that of RESPIRATION: like volition, the motive influence in respiration passes in a DIRECT LINE from one point of the nervous system to certain muscles; but as voluntary motion seems to originate in the cerebrum, so the respiratory motions originate in the medulla oblongata: like the voluntary motions, the motions of respirations are spontaneous; they continue, at least, after the eighth pair of nerves have been divided. The THIRD kind of muscular action in the animal economy is that termed involuntary: it depends upon the principle of irritability and requires the IMMEDIATE application of a stimulus to the nervo-muscular fibre itself. These three kinds of muscular motion are well known to physiologists; and I believe they are all which have been hitherto pointed out. There is, however, a FOURTH, which subsists, in part, after the voluntary and respiratory motions have ceased, by the removal of the cerebrum and medulla oblongata, and which is attached to the medulla spinalis, ceasing itself when this is removed, and leaving the irritability undiminished. In this kind of muscular motion the motive influence does not originate in any central part of the nervous system, but from a distance from that centre; it is neither spontaneous in its action nor direct in its course; it is, on the contrary, EXCITED by the application of appropriate stimuli, which are not, however, applied immediately to the muscular or nervo-muscular fibre, but to certain membraneous parts, whence the impression is carried through the medulla, REFLECTED and reconducted to the part impressed, or conducted to a part remote from it in which muscular contraction is effected.

"The first three modes of muscular action are known only by actual movements of muscular contractions. But the reflex function exists as a continuous muscular action, as a power presiding over organs not actually in a state of motion, preserving in some, as the glottis, an open, in others, as the sphincters, a closed form, and in the limbs a due degree of equilibrium or balanced muscular action - a function not, I think, hitherto recognized by physiologists.

The three kinds of muscular motion hitherto known may be distinguished in another way. The muscles of voluntary motion and of respiration may be excited by stimulating the nerves which supply them, in any part of their course, whether at their source as a part of the medulla oblongata or the medulla spinalis or exterior to the spinal canal: the muscles of involuntary motion are chiefly excited by the actual contact of stimuli. In the case of the reflex function alone the muscles are excited by a stimulus acting mediately and indirectly in a curved and reflex course, along superficial subcutaneous or submucous nerves proceeding from the medulla. The first three of these causes of muscular motion may act on detached limbs or muscles. The last requires the connection with the medulla to be preserved entire.

"All the kinds of muscular motion may be unduly excited, but the reflex function is peculiar in being excitable in two modes of action, not previously subsisting in the animal economy, as in the case of sneezing, coughing, vomiting, etc. The reflex function also admits of being permanently diminished or augmented and of taking on some other morbid forms, of which I shall treat hereafter.

"Before I proceed to the details of the experiments upon which this disposition rests, it may be well to point out several instances in illustration of the various sources of and the modes of muscular action which have been enumerated. None can be more familiar than the act of swallowing. Yet how complicated is the act! The apprehension of the food by the teeth and tongue, etc., is voluntary, and cannot, therefore, take place in an animal from which the cerebrum is removed. The transition of food over the glottis and along the middle and lower part of the pharynx depends upon the reflex action: it can take place in animals from which the cerebrum has been removed or the ninth pair of nerves divided; but it requires the connection with the medulla oblongata to be preserved entirely; and the actual contact of some substance which may act as a stimulus: it is attended by the accurate closure of the glottis and by the contraction of the pharynx. The completion of the act of deglutition is dependent upon the stimulus immediately impressed upon the muscular fibre of the oesophagus, and is the result of excited irritability.

"However plain these observations may have made the fact that there is a function of the nervous muscular system distinct from sensation, from the voluntary and respiratory motions, and from irritability, it is right, in every such inquiry as the present, that the statements and reasonings should be made with the experiment, as it were, actually before us. It has already been remarked that the voluntary and respiratory motions are spontaneous, not necessarily requiring the agency of a stimulus. If, then, an animal can be placed in such circumstances that such motions will certainly not take place, the power of moving remaining, it may be concluded that volition and the motive influence of respiration are annihilated. Now this is effected by removing the cerebrum and the medulla oblongata. These facts are fully proved by the experiments of Legallois and M. Flourens, and by several which I proceed to detail, for the sake of the opportunity afforded by doing so of stating the arguments most clearly.

"I divided the spinal marrow of a very lively snake between the second and third vertebrae. The movements of the animal were immediately before extremely vigorous and unintermitted. From the moment of the division of the spinal marrow it lay perfectly tranquil and motionless, with the exception of occasional gaspings and slight movements of the head. It became quite evident that this state of quiescence would continue indefinitely were the animal secured from all external impressions.

"Being now stimulated, the body began to move with great activity, and continued to do so for a considerable time, each change of position or situation bringing some fresh part of the surface of the animal into contact with the table or other objects and renewing the application of stimulants.

"At length the animal became again quiescent; and being carefully protected from all external impressions it moved no more, but died in the precise position and form which it had last assumed.

"It requires a little manoeuvre to perform this experiment successfully: the motions of the animal must be watched and slowly and cautiously arrested by opposing some soft substance, as a glove or cotton wool; they are by this means gradually lulled into quiescence. The slightest touch with a hard substance, the slightest stimulus, will, on the other hand, renew the movements on the animal in an active form. But that this phenomenon does not depend upon sensation is further fully proved by the facts that the position last assumed, and the stimuli, may be such as would be attended by extreme or continued pain, if the sensibility were undestroyed: in one case the animal remained partially suspended over the acute edge of the table; in others the infliction of punctures and the application of a lighted taper did not prevent the animal, still possessed of active powers of motion, from passing into a state of complete and permanent quiescence."

In summing up this long paper Hall concludes with this sentence: "The reflex function appears in a word to be the COMPlEMENT of the functions of the nervous system hitherto known."[2]

All these considerations as to nerve currents and nerve tracts becoming stock knowledge of science, it was natural that interest should become stimulated as to the exact character of these nerve tracts in themselves, and all the more natural in that the perfected microscope was just now claiming all fields for its own. A troop of observers soon entered upon the study of the nerves, and the leader here, as in so many other lines of microscopical research, was no other than Theodor Schwann. Through his efforts, and with the invaluable aid of such other workers as Remak, Purkinje, Henle, Muller, and the rest, all the mystery as to the general characteristics of nerve tracts was cleared away. It came to be known that in its essentials a nerve tract is a tenuous fibre or thread of protoplasm stretching between two terminal points in the organism, one of such termini being usually a cell of the brain or spinal cord, the other a distribution-point at or near the periphery - for example, in a muscle or in the skin. Such a fibril may have about it a protective covering, which is known as the sheath of Schwann; but the fibril itself is the essential nerve tract; and in many cases, as Remak presently discovered, the sheath is dispensed with, particularly in case of the nerves of the so-called sympathetic system.

This sympathetic system of ganglia and nerves, by-the-bye, had long been a puzzle to the physiologists. Its ganglia, the seeming centre of the system, usually minute in size and never very large, are found everywhere through the organism, but in particular are gathered into a long double chain which lies within the body cavity, outside the spinal column, and represents the sole nervous system of the non-vertebrated organisms. Fibrils from these ganglia were seen to join the cranial and spinal nerve fibrils and to accompany them everywhere, but what special function they subserved was long a mere matter of conjecture and led to many absurd speculations. Fact was not substituted for conjecture until about the year 1851, when the great Frenchman Claude Bernard conclusively proved that at least one chief function of the sympathetic fibrils is to cause contraction of the walls of the arterioles of the system, thus regulating the blood-supply of any given part. Ten years earlier Henle had demonstrated the existence of annular bands of muscle fibres in the arterioles, hitherto a much-mooted question, and several tentative explanations of the action of these fibres had been made, particularly by the brothers Weber, by Stilling, who, as early as 1840, had ventured to speak of "vaso-motor" nerves, and by Schiff, who was hard upon the same track at the time of Bernard's discovery. But a clear light was not thrown on the subject until Bernard's experiments were made in 1851. The experiments were soon after confirmed and extended by Brown-Sequard, Waller, Budge, and numerous others, and henceforth physiologists felt that they understood how the blood-supply of any given part is regulated by the nervous system.

In reality, however, they had learned only half the story, as Bernard himself proved only a few years later by opening up a new and quite unsuspected chapter. While experimenting in 1858 he discovered that there are certain nerves supplying the heart which, if stimulated, cause that organ to relax and cease beating. As the heart is essentially nothing more than an aggregation of muscles, this phenomenon was utterly puzzling and without precedent in the experience of physiologists. An impulse travelling along a motor nerve had been supposed to be able to cause a muscular contraction and to do nothing else; yet here such an impulse had exactly the opposite effect. The only tenable explanation seemed to be that this particular impulse must arrest or inhibit the action of the impulses that ordinarily cause the heart muscles to contract. But the idea of such inhibition of one impulse by another was utterly novel and at first difficult to comprehend. Gradually, however, the idea took its place in the current knowledge of nerve physiology, and in time it came to be understood that what happens in the case of the heart nerve-supply is only a particular case under a very general, indeed universal, form of nervous action. Growing out of Bernard's initial discovery came the final understanding that the entire nervous system is a mechanism of centres subordinate and centres superior, the action of the one of which may be counteracted and annulled in effect by the action of the other. This applies not merely to such physical processes as heart-beats and arterial contraction and relaxing, but to the most intricate functionings which have their counterpart in psychical processes as well. Thus the observation of the inhibition of the heart's action by a nervous impulse furnished the point of departure for studies that led to a better understanding of the modus operandi of the mind's activities than had ever previously been attained by the most subtle of psychologists.