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Scala vestibuli (perilyumh)<br />

Bone<br />

Tectorial<br />

membrane<br />

200Hz<br />

300Hz 100Hz 50Hz<br />

fl~~;t,·, "!<br />

Scala media<br />

(endolymh)<br />

Tectorial<br />

membrane<br />

Basilar membrane<br />

Fig. 9.5. Schematic diagram of<br />

shearing force created between<br />

the hair cells and the tectorial<br />

membrane as a result of basilar<br />

membrane displacement<br />

20<br />

25<br />

30<br />

Fig. 9.6. Theory of von Bekesy:<br />

dependence of the position of<br />

maximum vibration amplitude<br />

on the sound frequency<br />

Scala tympany (perilyumh)<br />

Fig. 9.4. A cross section of the cochlea<br />

section of the cochlea. Essentially the organ contains two fluidfilled<br />

chambers divided by a membrane, called the basilar membrane.<br />

This membrane does not, however, completely separate<br />

the two chambers. A small opening, called the helicotrema, is present<br />

at the distal or innermost part of the system. The basilar<br />

membrane contains the neural detecting celfs, which are small<br />

hair-like cells which are triggered by shear effects occurring<br />

between the fluid and the membrane. Thus, any lateral or<br />

sideways motion of the membrane with respect to the fluid is<br />

detected. The oval window is forced to vibrate by the action of the<br />

ossicles which, in turn, causes an oscillating pressure in the<br />

upper chamber. This pressure can be equilibrated by the membrane<br />

bulging downward. The bulging process produces the shearing<br />

effects which can be detected by the hairs which are located<br />

between the tectorial and basilar membranes (fig. 9.5). The nervous<br />

system then transmits the signal to the brain. According to<br />

von Bekesy's theory, different frequencies will cause a maximum<br />

vibration amplitude at different points along membrane (fig. 9.6).<br />

Birds. Hearing is particularly important when the birds rest or<br />

perch. It also facilitates communication over long distances in<br />

vegetative or arboreal environments where vision may be partially<br />

or completely occluded. Birds (like mammals) have an external<br />

ear and an ear canal which leads to a recessed ear drum. In many<br />

76<br />

birds, the outer opening of the ear canal is surrounded by specialized<br />

feathers in the form of a funnel which act as an efficient<br />

sound collector. The facial ruff of certain owls is shaped either<br />

like a single parabola or as two parabolas side by side, separated<br />

by the beak at the midline. Such a design increases the animal's<br />

acoustic sensitivity. Peak sensitivity for birds occurs at 2 kHz. One<br />

of the most interesting characteristics of a birds' auditory system is<br />

the ability to resolve the small differences that exist in the<br />

frequency of most sounds, called frequency modulation. Birds are<br />

able to detect a frequency change of only 10 - 15 Hz. These slow<br />

or rapid changes in acoustic frequency enable a bird to recognize a<br />

member of its own species. In birds, the ability to determine the<br />

exact location of a sound source depends on a variety of factors.<br />

The barn owl, for instance, is capable of orienting to a sound<br />

source in space with an error of less than 2 degrees.<br />

Fish have a labyrinth, or inner ear which extends their hearing<br />

to higher frequencies. Sound waves travel to chambers where<br />

small granules called otholits are located. These granules stimulate<br />

sensory hair cells and trigger action potentials in the neurons of<br />

the auditory nerve.<br />

The gas bladders of some fish also provide an area of variable<br />

density. Vibrations pass through the gas bladder and travel<br />

through a pathway of small bones called Weber ossicles. These<br />

serve to connect the gas bladder directly with the inner ear<br />

of the fish.<br />

77

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