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Muscle contraction

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increases, eventually reaching zero at some maximum velocity. The reverse holds true for when the muscle is stretched – force increases above isometric maximum, until finally reaching an absolute maximum. This intrinsic property of active muscle tissue plays a role in the active damping of joints that are actuated by simultaneously active opposing muscles. In such cases, the force-velocity profile enhances the force produced by the lengthening muscle at the expense of the shortening muscle. This favoring of whichever muscle returns the joint to equilibrium effectively increases the damping of the joint. Moreover, the strength of the damping increases with muscle force. The motor system can thus actively control joint damping via the simultaneous contraction (co-contraction) of opposing muscle groups.
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without any corresponding changes in muscle length, the muscle contraction is described as isometric. If the muscle length changes while muscle tension remains the same, then the muscle contraction is isotonic. In an isotonic contraction, the muscle length can either shorten to produce a concentric contraction or lengthen to produce an eccentric contraction. In natural movements that underlie locomotor activity, muscle contractions are multifaceted as they are able to produce changes in length and tension in a time-varying manner. Therefore, neither length nor tension is likely to remain constant when the muscle is active during locomotor activity.
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period, there is a rapid burst of energy use as measured by oxygen consumption. Within a few minutes of initiation, the calcium level markedly decreases, the 20 kDa myosin light chains' phosphorylation decreases, and energy use decreases; however, force in tonic smooth muscle is maintained. During contraction of muscle, rapidly cycling crossbridges form between activated actin and phosphorylated myosin, generating force. It is hypothesized that the maintenance of force results from dephosphorylated "latch-bridges" that slowly cycle and maintain force. A number of kinases such as
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end-plate potential. The voltage-gated ion channels of the sarcolemma next to the end plate open in response to the end plate potential. They are sodium and potassium specific and only allow one through. This wave of ion movements creates the action potential that spreads from the motor end plate in all directions. If action potentials stop arriving, then acetylcholine ceases to be released from the terminal bouton. The remaining acetylcholine in the synaptic cleft is either degraded by active
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response is contraction. Excitation–contraction coupling can be dysregulated in many diseases. Though excitation–contraction coupling has been known for over half a century, it is still an active area of biomedical research. The general scheme is that an action potential arrives to depolarize the cell membrane. By mechanisms specific to the muscle type, this depolarization results in an increase in cytosolic
678: 1395:(REEP5), functions to keep the normal morphology of junctional SR. Defects of junctional coupling can result from deficiencies of either of the two proteins. During the process of calcium-induced calcium release, RyR2s are activated by a calcium trigger, which is brought about by the flow of Ca through the L-type calcium channels. After this, cardiac muscle tends to exhibit 633:. The activated dihydropyridine receptors physically interact with ryanodine receptors to activate them via foot processes (involving conformational changes that allosterically activates the ryanodine receptors). As ryanodine receptors open, Ca is released from the sarcoplasmic reticulum into the local junctional space and diffuses into the bulk cytoplasm to cause a 1668:
animal moves forward. As the front end of the earthworm becomes anchored and the circular muscles in the anterior segments become relaxed, a wave of longitudinal muscle contractions passes backwards, which pulls the rest of animal's trailing body forward. These alternating waves of circular and longitudinal contractions is called
500:. It is the site in which a motor neuron transmits a signal to a muscle fiber to initiate muscle contraction. The sequence of events that results in the depolarization of the muscle fiber at the neuromuscular junction begins when an action potential is initiated in the cell body of a motor neuron, which is then propagated by 1017:
the action of the muscle itself or by an outside force), the maximum active tension generated decreases. This decrease is minimal for small deviations, but the tension drops off rapidly as the length deviates further from the ideal. Due to the presence of elastic proteins within a muscle cell (such as
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by maintaining turgidity of the earthworm. When the circular muscles in the anterior segments contract, the anterior portion of animal's body begins to constrict radially, which pushes the incompressible coelomic fluid forward and increasing the length of the animal. As a result, the front end of the
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Unlike single-unit smooth muscle cells, multiunit smooth muscle cells are found in the muscle of the eye and in the base of hair follicles. Multiunit smooth muscle cells contract by being separately stimulated by nerves of the autonomic nervous system. As such, they allow for fine control and gradual
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because the number of contractions in these muscles do not correspond (or synchronize) with the number of action potentials. For example, a wing muscle of a tethered fly may receive action potentials at a frequency of 3 Hz but it is able to beat at a frequency of 120 Hz. The high frequency
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The contractile activity of smooth muscle cells can be tonic (sustained) or phasic (transient) and is influenced by multiple inputs such as spontaneous electrical activity, neural and hormonal inputs, local changes in chemical composition, and stretch. This is in contrast to the contractile activity
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Force–velocity relationship: right of the vertical axis concentric contractions (the muscle is shortening), left of the axis eccentric contractions (the muscle is lengthened under load); power developed by the muscle in red. Since power is equal to force times velocity, the muscle generates no power
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Mitochondria also participate in Ca reuptake, ultimately delivering their gathered Ca to SERCA for storage in the sarcoplasmic reticulum. A few of the relaxation mechanisms (NCX, Ca2+ pumps and Ca2+ leak channels) move Ca2+ completely out of the cells as well. As Ca concentration declines to resting
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reverses polarity and its voltage quickly jumps from the resting membrane potential of -90mV to as high as +75mV as sodium enters. The membrane potential then becomes hyperpolarized when potassium exits and is then adjusted back to the resting membrane potential. This rapid fluctuation is called the
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slide past each other, pulling the Z-lines together. During an eccentric contraction, the myofilaments slide past each other the opposite way, though the actual movement of the myosin heads during an eccentric contraction is not known. Exercise featuring a heavy eccentric load can actually support a
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the joint at the end of a movement or otherwise control the repositioning of a load. This can occur involuntarily (e.g., when attempting to move a weight too heavy for the muscle to lift) or voluntarily (e.g., when the muscle is 'smoothing out' a movement or resisting gravity such as during downhill
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cells: autorhythmic and contractile. Autorhythmic cells do not contract, but instead set the pace of contraction for other cardiac muscle cells, which can be modulated by the autonomic nervous system. In contrast, contractile muscle cells (cardiomyocytes) constitute the majority of the heart muscle
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are sent to muscle fibers. Action potentials do not arrive at muscles synchronously, and, during a contraction, some fraction of the fibers in the muscle will be firing at any given time. In a typical circumstance, when humans are exerting their muscles as hard as they are consciously able, roughly
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Length-tension relationship relates the strength of an isometric contraction to the length of the muscle at which the contraction occurs. Muscles operate with greatest active tension when close to an ideal length (often their resting length). When stretched or shortened beyond this (whether due to
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increases, more motor units are excited in addition to larger ones, with the largest motor units having as much as 50 times the contractile strength as the smaller ones. As more and larger motor units are activated, the force of muscle contraction becomes progressively stronger. A concept known as
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one to two days after training. Exercise that incorporates both eccentric and concentric muscular contractions (i.e., involving a strong contraction and a controlled lowering of the weight) can produce greater gains in strength than concentric contractions alone. While unaccustomed heavy eccentric
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The cytoplasmic calcium binds to Troponin C, moving the tropomyosin complex off the actin binding site allowing the myosin head to bind to the actin filament. From this point on, the contractile mechanism is essentially the same as for skeletal muscle (above). Briefly, using ATP hydrolysis, the
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When the desired motion is accomplished, relaxation can be achieved quickly through numerous pathways. Relaxation is quickly achieved through a Ca buffer with various cytoplasmic proteins binding to Ca with very high affinity. These cytoplasmic proteins allow for quick relaxation in fast twitch
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Muscle contractions can be described based on two variables: force and length. Force itself can be differentiated as either tension or load. Muscle tension is the force exerted by the muscle on an object whereas a load is the force exerted by an object on the muscle. When muscle tension changes
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Termination of crossbridge cycling (and leaving the muscle in latch-state) occurs when myosin light chain phosphatase removes the phosphate groups from the myosin heads. Phosphorylation of the 20 kDa myosin light chains correlates well with the shortening velocity of smooth muscle. During this
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In 1952, the term excitation–contraction coupling was coined to describe the physiological process of converting an electrical stimulus to a mechanical response. This process is fundamental to muscle physiology, whereby the electrical stimulus is usually an action potential and the mechanical
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Eccentric contractions normally occur as a braking force in opposition to a concentric contraction to protect joints from damage. During virtually any routine movement, eccentric contractions assist in keeping motions smooth, but can also slow rapid movements such as a punch or throw. Part of
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Force–velocity relationship relates the speed at which a muscle changes its length (usually regulated by external forces, such as load or other muscles) to the amount of force that it generates. Force declines in a hyperbolic fashion relative to the isometric force as the shortening velocity
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release the neurotransmitters epinephrine and norepinephrine, which bind to adrenergic receptors that are also metabotropic. The exact effects on the smooth muscle depend on the specific characteristics of the receptor activated—both parasympathetic input and sympathetic input can be either
742:, thereby allowing myosin to bind to another actin molecule. Once attached, the ATP is hydrolyzed by myosin, which uses the released energy to move into the "cocked position" whereby it binds weakly to a part of the actin binding site. The remainder of the actin binding site is blocked by 792:. Myosin then releases ADP but still remains tightly bound to actin. At the end of the power stroke, ADP is released from the myosin head, leaving myosin attached to actin in a rigor state until another ATP binds to myosin. A lack of ATP would result in the rigor state characteristic of 783:
complex causes tropomyosin to slide over and unblock the remainder of the actin binding site. Unblocking the rest of the actin binding sites allows the two myosin heads to close and myosin to bind strongly to actin. The myosin head then releases the inorganic phosphate and initiates a
103:. A single motor neuron is able to innervate multiple muscle fibers, thereby causing the fibers to contract at the same time. Once innervated, the protein filaments within each skeletal muscle fiber slide past each other to produce a contraction, which is explained by the 254:
In eccentric contraction, the tension generated while isometric is insufficient to overcome the external load on the muscle and the muscle fibers lengthen as they contract. Rather than working to pull a joint in the direction of the muscle contraction, the muscle acts to
1659:, circular and longitudinal muscles cells form the body wall of these animals and are responsible for their movement. In an earthworm that is moving through a soil, for example, contractions of circular and longitudinal muscles occur reciprocally while the 620:
Excitation–contraction coupling (ECC) occurs when depolarization of skeletal muscles (usually through neural innervation) results in a muscle action potential. This action potential spreads across the muscle's surface and into the muscle fiber's network of
1021:) and extracellular matrix, as the muscle is stretched beyond a given length, there is an entirely passive tension, which opposes lengthening. Combined, there is a strong resistance to lengthening an active muscle far beyond the peak of active tension. 59:, muscle contraction does not necessarily mean muscle shortening because muscle tension can be produced without changes in muscle length, such as when holding something heavy in the same position. The termination of muscle contraction is followed by 709:
in 1954. Physiologically, this contraction is not uniform across the sarcomere; the central position of the thick filaments becomes unstable and can shift during contraction but this is countered by the actions of the elastic myofilament of
1244:. Unlike skeletal muscle cells, smooth muscle cells lack troponin, even though they contain the thin filament protein tropomyosin and other notable proteins – caldesmon and calponin. Thus, smooth muscle contractions are initiated by the 934:. A twitch is a single contraction and relaxation cycle produced by an action potential within the muscle fiber itself. The time between a stimulus to the motor nerve and the subsequent contraction of the innervated muscle is called the 1355:
In both skeletal and cardiac muscle excitation-contraction (E-C) coupling, depolarization conduction and Ca release processes occur. However, though the proteins involved are similar, they are distinct in structure and regulation. The
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levels, Ca2+ releases from Troponin C, disallowing cross bridge-cycling, causing the force to decline and relaxation to occur. Once relaxation has fully occurred, the muscle is able to contract again, thus fully resetting the cycle.
609:), while the RyRs reside across the SR membrane. The close apposition of a transverse tubule and two SR regions containing RyRs is described as a triad and is predominantly where excitation–contraction coupling takes place. 836:
is no longer present on the thin filament, the tropomyosin changes conformation back to its previous state so as to block the binding sites again. The myosin ceases binding to the thin filament, and the muscle relaxes. The
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Asynchronous muscles power flight in most insect species. a: Wings b: Wing joint c: Dorsoventral muscles power the upstroke d: Dorsolongitudinal muscles (DLM) power the downstroke. The DLMs are oriented out of the page.
318:) is nevertheless consumed, although less than would be consumed during a concentric contraction of the same force. For example, one expends more energy going up a flight of stairs than going down the same flight. 216:
In concentric contraction, muscle tension is sufficient to overcome the load, and the muscle shortens as it contracts. This occurs when the force generated by the muscle exceeds the load opposing its contraction.
130:(meaning that they are initiated by the smooth or heart muscle cells themselves instead of being stimulated by an outside event such as nerve stimulation), although they can be modulated by stimuli from the 1625:
concentration allows the troponin complex to dissociate from the actin filament thereby ending contraction. The heart relaxes, allowing the ventricles to fill with blood and begin the cardiac cycle again.
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sparks gives a cell-wide increase in cytoplasmic calcium concentration. The increase in cytosolic calcium following the flow of calcium through the cell membrane and sarcoplasmic reticulum is moderated by
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through a feedback loop with the grey matter. Other actions such as locomotion, breathing, and chewing have a reflex aspect to them: the contractions can be initiated either consciously or unconsciously.
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at either isometric force (due to zero velocity) or maximal velocity (due to zero force). The optimal shortening velocity for power generation is approximately one-third of maximum shortening velocity.
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of skeletal muscle cells, which relies on a single neural input. Some types of smooth muscle cells are able to generate their own action potentials spontaneously, which usually occur following a
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because they contain myofibrils that are thick and conspicuous. A remarkable feature of these muscles is that they do not require stimulation for each muscle contraction. Hence, they are called
145:, muscle contractions are multifaceted as they are able to produce changes in length and tension in a time-varying manner. Therefore, neither length nor tension is likely to remain the same in 1688:, possess obliquely striated muscles, which contain bands of thick and thin filaments that are arranged helically rather than transversely, like in vertebrate skeletal or cardiac muscles. In 987:
the size principle, allows for a gradation of muscle force during weak contraction to occur in small steps, which then become progressively larger when greater amounts of force are required.
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If another muscle action potential were to be produced before the complete relaxation of a muscle twitch, then the next twitch will simply sum onto the previous twitch, thereby producing a
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if muscle tension remains the same throughout the contraction. If the muscle length shortens, the contraction is concentric; if the muscle length lengthens, the contraction is eccentric.
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Enoka, Roger M.; Pearson, Keir G. (2013). "The motor unit and muscle action". In Kandel, Eric R.; Schwartz, James H.; Jessell, Thomas M.; Siegelbaum, Steven A.; Hudspeth, A. J. (eds.).
2364: 224:. This occurs throughout the length of the muscle, generating a force at the origin and insertion, causing the muscle to shorten and changing the angle of the joint. In relation to the 3270:
Guo, Ang; Zhang, Xiaoying; Iyer, Venkat Ramesh; Chen, Biyi; Zhang, Caimei; Kutschke, William J.; Weiss, Robert M.; Franzini-Armstrong, Clara; Song, Long-Sheng (19 August 2014).
1066:. Single-unit smooth muscle cells can be found in the gut and blood vessels. Because these cells are linked together by gap junctions, they are able to contract as a functional 693:
to contract. It is a cycle of repetitive events that cause a thin filament to slide over a thick filament and generate tension in the muscle. It was independently developed by
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Saladin, Kenneth S., Stephen J. Sullivan, and Christina A. Gan. Anatomy & Physiology: The Unity of Form and Function. 7th ed. New York: McGraw-Hill Education, 2015. Print.
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Finally, if the frequency of muscle action potentials increases such that the muscle contraction reaches its peak force and plateaus at this level, then the contraction is a
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Wei, Sheng; Guo, Ang; Chen, Biyi; Kutschke, William; Xie, Yu-Ping; Zimmerman, Kathy; Weiss, Robert M.; Anderson, Mark E.; Cheng, Heping; Song, Long-Sheng (20 August 2010).
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Saladin, Kenneth S., Stephen J. Sullivan, and Christina A. Gan. (2015). Anatomy & Physiology: The Unity of Form and Function. 7th ed. New York: McGraw-Hill Education.
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The mechanism for muscle contraction evaded scientists for years and requires continued research and updating. The sliding filament theory was independently developed by
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greater weight (muscles are approximately 40% stronger during eccentric contractions than during concentric contractions) and also results in greater muscular damage and
208:, the tension in the muscle remains constant despite a change in muscle length. This occurs when a muscle's force of contraction matches the total load on the muscle. 2400:"A randomised clinical trial of the efficacy of drop squats or leg extension/leg curl exercises to treat clinically diagnosed jumper's knee in athletes: pilot study" 962:
one-third of the fibers in each of those muscles will fire at once, though this ratio can be affected by various physiological and psychological factors (including
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between the motor neuron terminal and the neuromuscular junction of the skeletal muscle fiber. Acetylcholine diffuses across the synapse and binds to and activates
1240:(kDa) myosin light chains on amino acid residue-serine 19, enabling the molecular interaction of myosin and actin, and initiating contraction and activating the 115:. In skeletal muscles, muscle tension is at its greatest when the muscle is stretched to an intermediate length as described by the length-tension relationship. 1692:, the obliquely striated muscles can maintain tension over long periods without using too much energy. Bivalves use these muscles to keep their shells closed. 3868: 642: 2447:
Tassinary; Cacioppo (2000). "The Skeletomotor system: surface electromyography". In Cacioppo, John T.; Tassinary, Luois G.; Berntson, Gary G. (eds.).
2384: 3997: 1524:, which bind a large proportion of intracellular calcium. As a result, a large increase in total calcium leads to a relatively small rise in free 3642:
Crespo LM, Grantham CJ, Cannell MB (June 1990). "Kinetics, stoichiometry and role of the Na-Ca exchange mechanism in isolated cardiac myocytes".
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Cheng H, Lederer WJ, Cannell MB (October 1993). "Calcium sparks: elementary events underlying excitation-contraction coupling in heart muscle".
2800:"The positional stability of thick filaments in activated skeletal muscle depends on sarcomere length: evidence for the role of titin filaments" 1768:
discovered that the muscles of dead frogs' legs twitched when struck by an electrical spark. This was one of the first forays into the study of
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more than training with concentric contractions alone. However, exercise-induced muscle damage is also greater during lengthening contractions.
1441:. Although this Ca influx only count for about 10% of the Ca needed for activation, it is relatively larger than that of skeletal muscle. This 734:
is a myosin projection, consisting of two myosin heads, that extends from the thick filaments. Each myosin head has two binding sites: one for
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ions from the troponin. Thus, the tropomyosin-troponin complex again covers the binding sites on the actin filaments and contraction ceases.
665:(SERCA) actively pumps Ca back into the sarcoplasmic reticulum, resulting in a permanent relaxation until the next action potential arrives. 2747:
Huxley H, Hanson J (1954). "Changes in the cross-striations of muscle during contraction and stretch and their structural interpretation".
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Yao, Lei; Xie, Duanyang; Geng, Li; Shi, Dan; Huang, Jian; Wu, Yufei; Lv, Fei; Liang, Dandan; Li, Li; Liu, Yi; Li, Jun (3 February 2018).
1554:(SERCA) pump back into the sarcoplasmic reticulum ready for the next cycle to begin. Calcium is also ejected from the cell mainly by the 427:
because of their striped appearance under a microscope, which is due to the highly organized alternating pattern of A bands and I bands.
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An isometric contraction of a muscle generates tension without changing length. An example can be found when the muscles of the hand and
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For the contractions to happen, the muscle cells must rely on the change in action of two types of filaments: thin and thick filaments.
1376:(main RyR isoform in cardiac muscle) are not physically coupled in cardiac muscle, but face with each other by a junctional coupling. 4036: 2693:
Huxley AF, Niedergerke R (1954). "Structural Changes in Muscle During Contraction: Interference Microscopy of Living Muscle Fibres".
3272:"Overexpression of junctophilin-2 does not enhance baseline function but attenuates heart failure development after cardiac stress" 1577:
does not have to leave the cell entirely. At high heart rates, phospholamban is phosphorylated and deactivated thus taking most
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Excitation-contraction coupling in cardiac muscle cells occurs when an action potential is initiated by pacemaker cells in the
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exercise) as compared to concentric loading. When eccentric contractions are used in weight training, they are normally called
714:. This fine myofilament maintains uniform tension across the sarcomere by pulling the thick filament into a central position. 3781: 3705: 3126:"Tonic and phasic smooth muscle contraction is not regulated by the PKCα - CPI-17 pathway in swine stomach antrum and fundus" 2886: 2861: 2674: 2456: 2133: 2079: 2043: 1999: 1971: 1946: 1566:, serves as a brake for SERCA. At low heart rates, phospholamban is active and slows down the activity of the ATPase so that 1780:
that is called a calcium transient. This increase in calcium activates calcium-sensitive contractile proteins that then use
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to contract. In skeletal muscles, excitation–contraction coupling relies on a direct coupling between two key proteins, the
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Faulkner, JA (2003). "Terminology for contractions of muscles during shortening, while isometric, and during lengthening".
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Muscle contraction can also be described in terms of two variables: length and tension. In natural movements that underlie
17: 1321:, or G-protein coupled receptors that initiate a second messenger cascade. Conversely, postganglionic nerve fibers of the 240:
would change the angle of the joint in the opposite direction, straightening the arm and moving the hand towards the leg.
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ions into the sarcoplasmic reticulum creates a deficiency in the fluid around the myofibrils. This causes the removal of
796:. Once another ATP binds to myosin, the myosin head will again detach from actin and another cross-bridge cycle occurs. 731: 447:
contractions occur as a result of signals originating in the brain. The brain sends electrochemical signals through the
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if the muscle tension changes but the muscle length remains the same. In contrast, a muscle contraction is described as
2481: 1314: 379:(also known as jumper's knee or patellar tendonosis) have been shown to benefit from high-load eccentric contractions. 3443:"REEP5 (Receptor Accessory Protein 5) Acts as a Sarcoplasmic Reticulum Membrane Sculptor to Modulate Cardiac Function" 1301:
Although smooth muscle contractions are myogenic, the rate and strength of their contractions can be modulated by the
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during baseball involves reducing eccentric braking allowing a greater power to be developed throughout the movement.
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binding to the troponin complex that regulates myosin binding sites on actin like in skeletal and cardiac muscles.
2881:. Academic Press Series in Biomedical Engineering (1st ed.). New York, NY: Academic Press. pp. 239–248. 371:
Eccentric contractions are being researched for their ability to speed rehabilitation of weak or injured tendons.
4407: 4113: 1823: 1490: 1380: 1070:. Single-unit smooth muscle cells contract myogenically, which can be modulated by the autonomic nervous system. 80:. Together, these two filaments form myofibrils - the basic functional organelles in the skeletal muscle system. 4334: 1310: 88: 2365:"Effectiveness of physical therapy for Achilles tendinopathy: An evidence based review of eccentric exercises" 1542: 582: 4118: 4029: 2222:
Colliander EB, Tesch PA (1990). "Effects of eccentric and concentric muscle actions in resistance training".
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Unlike skeletal muscle, E-C coupling in cardiac muscle is thought to depend primarily on a mechanism called
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which generates a force of 2 pN. The power stroke moves the actin filament inwards, thereby shortening the
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of the tendon—the force generated by a 95% contraction of all fibers is sufficient to damage the body. In
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of the hand do not move, but muscles generate sufficient force to prevent the object from being dropped.
1772:, a field that still studies the electrical patterns and signals in tissues such as nerves and muscles. 4183: 2973:
Shwedyk, E.; Balasubramanian, R.; Scott, R. N. (1977). "A nonstationary model for the Electromyogram".
1818: 1322: 625:, depolarizing the inner portion of the muscle fiber. This activates dihydropyridine receptors in the 612: 391:
In vertebrate animals, there are three types of muscle tissues: 1) skeletal, 2) smooth, and 3) cardiac
2110:. Vol. 578 (4th ed.). Baltimore, Maryland: Lippincott Williams and Wilkins. pp. 37–56. 2035: 1233: 1059: 730:
Cross-bridge cycling is a sequence of molecular events that underlies the sliding filament theory. A
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are involved in eccentric contractions, but their mechanism is poorly understood in comparison to
2919:"A brief contraction has complex effects on summation of twitch pairs in human adductor pollicis" 1863: 1063: 1051:
Swellings called varicosities belonging to an autonomic neuron innervate the smooth muscle cells.
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Muscles undergoing heavy eccentric loading suffer greater damage when overloaded (such as during
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The Science of Common Things: A Familiar Explanation of the First Principles of Physical Science
3067:"Muscle co-contraction modulates damping and joint stability in a three-link biomechanical limb" 2514:"Ryanodine receptors: structure, expression, molecular details, and function in calcium release" 2512:
Lanner, Johanna T.; Georgiou, Dimitra K.; Joshi, Aditya D.; Hamilton, Susan L. (November 2010).
1803:. Their findings were published as two consecutive papers published in the 22 May 1954 issue of 1422:(the latter are not seen in all cardiac cell types) and the depolarisation causes extracellular 283:, the elbow starts the movement straight and then bends as the hand moves towards the shoulder. 275:, the elbow starts the movement while bent and then straightens as the hand moves away from the 3984:
Krans, J. L. (2010) The Sliding Filament Theory of Muscle Contraction. Nature Education 3(9):66
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Fabiato, A. (1983). "Calcium-induced calcium release from the cardiac sarcoplasmic reticulum".
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constitutes the majority of muscle mass in the body and is responsible for locomotor activity.
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The major constituent of thin filaments is a chain formed by helical coiling of two strands of
605:, (DHPRs). DHPRs are located on the sarcolemma (which includes the surface sarcolemma and the 1990:
Biewener, Andrew A. (2003). "Muscles and skeletons: The building blocks of animal movement".
1411: 1318: 747: 272: 2315:"Heavy-load eccentric calf muscle training for the treatment of chronic Achilles tendinosis" 2028: 799:
Cross-bridge cycling is able to continue as long as there are sufficient amounts of ATP and
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that constitute the flight muscles in these animals. These flight muscles are often called
1732: 1664: 1368:(main RyR isoform in skeletal muscle) to regulate Ca release in skeletal muscle, while the 1236:. The calcium-calmodulin-myosin light-chain kinase complex phosphorylates myosin on the 20 957:, the force exerted by the skeletal muscle is controlled by varying the frequency at which 931: 654: 594: 573:
or reabsorbed by the synaptic knob and none is left to replace the degraded acetylcholine.
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would cause the arm to bend at the elbow as the hand moved from the leg to the shoulder (a
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Sliding filament theory: A sarcomere in relaxed (above) and contracted (below) positions
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on the neuromuscular junction. Activation of the nicotinic receptor opens its intrinsic
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During a concentric contraction, a muscle is stimulated to contract according to the
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Levitan, Irwin; Kaczmarek, Leonard (19 August 2015). "Intercellular communication".
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Key proteins involved in cardiac calcium cycling and excitation-contraction coupling
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ions that are released from the sarcoplasmic reticulum. The elevation of cytosolic
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is detected by RyR2 in the membrane of the sarcoplasmic reticulum, which releases
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channel, causing sodium to rush in and potassium to trickle out. As a result, the
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Nikolaidis MG, Kyparos A, Spanou C, Paschalis V, Theodorou AA, Vrabas IS (2012).
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makes up the heart, which pumps blood. Skeletal and cardiac muscles are called
420: 123: 92: 2586: 1660: 641:
and causes a cell-wide increase in calcium giving rise to the upstroke of the
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Gardner, C.R. (1976). "The neuronal control of locomotion in the earthworm".
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1848: 1765: 1563: 1415: 1241: 1091: 1055: 927: 894: 637:. The action potential creates a near synchronous activation of thousands of 634: 545: 435: 412: 408: 400: 135: 119: 100: 74: 37: 3814: 3628: 3519: 3458: 3296: 1387:(JPH2) is essential to maintain this structure, as well as the integrity of 4274: 3967: 3941: 3910: 3830: 3776:(2nd ed.). Princeton, NJ: Princeton University Press. pp. 15–37. 3484: 3427: 3395: 3394:
Takeshima, H.; Komazaki, S.; Nishi, M.; Iino, M.; Kangawa, K. (July 2000).
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The strength of skeletal muscle contractions can be broadly separated into
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under the common theme "Structural Changes in Muscle During Contraction".
1614:
concentration in response to a large change in total calcium. The falling
982:, being more excitable than the larger ones, are stimulated first. As the 4311: 4134: 4091: 3745: 3339:"T-tubule remodeling during transition from hypertrophy to heart failure" 2994: 2476:(4th ed.). New York, NY: Oxford University Press. pp. 153–328. 2415: 1858: 1800: 1796: 1669: 1639: 1539:
myosin head pulls the actin filament toward the centre of the sarcomere.
1115: 908: 743: 706: 702: 464: 334: 233: 225: 52: 4216: 2688: 2686: 970:). This 'low' level of contraction is a protective mechanism to prevent 264:
involving both eccentric and concentric contractions appear to increase
4367: 3949: 3396:"Junctophilins: a novel family of junctional membrane complex proteins" 2275: 2258: 1700: 1652: 1282:
are believed to participate in the sustained phase of contraction, and
1271: 1229: 1170:
ions are also required for crossbridge cycling in smooth muscle cells.
1137:. These action potentials are generated by the influx of extracellular 979: 769: 646: 565: 396: 84: 56: 3700:(2nd ed.). Sunderland, MA: Sinauer Associates. pp. 681–698. 1588:
from the cytoplasm back into the sarcoplasmic reticulum. Once again,
629:, which are in close proximity to ryanodine receptors in the adjacent 504:
along its axon toward the neuromuscular junction. Once it reaches the
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Josephson, R. K.; Malamud, J. G.; Stokes, D. R. (15 September 2000).
3663: 2935: 2856:(5th ed.). New York, NY: McGraw-Hill Medical. pp. 768–789. 2768: 2714: 2683: 1873: 1760:
Electrodes touch a frog, and the legs twitch into the upward position
1745: 810:
in the cytoplasm. Termination of cross-bridge cycling can occur when
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to fuse with the plasma membrane, releasing acetylcholine into the
460: 419:, and other areas in the body that produce sustained contractions. 276: 4014: 2742: 2740: 2571:"The excitation-contraction coupling mechanism in skeletal muscle" 149:
that contract during locomotion. Contractions can be described as
1777: 1656: 1562:. Some calcium is also taken up by the mitochondria. An enzyme, 1334: 616:
Picture showing the different interactions within the ECC pathway
365: 311: 292: 189: 96: 2877:
Feher, Joseph (2012). "Chapter 3.4: Skeletal muscle mechanics".
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beating is made possible because the muscles are connected to a
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Excitation–contraction coupling (ECC) is the process by which a
4248: 4130: 4081: 2737: 1728: 1418:. The action potential travels along the surface membrane into 746:. With the ATP hydrolyzed, the cocked myosin head now contains 690: 557: 338: 284: 229: 77: 2569:
Calderón, Juan C.; Bolaños, Pura; Caputo, Carlo (March 2014).
1966:(7th ed.). San Francisco, CA: Pearson. pp. 377–416. 1941:(12th ed.). New York, NY: McGraw-Hill. pp. 250–291. 1603:
concentration, permitting a relatively small decrease in free
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Excitation-contraction coupling and cardiac contractile force
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i during excitation-contraction coupling in cardiac myocytes"
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Exercise Physiology: Human Bioenergetics and Its Applications
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system, which is driven to a natural frequency of vibration.
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A simplified image showing earthworm movement via peristalsis
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Following systole, intracellular calcium is taken up by the
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Alfredson, H; Pietilä, T; Jonsson, P; Lorentzon, R (1998).
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physiological response. This positive feedback is known as
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to enter the cell via L-type calcium channels and possibly
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release the neurotransmitter acetylcholine, which binds to
859:
ion concentration in the sarcoplasm. The active pumping of
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A History of the Theories of Aether and Electricity. Vol 1
3393: 2451:(Second ed.). Cambridge: Cambridge University Press. 2397: 1939:
Vander's Human Physiology: The Mechanisms of Body Function
1364:(RyRs) are distinct isoforms. Besides, DHPR contacts with 1081: 355:, moderate training may confer protection against injury. 73:, and thick filaments dominantly consist of chains of the 3726:
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Smith, Ian C.; Adam, Helen; Herzog, Walter (April 2020).
1724: 1720: 2490: 2128:(1st ed.). Boca Raton, FL: CRC Press. p. 113. 3613:(2nd ed.). Dordrecht: Kluwer Academic Publishers. 3064: 1672:, which underlies the creeping movement of earthworms. 1508:
sparks). The spatial and temporal summation of ~30,000
1121:
Sliding filaments in contracted and uncontracted states
111:, summation, or tetanus, depending on the frequency of 3885:"Excitation-Contraction Coupling in Muscular Response" 3796: 3123: 2667:
Anatomy and Physiology: The Unity of Form and Function
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influx causes a small local increase in intracellular
333:. During a concentric contraction, contractile muscle 3876: 3641: 2568: 1922: 1920: 1918: 1916: 1914: 1912: 1910: 1908: 1906: 1904: 1317:(mAChRs) on smooth muscle cells. These receptors are 358: 2101: 2099: 2097: 2095: 2093: 2091: 1634: 3772:Alexander, R. McNeill (2003). "Muscle, the motor". 3540: 3497: 2074:(1st ed.). New York, NY: Thieme. p. 113. 1326:excitatory (contractile) or inhibitory (relaxing). 657:, producing force and, in some situations, motion. 3336: 3015: 2027: 1936: 1901: 1350: 653:. This bond allows the actin filaments to perform 576: 3767: 3765: 3763: 3695: 3541:Cannell MB, Cheng H, Lederer WJ (November 1994). 2692: 2507: 2505: 2446: 2088: 1558:(NCX) and, to a lesser extent, a plasma membrane 1255:-activated phosphorylation of myosin rather than 848:ions leave the troponin molecule to maintain the 138:are similar to those in skeletal muscle tissues. 107:. The contraction produced can be described as a 4384: 3926:"Past, Present and Future Experiments on Muscle" 3269: 3183:Martonosi, Anthony N.; Pikula, Slawomir (2003). 2797: 2471: 2297: 2105: 2070:Sircar, Sabyasachi (2008). "Muscle elasticity". 1360:(DHPRs) are encoded by different genes, and the 302:Though the muscle is doing a negative amount of 27:Activation of tension-generating sites in muscle 3930:Philosophical Transactions: Biological Sciences 3719: 3717: 3182: 2916: 2221: 1985: 1983: 1955: 1024: 593:(SR) calcium release channel identified as the 3852:. New York: Ivison & Phinney. p. 290. 3760: 3691: 3689: 3124:Zhang, Y; Hermanson, ME; Eddinger, TJ (2013). 2879:Quantitative Human Physiology: An Introduction 2502: 2298:Brooks, G.A; Fahey, T.D.; White, T.P. (1996). 997: 463:, the signal to contract can originate in the 4232: 4030: 3440: 3185:"The network of calcium regulation in muscle" 2847: 2845: 2843: 2217: 2215: 2213: 2065: 2063: 2061: 2059: 2057: 2055: 2021: 2019: 2017: 2015: 2013: 2011: 1675: 1184:in smooth muscle cells are the extracellular 4010:Sliding Filament Model of Muscle Contraction 3714: 2851: 2746: 2658: 2465: 2362: 2250: 2119: 2117: 1980: 1962:Silverthorn, Dee Unglaub (2016). "Muscles". 1414:and conducted to all cells in the heart via 645:. The Ca released into the cytosol binds to 118:Unlike skeletal muscle, the contractions of 3917: 3686: 3234: 2975:IEEE Transactions on Biomedical Engineering 2791: 2142: 1961: 920:Three types of skeletal muscle contractions 825:back into the sarcoplasmic reticulum. When 459:several muscle fibers. In the case of some 4239: 4225: 4037: 4023: 3230: 3228: 2870: 2840: 2628:"SMOOTH MUSCLE CONTRACTION AND RELAXATION" 2518:Cold Spring Harbor Perspectives in Biology 2210: 2052: 2008: 1195:entering through calcium channels and the 672: 663:sarco/endoplasmic reticulum calcium-ATPase 4246: 3957: 3900: 3862: 3856: 3846:"How galvanic electricity was discovered" 3771: 3585: 3474: 3447:Journal of the American Heart Association 3370: 3313: 3295: 3200: 3159: 3149: 3100: 3082: 3041: 2934: 2906:(1st ed.). Elsevier. pp. 101–2. 2823: 2798:Horowits R, Podolsky RJ (November 1987). 2602: 2545: 2423: 2274: 2114: 945:. Summation can be achieved in two ways: 885:Gradation of skeletal muscle contractions 471: 279:. During an eccentric contraction of the 211: 134:. The mechanisms of contraction in these 3016:Gordon AM, Huxley AF, Julian FJ (1966). 2148: 1989: 1964:Human Physiology: An Integrated Approach 1755: 1699: 1638: 1541: 1333: 1046: 1028: 1007: 721: 676: 611: 475: 434: 386: 310:the muscle), chemical energy (of fat or 243: 177: 164: 31: 3723: 3225: 2901: 2664: 2322:The American Journal of Sports Medicine 1695: 1082:Mechanisms of smooth muscle contraction 522:influx into the terminal by way of the 271:During an eccentric contraction of the 199: 14: 4385: 3923: 3882: 3117: 2474:The Neuron: Cell and Molecular Biology 2190: 2188: 2069: 2025: 4220: 4018: 3843: 3009: 2910: 2876: 2123: 772:on the actin filaments. The troponin- 717: 364:training for rapid movements such as 3608: 2625: 1012:Muscle length versus isometric force 599:voltage-gated L-type calcium channel 480:Structure of neuromuscular junction. 4044: 2185: 1437:(NCX) during the early part of the 1159:. Like skeletal muscles, cytosolic 1058:can be divided into two subgroups: 236:). A concentric contraction of the 24: 3998:Animation: Myofilament Contraction 3978: 3738:10.1111/j.1469-185X.1976.tb01119.x 2626:Webb, R. Clinton (December 2003). 2236:10.1111/j.1748-1716.1990.tb08973.x 1552:sarco/endoplasmic reticulum ATPase 1315:muscarinic acetylcholine receptors 1296: 430: 359:Eccentric contractions in movement 228:, a concentric contraction of the 25: 4424: 4201:List of muscles of the human body 3991: 2030:The Physiology of Excitable Cells 1635:Circular and longitudinal muscles 1329: 1232:, which then binds and activates 554:nicotinic acetylcholine receptors 403:: skeletal, smooth, and cardiac. 2632:Advances in Physiology Education 2369:Isokinetics and Exercise Science 2072:Principles of Medical Physiology 1680:Invertebrates such as annelids, 1463:. The increase of intracellular 1114: 1100: 1042: 907: 893: 544:containing the neurotransmitter 508:, the action potential causes a 492:formed by the contact between a 351:contractions can easily lead to 3837: 3803:Journal of Experimental Biology 3799:"Asynchronous muscle: a primer" 3790: 3774:Principles of Animal Locomotion 3635: 3602: 3534: 3491: 3434: 3387: 3330: 3263: 3176: 3058: 2966: 2895: 2619: 2562: 2440: 2391: 2356: 2306: 2291: 2163:10.1152/japplphysiol.00280.2003 1824:calcium-induced calcium release 1629: 1491:calcium-induced calcium release 1381:calcium-induced calcium release 1372:(DHPR on cardiac myocytes) and 1351:Excitation-contraction coupling 577:Excitation–contraction coupling 439:Organization of skeletal muscle 260:walking). Over the short-term, 3237:American Journal of Physiology 3034:10.1113/jphysiol.1966.sp007909 2904:Textbook Of Medical Physiology 1311:parasympathetic nervous system 1173:The two sources for cytosolic 661:muscles. Although slower, the 524:voltage-gated calcium channels 13: 1: 3578:10.1016/S0006-3495(94)80677-0 3543:"Spatial non-uniformities in 3412:10.1016/s1097-2765(00)00003-4 3355:10.1161/CIRCRESAHA.109.212324 3249:10.1152/ajpcell.1983.245.1.C1 2151:Journal of Applied Physiology 1894: 382: 348:delayed onset muscle soreness 3924:Huxley, H. E. (April 2000). 3151:10.1371/journal.pone.0074608 2854:Principles of Neural Science 2449:Handbook of Psychophysiology 2334:10.1177/03635465980260030301 1393:receptor accessory protein 5 1025:Force-velocity relationships 689:describes a process used by 299:in concentric contractions. 169:Types of muscle contractions 89:skeletal muscle contractions 7: 2530:10.1101/cshperspect.a003996 2363:Satyendra L, Byl N (2006). 1812: 1307:Postganglionic nerve fibers 998:Length-tension relationship 585:in the muscle fiber causes 399:, there are three types of 314:, or temporarily stored in 10: 4429: 4184:Anatomical terms of muscle 3844:Wells, David Ames (1859). 3071:Frontiers in Neurorobotics 1819:Anatomical terms of motion 1784:to cause cell shortening. 1751: 1711:Insect wing § Muscles 1708: 1676:Obliquely striated muscles 1347:and are able to contract. 1323:sympathetic nervous system 1107:Smooth muscle contractions 1085: 1001: 247: 181: 4363: 4350: 4320: 4302: 4295: 4255: 4176: 4143: 4052: 2669:. New York: McGraw Hill. 2665:Saladin, Kenneth (2012). 2587:10.1007/s12551-013-0135-x 2302:. Mayfield Publishing Co. 1715:Advanced insects such as 1358:dihydropyridine receptors 1293:flux may be significant. 1234:myosin light-chain kinase 603:dihydropyridine receptors 583:muscular action potential 51:-generating sites within 3189:Acta Biochimica Polonica 3084:10.3389/fnbot.2011.00005 2987:10.1109/TBME.1977.326175 2644:10.1152/advan.00025.2003 1829:Cardiac action potential 1556:sodium-calcium exchanger 1435:sodium-calcium exchanger 1399:structures, rather than 1303:autonomic nervous system 976:multiple fiber summation 951:multiple fiber summation 443:Excluding reflexes, all 160: 132:autonomic nervous system 3815:10.1242/jeb.203.18.2713 3520:10.1126/science.8235594 3459:10.1161/JAHA.117.007205 3297:10.1073/pnas.1412729111 2923:Experimental Physiology 2196:"Types of contractions" 1864:In vitro muscle testing 1342:There are two types of 687:sliding filament theory 673:Sliding filament theory 222:sliding filament theory 105:sliding filament theory 4408:Musculoskeletal system 3942:10.1098/rstb.2000.0595 3202:10.18388/abp.2003_3711 2816:10.1083/jcb.105.5.2217 1761: 1706: 1644: 1592:moderate this fall in 1547: 1370:L-type calcium channel 1339: 1076:motor unit recruitment 1052: 1035: 1013: 984:strength of the signal 736:adenosine triphosphate 727: 682: 631:sarcoplasmic reticulum 617: 591:sarcoplasmic reticulum 571:acetylcholine esterase 486:neuromuscular junction 481: 472:Neuromuscular junction 440: 417:gastrointestinal tract 392: 306:, (work is being done 212:Concentric contraction 170: 41: 2381:10.3233/IES-2006-0223 1759: 1709:Further information: 1703: 1642: 1545: 1412:atrioventricular node 1337: 1086:Further information: 1074:responses, much like 1050: 1032: 1011: 1002:Further information: 914:Summation and tetanus 748:adenosine diphosphate 725: 680: 615: 479: 438: 390: 244:Eccentric contraction 178:Isometric contraction 168: 47:is the activation of 35: 3343:Circulation Research 2416:10.1136/bjsm.35.1.60 1741:asynchronous muscles 1733:asynchronous muscles 1696:Asynchronous muscles 1078:in skeletal muscle. 655:cross-bridge cycling 595:ryanodine receptor 1 502:saltatory conduction 297:cross-bridge cycling 206:isotonic contraction 200:Isotonic contraction 192:grip an object; the 18:Muscular contraction 4393:Exercise physiology 4355:End-plate potential 4340:Uterine contraction 4104:Fascial compartment 4003:21 May 2013 at the 3656:1990Natur.345..618C 3609:Bers, M.D. (2001). 3570:1994BpJ....67.1942C 3512:1993Sci...262..740C 3288:2014PNAS..11112240G 3282:(33): 12240–12245. 3142:2013PLoSO...874608Z 2761:1954Natur.173..973H 2707:1954Natur.173..971H 2575:Biophysical Reviews 2224:Acta Physiol. Scand 1889:Uterine contraction 1854:Hill's muscle model 1844:Exercise physiology 1391:. Another protein, 1362:ryanodine receptors 1135:slow wave potential 1131:pacemaker potential 1004:Hill's muscle model 964:Golgi tendon organs 955:frequency summation 947:frequency summation 377:patellar tendonitis 373:Achilles tendinitis 291:, and other z-line 4325:Muscle contraction 3883:Sandow, A (1952). 3698:Principles of Life 2276:10.1242/jeb.067470 2269:(Pt 10): 1615–25. 1762: 1707: 1645: 1548: 1493:and gives rise to 1340: 1088:Myogenic mechanism 1053: 1036: 1014: 728: 726:Cross-bridge cycle 718:Cross-bridge cycle 683: 627:terminal cisternae 618: 607:transverse tubules 482: 441: 397:vertebrate animals 393: 250:Eccentric training 184:Isometric exercise 171: 143:locomotor activity 45:Muscle contraction 42: 4380: 4379: 4376: 4375: 4214: 4213: 3936:(1396): 539–543. 3809:(18): 2713–2722. 3783:978-0-691-12634-0 3707:978-1-464-10947-8 2888:978-0-123-82163-8 2863:978-0-071-39011-8 2755:(4412): 973–976. 2701:(4412): 971–973. 2676:978-0-07-337825-1 2458:978-0-521-62634-7 2135:978-0-415-36953-4 2081:978-1-588-90572-7 2045:978-0-521-57421-1 2001:978-0-198-50022-3 1992:Animal Locomotion 1973:978-0-321-98122-6 1948:978-0-321-98122-6 1869:Lombard's paradox 1737:fibrillar muscles 1487:positive feedback 959:action potentials 930:, summation, and 643:calcium transient 542:synaptic vesicles 327:strength training 266:muscular strength 262:strength training 113:action potentials 61:muscle relaxation 16:(Redirected from 4420: 4300: 4299: 4241: 4234: 4227: 4218: 4217: 4206:Composite muscle 4039: 4032: 4025: 4016: 4015: 3972: 3971: 3961: 3921: 3915: 3914: 3904: 3880: 3874: 3873: 3872:, Nelson, London 3864:Whittaker, E. T. 3860: 3854: 3853: 3841: 3835: 3834: 3794: 3788: 3787: 3769: 3758: 3757: 3721: 3712: 3711: 3693: 3684: 3683: 3664:10.1038/345618a0 3650:(6276): 618–21. 3639: 3633: 3632: 3606: 3600: 3599: 3589: 3553: 3552: 3551: 3538: 3532: 3531: 3495: 3489: 3488: 3478: 3438: 3432: 3431: 3391: 3385: 3384: 3374: 3334: 3328: 3327: 3317: 3299: 3267: 3261: 3260: 3232: 3223: 3222: 3204: 3180: 3174: 3173: 3163: 3153: 3121: 3115: 3114: 3104: 3086: 3062: 3056: 3055: 3045: 3013: 3007: 3006: 2970: 2964: 2963: 2961: 2959: 2938: 2936:10.1113/ep088401 2914: 2908: 2907: 2899: 2893: 2892: 2874: 2868: 2867: 2849: 2838: 2837: 2827: 2795: 2789: 2788: 2769:10.1038/173973a0 2744: 2735: 2734: 2715:10.1038/173971a0 2690: 2681: 2680: 2662: 2656: 2655: 2623: 2617: 2616: 2606: 2566: 2560: 2559: 2549: 2509: 2500: 2497: 2488: 2487: 2469: 2463: 2462: 2444: 2438: 2437: 2427: 2395: 2389: 2388: 2383:. 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Huxley 1624: 1623: 1622: 1613: 1612: 1611: 1602: 1601: 1600: 1587: 1586: 1585: 1576: 1575: 1574: 1534: 1533: 1532: 1518: 1517: 1516: 1507: 1506: 1505: 1484: 1483: 1482: 1473: 1472: 1471: 1462: 1461: 1460: 1451: 1450: 1449: 1432: 1431: 1430: 1292: 1291: 1290: 1280:protein kinase C 1265: 1264: 1263: 1254: 1253: 1252: 1227: 1226: 1225: 1217:results in more 1216: 1215: 1214: 1205: 1204: 1203: 1194: 1193: 1192: 1183: 1182: 1181: 1169: 1168: 1167: 1158: 1157: 1156: 1147: 1146: 1145: 1118: 1104: 911: 897: 880: 879: 878: 869: 868: 867: 858: 857: 856: 847: 846: 845: 835: 834: 833: 820: 819: 818: 809: 808: 807: 782: 781: 780: 767: 766: 765: 699:Rolf Niedergerke 537: 536: 535: 519: 518: 517: 490:chemical synapse 147:skeletal muscles 95:as they require 21: 4428: 4427: 4423: 4422: 4421: 4419: 4418: 4417: 4403:Skeletal muscle 4398:Muscular system 4383: 4382: 4381: 4372: 4359: 4346: 4316: 4291: 4251: 4245: 4215: 4210: 4172: 4139: 4070:Skeletal muscle 4048: 4046:Muscular system 4043: 4005:Wayback Machine 3994: 3981: 3979:Further reading 3976: 3975: 3922: 3918: 3889:Yale J Biol Med 3881: 3877: 3861: 3857: 3842: 3838: 3795: 3791: 3784: 3770: 3761: 3722: 3715: 3708: 3694: 3687: 3640: 3636: 3621: 3607: 3603: 3550: 3548: 3547: 3546: 3544: 3539: 3535: 3506:(5134): 740–4. 3496: 3492: 3439: 3435: 3392: 3388: 3335: 3331: 3268: 3264: 3233: 3226: 3181: 3177: 3122: 3118: 3063: 3059: 3014: 3010: 2971: 2967: 2957: 2955: 2915: 2911: 2900: 2896: 2889: 2875: 2871: 2864: 2850: 2841: 2796: 2792: 2745: 2738: 2691: 2684: 2677: 2663: 2659: 2624: 2620: 2567: 2563: 2524:(11): a003996. 2510: 2503: 2498: 2491: 2484: 2470: 2466: 2459: 2445: 2441: 2404:Br J Sports Med 2396: 2392: 2387:on 9 July 2012. 2361: 2357: 2317: 2311: 2307: 2296: 2292: 2255: 2251: 2220: 2211: 2201: 2199: 2194: 2193: 2186: 2147: 2143: 2136: 2126:Muscle strength 2122: 2115: 2104: 2089: 2082: 2068: 2053: 2046: 2024: 2009: 2002: 1988: 1981: 1974: 1960: 1956: 1949: 1935: 1902: 1897: 1815: 1754: 1713: 1698: 1678: 1637: 1632: 1621: 1619: 1618: 1617: 1615: 1610: 1608: 1607: 1606: 1604: 1599: 1597: 1596: 1595: 1593: 1590:calcium buffers 1584: 1582: 1581: 1580: 1578: 1573: 1571: 1570: 1569: 1567: 1531: 1529: 1528: 1527: 1525: 1522:calcium buffers 1515: 1513: 1512: 1511: 1509: 1504: 1502: 1501: 1500: 1498: 1481: 1479: 1478: 1477: 1475: 1470: 1468: 1467: 1466: 1464: 1459: 1457: 1456: 1455: 1453: 1448: 1446: 1445: 1444: 1442: 1429: 1427: 1426: 1425: 1423: 1408:sinoatrial node 1353: 1332: 1299: 1297:Neuromodulation 1289: 1287: 1286: 1285: 1283: 1262: 1260: 1259: 1258: 1256: 1251: 1249: 1248: 1247: 1245: 1224: 1222: 1221: 1220: 1218: 1213: 1211: 1210: 1209: 1207: 1202: 1200: 1199: 1198: 1196: 1191: 1189: 1188: 1187: 1185: 1180: 1178: 1177: 1176: 1174: 1166: 1164: 1163: 1162: 1160: 1155: 1153: 1152: 1151: 1149: 1144: 1142: 1141: 1140: 1138: 1126: 1125: 1124: 1123: 1122: 1119: 1110: 1109: 1108: 1105: 1094: 1084: 1045: 1027: 1006: 1000: 924: 923: 922: 921: 917: 916: 915: 912: 903: 902: 901: 898: 887: 877: 875: 874: 873: 871: 866: 864: 863: 862: 860: 855: 853: 852: 851: 849: 844: 842: 841: 840: 838: 832: 830: 829: 828: 826: 823:actively pumped 817: 815: 814: 813: 811: 806: 804: 803: 802: 800: 779: 777: 776: 775: 773: 764: 762: 761: 760: 758: 755: 720: 675: 651:actin filaments 597:(RYR1) and the 579: 534: 532: 531: 530: 528: 516: 514: 513: 512: 510: 506:terminal bouton 474: 445:skeletal muscle 433: 431:Skeletal muscle 425:striated muscle 405:Skeletal muscle 385: 361: 323:muscle building 304:mechanical work 252: 246: 214: 202: 186: 180: 163: 124:cardiac muscles 28: 23: 22: 15: 12: 11: 5: 4426: 4416: 4415: 4410: 4405: 4400: 4395: 4378: 4377: 4374: 4373: 4371: 4370: 4364: 4361: 4360: 4358: 4357: 4351: 4348: 4347: 4345: 4344: 4343: 4342: 4337: 4332: 4321: 4318: 4317: 4315: 4314: 4309: 4303: 4297: 4293: 4292: 4290: 4289: 4288: 4287: 4282: 4277: 4267: 4261: 4259: 4253: 4252: 4247:Physiology of 4244: 4243: 4236: 4229: 4221: 4212: 4211: 4209: 4208: 4203: 4198: 4197: 4196: 4191: 4180: 4178: 4174: 4173: 4171: 4170: 4169: 4168: 4163: 4156:Pennate muscle 4153: 4147: 4145: 4141: 4140: 4138: 4137: 4128: 4127: 4126: 4121: 4116: 4111: 4101: 4100: 4099: 4094: 4089: 4079: 4078: 4077: 4072: 4067: 4065:Cardiac muscle 4056: 4054: 4050: 4049: 4042: 4041: 4034: 4027: 4019: 4013: 4012: 4007: 3993: 3992:External links 3990: 3989: 3988: 3985: 3980: 3977: 3974: 3973: 3916: 3895:(3): 176–201. 3875: 3855: 3836: 3789: 3782: 3759: 3713: 3706: 3685: 3634: 3619: 3601: 3564:(5): 1942–56. 3549: 3533: 3490: 3433: 3400:Molecular Cell 3386: 3349:(4): 520–531. 3329: 3262: 3224: 3175: 3116: 3057: 3008: 2981:(5): 417–424. 2965: 2929:(4): 676–689. 2909: 2894: 2887: 2869: 2862: 2839: 2810:(5): 2217–23. 2790: 2736: 2682: 2675: 2657: 2638:(4): 201–206. 2618: 2581:(1): 133–160. 2561: 2501: 2489: 2483:978-0199773893 2482: 2464: 2457: 2439: 2390: 2355: 2305: 2290: 2249: 2209: 2184: 2157:(2): 455–459. 2141: 2134: 2113: 2108:NMS Physiology 2087: 2080: 2051: 2044: 2007: 2000: 1979: 1972: 1954: 1947: 1899: 1898: 1896: 1893: 1892: 1891: 1886: 1881: 1876: 1871: 1866: 1861: 1856: 1851: 1846: 1841: 1836: 1831: 1826: 1821: 1814: 1811: 1770:bioelectricity 1753: 1750: 1697: 1694: 1677: 1674: 1661:coelomic fluid 1636: 1633: 1631: 1628: 1620: 1609: 1598: 1583: 1572: 1560:calcium ATPase 1530: 1514: 1503: 1495:calcium sparks 1480: 1469: 1458: 1447: 1428: 1385:Junctophilin-2 1352: 1349: 1344:cardiac muscle 1338:Cardiac muscle 1331: 1330:Cardiac muscle 1328: 1298: 1295: 1288: 1261: 1250: 1223: 1212: 1201: 1190: 1179: 1165: 1154: 1143: 1120: 1113: 1112: 1111: 1106: 1099: 1098: 1097: 1096: 1095: 1083: 1080: 1056:Smooth muscles 1044: 1041: 1026: 1023: 999: 996: 919: 918: 913: 906: 905: 904: 899: 892: 891: 890: 889: 888: 886: 883: 876: 865: 854: 843: 831: 816: 805: 778: 763: 753: 719: 716: 674: 671: 639:calcium sparks 601:identified as 578: 575: 550:synaptic cleft 533: 515: 473: 470: 449:nervous system 432: 429: 421:Cardiac muscle 401:muscle tissues 384: 381: 360: 357: 281:triceps muscle 245: 242: 213: 210: 201: 198: 182:Main article: 179: 176: 162: 159: 136:muscle tissues 97:synaptic input 26: 9: 6: 4: 3: 2: 4425: 4414: 4411: 4409: 4406: 4404: 4401: 4399: 4396: 4394: 4391: 4390: 4388: 4369: 4366: 4365: 4362: 4356: 4353: 4352: 4349: 4341: 4338: 4336: 4333: 4331: 4328: 4327: 4326: 4323: 4322: 4319: 4313: 4310: 4308: 4307:Hand strength 4305: 4304: 4301: 4298: 4294: 4286: 4283: 4281: 4278: 4276: 4273: 4272: 4271: 4268: 4266: 4263: 4262: 4260: 4258: 4254: 4250: 4242: 4237: 4235: 4230: 4228: 4223: 4222: 4219: 4207: 4204: 4202: 4199: 4195: 4192: 4190: 4187: 4186: 4185: 4182: 4181: 4179: 4175: 4167: 4164: 4162: 4159: 4158: 4157: 4154: 4152: 4149: 4148: 4146: 4142: 4136: 4132: 4129: 4125: 4122: 4120: 4117: 4115: 4112: 4110: 4107: 4106: 4105: 4102: 4098: 4095: 4093: 4090: 4088: 4085: 4084: 4083: 4080: 4076: 4075:Smooth muscle 4073: 4071: 4068: 4066: 4063: 4062: 4061: 4060:Muscle tissue 4058: 4057: 4055: 4051: 4047: 4040: 4035: 4033: 4028: 4026: 4021: 4020: 4017: 4011: 4008: 4006: 4002: 3999: 3996: 3995: 3986: 3983: 3982: 3969: 3965: 3960: 3955: 3951: 3947: 3943: 3939: 3935: 3931: 3927: 3920: 3912: 3908: 3903: 3898: 3894: 3890: 3886: 3879: 3871: 3870: 3865: 3859: 3851: 3847: 3840: 3832: 3828: 3824: 3820: 3816: 3812: 3808: 3804: 3800: 3793: 3785: 3779: 3775: 3768: 3766: 3764: 3755: 3751: 3747: 3743: 3739: 3735: 3731: 3727: 3720: 3718: 3709: 3703: 3699: 3692: 3690: 3681: 3677: 3673: 3669: 3665: 3661: 3657: 3653: 3649: 3645: 3638: 3630: 3626: 3622: 3620:9780792371571 3616: 3612: 3605: 3597: 3593: 3588: 3583: 3579: 3575: 3571: 3567: 3563: 3559: 3555: 3537: 3529: 3525: 3521: 3517: 3513: 3509: 3505: 3501: 3494: 3486: 3482: 3477: 3472: 3468: 3464: 3460: 3456: 3452: 3448: 3444: 3437: 3429: 3425: 3421: 3417: 3413: 3409: 3405: 3401: 3397: 3390: 3382: 3378: 3373: 3368: 3364: 3360: 3356: 3352: 3348: 3344: 3340: 3333: 3325: 3321: 3316: 3311: 3307: 3303: 3298: 3293: 3289: 3285: 3281: 3277: 3273: 3266: 3258: 3254: 3250: 3246: 3242: 3238: 3231: 3229: 3220: 3216: 3212: 3208: 3203: 3198: 3194: 3190: 3186: 3179: 3171: 3167: 3162: 3157: 3152: 3147: 3143: 3139: 3136:(9): e74608. 3135: 3131: 3127: 3120: 3112: 3108: 3103: 3098: 3094: 3090: 3085: 3080: 3076: 3072: 3068: 3061: 3053: 3049: 3044: 3039: 3035: 3031: 3028:(1): 170–92. 3027: 3023: 3019: 3012: 3004: 3000: 2996: 2992: 2988: 2984: 2980: 2976: 2969: 2954: 2950: 2946: 2942: 2937: 2932: 2928: 2924: 2920: 2913: 2905: 2898: 2890: 2884: 2880: 2873: 2865: 2859: 2855: 2848: 2846: 2844: 2835: 2831: 2826: 2821: 2817: 2813: 2809: 2805: 2801: 2794: 2786: 2782: 2778: 2774: 2770: 2766: 2762: 2758: 2754: 2750: 2743: 2741: 2732: 2728: 2724: 2720: 2716: 2712: 2708: 2704: 2700: 2696: 2689: 2687: 2678: 2672: 2668: 2661: 2653: 2649: 2645: 2641: 2637: 2633: 2629: 2622: 2614: 2610: 2605: 2600: 2596: 2592: 2588: 2584: 2580: 2576: 2572: 2565: 2557: 2553: 2548: 2543: 2539: 2535: 2531: 2527: 2523: 2519: 2515: 2508: 2506: 2496: 2494: 2485: 2479: 2475: 2468: 2460: 2454: 2450: 2443: 2435: 2431: 2426: 2421: 2417: 2413: 2409: 2405: 2401: 2394: 2386: 2382: 2378: 2374: 2370: 2366: 2359: 2351: 2347: 2343: 2339: 2335: 2331: 2327: 2323: 2316: 2309: 2301: 2294: 2286: 2282: 2277: 2272: 2268: 2264: 2260: 2253: 2245: 2241: 2237: 2233: 2229: 2225: 2218: 2216: 2214: 2198:. 31 May 2006 2197: 2191: 2189: 2180: 2176: 2172: 2168: 2164: 2160: 2156: 2152: 2145: 2137: 2131: 2127: 2120: 2118: 2109: 2102: 2100: 2098: 2096: 2094: 2092: 2083: 2077: 2073: 2066: 2064: 2062: 2060: 2058: 2056: 2047: 2041: 2037: 2032: 2031: 2022: 2020: 2018: 2016: 2014: 2012: 2003: 1997: 1993: 1986: 1984: 1975: 1969: 1965: 1958: 1950: 1944: 1940: 1933: 1931: 1929: 1927: 1925: 1923: 1921: 1919: 1917: 1915: 1913: 1911: 1909: 1907: 1905: 1900: 1890: 1887: 1885: 1882: 1880: 1877: 1875: 1872: 1870: 1867: 1865: 1862: 1860: 1857: 1855: 1852: 1850: 1849:Fasciculation 1847: 1845: 1842: 1840: 1837: 1835: 1832: 1830: 1827: 1825: 1822: 1820: 1817: 1816: 1810: 1808: 1807: 1802: 1798: 1794: 1790: 1785: 1783: 1779: 1773: 1771: 1767: 1766:Luigi Galvani 1758: 1749: 1747: 1742: 1738: 1734: 1730: 1726: 1722: 1718: 1712: 1702: 1693: 1691: 1687: 1683: 1673: 1671: 1666: 1665:hydroskeleton 1662: 1658: 1654: 1650: 1641: 1627: 1591: 1565: 1564:phospholamban 1561: 1557: 1553: 1544: 1540: 1536: 1523: 1496: 1492: 1488: 1440: 1439:plateau phase 1436: 1421: 1417: 1416:gap junctions 1413: 1409: 1404: 1402: 1398: 1394: 1390: 1386: 1382: 1377: 1375: 1371: 1367: 1363: 1359: 1348: 1345: 1336: 1327: 1324: 1320: 1316: 1312: 1308: 1304: 1294: 1281: 1277: 1273: 1267: 1243: 1242:myosin ATPase 1239: 1235: 1231: 1171: 1136: 1132: 1117: 1103: 1093: 1092:Myogenic tone 1089: 1079: 1077: 1071: 1069: 1065: 1061: 1057: 1049: 1043:Smooth muscle 1040: 1031: 1022: 1020: 1010: 1005: 995: 993: 988: 985: 981: 977: 973: 969: 968:Renshaw cells 965: 960: 956: 952: 948: 944: 939: 937: 936:latent period 933: 929: 910: 896: 882: 824: 797: 795: 791: 787: 786:power stroke, 771: 768:ions bind to 756: 749: 745: 741: 737: 733: 724: 715: 713: 708: 704: 700: 696: 695:Andrew Huxley 692: 688: 679: 670: 666: 664: 658: 656: 652: 648: 644: 640: 636: 635:calcium spark 632: 628: 624: 614: 610: 608: 604: 600: 596: 592: 588: 584: 574: 572: 567: 563: 559: 555: 551: 547: 546:acetylcholine 543: 539: 525: 521: 507: 503: 499: 495: 491: 487: 478: 469: 466: 462: 458: 454: 450: 446: 437: 428: 426: 422: 418: 414: 413:blood vessels 410: 409:Smooth muscle 406: 402: 398: 389: 380: 378: 374: 369: 367: 356: 354: 349: 344: 340: 336: 332: 328: 324: 319: 317: 313: 309: 305: 300: 298: 294: 290: 286: 282: 278: 274: 273:biceps muscle 269: 267: 263: 258: 251: 241: 239: 235: 231: 227: 223: 218: 209: 207: 197: 195: 191: 185: 175: 167: 158: 156: 152: 148: 144: 139: 137: 133: 129: 125: 121: 116: 114: 110: 106: 102: 101:motor neurons 98: 94: 90: 86: 81: 79: 76: 75:motor-protein 72: 67: 64: 62: 58: 54: 50: 46: 39: 38:smooth muscle 36:Depiction of 34: 30: 19: 4324: 4275:Eye movement 3933: 3929: 3919: 3892: 3888: 3878: 3867: 3858: 3849: 3839: 3806: 3802: 3792: 3773: 3732:(1): 25–52. 3729: 3725: 3697: 3647: 3643: 3637: 3610: 3604: 3561: 3557: 3536: 3503: 3499: 3493: 3450: 3446: 3436: 3406:(1): 11–22. 3403: 3399: 3389: 3346: 3342: 3332: 3279: 3275: 3265: 3243:(1): C1–14. 3240: 3236: 3192: 3188: 3178: 3133: 3129: 3119: 3074: 3070: 3060: 3025: 3021: 3011: 2978: 2974: 2968: 2956:. Retrieved 2926: 2922: 2912: 2903: 2897: 2878: 2872: 2853: 2807: 2804:J. Cell Biol 2803: 2793: 2752: 2748: 2698: 2694: 2666: 2660: 2635: 2631: 2621: 2578: 2574: 2564: 2521: 2517: 2473: 2467: 2448: 2442: 2407: 2403: 2393: 2385:the original 2375:(1): 71–80. 2372: 2368: 2358: 2328:(3): 360–6. 2325: 2321: 2308: 2299: 2293: 2266: 2263:J. Exp. Biol 2262: 2252: 2227: 2223: 2200:. Retrieved 2154: 2150: 2144: 2125: 2107: 2071: 2029: 1991: 1963: 1957: 1938: 1879:Rigor mortis 1804: 1786: 1774: 1763: 1740: 1736: 1714: 1679: 1663:serves as a 1646: 1630:Invertebrate 1549: 1537: 1405: 1378: 1354: 1341: 1319:metabotropic 1300: 1268: 1172: 1127: 1072: 1054: 1037: 1015: 991: 989: 975: 954: 950: 946: 942: 940: 935: 925: 798: 794:rigor mortis 785: 732:cross-bridge 729: 684: 667: 659: 619: 580: 498:muscle fiber 494:motor neuron 483: 453:motor neuron 442: 394: 370: 362: 353:overtraining 335:myofilaments 330: 320: 307: 301: 270: 253: 219: 215: 203: 187: 172: 140: 117: 82: 68: 65: 60: 53:muscle cells 44: 43: 29: 4312:Muscle tone 4135:Aponeurosis 4087:Superficial 3195:(1): 1–30. 2410:(1): 60–4. 2230:(1): 31–9. 1859:Hypnic jerk 1801:Jean Hanson 1797:Hugh Huxley 1670:peristalsis 1228:binding to 1060:single-unit 980:motor units 744:tropomyosin 707:Jean Hanson 703:Hugh Huxley 465:spinal cord 234:biceps curl 85:vertebrates 40:contraction 4387:Categories 4368:Myogenesis 4285:Locomotion 4161:Unipennate 3558:Biophys. J 3022:J. Physiol 1895:References 1653:earthworms 1272:rho kinase 1238:kilodalton 1230:calmodulin 1148:, and not 770:troponin C 647:Troponin C 587:myofibrils 566:sarcolemma 457:innervates 383:Vertebrate 257:decelerate 248:See also: 93:neurogenic 57:physiology 4413:Neurology 4330:Isometric 4194:Insertion 4166:Bipennate 3823:0022-0949 3467:2047-9980 3420:1097-2765 3363:1524-4571 3306:1091-6490 3211:0001-527X 3093:1662-5218 2953:211100581 2652:1043-4046 2595:1867-2450 2538:1943-0264 2202:2 October 1874:Myoclonus 1764:In 1780, 1686:nematodes 1420:T-tubules 1068:syncytium 1064:multiunit 943:summation 790:sarcomere 623:T-tubules 562:potassium 331:negatives 151:isometric 4335:Isotonic 4270:Movement 4265:Exercise 4257:Exertion 4151:Fusiform 4097:Visceral 4001:Archived 3968:10836507 3911:13015950 3866:(1951), 3831:10952872 3629:47659382 3485:29431104 3428:10949023 3381:20576937 3324:25092313 3219:12673344 3170:24058600 3130:PLOS ONE 3111:22275897 2945:32052487 2777:13165698 2723:13165697 2613:28509964 2556:20961976 2434:11157465 2350:30259362 2285:22539728 2179:28649208 2171:12851415 1839:Dystonia 1813:See also 1746:resonant 1731:possess 1690:bivalves 1682:mollusks 1651:such as 1649:annelids 1389:T-tubule 972:avulsion 750:(ADP) + 461:reflexes 366:pitching 293:proteins 277:shoulder 155:isotonic 128:myogenic 4249:muscles 4114:Forearm 3959:1692762 3950:3066716 3902:2599245 3754:9983649 3680:4348240 3672:2348872 3652:Bibcode 3596:7858131 3587:1225569 3566:Bibcode 3528:8235594 3508:Bibcode 3500:Science 3476:5850239 3372:2927862 3315:4143026 3284:Bibcode 3257:6346892 3161:3776813 3138:Bibcode 3102:3257849 3052:5921536 3043:1357553 3003:1770255 2958:5 April 2834:3680378 2825:2114850 2785:4180166 2757:Bibcode 2731:4275495 2703:Bibcode 2604:5425715 2547:2964179 2425:1724276 2342:9617396 2244:2275403 2036:323–335 1795:and by 1778:calcium 1752:History 1729:beetles 1657:leeches 992:tetanus 932:tetanus 701:and by 691:muscles 649:by the 540:causes 451:to the 312:glucose 238:triceps 190:forearm 49:tension 4189:Origin 4131:Tendon 4082:Fascia 4053:Tissue 3966:  3956:  3948:  3909:  3899:  3829:  3821:  3780:  3752:  3746:766843 3744:  3704:  3678:  3670:  3644:Nature 3627:  3617:  3594:  3584:  3526:  3483:  3473:  3465:  3426:  3418:  3379:  3369:  3361:  3322:  3312:  3304:  3255:  3217:  3209:  3168:  3158:  3109:  3099:  3091:  3050:  3040:  3001:  2995:892834 2993:  2951:  2943:  2885:  2860:  2832:  2822:  2783:  2775:  2749:Nature 2729:  2721:  2695:Nature 2673:  2650:  2611:  2601:  2593:  2554:  2544:  2536:  2480:  2455:  2432:  2422:  2348:  2340:  2283:  2242:  2177:  2169:  2132:  2078:  2042:  1998:  1970:  1945:  1806:Nature 1727:, and 1684:, and 1401:triads 1278:, and 928:twitch 900:Twitch 757:. Two 558:sodium 538:influx 526:. The 496:and a 415:, the 411:forms 339:myosin 285:Desmin 230:biceps 194:joints 120:smooth 109:twitch 78:myosin 4296:Other 4177:Other 4144:Shape 4119:Thigh 3946:JSTOR 3750:S2CID 3676:S2CID 3453:(3). 3077:: 5. 2999:S2CID 2949:S2CID 2781:S2CID 2727:S2CID 2346:S2CID 2318:(PDF) 2175:S2CID 1884:Spasm 1834:Cramp 1721:flies 1717:wasps 1485:in a 1276:DAPK3 1133:or a 1019:titin 953:. In 740:actin 712:titin 488:is a 455:that 343:actin 289:titin 226:elbow 161:Types 99:from 71:actin 55:. 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Index

Muscular contraction

smooth muscle
tension
muscle cells
physiology
actin
motor-protein
myosin
vertebrates
skeletal muscle contractions
neurogenic
synaptic input
motor neurons
sliding filament theory
twitch
action potentials
smooth
cardiac muscles
myogenic
autonomic nervous system
muscle tissues
locomotor activity
skeletal muscles
isometric
isotonic

Isometric exercise
forearm
joints

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