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had been changed through different types of structural mutations in the invertebrate deuterostomes and protostomes, and they argue that these structural changes in the genes allowed for a large divergence of muscle function and muscle formation in these species. Andrikou & Arnone were able to recognize not only any difference due to mutation in the genes found in vertebrates and invertebrates but also the integration of species-specific genes that could also cause divergence from the original gene regulatory network function. Thus, although a common muscle patterning system has been determined, they argue that this could be due to a more ancestral gene regulatory network being coopted several times across lineages with additional genes and mutations causing very divergent development of muscles. Thus it seems that the myogenic patterning framework may be an ancestral trait. However, Andrikou & Arnone explain that the basic muscle patterning structure must also be considered in combination with the
1088:. argue that there are only four common protein components that were present in all bilaterians muscle ancestors and that of these for necessary Z-disc components only an actin protein that they have already argued is an uninformative marker through its pleisiomorphic state is present in cnidarians. Through further molecular marker testing, Steinmetz et al. observe that non-bilaterians lack many regulatory and structural components necessary for bilaterians muscle formation and do not find any unique set of proteins to both bilaterians and cnidarians and ctenophores that are not present in earlier, more primitive animals such as the sponges and
795:
2038:
1018:. Schmid & Seipel argue that, even in bilaterians, not all muscle cells are derived from the mesendoderm: Their key examples are that in both the eye muscles of vertebrates and the muscles of spiralians, these cells derive from the ectodermal mesoderm, rather than the endodermal mesoderm. Furthermore, they argue that since myogenesis does occur in cnidarians with the help of the same molecular regulatory elements found in the specification of muscle cells in bilaterians, that there is evidence for a single origin for striated muscle.
1060:. also showed that the localization of this duplicated set of genes that serve both the function of facilitating the formation of striated muscle genes, and cell regulation and movement genes, were already separated into striated much and non-muscle MHC. This separation of the duplicated set of genes is shown through the localization of the striated much to the contractile vacuole in sponges, while the non-muscle much was more diffusely expressed during developmental cell shape and change. Steinmetz, Kraus,
331:
36:
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1080:. were not able to find even on unique striated muscle cell protein that was expressed in both cnidarians and bilaterians. Furthermore, the Z-disc seemed to have evolved differently even within bilaterians and there is a great deal of diversity of proteins developed even between this clade, showing a large degree of radiation for muscle cells. Through this divergence of the
1056:. Furthermore, they explain that the orthologues of the Myc genes that have been used to hypothesize the origin of striated muscle occurred through a gene duplication event that predates the first true muscle cells (meaning striated muscle), and they show that the Myc genes are present in the sponges that have contractile elements but no true muscle cells. Steinmetz, Kraus,
1076:. (2012) further argue for multiple origins of striated muscle in the metazoans by explaining that a key set of genes used to form the troponin complex for muscle regulation and formation in bilaterians is missing from the cnidarians and ctenophores, and 47 structural and regulatory proteins observed, Steinmetz, Kraus,
1108:
of the gene regulatory network in both invertebrate bilaterians and cnidarians. They argue that having this common, general regulatory circuit allowed for a high degree of divergence from a single well-functioning network. Andrikou & Arnone found that the orthologues of genes found in vertebrates
990:, meaning an organism with two germ layers, evolved secondarily, because of their observation of the lack of mesoderm or muscle found in most cnidarians and ctenophores. By comparing the morphology of cnidarians and ctenophores to bilaterians, Schmid & Seipel were able to conclude that there were
856:
When the action potential reaches the sarcoplasmic reticulum it triggers the release of Ca from the Ca channels. The Ca flows from the sarcoplasmic reticulum into the sarcomere with both of its filaments. This causes the filaments to start sliding and the sarcomeres to become shorter. This requires a
475:
that exist within the network of the sarcoplasmic reticulum, in which each T-tubule has two terminal cisternae on each side of it. The sarcoplasmic reticulum serves as a reservoir for calcium ions, so when an action potential spreads over the T-tubule, it signals the sarcoplasmic reticulum to release
1994:
Chal J, Oginuma M, Al
Tanoury Z, Gobert B, Sumara O, Hick A, Bousson F, Zidouni Y, Mursch C, Moncuquet P, Tassy O, Vincent S, Miyazaki A, Bera A, Garnier JM, Guevara G, Heston M, Kennedy L, Hayashi S, Drayton B, Cherrier T, Gayraud-Morel B, Gussoni E, Relaix F, Tajbakhsh S, Pourquié O (August 2015).
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present at different times during development. In contrast with the high level of gene family apparatuses structure, Andrikou and Arnone found that the cis-regulatory elements were not well conserved both in time and place in the network which could show a large degree of divergence in the formation
1099:
to look at how the hierarchy of genes and morphogens and another mechanism of tissue specification diverge and are similar among early deuterostomes and protostomes. By understanding not only what genes are present in all bilaterians but also the time and place of deployment of these genes, Andrikou
1068:
having this striated muscle marker present in the smooth muscle of the digestive tract. Thus, they argue that the pleisiomorphic trait of the separated orthologues of much cannot be used to determine the monophylogeny of muscle, and additionally argue that the presence of a striated muscle marker in
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and the endoderm. However, Schmid & Seipel (2005) counter skepticism – about whether the muscle cells found in ctenophores and cnidarians are "true" muscle cells – by considering that cnidarians develop through a medusa stage and polyp stage. They note that in the hydrozoans' medusa stage, there
540:
The cytoskeleton is what the rest of the cell builds off of and has two primary purposes; the first is to stabilize the topography of the intracellular components and the second is to help control the size and shape of the cell. While the first function is important for biochemical processes, the
1051:
elements are present in all metazoans not just the organisms that have been shown to have muscle cells. Thus, the usage of any of these structural or regulatory elements in determining whether or not the muscle cells of the cnidarians and ctenophores are similar enough to the muscle cells of the
1103:
In their paper, Andrikou & Arnone (2015) argue that to truly understand the evolution of muscle cells the function of transcriptional regulators must be understood in the context of other external and internal interactions. Through their analysis, Andrikou & Arnone found that there were
969:
trait that occurred concurrently with the development of the digestive and nervous systems of all animals, and that this origin can be traced to a single metazoan ancestor in which muscle cells are present. They argue that molecular and morphological similarities between the muscles cells in
1145:
The properties used for distinguishing fast, intermediate, and slow muscle fibers can be different for invertebrate flight and jump muscle. To further complicate this classification scheme, the mitochondrial content, and other morphological properties within a muscle fiber, can change in a
1029:
protein used to determine this single origin of striated muscle predate the formation of muscle cells. They use an example of the contractile elements present in the
Porifera, or sponges, that do truly lack this striated muscle containing this protein. Furthermore, Steinmetz, Kraus,
830:
to get from the surface to the interior of the myocyte, which is continuous within the cell membrane. Sarcoplasmic reticula are membranous bags that transverse tubules touch but remain separate from. These wrap themselves around each sarcomere and are filled with Ca.
1092:. Through this analysis, the authors conclude that due to the lack of elements that bilaterian muscles are dependent on for structure and usage, nonbilaterian muscles must be of a different origin with a different set of regulatory and structural proteins.
994:-like structures in the tentacles and gut of some species of cnidarians and the tentacles of ctenophores. Since this is a structure unique to muscle cells, these scientists determined based on the data collected by their peers that this is a marker for
1328:
838:, which triggers an action potential. With a singular neuromuscular junction, each muscle fiber receives input from just one somatic efferent neuron. Action potential in a somatic efferent neuron causes the release of the neurotransmitter
1114:
of muscle cells. Through this analysis, it seems that the myogenic GRN is an ancestral GRN with actual changes in myogenic function and structure possibly being linked to later coopts of genes at different times and places.
338:
Skeletal muscle cells are the individual contractile cells within a muscle and are more usually known as muscle fibers because of their longer threadlike appearance. Broadly there are two types of muscle fiber performing in
536:
by anchor fibers that are approximately 10 nm wide. These are generally located at the Z lines so that they form grooves and transverse tubules emanate. In cardiac myocytes, this forms a scalloped surface.
1038:
origin of striated muscle cell development through their analysis of morphological and molecular markers that are present in bilaterians and absent in cnidarians, ctenophores, and bilaterians. Steinmetz, Kraus,
1014:
is a layer of cells that separate from the distal side of the ectoderm, which forms the striated muscle cells in a way similar to that of the mesoderm; they call this third separated layer of cells the
655:
found in striated muscle cells. Although smooth muscle cells lack sarcomeres and myofibrils they do contain large amounts of the contractile proteins actin and myosin. Actin filaments are anchored by
1324:
861:
head. Very quickly Ca is actively transported back into the sarcoplasmic reticulum, which blocks the interaction between the thin and thick filament. This in turn causes the muscle cell to relax.
998:
similar to that observed in bilaterians. The authors also remark that the muscle cells found in cnidarians and ctenophores are often contested due to the origin of these muscle cells being the
982:
that there would be one ancestor in metazoans from which muscle cells derive. In this case, Schmid & Seipel argue that the last common ancestor of
Bilateria, Ctenophora and Cnidaria, was a
758:(the connective tissue investment that divides the muscle fascicles into individual fibers). To re-activate myogenesis, the satellite cells must be stimulated to differentiate into new fibers.
532:
Cardiac muscle like the skeletal muscle is also striated and the cells contain myofibrils, myofilaments, and sarcomeres as the skeletal muscle cell. The cell membrane is anchored to the cell's
944:
is highly debated: One view is that muscle cells evolved once, and thus all muscle cells have a single common ancestor. Another view is that muscles cells evolved more than once, and any
1832:
Betts, J. Gordon; Young, Kelly A.; Wise, James A.; Johnson, Eddie; Poe, Brandon; Kruse, Dean H.; Korol, Oksana; Johnson, Jody E.; Womble, Mark; Desaix, Peter (6 March 2013).
1342:
Betts, J. Gordon; Young, Kelly A.; Wise, James A.; Johnson, Eddie; Poe, Brandon; Kruse, Dean H.; Korol, Oksana; Johnson, Jody E.; Womble, Mark; Desaix, Peter (6 March 2013).
2447:
Anderson, M.; Finlayson, L.H. (1976). "The effect of exercise on the growth of mitochondria and myofibrils in the flight muscles of the Tsetse fly, Glossina morsitans".
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467:, forms a network around each myofibril of the muscle fiber. This network is composed of groupings of two dilated end-sacs called terminal cisternae, and a single
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in length. This is thousands of times shorter than skeletal muscle fibers. The diameter of their cells is also much smaller which removes the need for
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nervous systems. These systems act to increase and decrease, respectively, the rate of production of electrical impulses by the sinoatrial node.
2274:
Steinmetz, Patrick R.H.; Kraus, Johanna E.M.; Larroux, Claire; Hammel, Jörg U.; Amon-Hassenzahl, Annette; Houliston, Evelyn; et al. (2012).
2140:
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or structural similarities are due to convergent evolution, and the development of shared genes that predate the evolution of muscle – even the
517:. The cell membrane is covered by a lamina coat which is approximately 50 nm wide. The laminar coat is separable into two layers; the
1364:
1047:, the ability to couple myosin side chains phosphorylation to higher concentrations of the positive concentrations of calcium, and other
2110:
541:
latter is crucial in defining the surface-to-volume ratio of the cell. This heavily influences the potential electrical properties of
1325:"Does anyone know why skeletal muscle fibers have peripheral nuclei, but the cardiomyocytes not? What are the functional advantages?"
907:
754:. These satellite cells remain adjacent to a skeletal muscle fiber, situated between the sarcolemma and the basement membrane of the
1069:
the smooth muscle of this cnidarian shows a fundamental different mechanism of muscle cell development and structure in cnidarians.
1457:
Klein, CS; Marsh, GD; Petrella, RJ; Rice, CL (July 2003). "Muscle fiber number in the biceps brachii muscle of young and old men".
2524:
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Smooth muscle cells are spindle-shaped with wide middles, and tapering ends. They have a single nucleus and range from 30 to 200
853:, a term unique to muscle cells that refers to the cell membrane. This initiates an impulse that travels across the sarcolemma.
1280:
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in a young adult human male contains around 253,000 muscle fibers. Skeletal muscle fibers are the only muscle cells that are
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Andrikou, Carmen; Arnone, Maria Ina (1 May 2015). "Too many ways to make a muscle: Evolution of GRNs governing myogenesis".
491:
condition results from multiple myoblasts fusing to produce each muscle fiber, where each myoblast contributes one nucleus.
2910:
814:. This pulls the Z discs closer together in a process called the sliding filament mechanism. The contraction of all the
479:
In skeletal muscle, at the end of each muscle fiber, the outer layer of the sarcolemma combines with tendon fibers at the
464:
280:. Smooth muscle has no myofibrils or sarcomeres and is therefore non-striated. Smooth muscle cells have a single nucleus.
2941:
471:(transverse tubule), which bores through the cell and emerge on the other side; together these three components form the
1640:"Enhancement of force generated by individual myosin heads in skinned rabbit psoas muscle fibers at low ionic strength"
425:
to shorten the fiber length in a muscle contraction. The third type of myofilament is an elastic filament composed of
2431:
1423:
1263:
1343:
146:
1638:
Sugi, Haruo; Abe, T; Kobayashi, T; Chaen, S; Ohnuki, Y; Saeki, Y; Sugiura, S; Guerrero-Hernandez, Agustin (2013).
1804:
1137:. Vertebrate smooth muscle was found to have evolved independently from the skeletal and cardiac muscle types.
915:
766:
545:. Additionally, deviation from the standard shape and size of the cell can have a negative prognostic impact.
2611:
134:
1833:
1502:"With the greatest care, stromal interaction molecule (STIM) proteins verify what skeletal muscle is doing"
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is when a muscle moves under a load. Concentric contraction is when a muscle shortens and generates force.
301:
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results in the contraction of the whole muscle fiber. This contraction of the myocyte is triggered by the
444:. The smallest contractile unit in the fiber is called the sarcomere which is a repeating unit within two
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rate. These electrical impulses coordinate contraction throughout the remaining heart muscle via the
127:
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1906:"Loss of both GATA4 and GATA6 blocks cardiac myocyte differentiation and results in acardia in mice"
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originating from a single myoblast. The fusion of myoblasts is specific to skeletal muscle, and not
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1439:
1190:
762:
686:
122:
110:
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Myoblasts and their derivatives, including satellite cells, can now be generated in vitro through
409:. There are three types of myofilaments: thin, thick, and elastic that work together to produce a
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1997:"Differentiation of pluripotent stem cells to muscle fiber to model Duchenne muscular dystrophy"
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In another take on the argument, Andrikou & Arnone (2015) use the newly available data on
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2367:"Phylogenetic relationship of muscle tissues deduced from the superimposition of gene trees"
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2342:. Special Issue: Proceedings of the 3rd International Congress on Invertebrate Morphology.
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bilaterians to confirm a single lineage is questionable according to
Steinmetz, Kraus,
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340:
19:"Muscle fiber" and "Myofiber" redirect here. For protein structures inside cells, see
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In contrast to this argument for a single origin of muscle cells, Steinmetz, Kraus,
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evolutionary line. This indicates that these types of muscle developed in a common
819:
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are so-called because they have neither myofibrils nor sarcomeres and therefore no
2202:
Gash, Matthew C.; Kandle, Patricia F.; Murray, Ian V.; Varacallo, Matthew (2024).
1064:. found a similar pattern of localization in cnidarians except with the cnidarian
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to give rise to the different muscle cell types. Differentiation is regulated by
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525:. In between these two layers can be several different types of ions, including
483:. Within the muscle fiber pressed against the sarcolemma are multiply flattened
2960:
2936:
1118:
910:. Sinoatrial node activity is modulated, in turn, by nerve fibers of both the
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are skeletal muscle contractions that do not cause movement of the muscle. and
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large amount of ATP, as it is used in both the attachment and release of every
736:
682:
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calcium ions from the gated membrane channels to stimulate muscle contraction.
355:
250:
223:
208:
200:
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172:
2351:
2053:"Kindlin-2 is required for myocyte elongation and is essential for myogenesis"
1955:"The skeletal muscle satellite cell: the stem cell that came in from the cold"
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are made when myoblasts fuse together; muscle fibers therefore are cells with
330:
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1971:
1954:
1865:"Molecular mechanisms regulating myogenic determination and differentiation"
1100:& Arnone discuss a deeper understanding of the evolution of myogenesis.
2704:
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2400:
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2022:
1996:
1980:
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1043:. showed that the traditional morphological and regulatory markers such as
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732:
660:
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518:
445:
441:
437:
363:
254:
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2620:
2276:"Independent evolution of striated muscles in cnidarians and bilaterians"
1105:
987:
456:, the red pigment that stores oxygen until needed for muscular activity.
406:
375:
269:
219:
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2228:"Evolution of striated muscle: Jellyfish and the origin of triploblasty"
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Schmid & Seipel (2005) argue that the origin of muscle cells is a
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2014:
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The origin of true muscle cells is argued by other authors to be the
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1205:"The evolutionary origin of bilaterian smooth and striated myocytes"
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Zhao R, Watt AJ, Battle MA, Li J, Bandow BJ, Duncan SA (May 2008).
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Excitation of a myocyte causes depolarization at its synapses, the
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652:
468:
449:
440:. Thin filaments of actin are the light filaments that make up the
379:
212:
1881:
1864:
1794:"Healthy versus sick myocytes: metabolism, structure and function"
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which provides energy to the cell during heightened exercise, and
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941:
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577:
526:
387:
383:
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2502:
1953:
Zammit, PS; Partridge, TA; Yablonka-Reuveni, Z (November 2006).
810:, thin and thick filaments slide concerning each other by using
35:
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1952:
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each contributing a nucleus to the newly formed muscle cell or
176:
86:
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Smooth muscle cells control involuntary movements such as the
253:
in the walls of the heart chambers, and have a single central
234:. Skeletal muscle cells and cardiac muscle cells both contain
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2824:
2746:
2741:
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2731:
1044:
903:
714:
710:
426:
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98:
1551:"Skeletal muscle fibers count on nuclear numbers for growth"
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plays a role in developmental elongation during myogenesis.
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Myoblasts in skeletal muscle that do not form muscle fibers
261:, and when joined in a visible unit they are described as a
2783:
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2709:
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2640:
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2625:
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Dowling JJ, Vreede AP, Kim S, Golden J, Feldman EL (2008).
1993:
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52:
1738:
Bentzinger, CF; Wang, YX; Rudnicki, MA (1 February 2012).
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When the acetylcholine is released it diffuses across the
257:. Cardiac muscle cells are joined to neighboring cells by
2201:
1696:
1617:(6th ed.). New York: McGraw-Hill. pp. 403–405.
1418:(3rd ed.). New York: McGraw-Hill. pp. 244–246.
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are skeletal muscle contractions that do cause movement.
609:
2269:
2267:
2265:
2263:
1699:"The development of the myotendinous junction. A review"
1615:
Anatomy & Physiology: The Unity of Form and
Function
783:
1258:(3rd ed.). New York: McGraw-Hill. pp. 72–73.
509:
has several specialized regions, which may include the
2095:
2050:
1737:
1456:
386:. Fusion depends on muscle-specific proteins known as
2260:
1831:
1740:"Building muscle: molecular regulation of myogenesis"
1697:
Charvet, B; Ruggiero, F; Le
Guellec, D (April 2012).
1637:
1341:
1160:
List of human cell types derived from the germ layers
2221:
2219:
2217:
1165:
List of distinct cell types in the adult human body
436:, myosin forms the dark filaments that make up the
292:of a muscle cell gave rise to its terminology. The
1281:"Human skeletal muscle fiber type classifications"
1117:Evolutionarily, specialized forms of skeletal and
1025:. (2012) argue that molecular markers such as the
868:: isometric, isotonic, eccentric and concentric.
2446:
2214:
1279:Scott, W; Stevens, J; Binder-Macleod, SA (2001).
1140:
986:(an organism having three germ layers), and that
962:that gives rise to muscle cells in vertebrates).
3158:
2422:. New York, NY: John Wiley & Sons. pp.
1903:
1853:page 395, Biology, Fifth Edition, Campbell, 1999
413:. The thin myofilaments are filaments of mostly
316:. The sarcolemma receives and conducts stimuli.
2337:
1731:
1440:"Structure of Skeletal Muscle | SEER Training"
902:generate electrical impulses that control the
801:
2518:
2333:
2331:
2329:
2327:
2226:Seipel, Katja; Schmid, Volker (1 June 2005).
2103:"Structure, and Function of Skeletal Muscles"
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717:also play a role in myocyte differentiation.
2225:
2044:
1987:
1897:
1862:
1555:Seminars in Cell & Developmental Biology
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3177:Non-terminally differentiated (blast) cells
1959:Journal of Histochemistry and Cytochemistry
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628:, smooth muscle cells such as those of the
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1744:Cold Spring Harbor Perspectives in Biology
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1391:"Atrial structure, fibers, and conduction"
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826:of the myocyte. The action potential uses
334:Diagram of skeletal muscle fiber structure
34:
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1880:
1803:. Oxford University Press. Archived from
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1500:Cho, CH; Lee, KJ; Lee, EH (August 2018).
1499:
1296:
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1220:
1189:at the U.S. National Library of Medicine
1002:rather than the mesoderm or mesendoderm.
908:electrical conduction system of the heart
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572:. They are found in the walls of hollow
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1331:from the original on 19 September 2017.
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3159:
2183:from the original on 23 September 2015
1703:Muscles, Ligaments and Tendons Journal
1542:
1202:
1182:
1180:
548:
417:and the thick filaments are of mostly
207:. Muscle cells develop from embryonic
2506:
2413:
2384:10.1093/oxfordjournals.molbev.a026170
2155:
2143:from the original on 27 February 2015
2113:from the original on 15 February 2015
1450:
1203:Brunet, Thibaut; et al. (2016).
784:Muscle contraction in striated muscle
405:consisting of long protein chains of
2911:Connective tissue in skeletal muscle
2195:
2167:"Muscle Cell Anatomy & Function"
1822:
1549:Prasad, V; Millay, DP (8 May 2021).
1432:
1272:
2610:
1791:
1493:
1383:
1177:
16:Type of cell found in muscle tissue
13:
663:in sarcomeres) to the sarcolemma.
179:of an animal. In humans and other
14:
3193:
2532:
2479:
2164:
401:A striated muscle fiber contains
2485:
2420:Muscles and Their Neural Control
2204:"Physiology, Muscle Contraction"
2036:
147:Anatomical terms of microanatomy
2942:Excitation–contraction coupling
2371:Molecular Biology and Evolution
2125:
1946:
1856:
1847:
1631:
1121:predated the divergence of the
978:are similar enough to those of
849:and binds to a receptor on the
448:. The sarcoplasm also contains
312:in a muscle cell is termed the
304:of a muscle cell is termed the
296:in a muscle cell is termed the
1357:
1317:
1141:Invertebrate muscle cell types
666:
636:to stand erect in response to
249:Cardiac muscle cells form the
218:Skeletal muscle cells form by
1:
2365:OOta, S.; Saitou, N. (1999).
1170:
2139:. University of Washington.
1665:10.1371/journal.pone.0063658
1567:10.1016/j.semcdb.2021.04.015
1518:10.5483/bmbrep.2018.51.8.128
1414:Saladin, Kenneth S. (2011).
1254:Saladin, Kenneth S. (2011).
921:
465:smooth endoplasmic reticulum
354:A single muscle such as the
302:smooth endoplasmic reticulum
283:
199:is long and threadlike with
7:
2245:10.1016/j.ydbio.2005.03.032
1922:10.1016/j.ydbio.2008.03.013
1863:Perry R, Rudnick M (2000).
1756:10.1101/cshperspect.a008342
1153:
866:types of muscle contraction
802:Skeletal muscle contraction
778:
691:myogenic regulatory factors
620:and alter the shape of the
596:, and in the tracts of the
10:
3198:
2947:Sliding filament mechanism
2498:Structure of a Muscle Cell
2416:"8. Muscle cell diversity"
940:origin of muscle cells in
925:
887:
884:Cardiac muscle contraction
787:
670:
558:
552:
498:
434:striations of muscle bands
323:
171:, is a mature contractile
18:
3110:
3068:
3043:
2996:
2989:
2959:
2919:
2881:
2805:
2772:
2695:
2688:
2666:
2601:
2587:
2576:
2565:
2540:
2352:10.1016/j.jcz.2015.03.005
2133:"Muscle Fiber Excitation"
1133:sometime before 700
1034:. present evidence for a
343:, either as slow twitch (
145:
133:
121:
109:
97:
92:
82:
77:
33:
28:
2210:. StatPearls Publishing.
1444:training.seer.cancer.gov
1191:Medical Subject Headings
1097:gene regulatory networks
763:directed differentiation
616:dilate and contract the
487:; embryologically, this
463:, a specialized type of
429:, a very large protein.
230:) in a process known as
222:of myoblasts to produce
3140:Fukutin-related protein
2461:10.1002/jmor.1051500205
1972:10.1369/jhc.6r6995.2006
1344:"Cardiac muscle tissue"
1150:with exercise and age.
1135:million years ago (mya)
1111:cis regulatory elements
836:neuromuscular junctions
561:Basal electrical rhythm
505:The cell membrane of a
366:usually referred to as
183:there are three types:
40:General structure of a
3098:Sarcoplasmic reticulum
2927:Neuromuscular junction
2835:elastic filament/titin
2557:Vascular smooth muscle
2414:Hoyle, Graham (1983).
2137:courses.washington.edu
2107:courses.washington.edu
2070:10.1186/1471-2121-9-36
1298:10.1093/ptj/81.11.1810
870:Isometric contractions
812:adenosine triphosphate
798:
767:pluripotent stem cells
721:Skeletal muscle fibers
461:sarcoplasmic reticulum
335:
326:Skeletal muscle fibers
306:sarcoplasmic reticulum
244:striated muscle tissue
58:Neuromuscular junction
46:neuromuscular junction
2830:thick filament/myosin
2340:Zoologischer Anzeiger
2232:Developmental Biology
1801:oxfordjournals.org/en
926:Further information:
878:Eccentric contraction
874:isotonic contractions
797:
559:Further information:
481:myotendinous junction
423:slide over each other
370:. This occurs during
333:
320:Skeletal muscle cells
63:Skeletal muscle fiber
2494:at Wikimedia Commons
2002:Nature Biotechnology
1313:on 13 February 2015.
1084:, Steinmetz, Kraus,
864:There are four main
606:reproductive systems
495:Cardiac muscle cells
272:contractions in the
263:cardiac muscle fiber
197:skeletal muscle cell
195:(cardiomyocytes). A
42:skeletal muscle cell
2825:thin filament/actin
2811:(a, i, and h bands;
2300:10.1038/nature11180
2292:2012Natur.487..231S
1810:on 19 February 2015
1656:2013PLoSO...863658S
1613:Saladin, K (2012).
1222:10.7554/elife.19607
566:Smooth muscle cells
549:Smooth muscle cells
507:cardiac muscle cell
290:microscopic anatomy
1792:Ferrari, Roberto.
1459:Muscle & Nerve
1371:on 13 October 2015
1072:Steinmetz, Kraus,
828:transverse tubules
799:
790:Muscle contraction
752:myosatellite cells
592:, in the walls of
515:transverse tubules
411:muscle contraction
347:) or fast twitch (
341:muscle contraction
336:
259:intercalated discs
167:, also known as a
128:H2.00.05.0.00002
3167:Contractile cells
3152:
3151:
3148:
3147:
3106:
3105:
3060:Myosatellite cell
2976:Intercalated disc
2955:
2954:
2883:Connective tissue
2801:
2800:
2797:
2796:
2764:Synemin/desmuslin
2684:
2683:
2490:Media related to
2286:(7406): 231–234.
1624:978-0-07-337825-1
1471:10.1002/mus.10386
1291:(11): 1810–1816.
1066:N. vectensis
511:intercalated disc
161:
160:
156:
3189:
2994:
2993:
2777:
2693:
2692:
2676:Laminin, alpha 2
2608:
2607:
2599:
2598:
2585:
2584:
2574:
2573:
2527:
2520:
2513:
2504:
2503:
2489:
2473:
2472:
2444:
2438:
2437:
2411:
2405:
2404:
2386:
2362:
2356:
2355:
2335:
2322:
2321:
2311:
2271:
2258:
2257:
2247:
2223:
2212:
2211:
2199:
2193:
2192:
2190:
2188:
2182:
2174:www.austincc.edu
2171:
2165:Ziser, Stephen.
2162:
2153:
2152:
2150:
2148:
2129:
2123:
2122:
2120:
2118:
2099:
2093:
2092:
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2072:
2048:
2042:
2041:
2040:
2034:
2015:10.1038/nbt.3297
1991:
1985:
1984:
1974:
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1944:
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1404:
1402:
1400:
1395:
1387:
1381:
1380:
1378:
1376:
1367:. Archived from
1365:"Muscle tissues"
1361:
1355:
1354:
1352:
1350:
1339:
1333:
1332:
1321:
1315:
1314:
1309:. Archived from
1300:
1285:Physical Therapy
1276:
1270:
1269:
1251:
1245:
1244:
1242:
1224:
1200:
1194:
1184:
996:striated muscles
820:action potential
681:is an embryonic
659:(similar to the
638:cold temperature
576:, including the
203:and is called a
153:edit on Wikidata
150:
38:
26:
25:
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2096:
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1992:
1988:
1965:(11): 1177–91.
1951:
1947:
1902:
1898:
1861:
1857:
1852:
1848:
1838:
1836:
1834:"Smooth muscle"
1830:
1823:
1813:
1811:
1807:
1796:
1790:
1781:
1736:
1732:
1695:
1691:
1636:
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1266:
1252:
1248:
1201:
1197:
1185:
1178:
1173:
1156:
1143:
1119:cardiac muscles
1009:portion of the
934:
924:
916:parasympathetic
900:sinoatrial node
892:
890:Bowditch effect
886:
804:
792:
786:
781:
748:dedifferentiate
725:multiple nuclei
675:
669:
614:ciliary muscles
563:
557:
551:
543:excitable cells
503:
497:
328:
322:
286:
209:precursor cells
157:
73:
72:
24:
17:
12:
11:
5:
3195:
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3179:
3174:
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3150:
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3137:
3132:
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3100:
3095:
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3078:
3072:
3070:
3066:
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3063:
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3057:
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2987:
2986:
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2973:
2967:
2965:
2957:
2956:
2953:
2952:
2950:
2949:
2944:
2939:
2937:Muscle spindle
2934:
2929:
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2898:
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2813:z and m lines)
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2481:
2480:External links
2478:
2475:
2474:
2455:(2): 321–326.
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2377:(6): 856–867.
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1750:(2): a008342.
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896:cardiomyocytes
885:
882:
803:
800:
788:Main article:
785:
782:
780:
777:
737:cardiac muscle
687:differentiates
683:precursor cell
671:Main article:
668:
665:
553:Main article:
550:
547:
501:Cardiac muscle
499:Main article:
496:
493:
360:multinucleated
356:biceps brachii
324:Main article:
321:
318:
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251:cardiac muscle
224:multinucleated
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2057:BMC Cell Biol
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1181:
1176:
1166:
1163:
1161:
1158:
1157:
1151:
1149:
1138:
1136:
1132:
1128:
1124:
1120:
1115:
1112:
1107:
1101:
1098:
1093:
1091:
1087:
1083:
1079:
1075:
1070:
1067:
1063:
1059:
1055:
1050:
1046:
1042:
1037:
1033:
1028:
1024:
1019:
1017:
1012:
1008:
1003:
1001:
997:
993:
989:
985:
981:
977:
973:
968:
963:
961:
957:
956:
951:
947:
946:morphological
943:
939:
933:
929:
919:
917:
913:
909:
905:
901:
897:
891:
881:
879:
875:
871:
867:
862:
860:
854:
852:
848:
843:
841:
840:acetylcholine
837:
832:
829:
825:
824:cell membrane
821:
817:
813:
809:
796:
791:
776:
774:
770:
768:
764:
759:
757:
753:
749:
744:
742:
741:smooth muscle
738:
734:
730:
726:
722:
718:
716:
712:
708:
704:
700:
696:
692:
688:
684:
680:
674:
664:
662:
658:
654:
650:
645:
643:
639:
635:
631:
630:arrector pili
627:
623:
619:
615:
611:
607:
603:
599:
595:
594:blood vessels
591:
587:
583:
579:
575:
571:
567:
562:
556:
555:Smooth muscle
546:
544:
538:
535:
530:
528:
524:
523:lamina lucida
520:
516:
512:
508:
502:
492:
490:
489:multinucleate
486:
482:
477:
474:
470:
466:
462:
457:
455:
451:
447:
443:
439:
435:
430:
428:
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420:
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412:
408:
404:
399:
397:
393:
389:
385:
381:
377:
373:
369:
365:
361:
357:
352:
350:
346:
342:
332:
327:
317:
315:
311:
310:cell membrane
307:
303:
299:
295:
291:
281:
279:
275:
271:
266:
264:
260:
256:
252:
247:
245:
241:
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233:
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225:
221:
216:
214:
210:
206:
202:
198:
194:
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178:
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170:
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154:
148:
144:
141:
138:
136:
132:
129:
126:
124:
120:
117:
114:
112:
108:
105:
102:
100:
96:
91:
88:
85:
81:
76:
69:
66:
64:
61:
59:
56:
54:
51:
50:
47:
43:
37:
32:
27:
22:
3172:Animal cells
3154:
3054:
3004:Muscle fiber
2774:
2705:Dystrobrevin
2658:Dystroglycan
2452:
2448:
2442:
2419:
2409:
2374:
2370:
2360:
2343:
2339:
2283:
2279:
2238:(1): 14–26.
2235:
2231:
2207:
2197:
2185:. Retrieved
2173:
2145:. Retrieved
2136:
2127:
2115:. Retrieved
2106:
2097:
2060:
2056:
2046:
2009:(9): 962–9.
2006:
2000:
1989:
1962:
1958:
1948:
1916:(2): 614–9.
1913:
1909:
1899:
1872:
1869:Front Biosci
1868:
1858:
1849:
1837:. Retrieved
1812:. Retrieved
1805:the original
1800:
1747:
1743:
1733:
1709:(2): 53–63.
1706:
1702:
1692:
1647:
1643:
1633:
1614:
1558:
1554:
1544:
1509:
1505:
1495:
1462:
1458:
1452:
1443:
1434:
1415:
1409:
1397:. Retrieved
1385:
1375:29 September
1373:. Retrieved
1369:the original
1359:
1347:. Retrieved
1337:
1319:
1311:the original
1288:
1284:
1274:
1255:
1249:
1212:
1208:
1198:
1144:
1116:
1102:
1094:
1085:
1077:
1073:
1071:
1065:
1061:
1057:
1053:
1040:
1036:polyphyletic
1031:
1022:
1020:
1015:
1004:
967:monophyletic
964:
953:
938:evolutionary
935:
894:Specialized
893:
863:
855:
844:
833:
805:
771:
760:
745:
733:cell nucleus
731:, with each
719:
693:, including
676:
657:dense bodies
646:
564:
539:
534:cytoskeleton
531:
519:lamina densa
504:
478:
458:
431:
407:myofilaments
400:
395:
391:
353:
337:
288:The unusual
287:
267:
262:
248:
217:
205:muscle fiber
204:
168:
164:
162:
103:
3055:Muscle cell
3030:Myofilament
2849:Tropomyosin
2820:Myofilament
2621:Sarcoglycan
2187:12 February
2147:11 February
2117:13 February
1875:: D750–67.
1814:12 February
1506:BMB Reports
1465:(1): 62–8.
1106:orthologues
1090:amoebozoans
988:diploblasty
984:triploblast
980:bilaterians
912:sympathetic
808:contracting
727:, known as
667:Development
649:micrometers
598:respiratory
270:peristalsis
242:and form a
201:many nuclei
181:vertebrates
165:muscle cell
93:Identifiers
29:Muscle cell
3161:Categories
3125:Telethonin
3086:Sarcolemma
3081:Sarcoplasm
3014:extrafusal
3009:intrafusal
2971:Myocardium
2932:Motor unit
2906:Endomysium
2901:Perimysium
2789:Caveolin 3
2722:Syntrophin
2700:Dystrophin
2552:Calmodulin
2449:J. Morphol
2208:StatPearls
1171:References
1148:tsetse fly
1123:vertebrate
1104:conserved
976:Ctenophora
960:germ layer
932:Adaptation
888:See also:
851:sarcolemma
816:sarcomeres
756:endomysium
750:back into
673:Myogenesis
582:intestines
570:striations
403:myofibrils
372:myogenesis
314:sarcolemma
308:; and the
298:sarcoplasm
240:sarcomeres
236:myofibrils
232:myogenesis
3182:Myoblasts
3130:Dysferlin
3113:ungrouped
3035:Sarcomere
3021:Myofibril
2981:Nebulette
2891:Epimysium
2807:Sarcomere
2759:Dysbindin
2754:Syncoilin
2671:Sarcospan
2602:Membrane/
2589:Costamere
2393:0737-4038
1910:Dev. Biol
1593:234362466
1231:2050-084X
1127:arthropod
1027:myosin II
1016:ectocodon
928:Evolution
922:Evolution
822:over the
773:Kindlin-2
729:myonuclei
653:T-tubules
624:. In the
608:. In the
454:myoglobin
421:and they
396:myomerger
380:myoblasts
374:with the
368:myonuclei
362:with the
294:cytoplasm
284:Structure
274:esophagus
213:myoblasts
68:Myofibril
21:Myofibril
3120:Myotilin
3091:T-tubule
3051:Myoblast
2896:Fascicle
2856:Troponin
2775:related:
2578:Skeletal
2567:Striated
2492:Myocytes
2469:85719905
2401:10368962
2346:: 2–13.
2318:22763458
2254:15936326
2178:Archived
2141:Archived
2111:Archived
2089:18611274
2031:21241434
2023:26237517
1981:16899758
1940:18400219
1891:10966875
1774:22300977
1725:23738275
1684:23691080
1644:PLOS ONE
1585:33972174
1561:: 3–10.
1536:29898810
1487:20508198
1479:12811774
1329:Archived
1307:11694174
1187:Myocytes
1154:See also
1131:ancestor
1011:mesoderm
1007:endoderm
1000:ectoderm
992:myoblast
972:Cnidaria
955:mesoderm
950:mesoderm
779:Function
703:myogenin
679:myoblast
469:T-tubule
450:glycogen
392:myomaker
388:fusogens
228:syncytia
185:skeletal
104:myocytus
83:Location
3135:Fukutin
2961:Cardiac
2920:General
2840:nebulin
2424:293–299
2309:3398149
2288:Bibcode
2080:2478659
1931:2423416
1839:10 June
1765:3281568
1716:3666507
1675:3655179
1652:Bibcode
1576:9070318
1527:6130827
1240:5167519
958:is the
942:animals
898:in the
847:synapse
661:Z discs
602:urinary
586:bladder
578:stomach
527:calcium
446:Z bands
390:called
384:myotube
349:type II
278:stomach
255:nucleus
226:cells (
211:called
193:cardiac
175:in the
169:myocyte
116:D032342
78:Details
3111:Other/
3076:Desmin
2963:muscle
2580:muscle
2569:muscle
2544:muscle
2542:Smooth
2467:
2430:
2399:
2391:
2316:
2306:
2280:Nature
2252:
2087:
2077:
2063:: 36.
2029:
2021:
1979:
1938:
1928:
1889:
1772:
1762:
1723:
1713:
1682:
1672:
1621:
1591:
1583:
1573:
1534:
1524:
1485:
1477:
1422:
1399:5 June
1305:
1262:
1237:
1229:
1193:(MeSH)
1082:Z-disc
859:myosin
705:, and
632:cause
612:, the
604:, and
590:uterus
574:organs
513:, and
485:nuclei
473:triads
442:I band
438:A band
419:myosin
376:fusion
364:nuclei
345:type I
300:; the
220:fusion
191:, and
189:smooth
177:muscle
87:Muscle
3069:Other
3044:Cells
2997:Fiber
2465:S2CID
2181:(PDF)
2170:(PDF)
2027:S2CID
1808:(PDF)
1797:(PDF)
1589:S2CID
1483:S2CID
1394:(PDF)
1349:3 May
1215:: 1.
1209:eLife
1086:et al
1078:et al
1074:et al
1062:et al
1058:et al
1054:et al
1045:actin
1041:et al
1032:et al
1023:et al
952:(the
904:heart
806:When
715:GATA6
711:GATA4
685:that
427:titin
415:actin
151:[
140:67328
99:Latin
2990:Both
2784:NOS1
2651:SGCZ
2646:SGCG
2641:SGCE
2636:SGCD
2631:SGCB
2626:SGCA
2594:DAPC
2428:ISBN
2397:PMID
2389:ISSN
2314:PMID
2250:PMID
2189:2015
2149:2015
2119:2015
2085:PMID
2019:PMID
1977:PMID
1936:PMID
1887:PMID
1841:2021
1816:2015
1770:PMID
1721:PMID
1680:PMID
1619:ISBN
1581:PMID
1532:PMID
1475:PMID
1420:ISBN
1401:2021
1377:2015
1351:2021
1303:PMID
1260:ISBN
1227:ISSN
1049:MyHC
974:and
936:The
930:and
914:and
713:and
707:MRF4
699:Myf5
695:MyoD
642:fear
634:hair
626:skin
622:lens
618:iris
610:eyes
588:and
521:and
459:The
394:and
276:and
238:and
173:cell
111:MeSH
53:Axon
44:and
2612:DAP
2457:doi
2453:150
2379:doi
2348:doi
2344:256
2304:PMC
2296:doi
2284:487
2240:doi
2236:282
2075:PMC
2065:doi
2011:doi
1967:doi
1926:PMC
1918:doi
1914:317
1877:doi
1760:PMC
1752:doi
1711:PMC
1670:PMC
1660:doi
1571:PMC
1563:doi
1559:119
1522:PMC
1514:doi
1467:doi
1293:doi
1235:PMC
1217:doi
765:of
739:or
640:or
432:In
378:of
351:).
135:FMA
3163::
2747:G2
2742:G1
2737:B2
2732:B1
2463:.
2451:.
2426:.
2418:.
2395:.
2387:.
2375:16
2373:.
2369:.
2326:^
2312:.
2302:.
2294:.
2282:.
2278:.
2262:^
2248:.
2234:.
2230:.
2216:^
2206:.
2176:.
2172:.
2157:^
2135:.
2109:.
2105:.
2083:.
2073:.
2059:.
2055:.
2025:.
2017:.
2007:33
2005:.
1999:.
1975:.
1963:54
1961:.
1957:.
1934:.
1924:.
1912:.
1908:.
1885:.
1871:.
1867:.
1824:^
1799:.
1782:^
1768:.
1758:.
1746:.
1742:.
1719:.
1705:.
1701:.
1678:.
1668:.
1658:.
1646:.
1642:.
1601:^
1587:.
1579:.
1569:.
1557:.
1553:.
1530:.
1520:.
1510:51
1508:.
1504:.
1481:.
1473:.
1463:28
1461:.
1442:.
1327:.
1301:.
1289:81
1287:.
1283:.
1233:.
1225:.
1211:.
1207:.
1179:^
1125:/
842:.
769:.
743:.
709:.
701:,
697:,
677:A
644:.
600:,
584:,
580:,
529:.
398:.
265:.
246:.
215:.
187:,
163:A
123:TH
48::
3053:/
3028:/
2871:I
2866:C
2861:T
2809:/
2727:A
2715:B
2710:A
2614::
2591:/
2526:e
2519:t
2512:v
2471:.
2459::
2436:.
2403:.
2381::
2354:.
2350::
2320:.
2298::
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2256:.
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2061:9
2033:.
2013::
1983:.
1969::
1942:.
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1754::
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1565::
1538:.
1516::
1489:.
1469::
1446:.
1428:.
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1379:.
1353:.
1295::
1268:.
1243:.
1219::
1213:5
155:]
23:.
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