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!
       I band region are flexible and function as “elas-  force of cardiac muscle at rest is greater than
       tic bands” that counteract passive stretching of  that of skeletal muscle (! E1, 2).
       a muscle and influence its shortening velocity.  Skeletal muscle normally functions in the
                                       plateau region of its length–force curve,
       The extensibility of titin molecules, which can
       stretch to up to around ten times their normal length  whereas cardiac muscle tends to operate in the
       in skeletal muscle and somewhat less in cardiac  ascending limb (below L max) of its length–force
       muscle, is mainly due to frequent repetition of the  curve without a plateau (! C, E1, 2). Hence,
       PEVK motif (proline-glutamate-valine-lysine). In very  the ventricle responds to increased diastolic
       strong muscle extension, which represents the  filling loads by increasing its force develop-
    Nerve and Muscle, Physical Work  stretches, the more sudden and crude this type of  steeper curve (! E2).  2+  +
       steepest part of the resting extensibility curve (! D),
                                       ment (Frank–Starling mechanism; ! p. 204).
       globular chain elements called immunoglobulin C2
                                       In cardiac muscle, extension also affects
       domains also unfold. The quicker the muscle
                                                         2+
                                       troponin’s sensitivity to Ca , resulting in a
       “shock absorber” action will be.
                                        Action potentials in cardiac muscle are of
       The length (L) and force (F) or “tension” of a
                                       much longer duration than those in skeletal
       muscle are closely related (! C, E). The total
                                       muscle (! p. 59 A) because g K temporarily
       force of a muscle is the sum of its active force
                                       decreases and g Ca increases for 200 to 500 ms
       and its extension force at rest, as was ex-
                                       after rapid inactivation of Na channels. This
       plained above. Since the active force is deter-
                                       allows the slow influx of Ca , causing the ac-
       myosin interactions, it varies in accordance
                                       refractory period does not end until a contrac-
       with the initial sarcomere length (! C, D).
                                       tion has almost subsided (! p. 59 A). There-
       Skeletal muscle can develop maximum active
    2  mined by the magnitude of all potential actin-  tion potential to reach a plateau. As a result, the
                                       fore, tetanus cannot be evoked in cardiac
       (isometric) force (F 0) from its resting length  muscle.
       (L max; sarcomere length ca. 2 to 2.2 µm; ! C).  Unlike skeletal muscle, cardiac muscle has
       When the sarcomeres shorten (L ! L max), part  no motor units. Instead, the stimulus spreads
       of the thin filaments overlap, allowing only  across all myocardial fibers of the atria and
       forces smaller than F 0 to develop (! C). When  subsequently of the ventricles generating an
       L is 70% of L max (sarcomere length: 1.65µm),  all-or-none contraction  of both atria and,
       the thick filaments make contact with the Z  thereafter, both ventricles.
       disks, and F becomes even smaller. In addition,  In cardiac muscle but not in skeletal muscle,
       a greatly pre-extended muscle (L " L max) can  the duration of an action potential can change
       develop only restricted force, because the  the force of contraction, which is controlled by
       number of potentially available actin–myosin  the variable influx of Ca 2+  into the cell.
       bridges is reduced (! C). When extended to  The greater the force (load), the lower the
       130% or more of the L max, the extension force at  velocity of an (isotonic) contraction (see velo-
       rest becomes a major part of the total muscle  city–force diagram, F1). Maximal force and a
       force (! E).                    small amount of heat will develop if shorten-
         The length–force curve corresponds to the  ing does not occur. The maximal velocity (bi-
       cardiac pressure–volume diagram in which  ceps: ca. 7 m/s) and a lot of heat will develop in
       ventricular filling volume corresponds to  muscle without a stress load. Light loads can
       muscle length, and ventricular pressure corre-  therefore be picked up more quickly than
       sponds to muscle force; ! p. 202. Changes in  heavy loads (! F2). The total amount of energy
       the cytosolic Ca 2+  concentration can modify  consumed for work and heat is greater in
       the pressure–volume relationship by causing a  isotonic contractions than in isometric ones.
       change in contractility (! p. 203 B2).  Muscle power is the product of force and the
         Other important functional differences be-  shortening velocity: N · m · s – 1  = W (! F1,
       tween cardiac muscle and skeletal muscle are  colored areas).
       listed below (see also p. 59 A):
         Since skeletal muscle is more extensible
   68  than the cardiac muscle, the passive extension
       Despopoulos, Color Atlas of Physiology © 2003 Thieme
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