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Molecular Biology of the Cell by Bruce Alberts, Alexander Johnson, Julian Lewis, David Morgan, Martin Raff, Keith Roberts, Peter Walter by by Bruce Alberts, Alexander Johnson, Julian Lewis, David Morg

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600 Chapter 11: Membrane Transport of Small Molecules and the Electrical Properties of Membranes

lipid

bilayer

(A)

simple

diffusion

OUTSIDE

channelmediated

transportermediated

PASSIVE TRANSPORT

+ + + +

+ + + +

ENERGY

ACTIVE TRANSPORT

+ + + +

+ + + +

concentration

gradient

Figure 11–4 Different forms of

membrane transport and the influence

of the membrane. Passive transport

down a concentration gradient (or an

electrochemical gradient—see B below)

occurs spontaneously, by diffusion, either

through the lipid bilayer directly or through

channels or passive transporters. By

contrast, active transport requires an input

of metabolic energy and is always mediated

by transporters that pump the solute

against its concentration or electrochemical

gradient. (B) The electrochemical gradient

of a charged solute (an ion) affects its

transport. This gradient combines the

membrane potential and the concentration

gradient of the solute. The electrical and

chemical gradients can work additively to

increase the driving force on an ion across

the membrane (middle) or can work against

each other (right).

INSIDE

– – – –

– – – –

– – – –

– – – –

(B)

concentration

gradient (with no

membrane potential)

electrochemical gradient with

a membrane potential

As shown in Figure 11–4A, in addition to passive transport, cells need to be

able to actively pump certain solutes across the membrane “uphill,” against their

electrochemical gradients. Such active transport is mediated by transporters

whose pumping activity is directional because it is tightly coupled to a source of

metabolic energy, such as an ion gradient or ATP hydrolysis, as discussed later.

Transmembrane movement of small molecules mediated by transporters can be

either active or passive, whereas that mediated by channels is always passive (see

Figure 11–4A).

MBoC6 m11.04/11.04

Summary

Lipid bilayers are virtually impermeable to most polar molecules. To transport

small water-soluble molecules into or out of cells or intracellular membrane-enclosed

compartments, cell membranes contain various membrane transport proteins,

each of which is responsible for transferring a particular solute or class of

solutes across the membrane. There are two classes of membrane transport proteins—transporters

and channels. Both form protein pathways across the lipid

bilayer. Whereas transmembrane movement mediated by transporters can be

either active or passive, solute flow through channel proteins is always passive. Both

active and passive ion transport is influenced by the ion’s concentration gradient

and the membrane potential—that is, its electrochemical gradient.

TRANSPORTERS AND ACTIVE MEMBRANE

TRANSPORT

The process by which a transporter transfers a solute molecule across the lipid

bilayer resembles an enzyme–substrate reaction, and in many ways transporters

behave like enzymes. By contrast to ordinary enzyme–substrate reactions, however,

the transporter does not modify the transported solute but instead delivers it

unchanged to the other side of the membrane.

Each type of transporter has one or more specific binding sites for its solute

(substrate). It transfers the solute across the lipid bilayer by undergoing reversible

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