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          smooth muscle does not lose its contractile   function (e.g., smooth muscle in the walls
                                                  of blood vessels).
          ability. The reverse occurs upon emptying
  VetBooks.ir  of a visceral organ when the stretched
          muscle shortens back to its original length:
          all tension is lost at first but returns shortly.   Role and Sources of Calcium
          Plasticity is believed to be due to changes   As in skeletal muscle, contraction and
          in the arrangement or binding of the myo-  relaxation of smooth muscle are linked to
          sin and actin filaments upon stretching or   the Ca  concentration in the cytosol of
                                                         2+
          shortening.
                                                  smooth muscle cells. However, how this
                                                  concentration is regulated and the role of
          Contraction and Relaxation              Ca  in the contraction process is quite
                                                     2+
                                                  different in the two types of muscle. Some
          All skeletal muscle contraction depends   calcium is stored in the sarcoplasmic retic-
          on ACh release at a neuromuscular junc-  ulum of smooth muscle cells, but many
          tion and the generation and propagation   types of smooth muscle cells also contain a
          of action potentials on the cell membrane.   significant number of calcium channels
          This is not the case for smooth muscle.   in the outer cell membrane (Fig. 9‐9). These
          The stimuli that bring about smooth mus-  calcium channels may be either  voltage‐
          cle contraction and relaxation are quite   gated or ligand‐gated, and the smooth
          variable. Thus, smooth muscle is a much   muscle within a given organ may  have
          more functionally diverse tissue than   both  types  of  channels  or  primarily  only
          skeletal muscle.                        one type of channel. When these channels
            The contraction and relaxation of most   open in response to the appropriate stimu-
          smooth muscle are much slower events    lus (changes in membrane potential or
          than the rapid muscle twitch that is char-  presence of specific ligands), calcium can
          acteristic of skeletal muscle. This permits   diffuse into the cell to initiate contraction
          the maintenance of a relatively constant   (Fig.  9‐9). Receptors that are specific for
          pressure with the use of little cell energy.   many different ligands (e.g., hormones
          This is characteristic of smooth muscle in   and  neurotransmitters) are found on the
          organs that require a constant state of tone   smooth muscle of different organs. This is
          or some degree of contraction for normal   important functionally, because a given


                                  Extracellular fluid
                      Calcium                             Cell membrane of
                      channel    Ca 2+     Ca 2+         smooth muscle cell



                          Ca 2+   Ca + calmodulin
                                    2+
                                                      Other enzymes which
                                    Inactive kinase  dephosphorylate myosin
                          Ca 2+    Ca 2+  calmodulin
                                                    Phosphorylated
                                    Activated kinase           + actin
                                                    myosin
                       Sarcoplasmic
                        reticulum                      Contraction


          Figure 9-9.  Sources and role of calcium ions in the initiation and regulation of smooth muscle contrac-
          tion. Dotted lines associated with calcium ions indicate movement by diffusion. Solid lines associated
          with calcium ions indicate removal of calcium from the cytosol to promote relaxation. Relaxation can also
          be promoted by the action of other intracellular enzymes that dephosphorylate myosin.
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