46 Chapter 3
the oval window and the round window. The auditory
portion of the inner ear is the snail-shaped cochlea. It is
a mechanical-to-electrical transducer and a fre-
quency-selective analyzer, sending coded nerve
impulses to the brain. This is represented crudely in Fig.
3-5. A rough sketch of the cross section of the cochlea
is shown in Fig. 3-6. The cochlea, throughout its length
(about 35 mm if stretched out straight), is divided by
Reissner’s membrane and the basilar membrane into
three separate compartments—namely, the scala ves-
tibuli, the scala media, and the scala tympani. The scala
vestibuli and the scala tympani share the same fluid,
perilymph, through a small hole, the helicotrema, at the
apex; while the scala media contains another fluid,
endolymph, which contains higher density of potassium
ions facilitating the function of the hair cells. The basi-
lar membrane supports the Organ of Corti, which con-
tains the hair cells that convert the relative motion
between the basilar membrane and the tectorial mem-
brane into nerve pulses to the auditory nerve.
When an incident sound arrives at the inner ear, the
vibration of the stapes is transported into the scala
vestibuli through the oval window. Because the cochlear
fluid is incompressible, the round window connected to
the scala tympani vibrates accordingly. Thus, the vibra-
tion starts from the base of the cochlea, travels along the
scala vestimbuli, all the way to the apex, and then
through the helicotrema into the scala tympani, back to
the base, and eventually ends at the round window. This
establishes a traveling wave on the basilar membrane
for frequency analysis. Each location at the basilar
membrane is most sensitive to a particular
frequency—i.e., the characteristic frequency—although
it also responds to a relatively broad frequency band at
smaller amplitude. The basilar membrane is narrower
(0.04 mm) and stiffer near the base, and wider (0.5 mm)
and looser near the apex. (By contrast, when observed
from outside, the cochlea is wider at the base and
smaller at the apex.) Therefore, the characteristic
frequency decreases gradually and monotonically from
the base to the apex, as indicated in Fig. 3-5. The trav-
eling-wave phenomenon illustrated in Figs. 3-7 and 3-8
shows the vibration patterns—i.e., amplitude versus
location—for incident pure tones of different frequen-
cies. An interesting point in Fig. 3-8 is that the vibration
pattern is asymmetric, with a slow tail close to the base
(for high frequencies) and a steep edge close to the apex
(for low frequencies). Because of this asymmetry, it is
easier for the low frequencies to mask the high frequen-
cies than vice versa.
Within the Organ of Corti on the basilar membrane,
there are a row of inner hair cells (IHC), and three to
five rows of outer hair cells (OHC), depending on loca-
tion. There are about 1500 IHCs and about 3500 OHCs.
Each hair cell contains stereociliae (hairs) that vibrate
corresponding to the mechanical vibration in the fluid
around them. Because each location on the basilar
membrane is most sensitive to its own characteristic
frequency, the hair cells at the location also respond
most to its characteristic frequency. The IHCs are
sensory cells, like microphones, which convert mechan-
ical vibration into electrical signal—i.e., neural firings.
The OHCs, on the other hand, change their shapes
according to the control signal received from efferent
nerves. Their function is to give an extra gain or attenua-
tion, so that the output of the IHC is tuned to the
characteristic frequency much more sharply than the
IHC itself. Fig. 3-9 shows the tuning curve (output level
vs. frequency) for a particular location on the basilar
membrane with and without functioning OHCs. The
tuning curve is much broader with poor frequency selec-
tivity when the OHCs do not function. The OHCs make
our auditory system an active device, instead of a
passive microphone. Because the OHCs are active and
Figure 3-5. The mechanical system of the middle ear.
Figure 3-6. Cross-sectional sketch of the cochlea.
Eardrum
80 mm^2
Obstacles
Ratio
1.3 to3:1
3 mm^2
window
Cochlea Basilar
membrane
Brain
10 kHz4 kHz 1 kHz100 Hz
Auditory
nerve
Reissner's
membrane
Tectorial
membrane
Outer
hair
cells
Basilar
membrane
Inner
hair cells
Scala
vestibuli
Scala tympani
Scala
media