Module 25: Muscle Tissue

Lesson 2: Skeletal Muscle

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Mỗi bài học (lesson) bao gồm 4 phần chính: Thuật ngữ, Luyện Đọc, Luyện Nghe, và Bàn Luận.
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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.
The best-known feature of skeletal muscle is its ability to contract and cause movement. Skeletal muscles act not only to produce movement but also to stop movement, such as resisting gravity to maintain posture. Small, constant adjustments of the skeletal muscles are needed to hold a body upright or balanced in any position. Muscles also prevent excess movement of the bones and joints, maintaining skeletal stability and preventing skeletal structure damage or deformation. Joints can become misaligned or dislocated entirely by pulling on the associated bones; muscles work to keep joints stable. Skeletal muscles are located throughout the body at the openings of internal tracts to control the movement of various substances. These muscles allow functions, such as swallowing, urination, and defecation, to be under voluntary control. Skeletal muscles also protect internal organs (particularly abdominal and pelvic organs) by acting as an external barrier or shield to external trauma and by supporting the weight of the organs.

Skeletal muscles contribute to the maintenance of homeostasis in the body by generating heat. Muscle contraction requires energy, and when ATP is broken down, heat is produced. This heat is very noticeable during exercise, when sustained muscle movement causes body temperature to rise, and in cases of extreme cold, when shivering produces random skeletal muscle contractions to generate heat.

Each skeletal muscle is an organ that consists of various integrated tissues. These tissues include the skeletal muscle fibers, blood vessels, nerve fibers, and connective tissue. Each skeletal muscle has three layers of connective tissue (called “mysia”) that enclose it and provide structure to the muscle as a whole, and also compartmentalize the muscle fibers within the muscle (Figure 1). Each muscle is wrapped in a sheath of dense, irregular connective tissue called the epimysium, which allows a muscle to contract and move powerfully while maintaining its structural integrity. The epimysium also separates muscle from other tissues and organs in the area, allowing the muscle to move independently.

Inside each skeletal muscle, muscle fibers are organized into individual bundles, each called a fascicle, by a middle layer of connective tissue called the perimysium. This fascicular organization is common in muscles of the limbs; it allows the nervous system to trigger a specific movement of a muscle by activating a subset of muscle fibers within a bundle, or fascicle of the muscle. Inside each fascicle, each muscle fiber is encased in a thin connective tissue layer of collagen and reticular fibers called the endomysium. The endomysium contains the extracellular fluid and nutrients to support the muscle fiber. These nutrients are supplied via blood to the muscle tissue.

In skeletal muscles that work with tendons to pull on bones, the collagen in the three tissue layers (the mysia) intertwines with the collagen of a tendon. At the other end of the tendon, it fuses with the periosteum coating the bone. The tension created by contraction of the muscle fibers is then transferred through the mysia, to the tendon, and then to the periosteum to pull on the bone for movement of the skeleton. In other places, the mysia may fuse with a broad, tendon-like sheet called an aponeurosis, or to fascia, the connective tissue between skin and bones. The broad sheet of connective tissue in the lower back that the latissimus dorsi muscles (the “lats”) fuse into is an example of an aponeurosis.

Every skeletal muscle is also richly supplied by blood vessels for nourishment, oxygen delivery, and waste removal. In addition, every muscle fiber in a skeletal muscle is supplied by the axon branch of a somatic motor neuron, which signals the fiber to contract. Unlike cardiac and smooth muscle, the only way to functionally contract a skeletal muscle is through signaling from the nervous system.
Because skeletal muscle cells are long and cylindrical, they are commonly referred to as muscle fibers. Skeletal muscle fibers can be quite large for human cells, with diameters up to 100 μm and lengths up to 30 cm (11.8 in) in the Sartorius of the upper leg. During early development, embryonic myoblasts, each with its own nucleus, fuse with up to hundreds of other myoblasts to form the multinucleated skeletal muscle fibers. Multiple nuclei mean multiple copies of genes, permitting the production of the large amounts of proteins and enzymes needed for muscle contraction.

Some other terminology associated with muscle fibers is rooted in the Greek sarco, which means “flesh.” The plasma membrane of muscle fibers is called the sarcolemma, the cytoplasm is referred to as sarcoplasm, and the specialized smooth endoplasmic reticulum, which stores, releases, and retrieves calcium ions (Ca++) is called the sarcoplasmic reticulum (SR) (Figure 2). As will soon be described, the functional unit of a skeletal muscle fiber is the sarcomere, a highly organized arrangement of the contractile myofilaments actin (thin filament) and myosin (thick filament), along with other support proteins.
The striated appearance of skeletal muscle fibers is due to the arrangement of the myofilaments of actin and myosin in sequential order from one end of the muscle fiber to the other. Each packet of these microfilaments and their regulatory proteins, troponin and tropomyosin (along with other proteins) is called a sarcomere.

The sarcomere is the functional unit of the muscle fiber. The sarcomere itself is bundled within the myofibril that runs the entire length of the muscle fiber and attaches to the sarcolemma at its end. As myofibrils contract, the entire muscle cell contracts. Because myofibrils are only approximately 1.2 μm in diameter, hundreds to thousands (each with thousands of sarcomeres) can be found inside one muscle fiber. Each sarcomere is approximately 2 μm in length with a three-dimensional cylinder-like arrangement and is bordered by structures called Z-discs (also called Z-lines, because pictures are two-dimensional), to which the actin myofilaments are anchored (Figure 3). Because the actin and its troponin-tropomyosin complex (projecting from the Z-discs toward the center of the sarcomere) form strands that are thinner than the myosin, it is called the thin filament of the sarcomere. Likewise, because the myosin strands and their multiple heads (projecting from the center of the sarcomere, toward but not all to way to, the Z-discs) have more mass and are thicker, they are called the thick filament of the sarcomere.
Another specialization of the skeletal muscle is the site where a motor neuron’s terminal meets the muscle fiber—called the neuromuscular junction (NMJ). This is where the muscle fiber first responds to signaling by the motor neuron. Every skeletal muscle fiber in every skeletal muscle is innervated by a motor neuron at the NMJ. Excitation signals from the neuron are the only way to functionally activate the fiber to contract.
All living cells have membrane potentials, or electrical gradients across their membranes. The inside of the membrane is usually around -60 to -90 mV, relative to the outside. This is referred to as a cell’s membrane potential. Neurons and muscle cells can use their membrane potentials to generate electrical signals. They do this by controlling the movement of charged particles, called ions, across their membranes to create electrical currents. This is achieved by opening and closing specialized proteins in the membrane called ion channels. Although the currents generated by ions moving through these channel proteins are very small, they form the basis of both neural signaling and muscle contraction.

Both neurons and skeletal muscle cells are electrically excitable, meaning that they are able to generate action potentials. An action potential is a special type of electrical signal that can travel along a cell membrane as a wave. This allows a signal to be transmitted quickly and faithfully over long distances.

Although the term excitation-contraction coupling confuses or scares some students, it comes down to this: for a skeletal muscle fiber to contract, its membrane must first be “excited”—in other words, it must be stimulated to fire an action potential. The muscle fiber action potential, which sweeps along the sarcolemma as a wave, is “coupled” to the actual contraction through the release of calcium ions (Ca++) from the SR. Once released, the Ca++ interacts with the shielding proteins, forcing them to move aside so that the actin-binding sites are available for attachment by myosin heads. The myosin then pulls the actin filaments toward the center, shortening the muscle fiber.

In skeletal muscle, this sequence begins with signals from the somatic motor division of the nervous system. In other words, the “excitation” step in skeletal muscles is always triggered by signaling from the nervous system (Figure 4).

The motor neurons that tell the skeletal muscle fibers to contract originate in the spinal cord, with a smaller number located in the brainstem for activation of skeletal muscles of the face, head, and neck. These neurons have long processes, called axons, which are specialized to transmit action potentials long distances— in this case, all the way from the spinal cord to the muscle itself (which may be up to three feet away). The axons of multiple neurons bundle together to form nerves, like wires bundled together in a cable.

Signaling begins when a neuronal action potential travels along the axon of a motor neuron, and then along the individual branches to terminate at the NMJ. At the NMJ, the axon terminal releases a chemical messenger, or neurotransmitter, called acetylcholine (ACh). The ACh molecules diffuse across a minute space called the synaptic cleft and bind to ACh receptors located within the motor end-plate of the sarcolemma on the other side of the synapse. Once ACh binds, a channel in the ACh receptor opens and positively charged ions can pass through into the muscle fiber, causing it to depolarize, meaning that the membrane potential of the muscle fiber becomes less negative (closer to zero.) As the membrane depolarizes, another set of ion channels called voltage-gated sodium channels are triggered to open. Sodium ions enter the muscle fiber, and an action potential rapidly spreads (or “fires”) along the entire membrane to initiate excitation-contraction coupling.

Things happen very quickly in the world of excitable membranes (just think about how quickly you can snap your fingers as soon as you decide to do it). Immediately following depolarization of the membrane, it repolarizes, re-establishing the negative membrane potential. Meanwhile, the ACh in the synaptic cleft is degraded by the enzyme acetylcholinesterase (AChE) so that the ACh cannot rebind to a receptor and reopen its channel, which would cause unwanted extended muscle excitation and contraction.

Propagation of an action potential along the sarcolemma is the excitation portion of excitation-contraction coupling. Recall that this excitation actually triggers the release of calcium ions (Ca++) from its storage in the cell’s SR. For the action potential to reach the membrane of the SR, there are periodic invaginations in the sarcolemma, called T-tubules (“T” stands for “transverse”). You will recall that the diameter of a muscle fiber can be up to 100 μm, so these T-tubules ensure that the membrane can get close to the SR in the sarcoplasm. The arrangement of a T-tubule with the membranes of SR on either side is called a triad (Figure 5). The triad surrounds the cylindrical structure called a myofibril, which contains actin and myosin.

The T-tubules carry the action potential into the interior of the cell, which triggers the opening of calcium channels in the membrane of the adjacent SR, causing Ca++ to diffuse out of the SR and into the sarcoplasm. It is the arrival of Ca++ in the sarcoplasm that initiates contraction of the muscle fiber by its contractile units, or sarcomeres.

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.

Bundles of muscle fibers, called fascicles, are covered by the perimysium. Muscle fibers are covered by the endomysium.

A skeletal muscle fiber is surrounded by a plasma membrane called the sarcolemma, which contains sarcoplasm, the cytoplasm of muscle cells. A muscle fiber is composed of many fibrils, which give the cell its striated appearance.

The sarcomere, the region from one Z-line to the next Z-line, is the functional unit of a skeletal muscle fiber.

At the NMJ, the axon terminal releases ACh. The motor end-plate is the location of the ACh-receptors in the muscle fiber sarcolemma. When ACh molecules are released, they diffuse across a minute space called the synaptic cleft and bind to the receptors.

Narrow T-tubules permit the conduction of electrical impulses. The SR functions to regulate intracellular levels of calcium. Two terminal cisternae (where enlarged SR connects to the T-tubule) and one T-tubule comprise a triad—a “threesome” of membranes, with those of SR on two sides and the T-tubule sandwiched between them.

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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. Skeletal muscles contain connective tissue, blood vessels, and nerves.
  2. There are three layers of connective tissue: epimysium, perimysium, and endomysium.
  3. Skeletal muscle fibers are organized into groups called fascicles.
  4. Blood vessels and nerves enter the connective tissue and branch in the cell.
  5. Muscles attach to bones directly or through tendons or aponeuroses.
  6. Skeletal muscles maintain posture, stabilize bones and joints, control internal movement, and generate heat.
  7. Skeletal muscle fibers are long, multinucleated cells.
  8. The membrane of the cell is the sarcolemma; the cytoplasm of the cell is the sarcoplasm.
  9. The sarcoplasmic reticulum is a form of endoplasmic reticulum.
  10. Muscle fibers are composed of myofibrils.
  11. The striations are created by the organization of actin and myosin resulting in the banding pattern of myofibrils.
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