Skeletal Muscle Fibers – Key Vocabulary
Sarcomere Structure & Length–Tension Relationship
- Skeletal muscle fiber ≡ skeletal muscle cell; functional contractile unit is the sarcomere (distance between two Z discs).
- Diagram details (verbal):
- Black zig-zag lines = Z discs.
- Thick filament: parallel arrays of myosin molecules; four myosin heads drawn per side (eight total) for illustration.
- Thin filament: actin; instructor drew one short strand of F-actin for simplicity, but physiologically there are two intertwined strands that span the entire half-sarcomere.
- H zone = region between the innermost ends of two adjacent thin filaments (i.e., bare region of thick filament that lacks actin overlap).
- During contraction the H zone narrows or disappears as thin filaments slide toward the M line.
- Optimal length–tension relationship:
- Achieved when all possible cross-bridges form (all myosin heads have an actin binding site available).
- Characterized by a “goldilocks” H-zone width: not too wide (over-stretched) and not absent (over-shortened).
- Produces maximum force because every myosin head contributes.
- Deviation scenarios:
- Over-stretched fiber
- H zone very wide; only the terminal myosin heads reach actin → few cross-bridges.
- Potential for large shortening exists but cannot be realized because most heads cannot attach.
- Over-shortened fiber
- Thin filaments overlap each other; H zone already zero.
- No further shortening possible → force production depressed.
ATP Sources for Skeletal Muscle Contraction
- Four sequential/parallel energy systems (listed in approximate order of utilization):
- Residual ATP in the cytosol
- Pre-synthesised; supports < 1 s of activity.
- Substrate-level phosphorylation (phosphagen system)
- Primary reaction: \text{Creatine~phosphate} + \text{ADP} \rightarrow \text{Creatine} + \text{ATP}
- “Borrowing/lending” a phosphate; fast, anaerobic.
- Glycolysis
- First stage of “cellular respiration”; can operate with or without O_2.
- Rapid ATP delivery; produces pyruvate.
• Without O_2 → pyruvate reduced to lactate (lactic-acid fermentation) → contributes to fatigue. - When combined with 1 & 2, can fuel ~1 min of maximal effort.
- Oxidative phosphorylation (mitochondrial ATP production)
- Requires O_2, mitochondria, electron-transport chain, ATP synthase, proton motive force, chemiosmosis.
- Slow to ramp up but yields orders of magnitude more ATP than systems 1–3.
Mechanical vs Electrical Events: Twitch, Summation & Tetanus
- Twitch = one contraction–relaxation mechanical cycle of a single muscle fiber.
- Initiated by an action potential (AP) generated by a somatic motor neuron.
- Electromechanical sequence:
- Motor neuron releases ACh → binds nicotinic receptors.
- End-plate potential depolarises sarcolemma → AP propagates.
- DHP receptor conformational change → opens RyR (ryanodine) channels in sarcoplasmic reticulum (SR).
- Ca^{2+} floods cytosol → binds troponin → tropomyosin shifts → cross-bridge cycling begins.
- Timing:
• AP duration ≈ 1\text{–}2\,\text{ms}.
• Twitch duration varies 7\text{–}70\,\text{ms} (fast- vs slow-twitch fibers).
• Latent period: brief delay between AP onset and force generation.
- Summation (temporal summation)
- Multiple APs fired before complete relaxation → SR re-opens or remains open → [Ca^{2+}] stays elevated.
- Result: larger, merged mechanical responses; decreases time available for relaxation.
- Tetanus
- Mechanical state of sustained maximal tension.
- Incomplete (unfused) tetanus: slight relaxation between stimuli; force oscillates.
- Complete (fused) tetanus: no relaxation; plateau of maximal force.
- Key distinction:
• Summation = description of the rapid electrical stimuli.
• Tetanus = resulting mechanical phenomenon.
Motor Unit Recruitment
- Motor unit = single somatic motor neuron + all skeletal muscle fibers it innervates (could be 3–>1000s fibers).
- Recruitment = activating additional motor units to increase whole-muscle force output.
- Light task (e.g., lifting an Expo® marker) → few small motor units engaged.
- Heavy task (e.g., 30-lb dumbbell curl) → progressively larger/more motor units activated.
- Provides coarse-to-fine control of force.
Mechanisms to Increase Muscle Force
- Optimal length–tension relationship (cross-bridge availability + H-zone width).
- Recruitment of additional motor units (spatial summation).
- Summation → tetanus (temporal summation; maintains cytosolic Ca^{2+} to keep cross-bridges cycling).
Skeletal Muscle Fiber Types: Slow-Twitch vs Fast-Twitch
- Type I (Slow-twitch)
- Twitch time ≈ \sim 70\,\text{ms}.
- Reliance on oxidative phosphorylation.
- High mitochondrial density, rich capillary supply, abundant myoglobin → dark meat coloration.
- Smaller fiber diameter; typical of endurance athletes (e.g., distance runners).
- Type II (Fast-twitch)
- Twitch time ≈ \sim 7\,\text{ms}.
- Predominantly glycolytic; ATP produced anaerobically → rapid availability.
- Larger diameter; high concentration of glycolytic enzymes; fewer mitochondria & capillaries → white meat coloration.
- Contain faster ATPases:
• Myosin ATPase (cross-bridge cycling) & Ca²⁺-ATPase (SR calcium pump) work at higher rates, enabling rapid contraction/relaxation.
- (Intermediate/Type IIa fibers exist but were omitted for brevity.)
Integrative/Real-World Connections & Implications
- Stretching beyond optimal sarcomere length before heavy lifts can decrease force output despite feeling “loose.”
- Phosphagen & glycolytic systems dominate early in high-intensity sports (sprints, Olympic lifts), whereas oxidative phosphorylation powers long-duration events (marathons).
- Muscle tetanus in pathology (e.g., tetanus toxin) arises from spinal inhibition failure, not from voluntary summation, but the mechanical outcome—prolonged contraction—is analogous.
- Training adaptations:
• Resistance training ↑ fiber diameter (especially Type II) and can shift Type IIx → IIa phenotype.
• Endurance training ↑ mitochondrial biogenesis & capillarisation in Type I fibers. - Creatine supplementation targets the phosphagen system, enhancing substrate-level phosphorylation capacity (useful for repeated short bursts).