{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,24]],"date-time":"2026-06-24T06:46:41Z","timestamp":1782283601227,"version":"3.54.5"},"reference-count":58,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,6,24]],"date-time":"2026-06-24T00:00:00Z","timestamp":1782259200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Nutr."],"abstract":"<jats:sec>\n                    <jats:title>Introduction<\/jats:title>\n                    <jats:p>Beyond their role in chemosensory perception, taste receptors function as metabolic regulators extra-orally. While decreased taste receptor expression in gustatory tissue is linked to obesity-associated inflammation, we tested the hypothesis that taste receptor-inflammation interactions extend to extra-oral contexts.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods<\/jats:title>\n                    <jats:p>\n                      Obesity-associated metabolic syndrome-like phenotype was induced in female C57BL\/6 mice by administration of a cafeteria diet for 18\u202fweeks. The controls consumed a standard diet. Gene-expression levels of\n                      <jats:italic>Tas1r2<\/jats:italic>\n                      ,\n                      <jats:italic>Tas1r3<\/jats:italic>\n                      ,\n                      <jats:italic>CD36<\/jats:italic>\n                      ,\n                      <jats:italic>Ffar1<\/jats:italic>\n                      ,\n                      <jats:italic>GPR84<\/jats:italic>\n                      and\n                      <jats:italic>Ffar4<\/jats:italic>\n                      were analyzed in circumvallate papillae, lower jejunum, stomach and spleen using RT-qPCR. The number and area of fungiform papillae were assessed on the anterior tongue. Correlation network connectivity for taste receptor expression and fungiform papillae markers with inflammatory and cardiometabolic risk markers was calculated tissue-specifically.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results and discussion<\/jats:title>\n                    <jats:p>\n                      The cafeteria diet induced obesity, metabolic dysfunction, and systemic inflammation. Additionally, number and area of fungiform papillae were reduced, and\n                      <jats:italic>Tas1r2, CD36, Ffar1<\/jats:italic>\n                      and\n                      <jats:italic>GPR84<\/jats:italic>\n                      were downregulated in circumvallate papillae, whereas in jejunum only\n                      <jats:italic>CD36<\/jats:italic>\n                      ,\n                      <jats:italic>Ffar1,<\/jats:italic>\n                      and\n                      <jats:italic>GPR84<\/jats:italic>\n                      were downregulated following cafeteria diet. Gustatory sweet and fatty acid receptor expression correlated with dietary intake patterns. Exploratory network analysis indicated tissue-specific correlation patterns, with taste receptor-inflammation crosstalk predominantly appearing orally. In contrast, extra-oral sweet and fatty acid receptor expression showed weaker or absent correlations with systemic inflammatory markers under the conditions tested.\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.3389\/fnut.2026.1857562","type":"journal-article","created":{"date-parts":[[2026,6,24]],"date-time":"2026-06-24T05:47:52Z","timestamp":1782280072000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Taste receptor downregulation in diet-induced obesity reveals circumvallate papillae-specific taste-inflammatory networks"],"prefix":"10.3389","volume":"13","author":[{"given":"Isabella","family":"Kimmeswenger","sequence":"first","affiliation":[{"name":"Faculty of Chemistry, Institute of Physiological Chemistry, University of Vienna","place":["Vienna, Austria"]},{"name":"Vienna Doctoral School in Chemistry (DoSChem), University of Vienna","place":["Vienna, Austria"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Andrej","family":"Feje\u0161","sequence":"additional","affiliation":[{"name":"Faculty of Medicine, Institute of Molecular Biomedicine, Comenius University","place":["Bratislava, Slovakia"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Veronika","family":"Somoza","sequence":"additional","affiliation":[{"name":"Faculty of Chemistry, Institute of Physiological Chemistry, University of Vienna","place":["Vienna, Austria"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Katar\u00edna","family":"\u0160ebekov\u00e1","sequence":"additional","affiliation":[{"name":"Faculty of Medicine, Institute of Molecular Biomedicine, Comenius University","place":["Bratislava, Slovakia"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Barbara","family":"Lieder","sequence":"additional","affiliation":[{"name":"Faculty of Chemistry, Institute of Physiological Chemistry, University of Vienna","place":["Vienna, Austria"]},{"name":"Christian Doppler Laboratory for Taste Research, Faculty of Chemistry, University of Vienna","place":["Vienna, Austria"]},{"name":"Institute of Clinical Nutrition, University of Hohenheim","place":["Stuttgart, Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,6,24]]},"reference":[{"key":"ref1","doi-asserted-by":"publisher","first-page":"881177","DOI":"10.3389\/fnut.2022.881177","article-title":"Extra-Oral taste receptors-function, disease, and perspectives","volume":"9","author":"Behrens","year":"2022","journal-title":"Front Nutr"},{"key":"ref2","doi-asserted-by":"publisher","first-page":"1737","DOI":"10.1007\/s00394-017-1456-5","article-title":"Effect of GPR84 deletion on obesity and diabetes development in mice fed long chain or medium chain fatty acid rich diets","volume":"57","author":"Du Toit","year":"2018","journal-title":"Eur J Nutr"},{"key":"ref3","doi-asserted-by":"publisher","first-page":"2280","DOI":"10.2337\/db08-0307","article-title":"Gpr40 is expressed in enteroendocrine cells and mediates free fatty acid stimulation of incretin secretion","volume":"57","author":"Edfalk","year":"2008","journal-title":"Diabetes"},{"key":"ref4","doi-asserted-by":"publisher","first-page":"379","DOI":"10.1097\/MCO.0000000000000058","article-title":"Functional roles of the sweet taste receptor in oral and extraoral tissues","volume":"17","author":"Laffitte","year":"2014","journal-title":"Curr Opin Clin Nutr Metab Care"},{"key":"ref5","doi-asserted-by":"publisher","first-page":"1457","DOI":"10.4155\/fmc.15.75","article-title":"Biological characteristics and agonists of GPR120 (FFAR4) receptor: the present status of research","volume":"7","author":"Li","year":"2015","journal-title":"Future Med Chem"},{"key":"ref6","doi-asserted-by":"publisher","first-page":"281","DOI":"10.1146\/annurev-nutr-071812-161220","article-title":"Structure-function of CD36 and importance of fatty acid signal transduction in fat metabolism","volume":"34","author":"Pepino","year":"2014","journal-title":"Annu Rev Nutr"},{"key":"ref7","doi-asserted-by":"publisher","first-page":"15069","DOI":"10.1073\/pnas.0706890104","article-title":"Gut-expressed gustducin and taste receptors regulate secretion of glucagon-like peptide-1","volume":"104","author":"Jang","year":"2007","journal-title":"Proc Natl Acad Sci USA"},{"key":"ref8","doi-asserted-by":"publisher","first-page":"15075","DOI":"10.1073\/pnas.0706678104","article-title":"T1R3 and gustducin in gut sense sugars to regulate expression of Na+\u2212glucose cotransporter 1","volume":"104","author":"Margolskee","year":"2007","journal-title":"Proc Natl Acad Sci USA"},{"key":"ref9","doi-asserted-by":"publisher","first-page":"3532","DOI":"10.2337\/db13-0581","article-title":"Disordered control of intestinal sweet taste receptor expression and glucose absorption in type 2 diabetes","volume":"62","author":"Young","year":"2013","journal-title":"Diabetes"},{"key":"ref10","doi-asserted-by":"publisher","first-page":"5","DOI":"10.1111\/dom.13129","article-title":"Incretin hormones: their role in health and disease","volume":"20","author":"Nauck","year":"2018","journal-title":"Diabetes Obes Metab"},{"key":"ref11","doi-asserted-by":"publisher","first-page":"E464","DOI":"10.1152\/ajpendo.00163.2012","article-title":"Transformation of postingestive glucose responses after deletion of sweet taste receptor subunits or gastric bypass surgery","volume":"303","author":"Geraedts","year":"2012","journal-title":"Am J Physiol Endocrinol Metab"},{"key":"ref12","doi-asserted-by":"publisher","first-page":"89","DOI":"10.1016\/j.rvsc.2022.05.017","article-title":"Chemosensing of fat digestion by the expression pattern of GPR40, GPR120, CD36 and enteroendocrine profile in sheep","volume":"150","author":"Krishnan","year":"2022","journal-title":"Res Vet Sci"},{"key":"ref13","doi-asserted-by":"publisher","first-page":"101193","DOI":"10.1016\/j.plipres.2022.101193","article-title":"CD36 favours fat sensing and transport to govern lipid metabolism","volume":"88","author":"Li","year":"2022","journal-title":"Prog Lipid Res"},{"key":"ref14","doi-asserted-by":"publisher","first-page":"41","DOI":"10.1016\/j.plipres.2016.03.002","article-title":"Mechanism of fat taste perception: association with diet and obesity","volume":"63","author":"Liu","year":"2016","journal-title":"Prog Lipid Res"},{"key":"ref15","doi-asserted-by":"publisher","first-page":"11","DOI":"10.1186\/s12576-024-00904-9","article-title":"Satiety: a gut-brain-relationship","volume":"74","author":"Barakat","year":"2024","journal-title":"J Physiol Sci"},{"key":"ref16","doi-asserted-by":"publisher","first-page":"523","DOI":"10.1007\/s00210-007-0200-8","article-title":"Free fatty acids induce cholecystokinin secretion through GPR120","volume":"377","author":"Tanaka","year":"2008","journal-title":"Naunyn Schmiedeberg's Arch Pharmacol"},{"key":"ref17","doi-asserted-by":"publisher","first-page":"108215","DOI":"10.1016\/j.appet.2025.108215","article-title":"Weight loss induced by calorie restriction in mice is associated with a higher sweet taste response and increased fungiform papillae density","volume":"214","author":"Kimmeswenger","year":"2025","journal-title":"Appetite"},{"key":"ref18","doi-asserted-by":"publisher","first-page":"594","DOI":"10.3390\/nu14030594","article-title":"Long-term consumption of a sugar-sweetened soft drink in combination with a Western-type diet is associated with morphological and molecular changes of taste markers independent of body weight development in mice","volume":"14","author":"Lieder","year":"2022","journal-title":"Nutrients"},{"key":"ref19","doi-asserted-by":"publisher","first-page":"115","DOI":"10.1016\/j.peptides.2013.07.015","article-title":"Alteration of sweet taste in high-fat diet induced obese rats after 4 weeks treatment with exenatide","volume":"47","author":"Zhang","year":"2013","journal-title":"Peptides"},{"key":"ref20","doi-asserted-by":"publisher","first-page":"e2001959","DOI":"10.1371\/journal.pbio.2001959","article-title":"Inflammation arising from obesity reduces taste bud abundance and inhibits renewal","volume":"16","author":"Kaufman","year":"2018","journal-title":"PLoS Biol"},{"key":"ref21","doi-asserted-by":"publisher","first-page":"1475","DOI":"10.1038\/s41366-018-0303-y","article-title":"Obesity is associated with altered gene expression in human tastebuds","volume":"43","author":"Archer","year":"2019","journal-title":"Int J Obes"},{"key":"ref22","doi-asserted-by":"publisher","first-page":"29094","DOI":"10.1038\/srep29094","article-title":"Impact of obesity on taste receptor expression in extra-oral tissues: emphasis on hypothalamus and brainstem","volume":"6","author":"Herrera Moro Chao","year":"2016","journal-title":"Sci Rep"},{"key":"ref23","doi-asserted-by":"publisher","first-page":"92","DOI":"10.1016\/j.neubiorev.2020.11.003","article-title":"The cafeteria diet: a standardized protocol and its effects on behavior","volume":"122","author":"Lalanza","year":"2021","journal-title":"Neurosci Biobehav Rev"},{"key":"ref24","doi-asserted-by":"publisher","first-page":"101724","DOI":"10.1016\/j.molmet.2023.101724","article-title":"Disentangling the effects of obesity and high-fat diet on glucose homeostasis using a photoperiod induced obesity model implicates ectopic fat deposition as a key factor","volume":"73","author":"Yang","year":"2023","journal-title":"Mol Metab"},{"key":"ref25","doi-asserted-by":"publisher","first-page":"806","DOI":"10.1016\/j.immuni.2013.10.010","article-title":"The spleen in local and systemic regulation of immunity","volume":"39","author":"Bronte","year":"2013","journal-title":"Immunity"},{"key":"ref26","doi-asserted-by":"publisher","first-page":"606","DOI":"10.1038\/nri1669","article-title":"Structure and function of the spleen","volume":"5","author":"Mebius","year":"2005","journal-title":"Nat Rev Immunol"},{"key":"ref27","doi-asserted-by":"publisher","first-page":"e3000410","DOI":"10.1371\/journal.pbio.3000410","article-title":"The ARRIVE guidelines 2.0: updated guidelines for reporting animal research","volume":"18","author":"Du Percie Sert","year":"2020","journal-title":"PLoS Biol"},{"key":"ref28","doi-asserted-by":"publisher","first-page":"2402","DOI":"10.1210\/jcem.85.7.6661","article-title":"Quantitative insulin sensitivity check index: a simple, accurate method for assessing insulin sensitivity in humans","volume":"85","author":"Katz","year":"2000","journal-title":"J Clin Endocrinol Metab"},{"key":"ref29","doi-asserted-by":"publisher","first-page":"e45","DOI":"10.1093\/nar\/gkp045","article-title":"Amplification efficiency: linking baseline and bias in the analysis of quantitative PCR data","volume":"37","author":"Ruijter","year":"2009","journal-title":"Nucleic Acids Res"},{"key":"ref30","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1002\/(SICI)1096-9861(19990621)409:1<13::AID-CNE2>3.0.CO;2-O","article-title":"Alterations in size, number, and morphology of gustatory papillae and taste buds in BDNF null mutant mice demonstrate neural dependence of developing taste organs","volume":"409","author":"Mistretta","year":"1999","journal-title":"J Comp Neurol"},{"key":"ref31","doi-asserted-by":"publisher","first-page":"289","DOI":"10.1111\/j.2517-6161.1995.tb02031.x","article-title":"Controlling the false discovery rate: a practical and powerful approach to multiple testing","volume":"57","author":"Benjamini","year":"1995","journal-title":"J R Stat Soc Ser B Stat Methodol"},{"key":"ref32","doi-asserted-by":"publisher","first-page":"1109","DOI":"10.1038\/oby.2011.18","article-title":"Cafeteria diet is a robust model of human metabolic syndrome with liver and adipose inflammation: comparison to high-fat diet","volume":"19","author":"Sampey","year":"2011","journal-title":"Obesity (Silver Spring)"},{"key":"ref33","doi-asserted-by":"publisher","first-page":"5349","DOI":"10.1182\/bloodadvances.2021004438","article-title":"Anti-inflammatory treatment in MPN: targeting TNFR1 and TNFR2 in JAK2-V617F-induced disease","volume":"5","author":"M\u00fcller","year":"2021","journal-title":"Blood Adv"},{"key":"ref34","doi-asserted-by":"publisher","first-page":"475","DOI":"10.1016\/j.cytogfr.2015.07.004","article-title":"Interleukin-6: biology, signaling and strategies of blockade","volume":"26","author":"Schaper","year":"2015","journal-title":"Cytokine Growth Factor Rev"},{"key":"ref35","doi-asserted-by":"publisher","first-page":"101941","DOI":"10.1016\/j.berh.2024.101941","article-title":"TNFR2 signalling in inflammatory diseases","volume":"38","author":"Williams","year":"2024","journal-title":"Best Pract Res Clin Rheumatol"},{"key":"ref36","doi-asserted-by":"publisher","first-page":"739","DOI":"10.1038\/s41366-019-0429-6","article-title":"Taste loss with obesity in mice and men","volume":"44","author":"Kaufman","year":"2020","journal-title":"Int J Obes"},{"key":"ref37","doi-asserted-by":"publisher","first-page":"112294","DOI":"10.1016\/j.foodres.2022.112294","article-title":"Tongue papillae density and fat taster status\u2014a cardinal role on sweet and bitter taste perception among Indian population","volume":"163","author":"Ponnusamy","year":"2023","journal-title":"Food Res Int"},{"key":"ref38","doi-asserted-by":"publisher","first-page":"3062","DOI":"10.3390\/nu13093062","article-title":"Sprague Dawley rats gaining weight on a high energy diet exhibit damage to taste tissue even after return to a healthy diet","volume":"13","author":"Harnischfeger","year":"2021","journal-title":"Nutrients"},{"key":"ref39","doi-asserted-by":"publisher","first-page":"106","DOI":"10.3390\/j1010011","article-title":"Receptor regulation in taste: can diet influence how we perceive foods?","volume":"1","author":"Shahbandi","year":"2018","journal-title":"J"},{"key":"ref40","doi-asserted-by":"publisher","first-page":"53","DOI":"10.1016\/j.brainres.2010.02.026","article-title":"Nutritional status alters saccharin intake and sweet receptor mRNA expression in rat taste buds","volume":"1325","author":"Chen","year":"2010","journal-title":"Brain Res"},{"key":"ref41","doi-asserted-by":"publisher","first-page":"e111232","DOI":"10.1371\/journal.pone.0111232","article-title":"Effect of diet on preference and intake of sucrose in obese prone and resistant rats","volume":"9","author":"Duca","year":"2014","journal-title":"PLoS One"},{"key":"ref42","doi-asserted-by":"publisher","first-page":"381","DOI":"10.1016\/s0092-8674(01)00451-2","article-title":"Mammalian sweet taste receptors","volume":"106","author":"Nelson","year":"2001","journal-title":"Cell"},{"key":"ref43","doi-asserted-by":"publisher","first-page":"4692","DOI":"10.1073\/pnas.072090199","article-title":"Human receptors for sweet and umami taste","volume":"99","author":"Li","year":"2002","journal-title":"Proc Natl Acad Sci USA"},{"key":"ref44","doi-asserted-by":"publisher","first-page":"663","DOI":"10.1016\/j.acthis.2010.09.007","article-title":"Decreased expression of CD36 in circumvallate taste buds of high-fat diet induced obese rats","volume":"113","author":"Zhang","year":"2011","journal-title":"Acta Histochem"},{"key":"ref45","doi-asserted-by":"publisher","first-page":"687","DOI":"10.1016\/j.cell.2010.07.041","article-title":"GPR120 is an omega-3 fatty acid receptor mediating potent anti-inflammatory and insulin-sensitizing effects","volume":"142","author":"Da Oh","year":"2010","journal-title":"Cell"},{"key":"ref46","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1111\/j.1749-6632.1998.tb10560.x","article-title":"Dietary fat preferences are inversely correlated with peripheral gustatory fatty acid sensitivity a","volume":"855","author":"Gilbertson","year":"1998","journal-title":"Ann N Y Acad Sci"},{"key":"ref47","doi-asserted-by":"publisher","first-page":"145","DOI":"10.1017\/S0007114510000267","article-title":"Oral sensitivity to fatty acids, food consumption and BMI in human subjects","volume":"104","author":"Stewart","year":"2010","journal-title":"Br J Nutr"},{"key":"ref48","doi-asserted-by":"publisher","first-page":"3543","DOI":"10.3390\/nu14173543","article-title":"Chemosensory functions in patients with inflammatory bowel disease and their association with clinical disease activity","volume":"14","author":"Han","year":"2022","journal-title":"Nutrients"},{"key":"ref49","doi-asserted-by":"publisher","first-page":"267","DOI":"10.1016\/j.jobcr.2023.02.008","article-title":"Detection of SARS-CoV-2 IgG antibodies and inflammatory cytokines in saliva-a pilot study","volume":"13","author":"Wadhwa","year":"2023","journal-title":"J Oral Biol Craniofac Res"},{"key":"ref50","doi-asserted-by":"publisher","first-page":"1675","DOI":"10.1007\/s00405-020-05874-3","article-title":"Chemosensory function is decreased in rheumatoid arthritis","volume":"277","author":"Walliczek-Dworschak","year":"2020","journal-title":"Eur Arch Otorrinolaringol"},{"key":"ref51","doi-asserted-by":"publisher","first-page":"32785183","DOI":"10.3390\/ijerph17165792","article-title":"Correlation of PTC taste status with fungiform papillae count and body mass index in smokers and non-smokers of Eastern Province, Saudi Arabia","volume":"17","author":"Khan","year":"2020","journal-title":"Int J Environ Res Public Health"},{"key":"ref52","doi-asserted-by":"publisher","first-page":"148","DOI":"10.1186\/s12986-025-01050-8","article-title":"High physical activity is associated with decreased fungiform papillae area and number, elevated sucrose recognition thresholds, and increased IL-6 levels: an observational human study","volume":"22","author":"Kimmeswenger","year":"2025","journal-title":"Nutr Metab (Lond)"},{"key":"ref53","doi-asserted-by":"publisher","first-page":"284","DOI":"10.1002\/oby.22684","article-title":"Differential effects of diet and weight on taste responses in diet-induced obese mice","volume":"28","author":"Ahart","year":"2020","journal-title":"Obesity (Silver Spring)"},{"key":"ref54","doi-asserted-by":"publisher","first-page":"E566","DOI":"10.1152\/ajpendo.00331.2020","article-title":"Sex differences in fat taste responsiveness are modulated by estradiol","volume":"320","author":"Dahir","year":"2021","journal-title":"Am J Physiol Endocrinol Metab"},{"key":"ref55","doi-asserted-by":"publisher","first-page":"e13665","DOI":"10.1111\/obr.13665","article-title":"Sex differences in systemic inflammation and immune function in diet-induced obesity rodent models: a systematic review","volume":"25","author":"Braga Tibaes","year":"2024","journal-title":"Obes Rev"},{"key":"ref56","doi-asserted-by":"publisher","first-page":"105","DOI":"10.1186\/s13293-024-00677-1","article-title":"Sex differentially affects pro-inflammatory cell subsets in adipose tissue depots in a diet induced obesity model","volume":"15","author":"Schuetz","year":"2024","journal-title":"Biol Sex Differ"},{"key":"ref57","doi-asserted-by":"publisher","first-page":"828522","DOI":"10.3389\/fnut.2022.828522","article-title":"Sex and species differences in the development of diet-induced obesity and metabolic disturbances in rodents","volume":"9","author":"Maric","year":"2022","journal-title":"Front Nutr"},{"key":"ref58","doi-asserted-by":"publisher","first-page":"E559","DOI":"10.1152\/ajpendo.00036.2019","article-title":"Metabolic control via nutrient-sensing mechanisms: role of taste receptors and the gut-brain neuroendocrine axis","volume":"317","author":"Raka","year":"2019","journal-title":"Am J Physiol Endocrinol Metab"}],"container-title":["Frontiers in Nutrition"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fnut.2026.1857562\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,6,24]],"date-time":"2026-06-24T05:47:53Z","timestamp":1782280073000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fnut.2026.1857562\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,6,24]]},"references-count":58,"alternative-id":["10.3389\/fnut.2026.1857562"],"URL":"https:\/\/doi.org\/10.3389\/fnut.2026.1857562","relation":{},"ISSN":["2296-861X"],"issn-type":[{"value":"2296-861X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,6,24]]},"article-number":"1857562"}}