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Syncytium

A syncytium (/sɪnˈsɪʃiəm/; pl.: syncytia; from Greek: σύν syn "together" and κύτος kytos "box, i.e. cell") or symplasm is a multinucleate cell that can result from multiple cell fusions of uninuclear cells (i.e., cells with a single nucleus), in contrast to a coenocyte, which can result from multiple nuclear divisions without accompanying cytokinesis.[1] The muscle cell that makes up animal skeletal muscle is a classic example of a syncytium cell. The term may also refer to cells interconnected by specialized membranes with gap junctions, as seen in the heart muscle cells and certain smooth muscle cells, which are synchronized electrically in an action potential.

The field of embryogenesis uses the word syncytium to refer to the coenocytic blastoderm embryos of invertebrates, such as Drosophila melanogaster.[2]

Physiological examples edit

Protists edit

In protists, syncytia can be found in some rhizarians (e.g., chlorarachniophytes, plasmodiophorids, haplosporidians) and acellular slime moulds, dictyostelids (amoebozoans), acrasids (Excavata) and Haplozoon.

Plants edit

Some examples of plant syncytia, which result during plant development, include:

Fungi edit

A syncytium is the normal cell structure for many fungi. Most fungi of Basidiomycota exist as a dikaryon in which thread-like cells of the mycelium are partially partitioned into segments each containing two differing nuclei, called a heterokaryon.

Animals edit

Nerve net edit

The neurons which makes up the subepithelial nerve net in comb jellies (Ctenophora) are fused into a neural syncytium, consisting of a continuous plasma membrane instead of being connected through synapses.[6]

Skeletal muscle edit

A classic example of a syncytium is the formation of skeletal muscle. Large skeletal muscle fibers form by the fusion of thousands of individual muscle cells. The multinucleated arrangement is important in pathologic states such as myopathy, where focal necrosis (death) of a portion of a skeletal muscle fiber does not result in necrosis of the adjacent sections of that same skeletal muscle fiber, because those adjacent sections have their own nuclear material. Thus, myopathy is usually associated with such "segmental necrosis", with some of the surviving segments being functionally cut off from their nerve supply via loss of continuity with the neuromuscular junction.

Cardiac muscle edit

The syncytium of cardiac muscle is important because it allows rapid coordinated contraction of muscles along their entire length. Cardiac action potentials propagate along the surface of the muscle fiber from the point of synaptic contact through intercalated discs. Although a syncytium, cardiac muscle differs because the cells are not long and multinucleated. Cardiac tissue is therefore described as a functional syncytium, as opposed to the true syncytium of skeletal muscle.

Smooth muscle edit

Smooth muscle in the gastrointestinal tract is activated by a composite of three types of cells – smooth muscle cells (SMCs), interstitial cells of Cajal (ICCs), and platelet-derived growth factor receptor alpha (PDGFRα) that are electrically coupled and work together as an SIP functional syncytium.[7][8]

Osteoclasts edit

Certain animal immune-derived cells may form aggregate cells, such as the osteoclast cells responsible for bone resorption.

Placenta edit

Another important vertebrate syncytium is in the placenta of placental mammals. Embryo-derived cells that form the interface with the maternal blood stream fuse together to form a multinucleated barrier – the syncytiotrophoblast. This is probably important to limit the exchange of migratory cells between the developing embryo and the body of the mother, as some blood cells are specialized to be able to insert themselves between adjacent epithelial cells. The syncytial epithelium of the placenta does not provide such an access path from the maternal circulation into the embryo.

Glass sponges edit

Much of the body of Hexactinellid sponges is composed of syncitial tissue. This allows them to form their large siliceous spicules exclusively inside their cells.[9]

Tegument edit

The fine structure of the tegument in helminths is essentially the same in both the cestodes and trematodes. A typical tegument is 7–16 μm thick, with distinct layers. It is a syncytium consisting of multinucleated tissues with no distinct cell boundaries. The outer zone of the syncytium, called the "distal cytoplasm," is lined with a plasma membrane. This plasma membrane is in turn associated with a layer of carbohydrate-containing macromolecules known as the glycocalyx, that varies in thickness from one species to another. The distal cytoplasm is connected to the inner layer called the "proximal cytoplasm", which is the "cellular region or cyton or perikarya" through cytoplasmic tubes that are composed of microtubules. The proximal cytoplasm contains nuclei, endoplasmic reticulum, Golgi complex, mitochondria, ribosomes, glycogen deposits, and numerous vesicles.[10] The innermost layer is bounded by a layer of connective tissue known as the "basal lamina". The basal lamina is followed by a thick layer of muscle.[11]

Pathological examples edit

Viral infection edit

 
Syncytium caused by HSV-1 infection in Vero cells

Syncytia can also form when cells are infected with certain types of viruses, notably HSV-1, HIV, MeV, SARS-CoV-2, and pneumoviruses, e.g. respiratory syncytial virus (RSV). These syncytial formations create distinctive cytopathic effects when seen in permissive cells. Because many cells fuse together, syncytia are also known as multinucleated cells, giant cells, or polykaryocytes.[12] During infection, viral fusion proteins used by the virus to enter the cell are transported to the cell surface, where they can cause the host cell membrane to fuse with neighboring cells.

Reoviridae edit

Typically, the viral families that can cause syncytia are enveloped, because viral envelope proteins on the surface of the host cell are needed to fuse with other cells.[13] Certain members of the Reoviridae family are notable exceptions due to a unique set of proteins known as fusion-associated small transmembrane (FAST) proteins.[14] Reovirus induced syncytium formation is not found in humans, but is found in a number of other species and is caused by fusogenic orthoreoviruses. These fusogenic orthoreoviruses include reptilian orthoreovirus, avian orthoreovirus, Nelson Bay orthoreovirus, and baboon orthoreovirus.[15]

HIV edit

HIV infects Helper CD4+ T cells and makes them produce viral proteins, including fusion proteins. Then, the cells begin to display surface HIV glycoproteins, which are antigenic. Normally, a cytotoxic T cell will immediately come to "inject" lymphotoxins, such as perforin or granzyme, that will kill the infected T helper cell. However, if T helper cells are nearby, the gp41 HIV receptors displayed on the surface of the T helper cell will bind to other similar lymphocytes.[16] This makes dozens of T helper cells fuse cell membranes into a giant, nonfunctional syncytium, which allows the HIV virion to kill many T helper cells by infecting only one. It is associated with a faster progression of the disease[17]

Mumps edit

The mumps virus uses HN protein to stick to a potential host cell, then, the fusion protein allows it to bind with the host cell. The HN and fusion proteins are then left on the host cell walls, causing it to bind with neighbour epithelial cells.[18]

COVID-19 edit

Mutations within SARS-CoV-2 variants contain spike protein variants that can enhance syncytia formation.[19] The protease TMPRSS2 is essential for syncytia formation.[20] Syncytia can allow the virus to spread directly to other cells, shielded from neutralizing antibodies and other immune system components.[19] Syncytia formation in cells can be pathological to tissues.[19]

"Severe cases of COVID-19 are associated with extensive lung damage and the presence of infected multinucleated syncytial pneumocytes. The viral and cellular mechanisms regulating the formation of these syncytia are not well understood,"[21] but membrane cholesterol seems necessary.[22][23]

The syncytia appear to be long-lasting; the "complete regeneration" of the lungs after severe flu "does not happen" with COVID-19.[24]

See also edit

References edit

  1. ^ Daubenmire, R. F. (1936). "The Use of the Terms Coenocyte and Syncytium in Biology". Science. 84 (2189): 533–534. Bibcode:1936Sci....84..533D. doi:10.1126/science.84.2189.533. PMID 17806555.
  2. ^ Willmer, P. G. (1990). Invertebrate Relationships: Patterns in Animal Evolution. Cambridge University Press, Cambridge.
  3. ^ Płachno, B. J.; Swiątek, P. (2010). "Syncytia in plants: Cell fusion in endosperm—placental syncytium formation in Utricularia (Lentibulariaceae)". Protoplasma. 248 (2): 425–435. doi:10.1007/s00709-010-0173-1. PMID 20567861. S2CID 55445.
  4. ^ Tiwari, S. C.; Gunning, B. E. S. (1986). "Colchicine inhibits plasmodium formation and disrupts pathways of sporopollenin secretion in the anther tapetum ofTradescantia virginiana L". Protoplasma. 133 (2–3): 115. doi:10.1007/BF01304627. S2CID 24345281.
  5. ^ Murguía-Sánchez, G. (2002). "Embryo sac development in Vanroyenella plumosa, Podostemaceae". Aquatic Botany. 73 (3): 201–210. doi:10.1016/S0304-3770(02)00025-6.
  6. ^ Burkhardt, P.; Colgren, J.; Medhus, A.; Digel, L.; Naumann, B.; Soto-Angel, J. J.; Nordmann, E. L.; Sachkova, M. Y.; Kittelmann, M. (2023). "Syncytial nerve net in a ctenophore adds insights on the evolution of nervous systems". Science. 380 (6642): 293–297. Bibcode:2023Sci...380..293B. doi:10.1126/science.ade5645. PMID 37079688. S2CID 258239574.
  7. ^ Song, NN; Xu, WX (2016-10-25). "[Physiological and pathophysiological meanings of gastrointestinal smooth muscle motor unit SIP syncytium]". Sheng li xue bao: [Acta Physiologica Sinica]. 68 (5): 621–627. PMID 27778026.
  8. ^ Sanders, KM; Ward, SM; Koh, SD (July 2014). "Interstitial cells: regulators of smooth muscle function". Physiological Reviews. 94 (3): 859–907. doi:10.1152/physrev.00037.2013. PMC 4152167. PMID 24987007.
  9. ^ "Palaeos Metazoa: Porifera:Hexactinellida".
  10. ^ Gobert, Geoffrey N.; Stenzel, Deborah J.; McManus, Donald P.; Jones, Malcolm K. (December 2003). "The ultrastructural architecture of the adult Schistosoma japonicum tegument". International Journal for Parasitology. 33 (14): 1561–1575. doi:10.1016/s0020-7519(03)00255-8. ISSN 0020-7519. PMID 14636672.
  11. ^ Jerome), Bogitsh, Burton J. (Burton (2005). Human parasitology. Carter, Clint E. (Clint Earl), Oeltmann, Thomas N. Burlington, MA: Elsevier Academic Press. ISBN 0120884682. OCLC 769187741.{{cite book}}: CS1 maint: multiple names: authors list (link)
  12. ^ Albrecht, Thomas; Fons, Michael; Boldogh, Istvan; Rabson, Alan S. (1996-01-01). Baron, Samuel (ed.). Medical Microbiology (4th ed.). Galveston (TX): University of Texas Medical Branch at Galveston. ISBN 0963117211. PMID 21413282.
  13. ^ "ViralZone: Syncytium formation is induced by viral infection". viralzone.expasy.org. Retrieved 2016-12-16.
  14. ^ Salsman, Jayme; Top, Deniz; Boutilier, Julie; Duncan, Roy (2005-07-01). "Extensive Syncytium Formation Mediated by the Reovirus FAST Proteins Triggers Apoptosis-Induced Membrane Instability". Journal of Virology. 79 (13): 8090–8100. doi:10.1128/JVI.79.13.8090-8100.2005. ISSN 0022-538X. PMC 1143762. PMID 15956554.
  15. ^ Duncan, Roy; Corcoran, Jennifer; Shou, Jingyun; Stoltz, Don (2004-02-05). "Reptilian reovirus: a new fusogenic orthoreovirus species". Virology. 319 (1): 131–140. doi:10.1016/j.virol.2003.10.025. PMID 14967494.
  16. ^ Huerta, L.; López-Balderas, N.; Rivera-Toledo, E.; Sandoval, G.; Gȑmez-Icazbalceta, G.; Villarreal, C.; Lamoyi, E.; Larralde, C. (2009). "HIV-Envelope–Dependent Cell-Cell Fusion: Quantitative Studies". The Scientific World Journal. 9: 746–763. doi:10.1100/tsw.2009.90. PMC 5823155. PMID 19705036.
  17. ^ National Institutes of Health (2019-12-27). "Syncytium | Definition | AIDSinfo". Retrieved 2019-12-27.
  18. ^ "MUMPS, Mumps Virus, Mumps Infection". virology-online.com. Retrieved 2020-03-12.
  19. ^ a b c MaRajah MM, Bernier A, Buchrieser J, Schwartz O (2021). "The Mechanism and Consequences of SARS-CoV-2 Spike-Mediated Fusion and Syncytia Formation". Journal of Molecular Biology. 434 (6): 167280. doi:10.1016/j.jmb.2021.167280. PMC 8485708. PMID 34606831.
  20. ^ Chaves-Medina MJ, Gómez-Ospina JC, García-Perdomo HA (2021). "Molecular mechanisms for understanding the association between TMPRSS2 and beta coronaviruses SARS-CoV-2, SARS-CoV and MERS-CoV infection: scoping review". Archives of Microbiology. 204 (1): 77. doi:10.1007/s00203-021-02727-3. PMC 8709906. PMID 34953136.
  21. ^ Buchrieser, Julian; Dufloo, Jérémy; Hubert, Mathieu; Monel, Blandine; Planas, Delphine; Michael Rajah, Maaran; Planchais, Cyril; Porrot, Françoise; Guivel‐Benhassine, Florence; Van der Werf, Sylvie; Casartelli, Nicoletta; Mouquet, Hugo; Bruel, Timothée; Schwartz, Olivier (13 October 2020). "Syncytia formation by SARS‐CoV‐2 infected cells". The EMBO Journal. 39 (23): e106267. bioRxiv 10.1101/2020.07.14.202028. doi:10.15252/embj.2020106267. ISSN 0261-4189. PMC 7646020. PMID 33051876. As of 13 October 2020: accepted for publication and undergone full peer review but not copyedited, typeset, paginated, or proofread.
  22. ^ Sanders, David W.; Jumper, Chanelle C.; Ackerman, Paul J.; Bracha, Dan; Donlic, Anita; Kim, Hahn; Kenney, Devin; Castello-Serrano, Ivan; Suzuki, Saori; Tamura, Tomokazu; Tavares, Alexander H. (2020-12-14). "SARS-CoV-2 Requires Cholesterol for Viral Entry and Pathological Syncytia Formation". eLife. 10: 10:e65962. doi:10.7554/eLife.65962. PMC 8104966. PMID 33890572.
  23. ^ "SARS-CoV-2 needs cholesterol to invade cells and form mega cells". phys.org. Retrieved 2021-01-22.
  24. ^ Gallagher, James (23 October 2020). "Covid: Why is coronavirus so deadly?". BBC News.

syncytium, syncytium, syncytia, from, greek, σύν, together, κύτος, kytos, cell, symplasm, multinucleate, cell, that, result, from, multiple, cell, fusions, uninuclear, cells, cells, with, single, nucleus, contrast, coenocyte, which, result, from, multiple, nuc. A syncytium s ɪ n ˈ s ɪ ʃ i e m pl syncytia from Greek syn syn together and kytos kytos box i e cell or symplasm is a multinucleate cell that can result from multiple cell fusions of uninuclear cells i e cells with a single nucleus in contrast to a coenocyte which can result from multiple nuclear divisions without accompanying cytokinesis 1 The muscle cell that makes up animal skeletal muscle is a classic example of a syncytium cell The term may also refer to cells interconnected by specialized membranes with gap junctions as seen in the heart muscle cells and certain smooth muscle cells which are synchronized electrically in an action potential The field of embryogenesis uses the word syncytium to refer to the coenocytic blastoderm embryos of invertebrates such as Drosophila melanogaster 2 Contents 1 Physiological examples 1 1 Protists 1 2 Plants 1 3 Fungi 1 4 Animals 1 4 1 Nerve net 1 4 2 Skeletal muscle 1 4 3 Cardiac muscle 1 4 4 Smooth muscle 1 4 5 Osteoclasts 1 4 6 Placenta 1 4 7 Glass sponges 1 4 8 Tegument 2 Pathological examples 2 1 Viral infection 2 1 1 Reoviridae 2 1 2 HIV 2 1 3 Mumps 2 1 4 COVID 19 3 See also 4 ReferencesPhysiological examples editProtists edit In protists syncytia can be found in some rhizarians e g chlorarachniophytes plasmodiophorids haplosporidians and acellular slime moulds dictyostelids amoebozoans acrasids Excavata and Haplozoon Plants edit Some examples of plant syncytia which result during plant development include Developing endosperm 3 The non articulated laticifers The plasmodial tapetum 4 and The nucellar plasmodium of the family Podostemaceae 5 Fungi edit A syncytium is the normal cell structure for many fungi Most fungi of Basidiomycota exist as a dikaryon in which thread like cells of the mycelium are partially partitioned into segments each containing two differing nuclei called a heterokaryon Animals edit Nerve net edit The neurons which makes up the subepithelial nerve net in comb jellies Ctenophora are fused into a neural syncytium consisting of a continuous plasma membrane instead of being connected through synapses 6 Skeletal muscle edit A classic example of a syncytium is the formation of skeletal muscle Large skeletal muscle fibers form by the fusion of thousands of individual muscle cells The multinucleated arrangement is important in pathologic states such as myopathy where focal necrosis death of a portion of a skeletal muscle fiber does not result in necrosis of the adjacent sections of that same skeletal muscle fiber because those adjacent sections have their own nuclear material Thus myopathy is usually associated with such segmental necrosis with some of the surviving segments being functionally cut off from their nerve supply via loss of continuity with the neuromuscular junction Cardiac muscle edit The syncytium of cardiac muscle is important because it allows rapid coordinated contraction of muscles along their entire length Cardiac action potentials propagate along the surface of the muscle fiber from the point of synaptic contact through intercalated discs Although a syncytium cardiac muscle differs because the cells are not long and multinucleated Cardiac tissue is therefore described as a functional syncytium as opposed to the true syncytium of skeletal muscle Smooth muscle edit Smooth muscle in the gastrointestinal tract is activated by a composite of three types of cells smooth muscle cells SMCs interstitial cells of Cajal ICCs and platelet derived growth factor receptor alpha PDGFRa that are electrically coupled and work together as an SIP functional syncytium 7 8 Osteoclasts edit Certain animal immune derived cells may form aggregate cells such as the osteoclast cells responsible for bone resorption Placenta edit Another important vertebrate syncytium is in the placenta of placental mammals Embryo derived cells that form the interface with the maternal blood stream fuse together to form a multinucleated barrier the syncytiotrophoblast This is probably important to limit the exchange of migratory cells between the developing embryo and the body of the mother as some blood cells are specialized to be able to insert themselves between adjacent epithelial cells The syncytial epithelium of the placenta does not provide such an access path from the maternal circulation into the embryo Glass sponges edit Much of the body of Hexactinellid sponges is composed of syncitial tissue This allows them to form their large siliceous spicules exclusively inside their cells 9 Tegument edit The fine structure of the tegument in helminths is essentially the same in both the cestodes and trematodes A typical tegument is 7 16 mm thick with distinct layers It is a syncytium consisting of multinucleated tissues with no distinct cell boundaries The outer zone of the syncytium called the distal cytoplasm is lined with a plasma membrane This plasma membrane is in turn associated with a layer of carbohydrate containing macromolecules known as the glycocalyx that varies in thickness from one species to another The distal cytoplasm is connected to the inner layer called the proximal cytoplasm which is the cellular region or cyton or perikarya through cytoplasmic tubes that are composed of microtubules The proximal cytoplasm contains nuclei endoplasmic reticulum Golgi complex mitochondria ribosomes glycogen deposits and numerous vesicles 10 The innermost layer is bounded by a layer of connective tissue known as the basal lamina The basal lamina is followed by a thick layer of muscle 11 Pathological examples editViral infection edit nbsp Syncytium caused by HSV 1 infection in Vero cellsSyncytia can also form when cells are infected with certain types of viruses notably HSV 1 HIV MeV SARS CoV 2 and pneumoviruses e g respiratory syncytial virus RSV These syncytial formations create distinctive cytopathic effects when seen in permissive cells Because many cells fuse together syncytia are also known as multinucleated cells giant cells or polykaryocytes 12 During infection viral fusion proteins used by the virus to enter the cell are transported to the cell surface where they can cause the host cell membrane to fuse with neighboring cells Reoviridae edit Typically the viral families that can cause syncytia are enveloped because viral envelope proteins on the surface of the host cell are needed to fuse with other cells 13 Certain members of the Reoviridae family are notable exceptions due to a unique set of proteins known as fusion associated small transmembrane FAST proteins 14 Reovirus induced syncytium formation is not found in humans but is found in a number of other species and is caused by fusogenic orthoreoviruses These fusogenic orthoreoviruses include reptilian orthoreovirus avian orthoreovirus Nelson Bay orthoreovirus and baboon orthoreovirus 15 HIV edit HIV infects Helper CD4 T cells and makes them produce viral proteins including fusion proteins Then the cells begin to display surface HIV glycoproteins which are antigenic Normally a cytotoxic T cell will immediately come to inject lymphotoxins such as perforin or granzyme that will kill the infected T helper cell However if T helper cells are nearby the gp41 HIV receptors displayed on the surface of the T helper cell will bind to other similar lymphocytes 16 This makes dozens of T helper cells fuse cell membranes into a giant nonfunctional syncytium which allows the HIV virion to kill many T helper cells by infecting only one It is associated with a faster progression of the disease 17 Mumps edit The mumps virus uses HN protein to stick to a potential host cell then the fusion protein allows it to bind with the host cell The HN and fusion proteins are then left on the host cell walls causing it to bind with neighbour epithelial cells 18 COVID 19 edit Mutations within SARS CoV 2 variants contain spike protein variants that can enhance syncytia formation 19 The protease TMPRSS2 is essential for syncytia formation 20 Syncytia can allow the virus to spread directly to other cells shielded from neutralizing antibodies and other immune system components 19 Syncytia formation in cells can be pathological to tissues 19 Severe cases of COVID 19 are associated with extensive lung damage and the presence of infected multinucleated syncytial pneumocytes The viral and cellular mechanisms regulating the formation of these syncytia are not well understood 21 but membrane cholesterol seems necessary 22 23 The syncytia appear to be long lasting the complete regeneration of the lungs after severe flu does not happen with COVID 19 24 See also editAtrial syncytium Coenocyte Giant cell Heterokaryon Heterokaryosis Plasmodium life cycle Enteridium lycoperdon a plasmodial slime mould Syncytiotrophoblast XenophyophoreaReferences edit Daubenmire R F 1936 The Use of the Terms Coenocyte and Syncytium in Biology Science 84 2189 533 534 Bibcode 1936Sci 84 533D doi 10 1126 science 84 2189 533 PMID 17806555 Willmer P G 1990 Invertebrate Relationships Patterns in Animal Evolution Cambridge University Press Cambridge Plachno B J Swiatek P 2010 Syncytia in plants Cell fusion in endosperm placental syncytium formation in Utricularia Lentibulariaceae Protoplasma 248 2 425 435 doi 10 1007 s00709 010 0173 1 PMID 20567861 S2CID 55445 Tiwari S C Gunning B E S 1986 Colchicine inhibits plasmodium formation and disrupts pathways of sporopollenin secretion in the anther tapetum ofTradescantia virginiana L Protoplasma 133 2 3 115 doi 10 1007 BF01304627 S2CID 24345281 Murguia Sanchez G 2002 Embryo sac development in Vanroyenella plumosa Podostemaceae Aquatic Botany 73 3 201 210 doi 10 1016 S0304 3770 02 00025 6 Burkhardt P Colgren J Medhus A Digel L Naumann B Soto Angel J J Nordmann E L Sachkova M Y Kittelmann M 2023 Syncytial nerve net in a ctenophore adds insights on the evolution of nervous systems Science 380 6642 293 297 Bibcode 2023Sci 380 293B doi 10 1126 science ade5645 PMID 37079688 S2CID 258239574 Song NN Xu WX 2016 10 25 Physiological and pathophysiological meanings of gastrointestinal smooth muscle motor unit SIP syncytium Sheng li xue bao Acta Physiologica Sinica 68 5 621 627 PMID 27778026 Sanders KM Ward SM Koh SD July 2014 Interstitial cells regulators of smooth muscle function Physiological Reviews 94 3 859 907 doi 10 1152 physrev 00037 2013 PMC 4152167 PMID 24987007 Palaeos Metazoa Porifera Hexactinellida Gobert Geoffrey N Stenzel Deborah J McManus Donald P Jones Malcolm K December 2003 The ultrastructural architecture of the adult Schistosoma japonicum tegument International Journal for Parasitology 33 14 1561 1575 doi 10 1016 s0020 7519 03 00255 8 ISSN 0020 7519 PMID 14636672 Jerome Bogitsh Burton J Burton 2005 Human parasitology Carter Clint E Clint Earl Oeltmann Thomas N Burlington MA Elsevier Academic Press ISBN 0120884682 OCLC 769187741 a href Template Cite book html title Template Cite book cite book a CS1 maint multiple names authors list link Albrecht Thomas Fons Michael Boldogh Istvan Rabson Alan S 1996 01 01 Baron Samuel ed Medical Microbiology 4th ed Galveston TX University of Texas Medical Branch at Galveston ISBN 0963117211 PMID 21413282 ViralZone Syncytium formation is induced by viral infection viralzone expasy org Retrieved 2016 12 16 Salsman Jayme Top Deniz Boutilier Julie Duncan Roy 2005 07 01 Extensive Syncytium Formation Mediated by the Reovirus FAST Proteins Triggers Apoptosis Induced Membrane Instability Journal of Virology 79 13 8090 8100 doi 10 1128 JVI 79 13 8090 8100 2005 ISSN 0022 538X PMC 1143762 PMID 15956554 Duncan Roy Corcoran Jennifer Shou Jingyun Stoltz Don 2004 02 05 Reptilian reovirus a new fusogenic orthoreovirus species Virology 319 1 131 140 doi 10 1016 j virol 2003 10 025 PMID 14967494 Huerta L Lopez Balderas N Rivera Toledo E Sandoval G Gȑmez Icazbalceta G Villarreal C Lamoyi E Larralde C 2009 HIV Envelope Dependent Cell Cell Fusion Quantitative Studies The Scientific World Journal 9 746 763 doi 10 1100 tsw 2009 90 PMC 5823155 PMID 19705036 National Institutes of Health 2019 12 27 Syncytium Definition AIDSinfo Retrieved 2019 12 27 MUMPS Mumps Virus Mumps Infection virology online com Retrieved 2020 03 12 a b c MaRajah MM Bernier A Buchrieser J Schwartz O 2021 The Mechanism and Consequences of SARS CoV 2 Spike Mediated Fusion and Syncytia Formation Journal of Molecular Biology 434 6 167280 doi 10 1016 j jmb 2021 167280 PMC 8485708 PMID 34606831 Chaves Medina MJ Gomez Ospina JC Garcia Perdomo HA 2021 Molecular mechanisms for understanding the association between TMPRSS2 and beta coronaviruses SARS CoV 2 SARS CoV and MERS CoV infection scoping review Archives of Microbiology 204 1 77 doi 10 1007 s00203 021 02727 3 PMC 8709906 PMID 34953136 Buchrieser Julian Dufloo Jeremy Hubert Mathieu Monel Blandine Planas Delphine Michael Rajah Maaran Planchais Cyril Porrot Francoise Guivel Benhassine Florence Van der Werf Sylvie Casartelli Nicoletta Mouquet Hugo Bruel Timothee Schwartz Olivier 13 October 2020 Syncytia formation by SARS CoV 2 infected cells The EMBO Journal 39 23 e106267 bioRxiv 10 1101 2020 07 14 202028 doi 10 15252 embj 2020106267 ISSN 0261 4189 PMC 7646020 PMID 33051876 As of 13 October 2020 accepted for publication and undergone full peer review but not copyedited typeset paginated or proofread Sanders David W Jumper Chanelle C Ackerman Paul J Bracha Dan Donlic Anita Kim Hahn Kenney Devin Castello Serrano Ivan Suzuki Saori Tamura Tomokazu Tavares Alexander H 2020 12 14 SARS CoV 2 Requires Cholesterol for Viral Entry and Pathological Syncytia Formation eLife 10 10 e65962 doi 10 7554 eLife 65962 PMC 8104966 PMID 33890572 SARS CoV 2 needs cholesterol to invade cells and form mega cells phys org Retrieved 2021 01 22 Gallagher James 23 October 2020 Covid Why is coronavirus so deadly BBC News Portals nbsp Evolutionary biology nbsp Science nbsp Viruses Retrieved from https en wikipedia org w index php title Syncytium amp oldid 1186962908, wikipedia, wiki, book, books, library,

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