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Contraction and Excitation of Smooth Muscle

Contraction of Smooth Muscle


Composed of far smaller fibers
The same attractive forces cause contraction in smooth muscle as in skeletal muscle, but the internal physical arrangement
of smooth muscle fibers is very different.
• Types of Smooth Muscle
o Multi-unit smooth muscle
 Composed of discrete, separate muscle fibers. Each fiber operates independently of the others and
is innervated by a single nerve ending
o Unitary smooth muscle (syncytial/visceral smooth muscle)
 Mass of hundreds-thousands of smooth muscle fibers that contract together as a single unit (cell
membranes joined by gap junctions through which ions can flow freely from one muscle cell to
the next so that action potentials or simple ion flow without action potentials can travel from one
fiber to the next and cause the muscle fibers to contract together.)
 Major share of control is exerted by non-nervous stimuli
• Contractile Mechanism in Smooth Muscle
o Chemical Basis for Smooth Muscle Contraction
 Contains actin and myosin filaments but does not contain troponin complex so the mechanism for
control of contraction is different.
 Actin/myosin in smooth muscle interact in much the same way they do in skeletal muscle.
 Differences between smooth muscle and skeletal muscle:
• Physical organization
• Excitation-contraction coupling
• Control of the contractile process by calcium ions
• Duration of contraction
• Amount of energy required for contraction
o Physical Basis for Smooth Muscle Contraction
 Large numbers of actin filaments attached to dense bodies
 Myofilaments and side-polar cross-bridges
• Allows myosin to pull an actin filament in one direction on one side while
simultaneously pulling another actin filament in the opposite direction on the other side
 contraction as much as 80% of their length of muscle
o Comparison of Smooth Muscle Contraction and Skeletal Muscle Contraction
 Slow cycling of myosin cross bridges
• Cycling= attachment to actin, release from the actin and reattachment for the next cycle
• Cycling is much slower in smooth muscle
o The cross-bridge heads have far less ATPase activity than in skeletal muscle, so
the degradation of the ATP that energizes the movements of the cross-bridge
heads is greatly reduced  slows rate of cycling
• Fraction of time that cross-bridges remain attached is increased in smooth muscle
(determines the force of contraction)
 Energy required to sustain smooth muscle contraction
• Only one molecule of ATP required for each cycle, regardless of duration (smooth muscle
uses less ATP)
 Slowness of onset of contraction and relaxation of the total smooth muscle tissue
• Most smooth muscle contraction is prolonged tonic contraction (hours to days)
• Slow onset of contraction of smooth muscle as well as its prolonged contraction is caused
by the slowness of attachment and detachment of the cross-bridges with the actin
filaments.
 Force of muscle contraction
• Force of contraction of smooth muscle is often greater than that of skeletal muscle.
• Force of smooth muscle contraction results from the prolonged period of attachment of
the myosin cross-bridges to the actin filaments
 Latch mechanism for prolonged holding of contractions of smooth muscle
• It can maintain prolonged tonic contraction in smooth muscle for hours with little use of
energy
 Stress-relaxation of smooth muscle
• Ability to return to nearly its original force of contraction after it has been
elongated/shortened
• Except for short periods of time, these phenomena allow a hollow organ to maintain
about the same amount of pressure inside its lumen despite long-term, large changes in
volume
• Regulation of Contraction by Calcium Ions: smooth muscle does not contain troponin
o Combination of calcium ions with calmodulin—activation of myosin kinase and phosphorylation of the
myosin head
 Calcium ions bind with calmodulin and this unit then joins with and activates myosin kinase.
 Myosin kinase phosphorylates the regulatory chains of the myosin head. It can now bind
repeatedly with the actin filament causing muscle contraction.
o Cessation of contraction—role of myosin phosphatatase
 [Calcium ion] falls below a critical level  myosin phosphatase splits the phosphate from the
regulatory light chain
 Cycling stops, contraction ceases
o Possible mechanism for regulation of the latch phenomenon
Nervous and Hormonal Control of Smooth Muscle Contraction
Smooth muscle can be stimulated to contract by multiple types of signals (smooth muscle contains many types of receptor
proteins that can initiate the contractile process.)
• Neuromuscular junctions of smooth muscle
o Physiologic anatomy of smooth muscle neuromuscular junctions
 Autonomic nerve fibers that innervate smooth muscle (diffuse junctions: fibers secrete transmitter
substance into the matrix coating of the smooth muscle which then diffuses to the cells.)
 Nerve fibers often innervate only the outer layer of a many-layered mass. Muscle excitation
travels from this outer layer to the inner layers by action potential conduction in the muscle mass
or additional diffusion of the transmitter.
 Most of the terminal axons have multiple varicosities distributed along their axes
o Excitatory and Inhibitory transmitter substances secreted at the smooth muscle neuromuscular junction
 Acetylcholine and norepinephrine
 Receptor proteins: excitatory receptors and inhibitory receptors (type of receptor determines
whether smooth muscle is inhibited or excited and also determines which of the two transmitters
is effective in causing the excitation or inhibition)
• Membrane Potentials and Action Potentials in Smooth Muscle
o Membrane potentials in smooth muscle
 Normal resting state: intracellular potential is approximately -50 to -60 mV
o Action potentials in unitary smooth muscle
 Spike potentials
 Action potentials with plateaus
o Importance of calcium channels in generating the smooth muscle action potential
 Flow of calcium ions to the interior of the smooth muscle fiber is mainly responsible for action
potentials.
 Calcium channels open more slowly and remain open much longer than sodium channels 
prolonged plateau action potentials
 Calcium ions act directly on the smooth muscle contractile mechanism to cause contraction
o Slow wave potentials in unitary smooth muscle and spontaneous generation of action potentials
 Some smooth muscle is self-excitatory (action potentials arise within the smooth muscle cells
themselves without an extrinsic stimulus)
 When slow waves are strong enough, they can initiate action potentials and contraction can occur
o Excitation of visceral smooth muscle by muscle stretch
o Depolarization of multi-unit smooth muscle without action potentials
• Effect of Local Tissue Factors and Hormones to Cause Smooth Muscle Contraction without Action Potentials
o Smooth muscle contraction in response to local tissue chemical factors
 Lack of oxygen in the local tissues causes smooth muscle relaxation and therefore vasodilation.
 Excess carbon dioxide causes vasodilation.
 Increased H+ concentration causes vasodilation.
o Effects of hormones on smooth muscle contraction
 A hormone causes contraction of a smooth muscle when the muscle cell membrane contains
hormone-gated excitatory receptors for the respective hormone.
 The hormone cause sinhibition if the membrane contains inhibitory receptors for the hormone.
o Mechanisms of smooth muscle excitation or inhibition by hormones or local tissue factors
• Source of Calcium Ions that cause Contraction through the Cell Membrane and from the Sarcoplasmic Reticulum
o Role of the smooth muscle sarcoplasmic reticulum
 The SR is only slightly developed in most smooth muscle  almost all the calcium ions that
cause contraction enter the muscle cell from the extracellular fluid at the time of the action
potential/stimulus
 In general, the more extensive the sarcoplasmic reticulum in the smooth muscle fiber, the more
rapidly it contracts.
o Effect on smooth muscle contraction caused by changing of extracellular calcium ion concentration
 The force of contraction of smooth muscle usually is highly dependent on extracellular fluid
calcium ion concentration.
o A calcium pump is required to cause smooth muscle relaxation

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