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Essential Cell Biology 5th edition

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704 CHAPTER 20 Cell Communities: Tissues, Stem Cells, and Cancer

LUMEN

tracer

molecule

interacting

plasma membranes

CELL

1

CELL

2

CELL

3

tight

junction

tight

junction

intercellular

space

(A)

(B)

CELL

1

CELL

2

CELL

1

CELL

2

0.5 µm 0.5 µm

sealing

strands of

occludin and

claudin

proteins

Figure 20–22 Tight junctions allow epithelial cell sheets to serve as barriers to

solute diffusion. (A) Schematic drawing showing how a small, extracellular tracer

molecule (yellow) added on one side of an epithelial cell sheet cannot traverse the

tight junctions that seal adjacent cells together. (B) Electron micrographs of cells in an

epithelium where a small, extracellular tracer molecule (dark stain) has been added to

either the apical side (on the left) or the basolateral side (on the right); in both cases,

the tracer is stopped by the tight junction. (C) A simplified model of the structure

of a tight junction, showing how the cells are sealed together by branching strands

of transmembrane proteins (green), called claudins and occludins, in the plasma

membranes of the interacting cells. Each type of protein binds to the same type in

the apposed membrane (not shown). (B, courtesy of Daniel Friend, by permission of

E.L. Bearer.)

(C)

CELL

1

CELL

2

0.3 µm

that water-soluble molecules cannot easily leak between them. If a small

tracer molecule is added to one side of an epithelial cell sheet, it will usually

not pass beyond the tight junction (Figure 20–22A and B). The tight

junction is formed from proteins called claudins and occludins, which

ECB5 e20.23/20.23

are arranged in strands along the lines of the junction to create the seal

(Figure 20–22C). Without tight junctions to prevent leakage, the pumping

activities of absorptive cells such as those in the gut would be futile,

and the composition of the extracellular fluid would become the same on

both sides of the epithelium.

Tight junctions also play a key part in maintaining the polarity of the

individual epithelial cells in two ways. First, the tight junctions around

the apical region of each cell prevent diffusion of proteins in the plasma

membrane and so keep the contents of apical domain of the plasma

membrane separate—and different—from the basolateral domain (see

Figure 11−32). Second, in many epithelia, the tight junctions are sites

of assembly for the complexes of intracellular proteins that govern the

apico-basal polarity of the cell interior.

Cytoskeleton-linked Junctions Bind Epithelial Cells

Robustly to One Another and to the Basal Lamina

The cell junctions that hold an epithelium together by forming strong

mechanical attachments are of three main types. Adherens junctions and

desmosomes bind one epithelial cell to another, while hemidesmosomes

bind epithelial cells to the basal lamina. All of these junctions provide

mechanical strength to the epithelium by the same strategy: the proteins

that form the junctions span the plasma membrane and are linked inside

the cell to cytoskeletal filaments. In this way, the cytoskeletal filaments

are tied into a network that extends from cell to cell across the whole

expanse of the epithelial sheet.

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