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Non-cytolytic enterovirus
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== Introduction == Non-cytolytic enterovirus is an aberrant form of viral infection [[enterovirus B]] serotypes (such as [[Coxsackie B virus|coxsackievirus B]] and [[echovirus]]) can transmute into within the host, a transformation that allows a virus normally only capable of acute infections to create persistent ones. Molecular evidence for the existence of this form of infection dates back to at least 1990, but an understanding of the molecular mechanism behind the conversion of lytic to non-cytolytic enterovirus was only obtained in 2005, when Professor [[Nora Chapman]] figured out the mechanism. The transformation of normal [[lytic]] (wild-type) enterovirus into the non-cytolytic form results from mutations (specifically, deletions in the [[genome]]) the virus acquires in the host during the first weeks of acute [[infection]]. This altered form of enterovirus may then cause persistent intracellular infections which replicate very slowly without killing the cells the virus resides in. Non-cytolytic enterovirus infections it is thought may lead to cellular dysfunction, affect immune signaling, induce autoimmunity, or elicit a pro-inflammatory immune response. Non-cytolytic enterovirus is difficult for the immune system to eradicate, so it can persist for very long periods, and is associated with a range of chronic diseases. Persistent enteroviral infections have been found in: * The [[brain]], [[muscle]] and stomach tissues of ''myalgic encephalomyelitis / [[chronic fatigue syndrome]]'' (ME/CFS) patients; see [[List of enterovirus infection studies|enterovirus infection studies]]. * The heart muscle in murine and human coxsackievirus B (CVB) ''chronic [[myocarditis]]''.<ref name="Chapman2008a">{{Cite journal | last = Chapman | first = Nora M. | last2 = Kim | first2 = Kyung-Soo | last3 = Drescher | first3 = Kristen M. | last4 = Oka | first4 = Kuniyuki | last5 = Tracy | first5 = Steven | date = 2008-06-05 | title = 5' terminal deletions in the genome of a coxsackievirus B2 strain occurred naturally in human heart|url=https://www.ncbi.nlm.nih.gov/pubmed/18378272/|journal=Virology|volume=375|issue=2 | pages = 480–491|doi=10.1016/j.virol.2008.02.030|issn=0042-6822|pmc=2440640|pmid=18378272}}</ref><ref name="Chapman2008b">{{Cite journal | last = Chapman | first = N.M. | last2 = Kim | first2 = K.S. | date = 2008 | title = Persistent coxsackievirus infection: enterovirus persistence in chronic myocarditis and dilated cardiomyopathy|url=https://www.ncbi.nlm.nih.gov/pubmed/18357775|journal=Current Topics in Microbiology and Immunology|volume=323|pages=275–292|issn=0070-217X|pmid=18357775}}</ref> * The heart muscle in ''dilated cardiomyopathy'' (a sequelae of myocarditis).<ref name="Archard1987" /><ref name="Bowles1989">{{Cite journal | last = Bowles | first = N. E. | last2 = Rose | first2 = M.L. | last3 = Taylor | first3 = P. | last4 = Banner | first4 = N.R. | last5 = Morgan-Capner | first5 = P. | last6 = Cunningham | first6 = L. | last7 = Archard | first7 = L.C. | last8 = Yacoub | first8 = M.H. | date = Nov 1989 | title = End-stage dilated cardiomyopathy. Persistence of enterovirus RNA in myocardium at cardiac transplantation and lack of immune response|url=https://www.ncbi.nlm.nih.gov/pubmed/2553297|journal=Circulation|volume=80|issue=5|pages=1128–1136|issn=0009-7322|pmid=2553297}}</ref><ref name="Andreoletti1995">{{Cite journal | last = Andreoletti | first = L. | last2 = Wattre | first2 = P. | last3 = Decoene | first3 = C. | last4 = Lobert | first4 = P.E. | last5 = Dewilde | first5 = A. | last6 = Hober | first6 = D. | date = Nov 1995 | title = Detection of enterovirus-specific RNA sequences in explanted myocardium biopsy specimens from patients with dilated or ischemic cardiomyopathy|url=https://www.ncbi.nlm.nih.gov/pubmed/8589166|journal=Clinical Infectious Diseases: An Official Publication of the Infectious Diseases Society of America|volume=21|issue=5|pages=1315–1317|issn=1058-4838|pmid=8589166}}</ref><ref name="Andreoletti1996">{{Cite journal | last = Andreoletti | first = L. | last2 = Hober | first2 = D. | last3 = Decoene | first3 = C. | last4 = Copin | first4 = M.C. | last5 = Lobert | first5 = P.E. | last6 = Dewilde | first6 = A. | last7 = Stankowiac | first7 = C. | last8 = Wattre | first8 = P. | date = Jan 1996 | title = Detection of enteroviral RNA by polymerase chain reaction in endomyocardial tissue of patients with chronic cardiac diseases|url=https://www.ncbi.nlm.nih.gov/pubmed/8825711|journal=Journal of Medical Virology|volume=48|issue=1 | pages = 53–59|doi=10.1002/(SICI)1096-9071(199601)48:13.0.CO;2-K|issn=0146-6615|pmid=8825711}}</ref><ref name="Li2000">{{Cite journal | last = Li | first = Y. | last2 = Bourlet | first2 = T. | last3 = Andreoletti | first3 = L. | last4 = Mosnier | first4 = J.F. | last5 = Peng | first5 = T. | last6 = Yang | first6 = Y. | last7 = Archard | first7 = L.C. | last8 = Pozzetto | first8 = B. | last9 = Zhang | first9 = H. | date = 2000-01-25 | title = Enteroviral capsid protein VP1 is present in myocardial tissues from some patients with myocarditis or dilated cardiomyopathy|url=https://www.ncbi.nlm.nih.gov/pubmed/10645916|journal=Circulation|volume=101|issue=3|pages=231–234|issn=1524-4539|pmid=10645916}}</ref><ref name="Andreoletti2000">{{Cite journal | last = Andréoletti | first = L. | last2 = Bourlet | first2 = T. | last3 = Moukassa | first3 = D. | last4 = Rey | first4 = L. | last5 = Hot | first5 = D. | last6 = Li | first6 = Y. | last7 = Lambert | first7 = V. | last8 = Gosselin | first8 = B. | last9 = Mosnier | first9 = J.F. | date = Oct 2000 | title = Enteroviruses can persist with or without active viral replication in cardiac tissue of patients with end-stage ischemic or dilated cardiomyopathy|url=https://www.ncbi.nlm.nih.gov/pubmed/10979922|journal=The Journal of Infectious Diseases|volume=182|issue=4|pages=1222–1227|doi=10.1086/315818|issn=0022-1899|pmid=10979922}}</ref><ref name="Rey2001" /><ref name="Nguyen2013">{{Cite journal | last = Nguyen | first = Yohan | last2 = Renois | first2 = Fanny | last3 = Leveque | first3 = Nicolas | last4 = Giusti | first4 = Delphine | last5 = Picard-Maureau | first5 = Marcus | last6 = Bruneval | first6 = Patrick | last7 = Fornes | first7 = Paul | last8 = Andreoletti | first8 = Laurent | date = Jul 2013 | title = Virus detection and semiquantitation in explanted heart tissues of idiopathic dilated cardiomyopathy adult patients by use of PCR coupled with mass spectrometry analysis|url=https://www.ncbi.nlm.nih.gov/pubmed/23658274|journal=Journal of Clinical Microbiology|volume=51|issue=7|pages=2288–2294|doi=10.1128/JCM.00820-13|issn=1098-660X|pmc=3697695|pmid=23658274}}</ref><ref name="Bouin2016" /> * The [[cerebrospinal fluid]] and [[spinal cord]] tissue of [[amyotrophic lateral sclerosis]] (a motor neuron disease).<ref name="Vandenberghe2010">{{Cite journal | last = Vandenberghe | first = Nadia | last2 = Leveque | first2 = Nicolas | last3 = Corcia | first3 = Philippe | last4 = Brunaud-Danel | first4 = Véronique | last5 = Salort-Campana | first5 = Emmanuelle | last6 = Besson | first6 = Gérard | last7 = Tranchant | first7 = Christine | last8 = Clavelou | first8 = Pierre | last9 = Beaulieux | first9 = Frédérique | date = 2010-05-03 | title = Cerebrospinal fluid detection of enterovirus genome in ALS: a study of 242 patients and 354 controls|url=https://www.ncbi.nlm.nih.gov/pubmed/19900148|journal=Amyotrophic Lateral Sclerosis: Official Publication of the World Federation of Neurology Research Group on Motor Neuron Diseases|volume=11|issue=3|pages=277–282|doi=10.3109/17482960903262083|issn=1471-180X|pmid=19900148}}</ref><ref name="Woodall1994">{{Cite journal | last = Woodall | first = C. J. | last2 = Riding | first2 = M.H. | last3 = Graham | first3 = D.I. | last4 = Clements | first4 = G.B. | date = 1994-06-11 | title = Sequences specific for enterovirus detected in spinal cord from patients with motor neurone disease|url=https://www.ncbi.nlm.nih.gov/pubmed/8019310/|journal=BMJ (Clinical research ed.)|volume=308|issue=6943|pages=1541–1543|issn=0959-8138|pmc=2540520|pmid=8019310}}</ref><ref name="Xue2018">{{Cite journal | last = Xue | first = Yuan Chao | last2 = Feuer | first2 = Ralph | last3 = Cashman | first3 = Neil | last4 = Luo | first4 = Honglin | date = 2018 | title = Enteroviral Infection: The Forgotten Link to Amyotrophic Lateral Sclerosis?|url=https://www.frontiersin.org/articles/10.3389/fnmol.2018.00063/full|journal=Frontiers in Molecular Neuroscience|language=English|volume=11|doi=10.3389/fnmol.2018.00063|issn=1662-5099|pmc=5857577|pmid=29593492}}</ref> * The [[brainstem]] in [[Parkinson's disease]].<ref name="Dourmashkin2018">{{Cite journal | last = Dourmashkin | first = Robert R. | last2 = McCall | first2 = Sherman A. | last3 = Dourmashkin | first3 = Neil | last4 = Hannah | first4 = Matthew J. | date = 2018 | title = Virus-like particles and enterovirus antigen found in the brainstem neurons of Parkinson's disease|url=https://www.ncbi.nlm.nih.gov/pubmed/29899977|journal=F1000Research|volume=7|pages=302|doi=10.12688/f1000research.13626.2|issn=2046-1402|pmc=5968367|pmid=29899977}}</ref> * The salivary gland epithelial cells and lymphocyte infiltrates in primary [[Sjögren's syndrome]].<ref name="Trian2004">{{Cite journal | last = Triantafyllopoulou | first = Antigoni | last2 = Tapinos | first2 = Nikos | last3 = Moutsopoulos | first3 = Haralampos M. | date = Sep 2004 | title = Evidence for coxsackievirus infection in primary Sjögren's syndrome|url=https://www.ncbi.nlm.nih.gov/pubmed/15457458/|journal=Arthritis and Rheumatism|volume=50|issue=9|pages=2897–2902|doi=10.1002/art.20463|issn=0004-3591|pmid=15457458}}</ref> * The [[cerebrospinal fluid]] in [[post-polio syndrome]].<ref name="Muir1995">{{Cite journal | last = Muir | first = P. | last2 = Nicholson | first2 = F. | last3 = Sharief | first3 = M.K. | last4 = Thompson | first4 = E.J. | last5 = Cairns | first5 = N.J. | last6 = Lantos | first6 = P. | last7 = Spencer | first7 = G.T. | last8 = Kaminski | first8 = H.J. | last9 = Banatvala | first9 = J.E. | date = 1995-05-25 | title = Evidence for persistent enterovirus infection of the central nervous system in patients with previous paralytic poliomyelitis|url=https://www.ncbi.nlm.nih.gov/pubmed/7611631|journal=Annals of the New York Academy of Sciences|volume=753|pages=219–232|issn=0077-8923|pmid=7611631}}</ref> * The ''murine pancreas''<ref name="Tracy2015">{{Cite journal | last = Tracy | first = Steven | last2 = Smithee | first2 = Shane | last3 = Alhazmi | first3 = Abdulaziz | last4 = Chapman | first4 = Nora | date = Feb 2015 | title = Coxsackievirus can persist in murine pancreas by deletion of 5' terminal genomic sequences|url=https://www.ncbi.nlm.nih.gov/pubmed/25111164|journal=Journal of Medical Virology|volume=87|issue=2|pages=240–247|doi=10.1002/jmv.24039|issn=1096-9071|pmid=25111164}}</ref> and human pancreas,<ref name="Chehadeh2000">{{Cite journal | last = Chehadeh | first = W. | last2 = Kerr-Conte | first2 = J. | last3 = Pattou | first3 = F. | last4 = Alm | first4 = G. | last5 = Lefebvre | first5 = J. | last6 = Wattré | first6 = P. | last7 = Hober | first7 = D. | date = Nov 2000 | title = Persistent infection of human pancreatic islets by coxsackievirus B is associated with alpha interferon synthesis in beta cells|url=https://www.ncbi.nlm.nih.gov/pubmed/11024144|journal=Journal of Virology|volume=74|issue=21|pages=10153–10164|issn=0022-538X|pmid=11024144}}</ref><ref name="krogvold2015">{{Cite journal | last = Krogvold | first = Lars | last2 = Edwin | first2 = Bjørn | last3 = Buanes | first3 = Trond | last4 = Frisk | first4 = Gun | last5 = Skog | first5 = Oskar | last6 = Anagandula | first6 = Mahesh | last7 = Korsgren | first7 = Olle | last8 = Undlien | first8 = Dag | last9 = Eike | first9 = Morten C. | date = May 2015 | title = Detection of a low-grade enteroviral infection in the islets of langerhans of living patients newly diagnosed with type 1 diabetes|url=https://www.ncbi.nlm.nih.gov/pubmed/25422108|journal=Diabetes|volume=64|issue=5|pages=1682–1687|doi=10.2337/db14-1370|issn=1939-327X|pmid=25422108}}</ref> which has implications for type 1 [[diabetes]]. * The intestines in ileocecal [[Crohn's disease]].<ref name="Nystrom2013">{{Cite journal | last = Nyström | first = Niklas | last2 = Berg | first2 = Tove | last3 = Lundin | first3 = Elin | last4 = Skog | first4 = Oskar | last5 = Hansson | first5 = Inga | last6 = Frisk | first6 = Gun | last7 = Juko-Pecirep | first7 = Ivana | last8 = Nilsson | first8 = Mats | last9 = Gyllensten | first9 = Ulf | date = 2013-06-27 | title = Human enterovirus species B in ileocecal Crohn's disease|url=https://www.ncbi.nlm.nih.gov/pubmed/23804031|journal=Clinical and Translational Gastroenterology|volume=4| pages = e38|doi=10.1038/ctg.2013.7|issn=2155-384X|pmc=3696939|pmid=23804031}}</ref> * The heart valve tissues in valvular heart disease.<ref name="Burch1967">{{Cite journal | last = Burch | first = G. E. | last2 = Sun | first2 = S.C. | last3 = Colcolough | first3 = H.L. | last4 = Sohal | first4 = R.S. | last5 = DePasquale | first5 = N.P. | date = Jul 1967 | title = Coxsackie B viral myocarditis and valvulitis identified in routine autopsy specimens by immunofluorescent techniques|url=https://www.ncbi.nlm.nih.gov/pubmed/5338726|journal=American Heart Journal|volume=74|issue=1|pages=13–23|issn=0002-8703|pmid=5338726}}</ref><ref name="Li2002">{{Cite journal | last = Li | first = Y. | last2 = Pan | first2 = Z. | last3 = Ji | first3 = Y. | last4 = Peng | first4 = T. | last5 = Archard | first5 = L.C. | last6 = Zhang | first6 = H. | date = 2002-04-01 | title = Enterovirus replication in valvular tissue from patients with chronic rheumatic heart disease|url=https://academic.oup.com/eurheartj/article/23/7/567/457078|journal=European Heart Journal|language=en|volume=23|issue=7 | pages = 567–573|doi=10.1053/euhj.2001.2837|issn=0195-668X}}</ref> * The calf muscle in peripheral arterial disease.<ref name="Kim2013">{{Cite journal | last = Kim | first = Julian K. S. | last2 = Zhu | first2 = Zhen | last3 = Casale | first3 = George | last4 = Koutakis | first4 = Panagiotis | last5 = McComb | first5 = Rodney D. | last6 = Swanson | first6 = Stanley | last7 = Thompson | first7 = Jonathan | last8 = Miserlis | first8 = Dimitrios | last9 = Johanning | first9 = Jason M. | date = 2013-08-06 | title = Human enterovirus in the gastrocnemius of patients with peripheral arterial disease|url=https://www.ncbi.nlm.nih.gov/pubmed/23920231|journal=Journal of the American Heart Association|volume=2|issue=4| pages = e000082|doi=10.1161/JAHA.113.000082|issn=2047-9980|pmc=3828788|pmid=23920231}}</ref> * The stomach tissues in functional dyspepsia and chronic gastritis.<ref name="Chia2015" /> The persistent enterovirus infections found in the above diseases are in several cases (those shown in italics) explicitly demonstrated to be non-cytolytic. Several researchers including Prof [[Nora Chapman]], Prof Steven Tracy and Dr [[John Chia]] theorize that persistent non-cytolytic enterovirus infection may be a cause of ME/CFS. Persistent enterovirus is being investigated as a possible cause for the other above-listed diseases as well. Non-cytolytic enterovirus is also known as non-cytopathic enterovirus. [[Cytolytic]] refers to destruction of a cell by rupture of its membrane, and cytopathic signifies damage to a cell, typically by rupture. Thus the terms non-cytolytic and non-cytopathic indicate this mutated enterovirus does not usually destroy the cells in which it resides. Non-cytolytic enterovirus may also be referred to as defective enterovirus and terminally deleted enterovirus, terms which refer to the mutations (deletions) in its genome which render this virus defective and aberrant.
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