Module 25: Muscle Tissue

Lesson 4: Nervous System Control of Muscle Tension

Hệ Thần Kinh Kiểm Soát Trương Lực Cơ

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Dưới đây là danh sách những thuật ngữ Y khoa của module Muscle Tissue.
Khái quát được số lượng thuật ngữ sẽ xuất hiện trong bài đọc và nghe sẽ giúp bạn thoải mái tiêu thụ nội dung hơn. Sau khi hoàn thành nội dung đọc và nghe, bạn hãy quay lại đây và luyện tập (practice) để quen dần các thuật ngữ này. Đừng ép bản thân phải nhớ các thuật ngữ này vội vì bạn sẽ gặp và ôn lại danh sách này trong những bài học (lesson) khác của cùng một module.

Medical Terminology: Muscle Tissue

acetylcholine (ACh)
neurotransmitter that binds at a motor end-plate to trigger depolarization
actin
protein that makes up most of the thin myofilaments in a sarcomere muscle fiber
action potential
change in voltage of a cell membrane in response to a stimulus that results in transmission of an electrical signal; unique to neurons and muscle fibers
aerobic respiration
production of ATP in the presence of oxygen
angiogenesis
formation of blood capillary networks
aponeurosis
broad, tendon-like sheet of connective tissue that attaches a skeletal muscle to another skeletal muscle or to a bone
ATPase
enzyme that hydrolyzes ATP to ADP
atrophy
loss of structural proteins from muscle fibers
autorhythmicity
heart’s ability to control its own contractions
calmodulin
regulatory protein that facilitates contraction in smooth muscles
cardiac muscle
striated muscle found in the heart; joined to one another at intercalated discs and under the regulation of pacemaker cells, which contract as one unit to pump blood through the circulatory system. Cardiac muscle is under involuntary control.
concentric contraction
muscle contraction that shortens the muscle to move a load
contractility
ability to shorten (contract) forcibly
contraction phase
twitch contraction phase when tension increases
creatine phosphate
phosphagen used to store energy from ATP and transfer it to muscle
dense body
sarcoplasmic structure that attaches to the sarcolemma and shortens the muscle as thin filaments slide past thick filaments
depolarize
to reduce the voltage difference between the inside and outside of a cell’s plasma membrane (the sarcolemma for a muscle fiber), making the inside less negative than at rest
desmosome
cell structure that anchors the ends of cardiac muscle fibers to allow contraction to occur
eccentric contraction
muscle contraction that lengthens the muscle as the tension is diminished
elasticity
ability to stretch and rebound
endomysium
loose, and well-hydrated connective tissue covering each muscle fiber in a skeletal muscle
epimysium
outer layer of connective tissue around a skeletal muscle
excitability
ability to undergo neural stimulation
excitation-contraction coupling
sequence of events from motor neuron signaling to a skeletal muscle fiber to contraction of the fiber’s sarcomeres
extensibility
ability to lengthen (extend)
fascicle
bundle of muscle fibers within a skeletal muscle
fast glycolytic (FG)
muscle fiber that primarily uses anaerobic glycolysis
fast oxidative (FO)
intermediate muscle fiber that is between slow oxidative and fast glycolytic fibers
fibrosis
replacement of muscle fibers by scar tissue
glycolysis
anaerobic breakdown of glucose to ATP
graded muscle response
modification of contraction strength
hyperplasia
process in which one cell splits to produce new cells
hypertonia
abnormally high muscle tone
hypertrophy
addition of structural proteins to muscle fibers
hypotonia
abnormally low muscle tone caused by the absence of low-level contractions
intercalated disc
part of the sarcolemma that connects cardiac tissue, and contains gap junctions and desmosomes
isometric contraction
muscle contraction that occurs with no change in muscle length
isotonic contraction
muscle contraction that involves changes in muscle length
lactic acid
product of anaerobic glycolysis
latch-bridges
subset of a cross-bridge in which actin and myosin remain locked together
latent period
the time when a twitch does not produce contraction
motor end-plate
sarcolemma of muscle fiber at the neuromuscular junction, with receptors for the neurotransmitter acetylcholine
motor unit
motor neuron and the group of muscle fibers it innervates
muscle tension
force generated by the contraction of the muscle; tension generated during isotonic contractions and isometric contractions
muscle tone
low levels of muscle contraction that occur when a muscle is not producing movement
myoblast
muscle-forming stem cell
myofibril
long, cylindrical organelle that runs parallel within the muscle fiber and contains the sarcomeres
myogram
instrument used to measure twitch tension
myosin
protein that makes up most of the thick cylindrical myofilament within a sarcomere muscle fiber
myotube
fusion of many myoblast cells
neuromuscular junction (NMJ)
synapse between the axon terminal of a motor neuron and the section of the membrane of a muscle fiber with receptors for the acetylcholine released by the terminal
neurotransmitter
signaling chemical released by nerve terminals that bind to and activate receptors on target cells
oxygen debt
amount of oxygen needed to compensate for ATP produced without oxygen during muscle contraction
pacesetter cell
cell that triggers action potentials in smooth muscle
pericyte
stem cell that regenerates smooth muscle cells
perimysium
connective tissue that bundles skeletal muscle fibers into fascicles within a skeletal muscle
power stroke
action of myosin pulling actin inward (toward the M line)
pyruvic acid
product of glycolysis that can be used in aerobic respiration or converted to lactic acid
recruitment
increase in the number of motor units involved in contraction
relaxation phase
period after twitch contraction when tension decreases
sarcolemma
plasma membrane of a skeletal muscle fiber
sarcomere
longitudinally, repeating functional unit of skeletal muscle, with all of the contractile and associated proteins involved in contraction
sarcopenia
age-related muscle atrophy
sarcoplasm
cytoplasm of a muscle cell
sarcoplasmic reticulum (SR)
specialized smooth endoplasmic reticulum, which stores, releases, and retrieves Ca++
satellite cell
stem cell that helps to repair muscle cells
skeletal muscle
striated, multinucleated muscle that requires signaling from the nervous system to trigger contraction; most skeletal muscles are referred to as voluntary muscles that move bones and produce movement
slow oxidative (SO)
muscle fiber that primarily uses aerobic respiration
smooth muscle
nonstriated, mononucleated muscle in the skin that is associated with hair follicles; assists in moving materials in the walls of internal organs, blood vessels, and internal passageways
somites
blocks of paraxial mesoderm cells
stress-relaxation response
relaxation of smooth muscle tissue after being stretched
synaptic cleft
space between a nerve (axon) terminal and a motor end-plate
T-tubule
projection of the sarcolemma into the interior of the cell
tetanus
a continuous fused contraction
thick filament
the thick myosin strands and their multiple heads projecting from the center of the sarcomere toward, but not all to way to, the Z-discs
thin filament
thin strands of actin and its troponin-tropomyosin complex projecting from the Z-discs toward the center of the sarcomere
treppe
stepwise increase in contraction tension
triad
the grouping of one T-tubule and two terminal cisternae
tropomyosin
regulatory protein that covers myosin-binding sites to prevent actin from binding to myosin
troponin
regulatory protein that binds to actin, tropomyosin, and calcium
twitch
single contraction produced by one action potential
varicosity
enlargement of neurons that release neurotransmitters into synaptic clefts
visceral muscle
smooth muscle found in the walls of visceral organs
voltage-gated sodium channels
membrane proteins that open sodium channels in response to a sufficient voltage change, and initiate and transmit the action potential as Na+ enters through the channel
wave summation
addition of successive neural stimuli to produce greater contraction
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Dưới đây là các bài văn nằm ở bên trái. Ở bên phải là các bài luyện tập (practice) để đánh giá khả năng đọc hiểu của bạn. Sẽ khó khăn trong thời gian đầu nếu vốn từ vựng của bạn còn hạn chế, đặc biệt là từ vựng Y khoa. Hãy kiên nhẫn và đọc nhiều nhất có kể, lượng kiến thức tích tụ dần sẽ giúp bạn đọc thoải mái hơn.
To move an object, referred to as load, the sarcomeres in the muscle fibers of the skeletal muscle must shorten. The force generated by the contraction of the muscle (or shortening of the sarcomeres) is called muscle tension. However, muscle tension also is generated when the muscle is contracting against a load that does not move, resulting in two main types of skeletal muscle contractions: isotonic contractions and isometric contractions.

In isotonic contractions, where the tension in the muscle stays constant, a load is moved as the length of the muscle changes (shortens). There are two types of isotonic contractions: concentric and eccentric. A concentric contraction involves the muscle shortening to move a load. An example of this is the biceps brachii muscle contracting when a hand weight is brought upward with increasing muscle tension. As the biceps brachii contract, the angle of the elbow joint decreases as the forearm is brought toward the body. Here, the biceps brachii contracts as sarcomeres in its muscle fibers are shortening and cross-bridges form; the myosin heads pull the actin. An eccentric contraction occurs as the muscle tension diminishes and the muscle lengthens. In this case, the hand weight is lowered in a slow and controlled manner as the amount of cross-bridges being activated by nervous system stimulation decreases. In this case, as tension is released from the biceps brachii, the angle of the elbow joint increases. Eccentric contractions are also used for movement and balance of the body.

An isometric contraction occurs as the muscle produces tension without changing the angle of a skeletal joint. Isometric contractions involve sarcomere shortening and increasing muscle tension, but do not move a load, as the force produced cannot overcome the resistance provided by the load. For example, if one attempts to lift a hand weight that is too heavy, there will be sarcomere activation and shortening to a point, and ever-increasing muscle tension, but no change in the angle of the elbow joint. In everyday living, isometric contractions are active in maintaining posture and maintaining bone and joint stability. However, holding your head in an upright position occurs not because the muscles cannot move the head, but because the goal is to remain stationary and not produce movement. Most actions of the body are the result of a combination of isotonic and isometric contractions working together to produce a wide range of outcomes (Figure 1).

All of these muscle activities are under the exquisite control of the nervous system. Neural control regulates concentric, eccentric and isometric contractions, muscle fiber recruitment, and muscle tone. A crucial aspect of nervous system control of skeletal muscles is the role of motor units.
As you have learned, every skeletal muscle fiber must be innervated by the axon terminal of a motor neuron in order to contract. Each muscle fiber is innervated by only one motor neuron. The actual group of muscle fibers in a muscle innervated by a single motor neuron is called a motor unit. The size of a motor unit is variable depending on the nature of the muscle.

A small motor unit is an arrangement where a single motor neuron supplies a small number of muscle fibers in a muscle. Small motor units permit very fine motor control of the muscle. The best example in humans is the small motor units of the extraocular eye muscles that move the eyeballs. There are thousands of muscle fibers in each muscle, but every six or so fibers are supplied by a single motor neuron, as the axons branch to form synaptic connections at their individual NMJs. This allows for exquisite control of eye movements so that both eyes can quickly focus on the same object. Small motor units are also involved in the many fine movements of the fingers and thumb of the hand for grasping, texting, etc.

A large motor unit is an arrangement where a single motor neuron supplies a large number of muscle fibers in a muscle. Large motor units are concerned with simple, or “gross,” movements, such as powerfully extending the knee joint. The best example is the large motor units of the thigh muscles or back muscles, where a single motor neuron will supply thousands of muscle fibers in a muscle, as its axon splits into thousands of branches.

There is a wide range of motor units within many skeletal muscles, which gives the nervous system a wide range of control over the muscle. The small motor units in the muscle will have smaller, lower-threshold motor neurons that are more excitable, firing first to their skeletal muscle fibers, which also tend to be the smallest. Activation of these smaller motor units, results in a relatively small degree of contractile strength (tension) generated in the muscle. As more strength is needed, larger motor units, with bigger, higher-threshold motor neurons are enlisted to activate larger muscle fibers. This increasing activation of motor units produces an increase in muscle contraction known as recruitment. As more motor units are recruited, the muscle contraction grows progressively stronger. In some muscles, the largest motor units may generate a contractile force of 50 times more than the smallest motor units in the muscle. This allows a feather to be picked up using the biceps brachii arm muscle with minimal force, and a heavy weight to be lifted by the same muscle by recruiting the largest motor units.

When necessary, the maximal number of motor units in a muscle can be recruited simultaneously, producing the maximum force of contraction for that muscle, but this cannot last for very long because of the energy requirements to sustain the contraction. To prevent complete muscle fatigue, motor units are generally not all simultaneously active, but instead some motor units rest while others are active, which allows for longer muscle contractions. The nervous system uses recruitment as a mechanism to efficiently utilize a skeletal muscle.
When a skeletal muscle fiber contracts, myosin heads attach to actin to form cross-bridges followed by the thin filaments sliding over the thick filaments as the heads pull the actin, and this results in sarcomere shortening, creating the tension of the muscle contraction. The cross-bridges can only form where thin and thick filaments already overlap, so that the length of the sarcomere has a direct influence on the force generated when the sarcomere shortens. This is called the length-tension relationship.

The ideal length of a sarcomere to produce maximal tension occurs at 80 percent to 120 percent of its resting length, with 100 percent being the state where the medial edges of the thin filaments are just at the most-medial myosin heads of the thick filaments (Figure 2). This length maximizes the overlap of actin-binding sites and myosin heads. If a sarcomere is stretched past this ideal length (beyond 120 percent), thick and thin filaments do not overlap sufficiently, which results in less tension produced. If a sarcomere is shortened beyond 80 percent, the zone of overlap is reduced with the thin filaments jutting beyond the last of the myosin heads and shrinks the H zone, which is normally composed of myosin tails. Eventually, there is nowhere else for the thin filaments to go and the amount of tension is diminished. If the muscle is stretched to the point where thick and thin filaments do not overlap at all, no cross-bridges can be formed, and no tension is produced in that sarcomere. This amount of stretching does not usually occur, as accessory proteins and connective tissue oppose extreme stretching.
A single action potential from a motor neuron will produce a single contraction in the muscle fibers of its motor unit. This isolated contraction is called a twitch. A twitch can last for a few milliseconds or 100 milliseconds, depending on the muscle type. The tension produced by a single twitch can be measured by a myogram, an instrument that measures the amount of tension produced over time (Figure 3). Each twitch undergoes three phases. The first phase is the latent period, during which the action potential is being propagated along the sarcolemma and Ca++ ions are released from the SR. This is the phase during which excitation and contraction are being coupled but contraction has yet to occur. The contraction phase occurs next. The Ca++ ions in the sarcoplasm have bound to troponin, tropomyosin has shifted away from myosin-binding sites on actin, cross-bridges have formed, and sarcomeres are actively shortening to the point of peak tension. The last phase is the relaxation phase, when tension decreases as contraction stops. Ca++ ions are pumped out of the sarcoplasm into the SR, and cross-bridge cycling stops, returning the muscle fibers to their resting state.

Although a person can experience a muscle “twitch,” a single twitch does not produce any significant muscle activity in a living body. A series of action potentials to the muscle fibers is necessary to produce a muscle contraction that can produce work. A normal muscle contraction is more sustained, and it can be modified by input from the nervous system to produce varying amounts of force; this is called a graded muscle response. The frequency of action potentials (nerve impulses) from a motor neuron and the number of motor neurons transmitting action potentials both affect the tension produced in skeletal muscle.

The rate at which a motor neuron fires action potentials affects the tension produced in the skeletal muscle. If the fibers are stimulated while a previous twitch is still occurring, the second twitch will be stronger. This response is called wave summation, because the excitation-contraction coupling effects of successive motor neuron signaling are summed, or added together (Figure 4a). At the molecular level, summation occurs because the second stimulus triggers the release of more Ca++ ions, which become available to activate additional sarcomeres while the muscle is still contracting from the first stimulus. Summation results in greater contraction of the motor unit.

If the frequency of motor neuron signaling increases, summation and subsequent muscle tension in the motor unit continues to rise until it reaches a peak point. The tension at this point is about three to four times greater than the tension of a single twitch, a state referred to as incomplete tetanus. During incomplete tetanus, the muscle goes through quick cycles of contraction with a short relaxation phase for each. If the stimulus frequency is so high that the relaxation phase disappears completely, contractions become continuous in a process called complete tetanus (Figure 4b).

During tetanus, the concentration of Ca++ ions in the sarcoplasm allows virtually all of the sarcomeres to form cross-bridges and shorten, so that a contraction can continue uninterrupted (until the muscle fatigues and can no longer produce tension).
When a skeletal muscle has been dormant for an extended period and then activated to contract, with all other things being equal, the initial contractions generate about one-half the force of later contractions. The muscle tension increases in a graded manner that to some looks like a set of stairs. This tension increase is called treppe, a condition where muscle contractions become more efficient. It’s also known as the “staircase effect” (Figure 5).

It is believed that treppe results from a higher concentration of Ca++ in the sarcoplasm resulting from the steady stream of signals from the motor neuron. It can only be maintained with adequate ATP.
Skeletal muscles are rarely completely relaxed, or flaccid. Even if a muscle is not producing movement, it is contracted a small amount to maintain its contractile proteins and produce muscle tone. The tension produced by muscle tone allows muscles to continually stabilize joints and maintain posture.

Muscle tone is accomplished by a complex interaction between the nervous system and skeletal muscles that results in the activation of a few motor units at a time, most likely in a cyclical manner. In this manner, muscles never fatigue completely, as some motor units can recover while others are active.

The absence of the low-level contractions that lead to muscle tone is referred to as hypotonia, and can result from damage to parts of the central nervous system (CNS), such as the cerebellum, or from loss of innervations to a skeletal muscle, as in poliomyelitis. Hypotonic muscles have a flaccid appearance and display functional impairments, such as weak reflexes. Conversely, excessive muscle tone is referred to as hypertonia, accompanied by hyperreflexia (excessive reflex responses), often the result of damage to upper motor neurons in the CNS. Hypertonia can present with muscle rigidity (as seen in Parkinson’s disease) or spasticity, a phasic change in muscle tone, where a limb will “snap” back from passive stretching (as seen in some strokes).

OpenStax. (2022). Anatomy and Physiology 2e. Rice University. Retrieved June 15, 2023. ISBN-13: 978-1-711494-06-7 (Hardcover) ISBN-13: 978-1-711494-05-0 (Paperback) ISBN-13: 978-1-951693-42-8 (Digital). License: Attribution 4.0 International (CC BY 4.0). Access for free at openstax.org.

During isotonic contractions, muscle length changes to move a load. During isometric contractions, muscle length does not change because the load exceeds the tension the muscle can generate.

Sarcomeres produce maximal tension when thick and thin filaments overlap between about 80 percent to 120 percent.

A single muscle twitch has a latent period, a contraction phase when tension increases, and a relaxation phase when tension decreases. During the latent period, the action potential is being propagated along the sarcolemma. During the contraction phase, Ca++ ions in the sarcoplasm bind to troponin, tropomyosin moves from actin-binding sites, cross-bridges between actin and myosin form, and sarcomeres shorten. During the relaxation phase, tension decreases as Ca++ ions are pumped out of the sarcoplasm and cross-bridge cycling stops.

(a) The excitation-contraction coupling effects of successive motor neuron signaling is added together which is referred to as wave summation. The bottom of each wave, the end of the relaxation phase, represents the point of stimulus. (b) When the stimulus frequency is so high that the relaxation phase disappears completely, the contractions become continuous; this is called tetanus.

When muscle tension increases in a graded manner that looks like a set of stairs, it is called treppe. The bottom of each wave represents the point of stimulus.

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Dưới đây là video và các luyện tập (practice) của bài này. Nghe là một kĩ năng khó, đặc biệt là khi chúng ta chưa quen nội dung và chưa có nhạy cảm ngôn ngữ. Nhưng cứ đi thật chậm và đừng bỏ cuộc.
Xem video và cảm nhận nội dung bài. Bạn có thể thả trôi, cảm nhận dòng chảy ngôn ngữ và không nhất thiết phải hiểu toàn bộ bài. Bên dưới là script để bạn khái quát nội dụng và tra từ mới.
Script:
  1. The number of cross-bridges formed between actin and myosin determines the amount of tension produced by a muscle.
  2. The length of a sarcomere is optimal when the zone of overlap between thin and thick filaments is greatest.
  3. Muscles that are stretched or compressed too greatly do not produce maximal amounts of power.
  4. A motor unit is formed by a motor neuron and all of the muscle fibers that are innervated by that same motor neuron.
  5. A single contraction is called a twitch.
  6. A muscle twitch has a latent period, a contraction phase, and a relaxation phase.
  7. A graded muscle response allows variation in muscle tension.
  8. Summation occurs as successive stimuli are added together to produce a stronger muscle contraction.
  9. Tetanus is the fusion of contractions to produce a continuous contraction.
  10. Increasing the number of motor neurons involved increases the amount of motor units activated in a muscle, which is called recruitment.
  11. Muscle tone is the constant low-level contractions that allow for posture and stability.
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