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Smooth Muscle                   pendent and, in many cases, spontaneous (my-
                                       ogenic tonus). The second type, multi-unit
       Smooth muscle (SmM) consists of multiple  SmM, contracts primarily due to stimuli from
       layers of spindle-shaped cells. It is involved in  the autonomic nervous system (neurogenic
       the function of many organs (stomach, in-  tonus). This occurs in structures such as the
       testine, gall bladder, urinary bladder, uterus,  arterioles, spermatic ducts, iris, ciliary body,
       bronchi, eyes, etc.) and the blood vessels,  and the muscles at the roots of the hair. Since
       where it plays an important role in circulatory  these SmM cells generally are not connected
       control. SmM contains a special type of F-  by gap junctions, stimulation remains local-
    Nerve and Muscle, Physical Work  and no sarcomeres (nonstriated). It is there-  degree of depolarization (e.g., through stretch
       actin-tropomyosin and myosin II filaments
                                       ized, as in the motor units of the skeletal
                                       muscle.
       (! p. 60), but lacks troponin and myofibrils.
       Furthermore, it has no distinct tubular system
                                        Smooth muscle tonus is regulated by the
                                       or pacemaker cells) as well as by transmitter
       fore called smooth muscle because of this lack
                                       substances (e.g., acetylcholine or noradrena-
       of striation (see p. 59 A for further differences
                                       line) and numerous hormones (e.g., estrogens,
       in the muscle types). SmM filaments form a
                                       progesterone and oxytocin in the uterus and
       loose contractile apparatus arranged approxi-
                                       histamine, angiotensin II, adiuretin, serotonin
       mately longitudinally within the cell and at-
       tached to discoid plaques (see B for model),
                                       and bradykinin in vascular muscle). An in-
       which also provide a mechanical means for
                                       directly or indirectly increases the cytosolic
       shorten much more than striated muscle.
                                          concentration to more than 10
                                                                mol/L.
                                                              – 6
                                       Ca
                                        2+
    2  cell–cell binding of SmM. Smooth muscle can  crease in tonus will occur if any of these factors
         The membrane potential of the SmM cells of
                                            influx comes mainly from extracellu-
                                           2+
                                       The Ca
       many organs (e.g., the intestine) is not con-  lar sources, but a small portion comes from in-
       stant, but fluctuates rhythmically at a low  tracellular stores (! B1). Ca 2+  ions bind to cal-
       frequency (3 to 15 min ) and amplitude (10 to  modulin (CM) (! B2), and Ca -CM promotes
                                                         2+
                     – 1
       20 mV), producing slow waves. These waves  contraction in the following manner.
       trigger a burst of action potentials (spikes)  Regulation at myosin II (! B3): The Ca -CM
                                                                2+
       when they exceed a certain threshold poten-  complex activates myosin light chain kinase
       tial. The longer the slow wave remains above  (MLCK), which phosphorylates myosin’s regu-
       the threshold potential, the greater the num-  latory light chain (RLC) in a certain position,
       ber and frequency of the action potentials it  thereby enabling the myosin head to interact
       produces. A relatively sluggish contraction oc-  with actin (! B6).
       curs around 150 ms after a spike (! p. 59 A, left  Regulation at the actin level (! B4). The
       panel). Tetanus occurs at relatively low spike  Ca -CM complex also binds with caldesmon
                                        2+
       frequencies ( ! p. 66). Hence, SmM is con-  (CDM), which then detaches from the actin–
       stantly in a state of a more or less strong con-  tropomyosin complex, thus making it available
       traction (tonus or tone). The action potential of  for filament sliding (! B6). Phosphorylation of
       SmM cells of some organs has a plateau similar  CDM by protein kinase C (PK-C) also seems to
       to that of the cardiac action potential  be able to induce filament sliding (! B5).
       (! p. 59 A, middle panel).       Factors that lead to a reduction of tonus are:
         There are two types of smooth muscles  reduction of the cytosolic Ca 2+  concentration
       (! A). The cells of single-unit SmM are electri-  to less than 10 – 6  mol/L (! B7 ), phosphatase
       cally coupled with each other by gap junctions  activity (! B8), and PK-C if it phosphorylates
       (! pp. 18 and 50). Stimuli are passed along  another position on the RLC (! B9).
       from cell to cell in organs such as the stomach,  When length–force curves are recorded for
       intestine, gallbladder, urinary bladder, ureter,  smooth muscle, the curve shows that muscle
       uterus, and some types of blood vessels.  force decreases continuously while muscle
       Stimuli are generated autonomously from  length remains constant. This property of a
       within the SmM, partly by pacemaker cells). In  muscle is called plasticity.
   70  other words, the stimulus is innervation-inde-
       Despopoulos, Color Atlas of Physiology © 2003 Thieme
       All rights reserved. Usage subject to terms and conditions of license.
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