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Muscle contraction is the activation of
tension Tension may refer to: Science * Psychological stress * Tension (physics), a force related to the stretching of an object (the opposite of compression) * Tension (geology), a stress which stretches rocks in two opposite directions * Voltage or el ...
-generating sites within muscle cells. In
physiology Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a sub-discipline of biology, physiology focuses on how organisms, organ systems, individual organs, cells, and biomolecules carry out the chemical ...
, 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 muscle relaxation, which is a return of the muscle fibers to their low tension-generating state. For the contractions to happen, the muscle cells must rely on the interaction of two types of filaments which are the thin and thick filaments. Thin filaments are two strands of actin coiled around each, and thick filaments consist of mostly elongated proteins called myosin. Together, these two filaments form myofibrils which are important organelles in the skeletal muscle system. Muscle contraction can also be described based on two variables: length and tension. A muscle contraction is described as isometric if the muscle tension changes but the muscle length remains the same. In contrast, a muscle contraction is isotonic 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. In natural movements that underlie
locomotor activity Animal locomotion, in ethology, is any of a variety of methods that animals use to move from one place to another. Some modes of locomotion are (initially) self-propelled, e.g., running, swimming, jumping, flying, hopping, soaring and gliding. Th ...
, 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
skeletal muscle Skeletal muscles (commonly referred to as muscles) are organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other types of muscl ...
s that contract during locomotor activity. In
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
s, skeletal muscle contractions are
neurogenic In biology, the nervous system is the highly complex part of an animal that coordinates its actions and sensory information by transmitting signals to and from different parts of its body. The nervous system detects environmental changes ...
as they require synaptic input from
motor neurons A motor neuron (or motoneuron or efferent neuron) is a neuron whose cell body is located in the motor cortex, brainstem or the spinal cord, and whose axon (fiber) projects to the spinal cord or outside of the spinal cord to directly or indirectly ...
. 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
sliding filament theory The sliding filament theory explains the mechanism of muscle contraction based on muscle proteins that slide past each other to generate movement. According to the sliding filament theory, the myosin ( thick filaments) of muscle fibers slide past ...
. The contraction produced can be described as a
twitch Twitch may refer to: Biology * Muscle contraction ** Convulsion, rapid and repeated muscle contraction and relaxation ** Fasciculation, a small, local, involuntary muscle contraction ** Myoclonic twitch, a jerk usually caused by sudden muscle co ...
, summation, or tetanus, depending on the frequency of
action potentials An action potential occurs when the membrane potential of a specific cell location rapidly rises and falls. This depolarization then causes adjacent locations to similarly depolarize. Action potentials occur in several types of animal cells ...
. 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. Unlike skeletal muscle, the contractions of
smooth Smooth may refer to: Mathematics * Smooth function, a function that is infinitely differentiable; used in calculus and topology * Smooth manifold, a differentiable manifold for which all the transition maps are smooth functions * Smooth algebrai ...
and
cardiac muscle Cardiac muscle (also called heart muscle, myocardium, cardiomyocytes and cardiac myocytes) is one of three types of vertebrate muscle tissues, with the other two being skeletal muscle and smooth muscle. It is an involuntary, striated muscle th ...
s are
myogenic The myogenic mechanism is how arteries and arterioles react to an increase or decrease of blood pressure to keep the blood flow constant within the blood vessel. Myogenic response refers to a contraction initiated by the myocyte itself instead of ...
(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 autonomic nervous system. The mechanisms of contraction in these
muscle tissue Muscle tissue (or muscular tissue) is soft tissue that makes up the different types of muscles in most animals, and give the ability of muscles to contract. Muscle tissue is formed during embryonic development, in a process known as myogenesis. ...
s are similar to those in skeletal muscle tissues.


Types

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 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.


Isometric contraction

An isometric contraction of a muscle generates tension without changing length. An example can be found when the muscles of the
hand A hand is a prehensile, multi-fingered appendage located at the end of the forearm or forelimb of primates such as humans, chimpanzees, monkeys, and lemurs. A few other vertebrates such as the koala (which has two opposable thumbs on each "h ...
and
forearm The forearm is the region of the upper limb between the elbow and the wrist. The term forearm is used in anatomy to distinguish it from the arm, a word which is most often used to describe the entire appendage of the upper limb, but which in ...
grip an object; the
joint A joint or articulation (or articular surface) is the connection made between bones, ossicles, or other hard structures in the body which link an animal's skeletal system into a functional whole.Saladin, Ken. Anatomy & Physiology. 7th ed. McGraw ...
s of the hand do not move, but muscles generate sufficient force to prevent the object from being dropped.


Isotonic contraction

In
isotonic contraction In an isotonic contraction, tension remains the same, whilst the muscle's length changes. Isotonic contractions differ from isokinetic contractions in that in isokinetic contractions the muscle speed remains constant. While superficially identic ...
, 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.


Concentric contraction

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. During a concentric contraction, a muscle is stimulated to contract according to the
sliding filament theory The sliding filament theory explains the mechanism of muscle contraction based on muscle proteins that slide past each other to generate movement. According to the sliding filament theory, the myosin ( thick filaments) of muscle fibers slide past ...
. 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
elbow The elbow is the region between the arm and the forearm that surrounds the elbow joint. The elbow includes prominent landmarks such as the olecranon, the cubital fossa (also called the chelidon, or the elbow pit), and the lateral and the media ...
, a concentric contraction of the biceps would cause the
arm In human anatomy, the arm refers to the upper limb in common usage, although academically the term specifically means the upper arm between the glenohumeral joint (shoulder joint) and the elbow joint. The distal part of the upper limb between th ...
to bend at the elbow as the hand moved from the leg to the shoulder (a
biceps curl The term "biceps curl" refers to any of a number of weight training exercises that primarily targets the biceps brachii muscle. It may be performed using a barbell, dumbbell, resistance band, or other equipment. Overview The biceps curl mainly ...
). A concentric contraction of the
triceps The triceps, or triceps brachii (Latin for "three-headed muscle of the arm"), is a large muscle on the back of the upper limb of many vertebrates. It consists of 3 parts: the medial, lateral, and long head. It is the muscle principally responsibl ...
would change the angle of the joint in the opposite direction, straightening the arm and moving the hand towards the leg.


Eccentric contraction

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 decelerate 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 walking). Over the short-term,
strength training Strength training or resistance training involves the performance of physical exercises that are designed to improve strength and endurance. It is often associated with the lifting of weights. It can also incorporate a variety of training te ...
involving both eccentric and concentric contractions appear to increase
muscular strength Physical strength is the measure of a human's exertion of force on physical objects. Increasing physical strength is the goal of strength training. Overview An individual's physical strength is determined by two factors: the cross-sectional are ...
more than training with concentric contractions alone. However, exercise-induced muscle damage is also greater during lengthening contractions. During an eccentric contraction of the biceps muscle, the
elbow The elbow is the region between the arm and the forearm that surrounds the elbow joint. The elbow includes prominent landmarks such as the olecranon, the cubital fossa (also called the chelidon, or the elbow pit), and the lateral and the media ...
starts the movement while bent and then straightens as the hand moves away from the
shoulder The human shoulder is made up of three bones: the clavicle (collarbone), the scapula (shoulder blade), and the humerus (upper arm bone) as well as associated muscles, ligaments and tendons. The articulations between the bones of the shoulder mak ...
. During an eccentric contraction of the
triceps muscle The triceps, or triceps brachii (Latin for "three-headed muscle of the arm"), is a large muscle on the back of the upper limb of many vertebrates. It consists of 3 parts: the medial, lateral, and long head. It is the muscle principally responsib ...
, the elbow starts the movement straight and then bends as the hand moves towards the shoulder.
Desmin Desmin is a protein that in humans is encoded by the ''DES'' gene. Desmin is a muscle-specific, type III intermediate filament that integrates the sarcolemma, Z disk, and nuclear membrane in sarcomeres and regulates sarcomere architecture. Str ...
,
titin Titin (contraction for Titan protein) (also called connectin) is a protein that in humans is encoded by the ''TTN'' gene. Titin is a giant protein, greater than 1 µm in length, that functions as a molecular spring that is responsible for th ...
, and other z-line
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
s are involved in eccentric contractions, but their mechanism is poorly understood in comparison to
cross-bridge cycling Muscle contraction is the activation of Tension (physics), tension-generating sites within muscle cells. In physiology, muscle contraction does not necessarily mean muscle shortening because muscle tension can be produced without changes in muscl ...
in concentric contractions. Though the muscle is doing a negative amount of
mechanical work In physics, work is the energy transferred to or from an object via the application of force along a displacement. In its simplest form, for a constant force aligned with the direction of motion, the work equals the product of the force stre ...
, (work is being done ''on'' the muscle), chemical energy (of
fat In nutrition science, nutrition, biology, and chemistry, fat usually means any ester of fatty acids, or a mixture of such chemical compound, compounds, most commonly those that occur in living beings or in food. The term often refers spec ...
or
glucose Glucose is a simple sugar with the molecular formula . Glucose is overall the most abundant monosaccharide, a subcategory of carbohydrates. Glucose is mainly made by plants and most algae during photosynthesis from water and carbon dioxide, u ...
, or temporarily stored in ATP) 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. Muscles undergoing heavy eccentric loading suffer greater damage when overloaded (such as during muscle building or
strength training Strength training or resistance training involves the performance of physical exercises that are designed to improve strength and endurance. It is often associated with the lifting of weights. It can also incorporate a variety of training te ...
exercise) as compared to concentric loading. When eccentric contractions are used in weight training, they are normally called ''negatives''. During a concentric contraction, contractile muscle
myofilament Myofilaments are the three protein filaments of myofibrils in muscle cells. The main proteins involved are myosin, actin, and titin. Myosin and actin are the ''contractile proteins'' and titin is an elastic protein. The myofilaments act togethe ...
s of
myosin Myosins () are a superfamily of motor proteins best known for their roles in muscle contraction and in a wide range of other motility processes in eukaryotes. They are ATP-dependent and responsible for actin-based motility. The first myosin ...
and
actin Actin is a family of globular multi-functional proteins that form microfilaments in the cytoskeleton, and the thin filaments in muscle fibrils. It is found in essentially all eukaryotic cells, where it may be present at a concentration of over ...
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 greater weight (muscles are approximately 40% stronger during eccentric contractions than during concentric contractions) and also results in greater muscular damage and
delayed onset muscle soreness Delayed onset muscle soreness (DOMS) is the pain and stiffness felt in muscles after unaccustomed or strenuous exercise. The soreness is felt most strongly 24 to 72 hours after the exercise. It is thought to be caused by eccentric (lengthening) ex ...
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 contractions can easily lead to
overtraining Overtraining occurs when a person exceeds their body's ability to recover from strenuous exercise. Overtraining can be described as a point where a person may have a decrease in performance and plateauing as a result of failure to consistently perfo ...
, moderate training may confer protection against injury.


= Eccentric contractions in movement

= 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 training for rapid movements such as pitching during baseball involves reducing eccentric braking allowing a greater power to be developed throughout the movement. Eccentric contractions are being researched for their ability to speed rehabilitation of weak or injured tendons. Achilles tendinitis and
patellar tendonitis Patellar tendinitis, also known as jumper's knee, is an overuse injury of the tendon that straightens the knee. Symptoms include pain in the front of the knee. Typically the pain and tenderness is at the lower part of the kneecap, though the uppe ...
(also known as jumper's knee or patellar tendonosis) have been shown to benefit from high-load eccentric contractions.


Vertebrate

In vertebrate animals, there are three types of
muscle tissue Muscle tissue (or muscular tissue) is soft tissue that makes up the different types of muscles in most animals, and give the ability of muscles to contract. Muscle tissue is formed during embryonic development, in a process known as myogenesis. ...
s: skeletal, smooth, and cardiac.
Skeletal muscle Skeletal muscles (commonly referred to as muscles) are organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other types of muscl ...
constitutes the majority of muscle mass in the body and is responsible for locomotor activity. Smooth muscle forms blood vessels,
gastrointestinal tract The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organ (biology), organs of the digestive syste ...
, and other areas in the body that produce sustained contractions.
Cardiac muscle Cardiac muscle (also called heart muscle, myocardium, cardiomyocytes and cardiac myocytes) is one of three types of vertebrate muscle tissues, with the other two being skeletal muscle and smooth muscle. It is an involuntary, striated muscle th ...
make up the heart, which pumps blood. Skeletal and cardiac muscles are called
striated muscle Striations means a series of ridges, furrows or linear marks, and is used in several ways: * Glacial striation * Striation (fatigue), in material * Striation (geology), a ''striation'' as a result of a geological fault * Striation Valley, in An ...
because of their striped appearance under a microscope, which is due to the highly organized alternating pattern of A bands and I bands.


Skeletal muscle

Excluding reflexes, all
skeletal muscle Skeletal muscles (commonly referred to as muscles) are organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other types of muscl ...
s contractions occur as a result of signals originating in the
brain A brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as vision. It is the most complex organ in a v ...
. The brain sends electrochemical signals through the
nervous system In biology, the nervous system is the highly complex part of an animal that coordinates its actions and sensory information by transmitting signals to and from different parts of its body. The nervous system detects environmental changes th ...
to the
motor neuron A motor neuron (or motoneuron or efferent neuron) is a neuron whose cell body is located in the motor cortex, brainstem or the spinal cord, and whose axon (fiber) projects to the spinal cord or outside of the spinal cord to directly or indirectl ...
that innervates several muscle fibers. In the case of some reflexes, the signal to contract can originate in the
spinal cord The spinal cord is a long, thin, tubular structure made up of nervous tissue, which extends from the medulla oblongata in the brainstem to the lumbar region of the vertebral column (backbone). The backbone encloses the central canal of the spi ...
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 both consciously or unconsciously.


Neuromuscular junction

A neuromuscular junction is a
chemical synapse Chemical synapses are biological junctions through which neurons' signals can be sent to each other and to non-neuronal cells such as those in muscles or glands. Chemical synapses allow neurons to form circuits within the central nervous syste ...
formed by the contact between a
motor neuron A motor neuron (or motoneuron or efferent neuron) is a neuron whose cell body is located in the motor cortex, brainstem or the spinal cord, and whose axon (fiber) projects to the spinal cord or outside of the spinal cord to directly or indirectl ...
and a
muscle fiber A muscle cell is also known as a myocyte when referring to either a cardiac muscle cell (cardiomyocyte), or a smooth muscle cell as these are both small cells. A skeletal muscle cell is long and threadlike with many nuclei and is called a muscl ...
. 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
saltatory conduction In neuroscience, saltatory conduction () is the propagation of action potentials along myelinated axons from one node of Ranvier to the next node, increasing the conduction velocity of action potentials. The uninsulated nodes of Ranvier are th ...
along its axon toward the neuromuscular junction. Once it reaches the terminal bouton, the action potential causes a ion influx into the terminal by way of the
voltage-gated calcium channel Voltage-gated calcium channels (VGCCs), also known as voltage-dependent calcium channels (VDCCs), are a group of voltage-gated ion channels found in the membrane of excitable cells (''e.g.'', muscle, glial cells, neurons, etc.) with a permeabili ...
s. The influx causes
synaptic vesicles In a neuron, synaptic vesicles (or neurotransmitter vesicles) store various neurotransmitters that are released at the synapse. The release is regulated by a voltage-dependent calcium channel. Vesicles are essential for propagating nerve impul ...
containing the neurotransmitter
acetylcholine Acetylcholine (ACh) is an organic chemical that functions in the brain and body of many types of animals (including humans) as a neurotransmitter. Its name is derived from its chemical structure: it is an ester of acetic acid and choline. Part ...
to fuse with the plasma membrane, releasing acetylcholine into the
synaptic cleft Chemical synapses are biological junctions through which neurons' signals can be sent to each other and to non-neuronal cells such as those in muscles or glands. Chemical synapses allow neurons to form circuits within the central nervous syste ...
between the motor neuron terminal and the neuromuscular junction of the skeletal muscle fiber. Acetylcholine diffuses across the synapse and binds to and activates
nicotinic acetylcholine receptors Nicotinic acetylcholine receptors, or nAChRs, are receptor polypeptides that respond to the neurotransmitter acetylcholine. Nicotinic receptors also respond to drugs such as the agonist nicotine. They are found in the central and peripheral ...
on the neuromuscular junction. Activation of the nicotinic receptor opens its intrinsic
sodium Sodium is a chemical element with the symbol Na (from Latin ''natrium'') and atomic number 11. It is a soft, silvery-white, highly reactive metal. Sodium is an alkali metal, being in group 1 of the periodic table. Its only stable iso ...
/
potassium Potassium is the chemical element with the symbol K (from Neo-Latin ''kalium'') and atomic number19. Potassium is a silvery-white metal that is soft enough to be cut with a knife with little force. Potassium metal reacts rapidly with atmosphe ...
channel, causing sodium to rush in and potassium to trickle out. As a result, the
sarcolemma The sarcolemma (''sarco'' (from ''sarx'') from Greek; flesh, and ''lemma'' from Greek; sheath) also called the myolemma, is the cell membrane surrounding a skeletal muscle fiber or a cardiomyocyte. It consists of a lipid bilayer and a thin oute ...
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 end-plate potentialSaladin, 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. 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
acetylcholine esterase Acetylcholinesterase (HGNC symbol ACHE; EC 3.1.1.7; systematic name acetylcholine acetylhydrolase), also known as AChE, AChase or acetylhydrolase, is the primary cholinesterase in the body. It is an enzyme that catalyzes the breakdown of acet ...
or reabsorbed by the synaptic knob and none is left to replace the degraded acetylcholine.


Excitation–contraction coupling

Excitation–contraction coupling is the process by which a muscular action potential in the muscle fiber causes the
myofibril A myofibril (also known as a muscle fibril or sarcostyle) is a basic rod-like organelle of a muscle cell. Skeletal muscles are composed of long, tubular cells known as muscle fibers, and these cells contain many chains of myofibrils. Each myo ...
s to contract. In skeletal muscle, excitation–contraction coupling relies on a direct coupling between key proteins, the
sarcoplasmic reticulum The sarcoplasmic reticulum (SR) is a membrane-bound structure found within muscle cells that is similar to the smooth endoplasmic reticulum in other cells. The main function of the SR is to store calcium ions (Ca2+). Calcium ion levels are ke ...
(SR) calcium release channel (identified as the
ryanodine receptor 1 Ryanodine receptor 1 (RYR-1) also known as skeletal muscle calcium release channel or skeletal muscle-type ryanodine receptor is one of a class of ryanodine receptors and a protein found primarily in skeletal muscle. In humans, it is encoded by th ...
, RYR1) and voltage-gated L-type calcium channels (identified as dihydropyridine receptors, DHPRs). DHPRs are located on the sarcolemma (which includes the surface sarcolemma and the
transverse tubules T-tubules (transverse tubules) are extensions of the cell membrane that penetrate into the center of skeletal and cardiac muscle cells. With membranes that contain large concentrations of ion channels, transporters, and pumps, T-tubules permit ...
), 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. Excitation–contraction coupling occurs when depolarization of skeletal muscle cell results in a muscle action potential, which spreads across the cell surface and into the muscle fiber's network of
T-tubule T-tubules (transverse tubules) are extensions of the cell membrane that penetrate into the center of skeletal and cardiac muscle cells. With membranes that contain large concentrations of ion channels, transporters, and pumps, T-tubules permi ...
s, thereby depolarizing the inner portion of the muscle fiber. Depolarization of the inner portions activates dihydropyridine receptors in the terminal cisternae, which are in close proximity to ryanodine receptors in the adjacent
sarcoplasmic reticulum The sarcoplasmic reticulum (SR) is a membrane-bound structure found within muscle cells that is similar to the smooth endoplasmic reticulum in other cells. The main function of the SR is to store calcium ions (Ca2+). Calcium ion levels are ke ...
. 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 the ryanodine receptors open, is released from the sarcoplasmic reticulum into the local junctional space and diffuses into the bulk cytoplasm to cause a
calcium spark A calcium spark is the microscopic release of calcium ( Ca2+) from a store known as the sarcoplasmic reticulum (SR), located within muscle cells. This release occurs through an ion channel within the membrane of the SR, known as a ryanodine rec ...
. Note that the sarcoplasmic reticulum has a large
calcium buffering Calcium buffering describes the processes which help stabilise the concentration of free calcium ions within cells, in a similar manner to how pH buffers maintain a stable concentration of hydrogen ions. The majority of calcium ions within the cel ...
capacity partially due to a
calcium-binding protein Calcium-binding proteins are proteins that participate in calcium cell signalling pathways by binding to Ca2+, the calcium ion that plays an important role in many cellular processes. Calcium-binding proteins have specific domains that bind to cal ...
called
calsequestrin Calsequestrin is a calcium-binding protein that acts as a calcium buffer within the sarcoplasmic reticulum. The protein helps hold calcium in the cisterna of the sarcoplasmic reticulum after a muscle contraction, even though the concentration ...
. The near synchronous activation of thousands of
calcium sparks A calcium spark is the microscopic release of calcium ( Ca2+) from a store known as the sarcoplasmic reticulum (SR), located within muscle cells. This release occurs through an ion channel within the membrane of the SR, known as a ryanodine rece ...
by the action potential causes a cell-wide increase in calcium giving rise to the upstroke of the calcium transient. The released into the cytosol binds to
Troponin C Troponin C is a protein which is part of the troponin complex. It contains four calcium-binding EF hands, although different isoforms may have fewer than four functional calcium-binding subdomains. It is a component of thin filaments, along wi ...
by the actin filaments, to allow
cross-bridge cycling Muscle contraction is the activation of Tension (physics), tension-generating sites within muscle cells. In physiology, muscle contraction does not necessarily mean muscle shortening because muscle tension can be produced without changes in muscl ...
, producing force and, in some situations, motion. The sarco/endoplasmic reticulum calcium-ATPase (SERCA) actively pumps back into the sarcoplasmic reticulum. As declines back to resting levels, the force declines and relaxation occurs.


Sliding filament theory

The
sliding filament theory The sliding filament theory explains the mechanism of muscle contraction based on muscle proteins that slide past each other to generate movement. According to the sliding filament theory, the myosin ( thick filaments) of muscle fibers slide past ...
describes a process used by
muscle Skeletal muscles (commonly referred to as muscles) are organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other types of muscl ...
s 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
Andrew Huxley Sir Andrew Fielding Huxley (22 November 191730 May 2012) was an English physiologist and biophysicist. He was born into the prominent Huxley family. After leaving Westminster School in central London, he went to Trinity College, Cambridge ...
and
Rolf Niedergerke Rolf Nidergerke (30 April 1921 – 27 December 2011) was a German physiologist and physician, and one of the discoverers of the sliding filament theory of muscle contraction. He and Andrew Huxley, complimenting the independent works of Hugh Huxl ...
and by
Hugh Huxley Hugh Esmor Huxley MBE FRS (25 February 1924 – 25 July 2013) was a British molecular biologist who made important discoveries in the physiology of muscle. He was a graduate in physics from Christ's College, Cambridge. However, his education ...
and
Jean Hanson Emmeline Jean Hanson (14 November 1919 – 10 August 1973) was a biophysicist and zoologist known for her contributions to muscle research. Hanson gained her PhD in zoology from Bedford College, University of London before spending the majorit ...
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
titin Titin (contraction for Titan protein) (also called connectin) is a protein that in humans is encoded by the ''TTN'' gene. Titin is a giant protein, greater than 1 µm in length, that functions as a molecular spring that is responsible for th ...
. This fine myofilament maintains uniform tension across the sarcomere by pulling the thick filament into a central position.


=Cross-bridge cycle

= Cross-bridge cycling is a sequence of molecular events that underlies the sliding filament theory. A cross-bridge is a myosin projection, consisting of two myosin heads, that extends from the thick filaments. Each myosin head has two binding sites: one for
adenosine triphosphate Adenosine triphosphate (ATP) is an organic compound that provides energy to drive many processes in living cells, such as muscle contraction, nerve impulse propagation, condensate dissolution, and chemical synthesis. Found in all known forms o ...
(ATP) and another for actin. The binding of ATP to a myosin head detaches myosin from
actin Actin is a family of globular multi-functional proteins that form microfilaments in the cytoskeleton, and the thin filaments in muscle fibrils. It is found in essentially all eukaryotic cells, where it may be present at a concentration of over ...
, 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
tropomyosin Tropomyosin is a two-stranded alpha-helical, coiled coil protein found in actin-based cytoskeletons. Tropomyosin and the actin skeleton All organisms contain organelles that provide physical integrity to their cells. These type of organelles a ...
. With the ATP hydrolyzed, the cocked myosin head now contains adenosine diphosphate (ADP) + Pi. Two ions bind to
troponin C Troponin C is a protein which is part of the troponin complex. It contains four calcium-binding EF hands, although different isoforms may have fewer than four functional calcium-binding subdomains. It is a component of thin filaments, along wi ...
on the actin filaments. The troponin- 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 ''power stroke,'' which generates a force of 2 pN. The power stroke moves the actin filament inwards, thereby shortening the
sarcomere A sarcomere (Greek σάρξ ''sarx'' "flesh", μέρος ''meros'' "part") is the smallest functional unit of striated muscle tissue. It is the repeating unit between two Z-lines. Skeletal muscles are composed of tubular muscle cells (called mus ...
. 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
rigor mortis Rigor mortis (Latin: ''rigor'' "stiffness", and ''mortis'' "of death"), or postmortem rigidity, is the third stage of death. It is one of the recognizable signs of death, characterized by stiffening of the limbs of the corpse caused by chemic ...
. Once another ATP binds to myosin, the myosin head will again detach from actin and another cross-bridge cycle occurs. Cross-bridge cycling is able to continue as long as there are sufficient amounts of ATP and in the cytoplasm. Termination of cross-bridge cycling can occur when is actively pumped back into the sarcoplasmic reticulum. When 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 ions leave the troponin molecule in order to maintain the ion concentration in the sarcoplasm. The active pumping of ions into the sarcoplasmic reticulum creates a deficiency in the fluid around the myofibrils. This causes the removal of ions from the troponin. Thus, the tropomyosin-troponin complex again covers the binding sites on the actin filaments and contraction ceases.


Gradation of skeletal muscle contractions

The strength of skeletal muscle contractions can be broadly separated into twitch, summation, and
tetanus Tetanus, also known as lockjaw, is a bacterial infection caused by ''Clostridium tetani'', and is characterized by muscle spasms. In the most common type, the spasms begin in the jaw and then progress to the rest of the body. Each spasm usually ...
. 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 latent period, which usually takes about 10 ms and is caused by the time taken for nerve action potential to propagate, the time for chemical transmission at the neuromuscular junction, then the subsequent steps in excitation-contraction coupling. 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 ''summation''. Summation can be achieved in two ways: ''frequency summation'' and ''multiple fiber summation''. In frequency summation, the force exerted by the skeletal muscle is controlled by varying the frequency at which
action potential An action potential occurs when the membrane potential of a specific cell location rapidly rises and falls. This depolarization then causes adjacent locations to similarly depolarize. Action potentials occur in several types of animal cells, ...
s 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 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
Golgi tendon organ The Golgi tendon organ (GTO) (also called Golgi organ, tendon organ, neurotendinous organ or neurotendinous spindle) is a proprioceptor – a type of sensory receptor that senses changes in muscle tension. It lies at the interface between a musc ...
s and
Renshaw cell Renshaw cells are inhibitory interneurons found in the gray matter of the spinal cord, and are associated in two ways with an alpha motor neuron. * They receive an excitatory collateral from the alpha neuron's axon as they emerge from the motor ...
s). This 'low' level of contraction is a protective mechanism to prevent avulsion of the tendon—the force generated by a 95% contraction of all fibers is sufficient to damage the body. In multiple fiber summation, if the central nervous system sends a weak signal to contract a muscle, the smaller motor units, being more excitable than the larger ones, are stimulated first. As the strength of the signal 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 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. 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 ''tetanus''.


Length-tension relationship

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 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 titin) 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.


Force-velocity relationships

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 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.


Smooth muscle

Smooth muscles can be divided into two subgroups: single-unit and multiunit. 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
syncytium A syncytium (; plural syncytia; from Greek: σύν ''syn'' "together" and κύτος ''kytos'' "box, i.e. cell") or symplasm is a multinucleate cell which can result from multiple cell fusions of uninuclear cells (i.e., cells with a single nucleus) ...
. Single-unit smooth muscle cells contract myogenically, which can be modulated by the autonomic nervous system. 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 responses, much like
motor unit recruitment Motor unit recruitment is the activation of additional motor units to accomplish an increase in contractile strength in a muscle. A motor unit consists of one motor neuron and all of the muscle fibers it stimulates. All muscles consist of a nu ...
in skeletal muscle.


Mechanisms of smooth muscle contraction

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 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
pacemaker potential In the pacemaking cells of the heart (e.g., the sinoatrial node), the pacemaker potential (also called the pacemaker current) is the slow, positive increase in voltage across the cell's membrane (the membrane potential) that occurs between the en ...
or a
slow wave potential A slow-wave potential is a rhythmic electrophysiological event in the gastrointestinal tract. The normal conduction of slow waves is one of the key regulators of gastrointestinal motility. Slow waves are generated and propagated by a class of pa ...
. These action potentials are generated by the influx of extracellular , and not . Like skeletal muscles, cytosolic ions are also required for crossbridge cycling in smooth muscle cells. The two sources for cytosolic in smooth muscle cells are the extracellular entering through calcium channels and the ions that are released from the sarcoplasmic reticulum. The elevation of cytosolic results in more binding to
calmodulin Calmodulin (CaM) (an abbreviation for calcium-modulated protein) is a multifunctional intermediate calcium-binding messenger protein expressed in all eukaryotic cells. It is an intracellular target of the secondary messenger Ca2+, and the bind ...
, which then binds and activates
myosin light-chain kinase Myosin light-chain kinase also known as MYLK or MLCK is a serine/threonine-specific protein kinase that phosphorylates a specific myosin light chain, namely, the regulatory light chain of myosin II. General structural features While there ar ...
. The calcium-calmodulin-myosin light-chain kinase complex phosphorylates myosin on the 20
kilodalton The dalton or unified atomic mass unit (symbols: Da or u) is a non-SI unit of mass widely used in physics and chemistry. It is defined as of the mass of an unbound neutral atom of carbon-12 in its nuclear and electronic ground state and at ...
(kDa) myosin light chains on amino acid residue-serine 19, initiating contraction and activating the
myosin ATPase Myosin ATPase () is an enzyme with List of enzymes, systematic name ''ATP phosphohydrolase (actin-translocating)''. This enzyme catalysis, catalyses the following chemical reaction : ATP + H2O \rightleftharpoons ADP + phosphate ATP hydrolysis pr ...
. 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 -activated phosphorylation of myosin rather than binding to the troponin complex that regulates myosin binding sites on actin like in skeletal and cardiac muscles. 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 period, there is a rapid burst of energy utilization 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 utilization 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
rho kinase Rho-associated protein kinase (ROCK) is a kinase belonging to the AGC (PKA/ PKG/PKC) family of serine-threonine specific protein kinases. It is involved mainly in regulating the shape and movement of cells by acting on the cytoskeleton. ROCKs ( ...
,
DAPK3 Death-associated protein kinase 3 is an enzyme that in humans is encoded by the ''DAPK3'' gene. Function Death-associated protein kinase 3 (DAPK3) induces morphological changes in apoptosis when overexpressed in mammalian cells. These resul ...
, and protein kinase C are believed to participate in the sustained phase of contraction, and flux may be significant.


Neuromodulation

Although smooth muscle contractions are myogenic, the rate and strength of their contractions can be modulated by the autonomic nervous system.
Postganglionic nerve fibers In the autonomic nervous system, fibers from the ganglion to the effector organ are called postganglionic fibers. Neurotransmitters The neurotransmitters of postganglionic fibers differ: * In the parasympathetic division, neurons are ''cho ...
of
parasympathetic nervous system The parasympathetic nervous system (PSNS) is one of the three divisions of the autonomic nervous system, the others being the sympathetic nervous system and the enteric nervous system. The enteric nervous system is sometimes considered part of ...
release the neurotransmitter acetylcholine, which binds to
muscarinic acetylcholine receptors Muscarinic acetylcholine receptors, or mAChRs, are acetylcholine receptors that form G protein-coupled receptor complexes in the cell membranes of certain neurons and other cells. They play several roles, including acting as the main end-rece ...
(mAChRs) on smooth muscle cells. These receptors are
metabotropic A metabotropic receptor, also referred to by the broader term G-protein-coupled receptor, is a type of membrane receptor that initiates a number of metabolic steps to modulate cell activity. The nervous system utilizes two types of receptors: met ...
, or G-protein coupled receptors that initiate a second messenger cascade. Conversely, postganglionic nerve fibers of the
sympathetic nervous system The sympathetic nervous system (SNS) is one of the three divisions of the autonomic nervous system, the others being the parasympathetic nervous system and the enteric nervous system. The enteric nervous system is sometimes considered part of th ...
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 excitatory (contractile) or inhibitory (relaxing).


Cardiac muscle

There are two types of
cardiac muscle Cardiac muscle (also called heart muscle, myocardium, cardiomyocytes and cardiac myocytes) is one of three types of vertebrate muscle tissues, with the other two being skeletal muscle and smooth muscle. It is an involuntary, striated muscle th ...
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 and are able to contract.


Excitation-contraction coupling

In both skeletal and cardiac muscle excitation-contraction (E-C) coupling, depolarization conduction and Ca2+ release processes occur. However, though the proteins involved are similar, they are distinct in structure and regulation. The dihydropyridine receptors (DHPRs) are encoded by different genes, and the ryanodine receptors (RyRs) are distinct isoforms. Besides, DHPR contacts with RyR1 (main RyR isoform in skeletal muscle) to regulate Ca2+ release in skeletal muscle, while the L-type calcium channel (DHPR on cardiac myocytes) and RyR2 (main RyR isoform in cardiac muscle) are not physically coupled in cardiac muscle, but face with each other by a junctional coupling. Unlike skeletal muscle, E-C coupling in cardiac muscle is thought to depend primarily on a mechanism called calcium-induced calcium release, which is based on the junctional structure between T-tubule and sarcoplasmic reticulum. JPH2, Junctophilin-2 (JPH2) is essential to maintain this structure, as well as the integrity of
T-tubule T-tubules (transverse tubules) are extensions of the cell membrane that penetrate into the center of skeletal and cardiac muscle cells. With membranes that contain large concentrations of ion channels, transporters, and pumps, T-tubules permi ...
. Another protein, REEP5, receptor accessory protein 5 (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 Ca2+ through the L-type calcium channels. After this, cardiac muscle tends to exhibit diad structures, rather than Triad (anatomy), triads. Excitation-contraction coupling in cardiac muscle cells occurs when an action potential is initiated by pacemaker cells in the sinoatrial node or atrioventricular node and conducted to all cells in the heart via gap junctions. The action potential travels along the surface membrane into Cardiac striated muscle#T-tubules, T-tubules (the latter are not seen in all cardiac cell types) and the depolarisation causes extracellular to enter the cell via L-type calcium channels and possibly sodium-calcium exchanger (NCX) during the early part of the cardiac action potential#Phase 2, plateau phase. Although this Ca2+ influx only count for about 10% of the Ca2+ needed for activation, it is relatively larger than that of skeletal muscle. This influx causes a small local increase in intracellular . The increase of intracellular is detected by RyR2 in the membrane of the sarcoplasmic reticulum, which releases in a positive feedback physiological response. This positive feedback is known as calcium-induced calcium release and gives rise to
calcium sparks A calcium spark is the microscopic release of calcium ( Ca2+) from a store known as the sarcoplasmic reticulum (SR), located within muscle cells. This release occurs through an ion channel within the membrane of the SR, known as a ryanodine rece ...
( sparks). The spatial and temporal summation of ~30,000 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 Calcium buffering, calcium buffers, 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 . 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 myosin head pulls the actin filament toward the centre of the sarcomere. Following systole, intracellular calcium is taken up by the SERCA, sarco/endoplasmic reticulum ATPase (SERCA) pump back into the sarcoplasmic reticulum ready for the next cycle to begin. Calcium is also ejected from the cell mainly by the sodium-calcium exchanger (NCX) and, to a lesser extent, a plasma membrane calcium ATPase. Some calcium is also taken up by the mitochondria. An enzyme, phospholamban, serves as a brake for SERCA. At low heart rates, phospholamban is active and slows down the activity of the ATPase so that does not have to leave the cell entirely. At high heart rates, phospholamban is phosphorylated and deactivated thus taking most from the cytoplasm back into the sarcoplasmic reticulum. Once again, Calcium buffering, calcium buffers moderate this fall in concentration, permitting a relatively small decrease in free concentration in response to a large change in total calcium. The falling 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.


Invertebrate


Circular and longitudinal muscles

In annelids such as earthworms and leeches, 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 Coelom#Coelomic fluid, coelomic fluid serves as a Hydrostatic skeleton, hydroskeleton 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 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 peristalsis, which underlies the creeping movement of earthworms.


Obliquely striated muscles

Invertebrates such as annelids, Mollusca, mollusks, and nematodes, 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 Bivalvia, bivalves, the obliquely striated muscles can maintain tension over long periods without using too much energy. Bivalves use these muscles to keep their shells closed.


Asynchronous muscles

Advanced insects such as wasps, Fly, flies, bees, and beetles possess asynchronous muscles that constitute the flight muscles in these animals. These flight muscles are often called ''fibrillar muscles'' 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 ''asynchronous muscles'' 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 beating is made possible because the muscles are connected to a Resonance, resonant system, which is driven to a natural frequency of vibration.


History

In 1780, Luigi Galvani 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 bioelectricity, a field that still studies the electrical patterns and signals in tissues such as nerves and muscles. 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 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 calcium that is called a calcium transient. This increase in calcium activates calcium-sensitive contractile proteins that then use ATP to cause cell shortening. The mechanism for muscle contraction evaded scientists for years and requires continued research and updating. The sliding filament theory was independently developed by Andrew F. Huxley and
Rolf Niedergerke Rolf Nidergerke (30 April 1921 – 27 December 2011) was a German physiologist and physician, and one of the discoverers of the sliding filament theory of muscle contraction. He and Andrew Huxley, complimenting the independent works of Hugh Huxl ...
and by
Hugh Huxley Hugh Esmor Huxley MBE FRS (25 February 1924 – 25 July 2013) was a British molecular biologist who made important discoveries in the physiology of muscle. He was a graduate in physics from Christ's College, Cambridge. However, his education ...
and
Jean Hanson Emmeline Jean Hanson (14 November 1919 – 10 August 1973) was a biophysicist and zoologist known for her contributions to muscle research. Hanson gained her PhD in zoology from Bedford College, University of London before spending the majorit ...
. Their findings were published as two consecutive papers published in the 22 May 1954 issue of ''Nature (journal), Nature'' under the common theme "Structural Changes in Muscle During Contraction".


See also

* Anatomical terms of motion * calcium-induced calcium release * Cardiac action potential * Cramp * Dystonia * Exercise physiology * Fasciculation * Hill's muscle model * Hypnic jerk * In vitro muscle testing * Lombard's paradox * Myoclonus * Rigor mortis * Spasm * Uterine contraction


References


Further reading

* 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. * Krans, J. L. (2010) The Sliding Filament Theory of Muscle Contraction. Nature Education 3(9):66


External links


Sliding Filament Model of Muscle Contraction


{{DEFAULTSORT:Muscle Contraction Exercise physiology Muscular system Skeletal muscle Musculoskeletal system Neurology