Chapter X: Muscle (2)

Nothing so definitely removes muscle from the category of machines as its liability to fatigue. To speak of a muscle as tired is, of course, to transfer to an object a term which is applicable only to a phenomenon of consciousness; but it is necessary, unless a cumbrous expression is to be used, to designate thus the effect upon the muscle of prolonged activity. The petrol may be low in the tank, but the quantity burnt in the cylinder at each stroke is not reduced. If an isolated muscle is repeatedly stimulated by an electric current of a certain strength, the response which it makes improves for the first two or three induction shocks; then it begins to weaken. At each succeeding spasm the muscle shortens a trifle less than before. More remarkable than the diminution in the amount of work done by a muscle which is growing tired is the prolongation of the time taken both in contracting and in relaxing. Further, it has been shown that the fatigue which accompanies the contraction of an isolated muscle is not a condition dependent upon the shrinking of the store of energy which it possessed when it was first thrown into activity. Muscles undisturbed as to blood-supply, and contracting under the direction of the Will, also exhibit it. Speaking generally, it may be said that the tiring of muscle is not so much due to the exhaustion of its store of food as to accumulation of products of action. Vigour is restored to a tired muscle by passing through its bloodvessels a stream of salt-solution, which brings it no food, but washes away some of its waste. But the problem is far more complex than this. The machinery is not simply clogged with the products of its own activity. If the blood of a tired animal is injected into the vessels of one that is rested, the muscles of the latter exhibit the phenomena of fatigue. Evidently muscle is self-protective. During activity it prepares a “fatigue-substance” which poisons its own nerve-endings, making them worse conductors from nerve to muscle of the commands which descend from the brain. Not only does the fatigue-substance dull the nerve-endings in the particular muscle which has contracted, but, being distributed by the blood to the whole body, it produces a general effect. If the legs have been severely worked, they exhibit fatigue in the highest degree; but after a long walk the arms also are less ready and less capable than the state of their nutrition warrants.

The condition of stiffness experienced for a day or two after excessive exercise is due to various causes in combination. The fact that it may be remedied by encouraging the circulation through the muscles most affected, as by hot baths and massage, tempts us to assign it also in large measure to accumulation of products of action; but the means taken to reduce stiffness favour the nutrition of the muscles both by giving them more food and by carrying off their waste.

Equally remarkable with the self-protective disposition of muscle, which forbids it to give, except at the instance of increasingly urgent messages from the central nervous system, more than a part of the work of which it is capable, is its preparation for meeting an increased demand. It grows with use. Running increases the girth of the leg by developing especially the muscles of the calf. Raising weights enlarges the muscles of the shoulder and arm. Use-growth may reach inconvenient proportions. Nothing is more noticeable during the training of young athletes, whose nutritive responsiveness is at its height, than their liability to pass through a stage in which they are “muscle-bound.” Their legs grow bigger, but their pace falls off.

The development by means of exercises of a strong muscular system has received much attention during recent years. Our ancestors cultivated strength and agility in certain movements without paying much attention to the muscles by which the movements were performed. It is fashionable nowadays to lay stress upon the importance of maintaining an abundant musculature, because of its relation to general “fitness.” The balance between muscular activity and the organic functions which is observed by everyone who takes an active holiday proves beyond doubt that the nutritive condition of the various glands and of the heart and bloodvessels is in some degree dependent upon the condition of the muscles. Possibly they secrete into the blood other “messengers” in addition to fatigue-substance—messengers whose call wakes up the organs of digestion. The man who is so fortunate as to be able to use his muscles in the open air has no need of exercises in his bathroom. Failing out-of-door opportunities, much can be done by the systematic use of the various muscles working against resistance. It is alleged, and we are not disposed to dispute the justice of the contention, that movements made with the fullest degree of mental concurrence have a more rapid effect upon the growth of muscle than actions more or less unconscious. Muscle and nerve are parts of a single mechanism. It may be that fixing the attention on an exercise, and watching its performance in a looking-glass, aids the nutrition of muscles by increasing the influence of their nerves, possibly by improving the nutrition of their nerve-centre. Unfortunately, this is one of many theories which hardly come within the reach of a control experiment. Could one concentrate attention on the movements of the right arm, then absent-mindedly repeat them with equal vigour with the left, it might be possible to ascertain whether there is anything in this idea. Two other contentions with regard to the best way of performing movements, with a view to the promotion of muscular growth, appear to be justified by their results. Working against a moderate or light load is said to be more effective than putting muscles to a severe strain. A small number of maximal contractions, it is said, induce more rapid growth than many partial shortenings. According to this scheme, when a particular muscle needs strengthening, because in a certain action it is to be the chief performer, it is made to bring its two ends as near together as the plan of its attachments allows. Maximal shortening is apparently favourable to blood-supply and otherwise promotes nutrition.

=Tone.=—Hitherto we have spoken of quiescence and activity, as if muscle were doing nothing when not visibly contracting. A wrong impression may be engendered by these terms. Muscle is never idle. During sleep, and still more when a person is under the influence of anæsthetics, the muscles approach the condition of machines at rest. But again the language of the workshop is inapplicable. When a headless frog is hanging from a hook its legs are slightly bent. All its muscles are weakly contracted, if we understand by contraction a condition in which the length of muscle is less than it would be were it not alive. But the flexors are tenser than the extensors, hence the crooking of hip, knee, and ankle. If the sensory roots of the sciatic nerve are cut, the leg straightens out. So long as the nerve was intact the weight of the limbs acted as a stimulus to sensory nerve-endings, causing a reflex “tone” of the flexor muscles via the spinal cord. The tone of the extensor muscles was less because they were not stretched by the weight of the limbs. Every joint is under the influence of antagonistic muscles which are perpetually watching one another. When the limb is extended the flexors become anxious. When it is flexed the extensors get ready for a spring. Only when it is half flexed is there anything approaching to a truce. And this in most cases is the position of greatest comfort. But even when most at rest, muscles still possess a certain degree of tone. The tendency to shortening in one set causes it to pull against, and thereby increases the tone of, its opponents. When a muscle contracts it does not lift a loose bone. It has to overcome the tone of the muscles which would cause a movement in the opposite direction. And here another adjustment comes into play. The same gross stimulus which leads to the contraction of A starts impulses of a finer kind for B, directing it to relax its tone. We have seen how the heart and bloodvessels are under the influence of two sets of nerves of opposite sign—anabolic, diminishing irritability; katabolic, increasing it. All muscles are under similar management; but we can rarely detect the influence of the anabolic, inhibitory nerves, the brakes, because the katabolic display is overwhelmingly conspicuous. We must be content with two experimental demonstrations. An animal’s hamstrings have been cut; the flexor muscles of its thigh are therefore severed from their attachments below the knee. The tone of the extensor muscle keeps this joint extended. If now the pad of the foot be tickled, the flexor muscles contract, just as they would do if they were still able to carry out the reflex action of raising the foot. They cannot do this, because their tendons are divided; nevertheless, the knee bends owing to reflex relaxation of the extensor muscles. Still more striking evidence of reciprocal contraction and relaxation is afforded by the claw-muscles of a crayfish. A weak stimulus to its nerve causes the claw to set open; a stronger stimulus causes it to close. Both these movements are due, not to “contraction,” but to change of tone. Under certain conditions, a current passed through its abductor muscle, the claw being open at the time, causes closure by inhibiting the tone of this muscle. In this case the stimulus acts directly on the muscle, producing an effect which is opposite to the one we are accustomed to associate with stimulation; in place of contraction, relaxation.

Contraction of muscle, moving something, impresses one as a positive phenomenon. Relaxation seems negative—the undoing of contraction—and to a very large extent this attitude of mind is justified. Return of a muscle to its full length is due either to stretching by the weight it has lifted, or to the antagonism of other muscles. An isolated muscle lying on a pool of quicksilver does not return to its full length after it has contracted. But it is necessary to banish the machine idea. A machine gives out all the energy it has in store. Muscle is extremely parsimonious. No stimulus can induce it to part with more than a fraction of its energy. Recovery is as definite a function as disturbance. A machine starts when a crank is moved, stops when it is replaced. Muscle has a certain degree of automatism, although its tendency to act on its own account has been almost completely transferred to the governing nervous system. Muscle and nerve work together, and the efficiency of muscle depends upon the maintenance of its relations with its nerve. If the nerve is cut, the muscle atrophies. We will not stop to consider whether wasting may be properly attributed to disuse, or to vaso-motor changes. In its lowest form nervous influence shows itself in the regulation of the nutrition of muscle. A somewhat more forcible exhibition of control is seen in the regulation of tone. The maximum is reached when a wave of undoing which has passed down a nerve infects the protoplasm of muscle with the same tendency to disintegration. The muscle-substance explodes. The muscle shortens.

Remarkable evidence of the existence of muscle-tone is afforded by the =knee-jerk=. Place a person on an upright chair, with his legs crossed, muscles lax, foot hanging free. With a paper-knife or the end of a stethoscope, or even the hand used edgewise, tap the ligament which connects his knee-cap with his shin. The tap is instantly followed by a jerking forward of the foot. The deep muscles of the thigh, vastus, and crureus, have contracted. This phenomenon is easy to account for. When we are standing upright, the trunk is supported on three joints, of which one—the hip—is a perfect ball and socket, and the other two—knee and ankle—are of the same order so far as the absence of any provision for locking them is concerned. If the muscles on the front and the back of the leg did not constantly adjust our balance, by swaying the trunk forward when it falls back, and pulling it back when it sways forward, the joints of the leg would double up beneath us. A photographer knows how little confidence is to be placed in a man’s assertion that he is able to stand still. This see-saw of alternate contraction and relaxation is kept up by means of nerve-impulses which ascend from the nerve-endings surrounding the separate bundles of tendons, or from the Pacinian bodies which are found in abundance in the neighbourhood of tendons and ligaments, or from the elaborately twisted nerve-fibres found in muscle-spindles, or possibly from all three classes. Muscle and tendon are richly supplied with sense-organs susceptible to pressure and stretching. There is an abundance of nerve-endings to choose from. The slightest change in their tension, whether due to the muscle’s own contraction or to the action upon it of other muscles or weights, is recorded not only in the spinal cord, but also in the cortex of the cerebellum, and, if the contraction is an act of volition, in the cortex of the great brain. Although it was skin which was tapped, skin-nerves have nothing to do with the jerk. It was the result of the slight sudden stretching. In short, the tone-mechanism has been fooled. Notice the position of the leg. The knee is semiflexed; the foot is hanging free. There is nothing for the extensor muscles of the thigh to do. Now, if ever, they are justified in dozing. It is not to be wondered at that the sudden stretching of the ligament takes them off their guard, or that on waking they give a quite unreasonable start. The phenomenon is, as we asserted, easy to account for. It would also be easy to explain, if it were not for the extreme rapidity with which the jerk follows the tap. The interval is about one-hundredth of a second. This is thought to be too short to allow an impulse to ascend a sensory nerve, pass through the cord, and descend a motor nerve. It is true that these reflexes of adjustment must stand on a different level to other reflexes. The tone-impulses which cause them are incessantly patrolling to and fro from sense-organs to nerve-endings. The paths they follow must be the most open in the nervous system. Receptors and effectors must, in an electrician’s phrase, be incessantly switched on; or, to express the analogy more accurately, the flexor and extensor tone mechanisms are incessantly and reciprocally switching each other on and off. It must be confessed that it is very difficult to explain the knee-jerk if it be not a reflex action, but, as has been supposed, a direct response of the thigh muscles to their own stretching. The latter hypothesis does not appear to be reconcilable with its dependence upon the maintenance of the nervous connection of the muscles with the spinal cord. It cannot be elicited unless the “spinal arc” is intact. It ceases after the severance of either sensory or motor roots. Nor will it occur if the supply of blood to the lower end of the spinal cord has been cut off. Still more difficult is it to explain its extraordinary sympathy with everything that happens in the whole nervous system, if the impulses which cause it do not pass through the spinal cord. By a very simple mechanical arrangement it is possible to record the amplitude of the knee-jerk. The foot moves a lever which writes on a travelling surface. The jerk is elicited by the hammer of a clock strapped to the shin. In this way it is possible to extend the period of observation over several consecutive hours, the subject becoming completely oblivious of the movement his foot is making once a second, if it be screened from his view. In deep sleep the jerks stop; but the subject may doze, and still jerk follows tap. And the record made by his foot mirrors all the changes in his nervous system. If he clench his fist, the movement is reinforced, as it is when a child cries, a lamp is lighted, his ear itches. There is music in an adjoining room. His foot is the baton which beats _fortissimo_ to Wagner, and is lulled to _piano_ by the “Lieder ohne Wörte.” On a bright day this spinal pulse throbs gaily. It is indolent in dull, depressing weather. The knee-jerk is the physician’s guide to the condition of the nervous system.

=Elasticity of Muscles.=—Muscles are very extensible, and after stretching return to their original length. Their elasticity is a quality of great practical importance. It enables them to meet sudden resistance without rupture, as when a man alights from a height. At the moment when the feet touch ground elasticity dissipates the shock. The stretching of the muscles then leads reflexly to the increase of their tone. Here we see an advantage in the short reaction-time of the knee-jerk. Tone comes into play long before impulses generated by contact of the sole of the foot with the ground have had time to reach the brain, or even to induce reflex contraction through the spinal cord. The elasticity of the muscles is also of use in the performance of certain sudden actions. A pea is flicked across the room by pressing the thumbnail against the pad of a finger, or a finger against the thumb, and releasing it with a jerk.

An electrical change accompanies an impulse in its passage down a nerve, and a wave of contraction in its passage along a muscle. In 1788 Galvani observed that the hind-limbs of a frog, suspended by a metal hook to metal railings, twitched when the wind blew them against the bars. The hook passed through the lumbar plexus of nerves. He recognized that the cause of the twitch was the closing of a circuit. The birth of dynamic or galvanic electricity dates from this observation; and ever since this phenomenon was first observed the electric changes in nerve-muscle preparations made from frogs’ legs have been favourite subjects of research. Many observations with regard to nerve-conduction and muscle-contraction may be made, and many experiments performed, without special apparatus. A frog having been killed by cutting off its head, or by placing it beneath a tumbler with a wad of cotton-wool soaked in chloroform, the skin of the leg is removed, displaying the khaki-coloured muscles, bluish tendons, and bright white threads of nerve. A stretch of the largest nerve of the back of the thigh, the sciatic, is isolated. All the muscles of the thigh are then cut away and the bone nipped across just above the knee. The bones below the knee are removed, the superficial muscle of the calf, the gastrocnemius, being allowed to hang free, its bifid end attached to the fragment of thigh-bone. Its lower end terminating in the tendo Achillis, with its insertion into the prominence of the heel, is left intact. The bone is fixed in a clamp. A light lever made from a wooden spill is suspended from the tendo Achillis. The nerve may then be stimulated in various ways: by crushing in a pair of forceps, burning with a heated needle, touching with a drop of glycerin or a strong solution of salt. But of all methods of stimulation, the best is the current from an induction coil. Since it does not injure the nerve, it can be applied as often as may be desired. The amateur provided with an induction coil is in a position to study the relation between stimulus and response. He can vary the strength of the stimulus and vary the weight which the muscle has to lift. He can observe the progressive onset of fatigue, and otherwise gain much information regarding the behaviour of muscle as an isolated piece of apparatus.

It is the ambition of the expert to obtain absolutely correct records of the time-phases and of the changes in electric potential of nerve and muscle under varied experimental conditions. For this purpose he needs the finest apparatus which instrument-makers can furnish, and the knowledge and dexterity requisite for its employment. Consider, for example, the record of the change of form. A nerve-muscle preparation, obtained by the method already described, is arranged so that the point of the lever scratches on a rapidly travelling blackened surface. As the muscle contracts it makes a “tracing.” A tuning-fork vibrating at the rate of, say, 400 times a second also scratches a tracing on the same travelling-plate. It is easy to time the several phases of contraction and relaxation by comparing them with the undulations made by the tuning-fork. By means of an induction shock a single impulse is generated in the nerve and a single spasm evoked in the muscle. Our tracing shows that the spasm lasts about one-tenth of a second, and that about half this time is occupied by contraction, and half by relaxation. But the ascending curve is usually a little steeper than the descending curve, and the apex a little nearer to the commencement of ascent than to the termination of descent. An electric signal marked the instant at which the current was sent into the nerve. The time taken by the impulse in travelling from the spot where the electric current entered the nerve to its junction with the muscle can therefore be estimated. The contraction begins so much sooner or later, according as the shock is delivered nearer to, or farther from, the muscle. By shifting the electrodes up and down the nerve, the rate at which the impulse travels is directly measured. After the time that the impulse took in reaching the muscle has been allowed for, there still seems to be an interval before the muscle begins to shorten. This was termed the “latent period,” under the impression that some time is actually lost in turning the nerve-impulse into a muscle impulse. The impulse was supposed to be latent in the end-plates of the nerve. Various hypotheses were formulated as to the nature of the transformation. The progressive improvements in apparatus and methods is testified by the diminution in this latent period as given in the text-books of successive decades. It is now put at ¹/₄₀₀ second, and is regarded by most physiologists as a delay due to the inertia of the muscle. Owing to its elasticity, the molecular change in muscle does not immediately affect its shape. When the latent period appears to be longer—say ¹/₁₀₀ second—the balance is due to the inertia of the recording apparatus. Usually the curve shows a rise lasting ⁴/₁₀₀ second and a fall occupying ⁵/₁₀₀, due to the fact that inertia of muscle and apparatus delays the commencement of the rise, but does not hasten the termination of the fall.

When an impulse is generated artificially by an induction shock, a single spasm or twitch is the result; but in Nature contraction is never limited to a single twitch. Impulses descending from a motor nerve-cell to a muscle are always rhythmic. They follow at the rate of eight or ten a second in human nerves; and since in our muscles contraction and relaxation take longer than in a frog, a second impulse reaches the muscle before the effect of the first has passed away. The muscle has not had time to relax, when it is again called upon to contract. Hence a summation of contractions. The muscle continues to shorten until the maximum of contraction is reached. This condition is termed “tetanus,” to distinguish it from a single spasm. In fullest contraction the length of a muscle may be diminished by one-half, or even by two-thirds.

It would be impossible to treat of the =electrical phenomena= displayed by nerves and muscles without presupposing some acquaintance with the methods and laws of physics. As this is contrary to our understanding with our readers, we must be content with the statement of a few salient facts. At the moment when an impulse is passing along a nerve, or a wave of contraction along a muscle, the electric potential of the active part of the structure, whether nerve or muscle, is different from that of the not-acting parts on either side of it. A battery in its commonest form is a glass vessel containing sulphuric acid in which a plate of zinc and a plate of copper are immersed. The zinc is electro-positive as regards the copper. In a muscle the contracted portion is electro-positive as regards the parts uncontracted. The degree of positivity can be measured by connecting the muscle at two spots with the two wires of a galvanometer. When one wire makes contact with the contracted portion, and the other with a part which is not contracted, a current passes through the galvanometer, causing its needle to swing; and since the wave of contraction is not stationary, but passes down the muscle, the current is subsequently reversed. The wave, as it were, first tilts up one end, and then, passing on, tilts up the other, letting down the first. The contracted spot is electro-positive to the spot not contracted, and then the latter, contracting, becomes electro-positive to the former, which has relaxed. The needle of the galvanometer swings first to the left, then to the right. The importance of this method of investigation lies in the fact that the electric variation exactly represents, both in time and in intensity, the change which is occurring in nerve and in muscle. By following it, we can ascertain the rate at which an impulse travels down a nerve. We can determine its length and its “form.” Represented on paper, it is a wave. This wave travels in warm-blooded animals with the rapidity of 35 metres in a second. When it reaches a muscle, its rate—that is to say, the rate at which the wave of contraction invades the muscle—is 6 metres in a second. The time during which any particular level in the muscle remains contracted in a single spasm, under the influence of an artificial stimulus, is about 0·05 second. The length of the wave is 300 to 400 millimetres. These measurements give us a very clear idea of the events which occur in a nerve-muscle. An impulse picked up by a motor cell in the spinal cord runs down its axon—termed later a nerve-fibre—with great rapidity. Even the most distant muscle is reached in less than one-thirtieth of a second. From the end-plate of the nerve it travels in both directions along the muscle-fibre—or group of fibres, since each nerve divides into branchets for thirty to forty muscle-fibres—with reduced velocity. Every particle of each fibre rises and falls; but, seeing that the wave of contraction is much longer than the fibre, the whole fibre is in a state of contraction at the same time, although not with equal vigour throughout its whole length.

We cannot dismiss the further consideration of the electric phenomena of nerves and muscles without some inquiry into their meaning. It is evident that they are intimately related to the molecular changes which constitute an impulse. But at present the physics of the phenomena are beyond our grasp. We may speak in a general way of dissociation of ions; but we do not really know what is happening at the spot which is in a state of impulse. We cannot bring the transformation which it is undergoing into line with chemical and physical transformations which we understand. Probably the electrical phenomena which mark it are not peculiar to muscle and nerve. All living changes of state are of the same nature. Cellular activity, or protoplasmic activity, to use a better term, wherever it occurs, is accompanied by electrical change. But it so happens that nerve-substance and muscle-substance have a definite orientation which gives to the electric force a cumulative effect. In a liver-cell it is dispersed in all directions. In a muscle the change of potential at one particle is added to the change at the next, until the sum of all these changes, transmitted along the length of the fibre, is sufficiently large to deflect the needle of a galvanometer. Owing to its summation it attracts our attention.

Although they cannot tell the true significance of the electromotive change which marks the passage of an impulse, physiologists are in a much better position now than formerly to controvert certain popular misconceptions. There is no such thing as “nerve-force” in the vulgar sense. A nerve does not transmit energy to a muscle. The muscle obtains the energy which it dispenses when contracting from the foods with which the blood supplies it. The nerve transmits an excitation. Over-excitability is not a sign of strength, but of weakness. Nor is an impulse in a nerve an electric current. It may be generated by an electric shock, but a chemical stimulus is equally as effective. The slow rate at which it travels, as compared with electricity, puts it altogether out of comparison with an electric current. Its relatively rapid progress, on the other hand, equally excludes the hypothesis that it is a movement of ions, as that phenomenon is observed in solutions of salts.

What is the nature of the process by which energy is conveyed along a nerve? When speaking of the passage of impulses from receptors to the central nervous system, and through this to effectors, we have used the vague expression “molecular change,” to avoid the necessity of being more precise. But the problem is of such profound interest that we look with eagerness for any hint of the direction from which light will eventually be thrown upon it. Recent discoveries regarding the nature of electricity, combined with investigations at present in progress as to the physical constitution of proteid substances, give more than a hint. Hitherto the choice has lain between a chemical and a physical explanation; now the border-line between chemistry and physics, always wavering, has disappeared. The hypothesis that an impulse is a progression of chemical change has meant in the past that the “wave” was due to the oxidation of substances contained in nerve, with liberation of CO₂ and H₂O. Various considerations render such metabolism of the substance of which nerve-fibres are composed improbable. In the first place, nerve-cell bodies contain a store of material, tigroids (p. 320), which is recognizably drawn upon during nervous activity. It would appear, therefore, to be the tigroids, and not the substance of the nerve-fibre, which supply the energy transmitted along a nerve. Then, again, the axon of a nerve-fibre, enclosed as it is in a tube of fat, is peculiarly ill-placed for the reception of the nourishment which would be needed to make up for waste, if its metabolism be fluctuating and at times excessive. Nor have nerves more than a very meagre blood-supply. Secondly, observation does not give any support to the hypothesis of fluctuating metabolism. A nerve does not give off more CO₂ when active than when passive. Nor does it become acid. Thirdly, nerves, or, to be quite accurate, medullated nerves, are indefatigable. Their capacity for conduction is not diminished by previous use, as it would be were it dependent upon their reserve of nutriment. These various considerations rule out a “chemical” explanation of the old-fashioned type. It is premature to do more than outline the “physical” theory which seems destined to take its place; and the reader will perhaps forgive if, for the sake of clearness, the case is put with unjustifiable definiteness and simplicity. Proteid substances are constituted of clusters of molecules. The form of the clusters depends upon the salts (or, more precisely, the ions) with which they are associated, and the associations depend upon the electric charges which the ions carry. In resting nerve-protoplasm the clusters are small, and, since the total surface-area of a number of small spheres is greater than the surface-area of the same weight of matter when condensed into large spheres, there is, so to speak, more surface for the ions to cling to. Conversely, when the ions leave the small clusters, the latter are not protected from the influence of mutual attraction. They fuse into larger clusters. Fusion is carried to its extreme limits when a protein coagulates. A nerve-impulse is a “wave” of partial coagulation. The positive electricity generated in a cell-body by the metabolism of its tigroids repels the positively charged ions which cling to the nearest protoplasm-clusters in the axon. Like acrobats swinging from trapeze to trapeze, each flight of ions dispossesses the ions from the clusters in front of it; and in this way the disturbance progresses down the axon as an electric wave.

Thus we interpret the shadow cast by a theory of which either of several pioneers who are diligently climbing may at any time obtain a view. The conductivity of protoplasm (and what is true of its conductivity will be found to hold good equally for its irritability and changeableness of form) is due to the readiness with which its molecules enter into unstable associations with electrolytes. The instability of these associations is related to the tendency of the molecules to cluster. An impulse is passed along a nerve as a displacement of ions; the ions being transferred from one molecule, or group of molecules, to the next. Such an explanation of an impulse involves no chemical breakdown of nerve-substance during its passage along a nerve. It transfers the metabolism which liberates energy (reinforcing the impulses which have originated in sense-organs) to the nerve-cell bodies. It is based upon certain experimental data which appear to have been established; but, like all other hypotheses which are intended to account for physiological phenomena, this one must be brought to the test by varying the conditions under which impulses pass along nerves, and ascertaining whether the consequent alteration in the force, rate, and other attributes of the phenomena are in accordance with physical laws. In applying these tests to the activities of protoplasm, we are, however, met by an insuperable difficulty. The matter which transmits nerve-impulses is alive. We have no laboratory standards by which to judge whether the changes in conduction which are produced by changes in the conditions of the conductor are, or are not, consonant with physical theory. It is with protoplasm that we are dealing, and not with a mixture of proteins in solution. If we surround a nerve with nitrogen, it loses its conductivity in five hours, to recover it when oxygen replaces the neutral gas. This has been regarded as proving that metabolism of the nerve is necessary for the transmission of impulses. But conductivity is a phenomenon of life. Deprivation of oxygen for five hours must bring the nerve-substance to the verge of death. It might be argued that the retention by the nerve for so long a time of its power of conducting impulses shows that its metabolism is not a cause of the phenomenon. Again, it has been shown that warming the nerves of cold-blooded animals greatly increases the rapidity of conduction. It is more than doubled in the nerves of the “foot” of a slug-and a similar increase has been proved for the nerves of a frog-by a rise of temperature of 10° C. Reflecting on the results of this experiment, a physicist would exclaim: “Then an impulse is a wave of chemical change. A rise of 10° C. increases the rate of chemical processes from two to three times; whereas no known physical process is accelerated by more than 5 to 15 per cent.” But the physiologist remembers that a rise of temperature of 10° C. increases all the activities of a frog. He is hardly prepared to say that its greater vivacity may not be the expression of more rapid oxidation; but he sees no fore-ordained balance of vital enterprise and chemical change. He is, or ought to be, extremely suspicious of any explanation which appears to over-ride physical laws; yet, at the same time, he is aware that until he has more accurate knowledge regarding the constitution of protoplasm he will not be in a position to understand how physical laws apply. The protoplasmicity of protoplasm is increased by warmth. What change of molecular constitution does this imply?

The view that in a muscle molecular change gives rise to an electrical change, which in turn produces the change in form, has been very widely held. The hypothesis was based on observations which seemed to show that the electric variation travels a little ahead of the wave of contraction; but every improvement in recording apparatus has diminished this apparent want of synchronism. There can be little doubt but that the lagging behind of the wave of contraction is due to the inertia of the muscle and of the recording apparatus. Molecular change and electric variation are simultaneous. If this be true, the electric change cannot be regarded as the cause of the molecular change, in the sense, at any rate, in which they used to be considered as cause and effect.

The =power of muscle= varies as its cross-section. For human muscles the maximum lift amounts to from 7 to 10 kilogrammes for each square centimetre. This is a large figure, but it must be remembered that, owing to the arrangement of the bones as levers, most muscles act at a great mechanical disadvantage. The greater the difference in distance from the fulcrum between the point of application of the force and the point of incidence of the weight, when the force acts nearer to the fulcrum than the weight, the greater is the mechanical disadvantage. The greater also is the rapidity with which the weight is lifted. What is lost in strength is gained in swiftness. Contrast the slow steps of a negro, whose long heel separates the point of application of the power (tendo Achillis) from the fulcrum (the ankle-joint), with the springy movements of a European. A European needs, and as a rule has, a better developed calf, which allows him his more sprightly gait, without sacrificing his carrying power. Our preference for slender wrists and ankles is not purely æsthetic, unless we admit, as may be maintained, that all natural canons of taste rest upon utility. Slimness of joints means nimbleness. A few muscles act directly, without loss of power—as, for example, the masseter, which lifts the lower jaw (hence a grand capacity for cracking nuts)—but most muscles move levers of considerable length. Compare with the masseter the biceps and brachialis which lift the forearm. Their tendons are inserted into the radius and the ulna at a distance from the elbow-joint which is about one-tenth as great as the distance from it of a weight held in the hand. Their united cross-section is about 16 square centimetres: (16 × 10) / 10 = 16. One cannot hold out in the hand, the elbow being pressed against the side, so that these muscles alone are acting, a greater weight than 16 kilogrammes (34 pounds), although the muscles are exerting a traction ten times as great as this. The strength of muscle when pulling straight is well illustrated by the thick white mass in the centre of an oyster. It keeps the shell closed until a force equal to 1,300 times the animal’s weight has been applied. This muscle also affords a good illustration of the part played by reflex contraction in opposing stretching—the reaction by which tone is maintained. Anyone who inserts an instrument, such as the end of a screwdriver, between the slightly open valves of an oyster lying under water will find that he needs to give it an exceedingly smart twist if he would catch the muscle asleep. Stretching it causes a reaction proportional to the stretching force.

The fact that the output of energy by muscle is proportional, within certain limits, to the work to be done, is brought out even in laboratory experiments. A nerve-muscle preparation teaches that the amount of work is not a function of the stimulus. Within certain limits a stronger stimulus evokes a higher and stronger lift; but the stimulus remaining the same, the work done by muscle (_i.e._, the product of weight multiplied by height) is, up to a certain optimum, increased by increasing the weight. Often a very light load is not lifted as high by a nerve-muscle preparation as a slightly heavier one. No satisfactory theory of this reaction to load has yet been formulated. Explanations have been put forward, but they merely substitute one unknown for another, a not uncommon drawback to explanations.

Muscles are strongest when at their full physiological length. As he dips an oar into the water a man exerts the greatest force of which he is capable, provided that he is not guilty of “missing the beginning.” Hands over the stretcher, body between the knees, ankle, knee, hip, fully flexed, arms straight—all his strongest muscles are at their greatest physiological length. Rowing is an exercise which has no rival. Every muscle in the body, from little toe to little finger, comes into play under the conditions which suit it best. And not less admirable is the effect upon the abdominal muscles during recovery at the end of the stroke; and the rhythmic movement which encourages deep and measured respiration.

The greatest output of work is obtained when muscles contract against a progressively diminishing load. Towards the end of the lift the load must be small, if contraction is to be carried to its extreme limit. The provision for this is well seen in the case of the muscles of the arm when lifting a weight up to a position above the head. A portmanteau is held in the hand. Its handle is gripped by flexing the fingers. And here it may be noted that, since the range of movement of a muscle varies as its length, the thumb and fingers are not worked only by muscles contained in the palm of the hand. Fingers are bent and wrist flexed by muscles of which the origin is carried up even to the lower end of the humerus. As the portmanteau hangs by the side, biceps and brachialis are at their fullest length. Suppose it to be necessary to place it on a cab. These muscles begin the work under the best conditions. They could not, however, lift the portmanteau far did not the muscles of the shoulder displace the elbow from the side, so that at the end of their pull, the forearm being almost vertical, the muscles of the arm have little more to do than to move the hand inwards towards the head, in preparation for the extensor thrust. The secret of getting the greatest amount of work out of any particular muscle lies in securing for it the due co-operation of other muscles.

ELECTRIC ORGANS.

Muscle disperses energy in the forms of mechanical work, heat and electricity. Its structure, as already pointed out, is peculiarly favourable for the display of electromotive force. In certain fishes muscle is so modified as to give an electric discharge without developing mechanical work. The production of an electric change is a by-phenomenon of muscular activity. It becomes the sole function of an electric organ. If the skin be removed from the tail of a skate, a cylindrical column of brawny tissue about the size of a finger will be found embedded amongst the muscles near its root on either side. These are electric organs, although so weak that it is barely possible to feel the shock which they give in a live fish. The nearly allied Torpedo of the Mediterranean has far more powerful batteries. They are situate near its gills, occupying the whole thickness of the fish from skin to skin. When the back of a torpedo is pressed, it discharges a current of 30 volts, or even more. Still more violent are the shocks given by an eel—Gymnotus—which haunts the tributaries of the Amazon, a terror to all who have to cross their fords on foot; or the African fish, Malapterurus. The current which these animals develop attains an intensity of 200 volts. With the exception of those of Malapterurus, all electric organs are modified muscle, and closely similar in structure. The organs of Malapterurus appear to be modified glands. The skate’s electric organ may be taken as typical of the rest. When sliced with a knife, it is seen to be divided by firm connective tissue into minute chambers. These chambers are piled into hexagonal columns, which lie lengthwise in the organ (they are set dorso-ventrally in Torpedo). Each chamber contains a jelly-like substance which embeds an electric disc. The disc divides the chamber into a smaller anterior and a larger posterior compartment. Each chamber is supplied with several nerves which ramify into innumerable twigs on the front surface of the disc. The development of the disc must be considered for a moment if its structure is to be understood. It starts life looking as if it would grow into a voluntary muscle-fibre. A nerve joins it, forming an end-organ in the usual way. Then the end-organ increases its spread unduly, while the rest of the fibre fails to grow. The structure becomes toadstool-shaped, with the nerve arborizing on the seat of the stool. The front aspect of the disc, therefore, corresponds to a nerve-ending in a muscle. Its middle layer indicates clearly that the fibre makes an abortive attempt to develop cross-striation. It is laminated, the laminæ strangely contorted; in section they appear, not as plain lines, but as rows of dots, evidently a suggestion of longitudinal striation. The posterior layer of the disc consists of granular protoplasm drawn out as a number of short backwardly directed tongues, and one long process, the stem of the stool. No structure could be more suggestive of the function of the organ; but no one has as yet succeeded in catching the suggestion and pressing it into a definite explanation of the way in which it works. Certain physiologists, laying great stress on the fact that the functional connections between an electric organ and its nerves are not easily interrupted by the administration of curari, atropin, and other drugs, which block the passage of impulses from nerves to muscles, look upon the nerve-layer of the disc as the generator of electricity, and the rest as an accumulator or resonator, which stores, or exaggerates, the electric charge. Others consider that the portion of the disc which is altered muscle-fibre—the middle, or middle and posterior layers—generates the electromotive force, the nerve simply calling it into activity. All agree that a brief interval (about 0·003 second) elapses between the arrival of the nerve-impulse and the discharge of an electric shock. This “latent period” may be used as an argument in favour of either view. It would be in harmony with the general account which we have already given of protoplasm as a liberator of energy to suppose that a nerve-impulse, having reached a disc, immediately infects the protoplasm of the disc, inducing molecular commotion, and that the ions move in such directions as to disturb the electric equilibrium of the disc, its front surface becoming in relation to the back as zinc to copper in a battery. The current generated in the fish is in the direction from head to tail. It is certain that the change does not occur until an impulse reaches the organ. The organ is not charged by the nervous system during a period of inactivity, and then discharged by a releasing impulse. This is sufficiently evident from the fact that when a piece of the organ, with its nerve, is removed from the fish, although much sooner exhausted, it responds like a nerve-muscle preparation to repeated stimulation.

A shows the compartments into which septa of fibrous
tissue divide the organ. In the centre of each
compartment is a disc formed from a modified
muscle-fibre. Nerves ramify in abundance on its
anterior surface. B, a minute portion of a disc. At
the top are seen nerve-fibres in delicate nucleated
sheaths; then follow the nucleated layer with which
they come in contact, the contorted laminæ which
represent the striations of the muscle-fibre, the
granular nucleated substance of its posterior
layer, some connective tissue, a capillary
bloodvessel containing oval nucleated corpuscles.
In a tissue space, a single coarsely granular
leucocyte is to be seen.]

The usefulness of a torpedo’s electric organs is unmistakable. They are powerful enough to paralyse every animal that touches its back, whether foe or little fish suitable for food. But of what service is its feeble battery to a skate? This and the allied question as to the advantages which can have accrued to the ancestors of the torpedo who first began to change innocent muscle into a weapon of offence are usually answered by pointing to the liability of flat fish lying on the bottom of the sea to become resting-places of parasites, corallines, and other fixed growths. Very mild shocks would suffice to disturb the peace of would-be settlers. In the same way, the electric organs of fresh-water fish may, when rudimentary, have protected the skin from invasion by moulds.

LUMINOUS GLANDS.

If it be difficult, when considering the dispersal of energy as mechanical work, heat, or electricity, by living tissues, to bring the phenomena into line with those of which physics takes experimental cognizance, how are we to approach the problems involved in the generation of light? Yet the photogenic property of protoplasm is widely distributed. Protozoans and various other invertebrate animals cause the so-called phosphorescence of the sea. The abysmal depths of ocean are lighted by forests of luminous polyps, and traversed by fishes whose heads are furnished with lamps. By her own light the female glow-worm enables her winged mate to keep his tryst. Fireflies (Lampyrus) flash amongst the orange-trees of Italy, and blaze (Pyrophorus) beneath the mangoes of Ceylon.

Luminous organs vary too widely in structure to allow us to pick out, as in the case of electric organs, the features which are common to them all. In Pyrophorus the organ is a double mass of cylindrical cells near the tip of the abdomen. The cells are set vertically to the surface, and are supported by a tubular membrane. Their substance contains a kind of fat. Beneath them there is a layer of cells, not luminous, but evidently a part of the photogenic apparatus, containing chalky granules. The organ is well supplied with nerves and with respiratory tubes (tracheæ).

More interesting than its structure is the study of the peculiar character of the light which the organ emits. It gives a spectrum which extends from the red (beyond Fraunhofer’s line B of the solar spectrum) to the first blue rays (F). It shows no lines. Green rays appear only when the light is bright, and then they are the brightest of all the rays. The light is practically destitute of actinic or chemical rays. A photographic plate may be exposed for several minutes, almost without changing, to the light of a firefly bright enough to enable one to read with ease in a dark room; whereas light of equal brilliance from any other source would change it in the fraction of a second. Nor are heat-rays mixed with the light. Measurements show that the activity of the photogenic organs does not give rise to any greater rise of temperature than would occur in the case of any other gland.

The contrast between the emission of light by an animal and its production in any other manner is very striking when the physical evidence, or want of evidence, of what happens in the protoplasm which produces it is considered. The fact that no heat accompanies the light precludes us from attributing it to oxidation. If a firefly is enclosed in a vessel of oxygen, its lamp burns no brighter—clear evidence that its luminosity has nothing in common with the burning of a match or the glowing of a stick of phosphorus. Nor is the lamp put out when the insect is suddenly exposed to great cold (-100° C.). It continues to shine until the cold kills it. There is no relation between the luminosity of a firefly and the phenomenon termed “phosphorescence” by physicists. Sulphide of calcium—the substance used for rendering matchboxes visible in the dark—returns light which it has absorbed. A firefly’s power of emitting light is in no wise affected by keeping it for a long while in the dark.

Like all other events in vital chemistry, the generation of light by protoplasm is due to a process of fermentation. The luminous organs may be crushed, and the mixture of fermentable substance and ferment extracted with water. The extract is luminous. If an extract is prepared rapidly, and evaporated to dryness _in vacuo_, the residue glows when moistened with water. That two substances are present in the extract, one (luciferin) fermentable, the other (luciferase) a ferment, is proved by the following experiment: A certain quantity of extract is divided into two portions. One part (A) is allowed to glow until its capacity for emitting light is exhausted. The other portion (B), as soon as it is separated, is heated to 55° to kill the ferment. B still contains luciferin; A contains luciferase, although all its luciferin has been used up. Recombined, the extract is luminiferous.