{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,22]],"date-time":"2026-05-22T17:08:28Z","timestamp":1779469708134,"version":"3.53.1"},"reference-count":54,"publisher":"Oxford University Press (OUP)","issue":"3","license":[{"start":{"date-parts":[[2025,12,5]],"date-time":"2025-12-05T00:00:00Z","timestamp":1764892800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"NHMRC of Australia and the Channel 7 Children\u2019s Research Foundation"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,3,17]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Neutrophils are the first line of defense against fungal infection and other microbial pathogens. Through alternative pathway of complement activation in innate immunity, complement fragments (opsonin) generated promote phagocytosis of the yeast forms of fungal pathogens such as Candida albicans. However, when complement levels are limited, through consumption in disease and inflammation or where microbial pathogens employ mechanisms to degrade or inhibit complement components, phagocytosis may still proceed via innate receptors, in the absence of effective opsonisation. Mechanisms of innate phagocytosis of fungi via neutrophils remain ill-defined and controversial. We have addressed this issue and demonstrated that neutrophils show very little phagocytosis of zymosan. Interestingly, activation or priming of neutrophils with phorbol-12-myristate-13-acetate, N-formyl-methionyl-leucyl-phenylalanine, arachidonic acid, tumor necrosis factor, and granulocyte macrophage-colony stimulating factor led to significant phagocytosis of the fungi. The result also demonstrated that priming led to a significant increase in the expression of the neutrophil Dectin-1, a receptor for \u03b2-glucan on zymosan. The increased phagocytosis induced by priming could be blocked with an anti-Dectin-1 antibody as well as the soluble \u03b2-glucan, laminarin, suggesting the importance of Dectin-1 in the phagocytosis of zymosan. The priming mechanism for phagocytosis and upregulation of Dectin-1 was examined using pharmacological inhibitors of protein kinase C (PKC) and p38 MAP kinase activation. The results showed that priming occurs in a PKC- p38-dependent as well as a PKC-independent p38-dependent manner. The findings demonstrate a highly regulated innate immune mechanism that potentially rapidly tackles microbial pathogens invading the bloodstream and tissues.<\/jats:p>","DOI":"10.1093\/jimmun\/vkaf324","type":"journal-article","created":{"date-parts":[[2025,12,5]],"date-time":"2025-12-05T19:13:20Z","timestamp":1764962000000},"source":"Crossref","is-referenced-by-count":0,"title":["Innate recognition and phagocytosis of zymosan by human neutrophils require cell priming in a p38-dependent manner"],"prefix":"10.1093","volume":"215","author":[{"given":"Wei Da","family":"Ho","sequence":"first","affiliation":[{"name":"School of Biological Sciences, Faculty of Sciences, Engineering and Technology, University of Adelaide , Adelaide, SA,","place":["Australia"]},{"name":"Department of Immunopathology, SA Pathology at Women\u2019s and Children\u2019s Hospital, North Adelaide , SA,","place":["Australia"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1499-1736","authenticated-orcid":false,"given":"Khalida","family":"Perveen","sequence":"additional","affiliation":[{"name":"Department of Immunopathology, SA Pathology at Women\u2019s and Children\u2019s Hospital, North Adelaide , SA,","place":["Australia"]},{"name":"Robinson Research Institute, Adelaide Medical School and School of Biomedicine, Faculty of Health and Medical Science, University of Adelaide , Adelaide, SA,","place":["Australia"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alex","family":"Quach","sequence":"additional","affiliation":[{"name":"Department of Immunopathology, SA Pathology at Women\u2019s and Children\u2019s Hospital, North Adelaide , SA,","place":["Australia"]},{"name":"Robinson Research Institute, Adelaide Medical School and School of Biomedicine, Faculty of Health and Medical Science, University of Adelaide , Adelaide, SA,","place":["Australia"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Charles S","family":"Hii","sequence":"additional","affiliation":[{"name":"Department of Immunopathology, SA Pathology at Women\u2019s and Children\u2019s Hospital, North Adelaide , SA,","place":["Australia"]},{"name":"Robinson Research Institute, Adelaide Medical School and School of Biomedicine, Faculty of Health and Medical Science, University of Adelaide , Adelaide, SA,","place":["Australia"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Antonio","family":"Ferrante","sequence":"additional","affiliation":[{"name":"School of Biological Sciences, Faculty of Sciences, Engineering and Technology, University of Adelaide , Adelaide, SA,","place":["Australia"]},{"name":"Department of Immunopathology, SA Pathology at Women\u2019s and Children\u2019s Hospital, North Adelaide , SA,","place":["Australia"]},{"name":"Robinson Research Institute, Adelaide Medical School and School of Biomedicine, Faculty of Health and Medical Science, University of Adelaide , Adelaide, SA,","place":["Australia"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2025,12,5]]},"reference":[{"key":"2026052212171760500_vkaf324-B1","doi-asserted-by":"publisher","first-page":"293","DOI":"10.1080\/00362177985380431","article-title":"Requirement of heat-labile opsonins for maximal phagocytosis of Candida albicans","volume":"17","author":"Ferrante","year":"1979","journal-title":"Sabouraudia J Med Vet Mycol"},{"key":"2026052212171760500_vkaf324-B2","doi-asserted-by":"publisher","first-page":"1","DOI":"10.2741\/E1","article-title":"Complement in Candida albicans infections","volume":"1","author":"Rambach","year":"2009","journal-title":"Front Biosci (Elite Ed)"},{"key":"2026052212171760500_vkaf324-B3","doi-asserted-by":"publisher","DOI":"10.5772\/intechopen.80683","volume-title":"Candida Albicans","author":"Small","year":"2019"},{"key":"2026052212171760500_vkaf324-B4","doi-asserted-by":"publisher","first-page":"400","DOI":"10.1128\/MMBR.67.3.400-428.2003","article-title":"Candida albicans secreted aspartyl proteinases in virulence and pathogenesis","volume":"67","author":"Naglik","year":"2003","journal-title":"Microbiol Mol Biol Rev"},{"key":"2026052212171760500_vkaf324-B5","doi-asserted-by":"publisher","first-page":"275","DOI":"10.1111\/j.1600-0714.2009.00866.x","article-title":"Human oral keratinocyte E-cadherin degradation by Candida albicans and Candida glabrata","volume":"39","author":"P\u00e4rn\u00e4nen","year":"2010","journal-title":"J Oral Pathol Med"},{"key":"2026052212171760500_vkaf324-B6","doi-asserted-by":"publisher","first-page":"6633","DOI":"10.1128\/IAI.72.11.6633-6641.2004","article-title":"The hyphal and yeast forms of Candida albicans bind the complement regulator C4b-binding protein","volume":"72","author":"Meri","year":"2004","journal-title":"Infect Immun"},{"key":"2026052212171760500_vkaf324-B7","doi-asserted-by":"publisher","first-page":"5185","DOI":"10.1128\/IAI.70.9.5185-5192.2002","article-title":"The yeast Candida albicans binds complement regulators factor H and FHL-1","volume":"70","author":"Meri","year":"2002","journal-title":"Infect Immun"},{"key":"2026052212171760500_vkaf324-B8","doi-asserted-by":"publisher","first-page":"317","DOI":"10.1016\/j.it.2013.02.008","article-title":"Pattern recognition receptor function in neutrophils","volume":"34","author":"Thomas","year":"2013","journal-title":"Trends Immunol"},{"key":"2026052212171760500_vkaf324-B9","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1086\/375784","article-title":"Candida albicans phospholipomannan is sensed through Toll-like receptors","volume":"188","author":"Jouault","year":"2003","journal-title":"J Infect Dis"},{"key":"2026052212171760500_vkaf324-B10","doi-asserted-by":"publisher","first-page":"1642","DOI":"10.1172\/JCI27114","article-title":"Immune sensing of Candida albicans requires cooperative recognition of mannans and glucans by lectin and Toll-like receptors","volume":"116","author":"Netea","year":"2006","journal-title":"J Clin Invest"},{"key":"2026052212171760500_vkaf324-B11","doi-asserted-by":"publisher","first-page":"3059","DOI":"10.4049\/jimmunol.172.5.3059","article-title":"The contribution of the Toll-like\/IL-1 receptor superfamily to innate and adaptive immunity to fungal pathogens in vivo","volume":"172","author":"Bellocchio","year":"2004","journal-title":"J Immunol"},{"key":"2026052212171760500_vkaf324-B12","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.micinf.2003.09.020","article-title":"Toll-like receptor-2 is essential in murine defenses against Candida albicans infections","volume":"6","author":"Villam\u00f3n","year":"2004","journal-title":"Microbes Infect"},{"key":"2026052212171760500_vkaf324-B13","doi-asserted-by":"publisher","first-page":"679","DOI":"10.1093\/glycob\/cwn046","article-title":"Human and mouse macrophage-inducible C-type lectin (Mincle) bind Candida albicans","volume":"18","author":"Bugarcic","year":"2008","journal-title":"Glycobiology"},{"key":"2026052212171760500_vkaf324-B14","doi-asserted-by":"publisher","first-page":"e45781","DOI":"10.1371\/journal.pone.0045781","article-title":"Differential dependencies of monocytes and neutrophils on Dectin-1, Dectin-2 and complement for the recognition of fungal particles in inflammation","volume":"7","author":"McDonald","year":"2012","journal-title":"PLoS One"},{"key":"2026052212171760500_vkaf324-B15","doi-asserted-by":"publisher","first-page":"31","DOI":"10.1038\/ni1408","article-title":"Dectin-1 is required for \u03b2-glucan recognition and control of fungal infection","volume":"8","author":"Taylor","year":"2007","journal-title":"Nat Immunol"},{"key":"2026052212171760500_vkaf324-B16","doi-asserted-by":"publisher","first-page":"681","DOI":"10.1016\/j.immuni.2010.05.001","article-title":"Dectin-2 recognition of \u03b1-mannans and induction of Th17 cell differentiation is essential for host defense against Candida albicans","volume":"32","author":"Saijo","year":"2010","journal-title":"Immunity"},{"key":"2026052212171760500_vkaf324-B17","doi-asserted-by":"publisher","first-page":"1064","DOI":"10.1128\/IAI.01189-13","article-title":"Role of Dectin-2 for host defense against systemic infection with Candida glabrata","volume":"82","author":"Ifrim","year":"2014","journal-title":"Infect Immun"},{"key":"2026052212171760500_vkaf324-B18","doi-asserted-by":"publisher","first-page":"603","DOI":"10.1016\/j.chom.2011.10.009","article-title":"The \u03b2-glucan receptor Dectin-1 activates the integrin Mac-1 in neutrophils via vav protein signaling to promote Candida albicans clearance","volume":"10","author":"Li","year":"2011","journal-title":"Cell Host Microbe"},{"key":"2026052212171760500_vkaf324-B19","doi-asserted-by":"publisher","first-page":"38854","DOI":"10.1074\/jbc.M606542200","article-title":"Dectin-2 is a pattern recognition receptor for fungi that couples with the fc receptor \u03b3 chain to induce innate immune responses","volume":"281","author":"Sato","year":"2006","journal-title":"J Biol Chem"},{"key":"2026052212171760500_vkaf324-B20","doi-asserted-by":"publisher","first-page":"1042","DOI":"10.1110\/ps.072791207","article-title":"Structure of the fungal \u03b2-glucan-binding immune receptor Dectin-1: implications for function","volume":"16","author":"Brown","year":"2007","journal-title":"Protein Sci"},{"key":"2026052212171760500_vkaf324-B21","doi-asserted-by":"publisher","first-page":"467","DOI":"10.1002\/eji.200636653","article-title":"Dectin-1 promotes fungicidal activity of human neutrophils","volume":"37","author":"Kennedy","year":"2007","journal-title":"Eur J Immunol"},{"key":"2026052212171760500_vkaf324-B22","doi-asserted-by":"publisher","first-page":"575","DOI":"10.1016\/j.molimm.2009.09.018","article-title":"Complement receptor 3, not Dectin-1, is the major receptor on human neutrophils for \u03b2-glucan-bearing particles","volume":"47","author":"van Bruggen","year":"2009","journal-title":"Mol Immunol"},{"key":"2026052212171760500_vkaf324-B23","doi-asserted-by":"publisher","first-page":"e1254792","DOI":"10.1155\/2017\/1254792","article-title":"The application of dextran sedimentation as an initial step in neutrophil purification promotes their stimulation, due to the presence of monocytes","volume":"2017","author":"Quach","year":"2017","journal-title":"J Immunol Res."},{"key":"2026052212171760500_vkaf324-B24","doi-asserted-by":"publisher","first-page":"1038","DOI":"10.1097\/CCM.0b013e3181968fa0","article-title":"Leukocyte Dectin-1 expression is differentially regulated in fungal versus polymicrobial sepsis","volume":"37","author":"Ozment-Skelton","year":"2009","journal-title":"Crit Care Med"},{"key":"2026052212171760500_vkaf324-B25","doi-asserted-by":"publisher","first-page":"579","DOI":"10.1111\/mmi.12480","article-title":"Characterization of Aspergillus nidulans \u03b1-glucan synthesis: roles for two synthases and two amylases","volume":"91","author":"He","year":"2014","journal-title":"Mol Microbiol"},{"key":"2026052212171760500_vkaf324-B26","doi-asserted-by":"publisher","first-page":"341","DOI":"10.1016\/S0096-5332(08)60191-5","volume-title":"Advances in carbohydrate chemistry","author":"Neely","year":"1961"},{"key":"2026052212171760500_vkaf324-B27","doi-asserted-by":"crossref","first-page":"788","DOI":"10.1128\/iai.58.3.788-793.1990","article-title":"Tumor necrosis factor enhances neutrophil-mediated killing of Plasmodium falciparum","volume":"58","author":"Kumaratilake","year":"1990","journal-title":"Infect Immun"},{"key":"2026052212171760500_vkaf324-B28","doi-asserted-by":"publisher","first-page":"179","DOI":"10.1016\/0165-2478(90)90112-4","article-title":"Killing of Plasmodium falciparum by cytokine activated effector cells (neutrophils and macrophages)","volume":"25","author":"Ferrante","year":"1990","journal-title":"Immunol Lett"},{"key":"2026052212171760500_vkaf324-B29","doi-asserted-by":"crossref","first-page":"2115","DOI":"10.1128\/iai.57.7.2115-2122.1989","article-title":"Tumor necrosis factor alpha potentiates neutrophil antimicrobial activity: increased fungicidal activity against Torulopsis glabrata and Candida albicans and associated increases in oxygen radical production and lysosomal enzyme release","volume":"57","author":"Ferrante","year":"1989","journal-title":"Infect Immun"},{"key":"2026052212171760500_vkaf324-B30","doi-asserted-by":"publisher","first-page":"389","DOI":"10.1016\/0022-1759(78)90143-6","article-title":"A rapid one-step procedure for purification of mononuclear and polymorphonuclear leukocytes from human blood using a modification of the Hypaque\u2013Ficoll technique","volume":"24","author":"Ferrante","year":"1978","journal-title":"J Immunol Methods"},{"key":"2026052212171760500_vkaf324-B31","doi-asserted-by":"publisher","first-page":"245","DOI":"10.2144\/btn-2020-0003","article-title":"Phagocytosis assays with different pH-sensitive fluorescent particles and various readouts","volume":"68","author":"Lindner","year":"2020","journal-title":"BioTechniques"},{"key":"2026052212171760500_vkaf324-B32","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1128\/CDLI.7.1.9-13.2000","article-title":"Opsonization and phagocytosis of Plasmodium falciparum merozoites measured by flow cytometry","volume":"7","author":"Kumaratilake","year":"2000","journal-title":"Clin Diagn Lab Immunol"},{"key":"2026052212171760500_vkaf324-B33","doi-asserted-by":"publisher","first-page":"e0216465","DOI":"10.1371\/journal.pone.0216465","article-title":"Nrf2 downregulates zymosan-induced neutrophil activation and modulates migration","volume":"14","author":"Helou","year":"2019","journal-title":"PLoS One"},{"key":"2026052212171760500_vkaf324-B34","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1016\/j.jim.2011.06.027","article-title":"Development and validation of a flow cytometric method to evaluate phagocytosis of pHrodoTM BioParticles\u00ae by granulocytes in multiple species","volume":"390","author":"Neaga","year":"2013","journal-title":"J Immunol Methods"},{"key":"2026052212171760500_vkaf324-B35","doi-asserted-by":"publisher","first-page":"62","DOI":"10.1016\/0014-5793(89)81718-1","article-title":"Granulocyte-macrophage colony-stimulating factor (GM-CSF) primes the respiratory burst and stimulates protein biosynthesis in human neutrophils","volume":"256","author":"Edwards","year":"1989","journal-title":"FEBS Lett"},{"key":"2026052212171760500_vkaf324-B36","doi-asserted-by":"publisher","first-page":"137","DOI":"10.1111\/j.1365-2141.1992.tb08890.x","article-title":"Activation of human eosinophil and neutrophil functions by haematopoietic growth factors: comparisons of IL-1, IL-3, IL-5 and GM-CSF","volume":"80","author":"Fabian","year":"1992","journal-title":"Br J Haematol"},{"key":"2026052212171760500_vkaf324-B37","doi-asserted-by":"publisher","first-page":"260","DOI":"10.1016\/j.ejphar.2007.05.001","article-title":"Phospholipase D-dependent and -independent p38MAPK activation pathways are required for superoxide production and chemotactic induction, respectively, in rat neutrophils stimulated by fMLP","volume":"568","author":"Azuma","year":"2007","journal-title":"Eur J Pharmacol"},{"key":"2026052212171760500_vkaf324-B38","doi-asserted-by":"publisher","first-page":"3503","DOI":"10.1182\/blood.V89.10.3503.3503_3503_3521","article-title":"Granules of the human neutrophilic polymorphonuclear leukocyte","volume":"89","author":"Borregaard","year":"1997","journal-title":"Blood"},{"key":"2026052212171760500_vkaf324-B39","doi-asserted-by":"publisher","first-page":"195","DOI":"10.1016\/0309-1651(88)90096-3","article-title":"Effects of cytochalasin B, N-formyl peptide and plasma on polarisation, zeiosis (blebbing) and degranulation of polymorphonuclear leukocytes in suspension","volume":"12","author":"Bignold","year":"1988","journal-title":"Cell Biol Int Rep"},{"key":"2026052212171760500_vkaf324-B40","doi-asserted-by":"publisher","first-page":"52","DOI":"10.1289\/ehp.9610452","article-title":"Phospholipase A2 is involved in the mechanism of activation of neutrophils by polychlorinated biphenyls","volume":"104","author":"Tithof","year":"1996","journal-title":"Environ Health Perspect"},{"key":"2026052212171760500_vkaf324-B41","doi-asserted-by":"publisher","first-page":"2616","DOI":"10.4049\/jimmunol.171.5.2616","article-title":"Unique effect of arachidonic acid on human neutrophil TNF receptor expression: up-regulation involving protein kinase C, extracellular signal-regulated kinase, and phospholipase A2","volume":"171","author":"Moghaddami","year":"2003","journal-title":"J Immunol"},{"key":"2026052212171760500_vkaf324-B42","doi-asserted-by":"publisher","first-page":"33","DOI":"10.1038\/nri1745","article-title":"Dectin-1: a signalling non-TLR pattern-recognition receptor","volume":"6","author":"Brown","year":"2006","journal-title":"Nat Rev Immunol"},{"key":"2026052212171760500_vkaf324-B43","doi-asserted-by":"publisher","first-page":"159","DOI":"10.1016\/0022-1759(90)90206-B","article-title":"Preparative procedures of cooling and re-warming increase leukocyte integrin expression and function on neotrophils","volume":"128","author":"Forsyth","year":"1990","journal-title":"J Immunol Methods"},{"key":"2026052212171760500_vkaf324-B44","doi-asserted-by":"publisher","first-page":"2889","DOI":"10.3390\/cells11182889","article-title":"Neutrophil activated by the famous and potent PMA (Phorbol Myristate Acetate)","volume":"11","author":"Damascena","year":"2022","journal-title":"Cells"},{"key":"2026052212171760500_vkaf324-B45","doi-asserted-by":"publisher","first-page":"840510","DOI":"10.3389\/fimmu.2022.840510","article-title":"Neutrophils require activation to express functional cell-surface complement receptor immunoglobulin","volume":"13","author":"Small","year":"2022","journal-title":"Front Immunol"},{"key":"2026052212171760500_vkaf324-B46","doi-asserted-by":"publisher","first-page":"11","DOI":"10.1016\/S0022-1759(98)00049-0","article-title":"Effect of ex vivo storage on human peripheral blood neutrophil expression of CD11b and the stabilizing effects of Cyto-ChexTM","volume":"214","author":"Saxton","year":"1998","journal-title":"J Immunol Methods"},{"key":"2026052212171760500_vkaf324-B47","doi-asserted-by":"publisher","first-page":"207","DOI":"10.1016\/0035-9203(78)90067-6","article-title":"Phagocytic behaviour towards baker\u2019s yeast distinguishes pathogenic from non-pathogenic Naegleria","volume":"72","author":"Thong","year":"1978","journal-title":"Trans R Soc Trop Med Hyg"},{"key":"2026052212171760500_vkaf324-B48","doi-asserted-by":"crossref","first-page":"4821","DOI":"10.4049\/jimmunol.151.9.4821","article-title":"Killing of Staphylococcus aureus by tumor necrosis factor-alpha-activated neutrophils: the role of serum opsonins, integrin receptors, respiratory burst, and degranulation","volume":"151","author":"Ferrante","year":"1993","journal-title":"J Immunol"},{"key":"2026052212171760500_vkaf324-B49","doi-asserted-by":"publisher","first-page":"590","DOI":"10.1182\/blood-2014-01-551473","article-title":"Two independent killing mechanisms of Candida albicans by human neutrophils: evidence from innate immunity defects","volume":"124","author":"Gazendam","year":"2014","journal-title":"Blood"},{"key":"2026052212171760500_vkaf324-B50","doi-asserted-by":"publisher","first-page":"57","DOI":"10.1189\/jlb.69.1.57","article-title":"Neutrophil secretory vesicles are the intracellular reservoir for GPI-80, a protein with adhesion-regulating potential","volume":"69","author":"Dahlgren","year":"2001","journal-title":"J Leukoc Biol"},{"key":"2026052212171760500_vkaf324-B51","doi-asserted-by":"publisher","first-page":"4747","DOI":"10.1128\/iai.65.11.4747-4753.1997","article-title":"Azurophilic granules of human neutrophils contain CD14","volume":"65","author":"Rodeberg","year":"1997","journal-title":"Infect Immun"},{"key":"2026052212171760500_vkaf324-B52","doi-asserted-by":"publisher","first-page":"19","DOI":"10.1189\/jlb.3HI0716-293RR","article-title":"Frontline Science: tumor necrosis factor-\u03b1 stimulation and priming of human neutrophil granule exocytosis","volume":"102","author":"McLeish","year":"2017","journal-title":"J Leukoc Biol"},{"key":"2026052212171760500_vkaf324-B53","first-page":"1204","article-title":"Chemoattractant-regulated mobilization of a novel intracellular compartment in human neutrophils","volume":"237","author":"Borregaard","year":"1987","journal-title":"Science (1979)"},{"key":"2026052212171760500_vkaf324-B54","doi-asserted-by":"publisher","first-page":"4330","DOI":"10.1182\/blood.V88.11.4330.4330","article-title":"Demonstration of reversible priming of human neutrophils using platelet- activating factor","volume":"88","author":"Kitchen","year":"1996","journal-title":"Blood"}],"container-title":["The Journal of Immunology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/jimmunol\/article-pdf\/215\/3\/vkaf324\/65780448\/vkaf324.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/jimmunol\/article-pdf\/215\/3\/vkaf324\/65780448\/vkaf324.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,5,22]],"date-time":"2026-05-22T16:17:26Z","timestamp":1779466646000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/jimmunol\/article\/doi\/10.1093\/jimmun\/vkaf324\/8371966"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,12,5]]},"references-count":54,"journal-issue":{"issue":"3","published-print":{"date-parts":[[2026,3,17]]}},"URL":"https:\/\/doi.org\/10.1093\/jimmun\/vkaf324","relation":{},"ISSN":["0022-1767","1550-6606"],"issn-type":[{"value":"0022-1767","type":"print"},{"value":"1550-6606","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2026,3]]},"published":{"date-parts":[[2025,12,5]]},"article-number":"vkaf324"}}