Page:Encyclopædia Britannica, Ninth Edition, v. 7.djvu/614

592 592 EAR adapted for low sounds. In addition, increased tension of the membrane, by increasing the resistance, will diminish the intensity of vibrations. This is especially the case for sounds of low pitch. Helmholtz has also pointed out that the peculiar form of the membrana tympani in man has the effect of increasing the force of its vibrations at the expense of their amplitude. The vibrations of the membrana tympani are transmitted to the internal ear partly by the air which the middle ear or tympanum contains, and partly by the chain of bones, consisting of the malleus, incus, and stapes. Of these, transmission by the chain of bones is by far the most important. In birds and in the scaly amphibia, this chain is represented by a single rod-like ossicle, the columella, but in man the two membranes the membrana tympani and the membrane filling the fenestra ovalis are connected by a compound lever consisting of three bones, namely, the malleus, or hammer, inserted into the membrana tympani, the incus, or anvil, and the stapes, or stirrup, the base of which fits into the oval window. The lever thus formed has its fulcrum near the short process of the incus, which abuts against the tympanic wall ; the power is applied at the handle of the malleus, and the resistance is at the base of the stirrup. Both by direct experimental observation and by calculation from data supplied by measurement of the lengths of the arms of the lever, Helmholtz has shown that by this arrangement vibrations are diminished in extent in the ratio of 3 to 2, but are inversely increased in force. Considering the great resistance offered to excur sions of the stapes, such an arrangement must be advantageous. It must also be noted that in the transmis sion of vibrations of the membrana tympani to the fluid in the labyrinth or internal ear, through the oval window, the chain of ossicles vibrates as a whole and acts efficiently, although its length may be only a small fraction of the wave length of the sound transmitted. 3. Transmission in the Internal Ear. The internal ear is composed of the labyrinth, formed of the vestibule or central part, the semicircular canals, and the cochlea, each of which consists of an osseous and a membranous portion (see vol. i. p. 893). The osseous labyrinth may be regarded as an osseous mould in the petrous portion of the temporal bone, lined by tesselated endothelium, and con taining a small quantity of fluid called the perilymph. In this mould, partially surrounded by, and to some extent floating in, this fluid, there is the membranous labyrinth, in certain parts of which we find the terminal apparatus in connection with the auditory nerve, immersed in another fluid called the endolymph. The membranous labyrinth consists of a vestibular portion formed by two small sac- like dilatations, called the saccule and the utricle, the latter of which communicates with the semicircular canals by five openings. Each canal consists of a tube, bulging out at each extremity so as to form the so-called ampulla, in which, on a projecting ridge, called the crista acoustica, there are cells bearing or developed into long auditory hairs, which are to be regarded as the peripheral end-organs of the vestibular branches of the auditory nerve. The cochlear division of the membranous labyrinth consists of the dudus cochlearis, a tube of triangular form fitting in between the two cavities in the cochlea, called the scala vestibuli, because it commences in the vestibule, and the scala tympani, because it ends in the tympanum, at the round window. These two scalse communicate at the apex of the cochlea. The roof of the ductus cochlearis is formed by a thin membrane called the membrane of Reissner, while its floor consists of the basilar membrane, on which we find the remarkable organ of Corti, which constitutes the terminal organ of the cochlear division of the auditory nerve, and which is fully described in vol. i. p. 891. It is sufficient to state here that this organ consists essentially of an arrangement of epithelial cells bearing hairs which are in communication with the terminal filaments of this portion of the auditory nerve, and that groups of these hairs pass through holes in a closely investing mem brane, membrana reticidaris, which may be supposed to act as a damping apparatus, so as quickly to stop their movements. The ductus cochlearis and the two scake are filled with fluid. Sonorous vibrations may reach the fluid in the labyrinth by three different ways (1) by the osseous walls of the labyrinth, (2) by the air in the tym panum and the round window, and (3) by the base of the stapes inserted into the oval window. When the head is plunged into water, or brought into direct contact with any vibrating body, vibrations must be transmitted directly. Vibrations of the air in the mouth and in the nasal passages are also communicated directly to the walls of the cranium, and thus pass to the labyrinth. In like manner, we may experience peculiar auditive sensations, such as blowing, rubbing, and hissing sounds, due to muscular contraction or to the passage of blood in vessels close to the auditory organ. It has not been satisfactorily made out to what extent, if any, vibrations may be communicated to the fluid in the labyrinth by the round window. There can be no doubt, however, that ia ordinary hearing vibrations are communicated chiefly by the chain of bones. When the base of the stirrup is pushed into the oval window, the pressure in the labyrinth in creases, the impulse passes along the scala vestibuli to the scala tympani, and, as the only mobile part of the wall of the labyrinth is the membrane covering the round window, this membrane is forced outwards ; when the base of the stirrup passes outwards, a reverse action takes place. Thus the fluid of the labyrinth may receive a series of pulses or vibrations isochronous with the movements of the base of the stirrup, and these pulses affect the terminal apparatus in connection with the auditory nerve. Since the size of the membranous labyrinth is so small, measuring, in man, not more than ^ inch in length by chamber consisting partly of conduits of very irregular form, it is impossible to state accurately the course of vibrations transmitted to it by impulses communicated from the base of the stirrup. In the cochlea, vibrations must pass from the saccule along the scala vestibuli to tbe apex, thus affecting the membrane of Reissner, which forms its roof; then, passing through the opening at the apex (the helicotrema), they must descend by the scala tympani to the round window, and affect in their passage the membrana basilaris, on which the organ of Corti is situated. From the round window impulses must be reflected backwards, but how they affect the advancing impulses is not known. But the problem is even more complex when we take into account the fact that impulses are transmitted simultane ously to the utricle and to the semicircular canals com municating with it by five openings. The mode of action of these vibrations or impulses upon the nervous termina tions is still unknown ; but to appreciate critically the hypothesis which has been advanced to explain it, it is necessary, in the first place, to refer to some of the general characters of auditory sensation. 4. Certain conditions are necessary for excitation of the auditory nerve sufficient to produce a sensation. la the first place, the vibrations must have a certain amplitude : if too feeble, no impression will be produced. The minimum limit has been stated to be the sensation caused by the falling of a ball of pith, 1 millegramme in weight, upon a smooth surface, such as glass, from a height of 1 millimetre at a distance of 91 millimetres from the ear.
 * th inch in diameter at its widest part, and since it is a