{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T05:29:27Z","timestamp":1792042167399,"version":"4.4.0"},"reference-count":81,"publisher":"Springer Science and Business Media LLC","issue":"2","license":[{"start":{"date-parts":[[2024,5,10]],"date-time":"2024-05-10T00:00:00Z","timestamp":1715299200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2024,5,10]],"date-time":"2024-05-10T00:00:00Z","timestamp":1715299200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["J Physiol Biochem"],"published-print":{"date-parts":[[2025,5]]},"abstract":"<jats:title>Abstract<\/jats:title>\n          <jats:p>Obesity constitutes a global health epidemic which worsens the main leading death causes such as type 2 diabetes, cardiovascular diseases, and cancer. Changes in the metabolism in patients with obesity frequently lead to insulin resistance, along with hyperglycemia, dyslipidemia and low-grade inflammation, favoring a more aggressive tumor microenvironment. One of the hallmarks of cancer is the reprogramming of the energy metabolism, in which tumor cells change oxidative phosphorylation to aerobic glycolysis or \u201cWarburg effect\u201d. Aerobic glycolysis is faster than oxidative phosphorylation, but less efficient in terms of ATP production. To obtain sufficient ATP, tumor cells increase glucose uptake by the glucose transporters of the GLUT\/SLC2 family. The human glucose transporter GLUT12 was isolated from the breast cancer cell line MCF7. It is expressed in adipose tissue, skeletal muscle and small intestine, where insulin promotes its translocation to the plasma membrane. Moreover, GLUT12 over\u2010expression in mice increases the whole\u2010body insulin sensitivity. Thus, GLUT12 has been proposed as a second insulin\u2010responsive glucose transporter. In obesity, GLUT12 is downregulated and does not respond to insulin. In contrast, GLUT12 is overexpressed in human solid tumors such as breast, prostate, gastric, liver and colon. High glucose concentration, insulin, and hypoxia upregulate GLUT12 both in adipocytes and tumor cells. Inhibition of GLUT12 mediated Warburg effect suppresses proliferation, migration, and invasion of cancer cells and xenografted tumors. This review summarizes the up-to-date information about GLUT12 physiological role and its implication in obesity and cancer, opening new perspectives to consider this transporter as a therapeutic target.<\/jats:p>","DOI":"10.1007\/s13105-024-01028-9","type":"journal-article","created":{"date-parts":[[2024,5,10]],"date-time":"2024-05-10T06:01:43Z","timestamp":1715320903000},"page":"391-401","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":15,"title":["The glucose transporter GLUT12, a new actor in obesity and cancer"],"prefix":"10.1007","volume":"81","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-6674-0798","authenticated-orcid":false,"given":"Miguel","family":"Burgos","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4046-5756","authenticated-orcid":false,"given":"Eva","family":"Gil-Iturbe","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6228-7196","authenticated-orcid":false,"given":"Adri\u00e1n","family":"Idoate-Bay\u00f3n","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7610-1516","authenticated-orcid":false,"given":"Rosa","family":"Castilla-Madrigal","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2018-6434","authenticated-orcid":false,"given":"Maria J.","family":"Moreno-Aliaga","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7319-3451","authenticated-orcid":false,"given":"M. Pilar","family":"Lostao","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2024,5,10]]},"reference":[{"key":"1028_CR1","doi-asserted-by":"publisher","first-page":"30","DOI":"10.3109\/09687688.2010.508196","volume":"28","author":"LB Aerni-Flessner","year":"2011","unstructured":"Aerni-Flessner LB, Otu MC, Moley KH (2011) The amino acids upstream of NH(2)-terminal dileucine motif play a role in regulating the intracellular sorting of the class III transporters GLUT8 and GLUT12. Mol Membr Biol 28:30\u201341. https:\/\/doi.org\/10.3109\/09687688.2010.508196","journal-title":"Mol Membr Biol"},{"key":"1028_CR2","doi-asserted-by":"publisher","first-page":"2926","DOI":"10.1111\/febs.14577","volume":"285","author":"P-B Ancey","year":"2018","unstructured":"Ancey P-B, Contat C, Meylan E (2018) Glucose transporters in cancer \u2013 from tumor cells to the tumor microenvironment. FEBS J 285:2926\u20132943. https:\/\/doi.org\/10.1111\/febs.14577","journal-title":"FEBS J"},{"key":"1028_CR3","doi-asserted-by":"publisher","first-page":"315","DOI":"10.1002\/iub.315","volume":"62","author":"R Augustin","year":"2010","unstructured":"Augustin R (2010) The protein family of glucose transport facilitators: it\u2019s not only about glucose after all. IUBMB Life 62:315\u2013333. https:\/\/doi.org\/10.1002\/iub.315","journal-title":"IUBMB Life"},{"key":"1028_CR4","doi-asserted-by":"publisher","first-page":"C153","DOI":"10.1152\/ajpcell.00140.2022","volume":"324","author":"DL Baines","year":"2023","unstructured":"Baines DL, Vasiljevs S, Kalsi KK (2023) Getting sweeter: new evidence for glucose transporters in specific cell types of the airway? Am J Physiol Cell Physiol 324:C153\u2013C166. https:\/\/doi.org\/10.1152\/ajpcell.00140.2022","journal-title":"Am J Physiol Cell Physiol"},{"key":"1028_CR5","doi-asserted-by":"publisher","first-page":"124","DOI":"10.1016\/j.metabol.2015.10.007","volume":"65","author":"CC Barron","year":"2016","unstructured":"Barron CC, Bilan PJ, Tsakiridis T, Tsiani E (2016) Facilitative glucose transporters: implications for cancer detection, prognosis and treatment. Metabolism 65:124\u2013139. https:\/\/doi.org\/10.1016\/j.metabol.2015.10.007","journal-title":"Metabolism"},{"key":"1028_CR6","doi-asserted-by":"publisher","first-page":"20281","DOI":"10.1074\/jbc.274.29.20281","volume":"274","author":"A Barthel","year":"1999","unstructured":"Barthel A, Okino ST, Liao J et al (1999) Regulation of GLUT1 gene transcription by the serine\/threonine kinase Akt1. J Biol Chem 274:20281\u201320286. https:\/\/doi.org\/10.1074\/jbc.274.29.20281","journal-title":"J Biol Chem"},{"key":"1028_CR7","doi-asserted-by":"publisher","first-page":"C477","DOI":"10.1152\/ajpcell.00075.2007","volume":"293","author":"MU Baumann","year":"2007","unstructured":"Baumann MU, Zamudio S, Illsley NP (2007) Hypoxic upregulation of glucose transporters in BeWo choriocarcinoma cells is mediated by hypoxia-inducible factor-1. Am J Physiol Cell Physiol 293:C477-485. https:\/\/doi.org\/10.1152\/ajpcell.00075.2007","journal-title":"Am J Physiol Cell Physiol"},{"key":"1028_CR8","doi-asserted-by":"publisher","first-page":"107","DOI":"10.1016\/j.cell.2006.05.036","volume":"126","author":"K Bensaad","year":"2006","unstructured":"Bensaad K, Tsuruta A, Selak MA et al (2006) TIGAR, a p53-Inducible regulator of glycolysis and apoptosis. Cell 126:107\u2013120. https:\/\/doi.org\/10.1016\/j.cell.2006.05.036","journal-title":"Cell"},{"key":"1028_CR9","first-page":"1075","volume":"44","author":"GM Bormans","year":"2003","unstructured":"Bormans GM, Van Oosterwyck G, De Groot TJ et al (2003) Synthesis and biologic evaluation of (11)c-methyl-d-glucoside, a tracer of the sodium-dependent glucose transporters. J Nucl Med Off Publ Soc Nucl Med 44:1075\u20131081","journal-title":"J Nucl Med Off Publ Soc Nucl Med"},{"key":"1028_CR10","doi-asserted-by":"publisher","first-page":"267","DOI":"10.1038\/nrm782","volume":"3","author":"NJ Bryant","year":"2002","unstructured":"Bryant NJ, Govers R, James DE (2002) Regulated transport of the glucose transporter GLUT4. Nat Rev Mol Cell Biol 3:267\u2013277. https:\/\/doi.org\/10.1038\/nrm782","journal-title":"Nat Rev Mol Cell Biol"},{"key":"1028_CR11","doi-asserted-by":"publisher","first-page":"774","DOI":"10.1038\/sj.bjc.6605199","volume":"101","author":"JA Canner","year":"2009","unstructured":"Canner JA, Sobo M, Ball S et al (2009) MI-63: a novel small-molecule inhibitor targets MDM2 and induces apoptosis in embryonal and alveolar rhabdomyosarcoma cells with wild-type p53. Br J Cancer 101:774\u2013781. https:\/\/doi.org\/10.1038\/sj.bjc.6605199","journal-title":"Br J Cancer"},{"key":"1028_CR12","doi-asserted-by":"publisher","first-page":"104","DOI":"10.7150\/ijbs.75106","volume":"19","author":"B Cao","year":"2023","unstructured":"Cao B, Zhao R, Li H et al (2023) Inhibition of androgen receptor enhanced the anticancer effects of everolimus through targeting glucose transporter 12. Int J Biol Sci 19:104\u2013119. https:\/\/doi.org\/10.7150\/ijbs.75106","journal-title":"Int J Biol Sci"},{"key":"1028_CR13","doi-asserted-by":"publisher","first-page":"2035","DOI":"10.1002\/cncr.11293","volume":"97","author":"JD Chandler","year":"2003","unstructured":"Chandler JD, Williams ED, Slavin JL et al (2003) Expression and localization of GLUT1 and GLUT12 in prostate carcinoma. Cancer 97:2035\u20132042. https:\/\/doi.org\/10.1002\/cncr.11293","journal-title":"Cancer"},{"key":"1028_CR14","doi-asserted-by":"publisher","first-page":"841","DOI":"10.1007\/s11033-015-3858-x","volume":"42","author":"R Courtnay","year":"2015","unstructured":"Courtnay R, Ngo DC, Malik N et al (2015) Cancer metabolism and the Warburg effect: the role of HIF-1 and PI3K. Mol Biol Rep 42:841\u2013851. https:\/\/doi.org\/10.1007\/s11033-015-3858-x","journal-title":"Mol Biol Rep"},{"key":"1028_CR15","doi-asserted-by":"publisher","first-page":"35","DOI":"10.2174\/156720512799015037","volume":"9","author":"SM de la Monte","year":"2012","unstructured":"de la Monte SM (2012) Brain insulin resistance and deficiency as therapeutic targets in Alzheimer\u2019s disease. Curr Alzheimer Res 9:35\u201366. https:\/\/doi.org\/10.2174\/156720512799015037","journal-title":"Curr Alzheimer Res"},{"key":"1028_CR16","first-page":"481","volume":"50","author":"H Du","year":"2020","unstructured":"Du H, Liu Y, Yuan Y et al (2020) Distinct prognostic values of the mRNA expression of glucose transporters in non-small cell lung cancer. Ann Clin Lab Sci 50:481\u2013489","journal-title":"Ann Clin Lab Sci"},{"key":"1028_CR17","doi-asserted-by":"publisher","first-page":"313","DOI":"10.1530\/JOE-12-0032","volume":"214","author":"BC Fam","year":"2012","unstructured":"Fam BC, Rose LJ, Sgambellone R et al (2012) Normal muscle glucose uptake in mice deficient in muscle GLUT4. J Endocrinol 214:313\u2013327. https:\/\/doi.org\/10.1530\/JOE-12-0032","journal-title":"J Endocrinol"},{"issue":"Suppl 1","key":"1028_CR18","doi-asserted-by":"publisher","first-page":"S4","DOI":"10.1093\/gerona\/glu057","volume":"69","author":"C Franceschi","year":"2014","unstructured":"Franceschi C, Campisi J (2014) Chronic inflammation (inflammaging) and its potential contribution to age-associated diseases. J Gerontol A Biol Sci Med Sci 69(Suppl 1):S4-9. https:\/\/doi.org\/10.1093\/gerona\/glu057","journal-title":"J Gerontol A Biol Sci Med Sci"},{"key":"1028_CR19","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1615\/critrevimmunol.2013006833","volume":"34","author":"SK Garg","year":"2014","unstructured":"Garg SK, Delaney C, Shi H, Yung R (2014) Changes in adipose tissue macrophages and T cells during aging. Crit Rev Immunol 34:1\u201314. https:\/\/doi.org\/10.1615\/critrevimmunol.2013006833","journal-title":"Crit Rev Immunol"},{"key":"1028_CR20","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/nrdp.2017.34","volume":"3","author":"P Gonz\u00e1lez-Muniesa","year":"2017","unstructured":"Gonz\u00e1lez-Muniesa P, M\u00e1rtinez-Gonz\u00e1lez M-A, Hu FB et al (2017) Obesity. Nat Rev Dis Primer 3:1\u201318. https:\/\/doi.org\/10.1038\/nrdp.2017.34","journal-title":"Nat Rev Dis Primer"},{"key":"1028_CR21","doi-asserted-by":"publisher","first-page":"e13283","DOI":"10.1111\/apha.13283","volume":"226","author":"E Gil-Iturbe","year":"2019","unstructured":"Gil-Iturbe E, Arbones-Mainar JM, Moreno-Aliaga MJ, Lostao MP (2019) GLUT12 and adipose tissue: expression, regulation and its relation with obesity in mice. Acta Physiol Oxf Engl 226:e13283. https:\/\/doi.org\/10.1111\/apha.13283","journal-title":"Acta Physiol Oxf Engl"},{"key":"1028_CR22","doi-asserted-by":"publisher","first-page":"4396","DOI":"10.1002\/jcp.27231","volume":"234","author":"E Gil-Iturbe","year":"2019","unstructured":"Gil-Iturbe E, Castilla-Madrigal R, Barrenetxe J et al (2019) GLUT12 expression and regulation in murine small intestine and human Caco-2 cells. J Cell Physiol 234:4396\u20134408. https:\/\/doi.org\/10.1002\/jcp.27231","journal-title":"J Cell Physiol"},{"key":"1028_CR23","doi-asserted-by":"publisher","first-page":"798","DOI":"10.1007\/s12035-019-01743-1","volume":"57","author":"E Gil-Iturbe","year":"2020","unstructured":"Gil-Iturbe E, Solas M, Cuadrado-Tejedo M et al (2020) GLUT12 expression in brain of mouse models of Alzheimer\u2019s disease. Mol Neurobiol 57:798\u2013805. https:\/\/doi.org\/10.1007\/s12035-019-01743-1","journal-title":"Mol Neurobiol"},{"key":"1028_CR24","doi-asserted-by":"publisher","first-page":"957","DOI":"10.1139\/apnm-2019-0721","volume":"45","author":"E Gil-Iturbe","year":"2020","unstructured":"Gil-Iturbe E, F\u00e9lix-Soriano E, S\u00e1inz N et al (2020) Effect of aging and obesity on GLUT12 expression in small intestine, adipose tissue, muscle, and kidney and its regulation by docosahexaenoic acid and exercise in mice. Appl Physiol Nutr Metab Physiol Appl Nutr Metab 45:957\u2013967. https:\/\/doi.org\/10.1139\/apnm-2019-0721","journal-title":"Appl Physiol Nutr Metab Physiol Appl Nutr Metab"},{"key":"1028_CR25","doi-asserted-by":"publisher","first-page":"1426","DOI":"10.3390\/biom11101426","volume":"11","author":"E Gjermeni","year":"2021","unstructured":"Gjermeni E, Kirstein AS, Kolbig F et al (2021) Obesity-an update on the basic pathophysiology and review of recent therapeutic advances. Biomolecules 11:1426. https:\/\/doi.org\/10.3390\/biom11101426","journal-title":"Biomolecules"},{"key":"1028_CR26","doi-asserted-by":"publisher","first-page":"566","DOI":"10.1053\/plac.2002.0925","volume":"24","author":"NM Gude","year":"2003","unstructured":"Gude NM, Stevenson JL, Rogers S et al (2003) GLUT12 expression in human placenta in first trimester and term. Placenta 24:566\u2013570. https:\/\/doi.org\/10.1053\/plac.2002.0925","journal-title":"Placenta"},{"key":"1028_CR27","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1016\/j.placenta.2004.04.006","volume":"26","author":"NM Gude","year":"2005","unstructured":"Gude NM, Stevenson JL, Murthi P et al (2005) Expression of GLUT12 in the fetal membranes of the human placenta. Placenta 26:67\u201372. https:\/\/doi.org\/10.1016\/j.placenta.2004.04.006","journal-title":"Placenta"},{"key":"1028_CR28","doi-asserted-by":"publisher","first-page":"646","DOI":"10.1016\/j.cell.2011.02.013","volume":"144","author":"D Hanahan","year":"2011","unstructured":"Hanahan D, Weinberg RA (2011) Hallmarks of cancer: the next generation. Cell 144:646\u2013674. https:\/\/doi.org\/10.1016\/j.cell.2011.02.013","journal-title":"Cell"},{"key":"1028_CR29","doi-asserted-by":"publisher","first-page":"251","DOI":"10.1038\/nrm3311","volume":"13","author":"DG Hardie","year":"2012","unstructured":"Hardie DG, Ross FA, Hawley SA (2012) AMPK: a nutrient and energy sensor that maintains energy homeostasis. Nat Rev Mol Cell Biol 13:251\u2013262. https:\/\/doi.org\/10.1038\/nrm3311","journal-title":"Nat Rev Mol Cell Biol"},{"key":"1028_CR30","doi-asserted-by":"publisher","first-page":"77","DOI":"10.1016\/0304-419X(74)90008-0","volume":"355","author":"M Hatanaka","year":"1974","unstructured":"Hatanaka M (1974) Transport of sugars in tumor cell membranes. Biochim Biophys Acta BBA - Rev Cancer 355:77\u2013104. https:\/\/doi.org\/10.1016\/0304-419X(74)90008-0","journal-title":"Biochim Biophys Acta BBA - Rev Cancer"},{"key":"1028_CR31","doi-asserted-by":"publisher","first-page":"1155","DOI":"10.1007\/s00424-020-02411-3","volume":"472","author":"GD Holman","year":"2020","unstructured":"Holman GD (2020) Structure, function and regulation of mammalian glucose transporters of the SLC2 family. Pflugers Arch 472:1155\u20131175. https:\/\/doi.org\/10.1007\/s00424-020-02411-3","journal-title":"Pflugers Arch"},{"key":"1028_CR32","doi-asserted-by":"publisher","first-page":"4277","DOI":"10.1200\/JCO.2016.67.9712","volume":"34","author":"BD Hopkins","year":"2016","unstructured":"Hopkins BD, Goncalves MD, Cantley LC (2016) Obesity and cancer mechanisms: cancer metabolism. J Clin Oncol Off J Am Soc Clin Oncol 34:4277\u20134283. https:\/\/doi.org\/10.1200\/JCO.2016.67.9712","journal-title":"J Clin Oncol Off J Am Soc Clin Oncol"},{"key":"1028_CR33","doi-asserted-by":"publisher","first-page":"74","DOI":"10.1016\/j.semcancer.2017.03.001","volume":"43","author":"V Huber","year":"2017","unstructured":"Huber V, Camisaschi C, Berzi A et al (2017) Cancer acidity: An ultimate frontier of tumor immune escape and a novel target of immunomodulation. Semin Cancer Biol 43:74\u201389. https:\/\/doi.org\/10.1016\/j.semcancer.2017.03.001","journal-title":"Semin Cancer Biol"},{"key":"1028_CR34","doi-asserted-by":"publisher","first-page":"C775","DOI":"10.1152\/ajpcell.00279.2015","volume":"309","author":"ME Hubbi","year":"2015","unstructured":"Hubbi ME, Semenza GL (2015) Regulation of cell proliferation by hypoxia-inducible factors. Am J Physiol Cell Physiol 309:C775-782. https:\/\/doi.org\/10.1152\/ajpcell.00279.2015","journal-title":"Am J Physiol Cell Physiol"},{"key":"1028_CR35","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1530\/JOE-14-0539","volume":"224","author":"V Jim\u00e9nez-Amilburu","year":"2015","unstructured":"Jim\u00e9nez-Amilburu V, Jong-Raadsen S, Bakkers J et al (2015) GLUT12 deficiency during early development results in heart failure and a diabetic phenotype in zebrafish. J Endocrinol 224:1\u201315. https:\/\/doi.org\/10.1530\/JOE-14-0539","journal-title":"J Endocrinol"},{"key":"1028_CR36","doi-asserted-by":"publisher","first-page":"3990","DOI":"10.1172\/JCI129187","volume":"129","author":"CR Kahn","year":"2019","unstructured":"Kahn CR, Wang G, Lee KY (2019) Altered adipose tissue and adipocyte function in the pathogenesis of metabolic syndrome. J Clin Invest 129:3990\u20134000. https:\/\/doi.org\/10.1172\/JCI129187","journal-title":"J Clin Invest"},{"key":"1028_CR37","doi-asserted-by":"publisher","first-page":"151","DOI":"10.1038\/377151a0","volume":"377","author":"EB Katz","year":"1995","unstructured":"Katz EB, Stenbit AE, Hatton K et al (1995) Cardiac and adipose tissue abnormalities but not diabetes in mice deficient in GLUT4. Nature 377:151\u2013155. https:\/\/doi.org\/10.1038\/377151a0","journal-title":"Nature"},{"key":"1028_CR38","doi-asserted-by":"publisher","first-page":"4134","DOI":"10.1111\/cas.15940","volume":"114","author":"B Khaledian","year":"2023","unstructured":"Khaledian B, Thibes L, Shimono Y (2023) Adipocyte regulation of cancer stem cells. Cancer Sci 114:4134\u20134144. https:\/\/doi.org\/10.1111\/cas.15940","journal-title":"Cancer Sci"},{"key":"1028_CR39","doi-asserted-by":"publisher","first-page":"F205","DOI":"10.1152\/ajprenal.00237.2004","volume":"290","author":"KC Linden","year":"2006","unstructured":"Linden KC, DeHaan CL, Zhang Y et al (2006) Renal expression and localization of the facilitative glucose transporters GLUT1 and GLUT12 in animal models of hypertension and diabetic nephropathy. Am J Physiol Renal Physiol 290:F205-213. https:\/\/doi.org\/10.1152\/ajprenal.00237.2004","journal-title":"Am J Physiol Renal Physiol"},{"key":"1028_CR40","doi-asserted-by":"publisher","first-page":"91","DOI":"10.1007\/s00441-002-0661-5","volume":"311","author":"ML Macheda","year":"2003","unstructured":"Macheda ML, Williams ED, Best JD et al (2003) Expression and localisation of GLUT1 and GLUT12 glucose transporters in the pregnant and lactating rat mammary gland. Cell Tissue Res 311:91\u201397. https:\/\/doi.org\/10.1007\/s00441-002-0661-5","journal-title":"Cell Tissue Res"},{"key":"1028_CR41","doi-asserted-by":"publisher","first-page":"654","DOI":"10.1002\/jcp.20166","volume":"202","author":"ML Macheda","year":"2005","unstructured":"Macheda ML, Rogers S, Best JD (2005) Molecular and cellular regulation of glucose transporter (GLUT) proteins in cancer. J Cell Physiol 202:654\u2013662. https:\/\/doi.org\/10.1002\/jcp.20166","journal-title":"J Cell Physiol"},{"key":"1028_CR42","doi-asserted-by":"publisher","first-page":"455","DOI":"10.1080\/09687680701298143","volume":"24","author":"AR Manolescu","year":"2007","unstructured":"Manolescu AR, Augustin R, Moley K, Cheeseman C (2007) A highly conserved hydrophobic motif in the exofacial vestibule of fructose transporting SLC2A proteins acts as a critical determinant of their substrate selectivity. Mol Membr Biol 24:455\u2013463. https:\/\/doi.org\/10.1080\/09687680701298143","journal-title":"Mol Membr Biol"},{"key":"1028_CR43","doi-asserted-by":"publisher","first-page":"44","DOI":"10.1016\/j.aanat.2016.11.013","volume":"210","author":"FF Martins","year":"2017","unstructured":"Martins FF, Bargut TCL, Aguila MB, Mandarim-de-Lacerda CA (2017) Thermogenesis, fatty acid synthesis with oxidation, and inflammation in the brown adipose tissue of ob\/ob (-\/-) mice. Ann Anat Anat Anz Off Organ Anat Ges 210:44\u201351. https:\/\/doi.org\/10.1016\/j.aanat.2016.11.013","journal-title":"Ann Anat Anat Anz Off Organ Anat Ges"},{"key":"1028_CR44","doi-asserted-by":"publisher","unstructured":"Matsui C, Takatani-Nakase T, Maeda S, et al (2017) Potential roles of GLUT12 for glucose sensing and cellular migration in MCF-7 human breast cancer cells under high glucose conditions. Anticancer Res 37:6715\u20136722. https:\/\/doi.org\/10.21873\/anticanres.12130","DOI":"10.21873\/anticanres.12130"},{"key":"1028_CR45","doi-asserted-by":"publisher","first-page":"611","DOI":"10.1093\/jb\/mvaa090","volume":"168","author":"S Matsuo","year":"2020","unstructured":"Matsuo S, Hiasa M, Omote H (2020) Functional characterization and tissue localization of the facilitative glucose transporter GLUT12. J Biochem (Tokyo) 168:611\u2013620. https:\/\/doi.org\/10.1093\/jb\/mvaa090","journal-title":"J Biochem (Tokyo)"},{"key":"1028_CR46","doi-asserted-by":"publisher","unstructured":"Miyata H, Toyoda Y, Takada T, et al (2022) Identification of an exporter that regulates vitamin C supply from blood to the brain. iScience 25:103642. https:\/\/doi.org\/10.1016\/j.isci.2021.103642","DOI":"10.1016\/j.isci.2021.103642"},{"key":"1028_CR47","doi-asserted-by":"publisher","first-page":"181","DOI":"10.1007\/s13105-020-00745-1","volume":"76","author":"MJ Moreno-Aliaga","year":"2020","unstructured":"Moreno-Aliaga MJ, Villarroya F (2020) Nutritional and metabolic regulation of brown and beige adipose tissues. J Physiol Biochem 76:181\u2013184. https:\/\/doi.org\/10.1007\/s13105-020-00745-1","journal-title":"J Physiol Biochem"},{"key":"1028_CR48","first-page":"4285","volume":"57","author":"PM O\u2019Connor","year":"1997","unstructured":"O\u2019Connor PM, Jackman J, Bae I et al (1997) Characterization of the p53 tumor suppressor pathway in cell lines of the National Cancer Institute anticancer drug screen and correlations with the growth-inhibitory potency of 123 anticancer agents. Cancer Res 57:4285\u20134300","journal-title":"Cancer Res"},{"key":"1028_CR49","doi-asserted-by":"publisher","first-page":"679","DOI":"10.1111\/jop.13342","volume":"51","author":"R Paolini","year":"2022","unstructured":"Paolini R, Moore C, Matthyssen T et al (2022) Transcriptional regulation of glucose transporters in human oral squamous cell carcinoma cells. J Oral Pathol Med Off Publ Int Assoc Oral Pathol Am Acad Oral Pathol 51:679\u2013683. https:\/\/doi.org\/10.1111\/jop.13342","journal-title":"J Oral Pathol Med Off Publ Int Assoc Oral Pathol Am Acad Oral Pathol"},{"key":"1028_CR50","doi-asserted-by":"publisher","first-page":"455","DOI":"10.1038\/nrendo.2014.94","volume":"10","author":"J Park","year":"2014","unstructured":"Park J, Morley TS, Kim M et al (2014) Obesity and cancer\u2013mechanisms underlying tumour progression and recurrence. Nat Rev Endocrinol 10:455\u2013465. https:\/\/doi.org\/10.1038\/nrendo.2014.94","journal-title":"Nat Rev Endocrinol"},{"key":"1028_CR51","doi-asserted-by":"publisher","first-page":"933","DOI":"10.1007\/s11626-021-00635-w","volume":"57","author":"S Pehlivanoglu","year":"2021","unstructured":"Pehlivanoglu S, Sahan OB, Pehlivanoglu S, Aktas Kont K (2021) Epithelial mesenchymal transition regulator TWIST1 transcription factor stimulates glucose uptake through upregulation of GLUT1, GLUT3, and GLUT12 in vitro. In Vitro Cell Dev Biol Anim 57:933\u2013943. https:\/\/doi.org\/10.1007\/s11626-021-00635-w","journal-title":"In Vitro Cell Dev Biol Anim"},{"key":"1028_CR52","doi-asserted-by":"publisher","first-page":"664","DOI":"10.1016\/j.bbabio.2010.11.004","volume":"1807","author":"PL Prieto-Hontoria","year":"2011","unstructured":"Prieto-Hontoria PL, P\u00e9rez-Matute P, Fern\u00e1ndez-Galilea M et al (2011) Role of obesity-associated dysfunctional adipose tissue in cancer: a molecular nutrition approach. Biochim Biophys Acta 1807:664\u2013678. https:\/\/doi.org\/10.1016\/j.bbabio.2010.11.004","journal-title":"Biochim Biophys Acta"},{"key":"1028_CR53","doi-asserted-by":"publisher","first-page":"325","DOI":"10.1007\/s13105-012-0213-8","volume":"69","author":"J Pujol-Gim\u00e9nez","year":"2013","unstructured":"Pujol-Gim\u00e9nez J, Barrenetxe J, Gonz\u00e1lez-Muniesa P, Lostao MP (2013) The facilitative glucose transporter GLUT12: what do we know and what would we like to know? J Physiol Biochem 69:325\u2013333. https:\/\/doi.org\/10.1007\/s13105-012-0213-8","journal-title":"J Physiol Biochem"},{"key":"1028_CR54","doi-asserted-by":"publisher","first-page":"C1008","DOI":"10.1152\/ajpcell.00343.2014","volume":"308","author":"J Pujol-Gim\u00e9nez","year":"2015","unstructured":"Pujol-Gim\u00e9nez J, P\u00e9rez A, Reyes AM et al (2015) Functional characterization of the human facilitative glucose transporter 12 (GLUT12) by electrophysiological methods. Am J Physiol Cell Physiol 308:C1008-1022. https:\/\/doi.org\/10.1152\/ajpcell.00343.2014","journal-title":"Am J Physiol Cell Physiol"},{"key":"1028_CR55","doi-asserted-by":"publisher","first-page":"97","DOI":"10.3233\/JAD-132498","volume":"42","author":"J Pujol-Gimenez","year":"2014","unstructured":"Pujol-Gimenez J, Martisova E, Perez-Mediavilla A et al (2014) Expression of the glucose transporter GLUT12 in Alzheimer\u2019s disease patients. J Alzheimers Dis JAD 42:97\u2013101. https:\/\/doi.org\/10.3233\/JAD-132498","journal-title":"J Alzheimers Dis JAD"},{"key":"1028_CR56","doi-asserted-by":"publisher","first-page":"139","DOI":"10.7150\/jca.10429","volume":"6","author":"J Pujol-Gimenez","year":"2015","unstructured":"Pujol-Gimenez J, de Heredia FP, Idoate MA et al (2015) Could GLUT12 be a potential therapeutic target in cancer treatment? A preliminary report. J Cancer 6:139\u2013143. https:\/\/doi.org\/10.7150\/jca.10429","journal-title":"J Cancer"},{"key":"1028_CR57","doi-asserted-by":"publisher","first-page":"1478","DOI":"10.2337\/db11-0033","volume":"60","author":"SH Purcell","year":"2011","unstructured":"Purcell SH, Aerni-Flessner LB, Willcockson AR et al (2011) Improved insulin sensitivity by GLUT12 overexpression in mice. Diabetes 60:1478\u20131482. https:\/\/doi.org\/10.2337\/db11-0033","journal-title":"Diabetes"},{"key":"1028_CR58","doi-asserted-by":"publisher","first-page":"633","DOI":"10.1038\/nrendo.2017.90","volume":"13","author":"SM Reilly","year":"2017","unstructured":"Reilly SM, Saltiel AR (2017) Adapting to obesity with adipose tissue inflammation. Nat Rev Endocrinol 13:633\u2013643. https:\/\/doi.org\/10.1038\/nrendo.2017.90","journal-title":"Nat Rev Endocrinol"},{"key":"1028_CR59","doi-asserted-by":"publisher","first-page":"E733","DOI":"10.1152\/ajpendo.2002.282.3.E733","volume":"282","author":"S Rogers","year":"2002","unstructured":"Rogers S, Macheda ML, Docherty SE et al (2002) Identification of a novel glucose transporter-like protein-GLUT-12. Am J Physiol Endocrinol Metab 282:E733-738. https:\/\/doi.org\/10.1152\/ajpendo.2002.282.3.E733","journal-title":"Am J Physiol Endocrinol Metab"},{"key":"1028_CR60","doi-asserted-by":"publisher","first-page":"225","DOI":"10.1016\/s0304-3835(03)00010-7","volume":"193","author":"S Rogers","year":"2003","unstructured":"Rogers S, Docherty SE, Slavin JL et al (2003) Differential expression of GLUT12 in breast cancer and normal breast tissue. Cancer Lett 193:225\u2013233. https:\/\/doi.org\/10.1016\/s0304-3835(03)00010-7","journal-title":"Cancer Lett"},{"key":"1028_CR61","doi-asserted-by":"publisher","first-page":"53","DOI":"10.1016\/j.canlet.2020.09.012","volume":"495","author":"Y Shi","year":"2020","unstructured":"Shi Y, Zhang Y, Ran F et al (2020) Let-7a-5p inhibits triple-negative breast tumor growth and metastasis through GLUT12-mediated warburg effect. Cancer Lett 495:53\u201365. https:\/\/doi.org\/10.1016\/j.canlet.2020.09.012","journal-title":"Cancer Lett"},{"key":"1028_CR62","doi-asserted-by":"publisher","first-page":"G591","DOI":"10.1152\/ajpgi.00269.2014","volume":"308","author":"A Sideri","year":"2015","unstructured":"Sideri A, Stavrakis D, Bowe C et al (2015) Effects of obesity on severity of colitis and cytokine expression in mouse mesenteric fat. Potential role of adiponectin receptor 1. Am J Physiol-Gastrointest Liver Physiol 308:G591\u2013G604. https:\/\/doi.org\/10.1152\/ajpgi.00269.2014","journal-title":"Am J Physiol-Gastrointest Liver Physiol"},{"key":"1028_CR63","doi-asserted-by":"publisher","first-page":"1823","DOI":"10.7314\/apjcp.2014.15.4.1823","volume":"15","author":"K Song","year":"2014","unstructured":"Song K, Li M, Xu X-J et al (2014) HIF-1\u03b1 and GLUT1 gene expression is associated with chemoresistance of acute myeloid leukemia. Asian Pac J Cancer Prev APJCP 15:1823\u20131829. https:\/\/doi.org\/10.7314\/apjcp.2014.15.4.1823","journal-title":"Asian Pac J Cancer Prev APJCP"},{"key":"1028_CR64","doi-asserted-by":"publisher","first-page":"5833","DOI":"10.3390\/jcm10245833","volume":"10","author":"PJ Stanirowski","year":"2021","unstructured":"Stanirowski PJ, Szukiewicz D, Majewska A et al (2021) Differential expression of glucose transporter proteins GLUT-1, GLUT-3, GLUT-8 and GLUT-12 in the placenta of macrosomic, small-for-gestational-age and growth-restricted foetuses. J Clin Med 10:5833. https:\/\/doi.org\/10.3390\/jcm10245833","journal-title":"J Clin Med"},{"key":"1028_CR65","doi-asserted-by":"publisher","first-page":"560","DOI":"10.1111\/jdi.13680","volume":"13","author":"PJ Stanirowski","year":"2022","unstructured":"Stanirowski PJ, Szukiewicz D, Majewska A et al (2022) Placental expression of glucose transporters GLUT-1, GLUT-3, GLUT-8 and GLUT-12 in pregnancies complicated by gestational and type 1 diabetes mellitus. J Diabetes Investig 13:560\u2013570. https:\/\/doi.org\/10.1111\/jdi.13680","journal-title":"J Diabetes Investig"},{"key":"1028_CR66","doi-asserted-by":"publisher","first-page":"629","DOI":"10.1172\/JCI118833","volume":"98","author":"AE Stenbit","year":"1996","unstructured":"Stenbit AE, Burcelin R, Katz EB et al (1996) Diverse effects of Glut 4 ablation on glucose uptake and glycogen synthesis in red and white skeletal muscle. J Clin Invest 98:629\u2013634. https:\/\/doi.org\/10.1172\/JCI118833","journal-title":"J Clin Invest"},{"key":"1028_CR67","doi-asserted-by":"publisher","first-page":"3535","DOI":"10.1210\/jc.2009-0162","volume":"94","author":"CA Stuart","year":"2009","unstructured":"Stuart CA, Howell MEA, Zhang Y, Yin D (2009) Insulin-stimulated translocation of glucose transporter (GLUT) 12 parallels that of GLUT4 in normal muscle. J Clin Endocrinol Metab 94:3535\u20133542. https:\/\/doi.org\/10.1210\/jc.2009-0162","journal-title":"J Clin Endocrinol Metab"},{"key":"1028_CR68","doi-asserted-by":"publisher","first-page":"772","DOI":"10.1038\/nri1937","volume":"6","author":"H Tilg","year":"2006","unstructured":"Tilg H, Moschen AR (2006) Adipocytokines: mediators linking adipose tissue, inflammation and immunity. Nat Rev Immunol 6:772\u2013783. https:\/\/doi.org\/10.1038\/nri1937","journal-title":"Nat Rev Immunol"},{"key":"1028_CR69","doi-asserted-by":"publisher","first-page":"18175","DOI":"10.1073\/pnas.2006958117","volume":"117","author":"Y Toyoda","year":"2020","unstructured":"Toyoda Y, Takada T, Miyata H et al (2020) Identification of GLUT12\/SLC2A12 as a urate transporter that regulates the blood urate level in hyperuricemia model mice. Proc Natl Acad Sci 117:18175\u201318177. https:\/\/doi.org\/10.1073\/pnas.2006958117","journal-title":"Proc Natl Acad Sci"},{"key":"1028_CR70","doi-asserted-by":"publisher","first-page":"2067","DOI":"10.1016\/S0002-9440(10)64678-0","volume":"158","author":"DA Tweddle","year":"2001","unstructured":"Tweddle DA, Malcolm AJ, Cole M et al (2001) p53 cellular localization and function in neuroblastoma: evidence for defective G(1) arrest despite WAF1 induction in MYCN-amplified cells. Am J Pathol 158:2067\u20132077. https:\/\/doi.org\/10.1016\/S0002-9440(10)64678-0","journal-title":"Am J Pathol"},{"key":"1028_CR71","doi-asserted-by":"publisher","first-page":"1029","DOI":"10.1126\/science.1160809","volume":"324","author":"MG Vander Heiden","year":"2009","unstructured":"Vander Heiden MG, Cantley LC, Thompson CB (2009) Understanding the Warburg effect: the metabolic requirements of cell proliferation. Science 324:1029\u20131033. https:\/\/doi.org\/10.1126\/science.1160809","journal-title":"Science"},{"key":"1028_CR72","doi-asserted-by":"publisher","first-page":"5899","DOI":"10.1128\/MCB.21.17.5899-5912.2001","volume":"21","author":"MG Vander Heiden","year":"2001","unstructured":"Vander Heiden MG, Plas DR, Rathmell JC et al (2001) Growth factors can influence cell growth and survival through effects on glucose metabolism. Mol Cell Biol 21:5899\u20135912. https:\/\/doi.org\/10.1128\/MCB.21.17.5899-5912.2001","journal-title":"Mol Cell Biol"},{"key":"1028_CR73","doi-asserted-by":"publisher","first-page":"121","DOI":"10.1016\/j.bbadis.2012.09.013","volume":"1832","author":"AP Waller","year":"2013","unstructured":"Waller AP, George M, Kalyanasundaram A et al (2013) GLUT12 functions as a basal and insulin-independent glucose transporter in the heart. Biochim Biophys Acta 1832:121\u2013127. https:\/\/doi.org\/10.1016\/j.bbadis.2012.09.013","journal-title":"Biochim Biophys Acta"},{"key":"1028_CR74","doi-asserted-by":"publisher","first-page":"315","DOI":"10.1111\/j.1939-1676.2010.0674.x","volume":"25","author":"AP Waller","year":"2011","unstructured":"Waller AP, Burns TA, Mudge MC et al (2011) Insulin resistance selectively alters cell-surface glucose transporters but not their total protein expression in equine skeletal muscle. J Vet Intern Med 25:315\u2013321. https:\/\/doi.org\/10.1111\/j.1939-1676.2010.0674.x","journal-title":"J Vet Intern Med"},{"key":"1028_CR75","doi-asserted-by":"publisher","first-page":"1098","DOI":"10.1016\/j.bbadis.2011.02.007","volume":"1812","author":"AP Waller","year":"2011","unstructured":"Waller AP, Kohler K, Burns TA et al (2011) Naturally occurring compensated insulin resistance selectively alters glucose transporters in visceral and subcutaneous adipose tissues without change in AS160 activation. Biochim Biophys Acta 1812:1098\u20131103. https:\/\/doi.org\/10.1016\/j.bbadis.2011.02.007","journal-title":"Biochim Biophys Acta"},{"key":"1028_CR76","doi-asserted-by":"publisher","first-page":"453","DOI":"10.1530\/ERC-17-0051","volume":"25","author":"MA White","year":"2018","unstructured":"White MA, Tsouko E, Lin C et al (2018) GLUT12 promotes prostate cancer cell growth and is regulated by androgens and CaMKK2 signaling. Endocr Relat Cancer 25:453\u2013469. https:\/\/doi.org\/10.1530\/ERC-17-0051","journal-title":"Endocr Relat Cancer"},{"key":"1028_CR77","doi-asserted-by":"publisher","unstructured":"Wibmer AG, Becher T, Eljalby M, et al (2021) Brown adipose tissue is associated with healthier body fat distribution and metabolic benefits independent of regional adiposity. Cell Rep Med 2:100332. https:\/\/doi.org\/10.1016\/j.xcrm.2021.100332","DOI":"10.1016\/j.xcrm.2021.100332"},{"key":"1028_CR78","doi-asserted-by":"publisher","first-page":"917","DOI":"10.1210\/en.2007-0985","volume":"149","author":"AL Wilson-O\u2019Brien","year":"2008","unstructured":"Wilson-O\u2019Brien AL, Dehaan CL, Rogers S (2008) Mitogen-stimulated and rapamycin-sensitive glucose transporter 12 targeting and functional glucose transport in renal epithelial cells. Endocrinology 149:917\u2013924. https:\/\/doi.org\/10.1210\/en.2007-0985","journal-title":"Endocrinology"},{"key":"1028_CR79","doi-asserted-by":"publisher","first-page":"152","DOI":"10.1186\/1471-2148-10-152","volume":"10","author":"AL Wilson-O\u2019Brien","year":"2010","unstructured":"Wilson-O\u2019Brien AL, Patron N, Rogers S (2010) Evolutionary ancestry and novel functions of the mammalian glucose transporter (GLUT) family. BMC Evol Biol 10:152. https:\/\/doi.org\/10.1186\/1471-2148-10-152","journal-title":"BMC Evol Biol"},{"key":"1028_CR80","doi-asserted-by":"publisher","first-page":"823","DOI":"10.3390\/cancers14030823","volume":"14","author":"JE Zawacka-Pankau","year":"2022","unstructured":"Zawacka-Pankau JE (2022) The Role of p53 Family in Cancer. Cancers 14:823. https:\/\/doi.org\/10.3390\/cancers14030823","journal-title":"Cancers"},{"key":"1028_CR81","doi-asserted-by":"publisher","first-page":"41600","DOI":"10.1074\/jbc.M111.240812","volume":"286","author":"J Zawacka-Pankau","year":"2011","unstructured":"Zawacka-Pankau J, Grinkevich VV, H\u00fcnten S et al (2011) Inhibition of glycolytic enzymes mediated by pharmacologically activated p53: targeting Warburg effect to fight cancer. J Biol Chem 286:41600\u201341615. https:\/\/doi.org\/10.1074\/jbc.M111.240812","journal-title":"J Biol Chem"}],"container-title":["Journal of Physiology and Biochemistry"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s13105-024-01028-9.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s13105-024-01028-9\/fulltext.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s13105-024-01028-9.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,7,22]],"date-time":"2025-07-22T01:28:54Z","timestamp":1753147734000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s13105-024-01028-9"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2024,5,10]]},"references-count":81,"journal-issue":{"issue":"2","published-print":{"date-parts":[[2025,5]]}},"alternative-id":["1028"],"URL":"https:\/\/doi.org\/10.1007\/s13105-024-01028-9","relation":{},"ISSN":["1138-7548","1877-8755"],"issn-type":[{"value":"1138-7548","type":"print"},{"value":"1877-8755","type":"electronic"}],"subject":[],"published":{"date-parts":[[2024,5,10]]},"assertion":[{"value":"13 December 2023","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"1 May 2024","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"10 May 2024","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"Samples and data from patients included in the study were provided by the Biobank of the University of Navarra and were processed following standard operating procedures approved by the Ethical and Scientific Committees (protocol n. 2011.006mod1). The animal experiments were performed according to the national animal care guidelines and with the approval of the Ethics Committee for Animal Experimentation of the University of Navarra (protocol n. 113\u201315).","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethics approval"}},{"value":"Not applicable.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent to participate"}},{"value":"Not applicable.","order":4,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent for publication"}},{"value":"The authors declare no competing interest.","order":5,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}]}}