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142  Unit 1  Levels of Organization

	4  Figure 4–1  The Polarity of Epithelial Cells.  Many epithelial cells                          Intercellular connections involve either extensive areas of
                                                                                             opposing plasma membranes or specialized attachment sites,
    have functional specializations associated with an apical surface that                   discussed shortly. Large areas of opposing plasma membranes
    is exposed to either the external environment or internal spaces, and                    are interconnected by transmembrane proteins called cell ad-
    basolateral surfaces (base and sides) that interact with deeper tissues                  hesion molecules (CAMs), which bind to each other and to
    and adjacent cells. Often, the apical surface has microvilli; sometimes                  extracellular materials. For example, CAMs on the basolateral
    it has cilia. This functional polarity is also evident in the uneven distri-             surface of an epithelium help bind the cell to the underlying
    bution of organelles between the free surface (here, the top) and the                    basement membrane. The membranes of adjacent cells may
    basement membrane. In some epithelia, such as the lining of the kid-                     also be bonded by a thin layer of proteoglycans that contain
    ney tubules, mitochondria are concentrated near the base of the cell,                    polysaccharide derivatives known as glycosaminoglycans (GAGs),
    where energy is in high demand for the cell’s transport activities.                      most notably hyaluronan (hyaluronic acid).

                                         Cilia                                                    Cell junctions are specialized areas of the plasma mem-
                                                                                             brane that attach a cell to another cell or to extracellular materi-
                                         Microvilli                                          als. The three most common types of cell junctions are (1) tight
                                         Apical                                              junctions, (2) gap junctions, and (3) desmosomes (Figure 4–2a).
                                         surface
                                                                                                  Some epithelial functions require rapid intercellular com-
                                                                          Golgi              munication. At a gap junction (Figure 4–2b), two cells are held
                                                                          apparatus          together by two interlocking transmembrane proteins called
                                                                                             connexons. Each connexon is composed of six connexin proteins
                                                                          Nucleus            that form a cylinder with a central pore. Two aligned connexons
                                                                                             form a narrow passageway that lets small molecules and ions pass
                                                                          Mitochondria       from cell to cell. Gap junctions are common among epithelial
                                                                          Basement membrane  cells, where the movement of ions helps coordinate functions
                                                                          Basolateral        such as the beating of cilia. Gap junctions are also common in
                                                                          surfaces           other tissues. For example, gap junctions in cardiac muscle tissue
                                                                                             and smooth muscle tissue are essential in coordinating muscle
    or toxic compounds such as nicotine and carbon monoxide                                  cell contractions. A number of human diseases are caused by mu-
    in cigarette smoke injure the cilia or the epithelial cells, cili-                       tations in the genes that code for different connexin proteins.
    ary movement can stop. Without the protective upward flow of
    mucus, infection or disease become more likely.                                               At a tight junction (also known as an occluding junc-
                                                                                             tion), the lipid portions of the two plasma membranes are
    Maintaining the Integrity of Epithelia                                                   tightly bound together by interlocking membrane proteins
                                                                                             (Figure 4–2c). Inferior to the tight junctions, a continuous
    To be effective as a barrier, an epithelium must form a com-                             adhesion belt forms a band that encircles cells and binds them
    plete cover or lining. Three factors help maintain the physi-                            to their neighbors. The bands are attached to the microfila-
    cal integrity of an epithelium: (1) intercellular connections,                           ments of the terminal web. p. 97 This kind of attachment
    (2) attachment to the basement membrane, and (3) epithelial                              is so tight that tight junctions largely prevent water and solutes
    maintenance and repair.                                                                  from passing between the cells.

    Intercellular Connections                                                                     When the epithelium lines a tube, such as the intestinal
                                                                                             tract, the apical surfaces of the epithelial cells are exposed
    Cells in an epithelium are firmly attached to one another, and                           to the space inside the tube, a passageway called the lumen
    the epithelium as a unit is attached to extracellular fibers of                          (LOO-men). Tight junctions effectively isolate the contents of
    the superficial clear layer of the basement membrane. Many                               the lumen from the basolateral surfaces of the cell. For exam-
    cells in your body form permanent or temporary bonds with                                ple, tight junctions near the apical surfaces of cells that line
    other cells or extracellular material. Epithelial cells, however,                        the digestive tract help keep enzymes, acids, and wastes in the
    are specialists in intercellular connection.                                             lumen from reaching the basolateral surfaces and digesting or
                                                                                             otherwise damaging the underlying tissues and organs.

                                                                                                  Most epithelial cells are subject to mechanical stresses—
                                                                                             stretching, bending, twisting, or compression—so they must
                                                                                             have durable interconnections. At a desmosome (DEZ-mo. -so. m;
                                                                                             desmos, ligament + soma, body), CAMs and proteoglycans link
                                                                                             the opposing plasma membranes. Desmosomes are very strong
                                                                                             and can resist stretching and twisting.
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