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combination of C and E, or, in general, that no one note ever gives the effect of a combination or blend of notes higher and lower than itself. Homogeneous orange light gives the sensation of a blend of red and yellow; but there is nothing like this in the auditory sphere. In light, some wave-lengths give the effect of simple colors, as red and yellow; and other wave-lengths the effect of blends, as greenish yellow or bluish {233} green; but in sound, every wave-length gives a tone which seems just as elementary as any other.

There is nothing in auditory sensation to correspond to white, no simple sensation resulting from the combined action of all wave-lengths. Such a combination gives noise, but nothing that seems particularly simple. There is nothing auditory to correspond with black, for silence seems to be a genuine absence of sensation. There are no complementary tones like the complementary colors, no tones that destroy each other instead of blending. In a word, auditory sensation tallies with its stimulus much more closely than visual sensation does with its; and the main secret of this advantage of the sense of hearing is that it has a much larger number of elementary responses. Against the six elementary visual sensations are to be set auditory elements to the number of hundreds or thousands. From the fact that every distinguishable pitch gives a tone which seems as simple and unblended as any other, the conclusion would seem to be that each was an element; and this would mean thousands of elements. On the other hand, the fact that tones close together in pitch sound almost alike may mean that they have elements in common and are thus themselves compounds; but still there would undoubtedly be hundreds of elements.

Both sight and hearing are served by great armies of sense cells, but the two armies are organized on very different principles. In the retina, the sense cells are spread out in such a way that each is affected by light from one particular direction; and thus the retina gives excellent space information. But each retinal cell is affected by any light that happens to come from its particular direction. Every cone, in the central area of the retina, makes all the elementary visual responses and gives all the possible color sensations; so it is not strange that the number of visual {234} elements is small. On the other hand, the ear, having no sound lens, has no way of keeping separate the sounds from different directions (and accordingly gives only meager indications of the direction of sound); but its sense cells are so spread out as to be affected, some by sound of one wavelength, others by other wave-lengths. The different tones do not all come from the same sense cells. Some of the auditory cells give the low tones, others the medium tones, still others the high tones; and since there are thousands of cells, there may be thousands of elementary responses.

Theory of Hearing

The most famous theory of the action of the inner ear is the "piano theory" of Helmholtz. The foundation of the theory is the fact that the sense cells of the cochlea stand on the "basilar membrane", a long, narrow membrane, stretched between bony attachments at either side, and composed partly of fibers running crosswise, very much as the strings of a piano or harp are stretched between two side bars. If you imagine the strings of a piano to be the warp of a fabric and interwoven with crossing fibers, you have a fair idea of the structure of the basilar membrane, except for the fact that the "strings" of the basilar membrane do not differ in length anywhere like as much as the strings of the piano must differ in order to produce the whole range of notes. Now, a piano string can be thrown into "sympathetic vibration", as when you put on the "loud pedal" (remove the dampers from the strings) and then sing a note into the piano. You will find that the string of the pitch sung has been thrown into vibration by the action of the sound waves sung against it.

Now suppose the strings of the basilar membrane to be tuned to notes of all different pitches, within the range of {235} audible vibrations: then each string would be thrown into sympathetic vibration whenever waves of its own vibration rate reached it by way of the outer and middle ear; and the sense cells standing over the vibrating fibers would be shaken and excited. The theory is very attractive because it would account so nicely for the great number of elementary tone sensations (there are over 20,000 fibers or strings in the basilar membrane), as well as for various other facts of hearing--if we could only believe that the basilar membrane did vibrate in this simple manner, fiber by fiber. But (1) the fabric into which the strings of the membrane are woven would prevent their vibrating as freely and independently as the theory requires; (2) the strings do not differ in length a hundredth part of what they would need to differ in order to be tuned to all notes from the lowest to the highest, and there is no sign of differences in stretch or in loading of the strings to make up for their lack of difference in length; and (3) a little model of the basilar membrane, exposed to sound waves, is seen to be thrown into vibration, indeed, and into different forms of vibration for waves of different length, but not by any means into the simple sort of vibration demanded by the piano theory. This theory is accordingly too simple, but it probably points the way towards some truer, more complex, conception.

The fact that there are many elementary sensations of hearing is the chief reason why the art of tones is so much more elaborate than the art of color; for while painting might dispute with music as to which were the more highly developed art, painting depends on form as well as color, and there is no art of pure color at all comparable with music, which makes use simply of tones (and noises) with their combinations and sequences.

{236}

Senses of Bodily Movement

It is a remarkable fact that some parts of the inner ear are not connected with hearing at all, but with quite another sense, the existence of which was formerly unsuspected. The two groups of sense cells in the vestibule--the otolith organs--were formerly supposed to be the sense organ for noise; but noise now appears to be a compound of tones, and its organ, therefore, the cochlea. The semicircular canals, from their arrangement in three planes at right angles to each other, were once supposed to analyze the sound according to the direction from which it came; but no one could give anything but the vaguest idea of how they might do this, and besides the ear is now known to give practically no information regarding the direction of sound, except the one fact whether it comes from the right or left, which is given by the difference in the stimulation received by the two ears, and not by anything that exists in either ear taken alone.

The semicircular canals have been much studied by the physiologists. They found that injury to these structures brought lack of equilibrium and inability to walk, swim or fly in a straight course. If, for example, the horizontal canal in the left ear is destroyed, the animal continually deviates to the left as he advances, and so is forced into a "circus movement". They found that the compensatory movements normally made in reaction to a movement impressed on the animal from without were no longer made when the canals were destroyed. They found that something very much like these compensatory movements could be elicited by direct stimulation of the end-organs in the canals or of the sensory nerves leading from them. And they found that little currents of the liquid filling the canals acted as a stimulus to these end-organs and so aroused the {237} compensatory movements. They were thus led to accept a view that was originally suggested by the position of the canals in space.



Fig. 40.--How the sense cells in a semicircular canal are stimulated by a water current. This current is itself an inertia back-flow, resulting from a turning of the head in the opposite direction. (Figure text: water current, nerve to brain)

Each "semicircular" canal, itself considerably more than a semicircular tube, opens into the vestibule at each end and thus amounts to a complete circle. Therefore rotating the head must, by inertia, produce a back flow of the fluid contents of the canal, and this current, by bending the hairs of the sense cells in the canal, would stimulate them and give a sensation of rotation, or at least a sensory nerve impulse excited by the head rotation.

When a human subject is placed, blindfolded, in a chair that can be rotated without sound or jar, it is found that he can easily tell whenever you start to turn him in either direction. If you keep on turning him at a constant speed, he soon ceases to sense the movement, but if then you stop him, he says you are starting to turn him in the opposite {238} direction. He senses the beginning of the rotary movement because this causes the back flow through his canals; he ceases to sense the uniform movement because friction of the liquid in the slender canal soon abolishes the back flow by causing the liquid to move with the canal; and he senses the stopping of this movement because the liquid, again by inertia, continues to move in the direction it had been moving just before when it was keeping pace with the canal. Thus we see that there are conscious sensations of rotation from the canals, and that these give information of the starting or stopping of a rotation, though not of its steady continuance. Excessive stimulation of the canals gives the sensation of dizziness.

The otolith organs in the vestibule are probably excited, not by rotary movements, but by sudden startings and stoppings of rectilinear motion, as in an elevator; and also by the pull of gravity when the head is held in any position. They give information regarding the position and rectilinear movements of the head, as the canals do of rotary head movements. Both are important in maintaining equilibrium and motor efficiency.

The muscle sense is another sense of bodily movement; it was the "sixth sense", so bitterly fought in the middle of the last century by those who maintained that the five senses that were enough for our fathers ought to be enough for us, too. The question was whether the sense of touch did not account for all sensations of bodily movement. It was shown that there must be something besides the skin sense, because weights were better distinguished when "hefted" in the hand than when simply laid in the motionless palm; and it was shown that loss of skin sensation in an arm or leg interfered much less with the coördinated movements of the limb than did the loss of all the sensory nerves to the limb.



Fig. 41.--(From Cajal.) A "tendon spindle," very similar to the muscle spindle spoken of in the text, but found at the tendinous end of a muscle instead of embedded in the muscle substance itself, "a" indicates the tendon, and "e" the muscle fibers; "b" is a sensory axon, and "c" its end-brush about the spindle. Let the tendon become taut in muscular contraction, and the fine branches of the sensory axon will be squeezed and so stimulated.

Later, the crucial fact was established {239} that sense organs (the "muscle spindles") existed in the muscles and were connected with sensory nerve fibers; and that other sense organs existed in the

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