163. Vitality is characterized by incessant molecular movement, both of composition and decomposition, in the building up of structure and the liberation of energy. The life of every organism is a complex of changes, each of which directly or indirectly affects the statical and dynamical relations, each being the resultant of many co-operant forces. In the nourishment of every organite there is an accumulation of molecular tension, that is to say, stored-up energy in a latent state, ready to be expended in the activity of that organite; and this expenditure may take place in a steady flow, or in a sudden gush. The molecular movements under one aspect may be called convergent, or formative: they build the structure, and tend to the state of equilibrium which we call the statical condition of the organite, i. e. the condition in which it is not active, but ready to act. Perfect equilibrium is of course never attained, owing to the incessant molecular change: indeed Life is inconsistent with complete repose. Under another aspect the molecular movements may be called discharging: they constitute the dynamic condition of the organite, in which its functional activity appears. The energy is now diverted, liberated, and the surplus, over and above that which is absorbed in formation, instead of slowly dribbling off, gushes forth in a directed stream. The slow formation of a secretion in a gland-cell, and the discharge of that secretion, will illustrate this; or (if muscular tone be admitted) the incipient contraction of the chronic state, and the complete contraction of the dynamic state, may also be cited.
164. The discharge which follows excitation may thus be viewed as a directed quantity of molecular movement. Because it is always strictly relative to the energy of tension, and is inevitable when that tension attains a certain surplus over what is required in construction, there is a limit, 1°, to the growth and evolution of every organite, and every organism (comp. Problem I. § [118]), and, 2°, to its dynamical effect. When there is no surplus, the organite is incapable of discharge: it is then exhausted, i. e. will not respond to stimulus.
165. The speciality of nerve-tissue is its pre-eminence in directive energy. Like all other tissues, it grows, develops, and dies; but above all others it has what we call excitability, or readiness in discharging its energy in a directed stream. By its topographical distribution it plays the functional part of exciting the activity of other tissues: it transmits molecular disturbance from periphery to centre, from centre to centre, and from centre to muscles, vessels, and glands. When a muscle is excited it moves, and when a gland is excited it secretes; but these actions end, so to speak, with themselves; the muscle does not directly move any other muscle;[184] the gland does not directly excite any other gland. The nerve, on the contrary, has always a wide-spreading effect; it excites a centre which is continuous with other centres; and in exciting one muscle, usually excites a group. Hence the nervous system is that which binds the different organs into a dynamic unity. And Comparative Anatomy teaches that there is a parallelism between the development of this system and the efficient complexity of the organism. As the tissues become more and more specialized, and the organs more and more individualized, they would become more and more unsuited to the general service of the organism, were it not that a corresponding development of the nervous system brought a unifying mechanism.
The great instability of neurine, in other words, its high degree of tension, renders it especially apt to disturb the tension of other tissues. It is very variable; and this variability will have to be taken into account in explaining the restriction of discharges to particular centres. A good example of exaggerated tension is furnished by strychnine poisoning. The centres are then so readily excitable that a touch, or a puff of cold air on the skin, will determine convulsions. And it is worthy of remark that for some hours after this convulsive discharge the centres return to something like their normal state; and the animal may then be stroked, pinched, or blown upon without abnormal reactions. But during this interval the centres are slowly accumulating excess of tension from the poisoned blood; and at the close, convulsions will again follow the slightest stimulus. This alternation of exhaustion and recrudescence is noticed by Schröder van der Kolk in the periodicity of the phenomena exhibited in spinal disease.[185]
THE PROPAGATION OF EXCITATION.
166. Understanding, then, that the propagation of an excitation depends on the state of tension of the tissue, and always follows the line of least resistance, whichever that may be at the moment, we have to inquire whether the transmission takes place only in one direction, from periphery to centre in sensory nerves, and from centre to periphery in motor nerves? By most physiologists this is answered affirmatively. Indeed a special property has been assigned to each nerve, in virtue of this imaginary limitation of centripetal and centrifugal conduction. The “nerve-current” (accepted as a physical fact, and not simply a metaphor) is supposed to “flow” from the central cells along the motor nerve to the muscles; but by a strange oversight the current is also made to “flow” towards the central cells which are said to produce it! Now although the fact may be, and probably is, that normally the sensory nerve, being stimulated at its peripheral end, propagates the stimulation towards the centre, and the motor nerve propagates its central stimulation towards the periphery, the question whether each nerve is not capable of transmission in both directions is not thus answered. A priori it is irrational to assert that nerves fundamentally alike in composition and structure are unlike in properties; and we might as well suppose that a train of gunpowder could only be fired at one end, as to suppose that a nerve could only be excited at one end. And how does the evidence support this a priori conclusion? Dubois Reymond proved that each nerve conducted electricity in both directions; but as Neurility has not been satisfactorily shown to be identical with the electric current, this may not be considered decisive. Such a doubt does not hang over the following facts. M. Paul Bert, pursuing John Hunter’s curious experiments on animal grafting, has grafted the tail of a rat under the skin of the rat’s back, the tip of the tail being inserted under the skin, its base rising into the air, so that there is here an inversion of the normal position. In the course of time Sensibility gradually reappears in this grafted tail; and at the end of about twelve months the rat not only feels when the tail is pinched, but knows where the irritation lies, and turns round to bite the pincers.[186] Here we have a case of a sensory nerve reversed, yet transmitting stimulation from the base to the tip of the tail, instead of from the tip to the base, as in a normal organ. Vulpian and Philippeaux having divided two nerves, united the central end of the sensory nerve with the peripheral end of the motor nerve; when the organic union was complete, and each nerve was formed out of the halves of two different nerves, the effect of pinching one of these was to produce simultaneously pain and movement, showing that the excitation was transmitted upwards to the centre, and downwards to the muscles.[187] It may be compared with a train of gunpowder having a loaded cannon at one end and a bundle of straw at the other, when if a spark be dropped anywhere on this train, the flame runs along in both directions, explodes the cannon, and sets alight the straw.
167. Indeed we have only to remember the semi-liquid nature of the axis cylinder to see at once that it must conduct a wave of motion as readily in one direction as in another. A liquid transmits waves in any direction according to the initial impulse. There is consequently no reason for asserting that because the usual direction is centripetal in a sensory nerve, and centrifugal in a motor nerve, each nerve is incapable of transmitting excitations in both directions. And I think many phenomena are more intelligible on the assumption that neural transmission is in both directions. If the eye is fixed steadfastly on a particular color during some minutes, the retina becomes exhausted, and no longer responds to the stimulus of that color: here the stimulation is of course centripetal. But if instead of looking intently on the color, the mind (in complete absence of light) pictures it intently, this cerebral image is equally capable of exhausting the retina; and unless we believe that color is a cerebral, not a retinal phenomenon (which is my private opinion), we must accept this as proof of a centrifugal excitation of a sensory tract. Another illustration may be drawn from the muscular sense. There may be a few sensory fibres distributed to muscles; but even if the observations of Sachs[188] should be confirmed, I do not think that all muscle sensations can be assigned to these fibres, but that the so-called motor fibres must also co-operate. When a nerve acts upon a muscle, the muscle reacts on the nerve; and when a nerve acts on a centre, the centre reacts on the nerve. The agitation of the central tissue cannot leave the nerve which blends with it unaffected; the agitation of the muscular tissue must also by a reversal of the “current” affect its nerve. Laplace points out how the movement of the hand which holds a suspended chain is propagated along the chain to its terminus, and if when the chain is at rest we once more set that terminus in motion, the vibration will remount to the hand.[189] The contraction of a muscle will not only stimulate the sensory fibres distributed through it, but also, I conceive, stimulate the very motor fibres which caused the contraction, since these fibres blend with the muscle.[190]
168. To understand this, it is necessary to remember that the stimulation of a nerve does not arise[191] in the changed state of that nerve, but in the process of change, i. e. the disturbance of the tension. The duration of the stimulation is that of the changing process, and the intensity increases with the differential of the velocity of change. So that when a nerve which has been excited by a change of state returns to its former state, this return—being another change—is a new excitation. That it is not the changed state, but the change, which is operative, explains the fact noted by Brown Séquard: a frog poisoned by strychnine, when decapitated and all respiration destroyed, will remain motionless for days together, if carefully protected from all external excitation; but its nervous system is in such a state of tension all this time that the first touch produces general convulsions. Freusberg also notes that if a brainless frog be suspended by the lower jaw, and one foot be pinched, the other leg is moved at first, then quickly droops again, and remains at rest until the pincers are removed from the pinched foot, when suddenly all four legs are violently moved by the stimulation which the simple removal produces. Let us also add the well-known and significant fact that if a nerve be divided rapidly by a sharp razor, neither sensation nor motion is produced, because the intensity of a stimulus being, to speak mathematically, the function of the changing process, the duration of the process is in this case too brief. On the same ground the application of a stimulus will excite no movement, if the force be very slowly increased from zero to an intensity which will destroy the nerve; but at any stage a sudden increase will excite a movement.
169. We may group all the foregoing considerations in this formula:
Law I. Every neural process is due to a sudden disturbance of the molecular tension. The liberated energy is discharged along the lines of least resistance.