Muscle-Bound Notes
General Notes
· Anything that says myo, mys or sarco is referring to muscles
· muscle cells are elongated, hence the name muscle fibers
· muscle contraction depends on myofilaments
· of the three muscle types, we will be concentrating on skeletal muscles
Functions of the Muscle System
· movement
· maintains posture
· joint stabilization
· heat generation
Functional Characteristics of Muscles : distinctive characteristics of muscle tissue that allow it to perform its duties
· excitability (irritability)
· contractility
· extensability
· elasticity
Gross Anatomy of a Skeletal Muscle
· each muscle fiber is surrounded by a fine layer of areolar (loose) tissue called the endomysium
· several endomysium-wrapped muscle fibers are joined together in a bundle called a fascicle which is
· bound together by a dense fibrous tissue sheath called the perimysium
· fascicles are bundled together into the muscle itself, held together by the dense fibrous epimysium.
· superficial to the epimysium is another sheath of connective tissue called the fascia or muscle belly.
· all of these sheaths are continuous with one another, so that when muscle fibers contract, the forces are transmitted throughout the muscle.
· each muscle fiber is supplied with a nerve ending to control its activity
· each muscle is served by an artery and one or more veins
· attachments
· most muscles span joints and are attached to bones in at least two places
· when a muscle contracts, the muscle's insertion moves toward the muscle's origin
· muscle attachments are either direct or indirect
· direct attachments (fleshy) : the epimysium is fused to the periosteum or perichondrium
· indirect attachments : the connective tissue wrappings extend beyond the muscle as a tendon or aponeurosis
· tendon : connects the muscle to a skeletal element
· aponeurosis : connects the muscle to the fascia of other muscles
Microscopic Anatomy of a Muscle Fiber
· each skeletal muscle cell is a long cylindrical, multinucleated cell, a syncytium of hundreds of embryonic cells
· the sarcolemma is the plasma membrane of a muscle fiber
· sarcoplasm is similar to cytoplasm, but it has higher amounts of stored glycogen and a protein called myoglobin which binds oxygen
· sarcoplasmic reticulum is a specialized smooth endoplasmic reticulum of a muscle cell; it regulates the intracellular Ca2+ in the cell
· transverse (or "t") tubules are hollow, elongated tubes where the sarcolemma penetrates into the cell's interior; they help to evenly distribute the muscle impulse
· myofibrils are rodlike structures that are organized into sarcomeres
Sarcomeres and Muscle Contraction
· a sarcomere, or "unit of muscle," is a contractile unit and is defined as the region between two "z" lines
· a z-line is a midline interruption of an i-band; it is a coin-shaped protein that anchors thin (actin) filaments and connects each myofibril to the next
· the i-band is a light band in a striated muscle; it is light because only actin (thin) filaments are found in that area
· an a-band is the dark band in striated muscle; it is dark because of the thick filaments there
· the h-zone is a slightly lighter area in the middle of the a-band; it is lighter because it is the point where there is no overlap of thin and thick filaments; they are only visible in relaxed muscle for that reason
· in the middle of the h-zone is the m-line, which is a dark band where the thick filaments are joined
· the thin filament -- the actin filament -- is actually made up of three proteins: G-actin, tropomyosin and troponin
· G-actin contains the active sites to which the myosin heads will bind
· G-actin "beads" join together to form F-actin
· two strands of F-actin coil around each other
· two strands of tropomyosin coil around the two strands of F-actin; they block the active sites during relaxation
· troponin, a three-polypeptide complex, binds to actin and to tropomyosin, and helps to control the positioning of tropomyosin
· the thick filament -- the myosin filament -- has two distinctive features: the myosin tail and the myosin head or cross bridge
· the cross bridge is named for the fact that it links actin and myosin during contraction
· the myosin heads contain ATP binding sites and ATPase enzymes to generate the energy needed during contraction
Sliding Filament Mechanism
· when a muscle contracts, the individual sarcomeres shorten, but the filaments do not
· the sliding filament theory of contraction was first proposed in 1954 by Hugh Huxley
· he said that thin filaments slide past the thick ones so that they overlap to a greater degree during contraction
· during contraction the z-lines move closer together, the h-zone disappears, and the a-bands gets closer while unchanging in their length but the i-band gets smaller
· the cause of sliding filaments
· when intracellular levels of calcium are low, the muscle is relaxed and the active sites on the thin filaments are blocked
· when calcium ions become available, they bind to troponin, moving the tropomyosin molecule and exposing the active sites and the following occurs in rapid succession:
· cross bridges attach
· power stroke
· cross bridge detachment
· "cocking" of the myosin head
· sliding of the filaments occurs as long as there are calcium ions available
· as the SR pumps the calcium back, and the active sites are blocked, relaxation occurs
· the sliding filament mechanism is the best accepted model of muscle contraction, but there is still some controversy
· actin has ATPase activity and undergoes conformation changes, so what is their true role?
· how many ATP molecules are used? one per power stroke?
· rigor mortis illustrates how cross bridge detachment is ATP driven
· muscles stiffen 3-4 hours after death (due to release of calcium ions from the SR)
· peak rigidity is about 12 hours after death
· 48-60 hours after death, muscles have relaxed again -- because the actin and myosin are being broken down
· at death, ATP is no longer being made, and so when the actin and myosin bind, they can't be detached without ATP
· regulation of muscle contraction
· skeletal muscle cells are stimulated by motor neurons of the somatic nervous system which form a neuromuscular junction
· the neuromusclar junction is not a direct connection, rather it is a very close association separated by a synaptic cleft
· the motor end plate is a highly folded part of the sarcolemma that has a great number of ACh receptors in it.
· when a nerve impulse reaches the junction, ACh (acetylcholine) is released into the synaptic cleft, fusing with the receptors in the motor end plate, causing a muscle impulse, similar to a nerve impulse
· the muscle impulse is the result of depolarizing the sarcolemma
· the ACh is rapidly destroyed after being taken up by the receptors in order to allow another muscle impulse; acetylcholinesterase is responsible for this
· a shortage of acetylcholine receptors and blood containing antibodies to ACh receptors can result in myasthenia gravis
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· the muscle impulse moves along the sarcolemma, including down into the muscle itself through the t-tubules
· the impulse triggers the release of calcium ions from the SR into the sarcoplasm
· the released calcium ions bind to troponin, allowing the binding of cross bridges to the now-exposed actin active sites
· calcium is shuttled back into the SR through a continuously active ATP-dependent calcium pump
· when intracellular calcium levels drop too low to allow contraction, the tropomyosin blockage is reestablished and relaxation occurs
Skeletal Muscle Contraction
· the motor unit is one motor nerve and the hundreds of muscle fibers that it services
· when a motor neuron fires, all of the fibers it supplies are stimulated as above
· a motor unit may have as many as several hundred fibers associated with it, or as few as 4
· the more finely controlled muscles, such as those controlling the fingers or eyes, have fewer fibers in each motor unit
· the fibers of a motor unit are not clustered together, but spread throughout the muscle, so stimulation of a single motor unit causes a weak contraction of the entire muscle
· the response of a muscle to a single, brief, threshold stimulus is a muscle twitch
· a muscle twitch produces a brief spike on a myogram
· it may be strong or weak depending on the number of motor units involved
· the muscle contracts quickly and then relaxes
· this is not the way our muscles work, not normally
· graded responses -- variation in the degree of muscle contraction -- are how our muscles work
· two ways of grading muscle contraction in vivo : changing the speed of contraction and changing the number of motor units activated
· if two identical stimuli are delivered to a muscle in rapid succession, the second twitch will be stronger than the first, and on a myogram, it will appear to "ride" on the shoulders of the first contraction -- this is wave summation, and occurs when the muscle is not completely relaxed before being stimulated again
· if the stimulus is held constant and the relaxation time between twitches becomes shorter and shorter, resulting in a "disappearance" of relaxation on a myogram, the sustained contraction is called tetanus
· tetanus is a reflection of how muscle contraction typically works in the body because motor neurons deliver a volley of impulses rather than just one impulse at a time
· tetanus can not be sustained indefinitely -- eventually the muscle will become fatigued and unable to contract any longer
· recruitment is how a muscle can call more than one motor unit into contraction, resulting in multiple motor unit summation
· threshold stimulus is the stimulus that results in the first observable muscle contraction; maximal stimulus is the strongest stimulus that results in increasing contractile force and represents the point where all of the motor units are engaged
· muscle tone is the state of semi-contraction that all muscles are in; it keeps the muscles firm, healthy and ready to act, as well as stabilizing joints and maintain posture
· isotonic and isometric contractions
· sometimes a muscle contracts but does not shorten
· isotonic contraction involves shortening the muscle fiber and doing work (concentric); or lengthening the muscle fiber (eccentric)
· isometric contractions increase tension, but the muscle no longer changes length
· posture maintaining muscles work isometrically
· isometric contractions occur when a muscle attempts to move a load greater than the force it is able to develop
· muscle fatigue : the muscle is physiologically unable to contract; this results from a relative deficit of ATP or an excessive accumulation of lactic acid or ionic imbalances
· muscle fiber types
· red slow -- thin cells that contain slow-acting myosin ATPases; have a lot of myoglobin (hence their red color), abundant mitochondria, rich capillary supply; oxygen dependent, very fatigue resistant and have high endurance, but don't generate much power
· white fast -- large pale cells with little myoglobin, contain fast-acting myosin ATPases and contract rapidly; contain few mitochondria, large glycogenic reserves, tire quickly, but very powerful
· red fast -- also called intermediate fibers or pink fibers, have fast-acting myosin ATPases, and contract quickly, but are more like red slow in that they are oxygen dependent, have a high myoglobin content and fatigue slower than the white but faster than the red slow