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Essential Cell Biology 5th edition

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Ion Channels and the Membrane Potential

409

entry of

positively

charged ions

supporting cells

auditory

hair cells

tectorial membrane

CHANNEL

CLOSED

linking

filament

CHANNEL

OPEN

stereocilia

(A)

basilar membrane

auditory nerve fibers

STEREOCILIA NOT

TILTED

(B)

STEREOCILIA

TILTED

Figure 12–28 Mechanically-gated ion channels allow us to hear. (A) A section through the organ of Corti,

which runs the length of the cochlea, the auditory portion of the inner ear. Each auditory hair cell has a tuft of

spiky extensions called stereocilia projecting from its upper surface. The hair cells are embedded in an epithelial

sheet of supporting cells, which is sandwiched between the basilar membrane below and the tectorial membrane

above. (These are not lipid bilayer membranes but sheets of extracellular matrix.) (B) Sound vibrations cause the

basilar membrane to vibrate up ECB5 and down, E12.27/12.28

causing the stereocilia to tilt. Each stereocilium in the staggered array

of stereocilia on a hair cell is attached to the next, shorter stereocilium by a fine filament. The tilting stretches

the filaments, which pull open mechanically-gated ion channels in the stereocilium plasma membrane, allowing

positively charged ions to enter from the surrounding fluid (Movie 12.7). The influx of ions activates the hair cells,

which stimulate underlying nerve endings of the auditory nerve fibers that relay the auditory signal to the brain.

The hair-cell mechanism is astonishingly sensitive: the faintest sounds we can hear have been estimated to stretch

the filaments by an average of about 0.04 nm, which is less than the diameter of a hydrogen ion (Movie 12.8).

Voltage-gated Ion Channels Respond to the Membrane

Potential

Voltage-gated ion channels play a major role in propagating electrical

signals along all nerve cell extensions, such as those that relay signals

from our brain to our toe muscles. But voltage-gated ion channels are

present in many other cell types, too, including muscle cells, egg cells,

protozoans, and even plant cells, where they enable electrical signals

to travel from one part of the plant to another, as in the leaf-closing

response of a Mimosa pudica plant (Figure 12–29).

Voltage-gated ion channels have domains called voltage sensors that

are extremely sensitive to changes in the membrane potential: changes

(A) time 0 sec (B) 1 sec (C) 3 sec (D) 5 sec

Figure 12–29 Both mechanically-gated and voltage-gated ion channels underlie the leaf-closing response in the touch-sensitive

plant Mimosa pudica. (A) Resting leaf. (B–D) Successive leaflet closures in response to touch. A few seconds after the leaf on the left is

touched, its leaflets snap shut. The response involves the opening of mechanically-gated ion channels in touch-sensitive sensory cells,

which then pass a signal to cells containing voltage-gated ion channels, generating an electric impulse. When the impulse reaches

specialized hinge cells at the base of each leaflet, a ECB5 rapid loss E12.28/12.29

of water by these cells occurs, causing the leaflets to fold into a closed

conformation suddenly and progressively down the leaf stalk (Movie 12.9).

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