{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,7]],"date-time":"2026-10-07T05:41:44Z","timestamp":1791351704421,"version":"4.3.1"},"reference-count":62,"publisher":"Public Library of Science (PLoS)","issue":"11","license":[{"start":{"date-parts":[[2012,11,1]],"date-time":"2012-11-01T00:00:00Z","timestamp":1351728000000},"content-version":"unspecified","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/publicdomain\/zero\/1.0\/"}],"content-domain":{"domain":["www.plospathogens.org"],"crossmark-restriction":false},"short-container-title":["PLoS Pathog"],"DOI":"10.1371\/journal.ppat.1002998","type":"journal-article","created":{"date-parts":[[2012,11,1]],"date-time":"2012-11-01T21:25:26Z","timestamp":1351805126000},"page":"e1002998","update-policy":"https:\/\/doi.org\/10.1371\/journal.ppat.corrections_policy","source":"Crossref","is-referenced-by-count":221,"title":["Identification of a Novel Splice Variant Form of the Influenza A Virus M2 Ion Channel with an Antigenically Distinct Ectodomain"],"prefix":"10.1371","volume":"8","author":[{"given":"Helen M.","family":"Wise","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Edward C.","family":"Hutchinson","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Brett W.","family":"Jagger","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Amanda D.","family":"Stuart","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zi H.","family":"Kang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nicole","family":"Robb","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Louis M.","family":"Schwartzman","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"John C.","family":"Kash","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ervin","family":"Fodor","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Andrew E.","family":"Firth","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Julia R.","family":"Gog","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jeffery K.","family":"Taubenberger","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Paul","family":"Digard","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2012,11,1]]},"reference":[{"key":"ref1","doi-asserted-by":"crossref","first-page":"402","DOI":"10.1016\/j.cell.2009.01.029","article-title":"The influenza virus enigma","volume":"136","author":"R Salomon","year":"2009","journal-title":"Cell"},{"key":"ref2","doi-asserted-by":"crossref","first-page":"280","DOI":"10.1016\/j.micinf.2010.01.001","article-title":"Research and development of universal influenza vaccines","volume":"12","author":"L Du","year":"2010","journal-title":"Microbes Infect"},{"key":"ref3","doi-asserted-by":"crossref","first-page":"6280","DOI":"10.1016\/j.vaccine.2009.07.007","article-title":"M2e-based universal influenza A vaccine","volume":"27","author":"W Fiers","year":"2009","journal-title":"Vaccine"},{"key":"ref4","doi-asserted-by":"crossref","first-page":"8021","DOI":"10.1128\/JVI.00826-09","article-title":"A complicated message: Identification of a novel PB1-related protein translated from influenza A virus segment 2 mRNA","volume":"83","author":"HM Wise","year":"2009","journal-title":"J Virol"},{"key":"ref5","doi-asserted-by":"crossref","first-page":"199","DOI":"10.1126\/science.1222213","article-title":"An overlapping protein-coding region in influenza A virus segment 3 modulates the host response","volume":"337","author":"BW Jagger","year":"2012","journal-title":"Science"},{"key":"ref6","doi-asserted-by":"crossref","first-page":"1306","DOI":"10.1038\/nm1201-1306","article-title":"A novel influenza A virus mitochondrial protein that induces cell death","volume":"7","author":"W Chen","year":"2001","journal-title":"Nat Med"},{"key":"ref7","doi-asserted-by":"crossref","first-page":"7775","DOI":"10.1093\/nar\/gkr487","article-title":"Overlapping signals for translational regulation and packaging of influenza A virus segment 2","volume":"39","author":"HM Wise","year":"2011","journal-title":"Nucleic Acids Res"},{"key":"ref8","doi-asserted-by":"crossref","first-page":"3575","DOI":"10.1093\/nar\/8.16.3575","article-title":"Relationship between the messenger RNAs transcribed from two overlapping genes of influenza virus","volume":"8","author":"SC Inglis","year":"1980","journal-title":"Nucleic Acids Res"},{"key":"ref9","doi-asserted-by":"crossref","first-page":"475","DOI":"10.1016\/0092-8674(80)90484-5","article-title":"Sequence of interrupted and uninterrupted mRNAs and cloned DNA coding for the two overlapping nonstructural proteins of influenza virus","volume":"21","author":"RA Lamb","year":"1980","journal-title":"Cell"},{"key":"ref10","doi-asserted-by":"crossref","first-page":"2727","DOI":"10.1093\/nar\/9.12.2727","article-title":"Spliced and unspliced RNAs encoded by virion RNA segment 7 of influenza virus","volume":"9","author":"SC Inglis","year":"1981","journal-title":"Nucleic Acids Res"},{"key":"ref11","doi-asserted-by":"crossref","first-page":"4170","DOI":"10.1073\/pnas.78.7.4170","article-title":"Sequences of mRNAs derived from genome RNA segment 7 of influenza virus: colinear and interrupted mRNAs code for overlapping proteins","volume":"78","author":"RA Lamb","year":"1981","journal-title":"Proc Natl Acad Sci U S A"},{"key":"ref12","doi-asserted-by":"crossref","first-page":"179","DOI":"10.1023\/A:1008024909222","article-title":"A novel spliced transcript of influenza A\/WSN\/33 virus","volume":"17","author":"SR Shih","year":"1998","journal-title":"Virus Genes"},{"key":"ref13","doi-asserted-by":"crossref","first-page":"729","DOI":"10.1016\/0042-6822(81)90317-2","article-title":"Identification of a second protein (M2) encoded by RNA segment 7 of influenza virus","volume":"112","author":"RA Lamb","year":"1981","journal-title":"Virology"},{"key":"ref14","doi-asserted-by":"crossref","first-page":"6324","DOI":"10.1073\/pnas.92.14.6324","article-title":"The choice of alternative 5\u2032 splice sites in influenza virus M1 mRNA is regulated by the viral polymerase complex","volume":"92","author":"SR Shih","year":"1995","journal-title":"Proc Natl Acad Sci U S A"},{"key":"ref15","doi-asserted-by":"crossref","first-page":"10873","DOI":"10.1128\/JVI.00506-08","article-title":"Mutations at alternative 5\u2032 splice sites of M1 mRNA negatively affect influenza A virus viability and growth rate","volume":"82","author":"C Chiang","year":"2008","journal-title":"J Virol"},{"key":"ref16","doi-asserted-by":"crossref","first-page":"3097","DOI":"10.1099\/vir.0.2008\/004739-0","article-title":"The influenza A virus spliced messenger RNA M mRNA3 is not required for viral replication in tissue culture","volume":"89","author":"D Jackson","year":"2008","journal-title":"J Gen Virol"},{"key":"ref17","article-title":"The accumulation of influenza A virus segment 7 spliced mRNAs is regulated by the NS1 protein","author":"NC Robb","year":"2011","journal-title":"J Gen Virol"},{"key":"ref18","doi-asserted-by":"crossref","first-page":"32","DOI":"10.1016\/0042-6822(91)90115-R","article-title":"Influenza virus M2 integral membrane protein is a homotetramer stabilized by formation of disulfide bonds","volume":"183","author":"LJ Holsinger","year":"1991","journal-title":"Virology"},{"key":"ref19","doi-asserted-by":"crossref","first-page":"617","DOI":"10.1016\/0042-6822(91)90075-M","article-title":"Structural characteristics of the M2 protein of influenza A viruses: evidence that it forms a tetrameric channel","volume":"180","author":"RJ Sugrue","year":"1991","journal-title":"Virology"},{"key":"ref20","doi-asserted-by":"crossref","first-page":"591","DOI":"10.1038\/nature06531","article-title":"Structure and mechanism of the M2 proton channel of influenza A virus","volume":"451","author":"JR Schnell","year":"2008","journal-title":"Nature"},{"key":"ref21","doi-asserted-by":"crossref","first-page":"522","DOI":"10.1016\/j.bbamem.2010.04.015","article-title":"Influenza M2 proton channels","volume":"1808","author":"RM Pielak","year":"2011","journal-title":"Biochim Biophys Acta"},{"key":"ref22","doi-asserted-by":"crossref","first-page":"3021","DOI":"10.1002\/j.1460-2075.1985.tb04038.x","article-title":"The molecular basis of the specific anti-influenza action of amantadine","volume":"4","author":"AJ Hay","year":"1985","journal-title":"Embo J"},{"key":"ref23","doi-asserted-by":"crossref","first-page":"517","DOI":"10.1016\/0092-8674(92)90452-I","article-title":"Influenza virus M2 protein has ion channel activity","volume":"69","author":"LH Pinto","year":"1992","journal-title":"Cell"},{"key":"ref24","doi-asserted-by":"crossref","first-page":"61","DOI":"10.1099\/0022-1317-60-1-61","article-title":"Uncoating of a rimantadine-resistant variant of influenza virus in the presence of rimantadine","volume":"60","author":"AG Bukrinskaya","year":"1982","journal-title":"J Gen Virol"},{"key":"ref25","doi-asserted-by":"crossref","first-page":"3469","DOI":"10.1002\/j.1460-2075.1990.tb07555.x","article-title":"Specific structural alteration of the influenza haemagglutinin by amantadine","volume":"9","author":"RJ Sugrue","year":"1990","journal-title":"EMBO J"},{"key":"ref26","doi-asserted-by":"crossref","first-page":"10059","DOI":"10.1128\/JVI.01184-08","article-title":"The influenza virus M2 protein cytoplasmic tail interacts with the M1 protein and influences virus assembly at the site of virus budding","volume":"82","author":"BJ Chen","year":"2008","journal-title":"J Virol"},{"key":"ref27","doi-asserted-by":"crossref","first-page":"8765","DOI":"10.1128\/JVI.00853-10","article-title":"Tyrosines in the influenza A virus M2 protein cytoplasmic tail are critical for production of infectious virus particles","volume":"84","author":"ML Grantham","year":"2010","journal-title":"J Virol"},{"key":"ref28","doi-asserted-by":"crossref","first-page":"5233","DOI":"10.1128\/JVI.00049-06","article-title":"The cytoplasmic tail of the influenza A virus M2 protein plays a role in viral assembly","volume":"80","author":"K Iwatsuki-Horimoto","year":"2006","journal-title":"J Virol"},{"key":"ref29","doi-asserted-by":"crossref","first-page":"3595","DOI":"10.1128\/JVI.79.6.3595-3605.2005","article-title":"The influenza A virus M2 cytoplasmic tail is required for infectious virus production and efficient genome packaging","volume":"79","author":"MF McCown","year":"2005","journal-title":"J Virol"},{"key":"ref30","doi-asserted-by":"crossref","first-page":"8178","DOI":"10.1128\/JVI.00627-06","article-title":"Distinct domains of the influenza a virus M2 protein cytoplasmic tail mediate binding to the M1 protein and facilitate infectious virus production","volume":"80","author":"MF McCown","year":"2006","journal-title":"J Virol"},{"key":"ref31","doi-asserted-by":"crossref","first-page":"5078","DOI":"10.1128\/JVI.00119-10","article-title":"Influenza virus m2 ion channel protein is necessary for filamentous virion formation","volume":"84","author":"JS Rossman","year":"2010","journal-title":"J Virol"},{"key":"ref32","doi-asserted-by":"crossref","first-page":"902","DOI":"10.1016\/j.cell.2010.08.029","article-title":"Influenza virus M2 protein mediates ESCRT-independent membrane scission","volume":"142","author":"JS Rossman","year":"2010","journal-title":"Cell"},{"key":"ref33","doi-asserted-by":"crossref","first-page":"2023","DOI":"10.1083\/jcb.109.5.2023","article-title":"Transposition of domains between the M2 and HN viral membrane proteins results in polypeptides which can adopt more than one membrane orientation","volume":"109","author":"GD Parks","year":"1989","journal-title":"J Cell Biol"},{"key":"ref34","doi-asserted-by":"crossref","first-page":"777","DOI":"10.1016\/0092-8674(91)90507-U","article-title":"Topology of eukaryotic type II membrane proteins: importance of N-terminal positively charged residues flanking the hydrophobic domain","volume":"64","author":"GD Parks","year":"1991","journal-title":"Cell"},{"key":"ref35","doi-asserted-by":"crossref","first-page":"2449","DOI":"10.1128\/JVI.72.3.2449-2455.1998","article-title":"The M2 ectodomain is important for its incorporation into influenza A virions","volume":"72","author":"EK Park","year":"1998","journal-title":"J Virol"},{"key":"ref36","doi-asserted-by":"crossref","first-page":"1447","DOI":"10.1099\/vir.0.80727-0","article-title":"Generation of recombinant influenza A virus without M2 ion-channel protein by introduction of a point mutation at the 5\u2032 end of the viral intron","volume":"86","author":"TK Cheung","year":"2005","journal-title":"J Gen Virol"},{"key":"ref37","doi-asserted-by":"crossref","first-page":"11869","DOI":"10.1128\/JVI.01634-08","article-title":"Mutational analysis of cis-acting RNA signals in segment 7 of influenza A virus","volume":"82","author":"EC Hutchinson","year":"2008","journal-title":"J Virol"},{"key":"ref38","doi-asserted-by":"crossref","first-page":"5656","DOI":"10.1128\/JVI.75.12.5656-5662.2001","article-title":"Influenza A virus can undergo multiple cycles of replication without M2 ion channel activity","volume":"75","author":"T Watanabe","year":"2001","journal-title":"J Virol"},{"key":"ref39","doi-asserted-by":"crossref","first-page":"1391","DOI":"10.1128\/JVI.76.3.1391-1399.2002","article-title":"Influenza a virus M2 ion channel activity is essential for efficient replication in tissue culture","volume":"76","author":"M Takeda","year":"2002","journal-title":"J Virol"},{"key":"ref40","doi-asserted-by":"crossref","first-page":"168","DOI":"10.1016\/j.antiviral.2008.06.002","article-title":"Therapeutic potential of a fully human monoclonal antibody against influenza A virus M2 protein","volume":"80","author":"R Wang","year":"2008","journal-title":"Antiviral Res"},{"key":"ref41","doi-asserted-by":"crossref","first-page":"2762","DOI":"10.1128\/JVI.62.8.2762-2772.1988","article-title":"Influenza A virus M2 protein: monoclonal antibody restriction of virus growth and detection of M2 in virions","volume":"62","author":"SL Zebedee","year":"1988","journal-title":"J Virol"},{"key":"ref42","doi-asserted-by":"crossref","first-page":"2195","DOI":"10.1016\/j.molimm.2005.12.015","article-title":"Fine specificity and sequence of antibodies directed against the ectodomain of matrix protein 2 of influenza A virus","volume":"43","author":"M Zhang","year":"2006","journal-title":"Mol Immunol"},{"key":"ref43","doi-asserted-by":"crossref","first-page":"1617","DOI":"10.1128\/JVI.76.4.1617-1625.2002","article-title":"NS1 protein of influenza A virus down-regulates apoptosis","volume":"76","author":"OP Zhirnov","year":"2002","journal-title":"J Virol"},{"key":"ref44","doi-asserted-by":"crossref","first-page":"5542","DOI":"10.1128\/JVI.66.9.5542-5552.1992","article-title":"Expression of the influenza A virus M2 protein is restricted to apical surfaces of polarized epithelial cells","volume":"66","author":"PG Hughey","year":"1992","journal-title":"J Virol"},{"key":"ref45","doi-asserted-by":"crossref","first-page":"3795","DOI":"10.1093\/nar\/19.14.3795","article-title":"A reappraisal of non-consensus mRNA splice sites","volume":"19","author":"IJ Jackson","year":"1991","journal-title":"Nucleic Acids Res"},{"key":"ref46","doi-asserted-by":"crossref","first-page":"283","DOI":"10.1016\/0092-8674(86)90762-2","article-title":"Point mutations define a sequence flanking the AUG initiator codon that modulates translation by eukaryotic ribosomes","volume":"44","author":"M Kozak","year":"1986","journal-title":"Cell"},{"key":"ref47","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1016\/j.virusres.2004.02.028","article-title":"Efficient generation and growth of influenza virus A\/PR\/8\/34 from eight cDNA fragments","volume":"103","author":"E de Wit","year":"2004","journal-title":"Virus Res"},{"key":"ref48","doi-asserted-by":"crossref","first-page":"301","DOI":"10.1099\/vir.0.027086-0","article-title":"Antiserum against the conserved nine amino acid N-terminal peptide of influenza A virus matrix protein 2 is not immunoprotective","volume":"92","author":"M De Filette","year":"2011","journal-title":"J Gen Virol"},{"key":"ref49","doi-asserted-by":"crossref","first-page":"356","DOI":"10.2307\/1592550","article-title":"Sequence analysis of related low-pathogenic and highly pathogenic H5N2 avian influenza isolates from United States live bird markets and poultry farms from 1983 to 1989","volume":"44","author":"DL Suarez","year":"2000","journal-title":"Avian Dis"},{"key":"ref50","doi-asserted-by":"crossref","first-page":"311","DOI":"10.1016\/j.cimid.2008.01.004","article-title":"The history of avian influenza","volume":"32","author":"B Lupiani","year":"2009","journal-title":"Comp Immunol Microbiol Infect Dis"},{"key":"ref51","doi-asserted-by":"crossref","first-page":"920","DOI":"10.1128\/JVI.68.2.920-926.1994","article-title":"Rescue of vector-expressed fowl plague virus hemagglutinin in biologically active form by acidotropic agents and coexpressed M2 protein","volume":"68","author":"M Ohuchi","year":"1994","journal-title":"J Virol"},{"key":"ref52","doi-asserted-by":"crossref","first-page":"1379","DOI":"10.1101\/gad.1788009","article-title":"Current-generation high-throughput sequencing: deepening insights into mammalian transcriptomes","volume":"23","author":"BJ Blencowe","year":"2009","journal-title":"Genes Dev"},{"key":"ref53","doi-asserted-by":"crossref","first-page":"4143","DOI":"10.1128\/JVI.02606-10","article-title":"A Rab11- and microtubule-dependent mechanism for cytoplasmic transport of influenza A virus viral RNA","volume":"85","author":"MJ Amorim","year":"2011","journal-title":"J Virol"},{"key":"ref54","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1006\/viro.1997.8916","article-title":"The M1 and M2 proteins of influenza A virus are important determinants in filamentous particle formation","volume":"240","author":"PC Roberts","year":"1998","journal-title":"Virology"},{"key":"ref55","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1111\/j.1600-0854.2006.00507.x","article-title":"Nuclear export of influenza A virus mRNAs requires ongoing RNA polymerase II activity","volume":"8","author":"MJ Amorim","year":"2007","journal-title":"Traffic"},{"key":"ref56","doi-asserted-by":"crossref","first-page":"2280","DOI":"10.1099\/vir.0.82809-0","article-title":"Identification of the domains of the influenza A virus M1 matrix protein required for NP binding, oligomerization and incorporation into virions","volume":"88","author":"SL Noton","year":"2007","journal-title":"J Gen Virol"},{"key":"ref57","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1186\/1743-422X-3-63","article-title":"New low-viscosity overlay medium for viral plaque assays","volume":"3","author":"M Matrosovich","year":"2006","journal-title":"Virol J"},{"key":"ref58","first-page":"36","article-title":"Image processing with ImageJ","volume":"11","author":"MD Abramoff","year":"1996","journal-title":"Biophotonics International"},{"key":"ref59","doi-asserted-by":"crossref","first-page":"2331","DOI":"10.1099\/vir.0.022004-0","article-title":"Splicing of influenza A virus NS1 mRNA is independent of the viral NS1 protein","volume":"91","author":"NC Robb","year":"2010","journal-title":"J Gen Virol"},{"key":"ref60","doi-asserted-by":"crossref","first-page":"408","DOI":"10.1128\/JVI.75.1.408-419.2001","article-title":"Interaction of the influenza virus nucleoprotein with the cellular CRM1-mediated nuclear export pathway","volume":"75","author":"D Elton","year":"2001","journal-title":"J Virol"},{"key":"ref61","doi-asserted-by":"crossref","first-page":"1061","DOI":"10.1073\/pnas.86.3.1061","article-title":"Growth restriction of influenza A virus by M2 protein antibody is genetically linked to the M1 protein","volume":"86","author":"SL Zebedee","year":"1989","journal-title":"Proc Natl Acad Sci U S A"},{"key":"ref62","doi-asserted-by":"crossref","first-page":"5415","DOI":"10.1002\/j.1460-2075.1996.tb00925.x","article-title":"Novel exploitation of a nuclear function by influenza virus: the cellular SF2\/ASF splicing factor controls the amount of the essential viral M2 ion channel protein in infected cells","volume":"15","author":"SR Shih","year":"1996","journal-title":"Embo J"}],"container-title":["PLoS Pathogens"],"original-title":[],"language":"en","link":[{"URL":"http:\/\/dx.plos.org\/10.1371\/journal.ppat.1002998","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2020,7,15]],"date-time":"2020-07-15T11:24:32Z","timestamp":1594812272000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.ppat.1002998"}},"subtitle":[],"editor":[{"given":"Andrew","family":"Pekosz","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2012,11,1]]},"references-count":62,"journal-issue":{"issue":"11","published-online":{"date-parts":[[2012,11,1]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.ppat.1002998","relation":{},"ISSN":["1553-7374"],"issn-type":[{"value":"1553-7374","type":"electronic"}],"subject":[],"published":{"date-parts":[[2012,11,1]]}}}