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current-item">Muscle</span></div><div class="d-inline-block col-md-11 p-0"><h1 class="entry-title">Muscle</h1></div><div class="float-md-right position-sm-absolute pt-md-4"><g:sharetoclassroom size=32 url="https://www.biologyonline.com/tutorials/muscle"></g:sharetoclassroom></div><div class=st-container> <img fetchpriority=high width=640 height=461 src="data:image/svg+xml,%3Csvg%20xmlns='http://www.w3.org/2000/svg'%20viewBox='0%200%20640%20461'%3E%3C/svg%3E" data-src=https://www.biologyonline.com/wp-content/uploads/2020/02/Muscle-768x553.jpg class="st-img wp-post-image lazy" alt=Muscle decoding=async data-srcset="https://www.biologyonline.com/wp-content/uploads/2020/02/Muscle-768x553.jpg 768w, https://www.biologyonline.com/wp-content/uploads/2020/02/Muscle-233x168.jpg 233w, https://www.biologyonline.com/wp-content/uploads/2020/02/Muscle-800x576.jpg 800w, https://www.biologyonline.com/wp-content/uploads/2020/02/Muscle-1536x1106.jpg 1536w, https://www.biologyonline.com/wp-content/uploads/2020/02/Muscle.jpg 1980w" data-sizes="(max-width: 640px) 100vw, 640px"><p>The human muscle are comprised of cells specialized for generating force and movement</p></div><div id=ez-toc-container class="ez-toc-v2_0_68_1 counter-hierarchy ez-toc-counter ez-toc-custom ez-toc-container-direction"><p class=ez-toc-title>Table of Contents</p> <label for=ez-toc-cssicon-toggle-item-674fba6ebe0d6 class=ez-toc-cssicon-toggle-label><span class><span class=eztoc-hide style=display:none;>Toggle</span><span class=ez-toc-icon-toggle-span><svg style="fill: #999;color:#999" xmlns=http://www.w3.org/2000/svg class=list-377408 width=20px height=20px viewBox="0 0 24 24" fill=none><path d="M6 6H4v2h2V6zm14 0H8v2h12V6zM4 11h2v2H4v-2zm16 0H8v2h12v-2zM4 16h2v2H4v-2zm16 0H8v2h12v-2z" fill=currentColor></path></svg><svg style="fill: #999;color:#999" class=arrow-unsorted-368013 xmlns=http://www.w3.org/2000/svg width=10px height=10px viewBox="0 0 24 24" version=1.2 baseProfile=tiny><path d="M18.2 9.3l-6.2-6.3-6.2 6.3c-.2.2-.3.4-.3.7s.1.5.3.7c.2.2.4.3.7.3h11c.3 0 .5-.1.7-.3.2-.2.3-.5.3-.7s-.1-.5-.3-.7zM5.8 14.7l6.2 6.3 6.2-6.3c.2-.2.3-.5.3-.7s-.1-.5-.3-.7c-.2-.2-.4-.3-.7-.3h-11c-.3 0-.5.1-.7.3-.2.2-.3.5-.3.7s.1.5.3.7z"/></svg></span></span></label><input type=checkbox id=ez-toc-cssicon-toggle-item-674fba6ebe0d6 checked aria-label=Toggle><nav><ul class='ez-toc-list ez-toc-list-level-1 ' ><li class='ez-toc-page-1 ez-toc-heading-level-2'><a class="ez-toc-link ez-toc-heading-1" href=#Skeletal_Muscle title="Skeletal Muscle">Skeletal Muscle</a><ul class=ez-toc-list-level-3 ><li class=ez-toc-heading-level-3><a class="ez-toc-link ez-toc-heading-2" href=#Structure title=Structure>Structure</a></li><li class='ez-toc-page-1 ez-toc-heading-level-3'><a class="ez-toc-link ez-toc-heading-3" href=#Molecular_mechanisms_of_contraction title="Molecular mechanisms of contraction">Molecular mechanisms of contraction</a></li><li class='ez-toc-page-1 ez-toc-heading-level-3'><a class="ez-toc-link ez-toc-heading-4" href=#Sliding-filament_mechanism title="Sliding-filament mechanism">Sliding-filament mechanism</a></li><li class='ez-toc-page-1 ez-toc-heading-level-3'><a class="ez-toc-link ez-toc-heading-5" href=#Roles_of_troponin_tropomyosin_and_calcium title="Roles of troponin, tropomyosin and calcium">Roles of troponin, tropomyosin and calcium</a></li><li class='ez-toc-page-1 ez-toc-heading-level-3'><a class="ez-toc-link ez-toc-heading-6" href=#Sarcoplasmic_reticulum title="Sarcoplasmic reticulum">Sarcoplasmic reticulum</a></li><li class='ez-toc-page-1 ez-toc-heading-level-3'><a class="ez-toc-link ez-toc-heading-7" href=#Neuromuscular_junction title="Neuromuscular junction">Neuromuscular junction</a></li><li class='ez-toc-page-1 ez-toc-heading-level-3'><a class="ez-toc-link ez-toc-heading-8" href=#Mechanics_of_a_single_fiber_contraction title="Mechanics of a single fiber contraction">Mechanics of a single fiber contraction</a></li><li class='ez-toc-page-1 ez-toc-heading-level-3'><a class="ez-toc-link ez-toc-heading-9" href=#Twitch_contraction title="Twitch contraction">Twitch contraction</a></li><li class='ez-toc-page-1 ez-toc-heading-level-3'><a class="ez-toc-link ez-toc-heading-10" href=#Frequency-tension_relation title="Frequency-tension relation">Frequency-tension relation</a></li><li class='ez-toc-page-1 ez-toc-heading-level-3'><a class="ez-toc-link ez-toc-heading-11" href=#Length-tension_relation title="Length-tension relation">Length-tension relation</a></li><li class='ez-toc-page-1 ez-toc-heading-level-3'><a class="ez-toc-link ez-toc-heading-12" href=#Skeletal_muscle_energy_metabolism title="Skeletal muscle energy metabolism">Skeletal muscle energy metabolism</a></li><li class='ez-toc-page-1 ez-toc-heading-level-3'><a class="ez-toc-link ez-toc-heading-13" href=#Muscle_fatigue title="Muscle fatigue">Muscle fatigue</a></li><li class='ez-toc-page-1 ez-toc-heading-level-3'><a class="ez-toc-link ez-toc-heading-14" href=#Types_of_skeletal_muscle_fibers title="Types of skeletal muscle fibers">Types of skeletal muscle fibers</a></li><li class='ez-toc-page-1 ez-toc-heading-level-3'><a class="ez-toc-link ez-toc-heading-15" href=#Control_of_muscle_tension title="Control of muscle tension">Control of muscle tension</a></li><li class='ez-toc-page-1 ez-toc-heading-level-3'><a class="ez-toc-link ez-toc-heading-16" href=#Muscle_adaptation_to_exercise title="Muscle adaptation to exercise">Muscle adaptation to exercise</a></li><li class='ez-toc-page-1 ez-toc-heading-level-3'><a class="ez-toc-link ez-toc-heading-17" href=#Lever_action_of_muscles_and_bones title="Lever action of muscles and bones">Lever action of muscles and bones</a></li><li class='ez-toc-page-1 ez-toc-heading-level-3'><a class="ez-toc-link ez-toc-heading-18" href=#Skeletal_muscle_disease title="Skeletal muscle disease">Skeletal muscle disease</a></li></ul></li><li class='ez-toc-page-1 ez-toc-heading-level-2'><a class="ez-toc-link ez-toc-heading-19" href=#Smooth_Muscle title="Smooth Muscle">Smooth Muscle</a><ul class=ez-toc-list-level-3 ><li class=ez-toc-heading-level-3><a class="ez-toc-link ez-toc-heading-20" href=#Structure-2 title=Structure>Structure</a></li><li class='ez-toc-page-1 ez-toc-heading-level-3'><a class="ez-toc-link ez-toc-heading-21" href=#Contraction title=Contraction>Contraction</a></li><li class='ez-toc-page-1 ez-toc-heading-level-3'><a class="ez-toc-link ez-toc-heading-22" href=#Sources_of_cytosolic_calcium title="Sources of cytosolic calcium">Sources of cytosolic calcium</a></li><li class='ez-toc-page-1 ez-toc-heading-level-3'><a class="ez-toc-link ez-toc-heading-23" href=#Membrane_activation title="Membrane activation">Membrane activation</a></li></ul></li><li class='ez-toc-page-1 ez-toc-heading-level-2'><a class="ez-toc-link ez-toc-heading-24" href=#Quiz title=Quiz>Quiz</a><ul class=ez-toc-list-level-3 ><li class=ez-toc-heading-level-3><a class="ez-toc-link ez-toc-heading-25" href=#Send_Your_Results_Optional title="Send Your Results (Optional)">Send Your Results (Optional)</a></li></ul></li></ul></nav></div><p>Muscle cells are specialized to generate force and movement. There are three types of muscle tissue: (1) <em>skeletal muscle</em>, (2) <em>smooth muscle</em>, and (3) <em>cardiac muscle</em>.</p><div class='code-block code-block-8 code-block-block-8' style='margin: 8px 0; clear: both;'> <script language=javascript src="https://live.primis.tech/live/liveView.php?s=106005&cbuster=%%CACHEBUSTER%%"></script> </div><h2><span class=ez-toc-section id=Skeletal_Muscle></span>Skeletal Muscle<span class=ez-toc-section-end></span></h2><p>Attached to the bone and moves them by contraction. The contraction is initiated by neural impulses and is under voluntary control.</p><div class='ai-viewports ai-viewport-1 ai-insert-41-74336005' style='margin: 8px 0; clear: both;' data-insertion-position=prepend data-selector=.ai-insert-41-74336005 data-insertion-no-dbg data-code='PGRpdiBjbGFzcz0nY29kZS1ibG9jayBjb2RlLWJsb2NrLTQxIGNvZGUtYmxvY2stYmxvY2stNDEnIHN0eWxlPSdtYXJnaW46IDhweCAwOyBjbGVhcjogYm90aDsnPgo8IS0tIGFmdGVyIGV2ZXJ5IDQgcGFyYWdyYXBoIGRlc2t0b3AgcG9zdC0tPgo8ZGl2IGlkPSdtaWQ2MTInIGNsYXNzPSdhZHliIHNob3ctYWQtbGFiZWwgbXgtbWQtbjMnIGRhdGEtYWQtbWFwcGluZz0nbWlkJyBkYXRhLWFkLXNsb3Q9J0JPX2FydGljbGUnIGRhdGEtYWQtdGFyZ2V0aW5ncG9zPSAnbWlkJyA+PC9kaXY+PC9kaXY+Cg==' data-block=41></div><h3><span class=ez-toc-section id=Structure></span>Structure<span class=ez-toc-section-end></span></h3><p>A muscle consists of a number of muscle fibers bound together by connective tissue and it is attached to a bone by collagen bundles called tendons. A single muscle fiber is a multinucleated cell and is formed from myoblasts during development. After infancy, new fibers are formed only from undifferentiated satellite cells and any compensation for lost muscles in adulthood occurs mostly through an increase in the size of fibers. Skeletal muscle cells have longitudinal bundles called myofibrils in the cytoplasm. Each myofibril consists of a repeating unit called sarcomere. Each sarcomere has a band of thick filaments in the middle called the A band, and they are flanked on both sides by thin filaments. One end of the thin filaments is anchored to the Z line in the I band which separates adjacent sarcomeres and the other end partially overlap the thick filaments. Due to the banded pattern provided by thin and thick filaments, skeletal muscle is also called striated muscle. The central region of the sarcomere where there is no overlap between thin and thick filaments is called the H zone. A band called the M line, in center of the H zone, link the center regions of thick filaments. Titin protein fibers from the Z line are linked to the M line and the thick filaments. Space between overlapping thick and thin filaments is bridged by projections called cross bridges from the thick filaments. In a cross-section, each thick filament is surrounded by a hexagonal array of six thin filaments and each thin filament is surrounded by a triangular array of three thick filaments. Thick filaments contain the contractile protein, myosin. Thin filaments contain the contractile protein, actin and two other proteins, troponin and tropomyosin.</p><h3><span class=ez-toc-section id=Molecular_mechanisms_of_contraction></span>Molecular mechanisms of contraction<span class=ez-toc-section-end></span></h3><p>Each actin molecule contains a binding site for <a href=https://www.biologyonline.com/dictionary/myosin/ data-wpel-link=internal rel=follow>myosin</a>. Each myosin molecule contains a binding site for actin and one for ATP.</p><div class='ai-viewports ai-viewport-1 ai-insert-41-12298209' style='margin: 8px 0; clear: both;' data-insertion-position=prepend data-selector=.ai-insert-41-12298209 data-insertion-no-dbg data-code='PGRpdiBjbGFzcz0nY29kZS1ibG9jayBjb2RlLWJsb2NrLTQxIGNvZGUtYmxvY2stYmxvY2stNDEnIHN0eWxlPSdtYXJnaW46IDhweCAwOyBjbGVhcjogYm90aDsnPgo8IS0tIGFmdGVyIGV2ZXJ5IDQgcGFyYWdyYXBoIGRlc2t0b3AgcG9zdC0tPgo8ZGl2IGlkPSdtaWQ2MTMnIGNsYXNzPSdhZHliIHNob3ctYWQtbGFiZWwgbXgtbWQtbjMnIGRhdGEtYWQtbWFwcGluZz0nbWlkJyBkYXRhLWFkLXNsb3Q9J0JPX2FydGljbGUnIGRhdGEtYWQtdGFyZ2V0aW5ncG9zPSAnbWlkJyA+PC9kaXY+PC9kaXY+Cg==' data-block=41></div><h3><span class=ez-toc-section id=Sliding-filament_mechanism></span>Sliding-filament mechanism<span class=ez-toc-section-end></span></h3><p>Muscle contraction is produced by cross-bridge cycles. A cycle has 4 steps:</p><ol> <li>Energizing of myosin cross-bridge<br> A + M•ATP —> A + M*•ADP•Pi (ATP is energizer here)</li> <li>Attachment of cross-bridge to a thin filament<br> A + M*•ADP•Pi —> A•M*•ADP•Pi</li> <li>Movement of cross-bridge, producing tension<br> A•M*•ADP•Pi —> A•M + ADP + Pi</li> <li>Detachment of cross-bridge from thin filament<br> A•M + ATP —> A + M•ATP (ATP is modulator here)</li></ol><p>Movement of the cross-bridges make the overlapping thick and thin filaments slide past each other (they do not change in length) to produce a contraction.</p><h3><span class=ez-toc-section id=Roles_of_troponin_tropomyosin_and_calcium></span>Roles of troponin, tropomyosin and calcium<span class=ez-toc-section-end></span></h3><p>In a resting muscle fiber, the protein <a href=https://www.biologyonline.com/dictionary/tropomyosin/ data-wpel-link=internal rel=follow>tropomyosin</a>, held by troponin, covers the myosin-binding sites on actin and prevents cross bridges from making contact with actin. An action potential releases Ca<sup>2+</sup> from the sarcoplasmic reticulum of the cell and increases the cytosolic concentration of Ca<sup>2+</sup> in the muscle fiber. When Ca<sup>2+</sup> binds to troponin, it removes tropomyosin from the myosin-binding site and makes it possible for cross bridges to attach with actin.</p><div class='ai-viewports ai-viewport-1 ai-insert-41-15947065' style='margin: 8px 0; clear: both;' data-insertion-position=prepend data-selector=.ai-insert-41-15947065 data-insertion-no-dbg data-code='PGRpdiBjbGFzcz0nY29kZS1ibG9jayBjb2RlLWJsb2NrLTQxIGNvZGUtYmxvY2stYmxvY2stNDEnIHN0eWxlPSdtYXJnaW46IDhweCAwOyBjbGVhcjogYm90aDsnPgo8IS0tIGFmdGVyIGV2ZXJ5IDQgcGFyYWdyYXBoIGRlc2t0b3AgcG9zdC0tPgo8ZGl2IGlkPSdtaWQ2MTQnIGNsYXNzPSdhZHliIHNob3ctYWQtbGFiZWwgbXgtbWQtbjMnIGRhdGEtYWQtbWFwcGluZz0nbWlkJyBkYXRhLWFkLXNsb3Q9J0JPX2FydGljbGUnIGRhdGEtYWQtdGFyZ2V0aW5ncG9zPSAnbWlkJyA+PC9kaXY+PC9kaXY+Cg==' data-block=41></div><h3><span class=ez-toc-section id=Sarcoplasmic_reticulum></span>Sarcoplasmic reticulum<span class=ez-toc-section-end></span></h3><p>The <a href=https://www.biologyonline.com/dictionary/sarcoplasmic-reticulum/ data-wpel-link=internal rel=follow>sarcoplasmic reticulum</a> is homologous to <em>endoplasmic reticulum</em>. Forms a sleeve around each myofibril. It has enlarged regions called lateral sacs that store Ca<sup>2+</sup>, and a tubular structure called transverse (T) tubule. Depolarization of the plasma membrane leads to depolarization of the T-tubule and opens the Ca<sup>2+ </sup>channels of the lateral sacs. To end contraction, Ca<sup>2+</sup> is pumped back into the lateral sacs by active transport proteins called Ca<sup>2+</sup>-ATP-ases.</p><h3><span class=ez-toc-section id=Neuromuscular_junction></span>Neuromuscular junction<span class=ez-toc-section-end></span></h3><p>Nerve cells that innervate skeletal muscle fibers are called motor (somatic efferent) neurons. These neurons have cell bodies in the CNS and large, myelinated axons that can propagate action potentials at high velocities. At a muscle, the axon gives branches to many muscle fibers but each fiber gets only one branch. A motor neuron and the fibers it innervates are together called a motor unit. The region of muscle-fiber plasma membrane lying directly under the axon is called the motor endplate and the junction is called the neuromuscular junction. Axon terminals of a motor neuron contain vesicles filled with acetylcholine (ACh). An action potential opens Ca<sup>2+</sup> channels in the nerve plasma membrane and Ca<sup>2+</sup> diffuses in to enable these vesicles to fuse with the plasma membrane and release ACh in the extracellular space. ACh opens ion channels in the motor endplate and produces a depolarization called endplate potential (EPP). The enzyme acetylcholinesterase on the motor endplate breaks down ACh to close the ion channels and return the plate to resting potential.</p><div class='ai-viewports ai-viewport-1 ai-insert-41-12999362' style='margin: 8px 0; clear: both;' data-insertion-position=prepend data-selector=.ai-insert-41-12999362 data-insertion-no-dbg data-code='PGRpdiBjbGFzcz0nY29kZS1ibG9jayBjb2RlLWJsb2NrLTQxIGNvZGUtYmxvY2stYmxvY2stNDEnIHN0eWxlPSdtYXJnaW46IDhweCAwOyBjbGVhcjogYm90aDsnPgo8IS0tIGFmdGVyIGV2ZXJ5IDQgcGFyYWdyYXBoIGRlc2t0b3AgcG9zdC0tPgo8ZGl2IGlkPSdtaWQ2MTUnIGNsYXNzPSdhZHliIHNob3ctYWQtbGFiZWwgbXgtbWQtbjMnIGRhdGEtYWQtbWFwcGluZz0nbWlkJyBkYXRhLWFkLXNsb3Q9J0JPX2FydGljbGUnIGRhdGEtYWQtdGFyZ2V0aW5ncG9zPSAnbWlkJyA+PC9kaXY+PC9kaXY+Cg==' data-block=41></div><h3><span class=ez-toc-section id=Mechanics_of_a_single_fiber_contraction></span>Mechanics of a single fiber contraction<span class=ez-toc-section-end></span></h3><p>Force exerted on an object by a contracting muscle is called muscle tension, and the force exerted on the muscle by the object (weight) is called load. These are opposing forces and whether exertion of force leads to fiber shortening depends on relative magnitudes of tension and load. For fibers to shorten and move a load, tension must be greater than load. When a muscle develops tension but does not shorten, the contraction is called isometric (constant length). Seen when muscle supports a load in constant position or attempts to move a supported load that is greater than the tension. During such a contraction, the bound cross-bridges do not move. A contraction in which the muscle shortens while load remains constant is called isotonic (constant tension). In such a contraction, the cross-bridges bound to actin move, shortening the fibers. Before an isotonic shortening, there is a period of isometric contraction during which tension increases. A lengthening (eccentric) contraction occurs when a load on a muscle is greater than the tension and the load lengthens the muscle. Not an active process but a result of external force on the muscle. In a lengthening contraction, cross bridges are pulled toward the Z line.</p><h3><span class=ez-toc-section id=Twitch_contraction></span>Twitch contraction<span class=ez-toc-section-end></span></h3><p>Mechanical response of a single muscle fiber to a single action potential. Following the action potential, there is a latent period, before the tension in muscle fiber increases. Time interval from the beginning of tension development and peak tension is called contraction time. The latent period is longer in an isotonic twitch than in an isometric twitch while the duration of contraction is longer in an isometric twitch than in an isotonic twitch. The latent period is positively correlated while the velocity of shortening, duration of twitch and distance shortened are negatively correlated with load.</p><div class='ai-viewports ai-viewport-1 ai-insert-41-44889882' style='margin: 8px 0; clear: both;' data-insertion-position=prepend data-selector=.ai-insert-41-44889882 data-insertion-no-dbg data-code='PGRpdiBjbGFzcz0nY29kZS1ibG9jayBjb2RlLWJsb2NrLTQxIGNvZGUtYmxvY2stYmxvY2stNDEnIHN0eWxlPSdtYXJnaW46IDhweCAwOyBjbGVhcjogYm90aDsnPgo8IS0tIGFmdGVyIGV2ZXJ5IDQgcGFyYWdyYXBoIGRlc2t0b3AgcG9zdC0tPgo8ZGl2IGlkPSdtaWQ2MTYnIGNsYXNzPSdhZHliIHNob3ctYWQtbGFiZWwgbXgtbWQtbjMnIGRhdGEtYWQtbWFwcGluZz0nbWlkJyBkYXRhLWFkLXNsb3Q9J0JPX2FydGljbGUnIGRhdGEtYWQtdGFyZ2V0aW5ncG9zPSAnbWlkJyA+PC9kaXY+PC9kaXY+Cg==' data-block=41></div><h3><span class=ez-toc-section id=Frequency-tension_relation></span>Frequency-tension relation<span class=ez-toc-section-end></span></h3><p>An increase in muscle tension from successive action potentials is called summation and a maintained contraction in response to repetitive stimulation is called tetanus. If a tetanus oscillates, it is called unfused tetanus while a tetanus without oscillations is called fused tetanus.</p><h3><span class=ez-toc-section id=Length-tension_relation></span>Length-tension relation<span class=ez-toc-section-end></span></h3><p>Passive tension in a relaxed fiber increases with increased stretch due to the elongation of titin filaments. The magnitude of active tension during contraction depends on the passive tension of a fiber because the potential for cross-bridge formation is different at different overlaps between actin and myosin filaments. The length at which the fiber develops the greatest active tension is called optimal length, l0. In a relaxed state, the length of most fibers is near l0.</p><div class='ai-viewports ai-viewport-1 ai-insert-41-49165003' style='margin: 8px 0; clear: both;' data-insertion-position=prepend data-selector=.ai-insert-41-49165003 data-insertion-no-dbg data-code='PGRpdiBjbGFzcz0nY29kZS1ibG9jayBjb2RlLWJsb2NrLTQxIGNvZGUtYmxvY2stYmxvY2stNDEnIHN0eWxlPSdtYXJnaW46IDhweCAwOyBjbGVhcjogYm90aDsnPgo8IS0tIGFmdGVyIGV2ZXJ5IDQgcGFyYWdyYXBoIGRlc2t0b3AgcG9zdC0tPgo8ZGl2IGlkPSdtaWQ2MTcnIGNsYXNzPSdhZHliIHNob3ctYWQtbGFiZWwgbXgtbWQtbjMnIGRhdGEtYWQtbWFwcGluZz0nbWlkJyBkYXRhLWFkLXNsb3Q9J0JPX2FydGljbGUnIGRhdGEtYWQtdGFyZ2V0aW5ncG9zPSAnbWlkJyA+PC9kaXY+PC9kaXY+Cg==' data-block=41></div><h3><span class=ez-toc-section id=Skeletal_muscle_energy_metabolism></span>Skeletal muscle energy metabolism<span class=ez-toc-section-end></span></h3><p>Muscle glycogen is the major fuel at the initial stages of exercise and is followed by the utilization of blood glucose and fatty acids.</p><p>A muscle fiber generates ATP by:</p><ul> <li><em>Phosphorylation of ADP by creatine phosphate.</em> Source of ATP during the initial phase of contraction. Followed by the slower pathways of:</li> <li><em>Oxidative phosphorylation in mitochondria.</em></li> <li><em>Glycolysis in cytosol.</em> This pathway provides a majority of the energy and lactic acid is produced.<br> At the end of muscle activity, creatine phosphate and glycogen levels are restored by energy-dependent processes, leading to a continued elevated level of oxygen consumption, called oxygen debt, even after exercise is over.</li></ul><h3><span class=ez-toc-section id=Muscle_fatigue></span>Muscle fatigue<span class=ez-toc-section-end></span></h3><p>Decline in muscle tension as a result of previous contractile activity. A fatigued muscle also has decreased shortening velocity and slower rate of relaxation. Onset and rate of fatigue depend on the type of skeletal muscle and duration of contractile activity. If a fatigued muscle is allowed to rest, it recovers. The rate of recovery depends on the duration and intensity of the previous exercise. Fatigue is not due to low ATP since a fatigued muscle still has quite a high concentration of ATP. Fatigue may be an adaptation to prevent rigor that will result from very low ATP level. High-frequency fatigue accompanying high intensity, short-duration exercise is due to failure in the conduction of action potential in the T tubule. Recovery from such fatigue is rapid. Low-frequency fatigue seen with low intensity, long-duration exercise is due to the build-up of lactic acid, which changes the conformation of muscle proteins. Recovery from such fatigue is slow.</p><div class='ai-viewports ai-viewport-1 ai-insert-41-31548758' style='margin: 8px 0; clear: both;' data-insertion-position=prepend data-selector=.ai-insert-41-31548758 data-insertion-no-dbg data-code='PGRpdiBjbGFzcz0nY29kZS1ibG9jayBjb2RlLWJsb2NrLTQxIGNvZGUtYmxvY2stYmxvY2stNDEnIHN0eWxlPSdtYXJnaW46IDhweCAwOyBjbGVhcjogYm90aDsnPgo8IS0tIGFmdGVyIGV2ZXJ5IDQgcGFyYWdyYXBoIGRlc2t0b3AgcG9zdC0tPgo8ZGl2IGlkPSdtaWQ2MTgnIGNsYXNzPSdhZHliIHNob3ctYWQtbGFiZWwgbXgtbWQtbjMnIGRhdGEtYWQtbWFwcGluZz0nbWlkJyBkYXRhLWFkLXNsb3Q9J0JPX2FydGljbGUnIGRhdGEtYWQtdGFyZ2V0aW5ncG9zPSAnbWlkJyA+PC9kaXY+PC9kaXY+Cg==' data-block=41></div><h3><span class=ez-toc-section id=Types_of_skeletal_muscle_fibers></span>Types of skeletal muscle fibers<span class=ez-toc-section-end></span></h3><ol> <li><em>Fast fibers</em> contain myosin with high ATPase activity and therefore have high shortening velocity. Fatigue rapidly.</li> <li><em>Slow fibers</em> contain myosin with low ATPase activity and therefore have low shortening velocity. Fatigue slowly.</li> <li><em>Oxidative fibers</em> have numerous mitochondria and therefore a high capacity of oxidative phosphorylation. ATP production is dependent on bloodborne oxygen and fuel.</li></ol><p>It also contains an oxygen-binding protein called myoglobin, which increases the rate of oxygen diffusion into the fiber. This protein gives the fibers a red color and therefore these fibers are also called red muscle fibers. Glycolytic fibers have few mitochondria but a high concentration of glycolytic enzymes and glycogen. Have a pale color and therefore are also called white muscle fibers. Larger and have more thick and thin filaments and therefore can develop more tension. Fatigue rapidly.</p><p>Three types of skeletal fibers: (l) Slow-oxidative fibers, (2) Fast-oxidative fibers and (3) Fast-glycolytic fibers. Most muscles contain all three kinds of fibers.</p><h3><span class=ez-toc-section id=Control_of_muscle_tension></span>Control of muscle tension<span class=ez-toc-section-end></span></h3><p>Total tension a muscle can develop depends on:</p><ol> <li>Amount of tension developed by each fiber</li> <li>Number of fibers contracting at any time, which in turn depends on: (a) Number of fibers in each motor unit (motor unit size) and (b) Number of active motor units. Increasing the number of active motor units in a muscle, called recruitment, is achieved by increasing excitatory input to the motor neurons. Finer control of muscle tension is possible in muscles with small motor unit size.</li></ol><div class='ai-viewports ai-viewport-1 ai-insert-41-14091887' style='margin: 8px 0; clear: both;' data-insertion-position=prepend data-selector=.ai-insert-41-14091887 data-insertion-no-dbg data-code='PGRpdiBjbGFzcz0nY29kZS1ibG9jayBjb2RlLWJsb2NrLTQxIGNvZGUtYmxvY2stYmxvY2stNDEnIHN0eWxlPSdtYXJnaW46IDhweCAwOyBjbGVhcjogYm90aDsnPgo8IS0tIGFmdGVyIGV2ZXJ5IDQgcGFyYWdyYXBoIGRlc2t0b3AgcG9zdC0tPgo8ZGl2IGlkPSdtaWQ2MTknIGNsYXNzPSdhZHliIHNob3ctYWQtbGFiZWwgbXgtbWQtbjMnIGRhdGEtYWQtbWFwcGluZz0nbWlkJyBkYXRhLWFkLXNsb3Q9J0JPX2FydGljbGUnIGRhdGEtYWQtdGFyZ2V0aW5ncG9zPSAnbWlkJyA+PC9kaXY+PC9kaXY+Cg==' data-block=41></div><h3><span class=ez-toc-section id=Muscle_adaptation_to_exercise></span>Muscle adaptation to exercise<span class=ez-toc-section-end></span></h3><p>Increased amount of contractile activity (exercise) increases size (hypertrophy) of muscle fibers and capacity for ATP production. <em>Low-intensity exercise</em> affects oxidative fibers, increasing the number of mitochondria and capillaries. <em>High-intensity exercise</em> affects glycolytic fibers, increasing their diameter by an increased synthesis of actin and myosin filaments, and an increased synthesis of glycolytic enzymes.</p><h3><span class=ez-toc-section id=Lever_action_of_muscles_and_bones></span>Lever action of muscles and bones<span class=ez-toc-section-end></span></h3><p>A contracting muscle exerts a pulling force and moves the bones. If this leads to a limb bending at a joint, it is called flexion while if it leads to a limb straightening from a joint, it is called extension. Groups of muscles producing opposite movements at a joint are called antagonists.</p><div class='ai-viewports ai-viewport-1 ai-insert-41-25354184' style='margin: 8px 0; clear: both;' data-insertion-position=prepend data-selector=.ai-insert-41-25354184 data-insertion-no-dbg data-code='PGRpdiBjbGFzcz0nY29kZS1ibG9jayBjb2RlLWJsb2NrLTQxIGNvZGUtYmxvY2stYmxvY2stNDEnIHN0eWxlPSdtYXJnaW46IDhweCAwOyBjbGVhcjogYm90aDsnPgo8IS0tIGFmdGVyIGV2ZXJ5IDQgcGFyYWdyYXBoIGRlc2t0b3AgcG9zdC0tPgo8ZGl2IGlkPSdtaWQ2MjAnIGNsYXNzPSdhZHliIHNob3ctYWQtbGFiZWwgbXgtbWQtbjMnIGRhdGEtYWQtbWFwcGluZz0nbWlkJyBkYXRhLWFkLXNsb3Q9J0JPX2FydGljbGUnIGRhdGEtYWQtdGFyZ2V0aW5ncG9zPSAnbWlkJyA+PC9kaXY+PC9kaXY+Cg==' data-block=41></div><h3><span class=ez-toc-section id=Skeletal_muscle_disease></span>Skeletal muscle disease<span class=ez-toc-section-end></span></h3><p>Poliomyelitis leads to paralysis of skeletal muscles due to destruction of motor neurons by a virus. Muscular dystrophy is a progressive degeneration of skeletal and cardiac muscle fibers due to a low amount of the protein dystrophin that maintains the structural integrity of the plasma membrane during stretching. It is a sex-linked recessive disease. Myastheniagravis leads to muscle fatigue due to low number of ACh receptors on the motor endplate.</p><h2><span class=ez-toc-section id=Smooth_Muscle></span>Smooth Muscle<span class=ez-toc-section-end></span></h2><p>A smooth muscle lacks the banded structure of skeletal muscles. Controlled by the autonomic nervous system and is not under voluntary control.</p><div class='ai-viewports ai-viewport-1 ai-insert-41-83227409' style='margin: 8px 0; clear: both;' data-insertion-position=prepend data-selector=.ai-insert-41-83227409 data-insertion-no-dbg data-code='PGRpdiBjbGFzcz0nY29kZS1ibG9jayBjb2RlLWJsb2NrLTQxIGNvZGUtYmxvY2stYmxvY2stNDEnIHN0eWxlPSdtYXJnaW46IDhweCAwOyBjbGVhcjogYm90aDsnPgo8IS0tIGFmdGVyIGV2ZXJ5IDQgcGFyYWdyYXBoIGRlc2t0b3AgcG9zdC0tPgo8ZGl2IGlkPSdtaWQ2MjEnIGNsYXNzPSdhZHliIHNob3ctYWQtbGFiZWwgbXgtbWQtbjMnIGRhdGEtYWQtbWFwcGluZz0nbWlkJyBkYXRhLWFkLXNsb3Q9J0JPX2FydGljbGUnIGRhdGEtYWQtdGFyZ2V0aW5ncG9zPSAnbWlkJyA+PC9kaXY+PC9kaXY+Cg==' data-block=41></div><h3><span class=ez-toc-section id=Structure-2></span>Structure<span class=ez-toc-section-end></span></h3><p>A smooth muscle fiber is a spindle-shaped, uninucleate cell. Has the capacity to divide throughout life. The thick and thin filaments are not regularly aligned into sarcomeres but are oriented diagonally to the long axis of the cell. Thin filaments are attached to the plasma membrane or to cytoplasmic structures called dense bodies. The concentration of myosin is low and that of actin is high.</p><h3><span class=ez-toc-section id=Contraction></span>Contraction<span class=ez-toc-section-end></span></h3><p>Thin filaments do not have troponin. As cytosolic Ca<sup>2+</sup> concentration increases,</p><ol> <li>Calcium binds to calmodulin, a protein</li> <li>Calcium-calmodulin complex binds to the enzyme, myosin light-chain kinase and activates it</li> <li>The active kinase uses ATP to phosphorylate the myosin cross-bridges</li> <li>The phosphorylated cross-bridges bind to actin</li> <li>Myosin is dephosphorylated by myosin light-chain phosphatase</li></ol><p>ATPase activity is low and therefore cross-bridge cycling and muscle shortening is slow. Dissociation of dephosphorylated cross-bridges is slow and therefore the muscle can maintain tension with very low ATP consumption. Does not undergo fatigue during prolonged periods of activity. The range of muscle lengths over which tension can be developed is greater.</p><div class='ai-viewports ai-viewport-1 ai-insert-41-42796975' style='margin: 8px 0; clear: both;' data-insertion-position=prepend data-selector=.ai-insert-41-42796975 data-insertion-no-dbg data-code='PGRpdiBjbGFzcz0nY29kZS1ibG9jayBjb2RlLWJsb2NrLTQxIGNvZGUtYmxvY2stYmxvY2stNDEnIHN0eWxlPSdtYXJnaW46IDhweCAwOyBjbGVhcjogYm90aDsnPgo8IS0tIGFmdGVyIGV2ZXJ5IDQgcGFyYWdyYXBoIGRlc2t0b3AgcG9zdC0tPgo8ZGl2IGlkPSdtaWQ2MjInIGNsYXNzPSdhZHliIHNob3ctYWQtbGFiZWwgbXgtbWQtbjMnIGRhdGEtYWQtbWFwcGluZz0nbWlkJyBkYXRhLWFkLXNsb3Q9J0JPX2FydGljbGUnIGRhdGEtYWQtdGFyZ2V0aW5ncG9zPSAnbWlkJyA+PC9kaXY+PC9kaXY+Cg==' data-block=41></div><h3><span class=ez-toc-section id=Sources_of_cytosolic_calcium></span>Sources of cytosolic calcium<span class=ez-toc-section-end></span></h3><ul> <li>Sarcoplasmic reticulum – Does not have T-tubules.</li> <li>Extracellular calcium – Opening of Ca<sup>2+</sup> channels in the plasma membrane.</li></ul><p>Rate of Ca<sup>2+</sup> removal is slower with the result that a twitch lasts longer. Concentration of Ca<sup>2+</sup> in response to a single action potential is only sufficient enough to activate a proportion of cross-bridges. Therefore, tension in smooth muscle can be graded by varying cytosolic Ca<sup>2+</sup> concentration. A low-level tension is always maintained, even in the absence of external stimuli, and this is called a smooth-muscle tone.</p><h3><span class=ez-toc-section id=Membrane_activation></span>Membrane activation<span class=ez-toc-section-end></span></h3><p>The plasma membrane of smooth muscles receives both excitatory and inhibitory inputs and the contractile state of the muscle depends on the relative intensity of both. Some smooth muscle fibers generate action potentials spontaneously in the absence of any input. The potential change occurring during such spontaneous depolarization is called pacemaker potential. Smooth muscles do not have motor endplates. The postganglionic autonomic neuron divides into branches in the smooth muscle fibers, and each branch has a series of swollen regions called varicosities, which contain vesicles filled with neurotransmitter that is released when an action potential arrives. The same neurotransmitter can produce excitation in one fiber and inhibition in the other. Varicosities from a single axon may innervate several fibers and a single fiber may receive varicosities from both sympathetic and parasympathetic neurons.</p><p>Smooth muscle plasma membrane also binds hormones and respond to them. Paracrine agents, acidity, oxygen concentration, osmolarity, ion composition can also influence smooth muscle tension, providing a response mechanism to local factors.</p><p><strong>(1) Single-unit Smooth Muscle</strong></p><p>All the fibers undergo synchronous activity because adjacent fibers arc linked by gap junction d action potential occurring anywhere is propagated to all other cells. The whole muscle responds to stimulation as a single unit. Some of the fibers may be pacemaker cells and they can control the contraction of the entire muscle a majority of which may consist of non-pacemaker cells. The nerve terminals, therefore, need to be restricted only to the pacemaker cells.</p><p><strong>(2) Multi-unit Smooth Muscle</strong></p><p>Each fiber responds independently of its neighboring fibers because there are no connecting gap junctions.</p><p> </p><div style="text-align: center; vertical-align: top; padding: 10px; border: 1px solid #e2e2e2;"><iframe src=https://www.youtube.com/embed/VVL-8zr2hk4 width=560 height=315 frameborder=0 allowfullscreen=allowfullscreen></iframe></div><div style="vertical-align: top; padding: 10px; border: 1px solid #e2e2e2;">Credit: <a href=https://www.youtube.com/channel/UCsooa4yRKGN_zEE8iknghZA data-wpel-link=external target=_blank rel="nofollow external noopener noreferrer">TED-Ed</a></div> <script>if(window.qmn_quiz_data===undefined){window.qmn_quiz_data=new Object();}</script><script>window.qmn_quiz_data["83"]={"quiz_id":"83","quiz_name":"Muscle - Quiz","disable_answer":"0","ajax_show_correct":"0","progress_bar":"1","contact_info_location":"1","qpages":{"1":{"id":"1","quizID":"83","pagekey":"5fa0f85225db8","hide_prevbtn":"0"}},"skip_validation_time_expire":"1","timer_limit_val":0,"disable_scroll_next_previous_click":"0","disable_first_page":0,"enable_result_after_timer_end":"0","enable_quick_result_mc":"0","end_quiz_if_wrong":0,"form_disable_autofill":0,"disable_mathjax":0,"enable_quick_correct_answer_info":0,"quick_result_correct_answer_text":"Correct! You have selected correct answer.","quick_result_wrong_answer_text":"Wrong! You have selected wrong answer.","quiz_processing_message":"","quiz_limit_choice":"Limit of choice is reached.","not_allow_after_expired_time":0,"scheduled_time_end":false,"error_messages":{"email_error_text":"Not a valid e-mail address!","number_error_text":"This field must be a number!","incorrect_error_text":"The entered text is not correct!","empty_error_text":"Please complete all required fields!","url_error_text":"The entered URL is not valid!","minlength_error_text":"Required atleast %minlength% characters.","maxlength_error_text":"Maximum %maxlength% characters allowed.","recaptcha_error_text":"ReCaptcha is missing"},"first_page":false}</script><style id=qmn_quiz_template-css>.qsm-pagination{border:none !important}</style><div class='qsm-quiz-container qsm-quiz-container-83 qmn_quiz_container mlw_qmn_quiz quiz_theme_default '><form name=quizForm83 id=quizForm83 action=/tutorials/muscle method=POST class="qsm-quiz-form qmn_quiz_form mlw_quiz_form" novalidate enctype=multipart/form-data> <input type=hidden name=qsm_hidden_questions id=qsm_hidden_questions value> <input type=hidden name=qsm_nonce id=qsm_nonce_83 value=72926d0365> <input type=hidden name=qsm_unique_key id=qsm_unique_key_83 value=674fba6e49568><div id=mlw_error_message class="qsm-error-message qmn_error_message_section"></div> <span id=mlw_top_of_quiz></span><section class="qsm-page "><div class="quiz_section quiz_begin"><div class='qsm-before-message mlw_qmn_message_before'><h2><span class=ez-toc-section id=Quiz></span>Quiz<span class=ez-toc-section-end></span></h2><p><strong>Select the best answer.</strong></p></div></div><div class="quiz_section qsm-question-wrapper question-type-0 question-section-id-422 " data-qid=422><div class=mlw_qmn_new_question>1. The type of skeletal muscle fiber with high ATPase activity and fatigues rapidly</div><div class='mlw_qmn_question qsm_remove_bold' ><p></p></div><div class='qmn_radio_answers mlwRequiredRadio'><div class="qmn_mc_answer_wrap mrq_checkbox_class" id="question422-1 "> <input type=radio class='qmn_quiz_radio qmn-multiple-choice-input' name=question422 id=question422_1 value=0> <label class=qsm-input-label for=question422_1> Fast fibers </label></div><div class="qmn_mc_answer_wrap mrq_checkbox_class" id="question422-2 "> <input type=radio class='qmn_quiz_radio qmn-multiple-choice-input' name=question422 id=question422_2 value=1> <label class=qsm-input-label for=question422_2> Slow fibers </label></div><div class="qmn_mc_answer_wrap mrq_checkbox_class" id="question422-3 "> <input type=radio class='qmn_quiz_radio qmn-multiple-choice-input' name=question422 id=question422_3 value=2> <label class=qsm-input-label for=question422_3> Oxidative fibers </label></div><div class="qmn_mc_answer_wrap mrq_checkbox_class" id="question422-4 "> <input type=radio class='qmn_quiz_radio qmn-multiple-choice-input' name=question422 id=question422_4 value=3> <label class=qsm-input-label for=question422_4> Fast fibers and slow fibers </label></div> <input type=radio style="display: none;" name=question422 id=question422_none checked=checked value></div></div><div class="quiz_section qsm-question-wrapper question-type-0 question-section-id-423 " data-qid=423><div class=mlw_qmn_new_question>2. How does low-intensity exercise affect skeletal muscles?</div><div class='mlw_qmn_question qsm_remove_bold' ><p></p></div><div class='qmn_radio_answers mlwRequiredRadio'><div class="qmn_mc_answer_wrap mrq_checkbox_class" id="question423-1 "> <input type=radio class='qmn_quiz_radio qmn-multiple-choice-input' name=question423 id=question423_1 value=0> <label class=qsm-input-label for=question423_1> It increases the diameter of glycolytic fibers. </label></div><div class="qmn_mc_answer_wrap mrq_checkbox_class" id="question423-2 "> <input type=radio class='qmn_quiz_radio qmn-multiple-choice-input' name=question423 id=question423_2 value=1> <label class=qsm-input-label for=question423_2> It increases the number of capillaries and the mitochondria in muscle cells. </label></div><div class="qmn_mc_answer_wrap mrq_checkbox_class" id="question423-3 "> <input type=radio class='qmn_quiz_radio qmn-multiple-choice-input' name=question423 id=question423_3 value=2> <label class=qsm-input-label for=question423_3> It decreases the number of glycolytic fibers. </label></div><div class="qmn_mc_answer_wrap mrq_checkbox_class" id="question423-4 "> <input type=radio class='qmn_quiz_radio qmn-multiple-choice-input' name=question423 id=question423_4 value=3> <label class=qsm-input-label for=question423_4> It decreases the number of capillaries and the mitochondria in muscle cells. </label></div> <input type=radio style="display: none;" name=question423 id=question423_none checked=checked value></div></div><div class="quiz_section qsm-question-wrapper question-type-0 question-section-id-424 " data-qid=424><div class=mlw_qmn_new_question>3. Characterized by the lack the banded structure</div><div class='mlw_qmn_question qsm_remove_bold' ><p></p></div><div class='qmn_radio_answers mlwRequiredRadio'><div class="qmn_mc_answer_wrap mrq_checkbox_class" id="question424-1 "> <input type=radio class='qmn_quiz_radio qmn-multiple-choice-input' name=question424 id=question424_1 value=0> <label class=qsm-input-label for=question424_1> Skeletal muscle </label></div><div class="qmn_mc_answer_wrap mrq_checkbox_class" id="question424-2 "> <input type=radio class='qmn_quiz_radio qmn-multiple-choice-input' name=question424 id=question424_2 value=1> <label class=qsm-input-label for=question424_2> Smooth muscle </label></div><div class="qmn_mc_answer_wrap mrq_checkbox_class" id="question424-3 "> <input type=radio class='qmn_quiz_radio qmn-multiple-choice-input' name=question424 id=question424_3 value=2> <label class=qsm-input-label for=question424_3> Cardiac muscle </label></div><div class="qmn_mc_answer_wrap mrq_checkbox_class" id="question424-4 "> <input type=radio class='qmn_quiz_radio qmn-multiple-choice-input' name=question424 id=question424_4 value=3> <label class=qsm-input-label for=question424_4> Smooth muscle and cardiac muscle </label></div> <input type=radio style="display: none;" name=question424 id=question424_none checked=checked value></div></div><div class="quiz_section qsm-question-wrapper question-type-0 question-section-id-425 " data-qid=425><div class=mlw_qmn_new_question>4. Which of these muscles is under voluntary control?</div><div class='mlw_qmn_question qsm_remove_bold' ><p></p></div><div class='qmn_radio_answers mlwRequiredRadio'><div class="qmn_mc_answer_wrap mrq_checkbox_class" id="question425-1 "> <input type=radio class='qmn_quiz_radio qmn-multiple-choice-input' name=question425 id=question425_1 value=0> <label class=qsm-input-label for=question425_1> Skeletal muscle </label></div><div class="qmn_mc_answer_wrap mrq_checkbox_class" id="question425-2 "> <input type=radio class='qmn_quiz_radio qmn-multiple-choice-input' name=question425 id=question425_2 value=1> <label class=qsm-input-label for=question425_2> Smooth muscle </label></div><div class="qmn_mc_answer_wrap mrq_checkbox_class" id="question425-3 "> <input type=radio class='qmn_quiz_radio qmn-multiple-choice-input' name=question425 id=question425_3 value=2> <label class=qsm-input-label for=question425_3> Cardiac muscle </label></div><div class="qmn_mc_answer_wrap mrq_checkbox_class" id="question425-4 "> <input type=radio class='qmn_quiz_radio qmn-multiple-choice-input' name=question425 id=question425_4 value=3> <label class=qsm-input-label for=question425_4> All of the above </label></div> <input type=radio style="display: none;" name=question425 id=question425_none checked=checked value></div></div><div class="quiz_section qsm-question-wrapper question-type-0 question-section-id-426 " data-qid=426><div class=mlw_qmn_new_question>5. Which of the following is NOT true about the sarcomere?</div><div class='mlw_qmn_question qsm_remove_bold' ><p></p></div><div class='qmn_radio_answers mlwRequiredRadio'><div class="qmn_mc_answer_wrap mrq_checkbox_class" id="question426-1 "> <input type=radio class='qmn_quiz_radio qmn-multiple-choice-input' name=question426 id=question426_1 value=0> <label class=qsm-input-label for=question426_1> The thick filaments in the middle are called A bands </label></div><div class="qmn_mc_answer_wrap mrq_checkbox_class" id="question426-2 "> <input type=radio class='qmn_quiz_radio qmn-multiple-choice-input' name=question426 id=question426_2 value=1> <label class=qsm-input-label for=question426_2> A bands are flanked on both sides by thin filaments. </label></div><div class="qmn_mc_answer_wrap mrq_checkbox_class" id="question426-3 "> <input type=radio class='qmn_quiz_radio qmn-multiple-choice-input' name=question426 id=question426_3 value=2> <label class=qsm-input-label for=question426_3> The central region where there is no overlap between thin and thick filaments is called H zone </label></div><div class="qmn_mc_answer_wrap mrq_checkbox_class" id="question426-4 "> <input type=radio class='qmn_quiz_radio qmn-multiple-choice-input' name=question426 id=question426_4 value=3> <label class=qsm-input-label for=question426_4> Sarcomere is the fundamental unit of contraction in all types of muscles. </label></div> <input type=radio style="display: none;" name=question426 id=question426_none checked=checked value></div></div><div class=quiz_section><div class='qsm-after-message mlw_qmn_message_end'><h3><span class=ez-toc-section id=Send_Your_Results_Optional></span>Send Your Results (Optional)<span class=ez-toc-section-end></span></h3></div><div class="qsm_contact_div qsm-contact-type-text"> <span class='mlw_qmn_question qsm_question'>Your Name</span> <input type=text class name=contact_field_0 id=contact_field_0 value placeholder='Your Name'></div><div class="qsm_contact_div qsm-contact-type-email"> <span class='mlw_qmn_question qsm_question'>To Email</span> <input type=email class='mlwEmail ' name=contact_field_1 id=contact_field_1 value placeholder='To Email'></div></div></section> <input type=hidden name=qmn_question_list value=422Q423Q424Q425Q426Q><div id=mlw_error_message_bottom class="qsm-error-message qmn_error_message_section"></div> <input type=hidden name=qmn_all_questions_count id=qmn_all_questions_count value=5> <input type=hidden name=total_questions id=total_questions value=5> <input type=hidden name=timer id=timer value=0> <input type=hidden name=timer_ms id=timer_ms value=0> <input type=hidden class=qmn_quiz_id name=qmn_quiz_id id=qmn_quiz_id value=83> <input type=hidden name=complete_quiz value=confirmation></form></div><div style="display: none;" class="qsm-popup qsm-popup-slide" id=modal-4 aria-hidden=false><div class=qsm-popup__overlay tabindex=-1 data-micromodal-close><div class="qsm-popup__container qmn_quiz_container" role=dialog aria-modal=true><div class=qsm-popup__content><img class=lazy decoding=async src="data:image/svg+xml,%3Csvg%20xmlns='http://www.w3.org/2000/svg'%20viewBox='0%200%201%201'%3E%3C/svg%3E" data-src=https://www.biologyonline.com/wp-content/plugins/quiz-master-next/assets/clock.png alt=clock.png><p class=qsm-time-up-text> Time's up</p></div><footer class=qsm-popup__footer><button class="qsm-popup-secondary-button qmn_btn" data-micromodal-close aria-label="Close this dialog window" onclick=location.reload();>Cancel</button></footer></div></div></div><div class='heateor_sss_sharing_container heateor_sss_vertical_sharing' style='width:44px;right: -10px;top: 200px;-webkit-box-shadow:none;box-shadow:none;' data-heateor-sss-href=https://www.biologyonline.com/tutorials/muscle><div class=heateor_sss_sharing_ul><a aria-label=Facebook class=heateor_sss_facebook href="https://www.facebook.com/sharer/sharer.php?u=https%3A%2F%2Fwww.biologyonline.com%2Ftutorials%2Fmuscle" title=Facebook rel="nofollow noopener" target=_blank style=font-size:32px!important;box-shadow:none;display:inline-block;vertical-align:middle data-wpel-link=internal><span class=heateor_sss_svg style="background-color:#0765FE;width:40px;height:40px;margin:0;display:inline-block;opacity:1;float:left;font-size:32px;box-shadow:none;display:inline-block;font-size:16px;padding:0 4px;vertical-align:middle;background-repeat:repeat;overflow:hidden;padding:0;cursor:pointer;box-sizing:content-box"><svg style=display:block; 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2|CINK SLIM|CINK SLIM 2|CINK +|CINK KING|CINK PEAX|CINK SLIM|SUBLIM",iMobile:"i-mobile (IQ|i-STYLE|idea|ZAA|Hitz)",SimValley:"\\b(SP-80|XT-930|SX-340|XT-930|SX-310|SP-360|SP60|SPT-800|SP-120|SPT-800|SP-140|SPX-5|SPX-8|SP-100|SPX-8|SPX-12)\\b",Wolfgang:"AT-B24D|AT-AS50HD|AT-AS40W|AT-AS55HD|AT-AS45q2|AT-B26D|AT-AS50Q",Alcatel:"Alcatel",Nintendo:"Nintendo (3DS|Switch)",Amoi:"Amoi",INQ:"INQ",OnePlus:"ONEPLUS",GenericPhone:"Tapatalk|PDA;|SAGEM|\\bmmp\\b|pocket|\\bpsp\\b|symbian|Smartphone|smartfon|treo|up.browser|up.link|vodafone|\\bwap\\b|nokia|Series40|Series60|S60|SonyEricsson|N900|MAUI.*WAP.*Browser"},tablets:{iPad:"iPad|iPad.*Mobile",NexusTablet:"Android.*Nexus[\\s]+(7|9|10)",GoogleTablet:"Android.*Pixel C",SamsungTablet:"SAMSUNG.*Tablet|Galaxy.*Tab|SC-01C|GT-P1000|GT-P1003|GT-P1010|GT-P3105|GT-P6210|GT-P6800|GT-P6810|GT-P7100|GT-P7300|GT-P7310|GT-P7500|GT-P7510|SCH-I800|SCH-I815|SCH-I905|SGH-I957|SGH-I987|SGH-T849|SGH-T859|SGH-T869|SPH-P100|GT-P3100|GT-P3108|GT-P3110|GT-P5100|GT-P5110|GT-P6200|GT-P7320|GT-P7511|GT-N8000|GT-P8510|SGH-I497|SPH-P500|SGH-T779|SCH-I705|SCH-I915|GT-N8013|GT-P3113|GT-P5113|GT-P8110|GT-N8010|GT-N8005|GT-N8020|GT-P1013|GT-P6201|GT-P7501|GT-N5100|GT-N5105|GT-N5110|SHV-E140K|SHV-E140L|SHV-E140S|SHV-E150S|SHV-E230K|SHV-E230L|SHV-E230S|SHW-M180K|SHW-M180L|SHW-M180S|SHW-M180W|SHW-M300W|SHW-M305W|SHW-M380K|SHW-M380S|SHW-M380W|SHW-M430W|SHW-M480K|SHW-M480S|SHW-M480W|SHW-M485W|SHW-M486W|SHW-M500W|GT-I9228|SCH-P739|SCH-I925|GT-I9200|GT-P5200|GT-P5210|GT-P5210X|SM-T311|SM-T310|SM-T310X|SM-T210|SM-T210R|SM-T211|SM-P600|SM-P601|SM-P605|SM-P900|SM-P901|SM-T217|SM-T217A|SM-T217S|SM-P6000|SM-T3100|SGH-I467|XE500|SM-T110|GT-P5220|GT-I9200X|GT-N5110X|GT-N5120|SM-P905|SM-T111|SM-T2105|SM-T315|SM-T320|SM-T320X|SM-T321|SM-T520|SM-T525|SM-T530NU|SM-T230NU|SM-T330NU|SM-T900|XE500T1C|SM-P605V|SM-P905V|SM-T337V|SM-T537V|SM-T707V|SM-T807V|SM-P600X|SM-P900X|SM-T210X|SM-T230|SM-T230X|SM-T325|GT-P7503|SM-T531|SM-T330|SM-T530|SM-T705|SM-T705C|SM-T535|SM-T331|SM-T800|SM-T700|SM-T537|SM-T807|SM-P907A|SM-T337A|SM-T537A|SM-T707A|SM-T807A|SM-T237|SM-T807P|SM-P607T|SM-T217T|SM-T337T|SM-T807T|SM-T116NQ|SM-T116BU|SM-P550|SM-T350|SM-T550|SM-T9000|SM-P9000|SM-T705Y|SM-T805|GT-P3113|SM-T710|SM-T810|SM-T815|SM-T360|SM-T533|SM-T113|SM-T335|SM-T715|SM-T560|SM-T670|SM-T677|SM-T377|SM-T567|SM-T357T|SM-T555|SM-T561|SM-T713|SM-T719|SM-T813|SM-T819|SM-T580|SM-T355Y?|SM-T280|SM-T817A|SM-T820|SM-W700|SM-P580|SM-T587|SM-P350|SM-P555M|SM-P355M|SM-T113NU|SM-T815Y|SM-T585|SM-T285|SM-T825|SM-W708|SM-T835|SM-T830|SM-T837V|SM-T720|SM-T510|SM-T387V|SM-P610|SM-T290|SM-T515|SM-T590|SM-T595|SM-T725|SM-T817P|SM-P585N0|SM-T395|SM-T295|SM-T865|SM-P610N|SM-P615|SM-T970|SM-T380|SM-T5950|SM-T905|SM-T231|SM-T500|SM-T860",Kindle:"Kindle|Silk.*Accelerated|Android.*\\b(KFOT|KFTT|KFJWI|KFJWA|KFOTE|KFSOWI|KFTHWI|KFTHWA|KFAPWI|KFAPWA|WFJWAE|KFSAWA|KFSAWI|KFASWI|KFARWI|KFFOWI|KFGIWI|KFMEWI)\\b|Android.*Silk/[0-9.]+ like Chrome/[0-9.]+ (?!Mobile)",SurfaceTablet:"Windows NT [0-9.]+; ARM;.*(Tablet|ARMBJS)",HPTablet:"HP Slate (7|8|10)|HP ElitePad 900|hp-tablet|EliteBook.*Touch|HP 8|Slate 21|HP SlateBook 10",AsusTablet:"^.*PadFone((?!Mobile).)*$|Transformer|TF101|TF101G|TF300T|TF300TG|TF300TL|TF700T|TF700KL|TF701T|TF810C|ME171|ME301T|ME302C|ME371MG|ME370T|ME372MG|ME172V|ME173X|ME400C|Slider SL101|\\bK00F\\b|\\bK00C\\b|\\bK00E\\b|\\bK00L\\b|TX201LA|ME176C|ME102A|\\bM80TA\\b|ME372CL|ME560CG|ME372CG|ME302KL| K010 | K011 | K017 | K01E |ME572C|ME103K|ME170C|ME171C|\\bME70C\\b|ME581C|ME581CL|ME8510C|ME181C|P01Y|PO1MA|P01Z|\\bP027\\b|\\bP024\\b|\\bP00C\\b",BlackBerryTablet:"PlayBook|RIM Tablet",HTCtablet:"HTC_Flyer_P512|HTC Flyer|HTC Jetstream|HTC-P715a|HTC EVO View 4G|PG41200|PG09410",MotorolaTablet:"xoom|sholest|MZ615|MZ605|MZ505|MZ601|MZ602|MZ603|MZ604|MZ606|MZ607|MZ608|MZ609|MZ615|MZ616|MZ617",NookTablet:"Android.*Nook|NookColor|nook browser|BNRV200|BNRV200A|BNTV250|BNTV250A|BNTV400|BNTV600|LogicPD Zoom2",AcerTablet:"Android.*; \\b(A100|A101|A110|A200|A210|A211|A500|A501|A510|A511|A700|A701|W500|W500P|W501|W501P|W510|W511|W700|G100|G100W|B1-A71|B1-710|B1-711|A1-810|A1-811|A1-830)\\b|W3-810|\\bA3-A10\\b|\\bA3-A11\\b|\\bA3-A20\\b|\\bA3-A30|A3-A40",ToshibaTablet:"Android.*(AT100|AT105|AT200|AT205|AT270|AT275|AT300|AT305|AT1S5|AT500|AT570|AT700|AT830)|TOSHIBA.*FOLIO",LGTablet:"\\bL-06C|LG-V909|LG-V900|LG-V700|LG-V510|LG-V500|LG-V410|LG-V400|LG-VK810\\b",FujitsuTablet:"Android.*\\b(F-01D|F-02F|F-05E|F-10D|M532|Q572)\\b",PrestigioTablet:"PMP3170B|PMP3270B|PMP3470B|PMP7170B|PMP3370B|PMP3570C|PMP5870C|PMP3670B|PMP5570C|PMP5770D|PMP3970B|PMP3870C|PMP5580C|PMP5880D|PMP5780D|PMP5588C|PMP7280C|PMP7280C3G|PMP7280|PMP7880D|PMP5597D|PMP5597|PMP7100D|PER3464|PER3274|PER3574|PER3884|PER5274|PER5474|PMP5097CPRO|PMP5097|PMP7380D|PMP5297C|PMP5297C_QUAD|PMP812E|PMP812E3G|PMP812F|PMP810E|PMP880TD|PMT3017|PMT3037|PMT3047|PMT3057|PMT7008|PMT5887|PMT5001|PMT5002",LenovoTablet:"Lenovo TAB|Idea(Tab|Pad)( A1|A10| K1|)|ThinkPad([ ]+)?Tablet|YT3-850M|YT3-X90L|YT3-X90F|YT3-X90X|Lenovo.*(S2109|S2110|S5000|S6000|K3011|A3000|A3500|A1000|A2107|A2109|A1107|A5500|A7600|B6000|B8000|B8080)(-|)(FL|F|HV|H|)|TB-X103F|TB-X304X|TB-X304F|TB-X304L|TB-X505F|TB-X505L|TB-X505X|TB-X605F|TB-X605L|TB-8703F|TB-8703X|TB-8703N|TB-8704N|TB-8704F|TB-8704X|TB-8704V|TB-7304F|TB-7304I|TB-7304X|Tab2A7-10F|Tab2A7-20F|TB2-X30L|YT3-X50L|YT3-X50F|YT3-X50M|YT-X705F|YT-X703F|YT-X703L|YT-X705L|YT-X705X|TB2-X30F|TB2-X30L|TB2-X30M|A2107A-F|A2107A-H|TB3-730F|TB3-730M|TB3-730X|TB-7504F|TB-7504X|TB-X704F|TB-X104F|TB3-X70F|TB-X705F|TB-8504F|TB3-X70L|TB3-710F|TB-X704L",DellTablet:"Venue 11|Venue 8|Venue 7|Dell Streak 10|Dell Streak 7",YarvikTablet:"Android.*\\b(TAB210|TAB211|TAB224|TAB250|TAB260|TAB264|TAB310|TAB360|TAB364|TAB410|TAB411|TAB420|TAB424|TAB450|TAB460|TAB461|TAB464|TAB465|TAB467|TAB468|TAB07-100|TAB07-101|TAB07-150|TAB07-151|TAB07-152|TAB07-200|TAB07-201-3G|TAB07-210|TAB07-211|TAB07-212|TAB07-214|TAB07-220|TAB07-400|TAB07-485|TAB08-150|TAB08-200|TAB08-201-3G|TAB08-201-30|TAB09-100|TAB09-211|TAB09-410|TAB10-150|TAB10-201|TAB10-211|TAB10-400|TAB10-410|TAB13-201|TAB274EUK|TAB275EUK|TAB374EUK|TAB462EUK|TAB474EUK|TAB9-200)\\b",MedionTablet:"Android.*\\bOYO\\b|LIFE.*(P9212|P9514|P9516|S9512)|LIFETAB",ArnovaTablet:"97G4|AN10G2|AN7bG3|AN7fG3|AN8G3|AN8cG3|AN7G3|AN9G3|AN7dG3|AN7dG3ST|AN7dG3ChildPad|AN10bG3|AN10bG3DT|AN9G2",IntensoTablet:"INM8002KP|INM1010FP|INM805ND|Intenso Tab|TAB1004",IRUTablet:"M702pro",MegafonTablet:"MegaFon V9|\\bZTE V9\\b|Android.*\\bMT7A\\b",EbodaTablet:"E-Boda (Supreme|Impresspeed|Izzycomm|Essential)",AllViewTablet:"Allview.*(Viva|Alldro|City|Speed|All TV|Frenzy|Quasar|Shine|TX1|AX1|AX2)",ArchosTablet:"\\b(101G9|80G9|A101IT)\\b|Qilive 97R|Archos5|\\bARCHOS (70|79|80|90|97|101|FAMILYPAD|)(b|c|)(G10| Cobalt| TITANIUM(HD|)| Xenon| Neon|XSK| 2| XS 2| PLATINUM| CARBON|GAMEPAD)\\b",AinolTablet:"NOVO7|NOVO8|NOVO10|Novo7Aurora|Novo7Basic|NOVO7PALADIN|novo9-Spark",NokiaLumiaTablet:"Lumia 2520",SonyTablet:"Sony.*Tablet|Xperia Tablet|Sony Tablet S|SO-03E|SGPT12|SGPT13|SGPT114|SGPT121|SGPT122|SGPT123|SGPT111|SGPT112|SGPT113|SGPT131|SGPT132|SGPT133|SGPT211|SGPT212|SGPT213|SGP311|SGP312|SGP321|EBRD1101|EBRD1102|EBRD1201|SGP351|SGP341|SGP511|SGP512|SGP521|SGP541|SGP551|SGP621|SGP641|SGP612|SOT31|SGP771|SGP611|SGP612|SGP712",PhilipsTablet:"\\b(PI2010|PI3000|PI3100|PI3105|PI3110|PI3205|PI3210|PI3900|PI4010|PI7000|PI7100)\\b",CubeTablet:"Android.*(K8GT|U9GT|U10GT|U16GT|U17GT|U18GT|U19GT|U20GT|U23GT|U30GT)|CUBE U8GT",CobyTablet:"MID1042|MID1045|MID1125|MID1126|MID7012|MID7014|MID7015|MID7034|MID7035|MID7036|MID7042|MID7048|MID7127|MID8042|MID8048|MID8127|MID9042|MID9740|MID9742|MID7022|MID7010",MIDTablet:"M9701|M9000|M9100|M806|M1052|M806|T703|MID701|MID713|MID710|MID727|MID760|MID830|MID728|MID933|MID125|MID810|MID732|MID120|MID930|MID800|MID731|MID900|MID100|MID820|MID735|MID980|MID130|MID833|MID737|MID960|MID135|MID860|MID736|MID140|MID930|MID835|MID733|MID4X10",MSITablet:"MSI \\b(Primo 73K|Primo 73L|Primo 81L|Primo 77|Primo 93|Primo 75|Primo 76|Primo 73|Primo 81|Primo 91|Primo 90|Enjoy 71|Enjoy 7|Enjoy 10)\\b",SMiTTablet:"Android.*(\\bMID\\b|MID-560|MTV-T1200|MTV-PND531|MTV-P1101|MTV-PND530)",RockChipTablet:"Android.*(RK2818|RK2808A|RK2918|RK3066)|RK2738|RK2808A",FlyTablet:"IQ310|Fly Vision",bqTablet:"Android.*(bq)?.*\\b(Elcano|Curie|Edison|Maxwell|Kepler|Pascal|Tesla|Hypatia|Platon|Newton|Livingstone|Cervantes|Avant|Aquaris ([E|M]10|M8))\\b|Maxwell.*Lite|Maxwell.*Plus",HuaweiTablet:"MediaPad|MediaPad 7 Youth|IDEOS S7|S7-201c|S7-202u|S7-101|S7-103|S7-104|S7-105|S7-106|S7-201|S7-Slim|M2-A01L|BAH-L09|BAH-W09|AGS-L09|CMR-AL19",NecTablet:"\\bN-06D|\\bN-08D",PantechTablet:"Pantech.*P4100",BronchoTablet:"Broncho.*(N701|N708|N802|a710)",VersusTablet:"TOUCHPAD.*[78910]|\\bTOUCHTAB\\b",ZyncTablet:"z1000|Z99 2G|z930|z990|z909|Z919|z900",PositivoTablet:"TB07STA|TB10STA|TB07FTA|TB10FTA",NabiTablet:"Android.*\\bNabi",KoboTablet:"Kobo Touch|\\bK080\\b|\\bVox\\b Build|\\bArc\\b Build",DanewTablet:"DSlide.*\\b(700|701R|702|703R|704|802|970|971|972|973|974|1010|1012)\\b",TexetTablet:"NaviPad|TB-772A|TM-7045|TM-7055|TM-9750|TM-7016|TM-7024|TM-7026|TM-7041|TM-7043|TM-7047|TM-8041|TM-9741|TM-9747|TM-9748|TM-9751|TM-7022|TM-7021|TM-7020|TM-7011|TM-7010|TM-7023|TM-7025|TM-7037W|TM-7038W|TM-7027W|TM-9720|TM-9725|TM-9737W|TM-1020|TM-9738W|TM-9740|TM-9743W|TB-807A|TB-771A|TB-727A|TB-725A|TB-719A|TB-823A|TB-805A|TB-723A|TB-715A|TB-707A|TB-705A|TB-709A|TB-711A|TB-890HD|TB-880HD|TB-790HD|TB-780HD|TB-770HD|TB-721HD|TB-710HD|TB-434HD|TB-860HD|TB-840HD|TB-760HD|TB-750HD|TB-740HD|TB-730HD|TB-722HD|TB-720HD|TB-700HD|TB-500HD|TB-470HD|TB-431HD|TB-430HD|TB-506|TB-504|TB-446|TB-436|TB-416|TB-146SE|TB-126SE",PlaystationTablet:"Playstation.*(Portable|Vita)",TrekstorTablet:"ST10416-1|VT10416-1|ST70408-1|ST702xx-1|ST702xx-2|ST80208|ST97216|ST70104-2|VT10416-2|ST10216-2A|SurfTab",PyleAudioTablet:"\\b(PTBL10CEU|PTBL10C|PTBL72BC|PTBL72BCEU|PTBL7CEU|PTBL7C|PTBL92BC|PTBL92BCEU|PTBL9CEU|PTBL9CUK|PTBL9C)\\b",AdvanTablet:"Android.* 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Tab|\\bM6pro\\b|CT1020W|arc 10HD|\\bTP750\\b|\\bQTAQZ3\\b|WVT101|TM1088|KT107"},oss:{AndroidOS:"Android",BlackBerryOS:"blackberry|\\bBB10\\b|rim tablet os",PalmOS:"PalmOS|avantgo|blazer|elaine|hiptop|palm|plucker|xiino",SymbianOS:"Symbian|SymbOS|Series60|Series40|SYB-[0-9]+|\\bS60\\b",WindowsMobileOS:"Windows CE.*(PPC|Smartphone|Mobile|[0-9]{3}x[0-9]{3})|Windows Mobile|Windows Phone [0-9.]+|WCE;",WindowsPhoneOS:"Windows Phone 10.0|Windows Phone 8.1|Windows Phone 8.0|Windows Phone OS|XBLWP7|ZuneWP7|Windows NT 6.[23]; ARM;",iOS:"\\biPhone.*Mobile|\\biPod|\\biPad|AppleCoreMedia",iPadOS:"CPU OS 13",SailfishOS:"Sailfish",MeeGoOS:"MeeGo",MaemoOS:"Maemo",JavaOS:"J2ME/|\\bMIDP\\b|\\bCLDC\\b",webOS:"webOS|hpwOS",badaOS:"\\bBada\\b",BREWOS:"BREW"},uas:{Chrome:"\\bCrMo\\b|CriOS|Android.*Chrome/[.0-9]* (Mobile)?",Dolfin:"\\bDolfin\\b",Opera:"Opera.*Mini|Opera.*Mobi|Android.*Opera|Mobile.*OPR/[0-9.]+$|Coast/[0-9.]+",Skyfire:"Skyfire",Edge:"\\bEdgiOS\\b|Mobile Safari/[.0-9]* Edge",IE:"IEMobile|MSIEMobile",Firefox:"fennec|firefox.*maemo|(Mobile|Tablet).*Firefox|Firefox.*Mobile|FxiOS",Bolt:"bolt",TeaShark:"teashark",Blazer:"Blazer",Safari:"Version((?!\\bEdgiOS\\b).)*Mobile.*Safari|Safari.*Mobile|MobileSafari",WeChat:"\\bMicroMessenger\\b",UCBrowser:"UC.*Browser|UCWEB",baiduboxapp:"baiduboxapp",baidubrowser:"baidubrowser",DiigoBrowser:"DiigoBrowser",Mercury:"\\bMercury\\b",ObigoBrowser:"Obigo",NetFront:"NF-Browser",GenericBrowser:"NokiaBrowser|OviBrowser|OneBrowser|TwonkyBeamBrowser|SEMC.*Browser|FlyFlow|Minimo|NetFront|Novarra-Vision|MQQBrowser|MicroMessenger",PaleMoon:"Android.*PaleMoon|Mobile.*PaleMoon"},props:{Mobile:"Mobile/[VER]",Build:"Build/[VER]",Version:"Version/[VER]",VendorID:"VendorID/[VER]",iPad:"iPad.*CPU[a-z ]+[VER]",iPhone:"iPhone.*CPU[a-z ]+[VER]",iPod:"iPod.*CPU[a-z ]+[VER]",Kindle:"Kindle/[VER]",Chrome:["Chrome/[VER]","CriOS/[VER]","CrMo/[VER]"],Coast:["Coast/[VER]"],Dolfin:"Dolfin/[VER]",Firefox:["Firefox/[VER]","FxiOS/[VER]"],Fennec:"Fennec/[VER]",Edge:"Edge/[VER]",IE:["IEMobile/[VER];","IEMobile [VER]","MSIE [VER];","Trident/[0-9.]+;.*rv:[VER]"],NetFront:"NetFront/[VER]",NokiaBrowser:"NokiaBrowser/[VER]",Opera:[" OPR/[VER]","Opera Mini/[VER]","Version/[VER]"],"Opera Mini":"Opera Mini/[VER]","Opera Mobi":"Version/[VER]",UCBrowser:["UCWEB[VER]","UC.*Browser/[VER]"],MQQBrowser:"MQQBrowser/[VER]",MicroMessenger:"MicroMessenger/[VER]",baiduboxapp:"baiduboxapp/[VER]",baidubrowser:"baidubrowser/[VER]",SamsungBrowser:"SamsungBrowser/[VER]",Iron:"Iron/[VER]",Safari:["Version/[VER]","Safari/[VER]"],Skyfire:"Skyfire/[VER]",Tizen:"Tizen/[VER]",Webkit:"webkit[ /][VER]",PaleMoon:"PaleMoon/[VER]",SailfishBrowser:"SailfishBrowser/[VER]",Gecko:"Gecko/[VER]",Trident:"Trident/[VER]",Presto:"Presto/[VER]",Goanna:"Goanna/[VER]",iOS:" \\bi?OS\\b [VER][ ;]{1}",Android:"Android [VER]",Sailfish:"Sailfish [VER]",BlackBerry:["BlackBerry[\\w]+/[VER]","BlackBerry.*Version/[VER]","Version/[VER]"],BREW:"BREW [VER]",Java:"Java/[VER]","Windows Phone OS":["Windows Phone OS [VER]","Windows Phone [VER]"],"Windows Phone":"Windows Phone [VER]","Windows CE":"Windows CE/[VER]","Windows NT":"Windows NT [VER]",Symbian:["SymbianOS/[VER]","Symbian/[VER]"],webOS:["webOS/[VER]","hpwOS/[VER];"]},utils:{Bot:"Googlebot|facebookexternalhit|Google-AMPHTML|s~amp-validator|AdsBot-Google|Google Keyword Suggestion|Facebot|YandexBot|YandexMobileBot|bingbot|ia_archiver|AhrefsBot|Ezooms|GSLFbot|WBSearchBot|Twitterbot|TweetmemeBot|Twikle|PaperLiBot|Wotbox|UnwindFetchor|Exabot|MJ12bot|YandexImages|TurnitinBot|Pingdom|contentkingapp|AspiegelBot",MobileBot:"Googlebot-Mobile|AdsBot-Google-Mobile|YahooSeeker/M1A1-R2D2",DesktopMode:"WPDesktop",TV:"SonyDTV|HbbTV",WebKit:"(webkit)[ /]([\\w.]+)",Console:"\\b(Nintendo|Nintendo WiiU|Nintendo 3DS|Nintendo 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