Skeletal Muscle
Skeletal muscles are attached to
the skeleton by tendons.
Skeletal muscles typically contain
many, many muscle fibers.
Arranged in bundles with
connective tissue, blood vessels
and nerves.
-Bundles of muscle fibers
-Fiber – bundles of myofibrils
-Myofibrils
Sarcomeres (end to end)
-FUNCTIONAL UNIT
Sarcomere: Functional Unit
The sarcomere is composed of:
-Thick filaments called myosin,
(anchored in place by titin fibers)
-Thin filaments called actin,
(anchored to Z-lines)
Thin filament actually actin, toponin, tropomyosin 1 Myosin : 6 Actin
Globular actin in 2 strands to form a helix 1 Actin : 3 Myosin.
Tropomyosin sits in groove of actin helix
Troponin anchors tropomyosin to actin
Sarcomere: Functional Unit
Sarcomere structures in an electron micrograph.
- I Band – Actin
- A Band – Actin and Myosin
- H Zone - Myosin
Contraction: Sliding Filament Theory
Contraction (shortening):
Myosin binds to actin, and
slides it, pulling the Z-lines
closer together, and
reducing the width of the
I-bands.
****Note that filament
lengths have not
changed.
Contraction: Cross-Bridge Cycling
Contraction:
Myosin’s cross-bridges bind to actin;
The crossbridges then flex to slide actin.
(BINDING & BENDING of Myosin Head Group)
Contraction: Cross-Bridge Cycling
The thick filament called myosin is actually a polymer of myosin molecules.
-2 Myosin heavy chains
-4 Myosin light chains
Form globular head that acts as the flexible cross-bridge that binds both ATP
and actin.
Cross-Bridge Cycling requires ATP
1. The myosin-binding site on actin
becomes available, so the
energized cross-bridge binds.
The full
2. hydrolysis
4. Partial and departure
hydrolysis of
of ADP + Pi
the bound ATP
energizes causes the
or “re-cocks” flexing of
the bridge. the bound
cross-bridge.
3. Binding of a “new” ATP
to the cross-bridge
uncouples the bridge.
Calcium regulation of Cross-Bridge Cycling
In relaxed skeletal muscle,
tropomyosin blocks the cross-
bridge binding site on actin.
Contraction occurs when calcium
ions bind to troponin; this complex
then pulls tropomyosin away from
the cross-bridge binding site.
Excitation-Contraction Coupling
The latent period between excitation and development of tension in a skeletal
muscle includes the time needed to release Ca2+ from the sarcoplasmic
reticulum (SR), move tropomyosin, and cycle the cross-bridges.
Sarcoplasmic Reticulum: Calcium Storage/Release
- The transverse tubules bring action potentials into the interior of the skeletal
muscle fibers, so that the wave of depolarization passes close to the
sarcoplasmic reticulum, stimulating the release of calcium ions.
- The extensive meshwork of sarcoplasmic reticulum assures that when it
releases calcium ions they can readily diffuse to all of the troponin sites.
Sarcoplasmic Reticulum: Calcium Storage/Release
Passage of an action potential
along the transverse tubule opens
nearby voltage-gated calcium
channels, the “ryanodine
receptor,” located on the
sarcoplasmic reticulum, and
calcium ions released into the
cytosol bind to troponin.
The calcium-troponin complex
“pulls” tropomyosin off the
myosin-binding site of actin, thus
allowing the binding of the cross-
bridge, followed by its flexing to
slide the actin filament.
Innervation of Skeletal Muscle
-Voluntary
-Somatic Motor Neurons
-Motor Unit:
Motor neuron plus all the fibers it
innervates
-Multiple Motor Units / muscle
-Motor Unit Recruitment
Neuromuscular Junction
AP ACh release
ACh binds to N-ACh-R
Opening of Na+
Channels
Na+ entry
Graded potential
(end plate potential)
Local current flow
Opening of voltage
gated Na+ Channels
Muscle action potential
Propagation
Excitation-Contraction Coupling
Isotonic and Isometric Contraction
iso = same tonic = tension metric = length
Tension increases
rapidly and
dissipates slowly
Shortening occurs
slowly, only after
taking up elastic
tension; the
relaxing muscle
quickly returns to
its resting length.
Isotonic Contractions with different loads
Light loads are more rapidly
moved than heavy loads.
Twitch, Summation, Tetanus
Unfused tetanus: Fused tetanus:
Partial dissipation of No time for dissipation of elastic
elastic tension between tension between rapidly recurring
subsequent stimuli. stimuli.
Length-Tension Relationship
Optimal-length sarcomere:
lots of actin-myosin overlap
and plenty of room to slide.
Long sarcomere:
actin and myosin
Short sarcomere: do not overlap
actin filaments much, so little
lack room to slide, tension can be
so little tension can developed.
be developed.
Substrates for ATP production in skeletal muscle
In skeletal muscle, ATP production via substrate phosphorylation
is supplemented by the availability of creatine phosphate.
Skeletal muscle’s capacity to produce ATP via oxidative
phosphorylation is further supplemented by the availability
of molecular oxygen bound to intracellular myoglobin.
Muscle Fatigue
-In skeletal muscle, repetitive stimulation leads to fatigue, evident as reduced
tension.
-Rest overcomes fatigue, but fatigue will reoccur sooner if inadequate recovery
time passes.
-Conduction failure (build up of extracellular potassium ions in T-tubules)
-Lactic acid buildup (acidification of muscle alters protein activity)
-Inhibition of cross-bridge cycling (ADP and Pi build up)
Muscle Fiber Types
Skeletal muscle fibers differ by:
1) Maximal velocities of shortening (fast or slow)
Fast Fibers: Myosin with high ATPase actvity –Type II fibers
Slow Fibers: Myosin with low ATPase activity – Type I fibers
2) Major pathway of ATP formation (oxidative or glycolytic)
Oxidative Phosphorylation – oxidative fibers
Lots of blood vessels around fibers (O2 delivery)
Myoglobin (O2 binding protein)
RED muscle fibers
Glycolysis – glycolytic fibers
High concentration of enzymes for glycolysis
WHITE muscle fibers
-Slow Oxidative Fibers (Type I)
-Fast Oxidative Fibers (Type IIa)
-Fast Glycolytic Fibers (Type IIb)
Muscle Fiber Types
Slow-oxidative skeletal muscle responds well
to repetitive stimulation without becoming
fatigued; muscles of body posture are
examples.
Fast-oxidative skeletal muscle responds
quickly and to repetitive stimulation without
becoming fatigued; muscles used in walking
are examples.
Fast-glycolytic skeletal muscle is used for
quick bursts of strong activation, such as
muscles used to jump or to run a short sprint.
Muscle Fiber Types
Control of Muscle Tension
Controlled by 2 main factors:
1) Number of fibers contracting at any one time
-Number of fibers in each motor unit (motor unit size)
-Number of active motor units
2) Amount of tension developed by each fiber
-Action potential frequency
-Fiber length
-Fiber diameter
-Fatigue
Smooth Muscle
-Spindle Shaped
-Single nucleus
-Generally interconnected to form sheets
-Myosin,
-Actin
-Tropomyosin
-BUT NO troponin
-Thin filaments attached
to DENSE BODIES
Smooth Muscle Contraction
Smooth Muscle Contraction
Calcium ions play major regulatory roles in the contraction
of smooth, but the calcium that enters the cytosol
Stimulated smooth muscles binds to calmodulin, forming a
complex that activates the enzyme that phosphorylates
myosin, permitting its binding interactions with actin.
Sources of cytosolic calcium:
(1) Sarcoplasmic reticulum
-Much less SR than in skeletal muscle
-No T-tubules
-Depolarization or second messengers
(2) Extracellular calcium
-voltage-sensitive calcium channels
Smooth Muscle: Membrane Activation
1) Spontaneous electrical Activity:
i) Pacemaker Potentials:
ii) Slow Waves:
Rhythmic changes in the membrane potential of smooth muscles results in
rhythmic patterns of action potentials and therefore rhythmic contraction;
in the gut, neighboring cells use gap junctions to further coordinate these
rhythmic contractions.
Smooth Muscle: Membrane Activation
2) Neurotransmitters and Hormones
Smooth Muscle: Membrane Activation
Types of Smooth Muscle
Singleunit Smooth Muscle:
-Synchronous activity (electrical/mechanical)
-Gap Junctions
-Some pacemaker cells
-Often activated by stretch (eg stomach, GI tract)
Multiunit Smooth Muscle:
-Few to no gap junctions
-cells respond as individuals
-Rich neural innervation (control)
(eg airways, large arteries)
NB: Most smooth muscles are a mix of these two extremes.
Cardiac Muscle
Cells joined by Intercalated discs:
-desmosomes
-gap junctions
-Striated (sarcomeres)
-Involuntary
-pacemaker cells
-T-Tubules
-Sarcoplasmic Reticulum
Cardiac Muscle: Excitation-Contraction Coupling
Depolarization
Opening of voltage gated
calcium channels
-L-Type Ca2+ Channel
(modified DHP receptor)
Triggers calcium induced
release of calcium (CIRC)
from S.R.
(ryanodine receptors)
Contraction as in skeletal
muscle fibers
- L Type channel
“Long-lasting current”
Critical for proper cardiac
functioning.
Skeletal vs. Cardiac Action Potentials
Characteristics of Muscle
