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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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641

Intracellular Compartments

and Protein Sorting

chapter

12

Unlike a bacterium, which generally consists of a single intracellular compartment

surrounded by a plasma membrane, a eukaryotic cell is elaborately subdivided

into functionally distinct, membrane-enclosed compartments. Each

compartment, or organelle, contains its own characteristic set of enzymes and

other specialized molecules, and complex distribution systems transport specific

products from one compartment to another. To understand the eukaryotic cell, it

is essential to know how the cell creates and maintains these compartments, what

occurs in each of them, and how molecules move between them.

Proteins confer upon each compartment its characteristic structural and

functional properties. They catalyze the reactions that occur there and selectively

transport small molecules into and out of the compartment. For membrane-enclosed

organelles in the cytoplasm, proteins also serve as organelle-specific surface

markers that direct new deliveries of proteins and lipids to the appropriate

organelle.

An animal cell contains about 10 billion (10 10 ) protein molecules of perhaps

10,000 kinds, and the synthesis of almost all of them begins in the cytosol, the

space of the cytoplasm outside the membrane-enclosed organelles. Each newly

synthesized protein is then delivered specifically to the organelle that requires it.

The intracellular transport of proteins is the central theme of both this chapter

and the next. By tracing the protein traffic from one compartment to another, one

can begin to make sense of the otherwise bewildering maze of intracellular membranes.

In This Chapter

The Compartmentalization

of Cells

THE TRANSPORT OF

MOLECULES BETWEEN THE

NUCLEUS AND THE CYTOSOL

THE TRANSPORT OF PROTEINS

INTO MITOCHONDRIA AND

CHLOROPLASTS

PEROXISOMES

THE ENDOPLASMIC RETICULUM

The Compartmentalization of Cells

In this brief overview of the compartments of the cell and the relationships

between them, we organize the organelles conceptually into a small number of

discrete families, discuss how proteins are directed to specific organelles, and

explain how proteins cross organelle membranes.

All Eukaryotic Cells Have the Same Basic Set of Membraneenclosed

Organelles

Many vital biochemical processes take place in membranes or on their surfaces.

Membrane-bound enzymes, for example, catalyze lipid metabolism; and oxidative

phosphorylation and photosynthesis both require a membrane to couple the

transport of H + to the synthesis of ATP. In addition to providing increased membrane

area to host biochemical reactions, intracellular membrane systems form

enclosed compartments that are separate from the cytosol, thus creating functionally

specialized aqueous spaces within the cell. In these spaces, subsets of molecules

(proteins, reactants, ions) are concentrated to optimize the biochemical

reactions in which they participate. Because the lipid bilayer of cell membranes is

impermeable to most hydrophilic molecules, the membrane of an organelle must

contain membrane transport proteins to import and export specific metabolites.

Each organelle membrane must also have a mechanism for importing, and incorporating

into the organelle, the specific proteins that make the organelle unique.

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