{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,11]],"date-time":"2026-10-11T12:10:39Z","timestamp":1791720639865,"version":"4.3.4"},"reference-count":47,"publisher":"MDPI AG","issue":"2","license":[{"start":{"date-parts":[[2021,2,6]],"date-time":"2021-02-06T00:00:00Z","timestamp":1612569600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Nutrients"],"abstract":"<jats:p>The aim of this study was to investigate the underlying mechanism for the improvement of glucose tolerance following intake of high \u03b2-glucan barley (HGB) in terms of intestinal metabolism. C57BL\/6J male mice were fed a fatty diet supplemented with HGB corresponding to 5% of dietary fiber for 83 days. An oral glucose tolerance test was performed at the end of the experimental period. The concentration of short-chain fatty acids (SCFAs) in the cecum was analyzed by GC\u2013MS (gas chromatography\u2013mass spectrometry). The mRNA expression levels related to L cell function in the ileum were measured by real-time PCR. Glucagon-like peptide-1 (GLP-1) levels in the portal vein and cecal content were assessed by enzyme-linked immunosorbent assay. GLP-1-producing L cells of the ileum were quantified by immunohistochemistry. HGB intake improved glucose tolerance and increased the cecal levels of SCFAs, acetate, and propionate. The number of GLP-1-positive L cells in the HGB group was significantly higher than in the control group. GLP-1 levels in the portal vein and cecal GLP-1 pool size in the HGB group were significantly higher than the control group. In conclusion, we report improved glucose tolerance after HGB intake induced by an increase in L cell number and subsequent rise in GLP-1 secretion.<\/jats:p>","DOI":"10.3390\/nu13020527","type":"journal-article","created":{"date-parts":[[2021,2,7]],"date-time":"2021-02-07T14:07:40Z","timestamp":1612706860000},"page":"527","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":45,"title":["High \u03b2-Glucan Barley Supplementation Improves Glucose Tolerance by Increasing GLP-1 Secretion in Diet-Induced Obesity Mice"],"prefix":"10.3390","volume":"13","author":[{"given":"Sachina","family":"Suzuki","sequence":"first","affiliation":[{"name":"The Institute of Human Culture Studies, Otsuma Women\u2019s University, Chiyoda-ku, Tokyo 102-8357, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2852-7405","authenticated-orcid":false,"given":"Seiichiro","family":"Aoe","sequence":"additional","affiliation":[{"name":"The Institute of Human Culture Studies, Otsuma Women\u2019s University, Chiyoda-ku, Tokyo 102-8357, Japan"},{"name":"Studies in Human Life Sciences, Graduate School of Studies in Human Culture, Otsuma Women\u2019s University, Chiyoda-ku, Tokyo 102-8357, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2021,2,6]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"182","DOI":"10.1080\/07315724.2005.10719464","article-title":"Comparison of hormone and glucose responses of overweight women to barley and oats","volume":"24","author":"Behall","year":"2005","journal-title":"J. Am. Coll. Nutr."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"313","DOI":"10.1080\/07315724.2006.10719541","article-title":"Post-prandial responses to cereal products enriched with barley \u03b2-glucan","volume":"25","author":"Casiraghi","year":"2006","journal-title":"J. Am. Coll. Nutr."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"1666","DOI":"10.3945\/jn.112.161372","article-title":"Barley cultivar, kernel composition, and processing affect the glycemic index","volume":"142","author":"Aldughpassi","year":"2012","journal-title":"J. Nutr."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"1472","DOI":"10.1038\/ejcn.2010.178","article-title":"Beta-glucan from barley and its lipid-lowering capacity, a meta-analysis of randomized, controlled trials","volume":"64","author":"AbuMweis","year":"2010","journal-title":"Eur. J. Clin. Nutr."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"1185","DOI":"10.1093\/ajcn\/80.5.1185","article-title":"Diets containing barley significantly reduce lipids in mildly hypercholesterolemic men and women","volume":"80","author":"Behall","year":"2004","journal-title":"Am. J. Clin. Nutr."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"720","DOI":"10.3945\/jn.115.223206","article-title":"High-Molecular-Weight beta-Glucan Decreases Serum Cholesterol Differentially Based on the CYP7A1 rs3808607 Polymorphism in Mildly Hypercholesterolemic Adults","volume":"146","author":"Wang","year":"2016","journal-title":"J. Nutr."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"482","DOI":"10.1111\/j.1574-6941.2008.00478.x","article-title":"In vitro fermentation of oat and barley derived beta-glucans by human faecal microbiota","volume":"64","author":"Hughes","year":"2008","journal-title":"FEMS Microbiol. Ecol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"553","DOI":"10.1111\/j.1574-6941.2011.01074.x","article-title":"Nonstarch polysaccharide-degrading enzymes alter the microbial community and the fermentation patterns of barley cultivars and wheat products in an in vitro model of the porcine gastrointestinal tract","volume":"76","author":"Bindelle","year":"2011","journal-title":"FEMS Microbiol. Ecol."},{"key":"ref_9","doi-asserted-by":"crossref","unstructured":"Aoe, S., Yamanaka, C., Fuwa, M., Tamiya, T., Nakayama, Y., Miyoshi, T., and Kitazono, E. (2019). Effects of BARLEYmax and high-beta-glucan barley line on short-chain fatty acids production and microbiota from the cecum to the distal colon in rats. PLoS ONE, 14.","DOI":"10.1371\/journal.pone.0218118"},{"key":"ref_10","first-page":"19","article-title":"Effect of ingestion of high \u03b2-glucan barley breads with different particle size on postprandial blood glucose levels","volume":"21","author":"Kanamoto","year":"2017","journal-title":"Luminacoids Res."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"283","DOI":"10.4327\/jsnfs.71.283","article-title":"Effects of Various Blending Ratios of Rice and Waxy Barley on Postprandial Blood Glucose Levels","volume":"71","author":"Aoe","year":"2018","journal-title":"J. Jpn. Soc. Nutr. Food Sci."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1007\/s11130-007-0064-6","article-title":"Effect of high beta-glucan barley on serum cholesterol concentrations and visceral fat area in Japanese men\u2014A randomized, double-blinded, placebo-controlled trial","volume":"63","author":"Shimizu","year":"2008","journal-title":"Plant Foods Hum. Nutr."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.nut.2017.05.002","article-title":"Effects of high beta-glucan barley on visceral fat obesity in Japanese individuals: A randomized, double-blind study","volume":"42","author":"Aoe","year":"2017","journal-title":"Nutrition"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"325","DOI":"10.1007\/s11130-014-0437-6","article-title":"Effect of cooked white rice with high \u03b2-glucan barley on appetite and energy intake in healthy Japanese subjects: A randomized controlled trial","volume":"69","author":"Aoe","year":"2014","journal-title":"Plant Foods Hum. Nutr."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"46","DOI":"10.1186\/1475-2891-12-46","article-title":"Effects of indigestible carbohydrates in barley on glucose metabolism, appetite and voluntary food intake over 16 h in healthy adults","volume":"12","author":"Johansson","year":"2013","journal-title":"Nutr. J."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"817","DOI":"10.1093\/ajcn\/83.4.817","article-title":"Colonic fermentation of indigestible carbohydrates contributes to the second-meal effect","volume":"83","author":"Brighenti","year":"2006","journal-title":"Am. J. Clin. Nutr."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"1339","DOI":"10.1093\/ajcn\/35.6.1339","article-title":"Slow release dietary carbohydrate improves second meal tolerance","volume":"35","author":"Jenkins","year":"1982","journal-title":"Am. J. Clin. Nutr."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1041","DOI":"10.1093\/ajcn\/48.4.1041","article-title":"Secondmeal effect: Low-glycemic-index foods eaten at dinner improve subsequent breakfast glycemic response","volume":"48","author":"Wolever","year":"1988","journal-title":"Am. J. Clin. Nutr."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"1055","DOI":"10.1210\/me.2015-1007","article-title":"An Acetate-Specific GPCR, FFAR2, Regulates Insulin Secretion","volume":"29","author":"Priyadarshini","year":"2015","journal-title":"Mol. Endocrinol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"1848","DOI":"10.1016\/0016-5085(94)90831-1","article-title":"Glucagon like peptide 1: A newly discovered gastrointestinal hormone","volume":"107","author":"Holst","year":"1994","journal-title":"Gastroenterology"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"535","DOI":"10.2337\/diab.43.4.535","article-title":"Tissue and plasma concentrations of amidated and glycine-extended glucagonlike peptide I in humans","volume":"43","author":"Orskov","year":"1994","journal-title":"Diabetes"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"3948","DOI":"10.1210\/endo.137.9.8756571","article-title":"Dietary fiber modulates intestinal proglucagon messenger ribonucleic acid and postprandial secretion of glucagon-like peptide-1 and insulin in rats","volume":"137","author":"Reimer","year":"1996","journal-title":"Endocrinology"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"1923","DOI":"10.1093\/jn\/127.10.1923","article-title":"A physiological level of rhubarb fiber increases proglucagon gene expression and modulates intestinal glucose uptake in rats","volume":"127","author":"Reimer","year":"1997","journal-title":"J. Nutr."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"851","DOI":"10.1016\/S0271-5317(00)00169-X","article-title":"Proglucagon messenger ribonucleic acid and intestinal glucose uptake are modulated by fermentable fiber and food intake in diabetic rats","volume":"20","author":"Reimer","year":"2000","journal-title":"Nutr. Res."},{"key":"ref_25","doi-asserted-by":"crossref","unstructured":"Mio, K., Yamanaka, C., Matsuoka, T., Kobayashi, T., and Aoe, S. (2020). Effects of beta-glucan rich barley flour on glucose and lipid metabolism in the ileum, liver, and adipose tissues of high-fat diet induced-obesity model male mice analyzed by DNA microarray. Nutrients, 12.","DOI":"10.3390\/nu12113546"},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Miyamoto, J., Watanabe, K., Taira, S., Kasubuchi, M., Li, X., Irie, J., Itoh, H., and Kimura, I. (2018). Barley beta-glucan improves metabolic condition via short-chain fatty acids produced by gut microbial fermentation in high fat diet fed mice. PLoS ONE, 13.","DOI":"10.1371\/journal.pone.0196579"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"162","DOI":"10.1016\/j.nutres.2014.12.006","article-title":"Wholegrain barley beta-glucan fermentation does not improve glucose tolerance in rats fed a high-fat diet","volume":"35","author":"Belobrajdic","year":"2015","journal-title":"Nutr. Res."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"724","DOI":"10.1093\/jaoac\/78.3.724","article-title":"Determination of soluble and insoluble dietary fiber in psyllium containing cereal products","volume":"78","author":"Lee","year":"1995","journal-title":"J. AOAC Int."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1002\/jsfa.2740550215","article-title":"Measurement of (1\u20133), (1\u20134)-\u03b2-D-glucan in barley and oats: A streamlined enzymic procedure","volume":"55","author":"McCleary","year":"1991","journal-title":"J. Sci. Food Agric."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"232","DOI":"10.1038\/nature12331","article-title":"Treg induction by a rationally selected mixture of Clostridia strains from the human microbiota","volume":"500","author":"Atarashi","year":"2013","journal-title":"Nature"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"485","DOI":"10.1055\/s-2007-980008","article-title":"Plasma and intestinal concentrations of GIP and GLP-1 (7-36) amide during suckling and after weaning in pigs","volume":"27","author":"Knapper","year":"1995","journal-title":"Horm. Metab. Res."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"2173","DOI":"10.1017\/S0007114513001682","article-title":"Molecular weight of barley beta-glucan influences energy expenditure, gastric emptying and glycaemic response in human subjects","volume":"110","author":"Thondre","year":"2013","journal-title":"Br. J. Nutr."},{"key":"ref_33","first-page":"851362","article-title":"Beta glucan: Health benefits in obesity and metabolic syndrome","volume":"2012","author":"Cuda","year":"2012","journal-title":"J. Nutr. Metab."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"1056","DOI":"10.1002\/cche.10328","article-title":"Effects of barley \u03b2-glucan with various molecular weights partially hydrolyzed by endogenous \u03b2-glucanase on glucose tolerance and lipid metabolism in mice","volume":"97","author":"Mio","year":"2020","journal-title":"Cereal Chem."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"645","DOI":"10.1093\/ajcn\/87.3.645","article-title":"Effect of cereal test breakfasts differing in glycemic index and content of indigestible carbohydrates on daylong glucose tolerance in healthy subjects","volume":"87","author":"Nilsson","year":"2008","journal-title":"Am. J. Clin. Nutr."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"110637","DOI":"10.1016\/j.nut.2019.110637","article-title":"Consumption of a meal containing refined barley flour bread is associated with a lower postprandial blood glucose concentration after a second meal compared with one containing refined wheat flour bread in healthy Japanese, A randomized control trial","volume":"72","author":"Matsuoka","year":"2020","journal-title":"Nutrition"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"2534S","DOI":"10.1093\/jn\/137.11.2534S","article-title":"Glucagon-like peptide-1 and energy homeostasis","volume":"137","author":"Burcelin","year":"2007","journal-title":"J. Nutr."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"18","DOI":"10.1016\/j.mce.2007.06.003","article-title":"GLUT2 and the incretin receptors are involved in glucoseinduced incretin secretion","volume":"276","author":"Cani","year":"2007","journal-title":"Mol. Cell. Endocrinol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"4768","DOI":"10.1210\/en.2008-0180","article-title":"Brain glucagon-like peptide 1 signaling controls the onset of high-fat diet-induced insulin resistance and reduces energy expenditure","volume":"149","author":"Knauf","year":"2008","journal-title":"Endocrinology"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"2603","DOI":"10.2337\/db07-1788","article-title":"Role of central nervous system glucagon-like Peptide-1 receptors in enteric glucose sensing","volume":"57","author":"Knauf","year":"2008","journal-title":"Diabetes"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"1099","DOI":"10.1093\/jn\/128.7.1099","article-title":"Insulin, glucagon-like peptide 1, glucose-dependent insulinotropic polypeptide and insulin-like growth factor I as putative mediators of the hypolipidemic effect of oligofructose in rats","volume":"128","author":"Kok","year":"1998","journal-title":"J. Nutr."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"1523","DOI":"10.1038\/oby.2006.176","article-title":"Effects of resistant starch, a non-digestible fermentable fiber, on reducing body fat","volume":"14","author":"Keenan","year":"2006","journal-title":"Obesity"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"683","DOI":"10.1038\/oby.2006.77","article-title":"Peptide YY and proglucagon mRNA expression patterns and regulation in the gut","volume":"14","author":"Zhou","year":"2006","journal-title":"Obesity"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1136\/gut.28.2.171","article-title":"Effects of an elemental diet, inert bulk and different types of dietary fibre on the response of the intestinal epithelium to refeeding in the rat and relationship to plasma gastrin, enteroglucagon, and PYY concentrations","volume":"28","author":"Goodlad","year":"1987","journal-title":"Gut"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"221","DOI":"10.1136\/gut.28.Suppl.221","article-title":"Proliferative effects of \u2018fibre\u2019 on the intestinal epithelium epithelium: Relationship to gastrin, enteroglucagon and PYY","volume":"28","author":"Goodlad","year":"1987","journal-title":"Gut"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"117","DOI":"10.1146\/annurev.nutr.18.1.117","article-title":"Dietary fructans","volume":"18","author":"Roberfroid","year":"1998","journal-title":"Annu. Rev. Nutr."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"353","DOI":"10.1007\/s00441-005-0140-x","article-title":"Short-chain fatty acid receptor, GPR43, is expressed by enteroendocrine cells and mucosal mast cells in rat intestine","volume":"324","author":"Karaki","year":"2006","journal-title":"Cell Tissue Res."}],"container-title":["Nutrients"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2072-6643\/13\/2\/527\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T05:20:22Z","timestamp":1760160022000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2072-6643\/13\/2\/527"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,2,6]]},"references-count":47,"journal-issue":{"issue":"2","published-online":{"date-parts":[[2021,2]]}},"alternative-id":["nu13020527"],"URL":"https:\/\/doi.org\/10.3390\/nu13020527","relation":{},"ISSN":["2072-6643"],"issn-type":[{"value":"2072-6643","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,2,6]]}}}