In the auditory canal waves of sound are reflected from side to side until they reach the membrana tympani. From the obliquity in position and peculiar curvature of this membrane, most of the waves strike it nearly perpendicularly, and in the most advantageous direction.
2. Transmission in Middle Ear.—The middle ear is a small cavity, the walls of which are rigid with the exception of the portions consisting of the membrana tympani, and the membrane of the round window and of the apparatus filling the oval window. This cavity communicates with the pharynx by the Eustachian tube, which forms an air-tube between the pharynx and the tympanum for the purpose of regulating pressure on the membrana tympani. During rest the tube is open, but it is closed during the act of deglutition. As this action is frequently taking place, not only when food or drink is introduced, but when saliva is swallowed, it is evident that the pressure of the air in the tympanum will be kept in a state of equilibrium with that of the external air on the outer surface of the membrana tympani, and that thus the membrana tympani will be rendered independent of variations of atmospheric pressure such as occur when we descend in a diving bell or ascend in a balloon. By a forcible expiration, the oral and nasal cavities being closed, air may be driven into the tympanum, while a forcible inspiration (Valsalva’s experiment) will draw air from that cavity. In the first case, the membrana tympani will bulge outwards, in the second case inwards, and in both, from excessive stretching of the membrane, there will be partial deafness, especially for sounds of high pitch. Permanent occlusion of the tube is one of the most common causes of deafness.
The membrana tympani is capable of being set into vibration by a sound of any pitch included in the range of perceptible sounds. It responds exactly as to number of vibrations (pitch), intensity of vibrations (intensity), and complexity of vibration (quality or timbre). Consequently we can hear a sound of any given pitch, of a certain intensity, and in its own specific timbre or quality. Generally speaking, very high tones are heard more easily than low tones of the same intensity. As the membrana tympani is not only fixed by its margin to a ring or tube of bone, but is also adherent to the handle of the malleus, which follows its movements, its vibrations meet with considerable resistance. This diminishes the intensity of its vibrations, and prevents also the continued vibration of the membrane after an external pressure has ceased, so that a sound is not heard much longer than its physical cause lasts. The tension of the membrane may be affected (1) by differences of pressure on the two surfaces of the membrana tympani, as may occur during forcible expiration or inspiration, and (2) by muscular action, due to contraction of the tensor tympani muscle. This small muscle arises from the apex of the petrous temporal and the cartilage of the Eustachian tube, enters the tympanum at its anterior wall, and is inserted into the malleus near its root. The handle of the malleus is inserted between the layers of the membrana tympani, and, as the malleus and incus move round an axis passing through the neck of the malleus from before backwards, the action of the muscle is to pull the membrana tympani inwards towards the tympanic cavity in the form of a cone, the meridians of which are not straight but curved, with convexity outwards. When the muscle contracts, the handle of the malleus is drawn still farther inwards, and thus a greater tension of the tympanic membrane is produced. On relaxation of the muscle, the membrane returns to its position of equilibrium by its elasticity and by the elasticity of the chain of bones. This power of varying the tension of the membrane is an accommodating mechanism for receiving and transmitting sounds of different pitch. With different degrees of tension it will respond more readily to sounds of different pitch. Thus, when the membrane is tense, it will readily respond to high sounds, while relaxation will be the condition most adapted for low tones. 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.
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 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 is attached to a membrane covering the oval window. It must also be noted that in the transmission 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 fraction of the wave-length of the sound transmitted. The chain is a lever in which the handle of the malleus forms the long arm, the fulcrum is where the short process of the incus abuts against the wall of the tympanum, while the long process of the incus, carrying the stapes, forms the short arm. The mechanism is a lever of the second order. Measurements show that the ratio of the lengths of the two arms is as 1.5:1; the ratio of the resulting force at the stapes is therefore as 1:1.5; while the amplitudes of the movements at the tip of the handle of the malleus and the stapes is as 1.5:1. Hence, while there is a diminution in amplitude there is a gain in power, and thus the pressures are conveyed with great efficiency from the membrana tympani to the labyrinth, while the amplitude of the oscillation is diminished so as to be adapted to the small capacity of the labyrinth. As the drum-head is nearly twenty times greater in area than the membrane covering the oval window, with which the base of the stapes is connected, the energy of the movements of the membrana tympani is concentrated on an area twenty times smaller; hence the pressure is increased thirtyfold (1.5×20) when it acts at the base of the stapes. Experiments on the human ear have shown that the movement of greatest amplitude was at the tip of the handle of the malleus, 0.76 mm.; the movement of the tip of the long arm process of the incus was 0.21 mm.; while the greatest amplitude at the base of the stapes was only .0714 mm. Other observations have shown the movements at the stapes to have a still smaller amplitude, varying from 0.001 to 0.032 mm. With tones of feeble intensity the movements must be almost infinitesimal. There may also be very minute transverse movements at the base of the stapes.
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. The osseous labyrinth may be regarded as an osseous mould in the petrous portion of the temporal bone, lined by tesselated endothelium, and containing 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 connexion 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 acustica, there are cells bearing long auditory hairs, which are the peripheral end-organs of the vestibular branches of the auditory nerve. The cochlear division of the membranous labyrinth consists of the ductus 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 scalae 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
