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 cell
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 ...
s.
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
The term ''isometric'' comes from the Greek for "having equal measurement".
isometric may mean:
* Cubic crystal system, also called isometric crystal system
* Isometre, a rhythmic technique in music.
* "Isometric (Intro)", a song by Madeon from ...
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, biology, the nervous system is the Complex system, highly complex part of an animal that coordinates its Behavior, actions and Sense, sensory information by transmitting action potential, signals to and from different parts of its ...
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 con ...
, 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, c ...
. 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 algebraic ...
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 autonomic nervous system (ANS), formerly referred to as the vegetative nervous system, is a division of the peripheral nervous system that supplies viscera, internal organs, smooth muscle and glands. The autonomic nervous system is a control ...
. 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. Mu ...
s are similar to those in skeletal muscle tissues.
Types
![1015 Types of Contraction new](https://upload.wikimedia.org/wikipedia/commons/d/d4/1015_Types_of_Contraction_new.jpg)
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 anat ...
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 identica ...
, 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
The biceps or biceps brachii ( la, musculus biceps brachii, "two-headed muscle of the arm") is a large muscle that lies on the front of the upper arm between the shoulder and the elbow. Both heads of the muscle arise on the scapula and join ...
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 the ...
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 targ ...
). 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 t ...
, 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 stren ...
, (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, using ...
, 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
Muscle hypertrophy or muscle building involves a hypertrophy or increase in size of skeletal muscle through a growth in size of its component cells. Two factors contribute to hypertrophy: sarcoplasmic hypertrophy, which focuses more on increased ...
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
Achilles tendinitis, also known as achilles tendinopathy, occurs when the Achilles tendon, found at the back of the ankle, becomes sore. Achilles tendinopathy is accompanied by alterations in the tendon's structure and mechanical properties. The ...
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. Mu ...
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
Smooth muscle is an involuntary non-striated muscle, so-called because it has no sarcomeres and therefore no striations (''bands'' or ''stripes''). It is divided into two subgroups, single-unit and multiunit smooth muscle. Within single-unit mus ...
forms blood vessels
The blood vessels are the components of the circulatory system that transport blood throughout the human body. These vessels transport blood cells, nutrients, and oxygen to the tissues of the body. They also take waste and carbon dioxide away f ...
, 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 Ant ...
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
In biology, a reflex, or reflex action, is an involuntary, unplanned sequence or action and nearly instantaneous response to a Stimulus (physiology), stimulus.
Reflexes are found with varying levels of complexity in organisms with a nervous s ...
, 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
A neuromuscular junction (or myoneural junction) is a chemical synapse between a motor neuron and a muscle fiber.
It allows the motor neuron to transmit a signal to the muscle fiber, causing muscle contraction.
Muscles require innervation to ...
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 syst ...
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 permeabi ...
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 n ...
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 potential[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.] 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 myofib ...
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 Cell (biology), cells. The main function of the SR is to store calcium ions (Ca2+). Calcium in bio ...
(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 receptor
Cav1.1 also known as the calcium channel, voltage-dependent, L type, alpha 1S subunit, (CACNA1S), is a protein which in humans is encoded by the ''CACNA1S'' gene. It is also known as CACNL1A3 and the dihydropyridine receptor (DHPR, so named due ...
s, 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 permit ...
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 Cell (biology), cells. The main function of the SR is to store calcium ions (Ca2+). Calcium in bio ...
. 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 recep ...
. 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 Buffer solution, pH buffers maintain a stable concentration of hydrogen ions. The majority of calcium ion ...
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 calc ...
called calsequestrin
Calsequestrin is a calcium-binding protein that acts as a Calcium buffering, 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 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 recep ...
by the action potential causes a cell-wide increase in calcium giving rise to the upstroke of the calcium transient
Calcium is a chemical element with the Symbol (chemistry), symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties ...
. 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 wit ...
by the actin filaments
Microfilaments, also called actin filaments, are protein filaments in the cytoplasm of eukaryotic cells that form part of the cytoskeleton. They are primarily composed of polymers of actin, but are modified by and interact with numerous other pr ...
, 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 on ...
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 t ...
. 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 of ...
(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 ar ...
. With the ATP hydrolyzed, the cocked myosin head now contains adenosine diphosphate
Adenosine diphosphate (ADP), also known as adenosine pyrophosphate (APP), is an important organic compound in metabolism and is essential to the flow of energy in living cells. ADP consists of three important structural components: a sugar backbon ...
(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 wit ...
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 musc ...
. 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 unit
A motor unit is made up of a motor neuron and all of the skeletal muscle fibers innervated by the neuron's axon terminals, including the neuromuscular junctions between the neuron and the fibres. Groups of motor units often work together as a mot ...
s, 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 muscle
Smooth muscle is an involuntary non-striated muscle, so-called because it has no sarcomeres and therefore no striations (''bands'' or ''stripes''). It is divided into two subgroups, single-unit and multiunit smooth muscle. Within single-unit mus ...
s 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 num ...
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 end ...
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 systematic name ''ATP phosphohydrolase (actin-translocating)''. This enzyme catalyses the following chemical reaction
: ATP + H2O \rightleftharpoons ADP + phosphate
ATP hydrolysis provides energy for actomyosi ...
. 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
In cell biology, Protein kinase C, commonly abbreviated to PKC (EC 2.7.11.13), is a family of protein kinase enzymes that are involved in controlling the function of other proteins through the phosphorylation of hydroxyl groups of serine and t ...
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
The autonomic nervous system (ANS), formerly referred to as the vegetative nervous system, is a division of the peripheral nervous system that supplies viscera, internal organs, smooth muscle and glands. The autonomic nervous system is a control ...
. 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 ''choli ...
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 receptor
Ryanodine receptors (RyR for short) form a class of intracellular calcium channels in various forms of excitable animal tissue like muscles and neurons.
There are three major isoforms of the ryanodine receptor, which are found in different tissu ...
s (RyRs) are distinct isoforms. Besides, DHPR contacts with RyR1
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 t ...
(main RyR isoform in skeletal muscle) to regulate Ca2+ release in skeletal muscle, while the L-type calcium channel
The L-type calcium channel (also known as the dihydropyridine channel, or DHP channel) is part of the high-voltage activated family of voltage-dependent calcium channel.
"L" stands for long-lasting referring to the length of activation. This c ...
(DHPR on cardiac myocytes) and RyR2
Ryanodine receptor 2 (RYR2) is one of a class of ryanodine receptors and a protein found primarily in cardiac muscle. In humans, it is encoded by the ''RYR2'' gene. In the process of cardiac calcium-induced calcium release, RYR2 is the major media ...
(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
Calcium-induced calcium release (CICR) describes a biological process whereby calcium is able to activate calcium release from intracellular Ca2+ stores (e.g., endoplasmic reticulum or sarcoplasmic reticulum). Although CICR was first proposed for ...
, which is based on the junctional structure between T-tubule and sarcoplasmic reticulum. 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 permit ...
. Another protein, 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
Within the muscle tissue of animals and humans, contraction and relaxation of the muscle cells (myocytes) is a highly regulated and rhythmic process. In cardiomyocytes, or cardiac muscle cells, muscular contraction takes place due to movement at a ...
structures, rather than triads.
Excitation-contraction coupling in cardiac muscle cells occurs when an action potential is initiated by pacemaker cells in the sinoatrial node
The sinoatrial node (also known as the sinuatrial node, SA node or sinus node) is an oval shaped region of special cardiac muscle in the upper back wall of the right atrium made up of cells known as pacemaker cells. The sinus node is approximat ...
or atrioventricular node
The atrioventricular node or AV node electrically connects the heart's atria and ventricles to coordinate beating in the top of the heart; it is part of the electrical conduction system of the heart. The AV node lies at the lower back section of t ...
and conducted to all cells in the heart via gap junctions
Gap junctions are specialized intercellular connections between a multitude of animal cell-types. They directly connect the cytoplasm of two cells, which allows various molecules, ions and electrical impulses to directly pass through a regulate ...
. The action potential travels along the surface membrane into T-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 ...
(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
The sodium-calcium exchanger (often denoted Na+/Ca2+ exchanger, exchange protein, or NCX) is an antiporter membrane protein that removes calcium from cells. It uses the energy that is stored in the electrochemical gradient of sodium (Na+) by al ...
(NCX) during the early part of the plateau phase
The human sexual response cycle is a four-stage model of Physiological response, physiological responses to sexual stimulation, which, in order of their occurrence, are the Sexual arousal, excitement, plateau, Orgasm, orgasmic, and Refractory perio ...
. 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
Positive feedback (exacerbating feedback, self-reinforcing feedback) is a process that occurs in a feedback loop which exacerbates the effects of a small disturbance. That is, the effects of a perturbation on a system include an increase in the ...
physiological response. This positive feedback is known as calcium-induced calcium release
Calcium-induced calcium release (CICR) describes a biological process whereby calcium is able to activate calcium release from intracellular Ca2+ stores (e.g., endoplasmic reticulum or sarcoplasmic reticulum). Although CICR was first proposed for ...
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 recep ...
( 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 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 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
The sodium-calcium exchanger (often denoted Na+/Ca2+ exchanger, exchange protein, or NCX) is an antiporter membrane protein that removes calcium from cells. It uses the energy that is stored in the electrochemical gradient of sodium (Na+) by al ...
(NCX) and, to a lesser extent, a plasma membrane calcium ATPase
Ca2+ ATPase is a form of P-ATPase that transfers calcium after a muscle has contracted. The two kinds of calcium ATPase are:
*Plasma membrane Ca2+ ATPase (PMCA)
*Sarcoplasmic reticulum Ca2+ ATPase (SERCA)
Plasma membrane Ca2+ ATPase (PMCA)
Pl ...
. Some calcium is also taken up by the mitochondria. An enzyme, phospholamban
Phospholamban, also known as PLN or PLB, is a micropeptide protein that in humans is encoded by the ''PLN'' gene. Phospholamban is a 52-amino acid integral membrane protein that regulates the calcium (Ca2+) pump in cardiac muscle cells.
Funct ...
, 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 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
The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecolo ...
such as earthworms
An earthworm is a terrestrial invertebrate that belongs to the phylum Annelida. They exhibit a tube-within-a-tube body plan; they are externally segmented with corresponding internal segmentation; and they usually have setae on all segments. Th ...
and leeches
Leeches are segmented parasitism, parasitic or Predation, predatory worms that comprise the Class (biology), subclass Hirudinea within the phylum Annelida. They are closely related to the Oligochaeta, oligochaetes, which include the earthwor ...
, 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 coelomic fluid serves as a hydroskeleton
A hydrostatic skeleton, or hydroskeleton, is a flexible skeleton supported by fluid pressure. Hydrostatic skeletons are common among simple invertebrate organisms. While more advanced organisms can be considered hydrostatic, they are sometimes ref ...
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
Peristalsis ( , ) is a radially symmetrical contraction and relaxation of muscles that propagate in a wave down a tube, in an anterograde direction. Peristalsis is progression of coordinated contraction of involuntary circular muscles, which ...
, which underlies the creeping movement of earthworms.
Obliquely striated muscles
Invertebrates such as annelids, mollusks
Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000 extant species of molluscs are recognized. The number of fossil species is esti ...
, and nematode
The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-Parasitism, parasitic nematodes also known as eelworms. They are a diverse animal phylum inhab ...
s, 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 bivalves
Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, bival ...
, 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 insect
Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s such as wasp
A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. Th ...
s, flies
Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...
, bee
Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyly, monophyletic lineage within the ...
s, and beetle
Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
s possess asynchronous muscles
Asynchronous muscles are muscles in which there is no one-to-one relationship between electrical stimulation and mechanical contraction. These muscles are found in 75% of flying insects and have convergently evolved 7-10 times. Unlike their synchr ...
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 resonant
Resonance describes the phenomenon of increased amplitude that occurs when the frequency of an applied periodic force (or a Fourier component of it) is equal or close to a natural frequency of the system on which it acts. When an oscillatin ...
system, which is driven to a natural frequency of vibration.
History
In 1780, Luigi Galvani
Luigi Galvani (, also ; ; la, Aloysius Galvanus; 9 September 1737 – 4 December 1798) was an Italian physician, physicist, biologist and philosopher, who studied animal electricity. In 1780, he discovered that the muscles of dead frogs' legs ...
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
In developmental biology, bioelectricity refers to the regulation of cell, tissue, and organ-level patterning and behavior as the result of endogenous electrically mediated signaling. Cells and tissues of all types use ion fluxes to communicate e ...
, 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
Calcium is a chemical element with the symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties are most similar to ...
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
Sir Andrew Fielding Huxley (22 November 191730 May 2012) was an English physiology, physiologist and biophysics, biophysicist. He was born into the prominent Huxley family. After leaving Westminster School in central London, he went to Trinit ...
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
Nature, in the broadest sense, is the physics, physical world or universe. "Nature" can refer to the phenomenon, phenomena of the physical world, and also to life in general. The study of nature is a large, if not the only, part of science. ...
'' under the common theme "Structural Changes in Muscle During Contraction".
See also
* Anatomical terms of motion
Motion, the process of movement, is described using specific anatomical terms. Motion includes movement of organs, joints, limbs, and specific sections of the body. The terminology used describes this motion according to its direction relative ...
* calcium-induced calcium release
Calcium-induced calcium release (CICR) describes a biological process whereby calcium is able to activate calcium release from intracellular Ca2+ stores (e.g., endoplasmic reticulum or sarcoplasmic reticulum). Although CICR was first proposed for ...
* Cardiac action potential
The cardiac action potential is a brief change in voltage ( membrane potential) across the cell membrane of heart cells. This is caused by the movement of charged atoms (called ions) between the inside and outside of the cell, through proteins c ...
* Cramp
A cramp is a sudden, involuntary, painful skeletal muscle contraction or overshortening associated with electrical activity; while generally temporary and non-damaging, they can cause significant pain and a paralysis-like immobility of the aff ...
* Dystonia
Dystonia is a neurological hyperkinetic movement disorder in which sustained or repetitive muscle contractions result in twisting and repetitive movements or abnormal fixed postures. The movements may resemble a tremor. Dystonia is often inten ...
* Exercise physiology
Exercise physiology is the physiology of physical exercise. It is one of the allied health professions, and involves the study of the acute responses and chronic adaptations to exercise. Exercise physiologists are the highest qualified exercise ...
* Fasciculation
A fasciculation, or muscle twitch, is a spontaneous, involuntary muscle contraction and relaxation, involving fine muscle fibers. They are common, with as many as 70% of people experiencing them. They can be benign, or associated with more serio ...
* Hill's muscle model In biomechanics, Hill's muscle model refers to either Hill's equations for tetanized muscle contraction or to the 3-element model. They were derived by the famous physiologist Archibald Vivian Hill.
Equation to tetanized muscle
This is a popular ...
* Hypnic jerk
A hypnic jerk, hypnagogic jerk, sleep start, sleep twitch, myoclonic jerk, or night start is a brief and sudden involuntary contraction of the muscles of the body which occurs when a person is beginning to fall asleep, often causing the person t ...
* In vitro muscle testing
''In vitro'' muscle testing is a method used to characterize properties of living muscle tissue after removing it from an organism, which allows more extensive and precise quantification of its properties than ''in vivo'' testing. ''In vitro'' mus ...
* Lombard's paradox
Lombard's paradox describes a paradoxical muscular contraction in humans. When rising to stand from a sitting or squatting position, both the hamstrings and quadriceps contract at the same time, despite them being antagonists to each other.
The r ...
* Myoclonus
Myoclonus is a brief, involuntary, irregular (lacking rhythm) twitching of a muscle or a group of muscles, different from clonus, which is rhythmic or regular. Myoclonus (myo "muscle", clonic "jerk") describes a medical sign and, generally, is ...
* 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 ...
* Spasm
A spasm is a sudden involuntary contraction of a muscle, a group of muscles, or a hollow organ such as the bladder.
A spasmodic muscle contraction may be caused by many medical conditions, including dystonia. Most commonly, it is a muscle c ...
* Uterine contraction
Uterine contractions are muscle contractions of the uterine smooth muscle that occur during the menstrual cycle and labour. Uterine contractions occur throughout the menstrual cycle in the non-pregnant state and throughout gestation.
Throughout m ...
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