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Linguistics Encyclopedia.pdf

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The linguistics encyclopedia 58<br />

The outer ear includes the pinna and the external meatus—the visible cartilaginous<br />

structures—and the external auditory canal which terminates at the tympanic<br />

membrane or eardrum. The outer ear ‘collects’ auditory signals which arrive as sound<br />

waves or changing acoustic pressures propagated through the surrounding medium,<br />

usually air. The outer ear also serves as protection for the delicate middle ear, provides<br />

some amplification and assists in sound localization, i.e. in determining where a sound<br />

originates.<br />

The middle ear is bounded on one side by the tympanic membrane and on the other<br />

by a bony wall containing the cochlea of the inner ear. In addition to the tympanic<br />

membrane, the middle ear contains three ossicles; these are the malleus, incus, and<br />

stapes, a set of three tiny interconnected bones extending in a chain from the tympanic<br />

membrane to the oval window of the cochlea. The tympanic membrane vibrates in<br />

response to the sound waves impinging upon it; the ossicles greatly amplify these<br />

vibratory patterns by transferring pressure from a greater area, the tympanic membrane,<br />

to a much smaller one, the footplate of the stapes attached to the oval window of the<br />

cochlea.<br />

The inner ear contains the vestibule, the semicircular canals—which primarily<br />

affect balance—and the cochlea, a small coiled passage of decreasing diameter. Running<br />

the length of the cochlea are the scala tympani and scala vestibuli, two fluid-filled<br />

canals which are separated from the fluid-filled scala media or cochlear duct. The<br />

vibratory patterns of sound-pressure waves are transferred into hydraulic pressure waves<br />

which travel through the scala vestibuli and scala tympani and from the base to the apex<br />

of the scala media.<br />

One surface of the scala media contains a layer of fibres called the basilar<br />

membrane. This tapered membrane is narrow and taut at its base in the larger vestibular<br />

end of the cochlea, and wide and flaccid at its terminus or apex in the smaller apical<br />

portion of the cochlea. On one surface of the basilar membrane is the organ of Corti<br />

which contains thousands of inner and outer hair cells, each supporting a number of cilia<br />

or hairs. When the basilar membrane is displaced in response to the travelling waves<br />

propagating throughout it, the tectorial membrane near the outer edge of the organ of<br />

Corti also moves. It is believed that the shearing effect of the motion of these two<br />

membranes stimulates the cilia of the hair cells, thereby triggering a neural response in<br />

the auditory-receptor cells. These cells, in turn, relay electrochemical impulses to a fibre<br />

bundle called the auditory nerve, or the VIIIth cranial nerve. Information about the<br />

spatial representation of frequencies on the basilar membrane is preserved in the auditory<br />

nerve, which is thus said to have tonotopic organization.<br />

The precise nature of the information received on the basilar membrane and encoded<br />

in the auditory nerve has been a matter of much investigation. The fact that the basilar<br />

membrane changes in width and rigidity throughout its length means that the amplitudes<br />

of pressure waves peak at specific loci or places on the membrane. Hence, the peak<br />

amplitudes of low-frequency sounds occur at the wider and more flaccid apex while the<br />

peak amplitudes of high-frequency sounds occur at the narrower and tauter base, which<br />

can, however, also respond to low-frequency stimulation. This was demonstrated in a<br />

series of experiments conducted by von Békésy in the 1930s and 1940s (see von Békésy,<br />

1960).

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