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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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II

I

1035

PART

III IV V

Cells in Their Social Context

Cell Junctions and the

Extracellular Matrix

chapter

19

Of all the social interactions between cells in a multicellular organism, the most

fundamental are those that hold the cells together. Cells may be linked by direct

interactions, or they may be held together within the extracellular matrix, a complex

network of proteins and polysaccharide chains that the cells secrete. By one

means or another, cells must cohere if they are to form an organized multicellular

structure that can withstand and respond to the various external forces that try to

pull it apart.

The mechanisms of cohesion govern the architecture of the body—its shape,

its strength, and the arrangement of its different cell types. The making and breaking

of the attachments between cells and the modeling of the extracellular matrix

govern the way cells move within the organism, guiding them as the body grows,

develops, and repairs itself. Attachments to other cells and to extracellular matrix

control the orientation and behavior of the cell’s cytoskeleton, thereby allowing

cells to sense and respond to changes in the mechanical features of their environment.

Thus, the apparatus of cell junctions and the extracellular matrix is critical

for every aspect of the organization, function, and dynamics of multicellular

structures. Defects in this apparatus underlie an enormous variety of diseases.

The key features of cell junctions and the extracellular matrix are best illustrated

by considering two broad categories of tissues that are found in all animals

(Figure 19–1). Connective tissues, such as bone or tendon, are formed from an

extracellular matrix produced by cells that are distributed sparsely in the matrix.

It is the matrix—rather than the cells—that bears most of the mechanical stress to

which the tissue is subjected. Direct attachments between one cell and another

are relatively rare, but the cells have important attachments to the matrix. These

cell–matrix junctions link the cytoskeleton to the matrix, allowing the cells to move

through the matrix and monitor changes in its mechanical properties.

In epithelial tissues, such as the lining of the gut or the epidermal covering of

the skin, cells are tightly bound together into sheets called epithelia. The extracellular

matrix is less pronounced, consisting mainly of a thin mat called the basal

lamina (or basement membrane) underlying the sheet. Within the epithelium,

cells are attached to each other directly by cell–cell junctions, where cytoskeletal

filaments are anchored, transmitting stresses across the interiors of the cells, from

In This Chapter

CELL–CELL JUNCTIONS

THE EXTRACELLULAR MATRIX

OF ANIMALS

CELL–MATRIX JUNCTIONS

THE PLANT CELL WALL

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