{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,28]],"date-time":"2026-07-28T03:17:15Z","timestamp":1785208635892,"version":"3.55.0"},"reference-count":58,"publisher":"Oxford University Press (OUP)","issue":"5","license":[{"start":{"date-parts":[[2016,9,1]],"date-time":"2016-09-01T00:00:00Z","timestamp":1472688000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/pages\/standard-publication-reuse-rights"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2016,9,1]]},"abstract":"<jats:title>Abstract<\/jats:title>\n               <jats:p>Post-translational protein modifications such as citrullination have been linked to the breach of immune tolerance and clinical autoimmunity. Previous studies from our laboratory support this concept, demonstrating that autoantibodies targeting citrullinated isoforms of heat shock protein 90 (HSP90) are associated with rheumatoid arthritis complicated by interstitial lung disease. To further explore the relationship between citrullination and structural determinants of HSP90 immunogenicity, we employed a combination of ELISA-based epitope profiling, computational modeling, and mass-spectrometric sequencing of peptidylarginine deiminase (PAD)-modified protein. Remarkably, ELISAs involving selected citrullinated HSP90\u03b2\/\u03b1 peptides identified a key epitope corresponding to an internal Arg residue (R502 [HSP90\u03b2]\/R510 [HSP90\u03b1]) that is normally buried within the crystal structure of native\/unmodified HSP90. In vitro time\/dose-response experiments reveal an ordered pattern of PAD-mediated deimination events culminating in citrullination of R502\/R510. Conventional as well as scaled molecular dynamics simulations further demonstrate that citrullination of selected Arg residues leads to progressive disruption of HSP90 tertiary structure, promoting exposure of R502\/R510 to PAD modification and subsequent autoantibody binding. Consistent with this process, ELISAs incorporating variably deiminated HSP90 as substrate Ag indicate a direct relationship between the degree of citrullination and the level of ex vivo Ab recognition. Overall, these data support a novel structural paradigm whereby citrullination-induced shifts in protein structure generate cryptic epitopes capable of bypassing B cell tolerance in the appropriate genetic context.<\/jats:p>","DOI":"10.4049\/jimmunol.1600162","type":"journal-article","created":{"date-parts":[[2016,7,23]],"date-time":"2016-07-23T02:59:27Z","timestamp":1469242767000},"page":"1926-1936","source":"Crossref","is-referenced-by-count":30,"title":["Extensive Citrullination Promotes Immunogenicity of HSP90 through Protein Unfolding and Exposure of Cryptic Epitopes"],"prefix":"10.1093","volume":"197","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-0105-3844","authenticated-orcid":false,"given":"Timothy S","family":"Travers","sequence":"first","affiliation":[{"name":"Department of Computational and Systems Biology, University of Pittsburgh School of Medicine , Pittsburgh, PA 15213"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lisa","family":"Harlow","sequence":"additional","affiliation":[{"name":"Division of Rheumatology, Department of Medicine, University of Miami Miller School of Medicine , Miami, FL 33136"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4622-618X","authenticated-orcid":false,"given":"Ivan O","family":"Rosas","sequence":"additional","affiliation":[{"name":"Division of Pulmonary and Critical Care Medicine, Department of Medicine, Brigham and Women\u2019s Hospital , Boston, MA 02115"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Bernadette R","family":"Gochuico","sequence":"additional","affiliation":[{"name":"Medical Genetics Branch, National Human Genome Research Institute, National Institutes of Health , Bethesda, MD 20892"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0897-2272","authenticated-orcid":false,"given":"Ted R","family":"Mikuls","sequence":"additional","affiliation":[{"name":"Division of Rheumatology, Department of Medicine, University of Nebraska Medical Center , Omaha, NE 68198"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3759-647X","authenticated-orcid":false,"given":"Sanjoy K","family":"Bhattacharya","sequence":"additional","affiliation":[{"name":"Department of Ophthalmology, University of Miami Miller School of Medicine , Miami, FL 33136"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Carlos J","family":"Camacho","sequence":"additional","affiliation":[{"name":"Department of Computational and Systems Biology, University of Pittsburgh School of Medicine , Pittsburgh, PA 15213"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1387-0860","authenticated-orcid":false,"given":"Dana P","family":"Ascherman","sequence":"additional","affiliation":[{"name":"Division of Rheumatology, Department of Medicine, University of Miami Miller School of Medicine , Miami, FL 33136"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2016,9,1]]},"reference":[{"key":"2025030617474966900_r1","doi-asserted-by":"crossref","first-page":"590","DOI":"10.1038\/nature03724","article-title":"Cellular and genetic mechanisms of self tolerance and autoimmunity","volume":"435","author":"Goodnow","year":"2005","journal-title":"Nature"},{"key":"2025030617474966900_r2","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1016\/j.jaut.2013.05.003","article-title":"Rethinking mechanisms of autoimmune pathogenesis","volume":"45","author":"Pillai","year":"2013","journal-title":"J. Autoimmun."},{"key":"2025030617474966900_r3","doi-asserted-by":"crossref","first-page":"753","DOI":"10.1016\/j.coi.2004.09.001","article-title":"Post-translational modifications of self antigens: implications for autoimmunity","volume":"16","author":"Anderton","year":"2004","journal-title":"Curr. Opin. Immunol."},{"key":"2025030617474966900_r4","doi-asserted-by":"crossref","first-page":"19","DOI":"10.1002\/prca.201300068","article-title":"Immune response to chemically modified proteome","volume":"8","author":"Bhat","year":"2014","journal-title":"Proteomics Clin. Appl."},{"key":"2025030617474966900_r5","doi-asserted-by":"crossref","first-page":"443","DOI":"10.1016\/S1471-4906(01)01976-7","article-title":"Post-translational protein modifications in antigen recognition and autoimmunity","volume":"22","author":"Doyle","year":"2001","journal-title":"Trends Immunol."},{"key":"2025030617474966900_r6","doi-asserted-by":"crossref","first-page":"490","DOI":"10.1016\/j.autrev.2015.01.013","article-title":"Citrullination and autoimmunity","volume":"14","author":"Valesini","year":"2015","journal-title":"Autoimmun. Rev."},{"key":"2025030617474966900_r7","doi-asserted-by":"crossref","first-page":"2205","DOI":"10.1056\/NEJMra1004965","article-title":"The pathogenesis of rheumatoid arthritis","volume":"365","author":"McInnes","year":"2011","journal-title":"N. Engl. J. Med."},{"key":"2025030617474966900_r8","doi-asserted-by":"crossref","first-page":"1583","DOI":"10.1002\/art.27405","article-title":"Incidence and mortality of interstitial lung disease in rheumatoid arthritis: a population-based study","volume":"62","author":"Bongartz","year":"2010","journal-title":"Arthritis Rheum."},{"key":"2025030617474966900_r9","doi-asserted-by":"crossref","first-page":"443","DOI":"10.1513\/pats.200703-045MS","article-title":"Rheumatoid lung disease","volume":"4","author":"Brown","year":"2007","journal-title":"Proc. Am. Thorac. Soc."},{"key":"2025030617474966900_r10","doi-asserted-by":"crossref","first-page":"961","DOI":"10.1016\/j.clinbiochem.2006.06.004","article-title":"Anti-cyclic citrullinated peptide (CCP) antibodies in patients with long-standing rheumatoid arthritis and their relationship with extra-articular manifestations","volume":"39","author":"Korkmaz","year":"2006","journal-title":"Clin. Biochem."},{"key":"2025030617474966900_r11","doi-asserted-by":"crossref","first-page":"871","DOI":"10.1016\/j.berh.2007.05.003","article-title":"Mortality in established rheumatoid arthritis","volume":"21","author":"Naz","year":"2007","journal-title":"Best Pract. Res. Clin. Rheumatol."},{"key":"2025030617474966900_r12","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1080\/03009740310004621","article-title":"Epidemiology of extra-articular manifestations in rheumatoid arthritis","volume":"33","author":"Turesson","year":"2004","journal-title":"Scand. J. Rheumatol."},{"key":"2025030617474966900_r13","doi-asserted-by":"crossref","first-page":"907","DOI":"10.1016\/j.berh.2007.05.007","article-title":"Extra-articular manifestations and complications of rheumatoid arthritis","volume":"21","author":"Young","year":"2007","journal-title":"Best Pract. Res. Clin. Rheumatol."},{"key":"2025030617474966900_r14","doi-asserted-by":"crossref","first-page":"350","DOI":"10.1093\/rheumatology\/kel253","article-title":"Mortality in rheumatoid arthritis: increased in the early course of disease, in ischaemic heart disease and in pulmonary fibrosis","volume":"46","author":"Young","year":"2007","journal-title":"Rheumatology (Oxford)"},{"key":"2025030617474966900_r15","doi-asserted-by":"crossref","first-page":"300","DOI":"10.1097\/BOR.0b013e3282fbd22a","article-title":"Antibodies to citrullinated proteins in arthritis: pathology and promise","volume":"20","author":"Klareskog","year":"2008","journal-title":"Curr. Opin. Rheumatol."},{"key":"2025030617474966900_r16","first-page":"281","article-title":"Anti-cyclic citrullinated peptide antibody in rheumatoid arthritis: relation with disease aggressiveness","volume":"24","author":"del Val del Amo","year":"2006","journal-title":"Clin. Exp. Rheumatol."},{"key":"2025030617474966900_r17","doi-asserted-by":"crossref","first-page":"1090","DOI":"10.1136\/ard.2003.014233","article-title":"Prediction of radiological outcome in early rheumatoid arthritis in clinical practice: role of antibodies to citrullinated peptides (anti-CCP)","volume":"63","author":"Forslind","year":"2004","journal-title":"Ann. Rheum. Dis."},{"key":"2025030617474966900_r18","doi-asserted-by":"crossref","first-page":"1831","DOI":"10.1002\/1529-0131(200008)43:8<1831::AID-ANR19>3.0.CO;2-6","article-title":"The prognostic value of anti-cyclic citrullinated peptide antibody in patients with recent-onset rheumatoid arthritis","volume":"43","author":"Kroot","year":"2000","journal-title":"Arthritis Rheum."},{"key":"2025030617474966900_r19","doi-asserted-by":"crossref","first-page":"1472","DOI":"10.1002\/art.24827","article-title":"Anti-citrullinated peptide antibody assays and their role in the diagnosis of rheumatoid arthritis","volume":"61","author":"Aggarwal","year":"2009","journal-title":"Arthritis Rheum."},{"key":"2025030617474966900_r20","doi-asserted-by":"crossref","first-page":"266","DOI":"10.1016\/j.cca.2012.09.015","article-title":"Third generation anti-citrullinated peptide antibody assay is a sensitive marker in rheumatoid factor negative rheumatoid arthritis","volume":"414","author":"Swart","year":"2012","journal-title":"Clin. Chim. Acta"},{"key":"2025030617474966900_r21","doi-asserted-by":"crossref","first-page":"2569","DOI":"10.1084\/jem.20131241","article-title":"A molecular basis for the association of the HLA-DRB1 locus, citrullination, and rheumatoid arthritis","volume":"210","author":"Scally","year":"2013","journal-title":"J. Exp. Med."},{"key":"2025030617474966900_r22","doi-asserted-by":"crossref","first-page":"1117","DOI":"10.1002\/art.21739","article-title":"The HLA-DRB1 shared epitope alleles are primarily a risk factor for anti-cyclic citrullinated peptide antibodies and are not an independent risk factor for development of rheumatoid arthritis","volume":"54","author":"van der Helm-van Mil","year":"2006","journal-title":"Arthritis Rheum."},{"key":"2025030617474966900_r23","doi-asserted-by":"crossref","first-page":"141","DOI":"10.1038\/nrrheum.2012.237","article-title":"Genetics and epigenetics of rheumatoid arthritis","volume":"9","author":"Viatte","year":"2013","journal-title":"Nat. Rev. Rheumatol."},{"key":"2025030617474966900_r24","doi-asserted-by":"crossref","first-page":"38","DOI":"10.1002\/art.21575","article-title":"A new model for an etiology of rheumatoid arthritis: smoking may trigger HLA-DR (shared epitope)-restricted immune reactions to autoantigens modified by citrullination","volume":"54","author":"Klareskog","year":"2006","journal-title":"Arthritis Rheum."},{"key":"2025030617474966900_r25","doi-asserted-by":"crossref","first-page":"1072","DOI":"10.1136\/ard.2009.114694","article-title":"Silica exposure among male current smokers is associated with a high risk of developing ACPA-positive rheumatoid arthritis","volume":"69","author":"Stolt","year":"2010","journal-title":"Ann. Rheum. Dis."},{"key":"2025030617474966900_r26","doi-asserted-by":"crossref","first-page":"425","DOI":"10.1002\/art.22373","article-title":"The HLA-DRB1 shared epitope alleles differ in the interaction with smoking and predisposition to antibodies to cyclic citrullinated peptide","volume":"56","author":"van der Helm-van Mil","year":"2007","journal-title":"Arthritis Rheum."},{"key":"2025030617474966900_r27","doi-asserted-by":"crossref","first-page":"1488","DOI":"10.1136\/ard.2007.075192","article-title":"Smoking increases peptidylarginine deiminase 2 enzyme expression in human lungs and increases citrullination in BAL cells","volume":"67","author":"Makrygiannakis","year":"2008","journal-title":"Ann. Rheum. Dis."},{"key":"2025030617474966900_r28","doi-asserted-by":"crossref","first-page":"645","DOI":"10.1038\/nrrheum.2014.115","article-title":"Lungs, joints and immunity against citrullinated proteins in rheumatoid arthritis","volume":"10","author":"Catrina","year":"2014","journal-title":"Nat. Rev. Rheumatol."},{"key":"2025030617474966900_r29","doi-asserted-by":"crossref","first-page":"869","DOI":"10.1002\/art.37881","article-title":"Identification of citrullinated hsp90 isoforms as novel autoantigens in rheumatoid arthritis-associated interstitial lung disease","volume":"65","author":"Harlow","year":"2013","journal-title":"Arthritis Rheum."},{"key":"2025030617474966900_r30","doi-asserted-by":"crossref","first-page":"60","DOI":"10.1016\/j.clim.2014.08.004","article-title":"Anti-citrullinated heat shock protein 90 antibodies identified in bronchoalveolar lavage fluid are a marker of lung-specific immune responses","volume":"155","author":"Harlow","year":"2014","journal-title":"Clin. Immunol."},{"key":"2025030617474966900_r31","doi-asserted-by":"crossref","first-page":"30709","DOI":"10.1074\/jbc.271.48.30709","article-title":"Protein unfolding by peptidylarginine deiminase: substrate specificity and structural relationships of the natural substrates trichohyalin and filaggrin","volume":"271","author":"Tarcsa","year":"1996","journal-title":"J. Biol. Chem."},{"key":"2025030617474966900_r32","doi-asserted-by":"crossref","first-page":"5374","DOI":"10.1021\/bi9925569","article-title":"Deimination of myelin basic protein. 1. Effect of deimination of arginyl residues of myelin basic protein on its structure and susceptibility to digestion by cathepsin D","volume":"39","author":"Pritzker","year":"2000","journal-title":"Biochemistry"},{"key":"2025030617474966900_r33","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1016\/j.cccn.2004.07.016","article-title":"Autoantibodies to citrullinated proteins in rheumatoid arthritis: clinical performance and biochemical aspects of an RA-specific marker","volume":"350","author":"Nijenhuis","year":"2004","journal-title":"Clin. Chim. Acta"},{"key":"2025030617474966900_r34","doi-asserted-by":"crossref","first-page":"e20457","DOI":"10.1371\/journal.pone.0020457","article-title":"Myelin basic protein as a novel genetic risk factor in rheumatoid arthritis\u2014a genome-wide study combined with immunological analyses","volume":"6","author":"Terao","year":"2011","journal-title":"PLoS One"},{"key":"2025030617474966900_r35","doi-asserted-by":"crossref","first-page":"315","DOI":"10.1002\/art.1780310302","article-title":"The American Rheumatism Association 1987 revised criteria for the classification of rheumatoid arthritis","volume":"31","author":"Arnett","year":"1988","journal-title":"Arthritis Rheum."},{"key":"2025030617474966900_r36","first-page":"1480","article-title":"The association of race and ethnicity with disease expression in male US veterans with rheumatoid arthritis","volume":"34","author":"Mikuls","year":"2007","journal-title":"J. Rheumatol."},{"key":"2025030617474966900_r37","doi-asserted-by":"crossref","first-page":"159","DOI":"10.1001\/archinternmed.2007.59","article-title":"Progressive preclinical interstitial lung disease in rheumatoid arthritis","volume":"168","author":"Gochuico","year":"2008","journal-title":"Arch. Intern. Med."},{"key":"2025030617474966900_r38","doi-asserted-by":"crossref","first-page":"e19961","DOI":"10.1371\/journal.pone.0019961","article-title":"The hexameric structures of human heat shock protein 90","volume":"6","author":"Lee","year":"2011","journal-title":"PLoS One"},{"key":"2025030617474966900_r39","first-page":"11","article-title":"Optimal alignments in linear space","volume":"4","author":"Myers","year":"1988","journal-title":"Comput. Appl. Biosci."},{"key":"2025030617474966900_r40"},{"key":"2025030617474966900_r41","volume-title":"AMBER 2015.","author":"Case","year":"2015"},{"key":"2025030617474966900_r42","doi-asserted-by":"crossref","first-page":"3878","DOI":"10.1021\/ct400314y","article-title":"Routine microsecond molecular dynamics simulations with AMBER on GPUs. 2. Explicit solvent particle mesh Ewald","volume":"9","author":"Salomon-Ferrer","year":"2013","journal-title":"J. Chem. Theory Comput."},{"key":"2025030617474966900_r43","doi-asserted-by":"crossref","first-page":"3696","DOI":"10.1021\/acs.jctc.5b00255","article-title":"ff14SB: improving the accuracy of protein side chain and backbone parameters from ff99SB","volume":"11","author":"Maier","year":"2015","journal-title":"J. Chem. Theory Comput."},{"key":"2025030617474966900_r44","doi-asserted-by":"crossref","first-page":"e1003154","DOI":"10.1371\/journal.pcbi.1003154","article-title":"A systematic framework for molecular dynamics simulations of protein post-translational modifications","volume":"9","author":"Petrov","year":"2013","journal-title":"PLOS Comput. Biol."},{"key":"2025030617474966900_r45","doi-asserted-by":"crossref","first-page":"335","DOI":"10.1021\/ja00392a016","article-title":"Quantum and statistical mechanical studies of liquids. 10. Transferable intermolecular potential functions for water, alcohols, and ethers: application to liquid water","volume":"103","author":"Jorgensen","year":"1981","journal-title":"J. Am. Chem. Soc."},{"key":"2025030617474966900_r46","doi-asserted-by":"crossref","first-page":"10089","DOI":"10.1063\/1.464397","article-title":"Particle mesh Ewald: an N\u22c5log(N) method for Ewald sums in large systems","volume":"98","author":"Darden","year":"1993","journal-title":"J. Chem. Phys."},{"key":"2025030617474966900_r47","doi-asserted-by":"crossref","first-page":"786","DOI":"10.1002\/(SICI)1096-987X(199906)20:8<786::AID-JCC5>3.0.CO;2-B","article-title":"Improving efficiency of large time-scale molecular dynamics simulations of hydrogen-rich systems","volume":"20","author":"Feenstra","year":"1999","journal-title":"J. Comput. Chem."},{"key":"2025030617474966900_r48","doi-asserted-by":"crossref","first-page":"952","DOI":"10.1002\/jcc.540130805","article-title":"Settle: an analytical version of the SHAKE and RATTLE algorithm for rigid water models","volume":"13","author":"Miyamoto","year":"1992","journal-title":"J. Comput. Chem."},{"key":"2025030617474966900_r49","doi-asserted-by":"crossref","first-page":"327","DOI":"10.1016\/0021-9991(77)90098-5","article-title":"Numerical integration of the cartesian equations of motion of a system with constraints: molecular dynamics of n-alkanes","volume":"23","author":"Ryckaert","year":"1977","journal-title":"J. Comput. Phys."},{"key":"2025030617474966900_r50","doi-asserted-by":"crossref","first-page":"12759","DOI":"10.1021\/jp401587e","article-title":"Population based reweighting of scaled molecular dynamics","volume":"117","author":"Sinko","year":"2013","journal-title":"J. Phys. Chem. B"},{"key":"2025030617474966900_r51","doi-asserted-by":"crossref","first-page":"33","DOI":"10.1016\/0263-7855(96)00018-5","article-title":"VMD: visual molecular dynamics","volume":"14","author":"Humphrey","year":"1996","journal-title":"J. Mol. Graph."},{"key":"2025030617474966900_r52","doi-asserted-by":"crossref","first-page":"3084","DOI":"10.1021\/ct400341p","article-title":"PTRAJ and CPPTRAJ: software for processing and analysis of molecular dynamics trajectory data","volume":"9","author":"Roe","year":"2013","journal-title":"J. Chem. Theory Comput."},{"key":"2025030617474966900_r53","doi-asserted-by":"crossref","first-page":"e1000231","DOI":"10.1371\/journal.pcbi.1000231","article-title":"Structural and thermodynamic approach to peptide immunogenicity","volume":"4","author":"Camacho","year":"2008","journal-title":"PLoS Comput. Biol"},{"key":"2025030617474966900_r54","doi-asserted-by":"crossref","first-page":"146","DOI":"10.4172\/2155-9899.1000146","article-title":"Citrullination: a target for disease intervention in multiple sclerosis and other inflammatory diseases?","volume":"4","author":"Chirivi","year":"2013","journal-title":"J. Clin. Cell. Immunol."},{"key":"2025030617474966900_r55","doi-asserted-by":"crossref","first-page":"1532","DOI":"10.1126\/science.1092385","article-title":"Neutrophil extracellular traps kill bacteria","volume":"303","author":"Brinkmann","year":"2004","journal-title":"Science"},{"key":"2025030617474966900_r56","doi-asserted-by":"crossref","first-page":"178ra40","DOI":"10.1126\/scitranslmed.3005580","article-title":"NETs are a source of citrullinated autoantigens and stimulate inflammatory responses in rheumatoid arthritis","volume":"5","author":"Khandpur","year":"2013","journal-title":"Sci. Transl. Med."},{"key":"2025030617474966900_r57","doi-asserted-by":"crossref","first-page":"R191","DOI":"10.1186\/ar3520","article-title":"Detection of autoantibodies to citrullinated BiP in rheumatoid arthritis patients and pro-inflammatory role of citrullinated BiP in collagen-induced arthritis","volume":"13","author":"Shoda","year":"2011","journal-title":"Arthritis Res. Ther."},{"key":"2025030617474966900_r58","doi-asserted-by":"crossref","first-page":"e8625","DOI":"10.1371\/journal.pone.0008625","article-title":"Crystal structures of the ATPase domains of four human Hsp70 isoforms: HSPA1L\/Hsp70-hom, HSPA2\/Hsp70-2, HSPA6\/Hsp70B\u2032, and HSPA5\/BiP\/GRP78","volume":"5","author":"Wisniewska","year":"2010","journal-title":"PLoS One"}],"container-title":["The Journal of Immunology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/jimmunol\/article-pdf\/197\/5\/1926\/61580519\/1600162.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/jimmunol\/article-pdf\/197\/5\/1926\/61580519\/1600162.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,3,6]],"date-time":"2025-03-06T19:01:36Z","timestamp":1741287696000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/jimmunol\/article\/197\/5\/1926\/7974595"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2016,9,1]]},"references-count":58,"journal-issue":{"issue":"5","published-print":{"date-parts":[[2016,9,1]]}},"URL":"https:\/\/doi.org\/10.4049\/jimmunol.1600162","relation":{},"ISSN":["0022-1767","1550-6606"],"issn-type":[{"value":"0022-1767","type":"print"},{"value":"1550-6606","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2016,9]]},"published":{"date-parts":[[2016,9,1]]}}}