{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,5]],"date-time":"2026-11-05T19:48:14Z","timestamp":1793908094637,"version":"4.8.0"},"reference-count":112,"publisher":"MDPI AG","issue":"21","license":[{"start":{"date-parts":[[2025,10,26]],"date-time":"2025-10-26T00:00:00Z","timestamp":1761436800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["22166026"],"award-info":[{"award-number":["22166026"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32572578"],"award-info":[{"award-number":["32572578"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100005230","name":"Natural Science Foundation project of Chongqing Municipality","doi-asserted-by":"crossref","award":["CSTB2022NSCQ-MSX1634"],"award-info":[{"award-number":["CSTB2022NSCQ-MSX1634"]}],"id":[{"id":"10.13039\/501100005230","id-type":"DOI","asserted-by":"crossref"}]},{"name":"Cross-Innovation Open Project of Food Flavor and Health, Beijing Technology & Business University","award":["FFHCI-2025051"],"award-info":[{"award-number":["FFHCI-2025051"]}],"id":[{"id":"https:\/\/ror.org\/013e0zm98","id-type":"ROR","asserted-by":"crossref"}]},{"name":"Academic and Technical Leaders of Major Academic Disciplines of Jiangxi Province","award":["20243BCE51113"],"award-info":[{"award-number":["20243BCE51113"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Foods"],"abstract":"<jats:p>Butyrate is a key short-chain fatty acid (SCFA) essential for maintaining colon health, immune homeostasis, and metabolic balance. Its synthesis primarily depends on the fermentation of dietary substrates by the gut microbiota. In a healthy state, carbohydrate fermentation sustains the stability of the gut microbiota; however, in chronic diseases, the diversity of the microbiota decreases, and the metabolic pathway shifts from carbohydrate fermentation to protein fermentation, thereby inhibiting butyrate production. Polysaccharides and proteins play key roles in regulating butyrate synthesis. As fermentable carbon sources, polysaccharides promote the proliferation of probiotics, lower colonic pH, and inhibit anaerobic fermentation of proteins. However, excessive protein fermentation produces branched-chain fatty acid (BCFA), ammonia, phenols, and other metabolites that inhibit butyrate production. The structural characteristics of polysaccharides and the digestibility of proteins can determine the substrate selection of gut microbiota. This review systematically elucidates the biosynthetic pathways of butyrate in the gut and the host metabolic signaling pathways in which it participates, focusing on the interactions between dietary proteins and polysaccharides and their key regulatory mechanisms affecting butyrate production by the gut microbiota.<\/jats:p>","DOI":"10.3390\/foods14213649","type":"journal-article","created":{"date-parts":[[2025,10,29]],"date-time":"2025-10-29T02:23:28Z","timestamp":1761704608000},"page":"3649","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":28,"title":["A Review of Nutritional Regulation of Intestinal Butyrate Synthesis: Interactions Between Dietary Polysaccharides and Proteins"],"prefix":"10.3390","volume":"14","author":[{"given":"Meiyu","family":"Yuan","sequence":"first","affiliation":[{"name":"State Key Laboratory of Food Science and Resources, Engineering Research Center for Biomass Conversion, Ministry of Education, Nanchang University, Nanchang 330047, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kaili","family":"Gao","sequence":"additional","affiliation":[{"name":"College of Biological and Environmental Engineering, Jingdezhen University, Jingdezhen 334000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kaitao","family":"Peng","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Food Science and Resources, Engineering Research Center for Biomass Conversion, Ministry of Education, Nanchang University, Nanchang 330047, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1896-1171","authenticated-orcid":false,"given":"Shuang","family":"Bi","sequence":"additional","affiliation":[{"name":"School of Food and Health, Beijing Technology & Business University, Beijing 100048, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xian","family":"Cui","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Food Science and Resources, Engineering Research Center for Biomass Conversion, Ministry of Education, Nanchang University, Nanchang 330047, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0003-0435-2572","authenticated-orcid":false,"given":"Yuhuan","family":"Liu","sequence":"additional","affiliation":[{"name":"Chongqing Research Institute of Nanchang University, Chongqing 402660, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2025,10,26]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"100139","DOI":"10.1016\/j.medmic.2025.100139","article-title":"Short-chain fatty acids (SCFAs) in gut health: Implications for drug metabolism and therapeutics","volume":"25","author":"Sankarganesh","year":"2025","journal-title":"Med. Microecol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"e26991","DOI":"10.1016\/j.heliyon.2024.e26991","article-title":"Short-chain fatty acids in nonalcoholic fatty liver disease: New prospects for short-chain fatty acids as therapeutic targets","volume":"10","author":"Li","year":"2024","journal-title":"Heliyon"},{"key":"ref_3","doi-asserted-by":"crossref","unstructured":"Deleu, S., Machiels, K., Raes, J., Verbeke, K., and Vermeire, S. (2021). Short chain fatty acids and its producing organisms: An overlooked therapy for IBD?. EBioMedicine, 66.","DOI":"10.1016\/j.ebiom.2021.103293"},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"Matheus, V.A., Oliveira, R.B., Maschio, D.A., Tada, S.F.S., Soares, G.M., Mousovich-Neto, F., Costa, R.G., Mori, M.A., Barbosa, H.C.L., and Collares-Buzato, C.B. (2023). Butyrate restores the fat\/lean mass ratio balance and energy metabolism and reinforces the tight junction-mediated intestinal epithelial barrier in prediabetic mice independently of its anti-inflammatory and epigenetic actions. J. Nutr. Biochem., 120.","DOI":"10.1016\/j.jnutbio.2023.109409"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"168080","DOI":"10.1016\/j.cej.2025.168080","article-title":"Induction of degradation of fucoidan oligosaccharides via low-temperature plasma treatment for enhanced structural characteristics and bioactivities","volume":"523","author":"Yuan","year":"2025","journal-title":"Chem. Eng. J."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"524","DOI":"10.1016\/j.jconrel.2025.01.023","article-title":"Mechanism exploration of intestinal mucus penetration of nano-Se: Regulated by polysaccharides with different functional groups and molecular weights","volume":"379","author":"Dai","year":"2025","journal-title":"J. Control. Release"},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Rong, X., Zhu, L., and Shu, Q. (2025). Synergistic gut microbiome-mediated degradation of Astragalus membranaceus polysaccharides and Codonopsis pilosula polysaccharides into butyric acid: A metatranscriptomic analysis. Microbiol. Spectr., 13.","DOI":"10.1128\/spectrum.03039-24"},{"key":"ref_8","doi-asserted-by":"crossref","unstructured":"Rong, X., and Shu, Q. (2024). Modulating butyric acid-producing bacterial community abundance and structure in the intestine of immunocompromised mice with neutral polysaccharides extracted from Codonopsis pilosula. Int. J. Biol. Macromol., 278.","DOI":"10.1016\/j.ijbiomac.2024.134959"},{"key":"ref_9","doi-asserted-by":"crossref","unstructured":"Huang, Z., Hu, M., Peng, X., Wang, R., Song, X., and Yin, J. (2023). The protective effect of small black soybean (Vigna mungo L.) polysaccharide on acetic acid-induced gastric ulcer in SD rats and its impact on gut microbiota and metabolites. Food Biosci., 56.","DOI":"10.1016\/j.fbio.2023.103187"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1016\/j.ijbiomac.2022.09.011","article-title":"Synthesis, characterization, and evaluation of microwave-assisted fabricated selenylation Astragalus polysaccharides","volume":"221","author":"Yue","year":"2022","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"S130","DOI":"10.1080\/10408398.2018.1542587","article-title":"Nondigestible carbohydrates, butyrate, and butyrate-producing bacteria","volume":"59","author":"Fu","year":"2019","journal-title":"Crit. Rev. Food Sci. Nutr."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"124","DOI":"10.1016\/j.clnu.2024.09.045","article-title":"Soy protein \u03b2-conglycinin ameliorates pressure overload-induced heart failure by increasing short-chain fatty acid (SCFA)-producing gut microbiota and intestinal SCFAs","volume":"43","author":"Furukawa","year":"2024","journal-title":"Clin. Nutr."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"117123","DOI":"10.1016\/j.foodres.2025.117123","article-title":"Effects of pressure cooker treatment-induced protein oxidation of cereals on human gut microbiota using an in vitro fermentation model","volume":"220","author":"Chu","year":"2025","journal-title":"Food Res. Int."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"192","DOI":"10.1080\/1745039X.2017.1291202","article-title":"Interactive effects of protein and carbohydrates on production of microbial metabolites in the large intestine of growing pigs","volume":"71","author":"Taciak","year":"2017","journal-title":"Arch. Anim. Nutr."},{"key":"ref_15","doi-asserted-by":"crossref","unstructured":"Mirzaei, R., Dehkhodaie, E., Bouzari, B., Rahimi, M., Gholestani, A., Hosseini-Fard, S.R., Keyvani, H., Teimoori, A., and Karampoor, S. (2022). Dual role of microbiota-derived short-chain fatty acids on host and pathogen. Biomed. Pharmacother., 145.","DOI":"10.1016\/j.biopha.2021.112352"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"61","DOI":"10.1016\/j.clnu.2022.10.024","article-title":"Butyrate\u2019s role in human health and the current progress towards its clinical application to treat gastrointestinal disease","volume":"42","author":"Hodgkinson","year":"2023","journal-title":"Clin. Nutr."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"178","DOI":"10.1016\/j.tim.2023.07.014","article-title":"Beyond butyrate: Microbial fiber metabolism supporting colonic epithelial homeostasis","volume":"32","author":"Fagundes","year":"2024","journal-title":"Trends Microbiol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1455","DOI":"10.1016\/j.ajpath.2023.06.007","article-title":"The Role of Gut Microbiome-Derived Short-Chain Fatty Acid Butyrate in Hepatobiliary Diseases","volume":"193","author":"Pant","year":"2023","journal-title":"Am. J. Surg. Pathol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1111\/1462-2920.13589","article-title":"Formation of propionate and butyrate by the human colonic microbiota","volume":"19","author":"Louis","year":"2017","journal-title":"Environ. Microbiol."},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Salvi, P.S., and Cowles, R.A. (2021). Butyrate and the Intestinal Epithelium: Modulation of Proliferation and Inflammation in Homeostasis and Disease. Cells, 10.","DOI":"10.3390\/cells10071775"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"113944","DOI":"10.1016\/j.yexcr.2024.113944","article-title":"Effect of sodium butyrate treatment at the basolateral membranes on the tight junction barrier function via a monocarboxylate transporter in goat mammary epithelial cells","volume":"436","author":"Tsugami","year":"2024","journal-title":"Exp. Cell Res."},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Ota, S., Sakuraba, H., Hiraga, H., Yoshida, S., Satake, M., Akemoto, Y., Tanaka, N., Watanabe, R., Takato, M., and Murai, Y. (2020). Cyclosporine protects from intestinal epithelial injury by modulating butyrate uptake via upregulation of membrane monocarboxylate transporter 1 levels. Biochem. Biophys. Rep., 24.","DOI":"10.1016\/j.bbrep.2020.100811"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1007\/s00232-011-9340-3","article-title":"Characterization of Butyrate Uptake by Nontransformed Intestinal Epithelial Cell Lines","volume":"240","author":"Martel","year":"2011","journal-title":"J. Membr. Biol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"13293","DOI":"10.1074\/jbc.C400059200","article-title":"Functional identification of SLC5A8, a tumor suppressor down-regulated in colon cancer, as a Na(+)-coupled transporter for short-chain fatty acids","volume":"279","author":"Miyauchi","year":"2004","journal-title":"J. Biol. Chem."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1016\/j.pbj.2016.04.004","article-title":"Regulation of colonic epithelial butyrate transport: Focus on colorectal cancer","volume":"1","author":"Martel","year":"2016","journal-title":"Porto Biomed. J."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"733","DOI":"10.1053\/j.gastro.2024.05.014","article-title":"Gut Microbiota-Derived Butyrate Induces Epigenetic and Metabolic Reprogramming in Myeloid-Derived Suppressor Cells to Alleviate Primary Biliary Cholangitis","volume":"167","author":"Wang","year":"2024","journal-title":"Gastroenterology"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"156307","DOI":"10.1016\/j.cyto.2023.156307","article-title":"Taurocholate uptake by Caco-2 cells is inhibited by pro-inflammatory cytokines and butyrate","volume":"169","author":"Couto","year":"2023","journal-title":"Cytokine"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"323","DOI":"10.1038\/s41422-023-00914-z","article-title":"Molecular recognition and activation mechanism of short-chain fatty acid receptors FFAR2\/3","volume":"34","author":"Li","year":"2024","journal-title":"Cell Res."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1093\/advances\/nmx009","article-title":"Butyrate: A Double-Edged Sword for Health?","volume":"9","author":"Liu","year":"2018","journal-title":"Adv. Nutr."},{"key":"ref_30","doi-asserted-by":"crossref","unstructured":"Lee, D.-H., Kim, M.-T., and Han, J.-H. (2024). GPR41 and GPR43: From development to metabolic regulation. Biomed. Pharmacother., 175.","DOI":"10.1016\/j.biopha.2024.116735"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"157312","DOI":"10.1016\/j.phymed.2025.157312","article-title":"Zhenqi Fuzheng Granule targets the SCFAs-GPR109A axis to enhance PD-1 antibody efficacy via immunometabolic remodeling in colorectal cancer","volume":"148","author":"Guo","year":"2025","journal-title":"Phytomedicine"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"2392","DOI":"10.1182\/blood.2021010719","article-title":"Alloreactive T cells deficient of the short-chain fatty acid receptor GPR109A induce less graft-versus-host disease","volume":"139","author":"Docampo","year":"2022","journal-title":"Blood"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"2029","DOI":"10.1093\/jimmun\/vkaf069","article-title":"Butyrate receptor HCAR2\/GPR109A controls imiquimod-induced psoriasis-like skin inflammation","volume":"214","author":"Leko","year":"2025","journal-title":"J. Immunol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"14638","DOI":"10.1021\/acs.jafc.3c04417","article-title":"Butyrate Protects against \u03b3-d-Glutamyl-meso-diaminopimelic Acid-Induced Inflammatory Response and Tight Junction Disruption through Histone Deacetylase 3 Inhibition in Bovine Mammary Epithelial Cells","volume":"71","author":"Wang","year":"2023","journal-title":"J. Agric. Food Chem."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"124097","DOI":"10.1016\/j.carbpol.2025.124097","article-title":"Ideal dietary fiber model: Personalized gut microbiota modulation based on structure-function relationships","volume":"368","author":"Wang","year":"2025","journal-title":"Carbohydr. Polym."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"213","DOI":"10.1016\/j.tifs.2016.08.011","article-title":"Gut microbiota role in dietary protein metabolism and health-related outcomes: The two sides of the coin","volume":"57","author":"Portune","year":"2016","journal-title":"Trends Food Sci. Technol."},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"De Angelis, M., Ferrocino, I., Calabrese, F.M., De Filippis, F., Cavallo, N., Siragusa, S., Rampelli, S., Di Cagno, R., Rantsiou, K., and Vannini, L. (2020). Diet influences the functions of the human intestinal microbiome. Sci. Rep., 10.","DOI":"10.1038\/s41598-020-61192-y"},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Alvarez, B., Canil, O.F., Low, K.E., Hettle, A.G., Abbott, D.W., and Boraston, A.B. (2025). Analysis of chondroitin degradation by components of a Bacteroides caccae polysaccharide utilization locus. J. Biol. Chem., 301.","DOI":"10.1016\/j.jbc.2025.110354"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"106835","DOI":"10.1016\/j.jff.2025.106835","article-title":"Structural characteristics of oligosaccharides obtained from Undaria pinnatifida polysaccharides and their alleviation of colitis by modulating macrophage polarization and gut microbiota","volume":"128","author":"Yu","year":"2025","journal-title":"J. Funct. Foods"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"104606","DOI":"10.1016\/j.tifs.2024.104606","article-title":"The fate of dietary polysaccharides in the digestive tract","volume":"150","author":"Yang","year":"2024","journal-title":"Trends Food Sci. Technol."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.tifs.2019.03.005","article-title":"In vitro colonic fermentation of dietary fibers: Fermentation rate, short-chain fatty acid production and changes in microbiota","volume":"88","author":"Wang","year":"2019","journal-title":"Trends Food Sci. Technol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"11245","DOI":"10.1021\/acs.jafc.2c03091","article-title":"Study on the Effect of Molecular Weight on the Gut Microbiota Fermentation Properties of Blackberry Polysaccharides In Vitro","volume":"70","author":"Li","year":"2022","journal-title":"J. Agric. Food Chem."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1089\/jmf.2005.8.113","article-title":"Comparison of different fibers for in vitro production of short chain fatty acids by intestinal microflora","volume":"8","author":"Pylkas","year":"2005","journal-title":"J. Med. Food"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"123209","DOI":"10.1016\/j.carbpol.2024.123209","article-title":"Correlation between the structures of natural polysaccharides and their properties in regulating gut microbiota: Current understanding and beyond","volume":"352","author":"Keung","year":"2025","journal-title":"Carbohydr. Polym."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"127179","DOI":"10.1016\/j.foodchem.2020.127179","article-title":"Dietary galactosyl and mannosyl carbohydrates: In-vitro assessment of prebiotic effects","volume":"329","author":"Wei","year":"2020","journal-title":"Food Chem."},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Harris, H.C., Edwards, C.A., and Morrison, D.J. (2017). Impact of Glycosidic Bond Configuration on Short Chain Fatty Acid Production from Model Fermentable Carbohydrates by the Human Gut Microbiota. Nutrients, 9.","DOI":"10.3390\/nu9010026"},{"key":"ref_47","doi-asserted-by":"crossref","unstructured":"Zou, Y., Xue, W., Lin, X., Lv, M., Luo, G., Dai, Y., Sun, H., Liu, S.-W., Sun, C.-H., and Hu, T. (2021). Butyribacter intestini gen. nov., sp. nov., a butyric acid-producing bacterium of the family Lachnospiraceae isolated from human faeces, and reclassification of Acetivibrio ethanolgignens as Acetanaerobacter ethanolgignens gen. nov., comb. nov. Syst. Appl. Microbiol., 44.","DOI":"10.1016\/j.syapm.2021.126201"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"158","DOI":"10.1016\/j.jff.2014.12.044","article-title":"Influences of structures of galactooligosaccharides and fructooligosaccharides on the fermentation in vitro by human intestinal microbiota","volume":"13","author":"Li","year":"2015","journal-title":"J. Funct. Foods"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"386","DOI":"10.1017\/S0029665121003694","article-title":"Dietary fibre complexity and its influence on functional groups of the human gut microbiota","volume":"80","author":"Louis","year":"2021","journal-title":"Proc. Nutr. Soc."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"122535","DOI":"10.1016\/j.carbpol.2024.122535","article-title":"Oat \u03b2-(1 \u2192 3, 1 \u2192 4)-d-glucan alleviates food allergy-induced colonic injury in mice by increasing Lachnospiraceae abundance and butyrate production","volume":"344","author":"Zhang","year":"2024","journal-title":"Carbohydr. Polym."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"705","DOI":"10.1079\/BJN20051531","article-title":"Short-chain fatty acid formation in the hindgut of rats fed oligosaccharides varying in monomeric composition, degree of polymerisation and solubility","volume":"94","author":"Nilsson","year":"2005","journal-title":"Br. J. Nutr."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"2525482","DOI":"10.1080\/19490976.2025.2525482","article-title":"Lactulose selectively stimulates members of the gut microbiota, as determined by multi-modal activity-based sorting","volume":"17","author":"Rasoulimehrabani","year":"2025","journal-title":"Gut Microbes"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"610","DOI":"10.1038\/ismej.2017.196","article-title":"Specific substrate-driven changes in human faecal microbiota composition contrast with functional redundancy in short-chain fatty acid production","volume":"12","author":"Reichardt","year":"2018","journal-title":"ISME J."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"124490","DOI":"10.1016\/j.carbpol.2025.124490","article-title":"Inulin with different degrees of polymerization adopts different strategies to mitigate radiation-induced intestinal injury: Focus on intestinal epithelial or T cell homeostasis","volume":"371","author":"Ji","year":"2025","journal-title":"Carbohydr. Polym."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"3186","DOI":"10.1039\/D3FO04187E","article-title":"Protein combined with certain dietary fibers increases butyrate production in gut microbiota fermentation","volume":"15","author":"Jackson","year":"2024","journal-title":"Food Funct."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"100195","DOI":"10.1016\/j.fochx.2021.100195","article-title":"What we know about protein gut metabolites: Implications and insights for human health and diseases","volume":"13","year":"2022","journal-title":"Food Chem. X"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1016\/j.jare.2020.03.003","article-title":"Hydrogen sulfide toxicity in the gut environment: Meta-analysis of sulfate-reducing and lactic acid bacteria in inflammatory processes","volume":"27","author":"Kushkevych","year":"2021","journal-title":"J. Adv. Res."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"370","DOI":"10.1016\/j.aninu.2024.08.005","article-title":"Dietary high lipid and high plant-protein affected growth performance, liver health, bile acid metabolism and gut microbiota in groupers","volume":"19","author":"Xu","year":"2024","journal-title":"Anim. Nutr."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"1509","DOI":"10.1001\/jamainternmed.2019.2806","article-title":"Association of Animal and Plant Protein Intake with All-Cause and Cause-Specific Mortality in a Japanese Cohort","volume":"179","author":"Budhathoki","year":"2019","journal-title":"Jama Intern. Med."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"1173","DOI":"10.1001\/jamainternmed.2020.2790","article-title":"Association Between Plant and Animal Protein Intake and Overall and Cause-Specific Mortality","volume":"180","author":"Huang","year":"2020","journal-title":"JAMA Intern. Med."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"1164","DOI":"10.1016\/j.ajcnut.2024.03.006","article-title":"Dietary intake of plant- and animal-derived protein and incident cardiovascular diseases: The pan-European EPIC-CVD case\u2013cohort study","volume":"119","author":"Zheng","year":"2024","journal-title":"Am. J. Clin. Nutr."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1139\/apnm-2024-0294","article-title":"Effects of a paternal diet high in animal protein (casein) versus plant protein (pea protein with added methionine) on offspring metabolic and gut microbiota outcomes in rats","volume":"50","author":"Patterson","year":"2025","journal-title":"Appl. Physiol. Nutr. Metab."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"106911","DOI":"10.1016\/j.foodhyd.2021.106911","article-title":"The role of dietary proteins and carbohydrates in gut microbiome composition and activity: A review","volume":"120","author":"Peled","year":"2021","journal-title":"Food Hydrocoll."},{"key":"ref_64","doi-asserted-by":"crossref","unstructured":"Yang, Y., Wang, Z., Wang, R., Sui, X., Qi, B., Han, F., Li, Y., and Jiang, L. (2016). Secondary Structure and Subunit Composition of Soy Protein In Vitro Digested by Pepsin and Its Relation with Digestibility. BioMed Res. Int., 2016.","DOI":"10.1155\/2016\/5498639"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"105136","DOI":"10.1016\/j.ultsonch.2020.105136","article-title":"Application of ultrasound-assisted physical mixing treatment improves in vitro protein digestibility of rapeseed napin","volume":"67","author":"Pan","year":"2020","journal-title":"Ultrason. Sonochem."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"140950","DOI":"10.1016\/j.foodchem.2024.140950","article-title":"The structure\u2013function relationships and techno-functions of \u03b2-conglycinin","volume":"462","author":"Ashaolu","year":"2025","journal-title":"Food Chem."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1080\/10408398.2025.2452362","article-title":"Influence of dietary protein and fiber intake interactions on the human gut microbiota composition and function: A systematic review and network meta-analysis of randomized controlled trials","volume":"16","author":"Mak","year":"2025","journal-title":"Crit. Rev. Food Sci. Nutr."},{"key":"ref_68","doi-asserted-by":"crossref","unstructured":"Wang, L., Fan, J.-H., Yang, Y., Qiu, H.-W., Li, H.-H., Kang, W.-Y., and Lv, G.-P. (2025). Gut microbiota modulation by structurally diverse foodborne polysaccharides: In vitro fermentation and prebiotic activity evaluation. Food Biosci., 73.","DOI":"10.1016\/j.fbio.2025.107660"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"123772","DOI":"10.1016\/j.carbpol.2025.123772","article-title":"Human gut Bifidobacterium longum subsp. suillum is enriched in vitro by a pectic polysaccharide isolated from the flowers of Lilium lancifolium","volume":"364","author":"Li","year":"2025","journal-title":"Carbohydr. Polym."},{"key":"ref_70","doi-asserted-by":"crossref","unstructured":"Yu, J., An, N., Cui, Y., and Zhang, L. (2025). Structural characterization and immunomodulatory effects of Bifidobacterium bifidum biofilms enriched with extracellular polysaccharides and surface proteins. Int. J. Biol. Macromol., 328.","DOI":"10.1016\/j.ijbiomac.2025.147604"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"S-616","DOI":"10.1016\/S0016-5085(10)62839-9","article-title":"T2026 A Dietary Intervention With Arabinoxylan Oligosaccharides Reduces Colonic Protein Fermentation in Healthy Subjects: Results From Faecal Metabolite Fingerprint Analysis","volume":"138","author":"Hamer","year":"2010","journal-title":"Gastroenterology"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"1329","DOI":"10.1016\/j.ijbiomac.2020.11.087","article-title":"The role of mucin and oligosaccharides via cross-feeding activities by Bifidobacterium: A review","volume":"167","author":"Luo","year":"2021","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"237","DOI":"10.1016\/j.tifs.2022.04.010","article-title":"A critical review on interplay between dietary fibers and gut microbiota","volume":"124","author":"Ye","year":"2022","journal-title":"Trends Food Sci. Tech."},{"key":"ref_74","doi-asserted-by":"crossref","unstructured":"La Rosa, S.L., Ostrowski, M.P., de Le\u00f3n, A.V.-P., McKee, L.S., Larsbrink, J., Eijsink, V.G., Lowe, E.C., Martens, E.C., and Pope, P.B. (2022). Glycan processing in gut microbiomes. Curr. Opin. Microbiol., 67.","DOI":"10.1016\/j.mib.2022.102143"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"111942","DOI":"10.1016\/j.foodres.2022.111942","article-title":"Co-culture fermentations suggest cross-feeding among Bacteroides ovatus DSMZ 1896, Lactiplantibacillus plantarum WCFS1 and Bifidobacterium adolescentis DSMZ 20083 for utilizing dietary galactomannans","volume":"162","author":"Mary","year":"2022","journal-title":"Food Res. Int."},{"key":"ref_76","doi-asserted-by":"crossref","unstructured":"Gurry, T., Nguyen, L.T.T., Yu, X.Q., and Alm, E.J. (2021). Functional heterogeneity in the fermentation capabilities of the healthy human gut microbiota. PLoS ONE, 16.","DOI":"10.1371\/journal.pone.0254004"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"299","DOI":"10.1016\/j.scib.2023.11.060","article-title":"Glycan utilization properties govern the cross-feeding network in gut microbiota: A focus on Bifidobacteria","volume":"69","author":"Yu","year":"2024","journal-title":"Sci. Bull."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1016\/j.ijfoodmicro.2011.03.003","article-title":"Cross-feeding between bifidobacteria and butyrate-producing colon bacteria explains bifdobacterial competitiveness, butyrate production, and gas production","volume":"149","author":"Leroy","year":"2011","journal-title":"Int. J. Food Microbiol."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"76","DOI":"10.1016\/j.ijfoodmicro.2016.05.015","article-title":"Bifidobacterial inulin-type fructan degradation capacity determines cross-feeding interactions between bifidobacteria and Faecalibacterium prausnitzii","volume":"231","author":"Moens","year":"2016","journal-title":"Int. J. Food Microbiol."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"485","DOI":"10.1016\/j.chom.2023.03.016","article-title":"Cross-feeding in the gut microbiome: Ecology and mechanisms","volume":"31","author":"Culp","year":"2023","journal-title":"Cell Host Microbe"},{"key":"ref_81","doi-asserted-by":"crossref","unstructured":"Lin, L., Du, R., Wang, Y., Wu, Q., and Xu, Y. (2022). Regulation of auxotrophic lactobacilli growth by amino acid cross-feeding interaction. Int. J. Food Microbiol., 377.","DOI":"10.1016\/j.ijfoodmicro.2022.109769"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"138","DOI":"10.1016\/j.livsci.2010.06.046","article-title":"Effects of two protein and fibre sources on SCFA concentration in pig large intestine","volume":"133","author":"Taciak","year":"2010","journal-title":"Livest. Sci."},{"key":"ref_83","doi-asserted-by":"crossref","unstructured":"Wu, S., Bhat, Z.F., Gounder, R.S., Ahmed, I.A., Al-Juhaimi, F.Y., Ding, Y., and Bekhit, A.E.A. (2022). Effect of Dietary Protein and Processing on Gut Microbiota\u2014A Systematic Review. Nutrients, 14.","DOI":"10.3390\/nu14030453"},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"124228","DOI":"10.1016\/j.carbpol.2025.124228","article-title":"Release, digestion and fermentation properties of polyphenols bound to sea buckthorn polysaccharides: Mechanistic explorations from integrated in vitro and in vivo studies","volume":"368","author":"Wang","year":"2025","journal-title":"Carbohydr. Polym."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"66","DOI":"10.1016\/j.copbio.2019.10.008","article-title":"Resistant starch: Impact on the gut microbiome and health","volume":"61","author":"DeMartino","year":"2020","journal-title":"Curr. Opin. Biotechnol."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"46","DOI":"10.1038\/s41387-024-00305-2","article-title":"Dietary resistant starch enhances immune health of the kidney in diabetes via promoting microbially-derived metabolites and dampening neutrophil recruitment","volume":"14","author":"Snelson","year":"2024","journal-title":"Nutr. Diabetes"},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"2169","DOI":"10.1093\/carcin\/bgn173","article-title":"Butyrylated starch protects colonocyte DNA against dietary protein-induced damage in rats","volume":"29","author":"Bajka","year":"2008","journal-title":"Carcinog."},{"key":"ref_88","doi-asserted-by":"crossref","unstructured":"Wang, J., Ren, M., Fu, L., Liu, L., Zhang, X., Sun, Y., and Liu, L. (2024). Safflower Dietary Fiber Alleviates Functional Constipation in Rats via Regulating Gut Microbiota and Metabolism. Food Biosci., 62.","DOI":"10.1016\/j.fbio.2024.105218"},{"key":"ref_89","doi-asserted-by":"crossref","unstructured":"Mio, K., Ogawa, R., Tadenuma, N., and Aoe, S. (2022). Arabinoxylan as well as \u03b2-glucan in barley promotes GLP-1 secretion by increasing short-chain fatty acids production. Biochem. Biophys. Rep., 32.","DOI":"10.1016\/j.bbrep.2022.101343"},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"273","DOI":"10.1016\/j.psj.2020.09.089","article-title":"Microbiota populations and short-chain fatty acids production in cecum of immunosuppressed broilers consuming diets containing \u03b3-irradiated Astragalus polysaccharides","volume":"100","author":"Liu","year":"2021","journal-title":"Poult. Sci."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"4155","DOI":"10.7150\/thno.54476","article-title":"Pectin supplement significantly enhanced the anti-PD-1 efficacy in tumor-bearing mice humanized with gut microbiota from patients with colorectal cancer","volume":"11","author":"Zhang","year":"2021","journal-title":"Theranostics"},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"106062","DOI":"10.1016\/j.intimp.2019.106062","article-title":"Inulin alleviates inflammation of alcoholic liver disease via SCFAs-inducing suppression of M1 and facilitation of M2 macrophages in mice","volume":"78","author":"Wang","year":"2020","journal-title":"Int. Immunopharmacol."},{"key":"ref_93","doi-asserted-by":"crossref","unstructured":"Lou, Y., Wen, X., Song, S., Zeng, Y., Huang, L., Xie, Z., Shao, T., and Wen, C. (2024). Dietary pectin and inulin: A promising adjuvant supplement for collagen-induced arthritis through gut microbiome restoration and CD4+ T cell reconstitution. J. Nutr. Biochem., 133.","DOI":"10.1016\/j.jnutbio.2024.109699"},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"43","DOI":"10.1016\/j.jff.2018.10.020","article-title":"Cumulative effect of yeast extract and fructooligosaccharide supplementation on composition and metabolic activity of elderly colonic microbiota in vitro","volume":"52","author":"Doo","year":"2019","journal-title":"J. Funct. Foods"},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"110677","DOI":"10.1016\/j.foodres.2021.110677","article-title":"Enzymatically synthesised fructooligosaccharides from sugarcane syrup modulate the composition and short-chain fatty acid production of the human intestinal microbiota","volume":"149","author":"Rahim","year":"2021","journal-title":"Food Res. Int."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"1199","DOI":"10.1017\/S0007114516003068","article-title":"Partially hydrolysed guar gum ameliorates murine intestinal inflammation in association with modulating luminal microbiota and SCFA","volume":"116","author":"Takagi","year":"2016","journal-title":"Br. J. Nutr."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"427","DOI":"10.1021\/acs.jnatprod.0c01125","article-title":"A Purified Glucomannan Oligosaccharide from Amorphophallus konjac Improves Colonic Mucosal Barrier Function via Enhancing Butyrate Production and Histone Protein H3 and H4 Acetylation","volume":"84","author":"Cui","year":"2021","journal-title":"J. Nat. Prod."},{"key":"ref_98","doi-asserted-by":"crossref","unstructured":"Teng, Y., Li, J., Guo, J., Yan, C., Wang, A., and Xia, X. (2024). Alginate oligosaccharide improves 5-fluorouracil-induced intestinal mucositis by enhancing intestinal barrier and modulating intestinal levels of butyrate and isovalerate. Int. J. Biol. Macromol., 276.","DOI":"10.1016\/j.ijbiomac.2024.133699"},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"330","DOI":"10.1016\/j.bbi.2021.04.005","article-title":"Mannan oligosaccharide attenuates cognitive and behavioral disorders in the 5xFAD Alzheimer\u2019s disease mouse model via regulating the gut microbiota-brain axis","volume":"95","author":"Liu","year":"2021","journal-title":"Brain, Behav. Immun."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"749","DOI":"10.1016\/j.nutres.2014.08.006","article-title":"Long-term dietary pattern of fecal donor correlates with butyrate production and markers of protein fermentation during in vitro fecal fermentation","volume":"34","author":"Yang","year":"2014","journal-title":"Nutr. Res."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"2042","DOI":"10.1017\/S1751731120001044","article-title":"Effect of early antibiotic intervention on specific bacterial communities and immune parameters in the small intestine of growing pigs fed different protein level diets","volume":"14","author":"Zhang","year":"2020","journal-title":"Animal"},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"951","DOI":"10.1139\/cjm-2017-0242","article-title":"Low dietary protein and high carbohydrate infant formula affects the microbial ecology of the large intestine in neonatal rats","volume":"63","author":"Fan","year":"2017","journal-title":"Can. J. Microbiol."},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"102773","DOI":"10.1016\/j.psj.2023.102773","article-title":"The effects of arginine and branched-chain amino acid supplementation to reduced-protein diet on intestinal health, cecal short-chain fatty acid profiles, and immune response in broiler chickens challenged with Eimeria spp.","volume":"102","author":"Liu","year":"2023","journal-title":"Poult. Sci."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"1156","DOI":"10.1093\/jn\/128.7.1156","article-title":"Resistant Proteins Alter Cecal Short-Chain Fatty Acid Profiles in Rats Fed High Amylose Cornstarch","volume":"128","author":"Morita","year":"1998","journal-title":"J. Nutr."},{"key":"ref_105","doi-asserted-by":"crossref","unstructured":"Wu, L., An, R., Lan, T., Tang, Z., Xu, Y., Peng, X., Pang, J., Sun, W., Shi, B., and Tang, Q. (2024). Isocaloric diets with varying protein levels affected energy metabolism in young adult Sprague-Dawley rats via modifying the gut microbes: A lipid imbalance was brought on by a diet with a particularly high protein content. J. Nutr. Biochem., 124.","DOI":"10.1016\/j.jnutbio.2023.109534"},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"3569","DOI":"10.2527\/jas.2008-1241","article-title":"Effect of dietary level of protein and fiber on the productive performance and health status of piglets","volume":"87","author":"Hermes","year":"2009","journal-title":"J. Anim. Sci."},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"1062","DOI":"10.3945\/ajcn.110.002188","article-title":"High-protein, reduced-carbohydrate weight-loss diets promote metabolite profiles likely to be detrimental to colonic health","volume":"93","author":"Russell","year":"2011","journal-title":"Am. J. Clin. Nutr."},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"1005","DOI":"10.3945\/ajcn.117.158816","article-title":"Quantity and source of dietary protein influence metabolite production by gut microbiota and rectal mucosa gene expression: A randomized, parallel, double-blind trial in overweight humans","volume":"106","author":"Beaumont","year":"2017","journal-title":"Am. J. Clin. Nutr."},{"key":"ref_109","doi-asserted-by":"crossref","unstructured":"Tamura, K., Sasaki, H., Shiga, K., Miyakawa, H., and Shibata, S. (2019). The Timing Effects of Soy Protein Intake on Mice Gut Microbiota. Nutrients, 12.","DOI":"10.3390\/nu12010087"},{"key":"ref_110","doi-asserted-by":"crossref","unstructured":"Vidal-Lletj\u00f3s, S., Andriamihaja, M., Blais, A., Grauso, M., Lepage, P., Davila, A.-M., Viel, R., Gaudichon, C., Leclerc, M., and Blachier, F. (2019). Dietary Protein Intake Level Modulates Mucosal Healing and Mucosa-Adherent Microbiota in Mouse Model of Colitis. Nutrients, 11.","DOI":"10.3390\/nu11030514"},{"key":"ref_111","first-page":"G624","article-title":"Dual effects of a high-protein diet on DSS-treated mice during colitis resolution phase","volume":"311","author":"Lan","year":"2016","journal-title":"Am. J. Physiol. Liver Physiol."},{"key":"ref_112","doi-asserted-by":"crossref","unstructured":"Ijaz, M.U., Ahmed, M.I., Zou, X., Hussain, M., Zhang, M., Zhao, F., Xu, X., Zhou, G., and Li, C. (2018). Beef, Casein, and Soy Proteins Differentially Affect Lipid Metabolism, Triglycerides Accumulation and Gut Microbiota of High-Fat Diet-Fed C57BL\/6J Mice. Front. Microbiol., 9.","DOI":"10.3389\/fmicb.2018.02200"}],"container-title":["Foods"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2304-8158\/14\/21\/3649\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,29]],"date-time":"2025-10-29T05:23:33Z","timestamp":1761715413000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2304-8158\/14\/21\/3649"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,10,26]]},"references-count":112,"journal-issue":{"issue":"21","published-online":{"date-parts":[[2025,11]]}},"alternative-id":["foods14213649"],"URL":"https:\/\/doi.org\/10.3390\/foods14213649","relation":{},"ISSN":["2304-8158"],"issn-type":[{"value":"2304-8158","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,10,26]]}}}