{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,27]],"date-time":"2026-08-27T11:16:23Z","timestamp":1787829383825,"version":"build-2784847793"},"reference-count":61,"publisher":"MDPI AG","issue":"22","license":[{"start":{"date-parts":[[2025,11,14]],"date-time":"2025-11-14T00:00:00Z","timestamp":1763078400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"National Key Research and Development Program of China","award":["2022YFF1100100"],"award-info":[{"award-number":["2022YFF1100100"]}]},{"name":"Cross-Innovation Open Project of Food Flavor and Health, Beijing Technology & Business University","award":["FFHCI-2025010"],"award-info":[{"award-number":["FFHCI-2025010"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Nutrients"],"abstract":"<jats:p>Background: Obesity is a highly prevalent chronic disease characterized by excessive weight gain and fat accumulation. There is growing evidence that Lactiplantibacillus plantarum strains with bile salt hydrolase (BSH) activity are effective in preventing and alleviating obesity. Methods: Initially, we screened bacterial strains with high hydrolytic activity against glycochenodeoxycholic acid (GDCA), and constructed an isogenic bsh1 knockout mutant. Subsequently, male C57BL\/6J mice fed a high-fat diet (HFD) were randomly assigned to receive daily gavage of either the wild-type Lp20 (Lp20-WT) or the bsh1-deficient mutant (Lp20-\u0394bsh1) for 8 weeks. Serum cholesterol levels and histopathological changes in liver sections were monitored. Hepatic gene expression was quantified by RT-qPCR, and fecal bacterial communities were analyzed via 16S rRNA gene sequencing. These comprehensive assessments aimed to evaluate metabolic improvements and uncover the potential mechanisms behind the observed effects. Results:L. plantarum Lp20 hydrolyzed 91.62% of GDCA, exhibiting the highest bile-salt hydrolase (BSH) activity among tested isolates. Whole-genome sequencing and in-silico analyses mapped this activity to bsh1; gene deletion of bsh1 confirmed the role of bsh1 in GDCA hydrolysis. Daily gavage of the wild-type strain (Lp20-WT) to diet-induced obese mice markedly attenuated weight gain, reduced inguinal white adipose tissue and mesenteric fat mass, and lowered serum TC and LDL-C by 20.8% and 33.3%, respectively, while decreasing ALT and AST levels and reversing hepatic steatosis. In contrast, the bsh1-null mutant (Lp20-\u0394bsh1) failed to elicit any measurable metabolic benefit. Mechanistically, Lp20-WT upregulated rate-limiting bile-acid synthetic enzymes CYP7A1 and CYP27A1, thereby accelerating the catabolism of cholesterol into bile acids. Concurrently, it activated hepatic TGR5 and FXR signaling axes to modulate hepatic metabolism. Moreover, Lp20-WT restructured the gut microbiota by notably enhancing the abundance of beneficial bacteria such as norank_f__Muribaculaceae, Akkermansia, and Alistipes, while reducing the abundance of potentially harmful taxa, including norank_f__Desulfovibrionaceae, Dubosiella, and Mucispirillum. Conclusions: This study provides direct evidence of BSH\u2019s anti-obesity effects through gene deletion. Specifically, BSH lowers cholesterol by modulating hepatic bile-acid metabolism-related gene expression and altering the gut microbiota composition. However, the study is limited by a small sample size (n = 6), the use of male mice only, and its preclinical stage, indicating a need for further validation across diverse strains and human populations.<\/jats:p>","DOI":"10.3390\/nu17223555","type":"journal-article","created":{"date-parts":[[2025,11,14]],"date-time":"2025-11-14T10:07:03Z","timestamp":1763114823000},"page":"3555","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":6,"title":["Lactiplantibacillus plantarum Lp20 Alleviates High Fat Diet-Induced Obesity in Mice via Its Bile Salt Hydrolase Activity"],"prefix":"10.3390","volume":"17","author":[{"given":"Xiaoyue","family":"Bai","sequence":"first","affiliation":[{"name":"College of Food Science and Nutritional Engineering, China Agricultural University, Beijing 100083, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8332-753X","authenticated-orcid":false,"given":"Fangzhou","family":"Lu","sequence":"additional","affiliation":[{"name":"College of Food Science and Nutritional Engineering, China Agricultural University, Beijing 100083, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0005-3694-9327","authenticated-orcid":false,"given":"Yizhi","family":"Jing","sequence":"additional","affiliation":[{"name":"Key Laboratory of Precision Nutrition and Food Quality, Department of Nutrition and Health, China Agricultural University, Beijing 100190, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hui","family":"Wang","sequence":"additional","affiliation":[{"name":"College of Food Science and Nutritional Engineering, China Agricultural University, Beijing 100083, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Haidong","family":"Qian","sequence":"additional","affiliation":[{"name":"Key Laboratory of Precision Nutrition and Food Quality, Department of Nutrition and Health, China Agricultural University, Beijing 100190, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1175-4636","authenticated-orcid":false,"given":"Ming","family":"Zhang","sequence":"additional","affiliation":[{"name":"School of Food and Health, Beijing Technology and Business University, Beijing 100048, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8154-1540","authenticated-orcid":false,"given":"Zhengyuan","family":"Zhai","sequence":"additional","affiliation":[{"name":"College of Food Science and Nutritional Engineering, China Agricultural University, Beijing 100083, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yanling","family":"Hao","sequence":"additional","affiliation":[{"name":"Key Laboratory of Precision Nutrition and Food Quality, Department of Nutrition and Health, China Agricultural University, Beijing 100190, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2025,11,14]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"288","DOI":"10.1038\/s41574-019-0176-8","article-title":"Obesity: Global epidemiology and pathogenesis","volume":"15","year":"2019","journal-title":"Nat. Rev. Endocrinol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"136","DOI":"10.1038\/s41574-023-00932-2","article-title":"Emerging mechanisms of obesity-associated immune dysfunction","volume":"20","author":"Shaikh","year":"2024","journal-title":"Nat. Rev. Endocrinol."},{"key":"ref_3","unstructured":"Lobstein, T., Jackson-Leach, R., Powis, J., Brinsden, H., and Gray, M. (2023). World Obesity Atlas 2023, World Obesity Federation."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1016\/S2213-8587(14)70134-2","article-title":"Obesity, inflammation, and the gut microbiota","volume":"3","author":"Cox","year":"2015","journal-title":"Lancet Diabetes Endocrinol."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"1241214","DOI":"10.1126\/science.1241214","article-title":"Gut microbiota from twins discordant for obesity modulate metabolism in mice","volume":"341","author":"Ridaura","year":"2013","journal-title":"Science"},{"key":"ref_6","doi-asserted-by":"crossref","unstructured":"Michael, D.R., Jack, A.A., Masetti, G., Davies, T.S., Loxley, K.E., Kerry-Smith, J., Plummer, J.F., Marchesi, J.R., Mullish, B.H., and McDonald, J.A.K. (2020). A randomised controlled study shows supplementation of overweight and obese adults with lactobacilli and bifidobacteria reduces bodyweight and improves well-being. Sci. Rep., 10.","DOI":"10.1038\/s41598-020-60991-7"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"2374","DOI":"10.1039\/C5FO01396H","article-title":"Heat-killed and live Lactobacillus reuteri GMNL-263 exhibit similar effects on improving metabolic functions in high-fat diet-induced obese rats","volume":"7","author":"Hsieh","year":"2016","journal-title":"Food Funct."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"5024","DOI":"10.1039\/D0FO00439A","article-title":"The ameliorative effect of Lactobacillus plantarum Y44 oral administration on inflammation and lipid metabolism in obese mice fed with a high fat diet","volume":"11","author":"Liu","year":"2020","journal-title":"Food Funct."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"2300731","DOI":"10.1002\/mnfr.202300731","article-title":"A probiotic targets bile acids metabolism to alleviate ulcerative colitis by reducing conjugated bile acids","volume":"68","author":"Fan","year":"2024","journal-title":"Mol. Nutr. Food Res."},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Li, L., Liu, T., Gu, Y., Wang, X., Xie, R., Sun, Y., Wang, B., and Cao, H. (2022). Regulation of gut microbiota-bile acids axis by probiotics in inflammatory bowel disease. Front. Immunol., 13.","DOI":"10.3389\/fimmu.2022.974305"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"434","DOI":"10.3746\/pnf.2019.24.4.434","article-title":"Probiotic BSH Activity and Anti-Obesity Potential of Lactobacillus plantarum Strain TCI378 Isolated from Korean Kimchi","volume":"24","author":"Huang","year":"2019","journal-title":"Prev. Nutr. Food Sci."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"4315","DOI":"10.1039\/D1FO00260K","article-title":"Lactiplantibacillus plantarum H-87 prevents high-fat diet-induced obesity by regulating bile acid metabolism in C57BL\/6J mice","volume":"12","author":"Liang","year":"2021","journal-title":"Food Funct."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"7421","DOI":"10.1073\/pnas.1323599111","article-title":"Regulation of host weight gain and lipid metabolism by bacterial bile acid modification in the gut","volume":"111","author":"Joyce","year":"2014","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Zheng, Y., Zhang, S., Zhang, Z., Zhang, T., Teng, X., Xiao, G., and Huang, S. (2024). Isolation of Lactobacillus acidophilus strain and its anti-obesity effect in a diet induced obese murine model. Lett. Appl. Microbiol., 77.","DOI":"10.1093\/lambio\/ovae021"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"76","DOI":"10.1016\/j.tim.2022.08.003","article-title":"Gut microbiome-mediated mechanisms for reducing cholesterol levels: Implications for ameliorating cardiovascular disease","volume":"31","author":"Jia","year":"2023","journal-title":"Trends Microbiol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"669","DOI":"10.4161\/19490976.2014.969986","article-title":"Bacterial bile salt hydrolase in host metabolism: Potential for influencing gastrointestinal microbe-host crosstalk","volume":"5","author":"Joyce","year":"2014","journal-title":"Gut Microbes"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"678","DOI":"10.1002\/hep.23721","article-title":"Transgenic expression of cholesterol 7\u03b1-hydroxylase in the liver prevents high-fat diet\u2013induced obesity and insulin resistance in mice","volume":"52","author":"Li","year":"2010","journal-title":"Hepatology"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1016\/j.celrep.2014.02.032","article-title":"Microbiota modification with probiotics induces hepatic bile acid synthesis via downregulation of the Fxr-Fgf15 axis in mice","volume":"7","author":"Degirolamo","year":"2014","journal-title":"Cell Rep."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"392","DOI":"10.3109\/07388551.2014.889077","article-title":"Probiotics and the BSH-related cholesterol lowering mechanism: A Jekyll and Hyde scenario","volume":"35","author":"Choi","year":"2015","journal-title":"Crit. Rev. Biotechnol."},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Xie, C., Huang, W., Young, R.L., Jones, K.L., Horowitz, M., Rayner, C.K., and Wu, T. (2021). Role of Bile Acids in the Regulation of Food Intake, and Their Dysregulation in Metabolic Disease. Nutrients, 13.","DOI":"10.3390\/nu13041104"},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Foley, M.H., O\u2019Flaherty, S., Barrangou, R., and Theriot, C.M. (2019). Bile salt hydrolases: Gatekeepers of bile acid metabolism and host-microbiome crosstalk in the gastrointestinal tract. PLoS Pathog., 15.","DOI":"10.1371\/journal.ppat.1007581"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"629","DOI":"10.1016\/j.str.2023.02.014","article-title":"Characterization of the mechanism of bile salt hydrolase substrate specificity by experimental and computational analyses","volume":"31","author":"Karlov","year":"2023","journal-title":"Structure"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"161","DOI":"10.1017\/S0954422421000081","article-title":"Circulating bile acids as a link between the gut microbiota and cardiovascular health: Impact of prebiotics, probiotics and polyphenol-rich foods","volume":"35","author":"Pushpass","year":"2022","journal-title":"Nutr. Res. Rev."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"236","DOI":"10.1038\/s41579-022-00805-x","article-title":"Bile acids and the gut microbiota: Metabolic interactions and impacts on disease","volume":"21","author":"Collins","year":"2023","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"41","DOI":"10.1016\/j.cmet.2016.05.005","article-title":"Intestinal Crosstalk between Bile Acids and Microbiota and Its Impact on Host Metabolism","volume":"24","author":"Sayin","year":"2016","journal-title":"Cell Metab."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"167","DOI":"10.1007\/164_2019_232","article-title":"Nonsteroidal FXR Ligands: Current Status and Clinical Applications","volume":"256","author":"Gege","year":"2019","journal-title":"Handb. Exp. Pharmacol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"M622","DOI":"10.1111\/j.1750-3841.2011.02431.x","article-title":"All 4 bile salt hydrolase proteins are responsible for the hydrolysis activity in Lactobacillus plantarum ST-III","volume":"76","author":"Ren","year":"2011","journal-title":"J. Food Sci."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"162","DOI":"10.1007\/s12602-012-9101-3","article-title":"Bile Salt Hydrolase (Bsh) Activity Screening of Lactobacilli: In Vitro Selection of Indigenous Lactobacillus Strains with Potential Bile Salt Hydrolysing and Cholesterol-Lowering Ability","volume":"4","author":"Kumar","year":"2012","journal-title":"Probiotics Antimicrob. Proteins"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"391","DOI":"10.1016\/j.idairyj.2004.08.007","article-title":"Bile salt deconjugation ability, bile salt hydrolase activity and cholesterol co-precipitation ability of lactobacilli strains","volume":"15","author":"Liong","year":"2005","journal-title":"Int. Dairy J."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"725","DOI":"10.1039\/D1FO03143K","article-title":"Cholesterol-lowering effect of bile salt hydrolase from a Lactobacillus johnsonii strain mediated by FXR pathway regulation","volume":"13","author":"Zhu","year":"2022","journal-title":"Food Funct."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"1340","DOI":"10.1039\/D3FO04871C","article-title":"Cholesterol lowering in diet-induced hypercholesterolemic mice using Lactobacillus bile salt hydrolases with different substrate specificities","volume":"15","author":"Zhao","year":"2024","journal-title":"Food Funct."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"D233","DOI":"10.1093\/nar\/gkn663","article-title":"The Carbohydrate-Active EnZymes database (CAZy): An expert resource for Glycogenomics","volume":"37","author":"Cantarel","year":"2009","journal-title":"Nucleic Acids Res."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1038\/75556","article-title":"Gene ontology: Tool for the unification of biology. The Gene Ontology Consortium","volume":"25","author":"Ashburner","year":"2000","journal-title":"Nat. Genet."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"D277","DOI":"10.1093\/nar\/gkh063","article-title":"The KEGG resource for deciphering the genome","volume":"32","author":"Kanehisa","year":"2004","journal-title":"Nucleic Acids Res."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"D261","DOI":"10.1093\/nar\/gku1223","article-title":"Expanded microbial genome coverage and improved protein family annotation in the COG database","volume":"43","author":"Galperin","year":"2015","journal-title":"Nucleic Acids Res."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"D461","DOI":"10.1093\/nar\/gkaa1004","article-title":"The Transporter Classification Database (TCDB): 2021 update","volume":"49","author":"Saier","year":"2021","journal-title":"Nucleic Acids Res."},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Alikhan, N.F., Petty, N.K., Ben Zakour, N.L., and Beatson, S.A. (2011). BLAST Ring Image Generator (BRIG): Simple prokaryote genome comparisons. BMC Genom., 12.","DOI":"10.1186\/1471-2164-12-402"},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Huo, Y., Lu, X., Wang, X., Wang, X., Chen, L., Guo, H., Zhang, M., and Li, Y. (2020). Bifidobacterium animalis subsp. lactis A6 Alleviates Obesity Associated with Promoting Mitochondrial Biogenesis and Function of Adipose Tissue in Mice. Molecules, 25.","DOI":"10.3390\/molecules25071490"},{"key":"ref_39","first-page":"491","article-title":"Diagnosis of nonalcoholic fatty liver disease: The importance of pathology","volume":"39","author":"Tian","year":"2023","journal-title":"J. Clin. Hepatol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"1101","DOI":"10.1038\/nprot.2008.73","article-title":"Analyzing real-time PCR data by the comparative C(T) method","volume":"3","author":"Schmittgen","year":"2008","journal-title":"Nat. Protoc."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"852","DOI":"10.1038\/s41587-019-0209-9","article-title":"Reproducible, interactive, scalable and extensible microbiome data science using QIIME 2","volume":"37","author":"Bolyen","year":"2019","journal-title":"Nat. Biotechnol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"i884","DOI":"10.1093\/bioinformatics\/bty560","article-title":"fastp: An ultra-fast all-in-one FASTQ preprocessor","volume":"34","author":"Chen","year":"2018","journal-title":"Bioinformatics"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"2957","DOI":"10.1093\/bioinformatics\/btr507","article-title":"FLASH: Fast length adjustment of short reads to improve genome assemblies","volume":"27","author":"Salzberg","year":"2011","journal-title":"Bioinformatics"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"581","DOI":"10.1038\/nmeth.3869","article-title":"DADA2: High-resolution sample inference from Illumina amplicon data","volume":"13","author":"Callahan","year":"2016","journal-title":"Nat. Methods"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1128\/mSystems.00191-16","article-title":"Deblur rapidly resolves single-nucleotide community sequence patterns","volume":"2","author":"Amir","year":"2017","journal-title":"mSystems"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"4719","DOI":"10.1128\/AEM.00137-08","article-title":"Functional analysis of four bile salt hydrolase and penicillin acylase family members in Lactobacillus plantarum WCFS1","volume":"74","author":"Lambert","year":"2008","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_47","doi-asserted-by":"crossref","unstructured":"Wang, B., Kong, Q., Li, X., Zhao, J., Zhang, H., Chen, W., and Wang, G. (2020). A high-fat diet increases gut microbiota biodiversity and energy expenditure due to nutrient difference. Nutrients, 12.","DOI":"10.3390\/nu12103197"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"e13349","DOI":"10.1111\/obr.13349","article-title":"The dietary inflammatory index, obesity, type 2 diabetes, and cardiovascular risk factors and diseases","volume":"23","author":"Hariharan","year":"2022","journal-title":"Obes. Rev."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"e002519","DOI":"10.1136\/bmjph-2024-002519","article-title":"What are community perspectives and experiences around GLP-1 receptor agonist medications for weight loss? A cross-sectional survey study in the UK","volume":"3","author":"Auerbach","year":"2025","journal-title":"BMJ Public Health"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"P4530","DOI":"10.1016\/j.cell.2025.05.010","article-title":"A microbial amino-acid-conjugated bile acid, tryptophan-cholic acid, improves glucose homeostasis via the orphan receptor MRGPRE","volume":"188","author":"Lin","year":"2025","journal-title":"Cell"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"852","DOI":"10.1038\/s41586-024-07017-8","article-title":"Bile salt hydrolase acyltransferase activity expands bile acid diversity","volume":"626","author":"Guzior","year":"2024","journal-title":"Nature"},{"key":"ref_52","doi-asserted-by":"crossref","unstructured":"Prete, R., Long, S.L., Gallardo, A.L., Gahan, C.G., Corsetti, A., and Joyce, S.A. (2020). Beneficial bile acid metabolism from Lactobacillus plantarum of food origin. Sci. Rep., 10.","DOI":"10.1038\/s41598-020-58069-5"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"2610","DOI":"10.1038\/s41467-020-16438-8","article-title":"Large-scale genome-wide analysis links lactic acid bacteria from food with the gut microbiome","volume":"11","author":"Pasolli","year":"2020","journal-title":"Nat. Commun."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"13580","DOI":"10.1073\/pnas.0804437105","article-title":"Functional and comparative metagenomic analysis of bile salt hydrolase activity in the human gut microbiome","volume":"105","author":"Jones","year":"2008","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"S27","DOI":"10.1093\/femsre\/fux030","article-title":"Lifestyles in transition: Evolution and natural history of the genus Lactobacillus","volume":"41","author":"Duar","year":"2017","journal-title":"FEMS Microbiol. Rev."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"111208","DOI":"10.1016\/j.lwt.2021.111208","article-title":"Specific bile salt hydrolase genes in Lactobacillus plantarum AR113 and relationship with bile salt resistance","volume":"145","author":"Wang","year":"2021","journal-title":"LWT"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"412","DOI":"10.1038\/nrgastro.2016.85","article-title":"The role of the gut microbiota in NAFLD","volume":"13","author":"Leung","year":"2016","journal-title":"Nat. Rev. Gastroenterol. Hepatol."},{"key":"ref_58","first-page":"1287","article-title":"High-fat diet decreases activity of the oxidative phosphorylation complexes and causes nonalcoholic steatohepatitis in mice","volume":"7","author":"Grau","year":"2014","journal-title":"Dis. Model. Mech."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"1266","DOI":"10.1056\/NEJM200004273421707","article-title":"Evaluation of abnormal liver-enzyme results in asymptomatic patients","volume":"342","author":"Pratt","year":"2000","journal-title":"N. Engl. J. Med."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"1034","DOI":"10.1002\/hep.25740","article-title":"Mechanism of tissue-specific farnesoid X receptor in suppressing the expression of genes in bile-acid synthesis in mice","volume":"56","author":"Kong","year":"2012","journal-title":"Hepatology"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"18","DOI":"10.1016\/S1499-3872(14)60307-6","article-title":"Bile acid signaling through farnesoid X and TGR5 receptors in hepatobiliary and intestinal diseases","volume":"14","author":"Stanimirov","year":"2015","journal-title":"Hepatobiliary Pancreat. Dis. Int."}],"container-title":["Nutrients"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2072-6643\/17\/22\/3555\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,11,16]],"date-time":"2025-11-16T05:13:38Z","timestamp":1763270018000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2072-6643\/17\/22\/3555"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,11,14]]},"references-count":61,"journal-issue":{"issue":"22","published-online":{"date-parts":[[2025,11]]}},"alternative-id":["nu17223555"],"URL":"https:\/\/doi.org\/10.3390\/nu17223555","relation":{},"ISSN":["2072-6643"],"issn-type":[{"value":"2072-6643","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,11,14]]}}}