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Contraction and excitation of smooth muscle
Smaller fibers Principle of contraction Same as skeletal muscle in many aspects Attraction between myosin and actin Different arrangement of muscle fibers
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Types of smooth muscle Different among different organs Two types
Physical dimensions Organization (bundle/sheet) Responsiveness to different stimuli Innervation Function Two types Multi-unit Unitary (single-unit)
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Multi-unit smooth muscle
Composed of discrete muscle fibers Innervated by a single nerve ending Covered with basement membrane Separation of individual fiber Independent contraction Controlled by nerve signal No spontaneous contraction
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Unitary/single-unit smooth muscle
Mass of muscle cells contracting as one unit (syncytical smooth muscle) Aggregated into sheet or bundle Adhered together Membrane (force) Gap junctions (action potential) Critical for contraction Found in the wall of digestive, urinary, and reproductive tracts, and blood vessel
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Contractile process Chemical basis Actin and myosin filaments
No troponin complex Interaction of filaments Similar to that of skeletal muscle Activated by Ca ions Require ATP
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Contractile process Physical basis No stratiation Actin filaments
Attached to dense bodies Myosin filaments Interspersed among actin filaments Ratio of actin filaments to myosin filaments More actin in smooth muscle Greater ratio of fiber length
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Individual contractile unit
Large number of actin filaments (attached to two dense bodies) Single myosin fiber (half-way between two dense bodies) Dense body (Z-disc) No regularity Side-polar arrangement of myosin cross-bridges
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Individual contractile unit
Side-polar arrangement of myosin cross-bridges One side of myosin heads pull actin fiber in one direction Greater contraction (80 % of the length)
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Smooth muscle contraction
Comparison with skeletal muscle Prolonged tonic contraction Different chemical characteristics
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Smooth muscle contraction
Cycling of myosin cross-bridge Slower repeated attachment and release of actin by cross-bridge (1/10 to 1/300 of skeletal muscles) Increased time of interaction Lower concentrations of ATPase in the cross-bridge heads
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Smooth muscle contraction
Energy requirements Much less (1/10 to 1/300) compared to skeletal muscles Slow attachment and detachment of myosin heads Critical for energy conservation Indefinite contraction of visceral smooth muscles
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Smooth muscle contraction
Slower onset of contraction and relaxation Typical smooth muscle Contraction initiated 50 to 100 mSec after excitation Full contraction achieved 0.5 sec later Decline in contractile force 1 to 2 sec Total time for process 1 to 3 sec
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Smooth muscle contraction
Slower onset of contraction and relaxation 30 times slower than skeletal muscles Various type of smooth muscles Slower detachment and attachment of cross-bridges Slower response to elevation in Ca concentrations
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Prolonged holding of contraction
Force of contraction Greater maximum force of contraction 4 to 6 kg/cm2 compared to 3 to 4 kg/cm2 for skeletal muscles Prolonged attachment of myosin to actin Prolonged holding of contraction Latch mechanism Low energy consumption Continuous stimulation
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Stress-relaxation Ability to return to near the original force of contraction shortly after elongation or contraction Increased stretching of hollow organ Immediate rise in pressure Pressure returns to normal even when the organ is continuously stretched
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Reverse stress-relaxation
Sudden decrease in pressure Immediate decrease in pressure Pressure returns to normal immediately after the decrease in size
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Increase in size Increase in pressure Normal size Normal pressure Decreased size Normal pressure Increased size Normal pressure Decrease in size Decrease in pressure
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Role of Ca ion Initiating factor Increase in intracellular Ca
Rise in intracellular Ca Increase in intracellular Ca Nerve impulses Hormones Physical External environment
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Role of Ca ion No tropinins Different mechanisms
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Role of Ca ion Calmodulin as a regulatory protein
Binds Ca Activation of myosin kinase by Ca-calmodulin complex Phosphorylation of myosin light chain Regulatory chain within the mysoin head Interaction of myosin head with actin filament
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Cessation of contraction
Loss of Ca Myosin head must be dephosphorylated Myosin phosphatase Timing of cessation and relaxation Myosin phosphatase concentration dependent
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Source of Ca ions Less-developed SR compared to skeletal muscle
Influx of Ca ions fromextracellular fluid Concentration gradient (<10-7 M inside vs. > 10-3 M outside) Needs 200 to 300 mSec for diffusion (latent period) Contraction of smooth muscle is highly dependent on extracellular Ca
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Role of SR Sacroplasmic tubules in some large smooth muscle cells
Caveole (invagination of the membrane) Rudimentary analog of T-tubules Stimulates release of Ca by sacroplasmic tubules when the action potential reaches More extensive the sacroplasmic reticulum, rapid contraction of muscle
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Removal of Ca Relaxation of smooth muscle Ca pump
Move Ca from the inside to the outside or into sacroplasmic tubules/reticulum Slow-acting (longer contraction)
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Regulation of latch phenomenon
Mechanism Myosin kinase and myosin phosphatase Activation of both enzymes Increased velocity of contraction (increased cycle frequency) Inactivation of enzymes Decrease velocity of contraction Increased interaction between myosin head and actin filament (per cycle) Increased number of myosin heads interacting with actin filament (increased static force of contraction)
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Nervous and hormonal control
Skeletal muscle Exclusives stimulated by nerve fiber Smooth muscle Contains receptors for various stimuli Stimulatory Inhibitory
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Neural stimulation Neuromusclular junction
Innervated by autonomic nerve fibers Only on the top layer Excitation potentials travel toward the inner layer by action potential conduction or additional diffusion of neurotransmitter Nerve fibers branch diffusely on top of sheet No direct contact Diffusion junction (few nm to few mm away from muscle membrane)
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Formation of varicosities
No motor end plate Vesicles containing neurotransmitters Acetylcholine Norepinephrine Formation of contact junction Close proximity of varicosities to the muscle membrane (20-30 nm) Faster contraction compared to diffusion junction
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Release of excitatory and inhibitory substances
Acetylcholine Norepinephrine Secreted by different fibers (either/or, not both from the same fiber)
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Acetylcholine Norepinephrine Excitatory in some muscles
Inhibitory in other muscles Norepinephrine Excitatory in muscles where acetylcholine is an inhibitor Inhibitory in muscles where acetylcholine is a positive stimulator
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Differential responses to acetylcholine and norepinephrine
Types of receptors
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Membrane and action potential
Membrane potential -50 to -16 mV at resting state Action potential Similar to skeletal muscle in unitary smooth muscle Absent in multi-unit smooth muscles
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Action potential in unitary smooth muscle
Spike potential Action potential with plateau Initiated by factors other than neural signals
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Spike potential Same as the action potential in skeletal muscle
10 to 50 mSec in duration
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Action potential with plateaus
Delayed depolarization As much as 1 second Prolonged contraction of the muscle
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Role of Ca channels Numerous voltage-gated Ca channels
Few voltage-gated Na channels Action potential generated by the influx of Ca ions Ca channels open slower than Na channels and remain open longer Prolonged plateaus Ca ions more critical for smooth muscle contraction
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Slow wave potential Self-excitation of some smooth muscles
Absence of stimuli Slow wave potential Slow wave (pacemaker wave) Not action potential (no spreading) Localized phenomenon
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Cause of slow wave Importance Unknown Initiation of action potential
One or more action potential per each peak of slow wave Rhythmical contraction of smooth muscle Initiation of action potential by stretching
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Depolarization of multi-unit smooth muscle
Contraction of multi-unit smooth muscle Input from autonomic nerve fiber Binding of neurotransmitters to their receptors Depolarization No action potential Nerve fibers are too small to initiate action potential Junctional potential (local depolarization) Electronic transmission to other muscle fibers
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Role of tissue factors and hormones
Non-nervous factors Do not generate action potential Local tissue factors (often associated with smooth muscle surrounding blood vessels) Oxygen Carbon dioxide Hydrogen ions
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Hormones Requires presence of excitatory/inhibitory receptors Mechanisms Changes in membrane potentials Release of Ca ions from SR Removal of Ca ions Alteration of enzyme activity
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