{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,3]],"date-time":"2026-11-03T22:26:37Z","timestamp":1793744797622,"version":"4.6.1"},"reference-count":38,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2019,4,1]],"date-time":"2019-04-01T00:00:00Z","timestamp":1554076800000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100003407","name":"Miur","doi-asserted-by":"crossref","award":["PON projects 01_0129"],"award-info":[{"award-number":["PON projects 01_0129"]}],"id":[{"id":"10.13039\/501100003407","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100003407","name":"Miur","doi-asserted-by":"crossref","award":["Futuro in Ricerca\u201d RBFR12SJA8"],"award-info":[{"award-number":["Futuro in Ricerca\u201d RBFR12SJA8"]}],"id":[{"id":"10.13039\/501100003407","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["acta neuropathol commun"],"published-print":{"date-parts":[[2019,12]]},"DOI":"10.1186\/s40478-019-0707-5","type":"journal-article","created":{"date-parts":[[2019,4,1]],"date-time":"2019-04-01T10:03:34Z","timestamp":1554113014000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":89,"title":["AQP4ex is crucial for the anchoring of AQP4 at the astrocyte end-feet and for neuromyelitis optica antibody binding"],"prefix":"10.1186","volume":"7","author":[{"given":"Claudia","family":"Palazzo","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Cinzia","family":"Buccoliero","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maria Grazia","family":"Mola","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Pasqua","family":"Abbrescia","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Grazia Paola","family":"Nicchia","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maria","family":"Trojano","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5284-2934","authenticated-orcid":false,"given":"Antonio","family":"Frigeri","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2019,4,1]]},"reference":[{"key":"707_CR1","doi-asserted-by":"publisher","first-page":"113","DOI":"10.1083\/jcb.200106158","volume":"155","author":"ME Adams","year":"2001","unstructured":"Adams ME, Mueller HA, Froehner SC (2001) In vivo requirement of the alpha-syntrophin PDZ domain for the sarcolemmal localization of nNOS and aquaporin-4. J Cell Biol 155:113\u2013122. https:\/\/doi.org\/10.1083\/jcb.200106158","journal-title":"J Cell Biol"},{"key":"707_CR2","doi-asserted-by":"publisher","first-page":"999","DOI":"10.1016\/j.neuroscience.2004.08.049","volume":"129","author":"M Amiry-Moghaddam","year":"2004","unstructured":"Amiry-Moghaddam M, Frydenlund DS, Ottersen OP (2004) Anchoring of aquaporin-4 in brain: molecular mechanisms and implications for the physiology and pathophysiology of water transport. Neuroscience 129:999\u20131010. https:\/\/doi.org\/10.1016\/j.neuroscience.2004.08.049","journal-title":"Neuroscience"},{"key":"707_CR3","doi-asserted-by":"publisher","first-page":"2106","DOI":"10.1073\/pnas.0437946100","volume":"100","author":"M Amiry-Moghaddam","year":"2003","unstructured":"Amiry-Moghaddam M, Otsuka T, Hurn PD, Traystman RJ, Haug FM, Froehner SC, Adams ME, Neely JD, Agre P, Ottersen OP et al (2003) An alpha-syntrophin-dependent pool of AQP4 in astroglial end-feet confers bidirectional water flow between blood and brain. Proc Natl Acad Sci U S A 100:2106\u20132111. https:\/\/doi.org\/10.1073\/pnas.0437946100","journal-title":"Proc Natl Acad Sci U S A"},{"key":"707_CR4","doi-asserted-by":"publisher","first-page":"879","DOI":"10.1002\/glia.20347","volume":"53","author":"AD Bragg","year":"2006","unstructured":"Bragg AD, Amiry-Moghaddam M, Ottersen OP, Adams ME, Froehner SC (2006) Assembly of a perivascular astrocyte protein scaffold at the mammalian blood-brain barrier is dependent on alpha-syntrophin. Glia 53:879\u2013890. https:\/\/doi.org\/10.1002\/glia.20347","journal-title":"Glia"},{"key":"707_CR5","doi-asserted-by":"publisher","first-page":"28387","DOI":"10.1074\/jbc.M402604200","volume":"279","author":"NC Connors","year":"2004","unstructured":"Connors NC, Adams ME, Froehner SC, Kofuji P (2004) The potassium channel Kir4.1 associates with the dystrophin-glycoprotein complex via alpha-syntrophin in glia. J Biol Chem 279:28387\u201328392. https:\/\/doi.org\/10.1074\/jbc.M402604200","journal-title":"J Biol Chem"},{"key":"707_CR6","doi-asserted-by":"publisher","first-page":"35850","DOI":"10.1074\/jbc.M109.071670","volume":"284","author":"JM Crane","year":"2009","unstructured":"Crane JM, Bennett JL, Verkman AS (2009) Live cell analysis of aquaporin-4 m1\/m23 interactions and regulated orthogonal array assembly in glial cells. J Biol Chem 284:35850\u201335860. https:\/\/doi.org\/10.1074\/jbc.M109.071670","journal-title":"J Biol Chem"},{"key":"707_CR7","doi-asserted-by":"publisher","first-page":"16516","DOI":"10.1074\/jbc.M111.227298","volume":"286","author":"JM Crane","year":"2011","unstructured":"Crane JM, Lam C, Rossi A, Gupta T, Bennett JL, Verkman AS (2011) Binding affinity and specificity of neuromyelitis optica autoantibodies to aquaporin-4 M1\/M23 isoforms and orthogonal arrays. J Biol Chem 286:16516\u201316524. https:\/\/doi.org\/10.1074\/jbc.M111.227298","journal-title":"J Biol Chem"},{"key":"707_CR8","doi-asserted-by":"publisher","unstructured":"De Bellis M, Pisani F, Mola MG, Rosito S, Simone L, Buccoliero C, Trojano M, Nicchia GP, Svelto M, Frigeri A (2017) Translational readthrough generates new astrocyte AQP4 isoforms that modulate supramolecular clustering, glial endfeet localization, and water transport. Glia. https:\/\/doi.org\/10.1002\/glia.23126","DOI":"10.1002\/glia.23126"},{"key":"707_CR9","doi-asserted-by":"publisher","unstructured":"De Bellis M, Pisani F, Mola MG, Basco D, Catalano F, Nicchia GP, Svelto M, Frigeri A (2014) A novel human aquaporin-4 splice variant exhibits a dominant-negative activity: a new mechanism to regulate water permeability. Molecular biology of the cell 25:470\u2013480. https:\/\/doi.org\/10.1091\/mbc.E13-06-0331","DOI":"10.1091\/mbc.E13-06-0331"},{"key":"707_CR10","doi-asserted-by":"publisher","first-page":"1605","DOI":"10.1016\/j.cell.2014.04.033","volume":"157","author":"SM Eswarappa","year":"2014","unstructured":"Eswarappa SM, Potdar AA, Koch WJ, Fan Y, Vasu K, Lindner D, Willard B, Graham LM, DiCorleto PE, Fox PL (2014) Programmed translational readthrough generates antiangiogenic VEGF-ax. Cell 157:1605\u20131618. https:\/\/doi.org\/10.1016\/j.cell.2014.04.033","journal-title":"Cell"},{"issue":"Pt 9","key":"707_CR11","doi-asserted-by":"crossref","first-page":"2993","DOI":"10.1242\/jcs.108.9.2993","volume":"108","author":"A Frigeri","year":"1995","unstructured":"Frigeri A, Gropper MA, Umenishi F, Kawashima M, Brown D, Verkman AS (1995) Localization of MIWC and GLIP water channel homologs in neuromuscular, epithelial and glandular tissues. J Cell Sci 108(Pt 9):2993\u20133002","journal-title":"J Cell Sci"},{"key":"707_CR12","doi-asserted-by":"publisher","first-page":"90","DOI":"10.1096\/fj.00-0260com","volume":"15","author":"A Frigeri","year":"2001","unstructured":"Frigeri A, Nicchia GP, Nico B, Quondamatteo F, Herken R, Roncali L, Svelto M (2001) Aquaporin-4 deficiency in skeletal muscle and brain of dystrophic mdx mice. FASEB J 15:90\u201398. https:\/\/doi.org\/10.1096\/fj.00-0260com","journal-title":"FASEB J"},{"key":"707_CR13","doi-asserted-by":"publisher","first-page":"1120","DOI":"10.1096\/fj.01-0797fje","volume":"16","author":"A Frigeri","year":"2002","unstructured":"Frigeri A, Nicchia GP, Repetto S, Bado M, Minetti C, Svelto M (2002) Altered aquaporin-4 expression in human muscular dystrophies: a common feature? FASEB J 16:1120\u20131122. https:\/\/doi.org\/10.1096\/fj.01-0797fje","journal-title":"FASEB J"},{"key":"707_CR14","doi-asserted-by":"publisher","first-page":"695","DOI":"10.1172\/JCI2545","volume":"102","author":"A Frigeri","year":"1998","unstructured":"Frigeri A, Nicchia GP, Verbavatz JM, Valenti G, Svelto M (1998) Expression of aquaporin-4 in fast-twitch fibers of mammalian skeletal muscle. J Clin Invest 102:695\u2013703. https:\/\/doi.org\/10.1172\/JCI2545","journal-title":"J Clin Invest"},{"key":"707_CR15","doi-asserted-by":"publisher","first-page":"13609","DOI":"10.1073\/pnas.2235843100","volume":"100","author":"CS Furman","year":"2003","unstructured":"Furman CS, Gorelick-Feldman DA, Davidson KG, Yasumura T, Neely JD, Agre P, Rash JE (2003) Aquaporin-4 square array assembly: opposing actions of M1 and M23 isoforms. Proc Natl Acad Sci U S A 100:13609\u201313614. https:\/\/doi.org\/10.1073\/pnas.2235843100","journal-title":"Proc Natl Acad Sci U S A"},{"key":"707_CR16","doi-asserted-by":"publisher","first-page":"21","DOI":"10.1016\/j.jaut.2012.07.008","volume":"40","author":"R Iorio","year":"2013","unstructured":"Iorio R, Fryer JP, Hinson SR, Fallier-Becker P, Wolburg H, Pittock SJ, Lennon VA (2013) Astrocytic autoantibody of neuromyelitis optica (NMO-IgG) binds to aquaporin-4 extracellular loops, monomers, tetramers and high order arrays. J Autoimmun 40:21\u201327. https:\/\/doi.org\/10.1016\/j.jaut.2012.07.008","journal-title":"J Autoimmun"},{"key":"707_CR17","doi-asserted-by":"publisher","first-page":"129","DOI":"10.1186\/1742-2094-11-129","volume":"11","author":"S Jarius","year":"2014","unstructured":"Jarius S, Paul F, Fechner K, Ruprecht K, Kleiter I, Franciotta D, Ringelstein M, Pache F, Aktas O, Wildemann B (2014) Aquaporin-4 antibody testing: direct comparison of M1-AQP4-DNA-transfected cells with leaky scanning versus M23-AQP4-DNA-transfected cells as antigenic substrate. J Neuroinflammation 11:129. https:\/\/doi.org\/10.1186\/1742-2094-11-129","journal-title":"J Neuroinflammation"},{"key":"707_CR18","doi-asserted-by":"publisher","first-page":"505","DOI":"10.1146\/annurev-biophys-062215-010822","volume":"46","author":"F Jiang","year":"2017","unstructured":"Jiang F, Doudna JA (2017) CRISPR-Cas9 structures and mechanisms. Annu Rev Biophys 46:505\u2013529. https:\/\/doi.org\/10.1146\/annurev-biophys-062215-010822","journal-title":"Annu Rev Biophys"},{"key":"707_CR19","doi-asserted-by":"publisher","first-page":"2936","DOI":"10.1016\/j.bpj.2011.05.012","volume":"100","author":"BJ Jin","year":"2011","unstructured":"Jin BJ, Rossi A, Verkman AS (2011) Model of aquaporin-4 supramolecular assembly in orthogonal arrays based on heterotetrameric association of M1-M23 isoforms. Biophys J 100:2936\u20132945. https:\/\/doi.org\/10.1016\/j.bpj.2011.05.012","journal-title":"Biophys J"},{"key":"707_CR20","doi-asserted-by":"publisher","first-page":"2106","DOI":"10.1016\/S0140-6736(04)17551-X","volume":"364","author":"VA Lennon","year":"2004","unstructured":"Lennon VA, Wingerchuk DM, Kryzer TJ, Pittock SJ, Lucchinetti CF, Fujihara K, Nakashima I, Weinshenker BG (2004) A serum autoantibody marker of neuromyelitis optica: distinction from multiple sclerosis. Lancet 364:2106\u20132112. https:\/\/doi.org\/10.1016\/S0140-6736(04)17551-X","journal-title":"Lancet"},{"key":"707_CR21","doi-asserted-by":"publisher","first-page":"8928","DOI":"10.1093\/nar\/gku608","volume":"42","author":"G Loughran","year":"2014","unstructured":"Loughran G, Chou MY, Ivanov IP, Jungreis I, Kellis M, Kiran AM, Baranov PV, Atkins JF (2014) Evidence of efficient stop codon readthrough in four mammalian genes. Nucleic Acids Res 42:8928\u20138938. https:\/\/doi.org\/10.1093\/nar\/gku608","journal-title":"Nucleic Acids Res"},{"key":"707_CR22","doi-asserted-by":"publisher","first-page":"8219","DOI":"10.1021\/ac901526k","volume":"81","author":"MG Mola","year":"2009","unstructured":"Mola MG, Nicchia GP, Svelto M, Spray DC, Frigeri A (2009) Automated cell-based assay for screening of aquaporin inhibitors. Anal Chem 81:8219\u20138229. https:\/\/doi.org\/10.1021\/ac901526k","journal-title":"Anal Chem"},{"key":"707_CR23","doi-asserted-by":"publisher","first-page":"1543","DOI":"10.1152\/physrev.00011.2013","volume":"93","author":"EA Nagelhus","year":"2013","unstructured":"Nagelhus EA, Ottersen OP (2013) Physiological roles of aquaporin-4 in brain. Physiol Rev 93:1543\u20131562. https:\/\/doi.org\/10.1152\/physrev.00011.2013","journal-title":"Physiol Rev"},{"key":"707_CR24","doi-asserted-by":"publisher","first-page":"14108","DOI":"10.1073\/pnas.241508198","volume":"98","author":"JD Neely","year":"2001","unstructured":"Neely JD, Amiry-Moghaddam M, Ottersen OP, Froehner SC, Agre P, Adams ME (2001) Syntrophin-dependent expression and localization of Aquaporin-4 water channel protein. Proc Natl Acad Sci U S A 98:14108\u201314113. https:\/\/doi.org\/10.1073\/pnas.241508198","journal-title":"Proc Natl Acad Sci U S A"},{"key":"707_CR25","doi-asserted-by":"publisher","first-page":"2156","DOI":"10.1111\/j.1471-4159.2008.05302.x","volume":"105","author":"GP Nicchia","year":"2008","unstructured":"Nicchia GP, Cogotzi L, Rossi A, Basco D, Brancaccio A, Svelto M, Frigeri A (2008) Expression of multiple AQP4 pools in the plasma membrane and their association with the dystrophin complex. J Neurochem 105:2156\u20132165. https:\/\/doi.org\/10.1111\/j.1471-4159.2008.05302.x","journal-title":"J Neurochem"},{"key":"707_CR26","doi-asserted-by":"publisher","first-page":"1508","DOI":"10.1096\/fj.02-1183fje","volume":"17","author":"GP Nicchia","year":"2003","unstructured":"Nicchia GP, Frigeri A, Liuzzi GM, Svelto M (2003) Inhibition of aquaporin-4 expression in astrocytes by RNAi determines alteration in cell morphology, growth, and water transport and induces changes in ischemia-related genes. FASEB J 17:1508\u20131510. https:\/\/doi.org\/10.1096\/fj.02-1183fje","journal-title":"FASEB J"},{"key":"707_CR27","doi-asserted-by":"publisher","first-page":"1363","DOI":"10.1002\/glia.20855","volume":"57","author":"GP Nicchia","year":"2009","unstructured":"Nicchia GP, Mastrototaro M, Rossi A, Pisani F, Tortorella C, Ruggieri M, Lia A, Trojano M, Frigeri A, Svelto M (2009) Aquaporin-4 orthogonal arrays of particles are the target for neuromyelitis optica autoantibodies. Glia 57:1363\u20131373. https:\/\/doi.org\/10.1002\/glia.20855","journal-title":"Glia"},{"key":"707_CR28","doi-asserted-by":"publisher","first-page":"903","DOI":"10.1016\/j.neuroscience.2010.02.008","volume":"168","author":"GP Nicchia","year":"2010","unstructured":"Nicchia GP, Rossi A, Mola MG, Pisani F, Stigliano C, Basco D, Mastrototaro M, Svelto M, Frigeri A (2010) Higher order structure of aquaporin-4. Neuroscience 168:903\u2013914. https:\/\/doi.org\/10.1016\/j.neuroscience.2010.02.008","journal-title":"Neuroscience"},{"key":"707_CR29","doi-asserted-by":"publisher","first-page":"1674","DOI":"10.1096\/fj.04-3281fje","volume":"19","author":"GP Nicchia","year":"2005","unstructured":"Nicchia GP, Srinivas M, Li W, Brosnan CF, Frigeri A, Spray DC (2005) New possible roles for aquaporin-4 in astrocytes: cell cytoskeleton and functional relationship with connexin43. FASEB J 19:1674\u20131676. https:\/\/doi.org\/10.1096\/fj.04-3281fje","journal-title":"FASEB J"},{"key":"707_CR30","doi-asserted-by":"publisher","first-page":"171","DOI":"10.1523\/JNEUROSCI.17-01-00171.1997","volume":"17","author":"S Nielsen","year":"1997","unstructured":"Nielsen S, Nagelhus EA, Amiry-Moghaddam M, Bourque C, Agre P, Ottersen OP (1997) Specialized membrane domains for water transport in glial cells: high-resolution immunogold cytochemistry of aquaporin-4 in rat brain. J Neurosci 17:171\u2013180","journal-title":"J Neurosci"},{"key":"707_CR31","doi-asserted-by":"publisher","first-page":"265","DOI":"10.1038\/nrn3468","volume":"14","author":"MC Papadopoulos","year":"2013","unstructured":"Papadopoulos MC, Verkman AS (2013) Aquaporin water channels in the nervous system. Nat Rev Neurosci 14:265\u2013277. https:\/\/doi.org\/10.1038\/nrn3468","journal-title":"Nat Rev Neurosci"},{"key":"707_CR32","doi-asserted-by":"publisher","first-page":"e79185","DOI":"10.1371\/journal.pone.0079185","volume":"8","author":"F Pisani","year":"2013","unstructured":"Pisani F, Sparaneo A, Tortorella C, Ruggieri M, Trojano M, Mola MG, Nicchia GP, Frigeri A, Svelto M (2013) Aquaporin-4 autoantibodies in Neuromyelitis Optica: AQP4 isoform-dependent sensitivity and specificity. PLoS One 8:e79185. https:\/\/doi.org\/10.1371\/journal.pone.0079185","journal-title":"PLoS One"},{"key":"707_CR33","doi-asserted-by":"publisher","first-page":"1236","DOI":"10.1111\/jcmm.13401","volume":"22","author":"S Rosito","year":"2018","unstructured":"Rosito S, Nicchia GP, Palazzo C, Lia A, Buccoliero C, Pisani F, Svelto M, Trojano M, Frigeri A (2018) Supramolecular aggregation of aquaporin-4 is different in muscle and brain: correlation with tissue susceptibility in neuromyelitis optica. J Cell Mol Med 22:1236\u20131246. https:\/\/doi.org\/10.1111\/jcmm.13401","journal-title":"J Cell Mol Med"},{"key":"707_CR34","doi-asserted-by":"publisher","first-page":"559","DOI":"10.1083\/jcb.201308118","volume":"204","author":"AJ Smith","year":"2014","unstructured":"Smith AJ, Jin BJ, Ratelade J, Verkman AS (2014) Aggregation state determines the localization and function of M1- and M23-aquaporin-4 in astrocytes. J Cell Biol 204:559\u2013573. https:\/\/doi.org\/10.1083\/jcb.201308118","journal-title":"J Cell Biol"},{"key":"707_CR35","doi-asserted-by":"publisher","first-page":"2511","DOI":"10.1016\/j.bpj.2015.10.047","volume":"109","author":"AJ Smith","year":"2015","unstructured":"Smith AJ, Verkman AS (2015) Superresolution imaging of Aquaporin-4 cluster size in antibody-stained paraffin brain sections. Biophys J 109:2511\u20132522. https:\/\/doi.org\/10.1016\/j.bpj.2015.10.047","journal-title":"Biophys J"},{"key":"707_CR36","doi-asserted-by":"publisher","first-page":"1161","DOI":"10.1080\/14728222.2017.1398236","volume":"21","author":"AS Verkman","year":"2017","unstructured":"Verkman AS, Smith AJ, Phuan PW, Tradtrantip L, Anderson MO (2017) The aquaporin-4 water channel as a potential drug target in neurological disorders. Expert Opin Ther Targets 21:1161\u20131170. https:\/\/doi.org\/10.1080\/14728222.2017.1398236","journal-title":"Expert Opin Ther Targets"},{"key":"707_CR37","doi-asserted-by":"publisher","first-page":"17765","DOI":"10.1074\/jbc.M111.314468","volume":"287","author":"Y Yamaguchi","year":"2012","unstructured":"Yamaguchi Y, Hayashi A, Campagnoni CW, Kimura A, Inuzuka T, Baba H (2012) L-MPZ, a novel isoform of myelin P0, is produced by stop codon readthrough. J Biol Chem 287:17765\u201317776. https:\/\/doi.org\/10.1074\/jbc.M111.314468","journal-title":"J Biol Chem"},{"key":"707_CR38","doi-asserted-by":"publisher","first-page":"e110","DOI":"10.1212\/NXI.0000000000000110","volume":"2","author":"A Zekeridou","year":"2015","unstructured":"Zekeridou A, Lennon VA (2015) Aquaporin-4 autoimmunity. Neurol Neuroimmunol Neuroinflamm 2:e110. https:\/\/doi.org\/10.1212\/NXI.0000000000000110","journal-title":"Neurol Neuroimmunol Neuroinflamm"}],"container-title":["Acta Neuropathologica Communications"],"original-title":[],"language":"en","link":[{"URL":"http:\/\/link.springer.com\/content\/pdf\/10.1186\/s40478-019-0707-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"http:\/\/link.springer.com\/article\/10.1186\/s40478-019-0707-5\/fulltext.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"http:\/\/link.springer.com\/content\/pdf\/10.1186\/s40478-019-0707-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,9,14]],"date-time":"2022-09-14T19:57:33Z","timestamp":1663185453000},"score":1,"resource":{"primary":{"URL":"https:\/\/actaneurocomms.biomedcentral.com\/articles\/10.1186\/s40478-019-0707-5"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2019,4,1]]},"references-count":38,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2019,12]]}},"alternative-id":["707"],"URL":"https:\/\/doi.org\/10.1186\/s40478-019-0707-5","relation":{},"ISSN":["2051-5960"],"issn-type":[{"value":"2051-5960","type":"electronic"}],"subject":[],"published":{"date-parts":[[2019,4,1]]},"assertion":[{"value":"29 January 2019","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"21 March 2019","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"1 April 2019","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"This research was performed in compliance with institutional guidelines and approved by the appropriate institutional committees. All applicable international, national, and\/or institutional guidelines for the care and use of animals were followed.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethics approval and consent to participate"}},{"value":"Not applicable.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent for publication"}},{"value":"The authors declare that they have no competing interests.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}},{"value":"Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.","order":4,"name":"Ethics","group":{"name":"EthicsHeading","label":"Publisher\u2019s Note"}}],"article-number":"51"}}