Skeletal Muscle Anatomy


The surface electromyograph (SEMG) measures the electrical changes in the muscle fiber membrane preceding contraction, not muscle contraction itself. SEMG biofeedback is arguably the most widely used biofeedback modality due to its diverse applications ranging from neuromuscular to optimal performance. SEMG biofeedback plays an increasingly important role in optimal performance training for Olympic athletes. Graphic © studioloco/Shutterstock.com.

BCIA Blueprint Coverage


This unit discusses Descriptions of most commonly employed biofeedback modalities: SEMG (III-A 1-2) and Muscle anatomy and physiology; antagonistic and synergistic muscle groups (IV-A).

 
This unit covers Three Types of Muscles, the Skeletal Muscle System, Motor Units, Muscle Action Potentials, Types of Skeletal Muscle Fibers, the EMG Signal, Skeletal Muscle Contraction, the Stretch Reflex, the Tendon Reflex, Muscle Action, Skeletal Muscles of Clinical Interest (Face), and Skeletal Muscles of Clinical Interest (Front and Back).

Please click on the podcast icon below to hear a full-length lecture.



Three Types of Muscles


The human body contains three types of muscles: skeletal, smooth, and cardiac.



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Graphic © Designua/Shutterstock.com.





Skeletal muscles consist of cylindrical fibers that vary from a few centimeters to 30-40 cm (Tortora & Derrickson, 2021). Their major functions are motion, posture, producing heat, and protection. They are multinucleated with nuclei located at the periphery. They are striated (striped) due to alternating light and dark bands visible using a light microscope. Tendons usually attach skeletal muscles to bones. They are called skeletal muscles because most of them move the skeleton's bones. We control skeletal muscles voluntarily and involuntarily. Graphic © 2006 Jose Luis Calvo/Shutterstock.com.




            

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We can voluntarily control skeletal muscles through the somatic division of the peripheral nervous system because their contraction produces proprioceptive feedback. Contract a skeletal muscle like the biceps brachii and the elbow flexes. This limb movement provides the sensory information we need to adjust our arm's position consciously. Graphic © Kjpargeter/Shutterstock.com.



Most muscles are also controlled involuntarily. We are often unaware of the rhythmic movement of the diaphragm and the intercostal muscles that allow us to breathe. The monosynaptic stretch reflex, which maintains postural muscle tone and stabilizes limb position, also operates unconsciously.

Smooth muscles comprise nonstriated fibers. They are called smooth because they do not show banding under a light microscope. They are small spindle-shaped cells with a central nucleus. In structures like the intestinal wall, gap junctions connect separate fibers to produce strong, usually involuntary, contractions. In structures without gap junctions like the iris, smooth muscle fibers contract separately like skeletal muscles. Smooth muscle functions include airway and blood vessel constriction, nutrient movement through the digestive tract, and gallbladder and urinary bladder contraction (Tortora & Derrickson, 2021). Graphic © Choksawatdikorn/Shutterstock.com.




There are two kinds of smooth muscle: single-unit and multiunit smooth muscle (Tortora & Derrickson, 2021).

Single-unit smooth muscle comprises part of the walls of small arteries and veins and the stomach, intestines, uterus, and urinary bladder.

Multiunit smooth muscle is found in the walls of large arteries, airways to the lungs, the arrector pili muscles that move hair follicles, the muscle rings of the iris, and the ciliary body that focuses the lens.

Compared with skeletal muscles, smooth muscle contraction starts more gradually and persists longer. Smooth muscles can shorten and stretch more than other kinds of muscles. Smooth muscles usually operate involuntarily, and some tissues have an intrinsic rhythm (autorhythmicity). They are jointly controlled by the autonomic branch of the peripheral nervous system and the endocrine system.

Before biofeedback training, most patients could not voluntarily control smooth muscles since their contraction does not provide sufficient feedback. For example, blood vessel dilation or constriction in the fingers produces such "faint" feedback that we are ordinarily unaware of these changes. Biofeedback converts smooth muscle activity into a digital display, colored bar, or tone that a patient can use to voluntarily warm or cool the fingers.

Below is a BioGraph ® Infiniti display of peripheral blood flow that simultaneously shows blood volume pulse and temperature detected from the hand.





Temperature biofeedback illustrates a crucial concept. We can teach patients to control "involuntary" processes by supplementing inadequate natural feedback.

Cardiac muscle comprises most of the heart wall. Cardiac muscle fibers are branched and striated with one central nucleus (sometimes two). The ends of cardiac muscle fibers attach to intercalated discs comprising desmosomes and gap junctions. Desmosomes contribute structural integrity during forceful contractions. Gap junctions rapidly conduct muscle action potentials to ensure coordinated vigorous contractions. Cardiac muscle has crossed striations that allow the heart to eject blood from the four chambers of the heart (Tortora & Derrickson, 2021). Graphic © sciencepics/Shutterstock.com.




Cardiac muscle fibers contain the same actin and myosin filaments, bands, zones, and Z discs as skeletal muscles. Graphic © Jose Luis Calvo/Shutterstock.com.



          
Cardiac muscle contraction lasts 10-15 times longer than skeletal muscle contraction due to the gradual movement of calcium ions into the sarcoplasm (muscle fiber cytoplasm).

Skeletal muscle contraction only occurs when a motor neuron releases acetylcholine. In contrast, the sinoatrial and atrioventricular node pacemaker cells generate the heart rhythm. The autonomic branch of the peripheral nervous system and the endocrine system jointly adjust heart rate by acting on these pacemakers.

As with smooth muscle, most patients cannot voluntarily control the heart rhythm before biofeedback training. Biofeedback supplements cardiac muscle feedback to teach patients to slow their heart rate or reduce the frequency of abnormal rhythms like premature ventricular contractions (PVCs) when they are distressed (Tortora & Derrickson, 2021).


Skeletal Muscles Are Endocrine Glands


Human skeletal muscles are increasingly recognized as endocrine organs that secrete a variety of bioactive molecules known as exerkines during physical activity. These exerkines play significant roles in inter-organ communication and systemic physiological regulation. For instance, Growth and Differentiation Factor 15 (GDF15) is identified as a novel exerkine secreted by skeletal muscle during exercise, which promotes lipolysis in human adipose tissue, highlighting its role in energy metabolism (Laurens et al., 2020).

Additionally, skeletal muscles secrete other exerkines such as thymosin beta-4 (TMSB4X), which is upregulated during muscle contraction and has been implicated in cellular crosstalk, although its effects on metabolic disorders remain inconclusive (Gonzalez-Franquesa et al., 2021).

Furthermore, skeletal muscle-derived exerkines like apelin, kynurenic acid, and lactate have been shown to influence the progression of neurodegenerative diseases by mediating the muscle-brain axis, thus underscoring their therapeutic potential (Bian et al., 2024). This evidence collectively supports the concept of skeletal muscles functioning as glands that secrete exerkines, contributing to the regulation of various physiological processes and disease states.

Skeletal Muscle System


The skeletal muscle system consists of extrafusal muscle fibers and connective tissue. Important units of distance, signal frequency and strength, and time are summarized below.



Skeletal muscle fibers are striated (striped) due to alternating light and dark bands. These fibers run from 10-100 micrometers in diameter and 10 to 30 centimeters in length. The number of skeletal muscle fibers is determined before birth, and most survive across our lifespan (Tortora & Derrickson, 2021),

The sarcolemma, a plasma membrane, encloses individual fibers. Muscular tissue shares four properties: electrical excitability (producing muscle action potentials), contractility (contracting forcefully), extensibility (stretching), and elasticity (returning to its initial length and shape; Tortora & Derrickson, 2021).



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Filaments Comprising the Sarcomere

Each skeletal muscle fiber consists of cylindrical myofibrils (hundreds to thousands) that are ~ 2 μ in diameter and run the muscle fiber's length. Myofibrils enable muscle fibers to contract due to their overlapping thin and thick filaments, whose striations give muscle fibers a striped appearance (Tortora & Derrickson, 2021). Graphic © Alila Medical Media/Shutterstock.com.



Lateral thin filaments are 8 nm in diameter, 1-2 μ in length, and comprise the protein actin. They span the I band and transit part of the A band. Central thick filaments are 16 nm in diameter, 1-2 μ in length, and comprise the protein myosin. They connect at a sarcomere's M-line (M for middle). There are two thin for every thick filament where thin and thick filaments overlap. The filaments that comprise a myofibril do not extend a muscle fiber's length. Instead, they are stacked in compartments called sarcomeres separated by plate-shaped zones of dense protein called Z discs. Z-discs anchor thin filaments. Sarcomeres extend from Z disc to Z disc and are a muscle fiber's smallest contractile unit. Sarcomeres comprising a myofibril are aligned like boxcars (Marieb & Hoehn, 2019; Tortora & Derrickson, 2021).


Muscle Fiber


Principles of Anatomy and Physiology
, 12th edition by Gerard J. Tortora and Bryan H. Derrickson © 2009 John Wiley & Sons, Inc. Reproduced with permission of John Wiley & Sons, Inc.


Sarcomere Organization

A sarcomere consists of A, I, and H bands, and the M line.

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An A band runs the length of thick filaments and appears as a sarcomere's darker middle region. Thin and thick filaments are almost perfectly aligned and overlap at the ends of each A band. The lighter, less dense I band contains the remaining thin and no thick filaments. Z disks intersect its center. The interleaving of dark A and light I bands produces the banding visible in myofibrils and skeletal muscle fibers (Marieb & Hoehn, 2019; Tortora & Derrickson, 2021).




At the center of each A band is a thin H band comprising only thick filaments. The proteins that connect these filaments at the middle of the H band create the M line. Graphic © Designua/ Shutterstock.com.


Myofibril and Sarcomere Elasticity

Titin, a structural protein called connectin, links a Z disc (also called Z line) to the M line in the center of a sarcomere to stabilize the thick filament's position. The segment of the titin molecule attached to the Z disc can stretch more than four times its resting length.

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This ability to stretch may explain a myofibril's elasticity and a sarcomere's ability to return to its resting length after shortening or lengthening. It also clarifies how an A band can maintain its central position within a sarcomere (Tortora & Derrickson, 2021). Graphic © Akor86/Shutterstock.com.



α-Actinin and Myomesin Connections

The structural protein α-actinin is a component of Z discs that connects actin molecules of thin filaments to titin. Myomesin molecules form the M line at the center of a sarcomere. The M lines' proteins also attach to titin and link adjacent thick filaments. Graphic © Blamb/Shutterstock.com.



How Skeletal Muscles Generate Force

Muscle fibers generate force by pulling Z discs together, shortening the sarcomeres. This force, called active tension, moves bones at joints and resists active stretching due to external forces like gravity. Check out the YouTube video Muscle Contraction Process: Molecular Mechanism [3D Animation].

The Role of Connective Tissue

Connective tissue packages muscle fibers together and connects muscle fibers to bone (as well as skin, muscle, and deep fascia).

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Fascia, fibrous connective tissue, divides muscles into functional groups, aids muscle movement, transmits force to bones, encloses and supports blood vessels and nerve fibers, and secures organs in their place (Tortora & Derrickson, 2021).

Muscle fiber bundles are called fascicles. The muscle fibers within a fascicle are parallel to each other.
Graphic © sciencepics/Shutterstock.com.



Fascicle-tendon connections can assume five shapes: circular, fusiform (narrow at the ends and wide in the center), parallel, pennate (feather-shaped, unipennate, bipennate, and multipennate), or triangular (convergent; Tortora & Derrickson, 2021). Graphic courtesy of Wikimedia.



Tendons are dense fasciae that connect muscles to other structures. Connective tissue provides the skeletal muscle system's elasticity. Connective tissue elasticity produces passive tension, allowing muscle fibers to generate, sum, and transmit force (Tortora & Derrickson, 2021).

Motor Units

Skeletal muscle fibers are organized into motor units consisting of an alpha motor neuron and the extrafusal muscle fibers it controls.

In the diagram below, a motor neuron's axon branches to simultaneously innervate four extrafusal muscle fibers. This motor unit is designed for precise control instead of generating force.



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An alpha motor neuron's cell body lies in the ventral horns of the spinal cord. The axon, depicted in the photomicrograph below as a dark filament, may extend three feet until it synapses with muscle fibers. Separate motor neurons may innervate muscles that perform several actions. Graphic © Christopher Meade/Shutterstock.com.




                   

Alpha motor neurons, which conduct messages from the central nervous system to skeletal muscles, are called efferent nerves because they leave the central nervous system.

A motor unit contains from 3 to 3000 muscle fibers that contract completely or not at all. The average motor unit contains 150 fibers (Buchthal & Schmalbruch, 1980; Tortora & Derrickson, 2021). All motor unit muscle fibers share the same composition.

Smaller motor units perform precise movements. For example, the larynx, which controls speech, contains motor units with 1-2 muscle fibers. The extraocular muscles that move the eyes contain motor units with 10-20 muscle fibers. Larger motor units are designed to deliver power instead of precise control. The gastrocnemius muscle located in the calf contains whose motor units with 2000-3000 muscle fibers (Marieb & Hoehn, 2019).

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SEMG biofeedback uses an electromyograph to monitor the total electrical output in microvolts (millionths of a volt) from all the motor units firing near surface electrodes. Contraction strength and SEMG signal voltage are proportional to the number of recruited motor units.

The muscles comprising each motor unit are protected against exhaustion by a refractory period of about 1 millisecond, during which they lose their excitability. Cardiac muscle has a 250-millisecond refractory period (Tortora & Derrickson, 2021). Motor units control adjoining bundles of fibers to produce coordinated muscle contraction.

A skeletal muscle contains motor units that differ in the number and size of fibers. Motor unit firing is rotated to prevent fatigue and produce smooth movement. Motor units are recruited in order of size. Smaller units, which are highly excitable, are recruited first since they gradually build up tension. These units provide the precise motor control required in tasks like writing. Larger motor units are normally recruited last as additional force is needed to perform a bench press. In life-threatening emergencies, many motor units may be recruited at once as part of the fight-or-flight response (Tortora & Derrickson, 2021).

While the muscle fibers comprising an individual motor unit show an all-or-none response, a skeletal muscle can produce graded responses by activating different sets of motor units (Tortora & Derrickson, 2021).

Muscle Action Potentials


An alpha motor neuron divides into terminal branches that penetrate the muscle fiber membrane. The muscle fiber beneath a terminal branch is called a motor endplate. The terminal branch and motor end plate comprise the neuromuscular junction (NMJ). Graphic ©Katreryna Kon/Shutterstock.com.




When an action potential reaches the terminal branches, calcium ions move inside them. Vesicles containing ACh expel their contents into the NMJ. Each motor endplate contains 30-40 million acetylcholine (ACh) receptors. ACh release depolarizes the motor end plate, producing an end plate potential (EPP).





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The EPP triggers a depolarizing impulse called a muscle action potential (MAP) that travels sequentially along the sarcolemma into the network of T tubules, causing the terminal cisterns of the sarcoplasmic reticulum to release Ca2+ ions into the sarcoplasm to initiate muscle contraction (Tortora & Derrickson, 2021).

Skeletal muscle depolarization is the source of the SEMG signal and initiates muscle contraction.

The EMG measures muscle endplate potentials that initiate skeletal muscle contraction, but not contraction force. Dani S@unclebelang on fiverr.com drew the EMG Myth graphic © BioSource Software.



The only way to accurately record a muscle's unique activity is to place electrodes along its striations. Recording across a muscle's striations produces invalid measurements since it averages the activity of several muscles (Sherman, 2010).

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When two ACh molecules bind to a nicotinic ACh receptor on the motor end plate, a sodium channel opens, allowing sodium ions to enter the muscle fiber and then potassium ions to leave the fiber's interior. The inflow of positive sodium ions shifts the end plate's internal voltage from -80 to -90 millivolts to +50 to +75 millivolts producing a muscle action potential (MAP). This skeletal muscle fiber depolarization (positive shift in membrane potential) triggers muscle contraction (Hall & Hall, 2020).

Below is a NeXus-10 ® BioTrace+ display of the raw and integrated 100-500 Hz SEMG signal generously provided by John S. Anderson.




Below is a BioGraph ® Infiniti SEMG display.





The enzyme acetylcholine esterase (AChE) deactivates ACh to allow muscle fiber relaxation. The sodium-potassium pump restores the muscle fiber to a resting negative voltage so it can be depolarized again, resulting in a new contraction (Tortora & Derrickson, 2021).

Types of Skeletal Muscle Fibers


Skeletal muscle fibers vary in structure and function. Three fiber categories have been identified: slow oxidative, fast oxidative, and fast glycolytic. Skeletal muscles are composed of different proportions of these fibers depending on muscle action. However, the motor units that make up a muscle contain the same kind of fiber.



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Slow oxidative (SO) fibers are small red fibers are rich in myoglobin, mitochondria, and capillaries. Their capacity to produce ATP through oxidative metabolism is high. Since SO fibers split ATP at a slow rate, they contract slowly, work for extended periods, and strongly resist fatigue. Postural muscles contain a high proportion of SO fibers that enables continuous isometric contraction to resist gravity. SO fibers contribute frequencies from 20-90 Hz (Bolek et al., 2016).

Fast oxidative-glycolytic (FOG) fibers are medium-size red fibers are rich in myoglobin, mitochondria, and capillaries. They generate energy aerobically and anaerobically. Since FOG fibers rapidly split ATP, they contract quickly. These fibers show less resistance to fatigue than SO fibers and work no longer than 30 minutes. These fibers comprise a large percentage of a sprinter's leg muscles. Graphic © Peter Bernick/Shutterstock.com. FOG fibers contribute frequencies from 90-500 Hz (Bolek et al., 2016).





Fast glycolytic (FG) fibers are large white fibers are poor in myoglobin, mitochondria, and capillaries but contain extensive glycogen stores. FG fibers produce ATP using anaerobic metabolism. FG fibers rapidly split ATP and can contract more quickly than FOG fibers. Since they generate energy anaerobically, these fibers quickly fatigue and only work for a few minutes. Arm muscles contain a high proportion of these fibers (Fox & Rompolski, 2022). FG fibers contribute frequencies from 90-500 Hz (Bolek et al., 2016). Graphic © Jannarong/ Shutterstock.com.





SO fibers contain fewer skeletal muscles in their motor units than FOG and FG fibers, are used more frequently in daily activities, and are recruited earlier (Fox & Rompolski, 2022).


         
Exercise can change the properties but not the number of muscle fibers. Endurance exercises like running can slowly transform FG fibers into FOG fibers, increasing diameter, mitochondria, capillaries, and strength. In contrast, activities like weight lifting that require explosive force for brief periods increase FG fiber size and strength. Muscle enlargement (hypertrophy) produces a weight lifter's bulging muscles (Tortora & Derrickson, 2021). Graphic © Dmytro Zinkevych/Shutterstock.




Muscle biopsies of chronic pain patients raise the possibility that their postural muscles contain a disproportionate number of FG fibers that fatigue more rapidly than SO fibers. Increased vulnerability to fatigue could result in a buildup of metabolic byproducts (like lactate and hydrogen ions) that can stimulate pain receptors (Tortora & Derrickson, 2021). Graphic © George Rudy/Shutterstock.com.





When treating urinary incontinence with EMG biofeedback for the pelvic floor, clinicians must train pelvic floor muscle endurance (Laycock & Haslam, 2008) and quick flicks. These tonic and phasic contractions prevent leakage during events that increase intra-abdominal pressure, such as coughing, lifting, and sneezing (Lee & Kaufman 2017). Graphic © Alila Medical Media/Shutterstock.com.


The EMG Signal


The SEMG records muscle action potentials from skeletal motor units.



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The frequency range for surface recording is 2-1,000 Hz.

Tissues (skin, subcutaneous fat, muscle, and connective tissue) absorb the higher frequencies that can be detected within the muscle fibers by inserted electrodes (Stern, Ray, & Quigley, 2001).

Below is a BioGraph ® Infiniti SEMG FFT display of a frontales placement. Lower frequency amplitude increases with stronger muscle contraction and the energy distribution shifts toward the higher frequencies.






The amplitude or strength of an electromyogram reflects the number of active motor units, their firing rate, and distance from the electrodes.







A bandpass of 20-200 Hz covers the most critical SEMG frequencies (Andreassi, 2007). Slow-twitch fibers generate frequencies from 10-90 Hz, and fast-twitch fibers generate frequencies from 90-500 Hz.

The greatest concentration of power in a resting muscle lies between 10 and 150 Hz (Stern et al., 2001).

Strong muscle contraction shifts the distribution of SEMG power upwards and to the right toward higher frequencies.

Fatigue decreases motor unit firing rate and shifts the distribution of SEMG power to lower frequencies, lowering the median frequency. Thought Technology Ltd. generously shared the illustration below.


Fatigue


A wide bandpass is desirable because it better monitors a muscle's dynamic range. Failure to use the correct bandpass may make a muscle appear more relaxed than it is or miss fatigue entirely (Sherman, 2010).

Muscle Action Potentials Are Briefer Than The Muscle Contraction Period

Skeletal muscle fiber depolarization (which produces the SEMG signal) lasts 1 to 2 ms and ends before a muscle fiber starts to contract. The SEMG reflects muscle depolarization, not muscle contraction. The period of mechanical contraction is considerably longer and lasts from 10-100 ms.

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Calcium binds to troponin, myosin-actin cross-bridges form, and peak muscle fiber tension develops. The relaxation phase also lasts 10-100 ms. Calcium is actively transported back to the sarcoplasmic reticulum, myosin heads detach from actin, and tension declines (Tortora & Derrickson, 2021)

A muscle action potential depolarizes the interior of the muscle fiber. This membrane potential change releases stored calcium ions from the sarcoplasmic reticulum. The presence of calcium ions allows actin (thin) and myosin (thick) filaments to bind to each other forming cross-bridges. Simple contact will not shorten a muscle. Myosin must bind to actin, move it inward, break contact, and bind again. These power strokes use stored energy released by splitting ATP.

A single thick filament forms 600 cross-bridges that attach and detach from actin about five times per second. While some myosin heads perform power strokes, others are detached and preparing to form cross-bridges with actin. Sliding filament video © sciencepics/Shutterstock.com.







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Remember that actin is attached to the Z discs that define each sarcomere (muscle compartment). When myosin ratchets actin inward toward a sarcomere's center, the entire sarcomere shortens to up to 50 percent of its resting length. Sarcomere shortening produces the force of muscle contraction that can move or stabilize limbs. A sarcomere's average length is 2.5 micrometers. It can shorten to about 1.5 micrometers and stretch to about 3 micrometers. Graphic © Blamb/Shutterstock.com.









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Skeletal Muscle Contraction


Muscle contraction may be isometric or isotonic. Muscles produce tension with minimal fiber shortening during isometric contraction. For example, muscles do not appreciably shorten when you perform a plank due to the resistance of gravity. Posture results from continuous isometric contraction. SO fibers are specialized for this contraction due to their fatigue resistance.



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Isotonic contraction produces movement by exerting tension on an attached structure (bone). These contractions may be either concentric or eccentric. Performing curls involves concentric contraction, where fibers shorten as you pull the bar towards your torso. Graphic © ra2studio/Shutterstock.com.





In contrast, a squat involves eccentric contraction where fibers lengthen as you lower yourself (Tortora & Derrickson, 2021). Graphic © Anatol Misnikou/Shutterstock.com.




Muscles groups work together to perform actions like the series of flips shown below.





You contract at least nine muscle groups in a specific sequence to open your mouth. You activate a different sequence to close your mouth. SEMG biofeedback records the electrical activity of surface muscles. The SEMG usually monitors the activity of muscle groups instead of individual muscles.

Gene Promotes Muscle Strength

Physical activity activates a newly discovered C18ORF25 gene that increases muscle strength without always changing muscle size (Blazev et al., 2022). Graphic © Blue Andy/Shutterstock.com.


Stretch Reflex


Muscle spindles are stretch receptors that lie in parallel with skeletal muscle fibers. Muscle spindles allow us to restore muscle length when muscle fibers lengthen. The image below from Dr. J. H. H. Scott of Leicester University is from the first dorsal interosseus muscle of the hand.




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When their annulospiral receptors stretch as a muscle lengthens, they activate alpha motor neurons to strengthen muscle contraction to increase muscle tone. Physicians assess the monosynaptic stretch reflex when stretching the patellar tendon with a rubber mallet to elicit a knee jerk (Breedlove & Watson, 2020). Graphic © Aldona Griskeviciene/Shutterstock.com.




The graphic below that depicts a muscle spindle was retrieved from irunfar.com.
 

Tendon Reflex


Golgi tendon organs are force detectors that lie in series with skeletal muscle fibers. They protect muscles and tendons from injury due to forceful contractions. Graphic retrieved from firstclassmed.com.




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When excessive contraction threatens to damage muscle and tendon, they inhibit the responsible alpha motor neurons to prevent injury. This protective mechanism is called the tendon reflex (Breedlove & Watson, 2020). The graphic below that depicts the tendon reflex was retrieved from irunfar.com.

Muscle Action


Skeletal muscles attach to the articulating bones of a joint and transmit force through the tendons to bones and tissues like the skin. Muscle contraction moves the two bones that comprise a joint unequally and pulls one of the articulating bones at a joint toward the more stationary bone. A muscle tendon's attachment to the more stationary bone is called its origin, and its other tendon's attachment to the movable bone is called its insertion. Graphic © stihii/Shutterstock.com.



A spring on a door provides a good analogy. The end of the spring attached to the frame is the origin; the end attached to the door is the insertion. The origin is usually proximal; the insertion is distal (Tortora & Derrickson, 2021).

The bone where a muscle originates can move when the insertion is fixed. For example, when the tibia/foot is off the ground (open chain), the tibia moves on the femur. When the tibia/foot is fixed to the ground (closed chain), the femur moves on the tibia. Think of a knee extension machine versus performing a squat (Jadali, 2021).

The movement produced by a muscle's contraction is its action. The same muscle can perform several different actions. Some muscles can perform reverse muscle action (RMA) in which the origin and insertion switch to perform the opposite movement. For example, the gluteus maximus normally extends the hips. However, the gluteus maximus pulls the trunk upright when the trunk bends forwards. Graphic ©studioloco/Shutterstock.com.

Muscles that operate joints are arranged in opposing prime mover-antagonist pairs, often located on the opposite side of a bone or joint. A prime mover (agonist) contracts to perform an action while its antagonist stretches to allow the action. For example, when the biceps brachii (prime mover) contracts to flex the forearm at the elbow joint, the triceps brachii (antagonist) must relax, or else flexion will be prevented (Tortora & Derrickson, 2021).

When muscles are arranged in opposing pairs, the prime mover and antagonist can switch roles to perform different movements. For example, when you extend the forearm against a resistance, the triceps brachii becomes the prime mover and the biceps, the antagonist.

In the lower extremities, the extensor chain muscles are generally stronger than their flexor counterparts due to their role in propulsion during the gait cycle and combatting gravity (Jadali, 2021).

When prime mover and antagonist muscles contract concurrently in stroke patients, this can produce spasms and rigidity. The table below was adapted from Thought Technology Ltd.'s (2009) Basics of Surface Electromyography Applied to Physical Rehabilitation and Biomechanics.






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The next section describes synergists, fixators, flexors and extensors, abductors and adductors, levators and depressors, supinators, and pronators, dorsiflexors and plantar flexors, inverters and evertors, tensors, and rotators.

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Prime movers may cross other joints before attaching to the joint where it exerts its primary action. Synergists aid prime movers by stabilizing intermediate joints to prevent co-contraction (e.g., flexing your fingers without flexing the wrist) and unwanted movement. They are usually near the prime mover. For example, the deltoid and pectoralis major anchor both the arm and shoulder when the biceps brachii (agonist) contracts to flex the forearm (Tortora & Derrickson, 2021).




Muscles that act as fixators stabilize a prime mover's origin for more efficient action. By steadying a limb's proximal end, fixators support movements at the distal end. For example, the mobile scapula is the origin of several arm muscles and must be steadied for actions like abduction. Depending on their movement, the same muscles may act as prime movers, antagonists, synergists, or fixators (Tortora & Derrickson, 2021).

The quadriceps and tibialis anterior muscles of your leg contract together to dorsiflex (bend the foot upwards) during the forward swing phase of walking.
 
Flexors decrease the angle between two bones. Biceps brachii flexes and supinates (turns up) the forearm. Weightlifters flex forearms, not muscles.




Extensors
increase the angle between two bones. Triceps brachii extends the forearm. The motor control system coordinates synergist, flexor, and extensor contraction to produce the needed limb position.






Abductors
move a limb away from the center of the trunk or a body part. The deltoid abducts the arm.






Adductors
move a limb toward the center of the trunk or a body part. Pectoralis major adducts the arm.






Levators
produce upward movement. Levator scapulae elevates the shoulder blades. 






Depressors
produce downward movement. Latissiumus dorsi is the primary shoulder girdle and scapular (shoulder blade) depressor. Scapular depression is mainly due to the failure of scapular elevators and upward rotators.






Supinators
turn the palm upward (anteriorly). Supinator exposes the anterior side of the forearm.





Pronators
turn the palm downward (posteriorly). Pronator teres exposes the posterior side of the forearm.






Dorsiflexors
point the toes toward the shin (superiorly) through flexion at the ankle joint. Tibialis anterior dorsiflexes and inverts the foot during the swing phase of walking. Dorsiflexion is disrupted during a neuromuscular disorder called foot drop in which the toes drag on the ground.






Plantar flexors
point the toes downward (inferiorly) through extension at the ankle joint. The gastrocnemius/soleus complex, which acts on the Achilles tendon, is the body's primary plantar flexor. The grastrocnemius is considered the most important walking muscle (Jadali, 2021). 






Invertors
turn the sole of a foot inward. The tibialis posterior and tibialis anterior are the primary invertors. Posterior tibial tendonitis is a common orthopedic injury related to inversion (Jadali, 2021).
 





Evertors turn the sole of a foot outward. The fibularis longus and tertius muscles perform eversion. Peroneal tendonitis affects eversion (Jadali, 2021).

 




Tensors
make a body part more rigid. Tensor fasciae latae flexes and abducts the thigh.

 




Rotators move a bone around a longitudinal axis. Obturator externus laterally rotates the thigh (Tortora & Derrickson, 2021).

 





The complex movement sequences used in jumping rope integrate many muscle actions.


MUSCLE ACTIONS, SENSOR PLACEMENTS, AND CLINICAL APPLICATIONS


The next two sections provide a detailed description of muscle actions, sensor placements, and clinical applications. Although the BCIA Biofeedback Blueprint does not cover this content at this level of detail, it can be invaluable for applicants during their mentorship and for understanding research reports.

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Skeletal Muscles of Clinical Interest (Face)


Basmajian and Blumenstein (1980) and Neblett (2006) were the references for sensor placement. Tortora and Derrickson (2021) was the resource for muscle action.





               




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Frontales

Action: draws the scalp forward, raises eyebrows, and wrinkles the forehead

Sensor placement: locate the actives between the eyebrows and hairline

Clinical application: anxiety, stress profile, and tension-type headache





Temporalis

Action: elevates and retracts the mandible (jaw)

Sensor placement: locate actives above the zygomatic arch (horizontal bony ridge from temporomandibular joint to the cheek)

Clinical application: tension-type headache




A bilateral placement may be used with a wide bandpass to assess and treat headaches (Neblett, 2012).






Masseter

Action: elevates and protracts the mandible

Sensor placement: locate actives using the angle of the jaw as a landmark

Clinical application: temporomandibular disorders (TMD)






Zygomaticus

Action: draws the corner of the mouth upward and outward when you smile

Sensor placement: locate actives above and at approximately a 45o angle from the corner of the mouth

Clinical application: Bell's palsy and stroke








Orbicularis oculus

Action: closes the eyelids and wrinkles the forehead

Sensor placement: locate actives immediately below the center of each eye

Clinical application: Bell's palsy, blepharospasm, and stroke







Sternocleidomastoid (SCM)

Action: flexes the vertebral column and rotates the head to the opposite side (contract left SCM and head twists to the right)

Sensor placement: center the actives 50% of the distance from the mastoid (bulge behind the outer ear) to the medial end of the clavicle (collarbone), which places them below the jaw along a vertical line

Clinical application: torticollis






Cervical paraspinal (semispinalis capitis and splenius capitis)

Action: extends, flexes, and rotates the head

Sensor placement: locate the actives around C2 and C5 lateral and medial to the spine

Clinical application: neck and shoulder pain and tension-type headache








Upper trapezius

Action: rotates and elevates the scapula (shoulder blade), extends, flexes, and rotates the head and neck

Sensor placement: center the actives between C7 (seventh cervical vertebra) and the angle of the acromion (posterior to bony triangle at the top of the shoulder)

Clinical application: shoulder pain and tension-type headache




Bilateral monitoring of the upper trapezius with a wide bandpass may be used to assess and treat pain due to shoulder injury (Neblett, 2012).




Bilateral monitoring of the upper trapezius with a narrow bandpass may also be used to assess and treat pain due to shoulder injury (Neblett, 2012).






Bilateral monitoring of the cervical and upper trapezius muscles using a narrow or wide bandpass may be used to assess and treat headache and neck and upper back pain (Neblett, 2012).




Skeletal Muscles of Clinical Interest




     
         
               


Posterior deltoid

Action: abducts, flexes, extends, and rotates the arm

Sensor placement: locate the actives behind the angle of the acromion (posterior to the bony triangle at the top of the shoulder)






Lateral head of triceps brachii

Action: extends the elbow joint

Sensor placement: center the actives 50% of the distance between the angle of the acromion (posterior to bony triangle at the top of the shoulder) and the olecranon process (behind the elbow)







Biceps brachii

Action: flexes and supinates the forearm

Sensor placement: center the actives over the bulge in this muscle








Latissimus dorsi

Action: extends, adducts, rotates the arm, and moves the arm inferiorly and posteriorly

Sensor placement: center the actives under the inferior angle of the scapula (shoulder blade)






Extensor carpi ulnaris

Action: extends and adducts the hand at the wrist joint

Sensor placement: center the actives 1/3 of the distance between the posterior medial epicondyle and the posterior styloid process of the forearm






A flexor/extensor placement using a wide bandpass may be used to assess and treat crushing injury to the arm and hand (Neblett, 2012).






A web placement using a wide bandpass may also be used to assess and treat crushing injury to the arm and hand (Neblett, 2012).






Abductor pollicis brevis

Action: abducts the thumb and assists in opposition

Sensor placement: center the actives over the most prominent bulge at the proximal end of the thumb



Adductor pollicis

Action: adducts the thumb

Sensor placement: center the actives within a triangle that starts at the skin web


Vastus lateralis (VL) and vastus medialis (VMO)


Action: each muscle extends the leg at the knee joint

Sensor placement: locate the actives vertically within the oval bulges above the patella (kneecap) for the vastus lateralis (VL) and vastus medialis obliquus (VMO) sites, respectively.






Tibialis anterior

Action: dorsiflexes and inverts the foot

Sensor placement: center the actives vertically within an oval region around the tuberosity of the tibia (shin bone)

Clinical application: foot drop





 
Gastrocnemius

Action: plantar flexes the foot and flexes the knee joint

Sensor placement: center the actives vertically over the bulges of either head of this muscle





Erector spinae (sacrospinalis)

Action: maintains the erect position of the spine and extends the vertebral column

Sensor placement: locate active pairs lateral to the spine, above the iliac crest (rounded upper margins of the ilium bone above the buttocks) about L4 and below the bottom of the ribs at about L2. Clinicians should palpate these landmarks to achieve consistent lumbar placements

Clinical application: low back pain






Bilateral monitoring of the lumbar region using a wide bandpass may be used to assess and treat middle back pain (Neblett, 2012).








Glossary


α-actinin: structural Z disc protein that connects actin to titin molecules.

A band:
dark, central region of a sarcomere that runs the length of thick filaments and includes overlapping actin filaments.

abductors: muscles that move a limb away from the center of the trunk or a body part. For example, abductor pollicis brevis moves the thumb outward.

acetylcholine (ACh): the neurotransmitter released by alpha motor neurons at the neuromuscular junction depolarizes motor endplates producing a muscle action potential (MAP).

acetylcholine esterase (AChE): the enzyme that deactivates ACh to allow skeletal muscle fiber relaxation.

actin: contractile protein that is the main component of thin filaments; each actin molecule contains a binding site for myosin heads which form cross-bridges during muscle contraction.

active tension: force generated by muscle fibers when myosin filaments pull Z discs together, shortening the sarcomeres.

adductors: muscles that move a limb toward the center of the trunk or a body part. For example, adductor pollicis moves the thumb inward.

alpha motor neuron: motoneuron that innervates the skeletal muscles fibers that comprise its motor unit.

amplitude: the strength of an EMG signal that is measured in microvolts.

angle of the acromion: the site posterior to the bony triangle at the top of the shoulder.

annulospiral receptors: muscle spindle length receptors that stretch as a muscle lengthens and activate alpha motor neurons to strengthen muscle contraction to increase muscle tone.

antagonist: a muscle that opposes a prime mover's action and yields to its movement. For example, the triceps brachii (antagonist) opposes flexion by the biceps brachii (prime mover or agonist).

biceps brachii: a muscle that flexes and supinates the forearm. Active electrodes are centered over the bulge in this muscle.

C7: the seventh cervical vertebra.

cardiac muscle: muscle fibers that comprise most of the heart wall. These fibers have crossed striations that allow the heart to pump and contain the same actin and myosin filaments, bands, zones, and Z discs as skeletal muscles.

clavicle: collarbone.

concentric contraction: fibers shorten as your body moves upward, for example, a pull-up.

deltoid: synergist that along with the pectoralis major muscle anchors both the arm and shoulder when the biceps brachii (agonist) contracts to flex the forearm. Active electrodes are located behind the angle of the acromion (posterior to the bony triangle at the top of the shoulder) for the posterior deltoid.

depressors: muscles that produce downward movement. For example, the latissimus dorsi lowers the shoulder blades and shoulder girdle.

dorsiflexors: muscles that point the toes toward the shin (superiorly) through flexion at the ankle joint. For example, the tibialis anterior dorsiflexes and inverts the foot during the swing phase of walking.

eccentric contraction: fibers lengthen as you lower yourself, for example, when performing a squat.

efferent nerves: nerves like alpha motor neurons that leave the central nervous system.

erector spinae (sacrospinalis): a muscle that maintains the erect position of the spine and extends the vertebral column. Active pairs are located lateral to the spine, above the iliac crest (rounded upper margins of the ilium bone above the buttocks) about L4, and below the bottom of the ribs at about L2.

evertors: muscles that turn the sole of a foot outward. For example, fibularis longus/tertius are the primary eversion muscles.

exerkines: a group of signaling molecules released during exercise that plays a crucial role in mediating its systemic benefits, including improved metabolism, cardiovascular health, and cognition.

extensor carpi ulnaris: a muscle that extends and adducts the hand at the wrist joint.

extensor digitorum longus: a muscle that dorsiflexes, extends the toes, and everts the foot.

extensors: muscles that increase the angle between two bones. For example, the triceps brachii extends the forearm.

extrafusal fibers: skeletal muscle fibers that are striated (striped) due to alternating light and dark bands.

fascia: fibrous connective tissue that divides muscles into functional groups, aids muscle movement, transmits force to bones, encloses and supports blood vessels and nerve fibers, and secures organs in their place.

fascicles: muscle fiber bundles.

fast glycolytic (FG) fibers: white fibers are poor in myoglobin, mitochondria, and capillaries but contain extensive stores of glycogen. FG fibers produce ATP using anaerobic metabolism that cannot continuously supply needed ATP, causing these fibers to fatigue easily. They are also called fast-twitch A fibers.

fast oxidative-glycolytic (FOG) fibers: red fibers are rich in myoglobin, mitochondria, and capillaries. Their capacity to produce ATP through oxidative metabolism is high, and FOG fibers split ATP rapidly producing high contraction velocities. They show less resistance to fatigue than SO fibers. They are also called fast-twitch B fibers.

flexor digitorum longus: a muscle that plantar flexes, flexes toes, and inverts the foot.

flexors: muscles that decrease the angle between two bones. For example, the biceps brachii flexes and supinates (turns up) the forearm.

frontales: a muscle that draws the scalp forward, raises eyebrows, and wrinkles the forehead. The actives are located between the eyebrows and hairline.

gastrocnemius: a muscle that plantar flexes the foot and flexes the knee joint. Active electrodes are centered vertically over the bulges of either head of this muscle.

Golgi tendon organs: force detectors that lie in series with skeletal muscle fibers. When excessive contraction threatens to damage muscle and tendon, they inhibit the responsible alpha motor neurons to prevent injury. This protective mechanism is called the tendon reflex.

H bands: narrow central region in each A band containing thick filaments.

I bands: lighter, less dense region of a sarcomere that contains the remaining thin but no thick filaments and is bisected by a Z disc.

iliac crest: rounded upper margins of the ilium bone above the buttocks.

ilium: bone located above the buttocks.

insertion: the more movable bone at a joint.

invertors: muscles that turn the sole of a foot inward. For example, the tibialis posterior and tibialis anterior are the primary invertors.

isometric contraction: a contraction in which muscles produce tension with minimal fiber shortening.

isotonic contraction: a contraction in which muscles produce movement by exerting tension on an attached structure (bone).

lateral head of triceps brachii: a muscle that extends the elbow joint.

latissimus dorsi: a muscle that extends, adducts, medially rotates the arm, and moves the arm inferiorly and posteriorly. The actives are centered actives under the inferior angle of the scapula (shoulder blade).

levators: muscles that produce upward movement. For example, the levator scapulae elevates the shoulder blades.

M lines: central region of an H zone, located at the center of a sarcomere, that contains the proteins that connect thick filaments.

mandible: jaw.

masseter: a muscle that elevates and protracts the mandible. The actives are located using the angle of the jaw as a landmark.

mastoid: the bulge behind the outer ear.

medial epicondyle: the medial prominence of the bottom aspect of humerus located on the inside of the elbow.

monosynaptic stretch reflex: a reflex that maintains postural muscle tone and stabilizes limb position by correcting increases in muscle length.

motor unit: an alpha motor neuron and the muscle fibers it controls.

multiunit smooth muscle: muscle found in the walls of large arteries, airways to the lungs, the arrector pili muscles that move hair follicles, the muscle rings of the iris, and the ciliary body that focuses the lens.

muscle action: the movement produced by skeletal muscle contraction.

muscle action potential (MAP): a depolarizing impulse that travels along the sarcolemma and T tubules, producing the EMG signal and initiating skeletal muscle contraction.

muscle spindles: stretch receptors that lie in parallel with skeletal muscle fibers and trigger the monosynaptic stretch reflex.

myofibrils: each skeletal muscle fiber comprises hundreds to thousands of these units built from thin and thick filaments.

myofilaments: building blocks of a myofibril. Thin filaments are primarily composed of actin, and thick filaments are mainly composed of myosin.

myomesin: structural protein that comprises a sarcomere's M line, binds to titin, and connects adjacent thick filaments.

myosin: contractile protein that comprises thick filaments; a single molecule contains two heads that bind to sites on actin during muscle contraction.

neuromuscular junction (NMJ): the synapse between an alpha motor neuron and a skeletal muscle.

nicotinic ACh receptor: an ionotropic receptor on the skeletal muscle endplate binds two ACh molecules to allow sodium ions to enter the muscle fiber.

olecranon process: the bony projection behind the elbow.

orbicularis oculus: muscle that closes the eyelids and wrinkles the forehead. The actives are located immediately below the center of each eye.

origin: a muscle's attachment to the more stationary bone.

passive tension: the elasticity of connective tissue allows muscle fibers to produce, sum, and transmit force.

patella: kneecap.

peak frequency: the highest amplitude frequency in the SEMG signal.

pectoralis major: synergist that, along with the deltoid, anchors both the arm and shoulder when the biceps brachii (agonist) contracts to flex the forearm.

plantar flexors: muscles that point the toes downward (inferiorly) through extension at the ankle joint. For example, the gastrocnemius/soleus complex act on the Achilles tendon.

posterior deltoid: a muscle that abducts, flexes, extends, and rotates the arm. Active electrodes are located behind the angle of the acromion (posterior to the bony triangle at the top of the shoulder).

posterior styloid process: the rear aspect the projection from the medial and back part of the ulna (elbow bone).

power strokes: after actin and myosin filaments form cross bridges, myosin moves actin inward, breaks contact, binds to actin, and then repeats this process.

prime mover (agonist): a muscle directly responsible for producing a specific movement and is opposed by an antagonist. For example, when the biceps brachii (prime mover) contracts to flex the forearm at the elbow joint, the triceps brachii (antagonist) must relax, or else flexion will be prevented.

pronators: muscles that turn the palm downward (posteriorly). For example, pronator teres exposes the posterior side of the forearm.

proprioceptive feedback: information about body position and movement.

quadriceps: a muscle that, along with the tibialis anterior, dorsiflexes (bends the foot upwards) during the forward swing phase of walking.

refractory period: 5-millisecond interval during which skeletal muscles lose their excitability to prevent exhaustion.

reverse muscle action (RMA): the origin and insertion switch to perform the opposite movement.

rotators: muscles that move a bone around a longitudinal axis. For example, the obturator externus laterally rotates the thigh.

sarcolemma: the skeletal muscle fiber membrane.

sarcomeres: skeletal muscle compartments separated by dense zones called Z discs.

scapula: shoulder blade.

SEMG: surface electromyogram.

SEMG power: the energy of the surface electromyogram measured in microvolts.

single-unit smooth muscle: smooth muscle that comprises part of the walls of small arteries and veins, the stomach, intestines, uterus, and urinary bladder.

skeletal muscles: extrafusal muscles that move the bones of the skeleton and are striated (striped) muscles due to alternating light (I) and dark (A) bands.

slow oxidative (SO) fibers: red fibers rich in myoglobin, mitochondria, and capillaries with a high capacity to produce ATP through oxidative metabolism. Since SO fibers split ATP at a slow rate, contraction velocity is slow, and these fibers are highly resistant to fatigue. Postural muscles contain a high proportion of SO fibers. They are also called slow-twitch fibers.

smooth muscle: single-unit and multi-unit muscle fibers whose contraction starts more gradually and persists longer than skeletal muscle contraction. Smooth muscles can shorten and stretch more than other muscles and usually operate involuntarily. Some tissues have an intrinsic rhythm (autorhythmicity).

sodium-potassium pump: the mechanism that exchanges sodium ions for potassium ions to restore the muscle fiber to a resting negative voltage so that it can be depolarized anew, resulting in new contraction.

sternocleidomastoid (SCM): a muscle that flexes the vertebral column and rotates the head to the opposite side (contract left SCM and head twists to the right). Actives are centered 50% of the distance from the mastoid (bulge behind the outer ear) to the medial end of the clavicle (collarbone), which places them below the jaw along a vertical line.

striated: striped. For example, skeletal muscle fibers are striped due to thin (light) and thick (dark) filaments.

supinators: muscles that turn the palm upward (anteriorly). For example, the supinator exposes the anterior side of the forearm.

synergists: muscles that stabilize a joint to reduce the origin's interference with movement. For example, the deltoid and pectoralis major anchor both the arm and shoulder when the biceps brachii (agonist) contracts to flex the forearm.

temporalis: a muscle that elevates and retracts the mandible (jaw). Clinicians place active electrodes above the zygomatic arch (horizontal bony ridge from temporomandibular joint to the cheek).

tendon: connective tissue that connects muscle to bone and transmits the force of muscle contraction to the attached bones.

tendon reflex:
Golgi tendon organs are force detectors that lie in series with skeletal muscle fibers. When excessive contraction threatens to damage muscle and tendon, they inhibit the responsible alpha motor neurons to prevent injury.

tensors: muscles that make a body part more rigid. For example, the tensor fasciae latae flexes and abducts the thigh.

thick filaments: dark-colored filaments that are mainly composed of myosin.

thin filaments: light-colored filaments that are mainly composed of actin.

tibia: shinbone.

tibialis anterior: a muscle that dorsiflexes and inverts the foot. Actives are vertically centered within an oval region around the tuberosity of the tibia (shinbone).

titin: a structural protein that links a Z disc to the M line in the center of a sarcomere to stabilize the thick filament position and contribute to myofibril elasticity and extensibility.

triceps brachii (lateral head): a muscle that extends the elbow joint. Actives are centered 50% of the distance between the angle of the acromion (posterior to bony triangle at the top of the shoulder) and the olecranon process (behind the elbow).

tropomyosin: a regulatory protein component of thin filaments that prevents myosin heads from binding to actin when a muscle fiber is relaxed.

troponin: a regulatory protein component of thin filaments that changes shape when calcium ions bind, removing tropomyosin from covering myosin-binding sites to allow the formation of cross-bridges and muscle contraction.

upper trapezius: a muscle that rotates and elevates the scapula (shoulder blade), extends, flexes, and turns the head and neck. Active electrodes are centered between C7 (seventh cervical vertebra) and the angle of the acromion (posterior to bony triangle at the top of the shoulder).

vastus lateralis (VL) and vastus medialis (VMO): muscles that extend the leg at the knee joint. Clinicians position active electrodes vertically within the oval bulges above the patella (kneecap) for the vastus lateralis (VL) and vastus medialis obliquus (VMO) sites, respectively.

Z discs: thin, plate-shaped dense zones that separate sarcomeres.

zygomatic arch: the horizontal bony ridge from temporomandibular joint to the cheek.

zygomaticus: a muscle that draws the corner of the mouth upward and outward when you smile. Actives are located above and at approximately a 45o angle from the corner of the mouth.

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Assignment


Now that you have completed this module, identify the muscles you monitor and train in your clinical practice using the muscle diagrams shown above.

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