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Dr. Hoe See Ziau Department of Physiology Faculty of Medicine University of Malaya
Muscles in human body
Specialised excitable tissues ~ 50 % body weight Ability to contract Contractions provide movements Do work
Move body or limbs Push, pull or hold an external load or object Mix or move food through the gastrointestinal track Pump blood out of the heart to the blood vessels Contract uterus for birth of foetus Micturition and defaecation
Types of Muscle
Three types of muscle: 1. Skeletal muscle 2. Cardiac muscle 3. Smooth muscle
Types of Muscle Skeletal Muscle striated voluntary Cardiac Muscle striated involuntary Smooth Muscle non-striated involuntary .
Basic Characteristics of Muscle Tissues Excitability Response to stimuli Conductivity Able to conduct action potential Contractibility Able to shorten in length Extensibility Stretches when pulled Elasticity tends to return to original shape & length after contraction or extension .
Skeletal Muscle Attached to bones & moves skeleton Makes up 40% of BW in men and 32% of BW in women Main functions of skeletal muscle: Initiate movements Perform work Maintain posture Stabilise joints Generate heat .
Level of Organisation in Skeletal Muscle Skeletal Muscle (organ) (fascicle) (bundle of muscle fibres) Fascicle Muscle Fibre (cell) Myofibril Sarcomere (Thin – actin) (Thick .myosin) Filaments .
Membranes of Skeletal Muscle Muscle surrounded by epimysium Bundles of fibres (fascicles) surrounded by perimysium Muscle fibre surrounded by endomysium These connective tissues extend beyond the ends of muscle to form tendons that attach muscle to bones .
up – internal conduction system Myofibrils for contraction Sarcomeres – regular to 750.Skeletal Muscle Fibre Large. shape like Transverse tubules (T-tubules) cylinder 10 – 100 µm in diameter.75 m) in length (extend entire length of muscle) Multinucleated with arrangement of thin (actin) & thick (myosin) filaments Actin filaments interdigitate abundant of mitochondria Sarcolemma (cell membrane) Sarcoplasm (muscle cell with myosin filaments Appears striated under microscope cytoplasm) Sarcoplasmic reticulum (modified ER) .000 µm (0. elongated.
Structure of a Skeletal Muscle Fibre .
Electron Micrograph of Skeletal Muscle .
Sarcomere • The functional unit of skeletal muscle • Multi-protein complexes composed different filament systems: Thin filament system Thick filament system .
Sarcomere sarcomere .
Sarcomere Sarcomere .
and is the region where they do not overlap the thick filaments H zone The lighter area in the centre of A band where the thin filaments do not overlap with thick filaments M line Consists of supporting proteins that hold the thick filaments together vertically within each stack Z line Consists of supporting proteins that hold the thin filaments together vertically within each stack Area between two Z lines is called a sarcomere .Sarcomere A band (dark band) consists of a stacked set of thick filaments I band (light band) Consists of the array of thin filaments.
with a special binding site for attachment with myosin cross bridge Joined into two strands and twisted together to form the backbone of a thin filament Tropomyosin Threadlike proteins that lie end-to-end alongside the groove of the actin spiral Covers active sites of actin Troponin complex binds to actin & holds tropomyosin in place .Thin Filament Actin Spherical in shape.
Thin Filament .
Thin Filament Troponin Complex TnT – binds to tropomyosin TnC – binds to Ca2+ TnI – binds to actin .
Thick Filament Each thick filament is composed of several hundred myosin molecules packed together A single myosin protein looks like 2 golf clubs with shafts twisted about one another Myosin molecules have elongated tails & globular heads Heads form cross-bridges between thick and thin filaments during contraction .
Thick Filament Cross Bridges Each cross bridge has two important sites: An actin-binding site A myosin ATPase site .
Organisation of Actin and Myosin Thin filaments are arranged hexagonally around thick filaments Cross bridges Each thin filament is surrounded by 3 thick filaments Cross bridges project from each thick filament in all 6 directions toward the surrounding thin filaments .
Contraction of Muscle Fibres
Done by sliding actin filaments
Contraction of Muscle Fibres
Contraction of Muscle Fibres
Sliding Filament Theory
Contraction occurs by actin filaments sliding into
Actin filaments move, myosin filaments remain
Sarcomeres shortened Cause whole muscle to contract
moves tropomyosin & exposes actin active site .Contraction of Muscle Fibres Role of Calcium Ca2+ released from sarcoplasmic reticulum Ca2+ binds to troponin C Troponin turns.
form cross-bridge. move & produces powerful strokes Actin slides in – muscle fibre contracts Cross-bridge action continues while Ca2+ is present When action potential stops. Ca2+ is pumped back to SR Tropomyosin covers back actin’s active site Relaxation occurs .Contraction of Muscle Fibres Role of Calcium Myosin head binds to actin active site.
Contraction of Muscle Fibres Role of Calcium .
energy is stored within the cross bridge Mg2+ must be attached to ATP before ATPase 2 can split the ATP Ca2+ released on excitation. ADP and Pi remain attached to myosin. removes 1 3 inhibitory influence from actin → energised myosin cross bridge bind with actin Cross bridge bends and causes power stroke ADP and Pi are released after power stroke is completed 4 ATPase site is free for attachment of another ATP Attachment of new ATP permits detachment of cross bridge .Contraction of Muscle Fibres Role of ATP ATP split by myosin ATPase .
Contraction of Muscle Fibres .
Contraction of Muscle Fibres All the cross bridges’ power strokes are directed toward the centre of the sarcomere All 6 of the surrounding thin filaments on each end of the sarcomere are pulled inward simultaneously .
[Ca2+]i begins to rise This Ca2+ moves the regulatory proteins aside. actin and myosin remain bound in rigor complex Resulting in stiffness condition of dead muscles . permitting actin bind with the myosin cross bridges. which were already charged with ATP before death No fresh ATP available after death.Contraction of Muscle Fibres Rigor Mortis “Stiffness of death” – a generalised locking in place of skeletal muscle that begins 3 to 4 hours after death Following death.
Electrical Properties of Muscle Fibres Resting membrane potential: -90mV → action potential Maximum potential: +30mV Depolarisation is due to influx of Na+ Membrane potential (mV) When an adequate stimulus is given Time taken: 1 – 2 msec Absolute refractory period & relative -90 refractory period present Action potential results in muscle contraction .
Action Potential and Muscle Twitch Tension Latent period The delay between stimulation and the onset of contraction (a few msec) Contraction time The time from the onset of contraction until peak tension is developed (average ~ 50 msec) Relaxation time The time from peak tension until relaxation (~ 50 msec or more) A single contraction/relaxation -90 Membrane potential (mV) cycle is called a muscle twitch .
Excitation-Contraction Coupling Refers to the series of events linking muscle excitation (electrical events) to muscle contraction (mechanical events) Electrical events – presence of action potential Mechanical events – cross-bridge activity Electrical events come first before mechanical events Ca2+ is the link between excitation and contraction .
Excitation-Contraction Coupling Transverse Tubules (T tubules) The surface membrane at each junction of A band and I band dips into muscle fiber to form a T tubule Action potential on the surface membrane spreads down into the T tubule The presence of local action potential in T tubule induces permeability changes in the sarcoplasmic reticulum .
Excitation-Contraction Coupling Sarcoplasmic Reticulum (SR) Modified endoplasmic reticulum Consists of a fine network of interconnected compartments surrounding each myofibril Separate segments of SR are wrapped around each A band and each I band The ends of each segment expand to form lateral sacs. which store Ca2+ .
local depolarisation activates the voltage-gated dihydropyridine receptors in T tubule These activated receptors in turn trigger the opening of Ca2+release channels (alias ryanodine receptors) in adjacent lateral sacs of SR Ca2+ is released into the surrounding sarcoplasm .Excitation-Contraction Coupling Release of Ca2+ from SR When action potential is propagated down the T tubules.
Relaxation of Muscle Fibres When ACh is removed from the neuromuscular junction. the muscle fibre action potential ceases No longer a local potential in T tubules to trigger Ca2+ release Released Ca2+ is pumped back into the lateral sacs by Ca2+-ATPase pump Removal of sarcoplasmic Ca2+ allows the troponin-tropomyosin complex to slip back into its blocking position Actin and myosin are no longer able to bind at the cross bridges Thin filaments are able to return passively to their resting position Relaxation occurs .
Excitation-Contraction Coupling .
form cross-bride. When action potential stops. moves and produces power stroke 6. Ca2+ pumped back to SR 11. New ATP binds to myosin head. muscle fibre contracts resulting in contraction of whole muscle 7. detachment of the cross bridge 9. Causes SR to release Ca2+ into sarcoplasm 4. Ach released from the terminal of a motor neuron initiates an action potential in the muscle fibres 2. Actin slides in. Relaxation occurs . ADP and Pi are released after the power stroke is complete 8. Ca2+ binds to troponin. Cross-bridge action continues while Ca2+ is present 10. Tropomyosin covers back active sites 12.Excitation-Contraction Coupling and Relaxation Summary of Events 1. Myosin head binds to active site. exposing actin’s cross-bridge binding sites 5. Muscle action potential travels down T tubule 3.
Contraction of Whole Muscles Even when muscle are at rest. certain amount of tautness usually remain → muscle tone Results from a low rate of nerve impulses coming from the spinal cord To maintain a normal posture .
tension is created Gradation of whole muscle tension depends on The number of muscle fibres contracting within a muscle The tension developed by each contracting fibre .Contraction of Whole Muscles Whole muscles are groups of muscle fibres bundled together Muscle fibres in each muscle can function cooperatively to produce contractions of variable grades of strength When the whole muscles contract.
all the muscle fibres in that motor unit are stimulated to contract simultaneously Each muscle consists of a number of intermingled motor units .Motor Unit Each whole muscle is innervated by a number of different motor neurons One motor neuron innervates a number of muscle fibers Each muscle fiber is supplied by only one motor neuron A motor neuron plus all the muscle fibres it innervates is called a motor unit When a motor neuron is activated.
Motor Unit Precise control of movement determined by number and size of motor units The number of muscle fibres innervated by one motor neuron – innervation ratio The bigger the ratio of nerve to muscle fibres. the coarser the movement will be Examples: 1:4 – fine movements (external eye muscle) 1:200-300 – Coarse movements (back muscle) 1:150 – on average .
Motor Unit .
only a few of its motor units are activated For stronger and stronger contractions. more and more motor units are stimulated to contract → motor unit recruitment .Motor Unit Recruitment For a weak contraction of the whole muscle.
Motor Unit Recruitment At minimum stimulus strength only motor units with low threshold will contract At maximum stimulus strength all motor units contract ↑ strength of stimulus ↑ recruitment of motor units ↑ contraction .
Mechanical Properties of Skeletal Muscle Muscle Twitch A twitch is a single contraction/ relaxation cycle .
Mechanical Properties of Skeletal Muscle Muscle Twitch If the muscle has completely relax before the next stimulus takes place A second twitch of the same magnitude as the first occurs • When a muscle begins to contract. its initial strength of contraction may be as little as ½ of its strength 10 to 50 muscle twitches later • The strength of contraction increase to plateau (Treppe) Maximum tension (in treppe) .
the 2nd twitch is added on to the 1st twitch. resulting in summation Tetanus When the muscle is stimulated so rapidly that it does not have an opportunity to relax between stimuli. a maximal sustained contraction occurs → tetanus .Mechanical Properties of Skeletal Muscle Summation and Tetanus Summation If the muscle is restimulated before it has completely relaxed.
Mechanical Properties of Skeletal Muscle Summation and Tetanus When muscle is stimulated. Ca2+ is released from SR → cross- bridges → contraction When stimulation ceases. there is not enough time between successive stimulations to remove all the Ca2+ from the sarcoplasm → Ca2+ levels in the sarcoplasm increase → more active crossbridges → a stronger contraction → summation occurs . Ca2+ is pumped back into SR If the 2nd stimulation occur far enough apart in time for all the released Ca2+ from the 1st contractile response to be pumped back into SR → an identical twitch response occurs With rapid stimulation.
then become sustained → summation & tetanus . more & more motor unit will be activated → recruitment As frequency of stimulation increases gradually. contraction will increase more & more.Mechanical Properties of Skeletal Muscle Response of muscle to repeated stimulation As strength of stimulation increases gradually.
Mechanical Properties of Skeletal Muscle Muscle Fatigue When the stimulation is given repeatedly at a fast rate Contraction becomes weaker & weaker gradually Contraction becomes more irregular Until no contraction occur Fatigue occurs .
Mechanical Properties of Skeletal Muscle Fatigue Type Muscle fatigue Occurs when an exercising muscle can no longer respond to stimulation with the same degree of contractile activity Causes: Accumulation of lactic acid Depletion of energy stores supply of O2 and nutrients Central fatigue Occurs when the CNS no longer adequately activates the motor neurons supplying the working muscles Neuromuscular fatigue Depletion of acetylcholine .
Mechanical Properties of Skeletal Muscle Length-tension relationship At each determined muscle length: Without stimulation → passive tension With stimulation → total tension Active tension = total tension – passive tension .
Mechanical Properties of Skeletal Muscle Length-tension relationship In general. tetanic tension develop muscle length (within limit) initial For every muscle → an optimal length at which maximal force can be developed In the body. relaxed length of muscle are also the optimal length Capable of obtaining maximal tetanic contractions & maximal force .
Mechanical Properties of Skeletal Muscle Length-tension relationship .
Isotonic contraction Tension developed – constant Muscle length – changes For Body movement Moving an external load or object 2.Mechanical Properties of Skeletal Muscle Types of Contraction 1. Isometric contraction Muscle length – constant Tension developed – changes For Holding a load or object .
Mechanical Properties of Skeletal Muscle Isotonic Contraction The muscle length changes to move a load .
Mechanical Properties of Skeletal Muscle Isometric Contraction Tension in the muscle increases but the muscle fibres neither shortened or lengthened .
Mechanical Properties of Skeletal Muscle Isotonic Isometric .
Skeletal Muscle Metabolism Contraction-relaxation process requires ATP in three different steps: 1. Splitting of ATP by myosin ATPase provides energy for the power stroke of the cross bridge 2. The active transport of Ca2+ back into the SR during relaxation ATP must constantly be supplied for contractile activity to continue . Binding of fresh ATP molecule to myosin permits detachment of the bridge from actin filament at the end of power stroke 3.
Glycolysis Synthesis ATP in the absence of O2 Uses large amounts of stored glycogen and produces lactic acid in the process . Creatine phosphate Creatine phosphate + ADP Creatine + ATP First source for supplying additional ATP 2.Skeletal Muscle Metabolism Additional ATP is supplied by three pathways: 1. Oxidative phosphorylation Takes place within the muscle mitochondria if sufficient O2 is present Fueled by glucose and fatty acids Relatively slow because involves many steps 3.
Skeletal Muscle Metabolism .
Fast-oxidative (type IIa) fibres 3. Slow-oxidative (type I) fibres 2. Fast-glycolytic (type IIb) fibres .Types of Skeletal Muscle Fibres Three types of muscle fibres are classified by: The pathways they used for ATP synthesis Oxidative Glycolytic Fast Slow The Speed of their contraction 1.
Types of Skeletal Muscle Fibres Fast vs slow fibres Fast fibres have higher myosin ATPase (ATP-splitting) activity ATP is split More rapidly The rate at which energy is made available for cross-bridge cycling is faster Results in a fast twitch Fast fibres are activated by large-diameter motor neurons Slow fibres are activated by small-diameter motor neurons .
Types of Skeletal Muscle Fibres Oxidative vs glycolytic fibres Oxidative fibres have a greater capacity to form ATP More ATP is yielded from each nutrient molecule processed → does not readily deplete energy stores Does not result in lactic acid accumulation More resistant to fatigue • Oxidative fibres have a high myoglobin content → red fibres .
Types of Skeletal Muscle Fibres Most skeletal muscles contain a mixture of all three fibre types A single motor unit always contains one type or the other The percentage of each type determined by the type of activity for which the muscle is specialised .
Muscle Hypertrophy Enlargement (increase in diameter) of muscle Total mass of muscle increases Results from increased synthesis of actin and myosin filaments in each muscle fibre Occurs when the muscle undergoes regular bouts of anaerobic. high-intensity resistance training . short-duration.
Muscle Atrophy Muscle becomes smaller and weaker Total mass of muscle decreases Results from decrease of actin and myosin content Disuse atrophy Occurs when a muscle is not used for a long period of time even though the nerve supply is intact Denervation atrophy Occurs after the nerve supply to a muscle is lost .