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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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1044 Chapter 19: Cell Junctions and the Extracellular Matrix

(A)

folded

α-catenin

NO TENSION

cadherin

plasma membrane

CYTOSOL

(B)

CYTOSOL

extended

α-catenin

TENSION

attached cell

pulls on cadherin

Figure 19–12 Mechanotransduction

in an adherens junction. (A) Cell–cell

junctions are able to sense increased

tension and respond by strengthening

their actin linkages. Tension sensing is

thought to depend in part on α-catenin

(see Figure 19–10). (B) When actin

filaments are pulled from within the cell by

non-muscle myosin II, the resulting force

unfolds a domain in α-catenin, thereby

exposing an otherwise hidden binding site

for the adaptor protein vinculin. Vinculin

then promotes additional actin recruitment,

strengthening the linkages between the

junction and the cytoskeleton.

vinculin

actin

filament

myosin II pulls on actin

actin filaments beneath the plasma membranes of two interacting cells. In heart

muscle, they anchor the actin bundles of the contractile apparatus and act in parallel

with desmosomes to link the contractile cells end-to-end. But the prototypical

examples of adherens junctions occur in epithelia, where they often form a

continuous adhesion belt (or zonula adherens) just beneath the apical face of the

epithelium, encircling each of the interacting MBoC6 n19.202/19.12 cells in the sheet (Figure 19–13).

Within each cell, a contractile bundle of actin filaments and myosin II lies adjacent

to the adhesion belt, oriented parallel to the plasma membrane and tethered to

actin filaments

inside microvillus

LUMEN

microvilli extending

from apical surface

adhesion

belt

cadherins

tight junction

bundle of

actin filaments

lateral plasma

membranes of

adjacent

epithelial cells

basal surface

Figure 19–13 Adherens junctions

between epithelial cells in the small

intestine. These cells are specialized

for absorption of nutrients; at their apex,

facing the lumen of the gut, they have

many microvilli (protrusions that increase

the absorptive surface area). The adherens

junction takes the form of an adhesion

belt, encircling each of the interacting cells.

Its most obvious feature is a contractile

bundle of actin filaments running along

the cytoplasmic surface of the junctional

plasma membrane. The actin filament

bundles are tethered by intracellular

proteins to cadherins, which bind to

cadherins on the adjacent cell. In this way,

the actin filament bundles in adjacent cells

are tied together. For clarity, this drawing

does not show most of the other cell–cell

and cell–matrix junctions of epithelial cells

(see Figure 19–2).

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