{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,5]],"date-time":"2026-11-05T18:37:51Z","timestamp":1793903871100,"version":"4.8.0"},"reference-count":151,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2023,8,11]],"date-time":"2023-08-11T00:00:00Z","timestamp":1691712000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001659","name":"Deutsche Forschungsgemeinschaft","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100001659","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Immunol."],"abstract":"<jats:p>\n                    The human 25-kDa Lipocalin 2 (LCN2) was first identified and purified as a protein that in part is associated with gelatinase from neutrophils. This protein shows a high degree of sequence similarity with the deduced sequences of rat \u03b1\n                    <jats:sub>2<\/jats:sub>\n                    -microglobulin-related protein and the mouse protein 24p3. Based on its typical lipocalin fold, which consists of an eight-stranded, anti-parallel, symmetrical \u03b2-barrel fold structure it was initially thought that LCN2 is a circulating protein functioning as a transporter of small lipophilic molecules. However, studies in\n                    <jats:italic>Lcn2<\/jats:italic>\n                    null mice have shown that LCN2 has bacteriostatic properties and plays a key role in innate immunity by sequestering bacterial iron siderophores. Numerous reports have further shown that LCN2 is involved in the control of cell differentiation, energy expenditure, cell death, chemotaxis, cell migration, and many other biological processes. In addition, important roles for LCN2 in health and disease have been identified in\n                    <jats:italic>Lcn2<\/jats:italic>\n                    null mice and multiple molecular pathways required for regulation of\n                    <jats:italic>Lcn2<\/jats:italic>\n                    expression have been identified. Nevertheless, although six putative receptors for LCN2 have been proposed, there is a fundamental lack in understanding of how these cell-surface receptors transmit and amplify LCN2 to the cell. In the present review we summarize the current knowledge on LCN2 receptors and discuss inconsistencies, misinterpretations and false assumptions in the understanding of these potential LCN2 receptors.\n                  <\/jats:p>","DOI":"10.3389\/fimmu.2023.1229885","type":"journal-article","created":{"date-parts":[[2023,8,11]],"date-time":"2023-08-11T13:10:02Z","timestamp":1691759402000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":67,"title":["Lipocalin 2 receptors: facts, fictions, and myths"],"prefix":"10.3389","volume":"14","author":[{"given":"Sarah K.","family":"Schr\u00f6der","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Natalie","family":"Gasterich","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sabine","family":"Weiskirchen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ralf","family":"Weiskirchen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2023,8,11]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"11368","DOI":"10.1093\/nar\/16.23.1136","article-title":"The primary structure of rat alpha 2\u00b5 globulin-related protein","volume":"16","author":"Chan","year":"1988","journal-title":"Nucleic Acids Res"},{"key":"B2","article-title":"SV40-induced expression of mouse gene 24p3 involves a post-transcriptional mechanism","volume":"4","author":"Hraba-Renevey","year":"1989","journal-title":"Oncogene"},{"key":"B3","doi-asserted-by":"publisher","DOI":"10.1016\/S0021-9258(18)82217-7","article-title":"Isolation and primary structure of NGAL, a novel protein associated with human neutrophil gelatinase","volume":"268","author":"Kjeldsen","year":"1993","journal-title":"J Biol Chem"},{"key":"B4","unstructured":"Protein Data Bank (PDB)"},{"key":"B5","doi-asserted-by":"publisher","DOI":"10.1016\/0006-291x(82)91845-9","article-title":"Rapid selective stimulation by growth factors of the incorporation by BALB\/C 3T3 cells of [35S]methionine into a glycoprotein and five superinducible proteins","volume":"108","author":"Nilsen-Hamilton","year":"1982","journal-title":"Biochem Biophys Res Commun"},{"key":"B6","doi-asserted-by":"publisher","DOI":"10.1016\/0167-4889(91)90077-b","article-title":"Basic fibroblast growth factor induces 3T3 fibroblasts to synthesize and secrete a cyclophilin-like protein and beta 2-microglobulin","volume":"1095","author":"Davis","year":"1991","journal-title":"Biochim Biophys Acta"},{"key":"B7","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.270.38.22565","article-title":"Identification of a new acute phase protein","volume":"270","author":"Liu","year":"1995","journal-title":"J Biol Chem"},{"key":"B8","doi-asserted-by":"publisher","DOI":"10.1111\/j.1478-3231.2011.02495.x","article-title":"Induction of lipocalin-2 expression in acute and chronic experimental liver injury moderated by pro-inflammatory cytokines interleukin-1\u03b2 through nuclear factor-\u03baB activation","volume":"31","author":"Borkham-Kamphorst","year":"2011","journal-title":"Liver Int"},{"key":"B9","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbadis.2013.01.014","article-title":"Protective effects of lipocalin-2 (LCN2) in acute liver injury suggest a novel function in liver homeostasis","volume":"1832","author":"Borkham-Kamphorst","year":"2013","journal-title":"Biochim Biophys Acta"},{"key":"B10","doi-asserted-by":"publisher","DOI":"10.1016\/s1097-2765(02)00708-6","article-title":"The neutrophil lipocalin NGAL is a bacteriostatic agent that interferes with siderophore-mediated iron acquisition","volume":"10","author":"Goetz","year":"2002","journal-title":"Mol Cell"},{"key":"B11","doi-asserted-by":"publisher","DOI":"10.1016\/s1097-2765(02)00710-4","article-title":"An iron delivery pathway mediated by a lipocalin","volume":"10","author":"Yang","year":"2002","journal-title":"Mol Cell"},{"key":"B12","doi-asserted-by":"publisher","DOI":"10.1038\/s41579-019-0284-4","article-title":"Bacterial siderophores in community and host interactions","volume":"18","author":"Kramer","year":"2020","journal-title":"Nat Rev Microbiol"},{"key":"B13","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2005.10.027","article-title":"A cell-surface receptor for lipocalin 24p3 selectively mediates apoptosis and iron uptake","volume":"123","author":"Devireddy","year":"2005","journal-title":"Cell"},{"key":"B14","doi-asserted-by":"publisher","DOI":"10.1038\/nature03104","article-title":"Lipocalin 2 mediates an innate immune response to bacterial infection by sequestrating iron","volume":"432","author":"Flo","year":"2004","journal-title":"Nature"},{"key":"B15","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.0510847103","article-title":"Lipocalin 2-deficient mice exhibit increased sensitivity to Escherichia coli infection but not to ischemia-reperfusion injury","volume":"103","author":"Berger","year":"2006","journal-title":"Proc Natl Acad Sci USA"},{"key":"B16","doi-asserted-by":"publisher","DOI":"10.3390\/ijms222313156","article-title":"Depletion of Lipocalin 2 (LCN2) in mice leads to dysbiosis and persistent colonization with segmented filamentous bacteria","volume":"22","author":"Kl\u00fcber","year":"2021","journal-title":"Int J Mol Sci"},{"key":"B17","doi-asserted-by":"publisher","DOI":"10.1002\/hep.27930","article-title":"Lipocalin-2 (NGAL\/LCN2), a \"help-me\" signal in organ inflammation","volume":"63","author":"Asimakopoulou","year":"2016","journal-title":"Hepatology"},{"key":"B18","doi-asserted-by":"publisher","DOI":"10.1002\/(SICI)1098-2795(199704)46:4<507","article-title":"Uterocalin: a mouse acute phase protein expressed in the uterus around birth","volume":"46","author":"Liu","year":"1997","journal-title":"Mol Reprod Dev"},{"key":"B19","doi-asserted-by":"publisher","DOI":"10.1677\/joe.0.1620011","article-title":"Ovarian steroids regulate 24p3 expression in mouse uterus during the natural estrous cycle and the preimplantation period","volume":"162","author":"Huang","year":"1999","journal-title":"J Endocrinol"},{"key":"B20","doi-asserted-by":"publisher","DOI":"10.3390\/ijms24119280","article-title":"Expression analysis of lipocalin 2 (LCN2) in reproductive and non-reproductive tissues of Esr1-deficient mice","volume":"24","author":"Kessel","year":"2023","journal-title":"Int J Mol Sci"},{"key":"B21","doi-asserted-by":"publisher","DOI":"10.1242\/dev.105148","article-title":"Lipocalin 2 binds to membrane phosphatidylethanolamine to induce lipid raft movement in a PKA-dependent manner and modulates sperm maturation","volume":"141","author":"Watanabe","year":"2014","journal-title":"Development"},{"key":"B22","doi-asserted-by":"publisher","DOI":"10.1080\/19396368.2017.1355416","article-title":"Expression patterns and changes of the LCN2 gene in the testes of induced cryptorchidism and busulfan-treated mice","volume":"63","author":"Kang","year":"2017","journal-title":"Syst Biol Reprod Med"},{"key":"B23","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20191261","article-title":"Lipocalin-2 counteracts metabolic dysregulation in obesity and diabetes","volume":"217","author":"Mosialou","year":"2020","journal-title":"J Exp Med"},{"key":"B24","doi-asserted-by":"publisher","DOI":"10.1002\/oby.20507","article-title":"Lipocalin-2 increases fat oxidation in vitro and is correlated with energy expenditure in normal weight but not obese women","volume":"21","author":"Paton","year":"2013","journal-title":"Obes (Silver Spring)"},{"key":"B25","doi-asserted-by":"publisher","first-page":"e58949","DOI":"10.7554\/eLife.58949","article-title":"Lipocalin-2 is an anorexigenic signal in primates","volume":"9","author":"Petropoulou","year":"2020","journal-title":"Elife"},{"key":"B26","doi-asserted-by":"publisher","first-page":"2057","DOI":"10.1038\/s41467-021-22361-3","article-title":"Lipocalin 2 mediates appetite suppression during pancreatic cancer cachexia","volume":"12","author":"Olson","year":"2021","journal-title":"Nat Commun"},{"key":"B27","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbadis.2017.04.006","article-title":"Altered mitochondrial and peroxisomal integrity in lipocalin-2-deficient mice with hepatic steatosis","volume":"1863","author":"Asimakopoulou","year":"2017","journal-title":"Biochim Biophys Acta Mol Basis Dis"},{"key":"B28","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbalip.2014.07.017","article-title":"Lipocalin-2 (LCN2) regulates PLIN5 expression and intracellular lipid droplet formation in the liver","volume":"1842","author":"Asimakopoulou","year":"2014","journal-title":"Biochim Biophys Acta"},{"key":"B29","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0043696","article-title":"Siderocalin\/Lcn2\/NGAL\/24p3 does not drive apoptosis through gentisic acid mediated iron withdrawal in hematopoietic cell lines","volume":"7","author":"Correnti","year":"2012","journal-title":"PloS One"},{"key":"B30","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M115.685644","article-title":"Biochemical and structural characterization of the interaction between the siderocalin NGAL\/LCN2 (neutrophil gelatinase-associated lipocalin\/lipocalin 2) and the N-terminal domain of its endocytic receptor SLC22A17","volume":"291","author":"Cabedo Martinez","year":"2016","journal-title":"J Biol Chem"},{"key":"B31","doi-asserted-by":"publisher","DOI":"10.1002\/dvdy.24394","article-title":"LRP2, an auxiliary receptor that controls sonic hedgehog signaling in development and disease","volume":"245","author":"Christ","year":"2016","journal-title":"Dev Dyn"},{"key":"B32","doi-asserted-by":"publisher","DOI":"10.1038\/nrm778","article-title":"Megalin and cubilin: multifunctional endocytic receptors","volume":"3","author":"Christensen","year":"2002","journal-title":"Nat Rev Mol Cell Biol"},{"key":"B33","doi-asserted-by":"publisher","DOI":"10.1126\/science.abf7958","article-title":"Structure reveals the activation mechanism of the MC4 receptor to initiate satiation signaling","volume":"372","author":"Israeli","year":"2021","journal-title":"Science"},{"key":"B34","doi-asserted-by":"publisher","DOI":"10.1038\/s41422-021-00552-3","article-title":"Structural insights into ligand recognition and activation of the melanocortin-4 receptor","volume":"31","author":"Zhang","year":"2021","journal-title":"Cell Res"},{"key":"B35","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.91.21.9725","article-title":"Complete cloning and sequencing of rat gp330\/\"megalin,\" a distinctive member of the low density lipoprotein receptor gene family","volume":"91","author":"Saito","year":"1994","journal-title":"Proc Natl Acad Sci USA"},{"key":"B36","doi-asserted-by":"publisher","DOI":"10.1016\/S0021-9258(18)82452-8","article-title":"Molecular cloning, expression, and gene localization of a fourth melanocortin receptor","volume":"268","author":"Gantz","year":"1993","journal-title":"J Biol Chem"},{"key":"B37","doi-asserted-by":"publisher","DOI":"10.1016\/0014-5793(92)80820-7","article-title":"Molecular cloning and expression of the human melanocyte stimulating hormone receptor cDNA","volume":"309","author":"Chhajlani","year":"1992","journal-title":"FEBS Lett"},{"key":"B38","doi-asserted-by":"publisher","DOI":"10.1126\/science.1325670","article-title":"The cloning of a family of genes that encode the melanocortin receptors","volume":"257","author":"Mountjoy","year":"1992","journal-title":"Science"},{"key":"B39","doi-asserted-by":"publisher","DOI":"10.1016\/S0021-9258(18)53088-X","article-title":"Molecular cloning of a novel melanocortin receptor","volume":"268","author":"Gantz","year":"1993","journal-title":"J Biol Chem"},{"key":"B40","doi-asserted-by":"publisher","DOI":"10.1042\/bj2990367","article-title":"Molecular cloning, functional expression and pharmacological characterization of a mouse melanocortin receptor gene","volume":"299","author":"Desarnaud","year":"1994","journal-title":"Biochem J"},{"key":"B41","doi-asserted-by":"publisher","DOI":"10.1126\/science.1061075","article-title":"Induction of apoptosis by a secreted lipocalin that is transcriptionally regulated by IL-3 deprivation","volume":"293","author":"Devireddy","year":"2001","journal-title":"Science"},{"key":"B42","doi-asserted-by":"publisher","DOI":"10.1016\/j.febslet.2004.12.031","article-title":"The endocytic receptor megalin binds the iron transporting neutrophil-gelatinase-associated lipocalin with high affinity and mediates its cellular uptake","volume":"579","author":"Hvidberg","year":"2005","journal-title":"FEBS Lett"},{"key":"B43","doi-asserted-by":"publisher","DOI":"10.1038\/nature21697","article-title":"MC4R-dependent suppression of appetite by bone-derived lipocalin 2","volume":"543","author":"Mosialou","year":"2017","journal-title":"Nature"},{"key":"B44","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.93.16.8460","article-title":"Defective forebrain development in mice lacking gp330\/megalin","volume":"93","author":"Willnow","year":"1996","journal-title":"Proc Natl Acad Sci USA"},{"key":"B45","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M109959200","article-title":"Megalin deficiency offers protection from renal aminoglycoside accumulation","volume":"277","author":"Schmitz","year":"2002","journal-title":"J Biol Chem"},{"key":"B46","doi-asserted-by":"publisher","DOI":"10.1016\/s0092-8674(00)81865-6","article-title":"Targeted disruption of the melanocortin-4 receptor results in obesity in mice","volume":"88","author":"Huszar","year":"1997","journal-title":"Cell"},{"key":"B47","doi-asserted-by":"publisher","DOI":"10.1093\/hmg\/10.21.2397","article-title":"Functional variation of MC1R alleles from red-haired individuals","volume":"10","author":"Healy","year":"2001","journal-title":"Hum Mol Genet"},{"key":"B48","doi-asserted-by":"publisher","first-page":"97","DOI":"10.1038\/79254","article-title":"Inactivation of the mouse melanocortin-3 receptor results in increased fat mass and reduced lean body mass","volume":"26","author":"Chen","year":"2000","journal-title":"Nat Genet"},{"key":"B49","doi-asserted-by":"publisher","DOI":"10.2174\/1574888X18666230320091546","article-title":"LCN2 inhibits the BMP9-induced osteogenic differentiation through reducing Wnt\/\u03b2-catenin signaling via interacting with LRP6 in mouse embryonic fibroblasts","volume":"18","author":"Jiang","year":"2023","journal-title":"Curr Stem Cell Res Ther"},{"key":"B50","doi-asserted-by":"publisher","first-page":"297","DOI":"10.1042\/BJ20060836","article-title":"A novel alternative spliced variant of neutrophil gelatinase-associated lipocalin receptor in oesophageal carcinoma cells","volume":"403","author":"Fang","year":"2007","journal-title":"Biochem J"},{"key":"B51","doi-asserted-by":"publisher","DOI":"10.1016\/j.tips.2016.11.008","article-title":"Classification systems of secondary active transporters","volume":"38","author":"Perland","year":"2017","journal-title":"Trends Pharmacol Sci"},{"key":"B52","doi-asserted-by":"publisher","DOI":"10.1111\/febs.15531","article-title":"A guide to plasma membrane solute carrier proteins","volume":"288","author":"Pizzagalli","year":"2021","journal-title":"FEBS J"},{"key":"B53","doi-asserted-by":"publisher","DOI":"10.1124\/dmd.121.000702","article-title":"Emerging roles of the human solute carrier 22 family","volume":"50","author":"Yee","year":"2021","journal-title":"Drug Metab Dispos"},{"key":"B54","unstructured":"Gene: Slc22a12, MGI:1926225"},{"key":"B55","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gkac972","article-title":"The International Mouse Phenotyping Consortium: comprehensive knockout phenotyping underpinning the study of human disease","volume":"51","author":"Groza","year":"2023","journal-title":"Nucleic Acids Res"},{"key":"B56","doi-asserted-by":"publisher","first-page":"888","DOI":"10.1002\/hep.32569","article-title":"Lipocalin-2 activates hepatic stellate cells and promotes nonalcoholic steatohepatitis in high-fat diet-fed Ob\/Ob mice","volume":"77","author":"Kim","year":"2023","journal-title":"Hepatology"},{"key":"B57","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.57264","article-title":"How to measure and evaluate binding affinities","volume":"9","author":"Jarmoskaite","year":"2020","journal-title":"Elife"},{"key":"B58","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0016536","article-title":"Comparing the epidermal growth factor interaction with four different cell lines: intriguing effects imply strong dependency of cellular context","volume":"6","author":"Bj\u00f6rkelund","year":"2011","journal-title":"PloS One"},{"key":"B59","doi-asserted-by":"publisher","DOI":"10.1194\/jlr.M039545","article-title":"New horizons for lipoprotein receptors: communication by \u03b2-propellers","volume":"54","author":"Andersen","year":"2013","journal-title":"J Lipid Res"},{"key":"B60","doi-asserted-by":"publisher","DOI":"10.1016\/j.jacbts.2021.09.011","article-title":"Interplay of low-density lipoprotein receptors, LRPs, and lipoproteins in pulmonary hypertension","volume":"7","author":"Calvier","year":"2022","journal-title":"JACC Basic Transl Sci"},{"key":"B61","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms12973","article-title":"Disposal of iron by a mutant form of lipocalin 2","volume":"7","author":"Barasch","year":"2016","journal-title":"Nat Commun"},{"key":"B62","doi-asserted-by":"publisher","DOI":"10.1681\/ASN.V104685","article-title":"Evidence for an essential role of megalin in transepithelial transport of retinol","volume":"10","author":"Christensen","year":"1999","journal-title":"J Am Soc Nephrol"},{"key":"B63","doi-asserted-by":"publisher","DOI":"10.1016\/j.biochi.2021.09.008","article-title":"The diversity of lipocalin receptors","volume":"192","author":"Redl","year":"2022","journal-title":"Biochimie"},{"key":"B64","doi-asserted-by":"publisher","DOI":"10.1016\/S0002-9440(10)65238-8","article-title":"Megalin knockout mice as an animal model of low molecular weight proteinuria","volume":"155","author":"Leheste","year":"1999","journal-title":"Am J Pathol"},{"key":"B65","doi-asserted-by":"publisher","DOI":"10.1016\/S0021-9258(19)85224-9","article-title":"Glycoprotein 330, a member of the low density lipoprotein receptor family, binds lipoprotein lipase in vitro","volume":"268","author":"Kounnas","year":"1993","journal-title":"J Biol Chem"},{"key":"B66","doi-asserted-by":"publisher","DOI":"10.3389\/fnmol.2022.930526","article-title":"The role of lipocalin 2 in brain injury and recovery after ischemic and hemorrhagic stroke","volume":"15","author":"Zhang","year":"2022","journal-title":"Front Mol Neurosci"},{"key":"B67","doi-asserted-by":"publisher","DOI":"10.1359\/jbmr.081235","article-title":"Where Wnts went: the exploding field of Lrp5 and Lrp6 signaling in bone","volume":"24","author":"Williams","year":"2009","journal-title":"J Bone Miner Res"},{"key":"B68","doi-asserted-by":"publisher","DOI":"10.3389\/fmed.2019.00145","article-title":"The importance of melanocortin receptors and their agonists in pulmonary disease","volume":"6","author":"Moscowitz","year":"2019","journal-title":"Front Med (Lausanne)"},{"key":"B69","doi-asserted-by":"publisher","DOI":"10.3390\/biom12101407","article-title":"Ligands for melanocortin receptors: Beyond melanocyte-stimulating hormones and adrenocorticotropin","volume":"12","author":"Yuan","year":"2022","journal-title":"Biomolecules"},{"key":"B70","doi-asserted-by":"publisher","first-page":"26","DOI":"10.1186\/s12974-021-02078-2","article-title":"Activation of MC1R with BMS-470539 attenuates neuroinflammation via cAMP\/PKA\/Nurr1 pathway after neonatal hypoxic-ischemic brain injury in rats","volume":"18","author":"Yu","year":"2021","journal-title":"J Neuroinflamm"},{"key":"B71","doi-asserted-by":"publisher","first-page":"294","DOI":"10.1007\/s13311-019-00772-x","article-title":"Activation of melanocortin 1 receptor attenuates early brain injury in a rat model of subarachnoid hemorrhage via the suppression of neuroinflammation through AMPK\/TBK1\/NF-\u03baB pathway in rats","volume":"17","author":"Xu","year":"2020","journal-title":"Neurotherapeutics"},{"key":"B72","doi-asserted-by":"publisher","DOI":"10.1016\/j.nbd.2023.106044","article-title":"Role of lipocalin 2 in stroke","volume":"179","author":"Zhao","year":"2023","journal-title":"Neurobiol Dis"},{"key":"B73","doi-asserted-by":"publisher","DOI":"10.1210\/er.2009-0037","article-title":"The melanocortin-4 receptor: physiology, pharmacology, and pathophysiology","volume":"31","author":"Tao","year":"2010","journal-title":"Endocr Rev"},{"key":"B74","doi-asserted-by":"publisher","DOI":"10.3389\/fnmol.2023.1038341","article-title":"Melanocortin receptor 3 and 4 mRNA expression in the adult female Syrian hamster brain","volume":"16","author":"Hall","year":"2023","journal-title":"Front Mol Neurosci"},{"key":"B75","doi-asserted-by":"publisher","DOI":"10.1007\/s11010-011-0748-y","article-title":"Expression and analysis of two novel rat organic cation transporter homologs, SLC22A17 and SLC22A23","volume":"352","author":"Bennett","year":"2011","journal-title":"Mol Cell Biochem"},{"key":"B76","doi-asserted-by":"publisher","DOI":"10.3390\/ijms20215398","article-title":"Inverse regulation of lipocalin-2\/24p3 receptor\/SLC22A17 and lipocalin-2 expression by tonicity, NFAT5\/TonEBP and arginine vasopressin in mouse cortical collecting duct cells mCCD(cl.1): Implications for osmotolerance","volume":"20","author":"Probst","year":"2019","journal-title":"Int J Mol Sci"},{"key":"B77","doi-asserted-by":"publisher","DOI":"10.1002\/jnr.22779","article-title":"Regulation by lipocalin-2 of neuronal cell death, migration, and morphology","volume":"90","author":"Lee","year":"2012","journal-title":"J Neurosci Res"},{"key":"B78","doi-asserted-by":"publisher","DOI":"10.7150\/ijbs.3.100","article-title":"Mouse 24p3 protein has an effect on L929 cell viability","volume":"3","author":"Li","year":"2007","journal-title":"Int J Biol Sci"},{"key":"B79","doi-asserted-by":"publisher","DOI":"10.1007\/s12264-022-00906-w","article-title":"Lipocalin-2-mediated insufficient oligodendrocyte progenitor cell remyelination for white matter injury after subarachnoid hemorrhage via SCL22A17 receptor\/early growth response protein 1 signaling","volume":"38","author":"Li","year":"2022","journal-title":"Neurosci Bull"},{"key":"B80","doi-asserted-by":"publisher","DOI":"10.1016\/j.cellsig.2019.01.001","article-title":"Liver parenchymal cells lacking Lipocalin 2 (LCN2) are prone to endoplasmic reticulum stress and unfolded protein response","volume":"55","author":"Borkham-Kamphorst","year":"2019","journal-title":"Cell Signal"},{"key":"B81","doi-asserted-by":"publisher","DOI":"10.1007\/s10565-009-9119-9","article-title":"Lipocalin-2: pro- or anti-apoptotic","volume":"26","author":"Kehrer","year":"2010","journal-title":"Cell Biol Toxicol"},{"key":"B82","doi-asserted-by":"publisher","DOI":"10.1210\/endo.139.3.5978","article-title":"Megalin (gp330): a putative endocytic receptor for thyroglobulin (Tg)","volume":"139","author":"Zheng","year":"1998","journal-title":"Endocrinology"},{"key":"B83","article-title":"Chapter 18 - lipoprotein receptors","volume-title":"Biochemistry of Lipids, Lipoproteins and Membranes","author":"Kuhbandner","year":"2015"},{"key":"B84","doi-asserted-by":"publisher","DOI":"10.1152\/ajprenal.00259.2021","article-title":"Cubilin-, megalin-, and Dab2-dependent transcription revealed by CRISPR\/Cas9 knockout in kidney proximal tubule cells","volume":"322","author":"Long","year":"2022","journal-title":"Am J Physiol Renal Physiol"},{"key":"B85","doi-asserted-by":"publisher","first-page":"1933","DOI":"10.1016\/j.cell.2019.04.044","article-title":"Illuminating G-protein-coupling selectivity of GPCRs","volume":"177","author":"Inoue","year":"2019","journal-title":"Cell"},{"key":"B86","doi-asserted-by":"publisher","DOI":"10.3390\/ijms21041224","article-title":"Differential signaling profiles of MC4R mutations with three different ligands","volume":"21","author":"Paisdzior","year":"2020","journal-title":"Int J Mol Sci"},{"key":"B87","doi-asserted-by":"publisher","DOI":"10.1172\/JCI76348","article-title":"G(q\/11)\u03b1 and G(s)\u03b1 mediate distinct physiological responses to central melanocortins","volume":"126","author":"Li","year":"2016","journal-title":"J Clin Invest"},{"key":"B88","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.RA118.003450","article-title":"The stimulatory G protein Gs\u03b1 is required in melanocortin 4 receptor-expressing cells for normal energy balance, thermogenesis, and glucose metabolism","volume":"293","author":"Podyma","year":"2018","journal-title":"J Biol Chem"},{"key":"B89","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbadis.2017.04.010","article-title":"Biased signaling at neural melanocortin receptors in regulation of energy homeostasis","volume":"1863","author":"Yang","year":"2017","journal-title":"Biochim Biophys Acta Mol Basis Dis"},{"key":"B90","doi-asserted-by":"publisher","DOI":"10.1038\/srep05403","article-title":"Network based analyses of gene expression profile of LCN2 overexpression in esophageal squamous cell carcinoma","volume":"4","author":"Wu","year":"2014","journal-title":"Sci Rep"},{"key":"B91","unstructured":"The open access resource for human proteins"},{"key":"B92","doi-asserted-by":"publisher","DOI":"10.1038\/nbt1210-1248","article-title":"Towards a knowledge-based human protein atlas","volume":"28","author":"Uhlen","year":"2010","journal-title":"Nat Biotechnol"},{"key":"B93","doi-asserted-by":"publisher","DOI":"10.1177\/002215549704500306","article-title":"Tissue distribution of human gp330\/megalin, a putative Ca(2+)-sensing protein","volume":"45","author":"Lundgren","year":"1997","journal-title":"J Histochem Cytochem"},{"key":"B94","doi-asserted-by":"publisher","first-page":"69","DOI":"10.1016\/j.modgep.2005.04.014","article-title":"Overlapping expression patterns of the multiligand endocytic receptors cubilin and megalin in the CNS, sensory organs and developing epithelia of the rodent embryo","volume":"6","author":"Ass\u00e9mat","year":"2005","journal-title":"Gene Expr Patterns"},{"key":"B95","doi-asserted-by":"publisher","DOI":"10.1038\/ki.1986.250","article-title":"Energy expenditure in patients with chronic renal failure","volume":"30","author":"Monteon","year":"1986","journal-title":"Kidney Int"},{"key":"B96","doi-asserted-by":"publisher","DOI":"10.2337\/db09-1735","article-title":"Lipocalin-2 deficiency impairs thermogenesis and potentiates diet-induced insulin resistance in mice","volume":"59","author":"Guo","year":"2010","journal-title":"Diabetes"},{"key":"B97","doi-asserted-by":"publisher","DOI":"10.2337\/db09-1541","article-title":"Lipocalin-2 deficiency attenuates insulin resistance associated with aging and obesity","volume":"59","author":"Law","year":"2010","journal-title":"Diabetes"},{"key":"B98","doi-asserted-by":"publisher","DOI":"10.1016\/j.mce.2016.03.011","article-title":"Lipocalin 2 produces insulin resistance and can be upregulated by glucocorticoids in human adipose tissue","volume":"427","author":"Kamble","year":"2016","journal-title":"Mol Cell Endocrinol"},{"key":"B99","doi-asserted-by":"publisher","DOI":"10.1681\/ASN.V9101759","article-title":"Megalin is an endocytic receptor for insulin","volume":"9","author":"Orlando","year":"1998","journal-title":"J Am Soc Nephrol"},{"key":"B100","doi-asserted-by":"publisher","DOI":"10.1152\/ajprenal.00210.2018","article-title":"Increased megalin expression in early type 2 diabetes: role of insulin-signaling pathways","volume":"315","author":"Bryniarski","year":"2018","journal-title":"Am J Physiol Renal Physiol"},{"key":"B101","doi-asserted-by":"publisher","DOI":"10.1007\/s00125-020-05289-z","article-title":"Hypothalamic MC4R regulates glucose homeostasis through adrenaline-mediated control of glucose reabsorption via renal GLUT2 in mice","volume":"64","author":"de Souza Cordeiro","year":"2021","journal-title":"Diabetologia"},{"key":"B102","doi-asserted-by":"publisher","DOI":"10.1007\/s12010-022-04089-y","article-title":"Islet MC4R regulates PC1\/3 to improve insulin secretion in T2DM mice via the cAMP and \u03b2-arrestin-1 pathways","volume":"194","author":"Ni","year":"2022","journal-title":"Appl Biochem Biotechnol"},{"key":"B103","doi-asserted-by":"publisher","first-page":"15558","DOI":"10.1038\/s41598-020-72172-7","article-title":"Hepatic lipocalin 2 promotes liver fibrosis and portal hypertension","volume":"10","author":"Chen","year":"2020","journal-title":"Sci Rep"},{"key":"B104","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0218965","article-title":"Effects of lipocalin-2 on brain endothelial adhesion and permeability","volume":"14","author":"Du","year":"2019","journal-title":"PloS One"},{"key":"B105","doi-asserted-by":"publisher","DOI":"10.1369\/0022155414545224","article-title":"Controls for immunohistochemistry: the Histochemical Society's standards of practice for validation of immunohistochemical assays","volume":"62","author":"Hewitt","year":"2014","journal-title":"J Histochem Cytochem"},{"key":"B106","doi-asserted-by":"publisher","DOI":"10.3233\/JAD-2012-120319","article-title":"Enhanced brain amyloid-\u03b2 clearance by rifampicin and caffeine as a possible protective mechanism against Alzheimer's disease","volume":"31","author":"Qosa","year":"2012","journal-title":"J Alzheimers Dis"},{"key":"B107","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M111.308296","article-title":"Lipocalin-2 (24p3\/neutrophil gelatinase-associated lipocalin (NGAL)) receptor is expressed in distal nephron and mediates protein endocytosis","volume":"287","author":"Langelueddecke","year":"2012","journal-title":"J Biol Chem"},{"key":"B108","doi-asserted-by":"publisher","DOI":"10.3892\/etm.2018.6196","article-title":"Expression profiling of lipocalin-2 and 24p3 receptor in murine gonads at different developmental stages","volume":"16","author":"De La Chesnaye","year":"2018","journal-title":"Exp Ther Med"},{"key":"B109","doi-asserted-by":"publisher","DOI":"10.4049\/jimmunol.172.5.2731","article-title":"Of mice and not men: differences between mouse and human immunology","volume":"172","author":"Mestas","year":"2004","journal-title":"J Immunol"},{"key":"B110","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1222878110","article-title":"Inflammation and Host Response to Injury, Large Scale Collaborative Research Program. Genomic responses in mouse models poorly mimic human inflammatory diseases","volume":"110","author":"Seok","year":"2013","journal-title":"Proc Natl Acad Sci USA"},{"key":"B111","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1401965111","article-title":"Genomic responses in mouse models greatly mimic human inflammatory diseases","volume":"112","author":"Takao","year":"2015","journal-title":"Proc Natl Acad Sci USA"},{"key":"B112","doi-asserted-by":"publisher","first-page":"223","DOI":"10.1186\/s12967-019-1976-2","article-title":"Animal to human translation: a systematic scoping review of reported concordance rates","volume":"17","author":"Leenaars","year":"2019","journal-title":"J Transl Med"},{"key":"B113","unstructured":"blastp"},{"key":"B114","doi-asserted-by":"publisher","DOI":"10.1093\/bib\/bbr031","article-title":"When orthologs diverge between human and mouse","volume":"12","author":"Gharib","year":"2011","journal-title":"Brief Bioinform"},{"key":"B115","doi-asserted-by":"publisher","DOI":"10.1016\/0014-5793(92)81511-j","article-title":"A 25 kDa alpha 2-microglobulin-related protein is a component of the 125 kDa form of human gelatinase","volume":"314","author":"Triebel","year":"1992","journal-title":"FEBS Lett"},{"key":"B116","doi-asserted-by":"publisher","first-page":"29","DOI":"10.1016\/j.str.2004.10.009","article-title":"Siderocalin (Lcn 2) also binds carboxymycobactins, potentially defending against mycobacterial infections through iron sequestration","volume":"13","author":"Holmes","year":"2005","journal-title":"Structure"},{"key":"B117","doi-asserted-by":"publisher","first-page":"112260","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":"B118","doi-asserted-by":"publisher","DOI":"10.1046\/j.1432-1327.2001.02066.x","article-title":"The human neutrophil lipocalin supports the allosteric activation of matrix metalloproteinases","volume":"268","author":"Tschesche","year":"2001","journal-title":"Eur J Biochem"},{"key":"B119","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbamcr.2015.08.010","article-title":"Roles of NGAL and MMP-9 in the tumor microenvironment and sensitivity to targeted therapy","volume":"1863","author":"Candido","year":"2016","journal-title":"Biochim Biophys Acta"},{"key":"B120","doi-asserted-by":"publisher","DOI":"10.3390\/ijms21124365","article-title":"Biological functions and therapeutic potential of lipocalin 2 in cancer","volume":"21","author":"Santiago-S\u00e1nchez","year":"2020","journal-title":"Int J Mol Sci"},{"key":"B121","doi-asserted-by":"publisher","first-page":"21","DOI":"10.1186\/s12929-015-0129-z","article-title":"Identification of lipocalin-2 as a PKC\u03b4 phosphorylation substrate in neutrophils","volume":"22","author":"Weng","year":"2015","journal-title":"J BioMed Sci"},{"key":"B122","doi-asserted-by":"publisher","DOI":"10.1021\/bi991162e","article-title":"Glycosylation of natural human neutrophil gelatinase B and neutrophil gelatinase B-associated lipocalin","volume":"38","author":"Rudd","year":"1999","journal-title":"Biochemistry"},{"key":"B123","doi-asserted-by":"publisher","DOI":"10.1016\/j.carres.2010.07.024","article-title":"Structural characterization of glycoprotein NGAL, an early predictive biomarker for acute kidney injury","volume":"345","author":"Zhao","year":"2010","journal-title":"Carbohydr Res"},{"key":"B124","doi-asserted-by":"publisher","DOI":"10.3389\/fphar.2018.00426","article-title":"N-glycosylation of Lipocalin 2 is not required for secretion or exosome targeting","volume":"9","author":"Borkham-Kamphorst","year":"2018","journal-title":"Front Pharmacol"},{"key":"B125","doi-asserted-by":"publisher","first-page":"7","DOI":"10.1186\/s12950-016-0116-5","article-title":"Lipopolysaccharide-induced inflammation or unilateral ureteral obstruction yielded multiple types of glycosylated Lipocalin 2","volume":"13","author":"Fujiwara","year":"2016","journal-title":"J Inflamm (Lond)"},{"key":"B126","doi-asserted-by":"publisher","DOI":"10.1161\/JAHA.114.000837","article-title":"Deamidated lipocalin-2 induces endothelial dysfunction and hypertension in dietary obese mice","volume":"3","author":"Song","year":"2014","journal-title":"J Am Heart Assoc"},{"key":"B127","doi-asserted-by":"publisher","DOI":"10.3390\/cancers12082124","article-title":"Relation of neutrophil gelatinase-associated lipocalin overexpression to the resistance to apoptosis of tumor B cells in chronic lymphocytic leukemia","volume":"12","author":"Bauvois","year":"2020","journal-title":"Cancers (Basel)"},{"key":"B128","doi-asserted-by":"publisher","DOI":"10.1111\/j.1600-0609.2011.01712.x","article-title":"Lipocalin-2 is associated with modulation of disease phenotype in a patient with concurrent JAK2-V617F and BCR-ABL mutation","volume":"88","author":"Nadarajan","year":"2012","journal-title":"Eur J Haematol"},{"key":"B129","doi-asserted-by":"publisher","DOI":"10.3892\/ol_00000019","article-title":"Overexpression of neutrophil gelatinase-associated lipocalin and its receptor in colorectal carcinoma: significant correlation with cell differentiation and tumour invasion","volume":"1","author":"Lv","year":"2010","journal-title":"Oncol Lett"},{"key":"B130","doi-asserted-by":"publisher","DOI":"10.1158\/1078-0432.CCR-08-0420","article-title":"NGALR is overexpressed and regulated by hypomethylation in esophageal squamous cell carcinoma","volume":"14","author":"Cui","year":"2008","journal-title":"Clin Cancer Res"},{"key":"B131","doi-asserted-by":"publisher","first-page":"69","DOI":"10.1136\/jcp.2010.083907","article-title":"Neutrophil gelatinase-associated lipocalin and its receptor: independent prognostic factors of oesophageal squamous cell carcinoma","volume":"64","author":"Du","year":"2011","journal-title":"J Clin Pathol"},{"key":"B132","doi-asserted-by":"publisher","DOI":"10.1007\/s11060-010-0486-0","article-title":"NGAL and NGALR are frequently overexpressed in human gliomas and are associated with clinical prognosis","volume":"104","author":"Liu","year":"2011","journal-title":"J Neurooncol"},{"key":"B133","doi-asserted-by":"publisher","DOI":"10.3233\/jad-160330","article-title":"Neutrophil Gelatinase-Associated Lipocalin and its Receptors in Alzheimer's Disease (AD) Brain Regions: Differential Findings in AD with and without Depression","volume":"55","author":"Dekens","year":"2017","journal-title":"J Alzheimers Dis"},{"key":"B134","doi-asserted-by":"publisher","DOI":"10.1126\/science.aaz2193","article-title":"Cancer cells deploy lipocalin-2 to collect limiting iron in leptomeningeal metastasis","volume":"369","author":"Chi","year":"2020","journal-title":"Science"},{"key":"B135","doi-asserted-by":"publisher","DOI":"10.1016\/j.gene.2018.08.060","article-title":"The expression analysis of NGAL and NGALR in clear cell renal cell carcinoma","volume":"676","author":"Liu","year":"2018","journal-title":"Gene"},{"key":"B136","doi-asserted-by":"publisher","DOI":"10.3892\/ijmm.2011.613","article-title":"Increased expression of neutrophil gelatinase-associated lipocalin receptor by interleukin-1\u03b2 in human mesangial cells via MAPK\/ERK activation","volume":"27","author":"Mao","year":"2011","journal-title":"Int J Mol Med"},{"key":"B137","doi-asserted-by":"publisher","DOI":"10.1016\/j.canep.2012.05.012","article-title":"NGAL and NGALR overexpression in human hepatocellular carcinoma toward a molecular prognostic classification","volume":"36","author":"Zhang","year":"2012","journal-title":"Cancer Epidemiol"},{"key":"B138","doi-asserted-by":"publisher","DOI":"10.4162\/nrp.2016.10.1.115","article-title":"Genome-wide association study for the interaction between BMR and BMI in obese Korean women including overweight","volume":"10","author":"Lee","year":"2016","journal-title":"Nut Res Pract"},{"key":"B139","doi-asserted-by":"publisher","first-page":"499","DOI":"10.1089\/jmf.2018.4310","article-title":"Metabolomics associated with genome-wide association study related to the basal metabolic rate in overweight\/obese Korean women","volume":"22","author":"Kim","year":"2019","journal-title":"J Med Food"},{"key":"B140","doi-asserted-by":"publisher","DOI":"10.1038\/s41430-020-00823-z","article-title":"A genome-wide association study in Japanese identified one variant associated with a preference for a Japanese dietary pattern","volume":"75","author":"Suzuki","year":"2021","journal-title":"Eur J Clin Nutr"},{"key":"B141","doi-asserted-by":"publisher","DOI":"10.2217\/pgs.15.61","article-title":"Genetic variants in SLC22A17 and SLC22A7 are associated with anthracycline-induced cardiotoxicity in children","volume":"16","author":"Visscher","year":"2015","journal-title":"Pharmacogenomics"},{"key":"B142","doi-asserted-by":"publisher","DOI":"10.1016\/j.lfs.2021.119505","article-title":"Correlations between tumor mutation burden and immune infiltrates and their prognostic value in pancreatic cancer by bioinformatic analysis","volume":"277","author":"Cai","year":"2021","journal-title":"Life Sci"},{"key":"B143","doi-asserted-by":"publisher","DOI":"10.1016\/j.jid.2016.03.002","article-title":"Increased lipocalin-2 contributes to the pathogenesis of psoriasis by modulating neutrophil chemotaxis and cytokine secretion","volume":"136","author":"Shao","year":"2016","journal-title":"J Invest Dermatol"},{"key":"B144","doi-asserted-by":"publisher","first-page":"2194","DOI":"10.1016\/j.jid.2022.01.012","article-title":"LCN2 mediates skin inflammation in psoriasis through the SREBP2\u2013NLRC4 axis","volume":"142","author":"Ma","year":"2022","journal-title":"J Invest Dermatol"},{"key":"B145","doi-asserted-by":"publisher","DOI":"10.2147\/ott.s287811","article-title":"An iron metabolism-related SLC22A17 for the prognostic value of gastric cancer","volume":"13","author":"Wei","year":"2020","journal-title":"Onco Targets Ther"},{"key":"B146","doi-asserted-by":"publisher","first-page":"206","DOI":"10.1186\/s12885-020-6702-1","article-title":"Identification of immune cells and mRNA associated with prognosis of gastric cancer","volume":"20","author":"Wang","year":"2020","journal-title":"BMC Cancer"},{"key":"B147","doi-asserted-by":"publisher","DOI":"10.3389\/fgene.2021.642608","article-title":"Significance of tumor mutation burden combined with immune infiltrates in the progression and prognosis of advanced gastric cancer","volume":"12","author":"Guo","year":"2021","journal-title":"Front Genet"},{"key":"B148","doi-asserted-by":"publisher","DOI":"10.1016\/j.yexmp.2011.06.00","article-title":"Immunohistochemical detection of a specific receptor for lipocalin2 (solute carrier family 22 member 17, SLC22A17) and its prognostic significance in endometrial carcinoma","volume":"91","author":"Miyamoto","year":"2011","journal-title":"Exp Mol Pathol"},{"key":"B149","article-title":"Correlation of lipocalin-2 and its receptor expressions with preeclampsia","volume":"47","author":"Cao","year":"2012","journal-title":"Zhonghua Fu Chan Ke Za Zhi"},{"key":"B150","doi-asserted-by":"publisher","first-page":"287","DOI":"10.1007\/s12026-022-09352-2","article-title":"Lipocalin 2-not only a biomarker: a study of current literature and systematic findings of ongoing clinical trials","volume":"71","author":"Asaf","year":"2023","journal-title":"Immunol Res"},{"key":"B151","doi-asserted-by":"publisher","DOI":"10.1016\/S0140-6736(05)74811-X","article-title":"Neutrophil gelatinase-associated lipocalin (NGAL) as a biomarker for acute renal injury after cardiac surgery","volume":"365","author":"Mishra","year":"2005","journal-title":"Lancet"}],"container-title":["Frontiers in Immunology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fimmu.2023.1229885\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,8,11]],"date-time":"2023-08-11T13:10:11Z","timestamp":1691759411000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fimmu.2023.1229885\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,8,11]]},"references-count":151,"alternative-id":["10.3389\/fimmu.2023.1229885"],"URL":"https:\/\/doi.org\/10.3389\/fimmu.2023.1229885","relation":{},"ISSN":["1664-3224"],"issn-type":[{"value":"1664-3224","type":"electronic"}],"subject":[],"published":{"date-parts":[[2023,8,11]]},"article-number":"1229885"}}