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206       Organelles



             Structure and functions                          B. Microtubules
                                                              Only the cell’s microtubules are shown here.
             The cytoskeleton carries out three major
             tasks:                                           They radiate out in all directions from a center
                                                              near the nucleus, the centrosome.The tube-
             • It represents the cell’s mechanical scaffold-  shaped microtubules are constantly being
                ing, which gives it its typical shape and     synthesized and broken down at their (+)
                connects membranes and organelles to          ends. In the centriole, the (–) end is blocked
                each other. This scaffolding has dynamic      by associated proteins (see p. 204). The (+)
                properties; it is constantly being synthe-    end can also be stabilized by associated pro-
                sized and brokendowntomeet the cell’s         teins—e. g., when the microtubules have
                requirements and changing conditions.         reached the cytoplasmic membrane.
             • It acts as the motor for movement of animal       The microtubules are involved in defining
                cells. Not only muscle cells (see p. 332), but  theshape of thecelland also serveas guiding
                also cells of noncontractile tissues contain  tracks for the transport of organelles. To-
                many different motor proteins, which they     gether with associated proteins (dynein, kine-
                use to achieve coordinated and directed       sin), microtubules areableto carry outme-
                movement. Cell movement, shape changes        chanical work—e. g., during the transport of
                during growth, cytoplasmic streaming, and     mitochondria, the movement of cilia (hair-
                cell division are all made possible by com-   like cell protrusions in the lungs, intestinal
                ponents of the cytoskeleton.                  epithelium, and oviduct) and the beating of
             • It serves as a transport track within the cell.  the flagella of sperm. Microtubules also play a
                Organelles and other large protein com-       special role in the mitotic period of cell divi-
                plexes can move along the filaments with      sion (see p. 394).
                the help of the motor proteins.
                                                              C. Architecture
             A. Microfilaments and intermediate               Thecomplex structureand net-like densityof
             filaments
                                                              the cytoskeleton is illustrated here using
             The illustration schematically shows a detail    three examples in which the cytoskeletal
             of the microvilli of an intestinal epithelial cell  components are visualized with the help of
             as an example of the structure and function of   antibodies.
             the components of the cytoskeleton (see also
             C1).                                                1. The border of an intestinal epithelial cell
                Microfilaments of Factin traverse the mi-     is seen here (see also B). There are micro-
             crovilli in ordered bundles. The microfila-      filaments (a) passing from the interior of the
             ments are attached to each other by actin–as-    cell out into the microvilli. The filaments are
                                                              firmly held together by spectrin (b), an asso-
             sociated proteins, particularly fimbrin and vil-
             lin. Calmodulin and a myosin–like ATPase con-    ciated protein, and they are anchored to in-
             nect the microfilaments laterally to the         termediate filaments (c).
                                                                 2. Only microtubules are seeninthis fibro-
             plasma membrane. Fodrin, another microfila-
             ment–associated protein, anchors the actin       blast cell. They originate from the microtubule
                                                              organizing center (centrosome) and radiate
             fibers to each other at the base, as well as
             attaching them to the cytoplasmic membrane       out as far as the plasma membrane.
                                                                 3. Keratin filaments are visible here in an
             and to a network of intermediate filaments. In   epithelial cell. Keratin fibers belong to the
             this example, the microfilaments have a
             mainly static function. In other cases, actin is  group of intermediate filaments (see pp. 70,
                                                              204; d = nucleus).
             also involved in dynamic processes. These in-
             clude muscle contraction (see p. 332), cell
             movement, phagocytosis by immune cells,
             the formation of microspikes and lamellipo-
             dia (cellular extensions), and the acrosomal
             process during the fusion of sperm with the
             egg cell.



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