{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,4]],"date-time":"2026-11-04T17:39:40Z","timestamp":1793813980400,"version":"4.6.1"},"reference-count":94,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,7,20]],"date-time":"2026-07-20T00:00:00Z","timestamp":1784505600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Immunol."],"abstract":"<jats:sec>\n                    <jats:title>Introduction<\/jats:title>\n                    <jats:p>Monocytes play a critical role in regulating immune response to exogenous and endogenous challenge. Recent studies have shown exogenous stimuli can induce metabolic, epigenetic and phenotypic changes in monocytes that reprogramme or \u2018prime\u2019 them to react with an altered response to subsequent stimuli. Obesity is associated with immune dysfunction. However, the ability of endogenous compounds associated with obesity to prime human monocytes and thereby modulate subsequent response remains to be investigated.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods and results<\/jats:title>\n                    <jats:p>In this study, human monocytes from non-obese males were utilized. Monocytes exposed to primary exogenous stimuli (\u03b2-glucan), but not endogenous stimuli (oxLDL, leptin, 17\u03b2-estradiol), demonstrated an altered phenotype (increased CD11b, PD-L1 expression) in response to SK-MEL-28 melanoma cells. In contrast, monocytes primed with oxLDL, but not \u03b2-glucan, exhibited an increased rate of superoxide production in response to secondary stimulation with phorbol 12-myristate 13-acetate, and enhanced matrix metalloprotease 9 activity in response to secondary stimulation with lipopolysaccharide. Bulk RNAseq analysis of non-primed monocytes and monocytes primed with 17\u03b2-estradiol or oxLDL also showed sustained differential expression of genes related to immune function, metabolism and protein modification. Moreover, mass spectrometry analysis showed that monocytes primed with \u03b2-glucan or oxLDL exhibited sustained modifications to global DNA methylation.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Conclusions<\/jats:title>\n                    <jats:p>These results provide the first steps in understanding the functional consequences of monocyte reprogramming by obesity-related factors, and the potential pathways that govern them.<\/jats:p>\n                  <\/jats:sec>","DOI":"10.3389\/fimmu.2026.1825431","type":"journal-article","created":{"date-parts":[[2026,7,20]],"date-time":"2026-07-20T04:21:08Z","timestamp":1784521268000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":3,"title":["Priming of human monocytes by \u03b2-glucan and obesity-associated factors: modifications to global DNA methylation, gene expression, phenotype and function"],"prefix":"10.3389","volume":"17","author":[{"given":"Ben J.","family":"Topham","sequence":"first","affiliation":[{"name":"Mackenzie Cancer Research Group, Department of Pathology & Molecular Medicine, University of Otago Christchurch","place":["Christchurch, New Zealand"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Barry D.","family":"Hock","sequence":"additional","affiliation":[{"name":"Haematology Research Group, Department of Pathology & Molecular Medicine, University of Otago Christchurch","place":["Christchurch, New Zealand"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"George A. R.","family":"Wiggins","sequence":"additional","affiliation":[{"name":"Mackenzie Cancer Research Group, Department of Pathology & Molecular Medicine, University of Otago Christchurch","place":["Christchurch, New Zealand"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Louisa V.","family":"Ashby","sequence":"additional","affiliation":[{"name":"M\u0101tai H\u0101ora - Centre for Redox Biology and Medicine, Department of Pathology & Molecular Medicine, University of Otago Christchurch","place":["Christchurch, New Zealand"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nicholas J.","family":"Magon","sequence":"additional","affiliation":[{"name":"M\u0101tai H\u0101ora - Centre for Redox Biology and Medicine, Department of Pathology & Molecular Medicine, University of Otago Christchurch","place":["Christchurch, New Zealand"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Elisabeth","family":"Phillips","sequence":"additional","affiliation":[{"name":"Mackenzie Cancer Research Group, Department of Pathology & Molecular Medicine, University of Otago Christchurch","place":["Christchurch, New Zealand"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Emma K. C.","family":"Symonds","sequence":"additional","affiliation":[{"name":"Department of Surgery and Critical Care, University of Otago Wellington","place":["Wellington, New Zealand"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kirsty M.","family":"Danielson","sequence":"additional","affiliation":[{"name":"Department of Surgery and Critical Care, University of Otago Wellington","place":["Wellington, New Zealand"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Margaret J.","family":"Currie","sequence":"additional","affiliation":[{"name":"Mackenzie Cancer Research Group, Department of Pathology & Molecular Medicine, University of Otago Christchurch","place":["Christchurch, New Zealand"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,7,20]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"11","DOI":"10.1155\/2017\/9624760","article-title":"Macrophages as key drivers of cancer progression and metastasis","volume":"2017","author":"Nielsen","year":"2017","journal-title":"Mediators Inflammation"},{"key":"B2","doi-asserted-by":"publisher","first-page":"18486","DOI":"10.18632\/oncotarget.14474","article-title":"Targeting macrophage anti-tumor activity to suppress melanoma progression","volume":"8","author":"Wang","year":"2017","journal-title":"Oncotarget"},{"key":"B3","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-2613.2006.00478.x","article-title":"Tumour\u2010associated macrophages and melanoma tumourigenesis: integrating the complexity","volume":"87","author":"Hussein","year":"2006","journal-title":"Int J Exp Path"},{"key":"B4","doi-asserted-by":"publisher","DOI":"10.3390\/cancers13081946","article-title":"The role of macrophages in cancer development and therapy","volume":"13","author":"Cendrowicz","year":"2021","journal-title":"Cancers"},{"key":"B5","doi-asserted-by":"publisher","first-page":"1979","DOI":"10.3390\/cells11121979","article-title":"The good and the bad: monocytes\u2019 and macrophages\u2019 diverse functions in inflammation","volume":"11","author":"Austermann","year":"2022","journal-title":"Cells"},{"key":"B6","doi-asserted-by":"publisher","DOI":"10.1038\/s41573-019-0025-4","article-title":"Therapeutic targeting of trained immunity","volume":"18","author":"Mulder","year":"2019","journal-title":"Nat Rev Drug Discov"},{"key":"B7","doi-asserted-by":"publisher","DOI":"10.1161\/atvbaha.120.314212","article-title":"In vitro experimental model of trained innate immunity in human primary monocytes","volume":"23","author":"Bekkering","year":"2016","journal-title":"Clin Vaccine Immunol"},{"key":"B8","doi-asserted-by":"publisher","DOI":"10.1038\/s41577-020-0285-6","article-title":"Defining trained immunity and its role in health and disease","volume":"20","author":"Netea","year":"2020","journal-title":"Nat Rev Immunol"},{"key":"B9","doi-asserted-by":"publisher","DOI":"10.1038\/s41590-022-01388-8","article-title":"Inducing trained immunity in pro-metastatic macrophages to control tumor metastasis","volume":"24","author":"Ding","year":"2023","journal-title":"Nat Immunol"},{"key":"B10","doi-asserted-by":"publisher","first-page":"1250684","DOI":"10.1126\/science.1250684","article-title":"mTOR-and HIF-1\u03b1\u2013mediated aerobic glycolysis as metabolic basis for trained immunity","volume":"345","author":"Cheng","year":"2014","journal-title":"Science"},{"key":"B11","doi-asserted-by":"publisher","first-page":"32","DOI":"10.1016\/j.immuni.2020.10.011","article-title":"The intersection of epigenetics and metabolism in trained immunity","volume":"54","author":"Fanucchi","year":"2021","journal-title":"Immunity"},{"key":"B12","doi-asserted-by":"publisher","first-page":"23","DOI":"10.1038\/s41581-022-00633-5","article-title":"Trained immunity\u2014basic concepts and contributions to immunopathology","volume":"19","author":"Ochando","year":"2023","journal-title":"Nat Rev Nephrol"},{"key":"B13","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2021.02.011","article-title":"Understanding and treating the inflammatory adverse events of cancer immunotherapy","volume":"184","author":"Dougan","year":"2021","journal-title":"Cell"},{"key":"B14","doi-asserted-by":"publisher","DOI":"10.1017\/s0029665115004176","article-title":"Obesity-associated cancer: an immunological perspective","volume":"75","author":"Conroy","year":"2016","journal-title":"Proc Nutr Soc"},{"key":"B15","doi-asserted-by":"publisher","DOI":"10.1007\/s11912-010-0139-7","article-title":"Obesity and cancer risk: recent review and evidence","volume":"13","author":"Basen-Engquist","year":"2011","journal-title":"Curr Oncol Rep"},{"key":"B16","doi-asserted-by":"publisher","DOI":"10.1038\/s41574-023-00932-2","article-title":"Emerging mechanisms of obesity-associated immune dysfunction","volume":"20","author":"Shaikh","year":"2024","journal-title":"Nat Rev Endocrinol"},{"key":"B17","doi-asserted-by":"publisher","DOI":"10.1038\/s41574-019-0161-2","article-title":"The \u2018obesity paradox\u2019 in action with cancer immunotherapy","volume":"15","author":"Lysaght","year":"2019","journal-title":"Nat Rev Endocrinol"},{"key":"B18","doi-asserted-by":"publisher","DOI":"10.1038\/nrrheum.2016.209","article-title":"Leptin in the interplay of inflammation, metabolism and immune system disorders","volume":"13","author":"Abella","year":"2017","journal-title":"Nat Rev Rheumatol"},{"key":"B19","doi-asserted-by":"publisher","DOI":"10.1161\/atvbaha.114.303887","article-title":"Oxidized low-density lipoprotein induces long-term proinflammatory cytokine production and foam cell formation via epigenetic reprogramming of monocytes","volume":"34","author":"Bekkering","year":"2014","journal-title":"Arteriosclerosis Thrombosis Vasc Biol"},{"key":"B20","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2018.12.018","article-title":"The short chain fatty acid butyrate imprints an antimicrobial program in macrophages","volume":"50","author":"Schulthess","year":"2019","journal-title":"Immunity"},{"key":"B21","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1620910114","article-title":"Uric acid priming in human monocytes is driven by the AKT\u2013PRAS40 autophagy pathway","volume":"114","author":"Cri\u015fan","year":"2017","journal-title":"Proc Natl Acad Sci"},{"key":"B22","doi-asserted-by":"publisher","DOI":"10.1093\/cvr\/cvz137","article-title":"Aldosterone induces trained immunity: the role of fatty acid synthesis","volume":"116","author":"van der Heijden","year":"2020","journal-title":"Cardiovasc Res"},{"key":"B23","doi-asserted-by":"publisher","DOI":"10.1056\/nejm199602013340503","article-title":"Serum immunoreactive-leptin concentrations in normal-weight and obese humans","volume":"334","author":"Considine","year":"1996","journal-title":"N Engl J Med"},{"key":"B24","doi-asserted-by":"publisher","DOI":"10.1210\/jcem-48-4-633","article-title":"Increased estrogen production in obese men","volume":"48","author":"Schneider","year":"1979","journal-title":"J Clin Endocrinol Metab"},{"key":"B25","doi-asserted-by":"publisher","DOI":"10.1093\/ajcn\/83.1.30","article-title":"Circulating oxidized LDL is associated with increased waist circumference independent of body mass index in men and women","volume":"83","author":"Weinbrenner","year":"2006","journal-title":"Am J Clin Nutr"},{"key":"B26","doi-asserted-by":"publisher","DOI":"10.1038\/s41366-020-00731-z","article-title":"Body mass index, waist circumference, waist-to-hip ratio, and body fat in relation to health care use in the Canadian Longitudinal Study on Aging","volume":"45","author":"Andreacchi","year":"2021","journal-title":"Int J Obes"},{"key":"B27","doi-asserted-by":"publisher","DOI":"10.1038\/icb.2017.13","article-title":"Evaluation of trained immunity by \u03b2\u20101, 3 (d)\u2010glucan on murine monocytes in vitro and duration of response in vivo","volume":"95","author":"Garcia\u2010Valtanen","year":"2017","journal-title":"Immunol Cell Biol"},{"key":"B28","doi-asserted-by":"publisher","DOI":"10.2337\/diabetes.53.4.1068","article-title":"The metabolic syndrome, circulating oxidized LDL, and risk of myocardial infarction in well-functioning elderly people in the health, aging, and body composition cohort","volume":"53","author":"Holvoet","year":"2004","journal-title":"Diabetes"},{"key":"B29","doi-asserted-by":"publisher","DOI":"10.1002\/dmrr.1011","article-title":"Association between oxidized LDL, obesity and type 2 diabetes in a population\u2010based cohort, the Health, Aging and Body Composition Study","volume":"25","author":"Njajou","year":"2009","journal-title":"Diabetes\/Metabolism Res Rev"},{"key":"B30","doi-asserted-by":"publisher","DOI":"10.14814\/phy2.12177","article-title":"Leptin regulates CD 16 expression on human monocytes in a sex\u2010specific manner","volume":"2","author":"Cannon","year":"2014","journal-title":"Physiol Rep"},{"key":"B31","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.167.8.4593","article-title":"High-dose leptin activates human leukocytes via receptor expression on monocytes","volume":"167","author":"Zarkesh-Esfahani","year":"2001","journal-title":"J Immunol"},{"key":"B32","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.163.7.3758","article-title":"Estradiol coupling to human monocyte nitric oxide release is dependent on intracellular calcium transients: evidence for an estrogen surface receptor","volume":"163","author":"Stefano","year":"1999","journal-title":"J Immunol"},{"key":"B33","doi-asserted-by":"publisher","first-page":"16","DOI":"10.1016\/j.mce.2007.08.014","article-title":"17\u03b2-Estradiol utilizes the estrogen receptor to regulate CD16 expression in monocytes","volume":"279","author":"Kramer","year":"2007","journal-title":"Mol Cell Endocrinol"},{"key":"B34","doi-asserted-by":"publisher","DOI":"10.1210\/jc.2013-2605","article-title":"17\u03b2-Estradiol and estradiol fatty acyl esters and estrogen-converting enzyme expression in adipose tissue in obese men and women","volume":"98","author":"Wang","year":"2013","journal-title":"J Clin Endocrinol Metab"},{"key":"B35","unstructured":"Protocol for Charcoal-stripping FBS to Deplete Hormones"},{"key":"B36","doi-asserted-by":"publisher","DOI":"10.1016\/0005-2728(76)90130-4","article-title":"The kinetics of the reduction of cytochrome c by the superoxide anion radical","volume":"449","author":"Koppenol","year":"1976","journal-title":"Biochim Biophys Acta (BBA)-Bioenergetics"},{"key":"B37","doi-asserted-by":"publisher","DOI":"10.1016\/0006-3002(62)90073-2","article-title":"The extinction coefficient of cytochrome c","volume":"58","author":"Van Gelder","year":"1962","journal-title":"Biochim Biophys Acta"},{"key":"B38","article-title":"FastQC: a quality control tool for high throughput sequence data","volume":"2","author":"Andersson","year":"2010","journal-title":"Version 0.11"},{"key":"B39","doi-asserted-by":"publisher","DOI":"10.1093\/bioinformatics\/btu170","article-title":"Trimmomatic: a flexible trimmer for Illumina sequence data","volume":"30","author":"Bolger","year":"2014","journal-title":"Bioinformatics"},{"key":"B40","doi-asserted-by":"publisher","first-page":"15","DOI":"10.1093\/bioinformatics\/bts635","article-title":"STAR: ultrafast universal RNA-seq aligner","volume":"29","author":"Dobin","year":"2013","journal-title":"Bioinformatics"},{"key":"B41","unstructured":"Human Release 44 (GRCh38.p14)"},{"key":"B42","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/s13059-014-0550-8","article-title":"Moderated estimation of fold change and dispersion for RNA-seq data with DESeq2","volume":"15","author":"Love","year":"2014","journal-title":"Genome Biol"},{"key":"B43","doi-asserted-by":"publisher","DOI":"10.3389\/fonc.2021.709543","article-title":"Ascorbate inhibits proliferation and promotes myeloid differentiation in TP53-mutant leukemia","volume":"11","author":"Smith-D\u00edaz","year":"2021","journal-title":"Front Oncol"},{"key":"B44","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1202870109","article-title":"Bacille Calmette-Guerin induces NOD2-dependent nonspecific protection from reinfection via epigenetic reprogramming of monocytes","volume":"109","author":"Kleinnijenhuis","year":"2012","journal-title":"Proc Natl Acad Sci"},{"key":"B45","doi-asserted-by":"publisher","DOI":"10.1016\/s0021-9258(19)40138-5","article-title":"Assembly and activation of the NADPH: O2 oxidoreductase in human neutrophils after stimulation with phorbol myristate acetate","volume":"265","author":"Ambruso","year":"1990","journal-title":"J Biol Chem"},{"key":"B46","article-title":"Cytochrome c","volume-title":"Handbook of Methods for Oxygen Radical Research","author":"Fridovich","year":"1985"},{"key":"B47","doi-asserted-by":"publisher","first-page":"5856","DOI":"10.3390\/ijms25115856","article-title":"Trained immunity and trained tolerance: the case of Helicobacter pylori infection","volume":"25","author":"Dore","year":"2024","journal-title":"Int J Mol Sci"},{"key":"B48","doi-asserted-by":"publisher","DOI":"10.1089\/ars.2016.6695","article-title":"Epigenetic regulation of monocyte and macrophage function","volume":"25","author":"Hoeksema","year":"2016","journal-title":"Antioxidants Redox Signaling"},{"key":"B49","doi-asserted-by":"publisher","first-page":"23","DOI":"10.1038\/npp.2012.112","article-title":"DNA methylation and its basic function","volume":"38","author":"Moore","year":"2013","journal-title":"Neuropsychopharmacology"},{"key":"B50","doi-asserted-by":"publisher","DOI":"10.3390\/nu15092124","article-title":"Beta-glucans of cereals: functional and technological properties","volume":"15","author":"Lante","year":"2023","journal-title":"Nutrients"},{"key":"B51","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2022.872652","article-title":"Trained immunity enhances human monocyte function in aging and sepsis","volume":"13","author":"Gill","year":"2022","journal-title":"Front Immunol"},{"key":"B52","doi-asserted-by":"publisher","DOI":"10.1016\/s0735-1097(97)00324-0","article-title":"HMG-CoA reductase inhibitors decrease CD11b expression and CD11b-dependent adhesion of monocytes to endothelium and reduce increased adhesiveness of monocytes isolated from patients with hypercholesterolemia","volume":"30","author":"Weber","year":"1997","journal-title":"J Am Coll Cardiol"},{"key":"B53","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1500396113","article-title":"CD11b regulates obesity-induced insulin resistance via limiting alternative activation and proliferation of adipose tissue macrophages","volume":"112","author":"Zheng","year":"2015","journal-title":"Proc Natl Acad Sci"},{"key":"B54","doi-asserted-by":"publisher","DOI":"10.1242\/jcs.018036","article-title":"Integrin-dependent phagocytosis\u2013spreading from microadhesion to new concepts","volume":"121","author":"Dupuy","year":"2008","journal-title":"J Cell Sci"},{"key":"B55","doi-asserted-by":"publisher","DOI":"10.3389\/fmed.2018.00052","article-title":"CD11b activity modulates pathogenesis of lupus nephritis","volume":"5","author":"Khan","year":"2018","journal-title":"Front Med"},{"key":"B56","doi-asserted-by":"publisher","first-page":"5379","DOI":"10.1038\/s41467-018-07387-4","article-title":"Integrin CD11b activation drives anti-tumor innate immunity","volume":"9","author":"Schmid","year":"2018","journal-title":"Nat Commun"},{"key":"B57","doi-asserted-by":"publisher","first-page":"195","DOI":"10.1038\/s41568-022-00442-9","article-title":"Mechanisms of immune activation and regulation: lessons from melanoma","volume":"22","author":"Kalaora","year":"2022","journal-title":"Nat Rev Cancer"},{"key":"B58","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2019.02298","article-title":"The diverse function of PD-1\/PD-L pathway beyond cancer","volume":"10","author":"Qin","year":"2019","journal-title":"Front Immunol"},{"key":"B59","doi-asserted-by":"publisher","DOI":"10.1016\/j.immuni.2018.03.014","article-title":"Regulation and function of the PD-L1 checkpoint","volume":"48","author":"Sun","year":"2018","journal-title":"Immunity"},{"key":"B60","doi-asserted-by":"publisher","DOI":"10.1016\/j.xcrm.2024.101420","article-title":"PD-L1-expressing tumor-associated macrophages are immunostimulatory and associate with good clinical outcome in human breast cancer","volume":"5","author":"Wang","year":"2024","journal-title":"Cell Rep Med"},{"key":"B61","doi-asserted-by":"publisher","DOI":"10.1038\/s41577-020-0306-5","article-title":"A guide to cancer immunotherapy: from T cell basic science to clinical practice","volume":"20","author":"Waldman","year":"2020","journal-title":"Nat Rev Immunol"},{"key":"B62","doi-asserted-by":"publisher","DOI":"10.1038\/nm.4466","article-title":"High-dimensional single-cell analysis predicts response to anti-PD-1 immunotherapy","volume":"24","author":"Krieg","year":"2018","journal-title":"Nat Med"},{"key":"B63","doi-asserted-by":"publisher","DOI":"10.3390\/cancers12102966","article-title":"High PD-L1\/CD274 expression of monocytes and blood dendritic cells is a risk factor in lung cancer patients undergoing treatment with PD1 inhibitor therapy","volume":"12","author":"Riemann","year":"2020","journal-title":"Cancers"},{"key":"B64","doi-asserted-by":"publisher","DOI":"10.1007\/s00262-020-02686-6","article-title":"A high number of PD-L1+ CD14+ monocytes in peripheral blood is correlated with shorter survival in patients receiving immune checkpoint inhibitors","volume":"70","author":"Ando","year":"2021","journal-title":"Cancer Immunol Immunother"},{"key":"B65","doi-asserted-by":"publisher","DOI":"10.3390\/genes16040448","article-title":"G0S2 promotes PD-L1 expression in monocytes and influences the efficacy of PD-1 inhibitors in hepatocellular carcinoma","volume":"16","author":"Du","year":"2025","journal-title":"Genes"},{"key":"B66","doi-asserted-by":"publisher","DOI":"10.3390\/cancers13050986","article-title":"Oxidative stress in the tumor microenvironment and its relevance to cancer immunotherapy","volume":"13","author":"Aboelella","year":"2021","journal-title":"Cancers"},{"key":"B67","doi-asserted-by":"publisher","DOI":"10.3390\/cancers14071847","article-title":"MMP9: a tough target for targeted therapy for cancer","volume":"14","author":"Augoff","year":"2022","journal-title":"Cancers"},{"key":"B68","doi-asserted-by":"publisher","DOI":"10.3109\/08830185.2012.755176","article-title":"Reactive oxygen species in the immune system","volume":"32","author":"Yang","year":"2013","journal-title":"Int Rev Immunol"},{"key":"B69","doi-asserted-by":"publisher","DOI":"10.1038\/nchembio.85","article-title":"Reconciling the chemistry and biology of reactive oxygen species","volume":"4","author":"Winterbourn","year":"2008","journal-title":"Nat Chem Biol"},{"key":"B70","doi-asserted-by":"publisher","first-page":"5","DOI":"10.1038\/cmi.2014.89","article-title":"NADPH oxidases: an overview from structure to innate immunity-associated pathologies","volume":"12","author":"Panday","year":"2015","journal-title":"Cell Mol Immunol"},{"key":"B71","doi-asserted-by":"publisher","first-page":"245","DOI":"10.1152\/physrev.00044.2005","article-title":"The NOX family of ROS-generating NADPH oxidases: physiology and pathophysiology","volume":"87","author":"Bedard","year":"2007","journal-title":"Physiol Rev"},{"key":"B72","doi-asserted-by":"publisher","DOI":"10.1016\/j.ejmech.2020.112260","article-title":"Matrix metalloproteinase-9 (MMP-9) and its inhibitors in cancer: a minireview","volume":"194","author":"Mondal","year":"2020","journal-title":"Eur J Med Chem"},{"key":"B73","doi-asserted-by":"publisher","first-page":"14351","DOI":"10.1038\/s41598-023-41514-6","article-title":"TNF\u03b1 induces matrix metalloproteinase-9 expression in monocytic cells through ACSL1\/JNK\/ERK\/NF-kB signaling pathways","volume":"13","author":"Al-Roub","year":"2023","journal-title":"Sci Rep"},{"key":"B74","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M111.290676","article-title":"Classically activated macrophages use stable microtubules for matrix metalloproteinase-9 (MMP-9) secretion","volume":"287","author":"Hanania","year":"2012","journal-title":"J Biol Chem"},{"key":"B75","doi-asserted-by":"publisher","DOI":"10.1111\/febs.14671","article-title":"A mitochondrial ROS pathway controls matrix metalloproteinase 9 levels and invasive properties in RAS\u2010activated cancer cells","volume":"286","author":"Mori","year":"2019","journal-title":"FEBS J"},{"key":"B76","doi-asserted-by":"publisher","DOI":"10.1016\/j.esmoop.2024.104047","article-title":"PP353 SLC5A3 regulates chemoresistance via PINK1\/Parkin-dependent mitophagy in Pancreatic Cancer","volume":"10","author":"Kim","year":"2025","journal-title":"ESMO Open"},{"key":"B77","doi-asserted-by":"publisher","DOI":"10.1002\/j.1460-2075.1984.tb02294.x","article-title":"Human genes and pseudogenes for the 7SL RNA component of signal recognition particle","volume":"3","author":"Ullu","year":"1984","journal-title":"EMBO J"},{"key":"B78","doi-asserted-by":"publisher","DOI":"10.3389\/fgene.2022.898083","article-title":"Signal recognition particle in human diseases","volume":"13","author":"Kellogg","year":"2022","journal-title":"Front Genet"},{"key":"B79","doi-asserted-by":"publisher","first-page":"4781","DOI":"10.1038\/s41598-018-23226-4","article-title":"Evaluation of two main RNA-seq approaches for gene quantification in clinical RNA sequencing: polyA+ selection versus rRNA depletion","volume":"8","author":"Zhao","year":"2018","journal-title":"Sci Rep"},{"key":"B80","doi-asserted-by":"publisher","DOI":"10.1016\/j.isci.2025.112440","article-title":"Inhibition of acid or neutral sphingomyelinases differentially impacts RNA and protein cargo sorting to extracellular vesicles","volume":"28","author":"Padilla","year":"2025","journal-title":"iScience"},{"key":"B81","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2023.1123841","article-title":"The role of glutamate receptors in the regulation of the tumor microenvironment","volume":"14","author":"Koda","year":"2023","journal-title":"Front Immunol"},{"key":"B82","doi-asserted-by":"publisher","DOI":"10.1016\/j.bioorg.2021.104625","article-title":"A newly identified small molecular compound acts as a protein kinase inhibitor to suppress metastasis of colorectal cancer","volume":"107","author":"Qi","year":"2021","journal-title":"Bioorg Chem"},{"key":"B83","doi-asserted-by":"publisher","DOI":"10.1084\/jem.181.5.1929","article-title":"Molecular behavior adapts to context: heparanase functions as an extracellular matrix-degrading enzyme or as a T cell adhesion molecule, depending on the local pH","volume":"181","author":"Gilat","year":"1995","journal-title":"J Exp Med"},{"key":"B84","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.m806266200","article-title":"Heparanase stimulation of protease expression implicates it as a master regulator of the aggressive tumor phenotype in myeloma","volume":"283","author":"Purushothaman","year":"2008","journal-title":"J Biol Chem"},{"key":"B85","doi-asserted-by":"publisher","DOI":"10.1038\/nature09303","article-title":"Role of Tet proteins in 5mC to 5hmC conversion, ES-cell self-renewal and inner cell mass specification","volume":"466","author":"Ito","year":"2010","journal-title":"Nature"},{"key":"B86","doi-asserted-by":"publisher","DOI":"10.1126\/science.1170116","article-title":"Conversion of 5-methylcytosine to 5-hydroxymethylcytosine in mammalian DNA by MLL partner TET1","volume":"324","author":"Tahiliani","year":"2009","journal-title":"Science"},{"key":"B87","doi-asserted-by":"publisher","DOI":"10.1126\/science.1210597","article-title":"Tet proteins can convert 5-methylcytosine to 5-formylcytosine and 5-carboxylcytosine","volume":"333","author":"Ito","year":"2011","journal-title":"Science"},{"key":"B88","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gks155","article-title":"Recognition and potential mechanisms for replication and erasure of cytosine hydroxymethylation","volume":"40","author":"Hashimoto","year":"2012","journal-title":"Nucleic Acids Res"},{"key":"B89","doi-asserted-by":"publisher","first-page":"7","DOI":"10.1186\/s13148-022-01404-6","article-title":"Vitamin C boosts DNA demethylation in TET2 germline mutation carriers","volume":"15","author":"Taira","year":"2023","journal-title":"Clin Epigenet"},{"key":"B90","doi-asserted-by":"publisher","DOI":"10.1093\/jleuko\/qiad118","article-title":"The effect of leptin on trained innate immunity and on systemic inflammation in subjects with obesity","volume":"115","author":"Flores Gomez","year":"2024","journal-title":"J Leukocyte Biol"},{"key":"B91","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.1200857","article-title":"Molecular mechanisms responsible for the selective and low-grade induction of proinflammatory mediators in murine macrophages by lipopolysaccharide","volume":"189","author":"Maitra","year":"2012","journal-title":"J Immunol"},{"key":"B92","doi-asserted-by":"publisher","first-page":"57","DOI":"10.1016\/j.yexcr.2018.02.017","article-title":"Human serum alters cell culture behavior and improves spheroid formation in comparison to fetal bovine serum","volume":"365","author":"Heger","year":"2018","journal-title":"Exp Cell Res"},{"key":"B93","doi-asserted-by":"publisher","DOI":"10.1038\/s41368-021-00146-0","article-title":"From bulk, single-cell to spatial RNA sequencing","volume":"13","author":"Li","year":"2021","journal-title":"Int J Oral Sci"},{"key":"B94","doi-asserted-by":"crossref","DOI":"10.1007\/978-1-61779-316-5_18","article-title":"Combined chromatin immunoprecipitation and bisulfite methylation sequencing analysis","volume-title":"Epigenetics Protocols","author":"Li","year":"2011"}],"container-title":["Frontiers in Immunology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fimmu.2026.1825431\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,20]],"date-time":"2026-07-20T04:21:10Z","timestamp":1784521270000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fimmu.2026.1825431\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,20]]},"references-count":94,"alternative-id":["10.3389\/fimmu.2026.1825431"],"URL":"https:\/\/doi.org\/10.3389\/fimmu.2026.1825431","relation":{},"ISSN":["1664-3224"],"issn-type":[{"value":"1664-3224","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,20]]},"article-number":"1825431"}}