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Essential Cell Biology 5th edition

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408 CHAPTER 12 Transport Across Cell Membranes

Figure 12–27 Different types of gated

ion channels respond to different types

of stimuli. Depending on the type of

channel, the probability of gate opening is

controlled by (A) a change in the voltage

difference across the membrane,

(B) the binding of a chemical ligand to

the extracellular face of a channel,

(C) ligand binding to the intracellular face

of a channel, or (D) mechanical stress. In

the case of the voltage-gated channels,

positively charged amino acids (white

plus signs) in the channel’s voltage sensor

domains become attracted to negative

charges on the extracellular surface of the

depolarized plasma membrane, pulling

the channel into its open conformation.

CLOSED

resting

plasma

membrane

OPEN

depolarized

plasma

membrane

+ ++ ++ +

+ +

CYTOSOL

–– –

– – –

+ +

+ ++ ++ +

CYTOSOL

(extracellular

ligand)

(intracellular

ligand)

(A) voltagegated

(B) ligand-gated (C) ligand-gated (D)

mechanicallygated

indicates that the channel has moving parts and is snapping back and

forth between open and closed conformations as the channel is knocked

from one conformation to the other by the random thermal movements

ECB5 e12.26/12.27

of the molecules in its environment. Patch-clamp recording was the

first technique that could detect such conformational changes, and the

picture it paints—of a jerky piece of machinery subjected to constant

external buffeting—is now known to apply also to other proteins with

moving parts.

The activity of each ion channel is very much “all-or-none”: when an ion

channel is open, it is fully open; when it is closed, it is fully closed. That

raises a fundamental question: If ion channels randomly snap between

open and closed conformations even when conditions on each side of

the membrane are held constant, how can their state be regulated by

conditions inside or outside the cell? The answer is that when the appropriate

conditions change, the random behavior continues but with a

greatly changed bias: if the altered conditions tend to open the channel,

for example, the channel will now spend a much greater proportion of its

time in the open conformation, although it will not remain open continuously

(see Figure 12–26).

Different Types of Stimuli Influence the Opening and

Closing of Ion Channels

There are more than a hundred types of ion channels, and even simple

organisms can possess many different types. The human genome contains

80 genes that encode different but related K + channels alone. Ion

channels differ from one another primarily with respect to their ion selectivity—the

type of ions they allow to pass—and their gating—the conditions

that influence their opening and closing. For a voltage-gated channel,

the probability of being open is controlled by the membrane potential

(Figure 12–27A). For a ligand-gated channel, opening is controlled by

the binding of some molecule (a ligand) to the channel (Figure 12–27B

and C). For a mechanically-gated channel, opening is controlled by a

mechanical force applied to the channel (Figure 12–27D).

The auditory hair cells in the ear are an important example of cells that

depend on mechanically-gated channels. Sound vibrations pull the channels

open, causing ions to flow into the hair cells; this ion flow sets up

an electrical signal that is transmitted from the hair cell to the auditory

nerve, which then conveys the signal to the brain (Figure 12–28).

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