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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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1054 Chapter 19: Cell Junctions and the Extracellular Matrix

cytoplasm

smooth

endoplasmic

reticulum

desmotubule

plasma

membrane

endoplasmic

reticulum

cytosol

cell walls

of adjacent

plant cells

desmotubule

cell

wall

plasmodesmata

(A)

plasma membrane lining

plasmodesma, connecting

two adjacent cells

(B)

100 nm

Figure 19–27 Plasmodesmata. (A) The cytoplasmic channels of

plasmodesmata pierce the plant cell wall and connect cells in a plant

together. (B) Each plasmodesma is lined with plasma membrane that is

common to the two connected cells. It usually also contains a fine tubular

structure, the desmotubule, derived from smooth endoplasmic reticulum.

(C) Electron micrograph of a longitudinal section of a plasmodesma from

a water fern. The plasma membrane lines the pore and is continuous from

one cell to the next. Endoplasmic reticulum and its association with the

central desmotubule can also be seen. (D) A similar plasmodesma seen in

cross section. (C and D, from R. Overall, J. Wolfe and B.E.S. Gunning, in

Protoplasma 111, pp. 134–150. Heidelberg: Springer-Verlag, 1982.)

(C)

0.1 µm

is similar to the molecular-weight cutoff for gap junctions. As with gap junctions,

transport through plasmodesmata is regulated. Dye-injection experiments, for

example, show that there can be barriers to the movement of even low-molecularweight

molecules between certain cells, or groups of cells, that are connected by

apparently normal plasmodesmata; the mechanisms that restrict communication

in these cases are not understood.

(D)

cell

wall

desmotubule

plasma membrane

25 nm

Selectins Mediate Transient Cell–Cell Adhesions in the

Bloodstream

We now complete our overview of cell–cell junctions and adhesion by briefly

describing some of the more specialized adhesion MBoC6 m19.38/19.28 mechanisms used in some tissues.

In addition to those we have already discussed, at least three other superfamilies

of cell–cell adhesion proteins are important: the integrins, the selectins,

and the adhesive immunoglobulin (Ig) superfamily members. We shall discuss

integrins in more detail later: their main function is in cell–matrix adhesion, but a

few of them mediate cell–cell adhesion in specialized circumstances. Ca 2+ dependence

provides one simple way to distinguish among these classes of adhesion

proteins experimentally. Selectins, like cadherins and integrins, require Ca 2+ for

their adhesive function; Ig superfamily members do not.

Selectins are cell-surface carbohydrate-binding proteins (lectins) that mediate

a variety of transient cell–cell adhesion interactions in the bloodstream. Their

main role, in vertebrates at least, is in governing the traffic of white blood cells

into normal lymphoid organs and any inflamed tissues. White blood cells lead a

nomadic life, roving between the bloodstream and the tissues, and this necessitates

special adhesive behavior. The selectins control the binding of white blood

cells to the endothelial cells lining blood vessels, thereby enabling the blood cells

to migrate out of the bloodstream into a tissue.

Each selectin is a transmembrane protein with a conserved lectin domain that

binds to a specific oligosaccharide on another cell (Figure 19–28A). There are at

least three types: L-selectin on white blood cells, P-selectin on blood platelets and

on endothelial cells that have been locally activated by an inflammatory response,

and E-selectin on activated endothelial cells. In a lymphoid organ, such as a lymph

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