1120:
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
1099:. 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
806:
2049:
1029:. 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.
1071:. 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,
342:
47:
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1091:. 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
1067:. 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,
1087:. (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,
1119:
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
1001:, 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
867:
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
486:
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
2005:
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
1110:
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
1079:
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
1024:
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
551:
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
1062:
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
1114:
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
980:
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
1156:
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
1040:
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,
841:
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.
1103:. 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.
1005:-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
1339:
849:, 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
1125:
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.
349:
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
547:
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.
1049:
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,
1025:
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
666:
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
1335:
872:
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.
1009:
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
993:
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
769:(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.
543:
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
955:
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
1843:
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).
1353:
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).
2458:
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".
2188:
478:, 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
1401:
150:
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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
929:
nervous systems. These systems act to increase and decrease, respectively, the rate of production of electrical impulses by the sinoatrial node.
2285:
Steinmetz, Patrick R.H.; Kraus, Johanna E.M.; Larroux, Claire; Hammel, Jörg U.; Amon-Hassenzahl, Annette; Houliston, Evelyn; et al. (2012).
2151:
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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
528:. The cell membrane is covered by a lamina coat which is approximately 50 nm wide. The laminar coat is separable into two layers; the
1375:
1058:, the ability to couple myosin side chains phosphorylation to higher concentrations of the positive concentrations of calcium, and other
2121:
552:
latter is crucial in defining the surface-to-volume ratio of the cell. This heavily influences the potential electrical properties of
1336:"Does anyone know why skeletal muscle fibers have peripheral nuclei, but the cardiomyocytes not? What are the functional advantages?"
918:
765:. These satellite cells remain adjacent to a skeletal muscle fiber, situated between the sarcolemma and the basement membrane of the
1080:
the smooth muscle of this cnidarian shows a fundamental different mechanism of muscle cell development and structure in cnidarians.
1468:
Klein, CS; Marsh, GD; Petrella, RJ; Rice, CL (July 2003). "Muscle fiber number in the biceps brachii muscle of young and old men".
2535:
658:
Smooth muscle cells are spindle-shaped with wide middles, and tapering ends. They have a single nucleus and range from 30 to 200
864:, a term unique to muscle cells that refers to the cell membrane. This initiates an impulse that travels across the sarcolemma.
1291:
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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
2349:
Andrikou, Carmen; Arnone, Maria Ina (1 May 2015). "Too many ways to make a muscle: Evolution of GRNs governing myogenesis".
502:
condition results from multiple myoblasts fusing to produce each muscle fiber, where each myoblast contributes one nucleus.
2921:
825:. This pulls the Z discs closer together in a process called the sliding filament mechanism. The contraction of all the
490:
In skeletal muscle, at the end of each muscle fiber, the outer layer of the sarcolemma combines with tendon fibers at the
475:
291:. Smooth muscle has no myofibrils or sarcomeres and is therefore non-striated. Smooth muscle cells have a single nucleus.
2952:
482:(transverse tubule), which bores through the cell and emerge on the other side; together these three components form the
1651:"Enhancement of force generated by individual myosin heads in skinned rabbit psoas muscle fibers at low ionic strength"
436:
to shorten the fiber length in a muscle contraction. The third type of myofilament is an elastic filament composed of
2442:
1434:
1274:
1354:
157:
1649:
Sugi, Haruo; Abe, T; Kobayashi, T; Chaen, S; Ohnuki, Y; Saeki, Y; Sugiura, S; Guerrero-Hernandez, Agustin (2013).
1815:
1148:. Vertebrate smooth muscle was found to have evolved independently from the skeletal and cardiac muscle types.
926:
777:
556:. Additionally, deviation from the standard shape and size of the cell can have a negative prognostic impact.
2622:
145:
1844:
1513:"With the greatest care, stromal interaction molecule (STIM) proteins verify what skeletal muscle is doing"
891:
is when a muscle moves under a load. Concentric contraction is when a muscle shortens and generates force.
312:
829:
results in the contraction of the whole muscle fiber. This contraction of the myocyte is triggered by the
455:. The smallest contractile unit in the fiber is called the sarcomere which is a repeating unit within two
2528:
701:
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73:
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rate. These electrical impulses coordinate contraction throughout the remaining heart muscle via the
138:
2502:
956:
2048:
1917:"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
2588:
1450:
1201:
773:
697:
133:
121:
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Myoblasts and their derivatives, including satellite cells, can now be generated in vitro through
420:. There are three types of myofilaments: thin, thick, and elastic that work together to produce a
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571:
17:
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2008:"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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491:
2378:"Phylogenetic relationship of muscle tissues deduced from the superimposition of gene trees"
2113:
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2353:. Special Issue: Proceedings of the 3rd International Congress on Invertebrate Morphology.
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8:
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bilaterians to confirm a single lineage is questionable according to
Steinmetz, Kraus,
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838:
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800:
608:
421:
351:
30:"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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2012:
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evolutionary line. This indicates that these types of muscle developed in a common
830:
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are so-called because they have neither myofibrils nor sarcomeres and therefore no
2213:
Gash, Matthew C.; Kandle, Patricia F.; Murray, Ian V.; Varacallo, Matthew (2024).
1075:. found a similar pattern of localization in cnidarians except with the cnidarian
3014:
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to give rise to the different muscle cell types. Differentiation is regulated by
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195:
126:
52:
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1932:
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536:. In between these two layers can be several different types of ions, including
494:. Within the muscle fiber pressed against the sarcolemma are multiply flattened
2971:
2947:
1129:
921:. Sinoatrial node activity is modulated, in turn, by nerve fibers of both the
906:
883:
are skeletal muscle contractions that do not cause movement of the muscle. and
868:
large amount of ATP, as it is used in both the attachment and release of every
747:
693:
632:
628:
624:
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511:
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calcium ions from the gated membrane channels to stimulate muscle contraction.
366:
261:
234:
219:
211:
203:
183:
2362:
2064:"Kindlin-2 is required for myocyte elongation and is essential for myogenesis"
1966:"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
341:
3171:
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751:
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370:
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199:
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1982:
1965:
1876:"Molecular mechanisms regulating myogenic determination and differentiation"
1111:& Arnone discuss a deeper understanding of the evolution of myogenesis.
2715:
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2411:
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2099:
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2033:
2007:
1991:
1950:
1901:
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1054:. showed that the traditional morphological and regulatory markers such as
994:
977:
743:
671:
604:
544:
529:
456:
452:
448:
374:
265:
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2859:
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2631:
2287:"Independent evolution of striated muscles in cnidarians and bilaterians"
1116:
998:
467:, the red pigment that stores oxygen until needed for muscular activity.
417:
386:
280:
230:
2310:
2239:"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
163:
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2817:
2769:
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The origin of true muscle cells is argued by other authors to be the
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783:
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304:
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1216:"The evolutionary origin of bilaterian smooth and striated myocytes"
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1915:
Zhao R, Watt AJ, Battle MA, Li J, Bandow BJ, Duncan SA (May 2008).
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1002:
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Excitation of a myocyte causes depolarization at its synapses, the
713:
689:
663:
479:
460:
451:. Thin filaments of actin are the light filaments that make up the
390:
223:
1892:
1875:
1805:"Healthy versus sick myocytes: metabolism, structure and function"
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which provides energy to the cell during heightened exercise, and
3145:
2850:
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952:
857:
588:
537:
398:
394:
288:
2513:
1964:
Zammit, PS; Partridge, TA; Yablonka-Reuveni, Z (November 2006).
821:, thin and thick filaments slide concerning each other by using
46:
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1963:
1059:
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600:
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429:
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each contributing a nucleus to the newly formed muscle cell or
187:
97:
279:
Smooth muscle cells control involuntary movements such as the
264:
in the walls of the heart chambers, and have a single central
245:. Skeletal muscle cells and cardiac muscle cells both contain
2845:
2835:
2757:
2752:
2747:
2742:
1055:
914:
725:
721:
437:
425:
109:
1562:"Skeletal muscle fibers count on nuclear numbers for growth"
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plays a role in developmental elongation during myogenesis.
757:
Myoblasts in skeletal muscle that do not form muscle fibers
272:, and when joined in a visible unit they are described as a
2794:
2725:
2720:
2661:
2656:
2651:
2641:
2636:
2284:
2062:
Dowling JJ, Vreede AP, Kim S, Golden J, Feldman EL (2008).
2004:
717:
709:
705:
652:
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644:
636:
63:
1749:
Bentzinger, CF; Wang, YX; Rudnicki, MA (1 February 2012).
1289:
856:
When the acetylcholine is released it diffuses across the
268:. Cardiac muscle cells are joined to neighboring cells by
2212:
1707:
1628:(6th ed.). New York: McGraw-Hill. pp. 403–405.
1429:(3rd ed.). New York: McGraw-Hill. pp. 244–246.
887:
are skeletal muscle contractions that do cause movement.
620:
2280:
2278:
2276:
2274:
1710:"The development of the myotendinous junction. A review"
1626:
Anatomy & Physiology: The Unity of Form and
Function
794:
1269:(3rd ed.). New York: McGraw-Hill. pp. 72–73.
520:
has several specialized regions, which may include the
2106:
2061:
1748:
1467:
397:. Fusion depends on muscle-specific proteins known as
2271:
1842:
1751:"Building muscle: molecular regulation of myogenesis"
1708:
Charvet, B; Ruggiero, F; Le
Guellec, D (April 2012).
1648:
1352:
1171:
List of human cell types derived from the germ layers
2232:
2230:
2228:
1176:
List of distinct cell types in the adult human body
447:, myosin forms the dark filaments that make up the
303:of a muscle cell gave rise to its terminology. The
1292:"Human skeletal muscle fiber type classifications"
1128:Evolutionarily, specialized forms of skeletal and
1036:. (2012) argue that molecular markers such as the
879:: isometric, isotonic, eccentric and concentric.
2457:
2225:
1290:Scott, W; Stevens, J; Binder-Macleod, SA (2001).
1151:
997:(an organism having three germ layers), and that
973:that gives rise to muscle cells in vertebrates).
3169:
2433:. New York, NY: John Wiley & Sons. pp.
1914:
1864:page 395, Biology, Fifth Edition, Campbell, 1999
424:. The thin myofilaments are filaments of mostly
327:. The sarcolemma receives and conducts stimuli.
2348:
1742:
1451:"Structure of Skeletal Muscle | SEER Training"
913:generate electrical impulses that control the
812:
2529:
2344:
2342:
2340:
2338:
2237:Seipel, Katja; Schmid, Volker (1 June 2005).
2114:"Structure, and Function of Skeletal Muscles"
894:
728:also play a role in myocyte differentiation.
2236:
2055:
1998:
1908:
1873:
1566:Seminars in Cell & Developmental Biology
1559:
3188:Non-terminally differentiated (blast) cells
1970:Journal of Histochemistry and Cytochemistry
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639:, smooth muscle cells such as those of the
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1755:Cold Spring Harbor Perspectives in Biology
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1402:"Atrial structure, fibers, and conduction"
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1207:
837:of the myocyte. The action potential uses
345:Diagram of skeletal muscle fiber structure
45:
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2318:
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1981:
1940:
1891:
1814:. Oxford University Press. Archived from
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1674:
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1536:
1511:Cho, CH; Lee, KJ; Lee, EH (August 2018).
1510:
1307:
1249:
1231:
1200:at the U.S. National Library of Medicine
1013:rather than the mesoderm or mesendoderm.
919:electrical conduction system of the heart
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804:
583:. They are found in the walls of hollow
340:
330:
1838:
1836:
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1701:
1623:
1610:
1424:
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1342:from the original on 19 September 2017.
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505:
14:
3170:
2194:from the original on 23 September 2015
1714:Muscles, Ligaments and Tendons Journal
1553:
1213:
1193:
1191:
559:
428:and the thick filaments are of mostly
218:. Muscle cells develop from embryonic
2517:
2424:
2395:10.1093/oxfordjournals.molbev.a026170
2166:
2154:from the original on 27 February 2015
2124:from the original on 15 February 2015
1461:
1214:Brunet, Thibaut; et al. (2016).
795:Muscle contraction in striated muscle
416:consisting of long protein chains of
2922:Connective tissue in skeletal muscle
2206:
2178:"Muscle Cell Anatomy & Function"
1833:
1560:Prasad, V; Millay, DP (8 May 2021).
1443:
1283:
2621:
1802:
1504:
1394:
1188:
27:Type of cell found in muscle tissue
24:
674:in sarcomeres) to the sarcolemma.
190:of an animal. In humans and other
25:
3204:
2543:
2490:
2175:
412:A striated muscle fiber contains
2496:
2431:Muscles and Their Neural Control
2215:"Physiology, Muscle Contraction"
2047:
158:Anatomical terms of microanatomy
2953:Excitation–contraction coupling
2382:Molecular Biology and Evolution
2136:
1957:
1867:
1858:
1642:
1132:predated the divergence of the
989:are similar enough to those of
860:and binds to a receptor on the
459:. The sarcoplasm also contains
323:in a muscle cell is termed the
315:of a muscle cell is termed the
307:in a muscle cell is termed the
1368:
1328:
1152:Invertebrate muscle cell types
677:
647:to stand erect in response to
260:Cardiac muscle cells form the
229:Skeletal muscle cells form by
13:
1:
2376:OOta, S.; Saitou, N. (1999).
1181:
2150:. University of Washington.
1676:10.1371/journal.pone.0063658
1578:10.1016/j.semcdb.2021.04.015
1529:10.5483/bmbrep.2018.51.8.128
1425:Saladin, Kenneth S. (2011).
1265:Saladin, Kenneth S. (2011).
932:
476:smooth endoplasmic reticulum
365:A single muscle such as the
313:smooth endoplasmic reticulum
294:
210:is long and threadlike with
7:
2256:10.1016/j.ydbio.2005.03.032
1933:10.1016/j.ydbio.2008.03.013
1874:Perry R, Rudnick M (2000).
1767:10.1101/cshperspect.a008342
1164:
877:types of muscle contraction
813:Skeletal muscle contraction
789:
702:myogenic regulatory factors
631:and alter the shape of the
607:, and in the tracts of the
10:
3209:
2958:Sliding filament mechanism
2509:Structure of a Muscle Cell
2427:"8. Muscle cell diversity"
951:origin of muscle cells in
936:
898:
895:Cardiac muscle contraction
798:
681:
569:
563:
509:
445:striations of muscle bands
334:
182:, is a mature contractile
29:
3121:
3079:
3054:
3007:
3000:
2970:
2930:
2892:
2816:
2783:
2706:
2699:
2677:
2612:
2598:
2587:
2576:
2551:
2363:10.1016/j.jcz.2015.03.005
2144:"Muscle Fiber Excitation"
1144:sometime before 700
1045:. present evidence for a
354:, either as slow twitch (
156:
144:
132:
120:
108:
103:
93:
88:
44:
39:
2221:. StatPearls Publishing.
1455:training.seer.cancer.gov
1202:Medical Subject Headings
1108:gene regulatory networks
774:directed differentiation
627:dilate and contract the
498:; embryologically, this
474:, a specialized type of
440:, a very large protein.
241:) in a process known as
233:of myoblasts to produce
3151:Fukutin-related protein
2472:10.1002/jmor.1051500205
1983:10.1369/jhc.6r6995.2006
1355:"Cardiac muscle tissue"
1161:with exercise and age.
1146:million years ago (mya)
1122:cis regulatory elements
847:neuromuscular junctions
572:Basal electrical rhythm
516:The cell membrane of a
377:usually referred to as
194:there are three types:
51:General structure of a
3109:Sarcoplasmic reticulum
2938:Neuromuscular junction
2846:elastic filament/titin
2568:Vascular smooth muscle
2425:Hoyle, Graham (1983).
2148:courses.washington.edu
2118:courses.washington.edu
2081:10.1186/1471-2121-9-36
1309:10.1093/ptj/81.11.1810
881:Isometric contractions
823:adenosine triphosphate
809:
778:pluripotent stem cells
732:Skeletal muscle fibers
472:sarcoplasmic reticulum
346:
337:Skeletal muscle fibers
317:sarcoplasmic reticulum
255:striated muscle tissue
69:Neuromuscular junction
57:neuromuscular junction
2841:thick filament/myosin
2351:Zoologischer Anzeiger
2243:Developmental Biology
1812:oxfordjournals.org/en
937:Further information:
889:Eccentric contraction
885:isotonic contractions
808:
570:Further information:
492:myotendinous junction
434:slide over each other
381:. This occurs during
344:
331:Skeletal muscle cells
74:Skeletal muscle fiber
2505:at Wikimedia Commons
2013:Nature Biotechnology
1324:on 13 February 2015.
1095:, Steinmetz, Kraus,
875:There are four main
617:reproductive systems
506:Cardiac muscle cells
283:contractions in the
274:cardiac muscle fiber
208:skeletal muscle cell
206:(cardiomyocytes). A
53:skeletal muscle cell
2836:thin filament/actin
2822:(a, i, and h bands;
2311:10.1038/nature11180
2303:2012Natur.487..231S
1821:on 19 February 2015
1667:2013PLoSO...863658S
1624:Saladin, K (2012).
1233:10.7554/elife.19607
577:Smooth muscle cells
560:Smooth muscle cells
518:cardiac muscle cell
301:microscopic anatomy
1803:Ferrari, Roberto.
1470:Muscle & Nerve
1382:on 13 October 2015
1083:Steinmetz, Kraus,
839:transverse tubules
810:
801:Muscle contraction
763:myosatellite cells
603:, in the walls of
526:transverse tubules
422:muscle contraction
358:) or fast twitch (
352:muscle contraction
347:
270:intercalated discs
178:, also known as a
139:H2.00.05.0.00002
3178:Contractile cells
3163:
3162:
3159:
3158:
3117:
3116:
3071:Myosatellite cell
2987:Intercalated disc
2966:
2965:
2894:Connective tissue
2812:
2811:
2808:
2807:
2775:Synemin/desmuslin
2695:
2694:
2501:Media related to
2297:(7406): 231–234.
1635:978-0-07-337825-1
1482:10.1002/mus.10386
1302:(11): 1810–1816.
1077:N. vectensis
522:intercalated disc
172:
171:
167:
16:(Redirected from
3200:
3005:
3004:
2788:
2704:
2703:
2687:Laminin, alpha 2
2619:
2618:
2610:
2609:
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2584:
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2185:www.austincc.edu
2182:
2176:Ziser, Stephen.
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2110:
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2026:10.1038/nbt.3297
2002:
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1389:
1387:
1378:. Archived from
1376:"Muscle tissues"
1372:
1366:
1365:
1363:
1361:
1350:
1344:
1343:
1332:
1326:
1325:
1320:. Archived from
1311:
1296:Physical Therapy
1287:
1281:
1280:
1262:
1256:
1255:
1253:
1235:
1211:
1205:
1195:
1007:striated muscles
831:action potential
692:is an embryonic
670:(similar to the
649:cold temperature
587:, including the
214:and is called a
164:edit on Wikidata
161:
49:
37:
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1976:(11): 1177–91.
1962:
1958:
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1872:
1868:
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1859:
1849:
1847:
1845:"Smooth muscle"
1841:
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1189:
1184:
1167:
1154:
1130:cardiac muscles
1020:portion of the
945:
935:
927:parasympathetic
911:sinoatrial node
903:
901:Bowditch effect
897:
815:
803:
797:
792:
759:dedifferentiate
736:multiple nuclei
686:
680:
625:ciliary muscles
574:
568:
562:
554:excitable cells
514:
508:
339:
333:
297:
220:precursor cells
168:
84:
83:
35:
28:
23:
22:
15:
12:
11:
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2948:Muscle spindle
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2824:z and m lines)
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2491:External links
2489:
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2466:(2): 321–326.
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907:cardiomyocytes
896:
893:
814:
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799:Main article:
796:
793:
791:
788:
748:cardiac muscle
698:differentiates
694:precursor cell
682:Main article:
679:
676:
564:Main article:
561:
558:
512:Cardiac muscle
510:Main article:
507:
504:
371:multinucleated
367:biceps brachii
335:Main article:
332:
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262:cardiac muscle
235:multinucleated
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1221:
1217:
1210:
1203:
1199:
1194:
1192:
1187:
1177:
1174:
1172:
1169:
1168:
1162:
1160:
1149:
1147:
1143:
1139:
1135:
1131:
1126:
1123:
1118:
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1109:
1104:
1102:
1098:
1094:
1090:
1086:
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1078:
1074:
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1066:
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1048:
1044:
1039:
1035:
1030:
1028:
1023:
1019:
1014:
1012:
1008:
1004:
1000:
996:
992:
988:
984:
979:
974:
972:
968:
967:
962:
958:
957:morphological
954:
950:
944:
940:
930:
928:
924:
920:
916:
912:
908:
902:
892:
890:
886:
882:
878:
873:
871:
865:
863:
859:
854:
852:
851:acetylcholine
848:
843:
840:
836:
835:cell membrane
832:
828:
824:
820:
807:
802:
787:
785:
781:
779:
775:
770:
768:
764:
760:
755:
753:
752:smooth muscle
749:
745:
741:
737:
733:
729:
727:
723:
719:
715:
711:
707:
703:
699:
695:
691:
685:
675:
673:
669:
665:
661:
656:
654:
650:
646:
642:
641:arrector pili
638:
634:
630:
626:
622:
618:
614:
610:
606:
605:blood vessels
602:
598:
594:
590:
586:
582:
578:
573:
567:
566:Smooth muscle
557:
555:
549:
546:
541:
539:
535:
534:lamina lucida
531:
527:
523:
519:
513:
503:
501:
500:multinucleate
497:
493:
488:
485:
481:
477:
473:
468:
466:
462:
458:
454:
450:
446:
441:
439:
435:
431:
427:
423:
419:
415:
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408:
404:
400:
396:
392:
388:
384:
380:
376:
372:
368:
363:
361:
357:
353:
343:
338:
328:
326:
322:
321:cell membrane
318:
314:
310:
306:
302:
292:
290:
286:
282:
277:
275:
271:
267:
263:
258:
256:
252:
248:
244:
240:
236:
232:
227:
225:
221:
217:
213:
209:
205:
201:
197:
193:
189:
185:
181:
177:
165:
159:
155:
152:
149:
147:
143:
140:
137:
135:
131:
128:
125:
123:
119:
116:
113:
111:
107:
102:
99:
96:
92:
87:
80:
77:
75:
72:
70:
67:
65:
62:
61:
58:
54:
48:
43:
38:
33:
19:
3183:Animal cells
3165:
3065:
3015:Muscle fiber
2785:
2716:Dystrobrevin
2669:Dystroglycan
2463:
2459:
2453:
2430:
2420:
2385:
2381:
2371:
2354:
2350:
2294:
2290:
2249:(1): 14–26.
2246:
2242:
2218:
2208:
2196:. Retrieved
2184:
2156:. Retrieved
2147:
2138:
2126:. Retrieved
2117:
2108:
2071:
2067:
2057:
2020:(9): 962–9.
2017:
2011:
2000:
1973:
1969:
1959:
1927:(2): 614–9.
1924:
1920:
1910:
1883:
1880:Front Biosci
1879:
1869:
1860:
1848:. Retrieved
1823:. Retrieved
1816:the original
1811:
1758:
1754:
1744:
1720:(2): 53–63.
1717:
1713:
1703:
1658:
1654:
1644:
1625:
1569:
1565:
1555:
1520:
1516:
1506:
1473:
1469:
1463:
1454:
1445:
1426:
1420:
1408:. Retrieved
1396:
1386:29 September
1384:. Retrieved
1380:the original
1370:
1358:. Retrieved
1348:
1330:
1322:the original
1299:
1295:
1285:
1266:
1260:
1223:
1219:
1209:
1155:
1127:
1113:
1105:
1096:
1088:
1084:
1082:
1076:
1072:
1068:
1064:
1051:
1047:polyphyletic
1042:
1033:
1031:
1026:
1015:
978:monophyletic
975:
964:
949:evolutionary
946:
905:Specialized
904:
874:
866:
855:
844:
816:
782:
771:
756:
744:cell nucleus
742:, with each
730:
704:, including
687:
668:dense bodies
657:
575:
550:
545:cytoskeleton
542:
530:lamina densa
515:
489:
469:
442:
418:myofilaments
411:
406:
402:
364:
348:
299:The unusual
298:
278:
273:
259:
228:
216:muscle fiber
215:
179:
175:
173:
114:
3066:Muscle cell
3041:Myofilament
2860:Tropomyosin
2831:Myofilament
2632:Sarcoglycan
2198:12 February
2158:11 February
2128:13 February
1886:: D750–67.
1825:12 February
1517:BMB Reports
1476:(1): 62–8.
1117:orthologues
1101:amoebozoans
999:diploblasty
995:triploblast
991:bilaterians
923:sympathetic
819:contracting
738:, known as
678:Development
660:micrometers
609:respiratory
281:peristalsis
253:and form a
212:many nuclei
192:vertebrates
176:muscle cell
104:Identifiers
40:Muscle cell
3172:Categories
3136:Telethonin
3097:Sarcolemma
3092:Sarcoplasm
3025:extrafusal
3020:intrafusal
2982:Myocardium
2943:Motor unit
2917:Endomysium
2912:Perimysium
2800:Caveolin 3
2733:Syntrophin
2711:Dystrophin
2563:Calmodulin
2460:J. Morphol
2219:StatPearls
1182:References
1159:tsetse fly
1134:vertebrate
1115:conserved
987:Ctenophora
971:germ layer
943:Adaptation
899:See also:
862:sarcolemma
827:sarcomeres
767:endomysium
761:back into
684:Myogenesis
593:intestines
581:striations
414:myofibrils
383:myogenesis
325:sarcolemma
319:; and the
309:sarcoplasm
251:sarcomeres
247:myofibrils
243:myogenesis
3193:Myoblasts
3141:Dysferlin
3124:ungrouped
3046:Sarcomere
3032:Myofibril
2992:Nebulette
2902:Epimysium
2818:Sarcomere
2770:Dysbindin
2765:Syncoilin
2682:Sarcospan
2613:Membrane/
2600:Costamere
2404:0737-4038
1921:Dev. Biol
1604:234362466
1242:2050-084X
1138:arthropod
1038:myosin II
1027:ectocodon
939:Evolution
933:Evolution
833:over the
784:Kindlin-2
740:myonuclei
664:T-tubules
635:. In the
619:. In the
465:myoglobin
432:and they
407:myomerger
391:myoblasts
385:with the
379:myonuclei
373:with the
305:cytoplasm
295:Structure
285:esophagus
224:myoblasts
79:Myofibril
32:Myofibril
3131:Myotilin
3102:T-tubule
3062:Myoblast
2907:Fascicle
2867:Troponin
2786:related:
2589:Skeletal
2578:Striated
2503:Myocytes
2480:85719905
2412:10368962
2357:: 2–13.
2329:22763458
2265:15936326
2189:Archived
2152:Archived
2122:Archived
2100:18611274
2042:21241434
2034:26237517
1992:16899758
1951:18400219
1902:10966875
1785:22300977
1736:23738275
1695:23691080
1655:PLOS ONE
1596:33972174
1572:: 3–10.
1547:29898810
1498:20508198
1490:12811774
1340:Archived
1318:11694174
1198:Myocytes
1165:See also
1142:ancestor
1022:mesoderm
1018:endoderm
1011:ectoderm
1003:myoblast
983:Cnidaria
966:mesoderm
961:mesoderm
790:Function
714:myogenin
690:myoblast
480:T-tubule
461:glycogen
403:myomaker
399:fusogens
239:syncytia
196:skeletal
115:myocytus
94:Location
3146:Fukutin
2972:Cardiac
2931:General
2851:nebulin
2435:293–299
2320:3398149
2299:Bibcode
2091:2478659
1942:2423416
1850:10 June
1776:3281568
1727:3666507
1686:3655179
1663:Bibcode
1587:9070318
1538:6130827
1251:5167519
969:is the
953:animals
909:in the
858:synapse
672:Z discs
613:urinary
597:bladder
589:stomach
538:calcium
457:Z bands
401:called
395:myotube
360:type II
289:stomach
266:nucleus
237:cells (
222:called
204:cardiac
186:in the
180:myocyte
127:D032342
89:Details
18:Myocyte
3122:Other/
3087:Desmin
2974:muscle
2591:muscle
2580:muscle
2555:muscle
2553:Smooth
2478:
2441:
2410:
2402:
2327:
2317:
2291:Nature
2263:
2098:
2088:
2074:: 36.
2040:
2032:
1990:
1949:
1939:
1900:
1783:
1773:
1734:
1724:
1693:
1683:
1632:
1602:
1594:
1584:
1545:
1535:
1496:
1488:
1433:
1410:5 June
1316:
1273:
1248:
1240:
1204:(MeSH)
1093:Z-disc
870:myosin
716:, and
643:cause
623:, the
615:, and
601:uterus
585:organs
524:, and
496:nuclei
484:triads
453:I band
449:A band
430:myosin
387:fusion
375:nuclei
356:type I
311:; the
231:fusion
202:, and
200:smooth
188:muscle
98:Muscle
3080:Other
3055:Cells
3008:Fiber
2476:S2CID
2192:(PDF)
2181:(PDF)
2038:S2CID
1819:(PDF)
1808:(PDF)
1600:S2CID
1494:S2CID
1405:(PDF)
1360:3 May
1226:: 1.
1220:eLife
1097:et al
1089:et al
1085:et al
1073:et al
1069:et al
1065:et al
1056:actin
1052:et al
1043:et al
1034:et al
963:(the
915:heart
817:When
726:GATA6
722:GATA4
696:that
438:titin
426:actin
162:[
151:67328
110:Latin
3001:Both
2795:NOS1
2662:SGCZ
2657:SGCG
2652:SGCE
2647:SGCD
2642:SGCB
2637:SGCA
2605:DAPC
2439:ISBN
2408:PMID
2400:ISSN
2325:PMID
2261:PMID
2200:2015
2160:2015
2130:2015
2096:PMID
2030:PMID
1988:PMID
1947:PMID
1898:PMID
1852:2021
1827:2015
1781:PMID
1732:PMID
1691:PMID
1630:ISBN
1592:PMID
1543:PMID
1486:PMID
1431:ISBN
1412:2021
1388:2015
1362:2021
1314:PMID
1271:ISBN
1238:ISSN
1060:MyHC
985:and
947:The
941:and
925:and
724:and
718:MRF4
710:Myf5
706:MyoD
653:fear
645:hair
637:skin
633:lens
629:iris
621:eyes
599:and
532:and
470:The
405:and
287:and
249:and
184:cell
122:MeSH
64:Axon
55:and
2623:DAP
2468:doi
2464:150
2390:doi
2359:doi
2355:256
2315:PMC
2307:doi
2295:487
2251:doi
2247:282
2086:PMC
2076:doi
2022:doi
1978:doi
1937:PMC
1929:doi
1925:317
1888:doi
1771:PMC
1763:doi
1722:PMC
1681:PMC
1671:doi
1582:PMC
1574:doi
1570:119
1533:PMC
1525:doi
1478:doi
1304:doi
1246:PMC
1228:doi
776:of
750:or
651:or
443:In
389:of
362:).
146:FMA
3174::
2758:G2
2753:G1
2748:B2
2743:B1
2474:.
2462:.
2437:.
2429:.
2406:.
2398:.
2386:16
2384:.
2380:.
2337:^
2323:.
2313:.
2305:.
2293:.
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2259:.
2245:.
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2217:.
2187:.
2183:.
2168:^
2146:.
2120:.
2116:.
2094:.
2084:.
2070:.
2066:.
2036:.
2028:.
2018:33
2016:.
2010:.
1986:.
1974:54
1972:.
1968:.
1945:.
1935:.
1923:.
1919:.
1896:.
1882:.
1878:.
1835:^
1810:.
1793:^
1779:.
1769:.
1757:.
1753:.
1730:.
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1653:.
1612:^
1598:.
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1541:.
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1515:.
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1300:81
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688:A
655:.
611:,
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540:.
409:.
276:.
257:.
226:.
198:,
174:A
134:TH
59::
3064:/
3039:/
2882:I
2877:C
2872:T
2820:/
2738:A
2726:B
2721:A
2625::
2602:/
2537:e
2530:t
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1527::
1500:.
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1457:.
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1306::
1279:.
1254:.
1230::
1224:5
166:]
34:.
20:)
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