{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T05:19:32Z","timestamp":1792041572912,"version":"4.4.0"},"reference-count":192,"publisher":"MDPI AG","issue":"1","license":[{"start":{"date-parts":[[2025,12,28]],"date-time":"2025-12-28T00:00:00Z","timestamp":1766880000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Recovery and Resilience Facility of the European Union"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Nutrients"],"abstract":"<jats:p>Iron constitutes an essential micronutrient in living organisms. All iron is absorbed into the body through dietary intake, except for exogenous therapeutic sources. Dietary iron is typically categorized as either heme or nonheme iron. Nonheme iron is essential for regulating iron in the body, as it exists in various forms, including soluble iron, storage iron within ferritin, and iron found in the catalytic centers of a wide range of proteins. Iron homeostasis is carefully managed to ensure that sufficient iron is available for critical biological processes while preventing the harmful effects that can arise from excess iron. The small peptide hormone hepcidin is the main regulator of iron homeostasis. Hepcidin and other iron regulatory molecules are regulated by various signaling pathways, such as IL-6\/JAK-STAT, BMP\/SMAD, and MAPK. Alterations in regulatory pathways may occur in response to iron accumulation or deficiency. Iron overload in the body can activate JAK\/STAT, BMP\/SMAD and MAPK pathways, leading to the initiation hepcidin synthesis. Conversely, in iron deficiency, as in hypoxic conditions or EPO-mediated signaling pathways, HAMP synthesis in the nucleus is reduced. Thus, this review provides an update on the possible regulatory pathways that play a role in iron regulation and may be potential therapeutic targets.<\/jats:p>","DOI":"10.3390\/nu18010109","type":"journal-article","created":{"date-parts":[[2025,12,28]],"date-time":"2025-12-28T23:54:36Z","timestamp":1766966076000},"page":"109","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":9,"title":["Iron, the Essential Micronutrient: A Comprehensive Review of Regulatory Pathways of Iron Metabolism"],"prefix":"10.3390","volume":"18","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-9315-082X","authenticated-orcid":false,"given":"Adrienn","family":"Horv\u00e1th","sequence":"first","affiliation":[{"name":"Department of Pharmaceutical Biology, Faculty of Pharmacy, University of P\u00e9cs, H-7624 P\u00e9cs, Hungary"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kitti","family":"Tam\u00e1si","sequence":"additional","affiliation":[{"name":"Department of Pharmaceutical Biology, Faculty of Pharmacy, University of P\u00e9cs, H-7624 P\u00e9cs, Hungary"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3340-0102","authenticated-orcid":false,"given":"Ram\u00f3na","family":"Pap","sequence":"additional","affiliation":[{"name":"Department of Pharmaceutical Biology, Faculty of Pharmacy, University of P\u00e9cs, H-7624 P\u00e9cs, Hungary"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5374-6680","authenticated-orcid":false,"given":"Gergely","family":"J\u00e1nosa","sequence":"additional","affiliation":[{"name":"Department of Pharmaceutical Biology, Faculty of Pharmacy, University of P\u00e9cs, H-7624 P\u00e9cs, Hungary"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5012-3242","authenticated-orcid":false,"given":"Edina","family":"Pandur","sequence":"additional","affiliation":[{"name":"Department of Pharmaceutical Biology, Faculty of Pharmacy, University of P\u00e9cs, H-7624 P\u00e9cs, Hungary"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2025,12,28]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1016\/j.cell.2010.06.028","article-title":"Two to Tango: Regulation of Mammalian Iron Metabolism","volume":"142","author":"Hentze","year":"2010","journal-title":"Cell"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"2012","DOI":"10.1111\/j.1742-4658.2009.06937.x","article-title":"A-1 Antitrypsin Binds Preprohepcidin Intracellularly and Prohepcidin in the Serum","volume":"276","author":"Pandur","year":"2009","journal-title":"FEBS J."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1016\/j.tcb.2006.12.003","article-title":"Iron Uptake and Metabolism in the New Millennium","volume":"17","author":"Dunn","year":"2007","journal-title":"Trends Cell Biol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"271","DOI":"10.1038\/s41392-024-01969-z","article-title":"Iron Homeostasis and Ferroptosis in Human Diseases: Mechanisms and Therapeutic Prospects","volume":"9","author":"Ru","year":"2024","journal-title":"Signal Transduct. Target. Ther."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"393","DOI":"10.1016\/j.fertnstert.2016.04.002","article-title":"Endometrial Expression and in Vitro Modulation of the Iron Transporter Divalent Metal Transporter-1: Implications for Endometriosis","volume":"106","author":"Devoto","year":"2016","journal-title":"Fertil. Steril."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1016\/j.gene.2005.07.020","article-title":"Hepcidin: Iron-Hormone and Anti-Microbial Peptide","volume":"364","author":"Muckenthaler","year":"2005","journal-title":"Gene"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"301","DOI":"10.1042\/BJ20121466","article-title":"Prohepcidin Binds to the HAMP Promoter and Autoregulates Its Own Expression","volume":"451","author":"Pandur","year":"2013","journal-title":"Biochem. J."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"5195","DOI":"10.1182\/blood-2007-09-111567","article-title":"Hemojuvelin Regulates Hepcidin Expression via a Selective Subset of BMP Ligands and Receptors Independently of Neogenin","volume":"111","author":"Xia","year":"2008","journal-title":"Blood"},{"key":"ref_9","doi-asserted-by":"crossref","unstructured":"Willemetz, A., Beatty, S., Richer, E., Rubio, A., Auriac, A., Milkereit, R.J., Thibaudeau, O., Vaulont, S., Malo, D., and Canonne-Hergaux, F. (2017). Iron- and Hepcidin-Independent Downregulation of the Iron Exporter Ferroportin in Macrophages during Salmonella Infection. Front. Immunol., 8.","DOI":"10.3389\/fimmu.2017.00498"},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Pandur, E., Varga, E., Tam\u00e1si, K., Pap, R., Nagy, J., and Sipos, K. (2018). Effect of Inflammatory Mediators Lipopolysaccharide and Lipoteichoic Acid on Iron Metabolism of Differentiated SH-SY5Y Cells Alters in the Presence of BV-2 Microglia. Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20010017"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"985","DOI":"10.1007\/s10571-019-00694-4","article-title":"Fractalkine Induces Hepcidin Expression of BV-2 Microglia and Causes Iron Accumulation in SH-SY5Y Cells","volume":"39","author":"Pandur","year":"2019","journal-title":"Cell. Mol. Neurobiol."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"112583","DOI":"10.1016\/j.yexcr.2021.112583","article-title":"Fractalkine Enhances Endometrial Receptivity and Activates Iron Transport towards Trophoblast Cells in an in Vitro Co-Culture System of HEC-1A and JEG-3 Cells","volume":"403","author":"Pandur","year":"2021","journal-title":"Exp. Cell Res."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"1830","DOI":"10.1111\/j.1471-4159.1989.tb07264.x","article-title":"Increased Nigral Iron Content and Alterations in Other Metal Ions Occurring in Brain in Parkinson\u2019s Disease","volume":"52","author":"Dexter","year":"1989","journal-title":"J. Neurochem."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"725370","DOI":"10.1155\/2015\/725370","article-title":"Inflammation, Iron, Energy Failure, and Oxidative Stress in the Pathogenesis of Multiple Sclerosis","volume":"2015","author":"Haider","year":"2015","journal-title":"Oxid. Med. Cell. Longev."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"1721","DOI":"10.1152\/physrev.00008.2013","article-title":"Systemic Iron Homeostasis","volume":"93","author":"Ganz","year":"2013","journal-title":"Physiol. Rev."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1182\/asheducation-2009.1.207","article-title":"Molecular Mechanisms of Normal Iron Homeostasis","volume":"2009","author":"Zhang","year":"2009","journal-title":"Hematology"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"299","DOI":"10.1016\/S1097-2765(00)80425-6","article-title":"A Novel Duodenal Iron-Regulated Transporter, IREG1, Implicated in the Basolateral Transfer of Iron to the Circulation","volume":"5","author":"McKie","year":"2000","journal-title":"Mol. Cell"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"104","DOI":"10.1016\/j.biosystems.2008.12.003","article-title":"New Insights into the Human Body Iron Metabolism Analyzed by a Petri Net Based Approach","volume":"96","author":"Sackmann","year":"2009","journal-title":"Biosystems"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"1037","DOI":"10.1172\/JCI0215686","article-title":"The Gene Encoding the Iron Regulatory Peptide Hepcidin Is Regulated by Anemia, Hypoxia, and Inflammation","volume":"110","author":"Nicolas","year":"2002","journal-title":"J. Clin. Investig."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"2090","DOI":"10.1126\/science.1104742","article-title":"Hepcidin Regulates Cellular Iron Efflux by Binding to Ferroportin and Inducing Its Internalization","volume":"306","author":"Nemeth","year":"2004","journal-title":"Science"},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Charlebois, E., and Pantopoulos, K. (2023). Nutritional Aspects of Iron in Health and Disease. Nutrients, 15.","DOI":"10.20944\/preprints202304.0859.v1"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"169","DOI":"10.1097\/MOH.0b013e3282f73335","article-title":"Iron Regulation and Erythropoiesis","volume":"15","author":"Nemeth","year":"2008","journal-title":"Curr. Opin. Hematol."},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Saunders, A.V., Craig, W.J., Baines, S.K., and Posen, J.S. (2013). Iron and Vegetarian Diets. Med. J. Aust., 199.","DOI":"10.5694\/mja11.11494"},{"key":"ref_24","doi-asserted-by":"crossref","unstructured":"Mascitelli, L., Goldstein, M.R., and Zacharski, L.R. (2015). The Mediterranean Diet and Body Iron Stores. The Mediterranean Diet, Elsevier.","DOI":"10.1016\/B978-0-12-407849-9.00024-5"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"1832","DOI":"10.1056\/NEJMra1401038","article-title":"Iron-Deficiency Anemia","volume":"372","author":"Camaschella","year":"2015","journal-title":"N. Engl. J. Med."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"e108","DOI":"10.1002\/hem3.108","article-title":"Recommendations for Diagnosis, Treatment, and Prevention of Iron Deficiency and Iron Deficiency Anemia","volume":"8","author":"Iolascon","year":"2024","journal-title":"HemaSphere"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"102911","DOI":"10.1016\/j.cpcardiol.2024.102911","article-title":"Thalassemia and Iron Overload Cardiomyopathy: Pathophysiological Insights, Clinical Implications, and Management Strategies","volume":"50","author":"Ansharullah","year":"2025","journal-title":"Curr. Probl. Cardiol."},{"key":"ref_28","unstructured":"Moustarah, F., and Daley, S.F. (2025). Dietary Iron, StatPearls."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"803","DOI":"10.1016\/j.dld.2013.02.019","article-title":"Iron Deficiency: From Diagnosis to Treatment","volume":"45","author":"Polin","year":"2013","journal-title":"Dig. Liver Dis."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"567","DOI":"10.1056\/NEJM199409013310902","article-title":"Efficacy of Deferoxamine in Preventing Complications of Iron Overload in Patients with Thalassemia Major","volume":"331","author":"Brittenham","year":"1994","journal-title":"N. Engl. J. Med."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"938","DOI":"10.1046\/j.1365-2141.2003.04297.x","article-title":"Monitoring Long-term Efficacy of Iron Chelation Therapy by Deferiprone and Desferrioxamine in Patients with \u0392-thalassaemia Major: Application of SQUID Biomagnetic Liver Susceptometry","volume":"121","author":"Fischer","year":"2003","journal-title":"Br. J. Haematol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1016\/j.bcmd.2015.11.002","article-title":"Optimising Iron Chelation Therapy with Deferasirox for Non-Transfusion-Dependent Thalassaemia Patients: 1-Year Results from the THETIS Study","volume":"57","author":"Taher","year":"2016","journal-title":"Blood Cells Mol. Dis."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"49","DOI":"10.4103\/0976-0105.177706","article-title":"Quality of Life, Clinical Effectiveness, and Satisfaction in Patients with Beta Thalassemia Major and Sickle Cell Anemia Receiving Deferasirox Chelation Therapy","volume":"7","author":"Senol","year":"2016","journal-title":"J. Basic Clin. Pharm."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"4911205","DOI":"10.1155\/2022\/4911205","article-title":"Iron Chelators in Treatment of Iron Overload","volume":"2022","author":"Entezari","year":"2022","journal-title":"J. Toxicol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1016\/j.tips.2014.01.004","article-title":"The Pathophysiology and Pharmacology of Hepcidin","volume":"35","author":"Ruchala","year":"2014","journal-title":"Trends Pharmacol. Sci."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"92","DOI":"10.1111\/ijlh.12358","article-title":"Hepcidin and Iron Disorders: New Biology and Clinical Approaches","volume":"37","author":"Arezes","year":"2015","journal-title":"Int. J. Lab. Hematol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"126","DOI":"10.3945\/an.116.013961","article-title":"Regulation of the Iron Homeostatic Hormone Hepcidin","volume":"8","author":"Sangkhae","year":"2017","journal-title":"Adv. Nutr."},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Saad, H.K.M., Abd Rahman, A.A., Ab Ghani, A.S., Taib, W.R.W., Ismail, I., Johan, M.F., Al-Wajeeh, A.S., and Al-Jamal, H.A.N. (2022). Activation of STAT and SMAD Signaling Induces Hepcidin Re-Expression as a Therapeutic Target for \u03b2-Thalassemia Patients. Biomedicines, 10.","DOI":"10.3390\/biomedicines10010189"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1016\/j.cmet.2007.11.016","article-title":"The Transferrin Receptor Modulates Hfe-Dependent Regulation of Hepcidin Expression","volume":"7","author":"Schmidt","year":"2008","journal-title":"Cell Metab."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"2473","DOI":"10.1089\/ars.2011.4271","article-title":"Mammalian Iron Homeostasis in Health and Disease: Uptake, Storage, Transport, and Molecular Mechanisms of Action","volume":"18","author":"Lawen","year":"2013","journal-title":"Antioxid. Redox Signal."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1172\/JCI11830","article-title":"NF-\u039aB: A Key Role in Inflammatory Diseases","volume":"107","author":"Tak","year":"2001","journal-title":"J. Clin. Investig."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"9891689","DOI":"10.34133\/2022\/9891689","article-title":"CCAAT\/Enhancer-Binding Proteins in Fibrosis: Complex Roles Beyond Conventional Understanding","volume":"2022","author":"Wang","year":"2022","journal-title":"Research"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"1224","DOI":"10.1007\/s11064-021-03322-0","article-title":"IL-6 Regulates Hepcidin Expression Via the BMP\/SMAD Pathway by Altering BMP6, TMPRSS6 and TfR2 Expressions at Normal and Inflammatory Conditions in BV2 Microglia","volume":"46","author":"Varga","year":"2021","journal-title":"Neurochem. Res."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"801","DOI":"10.1016\/j.jhep.2007.12.021","article-title":"2-Oxoglutarate-Dependent Oxygenases Control Hepcidin Gene Expression","volume":"48","author":"Braliou","year":"2008","journal-title":"J. Hepatol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"4090","DOI":"10.1074\/jbc.M110.173096","article-title":"Regulation of Type II Transmembrane Serine Proteinase TMPRSS6 by Hypoxia-Inducible Factors","volume":"286","author":"Lakhal","year":"2011","journal-title":"J. Biol. Chem."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"6561","DOI":"10.1074\/jbc.M413248200","article-title":"Hypoxia-Enhanced Expression of the Proprotein Convertase Furin Is Mediated by Hypoxia-Inducible Factor-1","volume":"280","author":"McMahon","year":"2005","journal-title":"J. Biol. Chem."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"291","DOI":"10.1002\/hep.25615","article-title":"Inhibition of Hepcidin Transcription by Growth Factors","volume":"56","author":"Goodnough","year":"2012","journal-title":"Hepatology"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"398","DOI":"10.1016\/j.gene.2012.09.060","article-title":"Estrogen Regulates Iron Homeostasis through Governing Hepatic Hepcidin Expression via an Estrogen Response Element","volume":"511","author":"Hou","year":"2012","journal-title":"Gene"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"1473","DOI":"10.1182\/blood-2018-06-857995","article-title":"Erythroferrone Inhibits the Induction of Hepcidin by BMP6","volume":"132","author":"Arezes","year":"2018","journal-title":"Blood"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"112","DOI":"10.1016\/j.jnutbio.2012.02.017","article-title":"Glucose Acts as a Regulator of Serum Iron by Increasing Serum Hepcidin Concentrations","volume":"24","author":"Aigner","year":"2013","journal-title":"J. Nutr. Biochem."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"1271","DOI":"10.1172\/JCI200420945","article-title":"IL-6 Mediates Hypoferremia of Inflammation by Inducing the Synthesis of the Iron Regulatory Hormone Hepcidin","volume":"113","author":"Nemeth","year":"2004","journal-title":"J. Clin. Investig."},{"key":"ref_52","doi-asserted-by":"crossref","unstructured":"Vogt, A.-C.S., Arsiwala, T., Mohsen, M., Vogel, M., Manolova, V., and Bachmann, M.F. (2021). On Iron Metabolism and Its Regulation. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms22094591"},{"key":"ref_53","doi-asserted-by":"crossref","unstructured":"Pandur, E., Tam\u00e1si, K., Pap, R., J\u00e1nosa, G., and Sipos, K. (2022). Modulatory Effects of Fractalkine on Inflammatory Response and Iron Metabolism of Lipopolysaccharide and Lipoteichoic Acid-Activated THP-1 Macrophages. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms23052629"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"222","DOI":"10.1016\/j.molimm.2017.02.018","article-title":"The Role of Kupffer Cells in Hepatic Diseases","volume":"85","author":"Li","year":"2017","journal-title":"Mol. Immunol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"10275","DOI":"10.1074\/jbc.M116.770974","article-title":"Interleukin-1\u03b2 (IL-1\u03b2) Transcriptionally Activates Hepcidin by Inducing CCAAT Enhancer-Binding Protein \u03b4 (C\/EBP\u03b4) Expression in Hepatocytes","volume":"292","author":"Kanamori","year":"2017","journal-title":"J. Biol. Chem."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"9263","DOI":"10.1073\/pnas.0403108101","article-title":"The IL-6- and Lipopolysaccharide-Induced Transcription of Hepcidin in HFE-, Transferrin Receptor 2-, and \u03b2 2 -Microglobulin-Deficient Hepatocytes","volume":"101","author":"Lee","year":"2004","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"521","DOI":"10.3748\/wjg.v25.i5.521","article-title":"Iron and Liver Fibrosis: Mechanistic and Clinical Aspects","volume":"25","author":"Mehta","year":"2019","journal-title":"World J. Gastroenterol."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"4946","DOI":"10.1073\/pnas.88.11.4946","article-title":"Iron-Independent Induction of Ferritin H Chain by Tumor Necrosis Factor","volume":"88","author":"Miller","year":"1991","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"700","DOI":"10.1074\/jbc.270.2.700","article-title":"Induction of Ferritin Synthesis by Oxidative Stress","volume":"270","author":"Cairo","year":"1995","journal-title":"J. Biol. Chem."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"2137","DOI":"10.1016\/j.biocel.2004.02.007","article-title":"Transferrin Receptor 1","volume":"36","author":"Aisen","year":"2004","journal-title":"Int. J. Biochem. Cell Biol."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"544","DOI":"10.1016\/j.jhep.2005.07.028","article-title":"Regulatory Effects of Tumor Necrosis Factor-Alpha and Interleukin-6 on HAMP Expression in Iron Loaded Rat Hepatocytes","volume":"44","author":"Dzikaite","year":"2006","journal-title":"J. Hepatol."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"725","DOI":"10.1080\/00365520212494","article-title":"Iron Overload Decreases the Protective Effect of Tumour Necrosis Factor-\u03b1 on Rat Hepatocytes Exposed to Oxidative Stress","volume":"37","author":"Hagen","year":"2002","journal-title":"Scand. J. Gastroenterol."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"1090","DOI":"10.1016\/j.bcp.2006.07.003","article-title":"Mechanisms of Crosstalk between TNF-Induced NF-\u039aB and JNK Activation in Hepatocytes","volume":"72","author":"Wullaert","year":"2006","journal-title":"Biochem. Pharmacol."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"3034","DOI":"10.3390\/nu5083034","article-title":"Out of Balance\u2014Systemic Iron Homeostasis in Iron-Related Disorders","volume":"5","author":"Steinbicker","year":"2013","journal-title":"Nutrients"},{"key":"ref_65","doi-asserted-by":"crossref","unstructured":"Core, A.B., Canali, S., and Babitt, J.L. (2014). Hemojuvelin and Bone Morphogenetic Protein (BMP) Signaling in Iron Homeostasis. Front. Pharmacol., 5.","DOI":"10.3389\/fphar.2014.00104"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"531","DOI":"10.1038\/ng1777","article-title":"Bone Morphogenetic Protein Signaling by Hemojuvelin Regulates Hepcidin Expression","volume":"38","author":"Babitt","year":"2006","journal-title":"Nat. Genet."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"1052","DOI":"10.1016\/j.jhep.2012.06.015","article-title":"The Hemochromatosis Proteins HFE, TfR2, and HJV Form a Membrane-Associated Protein Complex for Hepcidin Regulation","volume":"57","author":"Hentze","year":"2012","journal-title":"J. Hepatol."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"1737","DOI":"10.1002\/hep.22180","article-title":"The Role of Hfe in Transferrin-Bound Iron Uptake by Hepatocytes","volume":"47","author":"Chua","year":"2008","journal-title":"Hepatology"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"332","DOI":"10.1182\/blood.2022018740","article-title":"Transferrin Receptor 1: Keeper of HFE","volume":"141","author":"Parrow","year":"2023","journal-title":"Blood"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"1949","DOI":"10.1182\/blood.V98.6.1949","article-title":"Regulation of Expression of Murine Transferrin Receptor 2","volume":"98","author":"Kawabata","year":"2001","journal-title":"Blood"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"217","DOI":"10.1016\/j.cmet.2009.01.010","article-title":"Interaction of the Hereditary Hemochromatosis Protein HFE with Transferrin Receptor 2 Is Required for Transferrin-Induced Hepcidin Expression","volume":"9","author":"Gao","year":"2009","journal-title":"Cell Metab."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"4224","DOI":"10.1182\/blood-2011-03-339952","article-title":"Perturbation of Hepcidin Expression by BMP Type I Receptor Deletion Induces Iron Overload in Mice","volume":"118","author":"Steinbicker","year":"2011","journal-title":"Blood"},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"11197","DOI":"10.1074\/jbc.M608441200","article-title":"Restoration of Liver Mass after Injury Requires Proliferative and Not Embryonic Transcriptional Patterns","volume":"282","author":"Otu","year":"2007","journal-title":"J. Biol. Chem."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"4362","DOI":"10.1091\/mbc.e09-07-0539","article-title":"The Transforming Growth Factor-\u03b2 Type III Receptor Mediates Distinct Subcellular Trafficking and Downstream Signaling of Activin-like Kinase (ALK)3 and ALK6 Receptors","volume":"20","author":"Lee","year":"2009","journal-title":"Mol. Biol. Cell"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"966","DOI":"10.1093\/ndt\/gfaa203","article-title":"The Role of Activin: The Other Side of Chronic Kidney Disease\u2013Mineral Bone Disorder?","volume":"36","author":"Cianciolo","year":"2021","journal-title":"Nephrol. Dial. Transplant."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"e00192","DOI":"10.1042\/BSR20150014","article-title":"Hepcidin: Regulation of the Master Iron Regulator","volume":"35","author":"Rishi","year":"2015","journal-title":"Biosci. Rep."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"77","DOI":"10.1146\/annurev-nutr-071813-105646","article-title":"Bone Morphogenetic Proteins as Regulators of Iron Metabolism","volume":"34","author":"Parrow","year":"2014","journal-title":"Annu. Rev. Nutr."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"76","DOI":"10.1016\/j.bbadis.2012.08.013","article-title":"Smad6 and Smad7 Are Co-Regulated with Hepcidin in Mouse Models of Iron Overload","volume":"1832","author":"Sparla","year":"2013","journal-title":"Biochim. Biophys. Acta. Mol. Basis Dis."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"4099","DOI":"10.1074\/jbc.M111.310789","article-title":"BMPER Protein Is a Negative Regulator of Hepcidin and Is Up-Regulated in Hypotransferrinemic Mice","volume":"287","author":"Patel","year":"2012","journal-title":"J. Biol. Chem."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"12547","DOI":"10.1074\/jbc.M608788200","article-title":"Evidence That Inhibition of Hemojuvelin Shedding in Response to Iron Is Mediated through Neogenin","volume":"282","author":"Zhang","year":"2007","journal-title":"J. Biol. Chem."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"14122","DOI":"10.1074\/jbc.M410034200","article-title":"DRAGON, a Bone Morphogenetic Protein Co-Receptor","volume":"280","author":"Samad","year":"2005","journal-title":"J. Biol. Chem."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"29820","DOI":"10.1074\/jbc.M503511200","article-title":"Repulsive Guidance Molecule (RGMa), a DRAGON Homologue, Is a Bone Morphogenetic Protein Co-Receptor","volume":"280","author":"Babitt","year":"2005","journal-title":"J. Biol. Chem."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"33885","DOI":"10.1074\/jbc.M506207200","article-title":"Interaction of Hemojuvelin with Neogenin Results in Iron Accumulation in Human Embryonic Kidney 293 Cells","volume":"280","author":"Zhang","year":"2005","journal-title":"J. Biol. Chem."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"12322","DOI":"10.1074\/jbc.M116.721191","article-title":"Neogenin Facilitates the Induction of Hepcidin Expression by Hemojuvelin in the Liver","volume":"291","author":"Zhao","year":"2016","journal-title":"J. Biol. Chem."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"840","DOI":"10.3324\/haematol.2008.001867","article-title":"Matriptase-2 (TMPRSS6): A Proteolytic Regulator of Iron Homeostasis","volume":"94","author":"Ramsay","year":"2009","journal-title":"Haematologica"},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"21048","DOI":"10.1074\/jbc.M000636200","article-title":"Role of Hypoxia-Inducible Factor-1 in Transcriptional Activation of Ceruloplasmin by Iron Deficiency","volume":"275","author":"Mukhopadhyay","year":"2000","journal-title":"J. Biol. Chem."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"24142","DOI":"10.1074\/jbc.274.34.24142","article-title":"Transferrin Receptor Induction by Hypoxia","volume":"274","author":"Tacchini","year":"1999","journal-title":"J. Biol. Chem."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"3945","DOI":"10.1007\/s00018-012-1031-4","article-title":"Regulation of Iron Homeostasis by MicroRNAs","volume":"69","author":"Castoldi","year":"2012","journal-title":"Cell. Mol. Life Sci."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"8725","DOI":"10.1073\/pnas.88.19.8725","article-title":"Cell-Type-Specific and Hypoxia-Inducible Expression of the Human Erythropoietin Gene in Transgenic Mice","volume":"88","author":"Semenza","year":"1991","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"4635","DOI":"10.1172\/JCI63924","article-title":"Hypoxia-Inducible Factor Regulates Hepcidin via Erythropoietin-Induced Erythropoiesis","volume":"122","author":"Liu","year":"2012","journal-title":"J. Clin. Investig."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1038\/emm.2004.1","article-title":"Hypoxia-Inducible Factor (HIF-1)\u03b1: Its Protein Stability and Biological Functions","volume":"36","author":"Lee","year":"2004","journal-title":"Exp. Mol. Med."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"343","DOI":"10.1038\/nrm1366","article-title":"Oxygen Sensing by HIF Hydroxylases","volume":"5","author":"Schofield","year":"2004","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"1926","DOI":"10.1172\/JCI31370","article-title":"Regulation of Iron Homeostasis by the Hypoxia-Inducible Transcription Factors (HIFs)","volume":"117","author":"Peyssonnaux","year":"2007","journal-title":"J. Clin. Investig."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"2253","DOI":"10.1182\/blood-2002-02-0629","article-title":"Cellular Response to Hypoxia Involves Signaling via Smad Proteins","volume":"101","author":"Zhang","year":"2003","journal-title":"Blood"},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"399","DOI":"10.1016\/j.cmet.2005.10.010","article-title":"A Role of SMAD4 in Iron Metabolism through the Positive Regulation of Hepcidin Expression","volume":"2","author":"Wang","year":"2005","journal-title":"Cell Metab."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"4888","DOI":"10.1002\/jcp.30247","article-title":"Erythroferrone Structure, Function, and Physiology: Iron Homeostasis and Beyond","volume":"236","author":"Srole","year":"2021","journal-title":"J. Cell. Physiol."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"453","DOI":"10.1182\/blood.2019002620","article-title":"Erythroferrone Lowers Hepcidin by Sequestering BMP2\/6 Heterodimer from Binding to the BMP Type I Receptor ALK3","volume":"135","author":"Wang","year":"2020","journal-title":"Blood"},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"678","DOI":"10.1038\/ng.2996","article-title":"Identification of Erythroferrone as an Erythroid Regulator of Iron Metabolism","volume":"46","author":"Kautz","year":"2014","journal-title":"Nat. Genet."},{"key":"ref_99","doi-asserted-by":"crossref","unstructured":"Ikeda, Y., Tajima, S., Izawa-Ishizawa, Y., Kihira, Y., Ishizawa, K., Tomita, S., Tsuchiya, K., and Tamaki, T. (2012). Estrogen Regulates Hepcidin Expression via GPR30-BMP6-Dependent Signaling in Hepatocytes. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0040465"},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"138","DOI":"10.1016\/j.mce.2006.12.010","article-title":"GPR30: A G Protein-Coupled Receptor for Estrogen","volume":"265\u2013266","author":"Prossnitz","year":"2007","journal-title":"Mol. Cell. Endocrinol."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"934","DOI":"10.1016\/j.cellsig.2015.01.017","article-title":"Estrogen Contributes to Regulating Iron Metabolism through Governing Ferroportin Signaling via an Estrogen Response Element","volume":"27","author":"Qian","year":"2015","journal-title":"Cell. Signal."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"14","DOI":"10.1016\/j.pharmthera.2008.09.006","article-title":"PGRMC1 (Progesterone Receptor Membrane Component 1): A Targetable Protein with Multiple Functions in Steroid Signaling, P450 Activation and Drug Binding","volume":"121","author":"Rohe","year":"2009","journal-title":"Pharmacol. Ther."},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"389","DOI":"10.1172\/JCI83831","article-title":"Progesterone Receptor Membrane Component-1 Regulates Hepcidin Biosynthesis","volume":"126","author":"Li","year":"2016","journal-title":"J. Clin. Investig."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"280","DOI":"10.1111\/acel.12052","article-title":"Testosterone Administration Inhibits Hepcidin Transcription and Is Associated with Increased Iron Incorporation into Red Blood Cells","volume":"12","author":"Guo","year":"2013","journal-title":"Aging Cell"},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"683","DOI":"10.1002\/hep.26648","article-title":"Testosterone Perturbs Systemic Iron Balance Through Activation of Epidermal Growth Factor Receptor Signaling in the Liver and Repression of Hepcidin","volume":"59","author":"Latour","year":"2014","journal-title":"Hepatology"},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"263","DOI":"10.1309\/WWNE-KW2C-4KTW-PTJ5","article-title":"Histologic Features of the Liver in Insulin Resistance\u2013Associated Iron Overload","volume":"116","author":"Turlin","year":"2001","journal-title":"Am. J. Clin. Pathol."},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"123","DOI":"10.1016\/j.jhep.2008.03.011","article-title":"Hepcidin and Iron-Related Gene Expression in Subjects with Dysmetabolic Hepatic Iron Overload","volume":"49","author":"Barisani","year":"2008","journal-title":"J. Hepatol."},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"385","DOI":"10.1002\/dvg.20513","article-title":"Abnormal Glucose Metabolism in Heterozygous Mutant Mice for a Type I Receptor Required for BMP Signaling","volume":"47","author":"Scott","year":"2009","journal-title":"Genesis"},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1016\/j.cmet.2007.01.009","article-title":"BMP4-BMPR1A Signaling in \u03b2 Cells Is Required for and Augments Glucose-Stimulated Insulin Secretion","volume":"5","author":"Goulley","year":"2007","journal-title":"Cell Metab."},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"1282","DOI":"10.1007\/s00125-015-3568-x","article-title":"Bone Morphogenetic Protein 4 Inhibits Insulin Secretion from Rodent Beta Cells through Regulation of Calbindin1 Expression and Reduced Voltage-Dependent Calcium Currents","volume":"58","author":"Christensen","year":"2015","journal-title":"Diabetologia"},{"key":"ref_111","doi-asserted-by":"crossref","unstructured":"Gao, Q., Huan, C., Jia, Z., Cao, Q., Yuan, P., Li, X., Wang, C., Mao, Z., and Huo, W. (2024). SOCS3 Methylation Partially Mediated the Association of Exposure to Triclosan but Not Triclocarban with Type 2 Diabetes Mellitus: A Case-Control Study. Int. J. Mol. Sci., 25.","DOI":"10.3390\/ijms252212113"},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"1506","DOI":"10.2337\/db13-1195","article-title":"Hepcidin Is Directly Regulated by Insulin and Plays an Important Role in Iron Overload in Streptozotocin-Induced Diabetic Rats","volume":"63","author":"Wang","year":"2014","journal-title":"Diabetes"},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"4779","DOI":"10.7150\/thno.32543","article-title":"The MiR-561-5p\/CX 3 CL1 Signaling Axis Regulates Pulmonary Metastasis in Hepatocellular Carcinoma Involving CX 3 CR1 + Natural Killer Cells Infiltration","volume":"9","author":"Chen","year":"2019","journal-title":"Theranostics"},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"2611","DOI":"10.3390\/nu5072611","article-title":"Influence of MicroRNA on the Maintenance of Human Iron Metabolism","volume":"5","author":"Davis","year":"2013","journal-title":"Nutrients"},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"895","DOI":"10.1016\/j.cmet.2012.04.021","article-title":"Iron Homeostasis Regulates the Activity of the MicroRNA Pathway through Poly(C)-Binding Protein 2","volume":"15","author":"Li","year":"2012","journal-title":"Cell Metab."},{"key":"ref_116","doi-asserted-by":"crossref","first-page":"1386","DOI":"10.1172\/JCI44883","article-title":"The Liver-Specific MicroRNA MiR-122 Controls Systemic Iron Homeostasis in Mice","volume":"121","author":"Castoldi","year":"2011","journal-title":"J. Clin. Investig."},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"34110","DOI":"10.1074\/jbc.M112.356717","article-title":"Micromanaging Iron Homeostasis","volume":"287","author":"Yoshioka","year":"2012","journal-title":"J. Biol. Chem."},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"167748","DOI":"10.1016\/j.bbadis.2025.167748","article-title":"Hepatic MicroRNA-320 Restrains Ferroptosis to Mitigate Acute-on-Chronic Alcohol-Induced Liver Injury","volume":"1871","author":"Li","year":"2025","journal-title":"Biochim. Biophys. Acta Mol. Basis Dis."},{"key":"ref_119","doi-asserted-by":"crossref","first-page":"1244","DOI":"10.3324\/haematol.2009.020685","article-title":"Regulation of Divalent Metal Transporter 1 (DMT1) Non-IRE Isoform by the MicroRNA Let-7d in Erythroid Cells","volume":"95","author":"Andolfo","year":"2010","journal-title":"Haematologica"},{"key":"ref_120","doi-asserted-by":"crossref","unstructured":"Sangokoya, C., Doss, J.F., and Chi, J.-T. (2013). Iron-Responsive MiR-485-3p Regulates Cellular Iron Homeostasis by Targeting Ferroportin. PLoS Genet., 9.","DOI":"10.1371\/journal.pgen.1003408"},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1186\/s13098-022-00919-5","article-title":"Alterations of the Expression Levels of Glucose, Inflammation, and Iron Metabolism Related MiRNAs and Their Target Genes in the Hypothalamus of STZ-Induced Rat Diabetes Model","volume":"14","author":"Pandur","year":"2022","journal-title":"Diabetol. Metab. Syndr."},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"21199","DOI":"10.1038\/s41598-025-04730-w","article-title":"MiR-125b-5p Ameliorates Ventilator-Induced Lung Injury in Rats by Suppressing Ferroptosis via the Regulation of the Keap1\/Nrf2\/GPX4 Signaling Pathway","volume":"15","author":"Zhang","year":"2025","journal-title":"Sci. Rep."},{"key":"ref_123","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1007\/s10495-023-01890-w","article-title":"The Interplay of MiRNAs and Ferroptosis in Diseases Related to Iron Overload","volume":"29","author":"Jin","year":"2024","journal-title":"Apoptosis"},{"key":"ref_124","first-page":"23","article-title":"Curcumenol Triggered Ferroptosis in Lung Cancer Cells via LncRNA H19\/MiR-19b-3p\/FTH1 Axis","volume":"13","author":"Zhang","year":"2022","journal-title":"Bioact. Mater."},{"key":"ref_125","doi-asserted-by":"crossref","first-page":"26137","DOI":"10.18632\/aging.203801","article-title":"MiR-545 Promotes Colorectal Cancer by Inhibiting Transferring in the Non-Normal Ferroptosis Signaling","volume":"13","author":"Zheng","year":"2021","journal-title":"Aging"},{"key":"ref_126","doi-asserted-by":"crossref","first-page":"e13235","DOI":"10.1111\/acel.13235","article-title":"Targeting MiR-124\/Ferroportin Signaling Ameliorated Neuronal Cell Death through Inhibiting Apoptosis and Ferroptosis in Aged Intracerebral Hemorrhage Murine Model","volume":"19","author":"Bao","year":"2020","journal-title":"Aging Cell"},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"234","DOI":"10.1007\/s10238-025-01766-0","article-title":"MicroRNAs at the Crossroads of Exercise and Ferroptosis: A Regulatory Bridge","volume":"25","author":"Lin","year":"2025","journal-title":"Clin. Exp. Med."},{"key":"ref_128","doi-asserted-by":"crossref","first-page":"319","DOI":"10.3390\/pathophysiology29020023","article-title":"Serum Ferritin in Metabolic Syndrome\u2014Mechanisms and Clinical Applications","volume":"29","author":"Srivastav","year":"2022","journal-title":"Pathophysiology"},{"key":"ref_129","doi-asserted-by":"crossref","unstructured":"Gonz\u00e1lez-Dom\u00ednguez, \u00c1., Visiedo-Garc\u00eda, F.M., Dom\u00ednguez-Riscart, J., Gonz\u00e1lez-Dom\u00ednguez, R., Mateos, R.M., and Lechuga-Sancho, A.M. (2020). Iron Metabolism in Obesity and Metabolic Syndrome. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21155529"},{"key":"ref_130","doi-asserted-by":"crossref","unstructured":"Wang, J.-W., Jin, C.-H., Ke, J.-F., Ma, Y.-L., Wang, Y.-J., Lu, J.-X., Li, M.-F., and Li, L.-X. (2022). Serum Iron Is Closely Associated with Metabolic Dysfunction-Associated Fatty Liver Disease in Type 2 Diabetes: A Real-World Study. Front. Endocrinol., 13.","DOI":"10.3389\/fendo.2022.942412"},{"key":"ref_131","doi-asserted-by":"crossref","unstructured":"Abu AlSel, B.T., Mahmoud, A.A., Hamed, E.O., Hakim, N.A., Sindi, A.A.A., Jawad, N.M.M., Gusti, A.M.T., Fawzy, M.S., and Abd El-Fadeal, N.M. (2024). Iron Homeostasis-Related Parameters and Hepcidin\/Ferritin Ratio: Emerging Sex-Specific Predictive Markers for Metabolic Syndrome. Metabolites, 14.","DOI":"10.3390\/metabo14090473"},{"key":"ref_132","doi-asserted-by":"crossref","first-page":"100872","DOI":"10.1016\/j.mam.2020.100872","article-title":"Regulation of Iron Homeostasis: Lessons from Mouse Models","volume":"75","author":"Altamura","year":"2020","journal-title":"Mol. Aspects Med."},{"key":"ref_133","doi-asserted-by":"crossref","first-page":"738","DOI":"10.2353\/ajpath.2008.070097","article-title":"Iron Depletion by Deferoxamine Up-Regulates Glucose Uptake and Insulin Signaling in Hepatoma Cells and in Rat Liver","volume":"172","author":"Dongiovanni","year":"2008","journal-title":"Am. J. Pathol."},{"key":"ref_134","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1111\/eci.12032","article-title":"Iron Homeostasis in the Metabolic Syndrome","volume":"43","author":"Datz","year":"2013","journal-title":"Eur. J. Clin. Investig."},{"key":"ref_135","doi-asserted-by":"crossref","first-page":"1374","DOI":"10.1093\/ajcn\/87.5.1374","article-title":"Pathways Underlying Iron Accumulation in Human Nonalcoholic Fatty Liver Disease","volume":"87","author":"Aigner","year":"2008","journal-title":"Am. J. Clin. Nutr."},{"key":"ref_136","doi-asserted-by":"crossref","first-page":"1219","DOI":"10.1016\/j.jhep.2009.01.029","article-title":"Iron Absorption in Dysmetabolic Iron Overload Syndrome Is Decreased and Correlates with Increased Plasma Hepcidin","volume":"50","author":"Ruivard","year":"2009","journal-title":"J. Hepatol."},{"key":"ref_137","doi-asserted-by":"crossref","first-page":"130","DOI":"10.1186\/s10020-022-00558-6","article-title":"The Role of Iron Metabolism in Chronic Diseases Related to Obesity","volume":"28","author":"Qiu","year":"2022","journal-title":"Mol. Med."},{"key":"ref_138","doi-asserted-by":"crossref","first-page":"150","DOI":"10.1111\/j.1467-789X.2011.00938.x","article-title":"The Relationship between Obesity and Hypoferraemia in Adults: A Systematic Review","volume":"13","author":"Cheng","year":"2012","journal-title":"Obes. Rev."},{"key":"ref_139","doi-asserted-by":"crossref","first-page":"1081","DOI":"10.1111\/obr.12323","article-title":"Obesity and Iron Deficiency: A Quantitative Meta-analysis","volume":"16","author":"Zhao","year":"2015","journal-title":"Obes. Rev."},{"key":"ref_140","doi-asserted-by":"crossref","first-page":"e306","DOI":"10.1016\/j.lpm.2017.05.036","article-title":"Dysmetabolic Iron Overload Syndrome (DIOS)","volume":"46","author":"Deugnier","year":"2017","journal-title":"Presse Med."},{"key":"ref_141","doi-asserted-by":"crossref","first-page":"424","DOI":"10.1158\/1055-9965.EPI-08-0725","article-title":"Iron Overload Is Associated with Hepatic Oxidative Damage to DNA in Nonalcoholic Steatohepatitis","volume":"18","author":"Fujita","year":"2009","journal-title":"Cancer Epidemiol. Biomarkers Prev."},{"key":"ref_142","doi-asserted-by":"crossref","unstructured":"Smita, R.M., Shuvo, A.P.R., Raihan, S., Jahan, R., Simin, F.A., Rahman, A., Biswas, S., Salem, L., and Sagor, M.A.T. (2022). The Role of Mineral Deficiencies in Insulin Resistance and Obesity. Curr. Diabetes Rev., 18.","DOI":"10.2174\/1573399818666211117104626"},{"key":"ref_143","doi-asserted-by":"crossref","unstructured":"Kasai, S., Mimura, J., Ozaki, T., and Itoh, K. (2018). Emerging Regulatory Role of Nrf2 in Iron, Heme, and Hemoglobin Metabolism in Physiology and Disease. Front. Vet. Sci., 5.","DOI":"10.3389\/fvets.2018.00242"},{"key":"ref_144","doi-asserted-by":"crossref","unstructured":"Saha, S., Buttari, B., Panieri, E., Profumo, E., and Saso, L. (2020). An Overview of Nrf2 Signaling Pathway and Its Role in Inflammation. Molecules, 25.","DOI":"10.3390\/molecules25225474"},{"key":"ref_145","doi-asserted-by":"crossref","first-page":"1169","DOI":"10.1152\/physrev.00023.2017","article-title":"The KEAP1-NRF2 System: A Thiol-Based Sensor-Effector Apparatus for Maintaining Redox Homeostasis","volume":"98","author":"Yamamoto","year":"2018","journal-title":"Physiol. Rev."},{"key":"ref_146","doi-asserted-by":"crossref","first-page":"1247","DOI":"10.1111\/1440-1681.12665","article-title":"Bone Morphogenetic Protein 6\u2014A Possible New Player in Pathophysiology of Heart Failure","volume":"43","author":"Banach","year":"2016","journal-title":"Clin. Exp. Pharmacol. Physiol."},{"key":"ref_147","doi-asserted-by":"crossref","first-page":"2672","DOI":"10.1073\/pnas.1821022116","article-title":"Ferroptosis as a Target for Protection against Cardiomyopathy","volume":"116","author":"Fang","year":"2019","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_148","doi-asserted-by":"crossref","first-page":"486","DOI":"10.1161\/CIRCRESAHA.120.316509","article-title":"Loss of Cardiac Ferritin H Facilitates Cardiomyopathy via Slc7a11-Mediated Ferroptosis","volume":"127","author":"Fang","year":"2020","journal-title":"Circ. Res."},{"key":"ref_149","doi-asserted-by":"crossref","first-page":"598","DOI":"10.1016\/j.ebiom.2019.08.014","article-title":"New Insights into the Role of Iron in Inflammation and Atherosclerosis","volume":"47","author":"Cornelissen","year":"2019","journal-title":"EBioMedicine"},{"key":"ref_150","doi-asserted-by":"crossref","first-page":"126882","DOI":"10.1016\/j.jtemb.2021.126882","article-title":"Triad Role of Hepcidin, Ferroportin, and Nrf2 in Cardiac Iron Metabolism: From Health to Disease","volume":"69","author":"Jayakumar","year":"2022","journal-title":"J. Trace Elem. Med. Biol."},{"key":"ref_151","doi-asserted-by":"crossref","first-page":"1246","DOI":"10.1002\/2211-5463.70048","article-title":"Iron-dependent lysosomal LDL oxidation induces the expression of scavenger receptor A in human THP-1 monocytes","volume":"15","author":"Buchal","year":"2025","journal-title":"FEBS Open Bio."},{"key":"ref_152","doi-asserted-by":"crossref","first-page":"782","DOI":"10.1038\/s41419-021-04054-3","article-title":"Ferroptosis: The Potential Value Target in Atherosclerosis","volume":"12","author":"Ouyang","year":"2021","journal-title":"Cell Death Dis."},{"key":"ref_153","doi-asserted-by":"crossref","unstructured":"Medina-Leyte, D.J., Zepeda-Garc\u00eda, O., Dom\u00ednguez-P\u00e9rez, M., Gonz\u00e1lez-Garrido, A., Villarreal-Molina, T., and Jacobo-Albavera, L. (2021). Endothelial Dysfunction, Inflammation and Coronary Artery Disease: Potential Biomarkers and Promising Therapeutical Approaches. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms22083850"},{"key":"ref_154","doi-asserted-by":"crossref","first-page":"946","DOI":"10.1111\/jdi.13216","article-title":"Iron Metabolism and Type 2 Diabetes Mellitus: A Meta-analysis and Systematic Review","volume":"11","author":"Liu","year":"2020","journal-title":"J. Diabetes Investig."},{"key":"ref_155","doi-asserted-by":"crossref","unstructured":"Zhao, Z., Li, S., Liu, G., Yan, F., Ma, X., Huang, Z., and Tian, H. (2012). Body Iron Stores and Heme-Iron Intake in Relation to Risk of Type 2 Diabetes: A Systematic Review and Meta-Analysis. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0041641"},{"key":"ref_156","doi-asserted-by":"crossref","unstructured":"Ruze, R., Liu, T., Zou, X., Song, J., Chen, Y., Xu, R., Yin, X., and Xu, Q. (2023). Obesity and Type 2 Diabetes Mellitus: Connections in Epidemiology, Pathogenesis, and Treatments. Front. Endocrinol., 14.","DOI":"10.3389\/fendo.2023.1161521"},{"key":"ref_157","doi-asserted-by":"crossref","first-page":"978","DOI":"10.4239\/wjd.v6.i7.978","article-title":"Serum Hepcidin Concentrations and Type 2 Diabetes","volume":"6","author":"Aregbesola","year":"2015","journal-title":"World J. Diabetes"},{"key":"ref_158","doi-asserted-by":"crossref","first-page":"e06429","DOI":"10.1016\/j.heliyon.2021.e06429","article-title":"Body Weight and Its Influence on Hepcidin Levels in Patients with Type 2 Diabetes: A Systematic Review and Meta-Analysis of Clinical Studies","volume":"7","author":"Ndevahoma","year":"2021","journal-title":"Heliyon"},{"key":"ref_159","doi-asserted-by":"crossref","first-page":"2681","DOI":"10.1093\/eurheartj\/ehz112","article-title":"Atherosclerosis Is Aggravated by Iron Overload and Ameliorated by Dietary and Pharmacological Iron Restriction","volume":"41","author":"Vinchi","year":"2020","journal-title":"Eur. Heart J."},{"key":"ref_160","doi-asserted-by":"crossref","first-page":"RV17060","DOI":"10.5551\/jat.RV17060","article-title":"Crosstalk between Iron and Arteriosclerosis","volume":"29","author":"Naito","year":"2022","journal-title":"J. Atheroscler. Thromb."},{"key":"ref_161","doi-asserted-by":"crossref","first-page":"1567S","DOI":"10.3945\/ajcn.117.155812","article-title":"Iron Homeostasis during Pregnancy","volume":"106","author":"Fisher","year":"2017","journal-title":"Am. J. Clin. Nutr."},{"key":"ref_162","doi-asserted-by":"crossref","unstructured":"Zych-Krekora, K., Sylwestrzak, O., and Krekora, M. (2025). The Critical Role of Iron in Pregnancy, Puerperium, and Fetal Development. J. Clin. Med., 14.","DOI":"10.3390\/jcm14103482"},{"key":"ref_163","doi-asserted-by":"crossref","first-page":"625","DOI":"10.1172\/JCI127341","article-title":"Effects of Maternal Iron Status on Placental and Fetal Iron Homeostasis","volume":"130","author":"Sangkhae","year":"2019","journal-title":"J. Clin. Investig."},{"key":"ref_164","doi-asserted-by":"crossref","first-page":"1145","DOI":"10.1097\/01.AOG.0000238333.37423.ab","article-title":"Iron Intake and Risk of Ovulatory Infertility","volume":"108","author":"Chavarro","year":"2006","journal-title":"Obstet. Gynecol."},{"key":"ref_165","doi-asserted-by":"crossref","first-page":"61","DOI":"10.1080\/10715762.2023.2191813","article-title":"Iron-Loaded Extracellular Vesicles: Angel or Demon?","volume":"57","author":"Chen","year":"2023","journal-title":"Free Radic. Res."},{"key":"ref_166","doi-asserted-by":"crossref","first-page":"261","DOI":"10.1146\/annurev-med-043021-032816","article-title":"Hepcidin and Iron in Health and Disease","volume":"74","author":"Nemeth","year":"2023","journal-title":"Annu. Rev. Med."},{"key":"ref_167","doi-asserted-by":"crossref","first-page":"125","DOI":"10.1111\/j.1749-6632.2000.tb06223.x","article-title":"Anemia in Pregnancy","volume":"900","author":"Sifakis","year":"2000","journal-title":"Ann. N. Y. Acad. Sci."},{"key":"ref_168","doi-asserted-by":"crossref","first-page":"109","DOI":"10.1111\/tme.13125","article-title":"Understanding Hepcidin for Iron Management in Pregnancy","volume":"35","author":"Rosson","year":"2025","journal-title":"Transfus. Med."},{"key":"ref_169","doi-asserted-by":"crossref","unstructured":"Pap, R., Montsk\u00f3, G., J\u00e1nosa, G., Sipos, K., Kov\u00e1cs, G.L., and Pandur, E. (2020). Fractalkine Regulates HEC-1A\/JEG-3 Interaction by Influencing the Expression of Implantation-Related Genes in an In Vitro Co-Culture Model. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21093175"},{"key":"ref_170","doi-asserted-by":"crossref","first-page":"1756","DOI":"10.1089\/ars.2017.7176","article-title":"The Roles of NRF2 in Modulating Cellular Iron Homeostasis","volume":"29","author":"Kerins","year":"2018","journal-title":"Antioxid. Redox Signal"},{"key":"ref_171","doi-asserted-by":"crossref","unstructured":"Pandur, E., Pap, R., J\u00e1nosa, G., Horv\u00e1th, A., and Sipos, K. (2023). The Role of Fractalkine in the Regulation of Endometrial Iron Metabolism in Iron Deficiency. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms24129917"},{"key":"ref_172","doi-asserted-by":"crossref","unstructured":"Porras, C.A., and Rouault, T.A. (2022). Iron Homeostasis in the CNS: An Overview of the Pathological Consequences of Iron Metabolism Disruption. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms23094490"},{"key":"ref_173","doi-asserted-by":"crossref","unstructured":"Liu, H.T., Shie, J.H., Chen, S.H., Wang, Y.S., and Kuo, H.C. (2012). Differences in Mast Cell Infiltration, E-Cadherin, and Zonula Occludens-1 Expression between Patients with Overactive Bladder and Interstitial Cystitis\/Bladder Pain Syndrome. Urology, 80.","DOI":"10.1016\/j.urology.2012.01.047"},{"key":"ref_174","doi-asserted-by":"crossref","first-page":"1341","DOI":"10.1093\/brain\/awaa089","article-title":"Relationship between Cortical Iron and Tau Aggregation in Alzheimer\u2019s Disease","volume":"143","author":"Spotorno","year":"2020","journal-title":"Brain"},{"key":"ref_175","doi-asserted-by":"crossref","unstructured":"Cerasuolo, M., Di Meo, I., Auriemma, M.C., Trojsi, F., Maiorino, M.I., Cirillo, M., Esposito, F., Polito, R., Colangelo, A.M., and Paolisso, G. (2023). Iron and Ferroptosis More than a Suspect: Beyond the Most Common Mechanisms of Neurodegeneration for New Therapeutic Approaches to Cognitive Decline and Dementia. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms24119637"},{"key":"ref_176","doi-asserted-by":"crossref","first-page":"1761","DOI":"10.1212\/01.WNL.0000068021.13945.7F","article-title":"Parkinson\u2019s Disease Risks Associated with Dietary Iron, Manganese, and Other Nutrient Intakes","volume":"60","author":"Powers","year":"2003","journal-title":"Neurology"},{"key":"ref_177","doi-asserted-by":"crossref","first-page":"1381","DOI":"10.1093\/aje\/kwn273","article-title":"Dietary Iron Intake and Risk of Parkinson\u2019s Disease","volume":"168","author":"Logroscino","year":"2008","journal-title":"Am. J. Epidemiol."},{"key":"ref_178","doi-asserted-by":"crossref","first-page":"34","DOI":"10.1016\/j.neuint.2017.01.004","article-title":"Ferroptosis and Cell Death Mechanisms in Parkinson\u2019s Disease","volume":"104","author":"Guiney","year":"2017","journal-title":"Neurochem. Int."},{"key":"ref_179","doi-asserted-by":"crossref","unstructured":"Khan, M.S., Nasiripour, S., and Bopassa, J.C. (2025). Parkinson Disease Signaling Pathways, Molecular Mechanisms, and Potential Therapeutic Strategies: A Comprehensive Review. Int. J. Mol. Sci., 26.","DOI":"10.3390\/ijms26136416"},{"key":"ref_180","doi-asserted-by":"crossref","first-page":"103787","DOI":"10.1016\/j.redox.2025.103787","article-title":"Decoding Parkinson\u2019s Disease: The Interplay of Cell Death Pathways, Oxidative Stress, and Therapeutic Innovations","volume":"85","author":"Liu","year":"2025","journal-title":"Redox Biol."},{"key":"ref_181","doi-asserted-by":"crossref","first-page":"278","DOI":"10.1038\/s41589-019-0462-8","article-title":"Redox Lipid Reprogramming Commands Susceptibility of Macrophages and Microglia to Ferroptotic Death","volume":"16","author":"Kapralov","year":"2020","journal-title":"Nat. Chem. Biol."},{"key":"ref_182","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1186\/s40478-021-01126-5","article-title":"Iron Loading Is a Prominent Feature of Activated Microglia in Alzheimer\u2019s Disease Patients","volume":"9","author":"Kenkhuis","year":"2021","journal-title":"Acta Neuropathol. Commun."},{"key":"ref_183","doi-asserted-by":"crossref","first-page":"541","DOI":"10.1111\/jnc.12244","article-title":"Inflammation Alters the Expression of DMT, FPN1 and Hepcidin, and It Causes Iron Accumulation in Central Nervous System Cells","volume":"126","author":"Urrutia","year":"2013","journal-title":"J. Neurochem."},{"key":"ref_184","doi-asserted-by":"crossref","first-page":"258","DOI":"10.1016\/j.molmed.2022.02.003","article-title":"Ferroptosis in Parkinson\u2019s Disease: Glia\u2013Neuron Crosstalk","volume":"28","author":"Wang","year":"2022","journal-title":"Trends Mol. Med."},{"key":"ref_185","doi-asserted-by":"crossref","first-page":"107179","DOI":"10.1016\/j.intimp.2020.107179","article-title":"Ferrostatin-1 Alleviates Angiotensin II (Ang II)- Induced Inflammation and Ferroptosis in Astrocytes","volume":"90","author":"Li","year":"2021","journal-title":"Int. Immunopharmacol."},{"key":"ref_186","doi-asserted-by":"crossref","first-page":"215","DOI":"10.14336\/AD.2016.0811","article-title":"Serum Hepcidin Levels, Iron Dyshomeostasis and Cognitive Loss in Alzheimer\u2019s Disease","volume":"8","author":"Sternberg","year":"2017","journal-title":"Aging Dis."},{"key":"ref_187","doi-asserted-by":"crossref","first-page":"85","DOI":"10.1016\/j.jns.2019.06.008","article-title":"Clinical Utility of Serum Hepcidin and Iron Profile Measurements in Alzheimer\u2019s Disease","volume":"403","author":"Kweon","year":"2019","journal-title":"J. Neurol. Sci."},{"key":"ref_188","doi-asserted-by":"crossref","first-page":"42","DOI":"10.1186\/s13195-025-01696-9","article-title":"Plasma Hepcidin as a Potential Informative Biomarker of Alzheimer Disease and Vascular Dementia","volume":"17","author":"Delaby","year":"2025","journal-title":"Alzheimers. Res. Ther."},{"key":"ref_189","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1186\/s12967-018-1399-5","article-title":"Hepcidin, an Emerging and Important Player in Brain Iron Homeostasis","volume":"16","author":"Vela","year":"2018","journal-title":"J. Transl. Med."},{"key":"ref_190","doi-asserted-by":"crossref","first-page":"e2676","DOI":"10.1038\/cddis.2017.93","article-title":"Astrocyte Hepcidin Is a Key Factor in LPS-Induced Neuronal Apoptosis","volume":"8","author":"You","year":"2017","journal-title":"Cell Death Dis."},{"key":"ref_191","doi-asserted-by":"crossref","first-page":"100042","DOI":"10.1016\/j.jnrt.2023.100042","article-title":"The Effect of CX3CL1\/ CX3CR1 Signal Axis on Microglia in Central Nervous System Diseases","volume":"11","author":"Zhao","year":"2023","journal-title":"J. Neurorestoratology"},{"key":"ref_192","doi-asserted-by":"crossref","first-page":"104930","DOI":"10.1016\/j.phrs.2020.104930","article-title":"Fractalkine (CX3CL1) Signaling and Neuroinflammation in Parkinson\u2019s Disease: Potential Clinical and Therapeutic Implications","volume":"158","author":"Angelopoulou","year":"2020","journal-title":"Pharmacol. Res."}],"container-title":["Nutrients"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2072-6643\/18\/1\/109\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,12,31]],"date-time":"2025-12-31T13:37:35Z","timestamp":1767188255000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2072-6643\/18\/1\/109"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,12,28]]},"references-count":192,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,1]]}},"alternative-id":["nu18010109"],"URL":"https:\/\/doi.org\/10.3390\/nu18010109","relation":{},"ISSN":["2072-6643"],"issn-type":[{"value":"2072-6643","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,12,28]]}}}