{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,29]],"date-time":"2026-09-29T04:01:09Z","timestamp":1790654469152,"version":"4.1.0"},"reference-count":73,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,9,8]],"date-time":"2026-09-08T00:00:00Z","timestamp":1788825600000},"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. Pharmacol."],"abstract":"<jats:p>Metabolic inflammation is a chronic, low-grade inflammatory state sustained by metabolic stress, organelle dysfunction, immune activation, and gut-derived signals. Because these processes are closely interconnected, single-target interventions may be insufficient. Berberine, a plant-derived isoquinoline alkaloid, has attracted attention for its broad metabolic and anti-inflammatory effects. In this review, we summarize evidence that berberine regulates several interacting pathways, including cellular energy sensing, glucose and lipid metabolism, mitochondrial quality control, redox and inflammatory signaling, and gut\u2013liver\u2013microbiota communication. We also discuss circadian regulation as a relevant but still underexplored aspect of berberine action, focusing on BMAL1\/CLOCK- and REV-ERB\u03b1-related metabolic and inflammatory networks. Current clinical evidence suggests modest benefits for glycemic control, insulin resistance, dyslipidemia, and selected features of steatotic liver disease. However, formulation variability, heterogeneous trial designs, and limited long-term outcome data remain important barriers. Overall, berberine may serve as an adjunctive metabolic modulator, but standardized preparations and time-aware clinical studies are needed.<\/jats:p>","DOI":"10.3389\/fphar.2026.1935151","type":"journal-article","created":{"date-parts":[[2026,9,8]],"date-time":"2026-09-08T08:02:15Z","timestamp":1788854535000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":1,"title":["Berberine as a multitarget modulator of metabolic inflammation: molecular mechanisms, circadian regulation, and translational perspectives"],"prefix":"10.3389","volume":"17","author":[{"given":"Jiaxuan","family":"Xie","sequence":"first","affiliation":[{"name":"Graduate School, Shanghai University of Traditional Chinese Medicine","place":["Shanghai, China"]},{"name":"School of Basic Medical Sciences, Shanghai University of Medicine and Health Sciences","place":["Shanghai, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Junbo","family":"Cao","sequence":"additional","affiliation":[{"name":"Graduate School, Shanghai University of Traditional Chinese Medicine","place":["Shanghai, China"]},{"name":"School of Basic Medical Sciences, Shanghai University of Medicine and Health Sciences","place":["Shanghai, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shuying","family":"Zhang","sequence":"additional","affiliation":[{"name":"School of Clinical Medicine, Shanghai University of Medicine and Health Sciences","place":["Shanghai, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jiayang","family":"Zhang","sequence":"additional","affiliation":[{"name":"School of Basic Medical Sciences, Shanghai University of Medicine and Health Sciences","place":["Shanghai, China"]},{"name":"Health Science Center, East China Normal University","place":["Shanghai, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Guanlin","family":"Wu","sequence":"additional","affiliation":[{"name":"School of Clinical Medicine, Shanghai University of Medicine and Health Sciences","place":["Shanghai, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xinyu","family":"Qiu","sequence":"additional","affiliation":[{"name":"School of Clinical Medicine, Shanghai University of Medicine and Health Sciences","place":["Shanghai, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lihong","family":"Chen","sequence":"additional","affiliation":[{"name":"Health Science Center, East China Normal University","place":["Shanghai, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Guangrui","family":"Yang","sequence":"additional","affiliation":[{"name":"Graduate School, Shanghai University of Traditional Chinese Medicine","place":["Shanghai, China"]},{"name":"School of Basic Medical Sciences, Shanghai University of Medicine and Health Sciences","place":["Shanghai, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,9,8]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"816","DOI":"10.1007\/s11033-026-11967-9","article-title":"Berberine and its structural diversity: structure-activity relationships, molecular mechanisms, and drug discovery opportunities","volume":"53","author":"Albukhari","year":"2026","journal-title":"Mol. Biol. Rep."},{"key":"B2","doi-asserted-by":"publisher","first-page":"e18924","DOI":"10.7717\/peerj.18924","article-title":"Berberine-induced browning and energy metabolism: mechanisms and implications","volume":"13","author":"Alpaslan A\u011fa\u00e7diken","year":"2025","journal-title":"PeerJ"},{"key":"B3","doi-asserted-by":"publisher","first-page":"1575","DOI":"10.1002\/jcb.29392","article-title":"Therapeutic and biological activities of berberine: the involvement of Nrf2 signaling pathway","volume":"121","author":"Ashrafizadeh","year":"2020","journal-title":"J. Cell Biochem."},{"key":"B4","doi-asserted-by":"publisher","first-page":"2284","DOI":"10.3390\/nu16142284","article-title":"Comparative effects of Gymnema sylvestre and berberine on adipokines, body composition, and metabolic parameters in Obese patients: a randomized study","volume":"16","author":"Bandala","year":"2024","journal-title":"Nutrients"},{"key":"B5","doi-asserted-by":"publisher","first-page":"e70317","DOI":"10.1111\/cbdd.70317","article-title":"Synergistic potential of berberine and metformin: unlocking enhanced anti-diabetic efficacy","volume":"107","author":"Bhushan","year":"2026","journal-title":"Chem. Biol. Drug Des."},{"key":"B6","doi-asserted-by":"publisher","first-page":"42","DOI":"10.1186\/s13073-016-0303-2","article-title":"Impact of the gut microbiota on inflammation, obesity, and metabolic disease","volume":"8","author":"Boulang\u00e9","year":"2016","journal-title":"Genome Med."},{"key":"B7","doi-asserted-by":"publisher","first-page":"1751887","DOI":"10.3389\/fmicb.2026.1751887","article-title":"Berberine-gut microbiota interactions based on CiteSpace: current research status and hotspots","volume":"17","author":"Chen","year":"2026","journal-title":"Front. Microbiol."},{"key":"B8","doi-asserted-by":"publisher","first-page":"541","DOI":"10.1016\/j.jpha.2021.10.003","article-title":"Interactions between gut microbiota and berberine, a necessary procedure to understand the mechanisms of berberine","volume":"12","author":"Cheng","year":"2022","journal-title":"J. Pharm. Anal."},{"key":"B9","doi-asserted-by":"publisher","first-page":"27","DOI":"10.1007\/978-3-319-41334-1_2","article-title":"Berberine and its role in chronic disease","volume":"928","author":"Cicero","year":"2016","journal-title":"Adv. Exp. Med. Biol."},{"key":"B10","doi-asserted-by":"publisher","first-page":"312","DOI":"10.2174\/0127724328250471231222094648","article-title":"Berberine: a multi-target natural PCSK9 inhibitor with the potential to treat diabetes, Alzheimer\u2019s, cancer and cardiovascular disease","volume":"19","author":"Coppinger","year":"2024","journal-title":"Curr. Rev. Clin. Exp. Pharmacol."},{"key":"B11","doi-asserted-by":"publisher","first-page":"8485","DOI":"10.1007\/s00210-024-03199-0","article-title":"Research progress on pharmacological effects and bioavailability of berberine","volume":"397","author":"Cui","year":"2024","journal-title":"Naunyn Schmiedeb. Arch. Pharmacol."},{"key":"B12","doi-asserted-by":"publisher","first-page":"1381","DOI":"10.7150\/ijbs.65476","article-title":"Berberine ameliorates DSS-induced intestinal mucosal barrier dysfunction through microbiota-dependence and Wnt\/\u03b2-catenin pathway","volume":"18","author":"Dong","year":"2022","journal-title":"Int. J. Biol. Sci."},{"key":"B13","doi-asserted-by":"publisher","first-page":"121984","DOI":"10.1016\/j.jep.2026.121984","article-title":"Huanglian Wendan decoction attenuates hepatic inflammation and lipogenesis via inhibition of the NF-\u03baB\/HDAC1\/SREBP-1c axis","volume":"370","author":"Fan","year":"2026","journal-title":"J. Ethnopharmacol."},{"key":"B14","doi-asserted-by":"publisher","first-page":"267","DOI":"10.1038\/nrrheum.2017.37","article-title":"Metabolic regulation of inflammation","volume":"13","author":"Gaber","year":"2017","journal-title":"Nat. Rev. Rheumatol."},{"key":"B15","doi-asserted-by":"publisher","first-page":"943706","DOI":"10.1155\/2012\/943706","article-title":"Inflammation and oxidative stress in obesity, metabolic syndrome, and diabetes","volume":"2012","author":"Galassetti","year":"2012","journal-title":"Exp. Diabetes Res."},{"key":"B16","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1007\/s00228-011-1108-2","article-title":"Repeated administration of berberine inhibits cytochromes P450 in humans","volume":"68","author":"Guo","year":"2012","journal-title":"Eur. J. Clin. Pharmacol."},{"key":"B17","doi-asserted-by":"publisher","first-page":"104722","DOI":"10.1016\/j.phrs.2020.104722","article-title":"Berberine pharmacology and the gut microbiota: a hidden therapeutic link","volume":"155","author":"Habtemariam","year":"2020","journal-title":"Pharmacol. Res."},{"key":"B18","doi-asserted-by":"publisher","first-page":"157420","DOI":"10.1016\/j.phymed.2025.157420","article-title":"Network pharmacology of natural alkaloids: rewiring the vicious cycle of mitochondrial dysfunction","volume":"148","author":"He","year":"2025","journal-title":"Phytomedicine"},{"key":"B19","first-page":"20","article-title":"Berberine: a traditional natural product with novel biological activities","volume":"26","author":"Hou","year":"2020","journal-title":"Altern. Ther. Health Med."},{"key":"B20","doi-asserted-by":"publisher","first-page":"1745","DOI":"10.1002\/ptr.5693","article-title":"Berberis vulgaris and berberine: an update review","volume":"30","author":"Imenshahidi","year":"2016","journal-title":"Phytother. Res."},{"key":"B21","doi-asserted-by":"publisher","first-page":"549","DOI":"10.1016\/j.chmed.2023.02.007","article-title":"Antioxidant and antiarthritic potential of berberine: in vitro and in vivo studies","volume":"15","author":"Jain","year":"","journal-title":"Chin. Herbal Medicines"},{"key":"B22","doi-asserted-by":"publisher","first-page":"5775","DOI":"10.55522\/jmpas.v12i3.4800","article-title":"Pharmacognostic and phytochemical assay of Berberisaristata DC roots","volume":"12","author":"Jain","year":"","journal-title":"J. Med. Pharm. Allied Sci."},{"key":"B23","doi-asserted-by":"publisher","first-page":"6862","DOI":"10.3390\/ijms22136862","article-title":"Recent progress in metabolic syndrome research and therapeutics","volume":"22","author":"Kao","year":"2021","journal-title":"Int. J. Mol. Sci."},{"key":"B24","doi-asserted-by":"publisher","first-page":"435","DOI":"10.62946\/ijmphs\/2.4.435-446","article-title":"Herbal isolates and their role in the treatment of liver fibrosis: an updated review: herbal isolates and their role in the treatment of liver fibrosis","volume":"2","author":"Kashyap","year":"2025","journal-title":"Int. J. Med. Pharm. Health Sci."},{"key":"B25","doi-asserted-by":"publisher","first-page":"1344","DOI":"10.1038\/nm1135","article-title":"Berberine is a novel cholesterol-lowering drug working through a unique mechanism distinct from statins","volume":"10","author":"Kong","year":"2004","journal-title":"Nat. Med."},{"key":"B26","doi-asserted-by":"publisher","first-page":"477","DOI":"10.1186\/s40001-025-02738-6","article-title":"Berberine as a multi-target therapeutic agent for obesity: from pharmacological mechanisms to clinical evidence","volume":"30","author":"Kong","year":"2025","journal-title":"Eur. J. Med. Res."},{"key":"B27","doi-asserted-by":"publisher","first-page":"154288","DOI":"10.1016\/j.phymed.2022.154288","article-title":"Biodistribution and pharmacokinetic profile of berberine and its metabolites in hepatocytes","volume":"104","author":"Liu","year":"2022","journal-title":"Phytomedicine"},{"key":"B28","doi-asserted-by":"publisher","first-page":"1890","DOI":"10.3390\/ph18121890","article-title":"Berberine: a rising star in the management of type 2 diabetes-novel insights into its anti-inflammatory, metabolic, and epigenetic mechanisms","volume":"18","author":"Liu","year":"2025","journal-title":"Pharm. (Basel)"},{"key":"B29","doi-asserted-by":"publisher","first-page":"1100","DOI":"10.1124\/dmd.115.063966","article-title":"The effect of oxidation on berberine-mediated CYP1 inhibition: oxidation behavior and metabolite-mediated inhibition","volume":"43","author":"Lo","year":"2015","journal-title":"Drug Metab. Dispos."},{"key":"B30","doi-asserted-by":"publisher","first-page":"297","DOI":"10.1007\/s43440-021-00348-7","article-title":"Berberine and its derivatives represent as the promising therapeutic agents for inflammatory disorders","volume":"74","author":"Lu","year":"2022","journal-title":"Pharmacol. Rep."},{"key":"B31","doi-asserted-by":"publisher","first-page":"117880","DOI":"10.1016\/j.bmc.2024.117880","article-title":"Current development and structure-activity relationship study of berberine derivatives","volume":"112","author":"Luo","year":"2024","journal-title":"Bioorg Med. Chem."},{"key":"B32","doi-asserted-by":"publisher","first-page":"7898","DOI":"10.3390\/ijms24097898","article-title":"Mechanisms of oxidative stress in metabolic syndrome","volume":"24","author":"Masenga","year":"2023","journal-title":"Int. J. Mol. Sci."},{"key":"B33","doi-asserted-by":"publisher","first-page":"10263","DOI":"10.2147\/ijn.S475320","article-title":"Nanocarrier-assisted delivery of berberine promotes diabetic alveolar bone regeneration by scavenging ROS and improving mitochondrial dysfunction","volume":"19","author":"Ming","year":"2024","journal-title":"Int. J. Nanomed."},{"key":"B34","doi-asserted-by":"publisher","first-page":"52","DOI":"10.62946\/ijmphs\/1.2.52-64","article-title":"Interleukin-6: a multifunctional target cytokine for treatment of rheumatoid arthritis","volume":"1","author":"Mishra","year":"2024","journal-title":"Int. J. Med. Pharm. Health Sci."},{"key":"B35","doi-asserted-by":"publisher","first-page":"225","DOI":"10.1186\/s12967-024-05011-2","article-title":"The clinical efficacy and safety of berberine in the treatment of non-alcoholic fatty liver disease: a meta-analysis and systematic review","volume":"22","author":"Nie","year":"2024","journal-title":"J. Transl. Med."},{"key":"B36","doi-asserted-by":"publisher","first-page":"713","DOI":"10.3390\/toxins12110713","article-title":"Biological activity of Berberine-A summary update","volume":"12","author":"Och","year":"2020","journal-title":"Toxins (Basel)"},{"key":"B37","doi-asserted-by":"publisher","first-page":"1351","DOI":"10.3390\/molecules27041351","article-title":"Berberine, a herbal metabolite in the metabolic syndrome: the risk factors, course, and consequences of the disease","volume":"27","author":"Och","year":"2022","journal-title":"Molecules"},{"key":"B38","doi-asserted-by":"publisher","first-page":"106126","DOI":"10.1016\/j.phrs.2022.106126","article-title":"Natural isoquinoline alkaloids: pharmacological features and multi-target potential for complex diseases","volume":"177","author":"Plazas","year":"2022","journal-title":"Pharmacol. Res."},{"key":"B39","doi-asserted-by":"publisher","first-page":"21","DOI":"10.3810\/pgm.2012.01.2514","article-title":"Metabolic syndrome: definition and therapeutic implications","volume":"124","author":"Prasad","year":"2012","journal-title":"Postgrad. Med."},{"key":"B40","doi-asserted-by":"publisher","first-page":"3646","DOI":"10.1111\/bph.14935","article-title":"Berberine protects against diabetic kidney disease via promoting PGC-1\u03b1-regulated mitochondrial energy homeostasis","volume":"177","author":"Qin","year":"2020","journal-title":"Br. J. Pharmacol."},{"key":"B41","doi-asserted-by":"publisher","first-page":"112422","DOI":"10.1016\/j.intimp.2024.112422","article-title":"Berberine, a natural alkaloid: advances in its pharmacological effects and mechanisms in the treatment of autoimmune diseases","volume":"137","author":"Qin","year":"2024","journal-title":"Int. Immunopharmacol."},{"key":"B42","doi-asserted-by":"publisher","first-page":"70259","DOI":"10.3791\/70259","article-title":"Traditional Chinese medicine modulation of the gut-liver axis: from barrier disruption to inflammasome activation in MASLD\/MASH","volume":"230","author":"Ren","year":"2026","journal-title":"J. Vis. Exp."},{"key":"B43","doi-asserted-by":"publisher","first-page":"313","DOI":"10.1017\/s0029665118002872","article-title":"Dietary modulation of energy homoeostasis and metabolic-inflammation","volume":"78","author":"Roche","year":"2019","journal-title":"Proc. Nutr. Soc."},{"key":"B44","doi-asserted-by":"publisher","first-page":"335","DOI":"10.1301\/002966402320583398","article-title":"The metabolic syndrome: where are we and where do we go?","volume":"60","author":"Roth","year":"2002","journal-title":"Nutr. Rev."},{"key":"B45","doi-asserted-by":"publisher","first-page":"e21233","DOI":"10.1016\/j.heliyon.2023.e21233","article-title":"Addressing the preventive and therapeutic perspective of berberine against diabetes","volume":"9","author":"Shrivastava","year":"2023","journal-title":"Heliyon"},{"key":"B46","doi-asserted-by":"publisher","first-page":"564","DOI":"10.1007\/s11684-019-0724-6","article-title":"Biological properties and clinical applications of berberine","volume":"14","author":"Song","year":"2020","journal-title":"Front. Med."},{"key":"B47","doi-asserted-by":"publisher","first-page":"126177","DOI":"10.1016\/j.ijpharm.2025.126177","article-title":"Improving the oral bioavailability of berberine: a crystal engineering approach","volume":"685","author":"Surampudi","year":"2025","journal-title":"Int. J. Pharm."},{"key":"B48","doi-asserted-by":"publisher","first-page":"110691","DOI":"10.1016\/j.diabres.2023.110691","article-title":"Association between obesity, inflammation and insulin resistance: insights into signaling pathways and therapeutic interventions","volume":"200","author":"Suren Garg","year":"2023","journal-title":"Diabetes Res. Clin. Pract."},{"key":"B49","doi-asserted-by":"publisher","first-page":"27","DOI":"10.1208\/s12249-025-03260-y","article-title":"Overcoming bioavailability barriers: optimized hybrid nanocarrier system to amplify berberine absorption via Peyer\u2019s patches","volume":"27","author":"Thool","year":"2025","journal-title":"AAPS PharmSciTech"},{"key":"B50","doi-asserted-by":"publisher","first-page":"139","DOI":"10.1080\/03602532.2017.1306544","article-title":"The metabolism of berberine and its contribution to the pharmacological effects","volume":"49","author":"Wang","year":"2017","journal-title":"Drug Metab. Rev."},{"key":"B51","doi-asserted-by":"publisher","first-page":"971","DOI":"10.2174\/1871530320666200910105612","article-title":"Berberine as a potential multi-target agent for metabolic diseases: a review of investigations for berberine","volume":"21","author":"Wang","year":"2021","journal-title":"Endocr. Metab. Immune Disord. Drug Targets"},{"key":"B52","doi-asserted-by":"publisher","first-page":"155268","DOI":"10.1016\/j.phymed.2023.155268","article-title":"Berberine promotes lacteal junction zippering and ameliorates diet-induced obesity through the RhoA\/ROCK signaling pathway","volume":"124","author":"Wang","year":"2024","journal-title":"Phytomedicine"},{"key":"B53","doi-asserted-by":"publisher","first-page":"4586","DOI":"10.19540\/j.cnki.cjcmm.20240611.706","article-title":"Mechanism of berberine in improving adipocytic IR by mediating BMAL1:CLOCK complex and regulating glucose and lipid metabolism","volume":"49","author":"Wang","year":"2024","journal-title":"Zhongguo Zhong Yao Za Zhi"},{"key":"B54","doi-asserted-by":"publisher","first-page":"1071","DOI":"10.3350\/cmh.2025.0145","article-title":"HTD1801 demonstrates promising potential for histologic improvements in metabolic dysfunction-associated steatohepatitis in both a preclinical and phase 2 study","volume":"31","author":"Wong","year":"2025","journal-title":"Clin. Mol. Hepatol."},{"key":"B55","doi-asserted-by":"publisher","first-page":"567","DOI":"10.1007\/s00228-005-0952-3","article-title":"Effects of berberine on the blood concentration of cyclosporin A in renal transplanted recipients: clinical and pharmacokinetic study","volume":"61","author":"Wu","year":"2005","journal-title":"Eur. J. Clin. Pharmacol."},{"key":"B56","first-page":"1459","article-title":"Moxibustion benignantly regulates circadian rhythm of REV-ERB\u03b1 in RA rats","volume":"12","author":"Wu","year":"2020","journal-title":"Am. J. Transl. Res."},{"key":"B57","doi-asserted-by":"publisher","first-page":"110984","DOI":"10.1016\/j.biopha.2020.110984","article-title":"Therapeutic effect of berberine on metabolic diseases: both pharmacological data and clinical evidence","volume":"133","author":"Xu","year":"2021","journal-title":"Biomed. and Pharmacother."},{"key":"B58","doi-asserted-by":"publisher","first-page":"e13724","DOI":"10.1111\/obr.13724","article-title":"Inflammation-mediated metabolic regulation in adipose tissue","volume":"25","author":"Xu","year":"2024","journal-title":"Obes. Rev."},{"key":"B59","doi-asserted-by":"publisher","first-page":"538","DOI":"10.1186\/s12951-024-02804-x","article-title":"Berberine and magnolol exert cooperative effects on ulcerative colitis in mice by self-assembling into carrier-free nanostructures","volume":"22","author":"Xu","year":"2024","journal-title":"J. Nanobiotechnol."},{"key":"B60","doi-asserted-by":"publisher","first-page":"110069","DOI":"10.1016\/j.jnutbio.2025.110069","article-title":"Berberine alleviates metabolic dysfunction-associated steatohepatitis by enhancing the abundance of Akkermansia muciniphila","volume":"146","author":"Xu","year":"2025","journal-title":"J. Nutr. Biochem."},{"key":"B61","doi-asserted-by":"publisher","first-page":"4293","DOI":"10.19540\/j.cnki.cjcmm.20250410.401","article-title":"Hypoglycemic effect and mechanism of berberine in vitro based on regulation of BMAL1:CLOCK complex involved in hepatic glycolysis, glucose oxidation a nd gluconeogenesis to improve energy metabolism","volume":"50","author":"Xu","year":"2025","journal-title":"Zhongguo Zhong Yao Za Zhi"},{"key":"B62","doi-asserted-by":"publisher","first-page":"253","DOI":"10.1142\/S0192415X24500113","article-title":"Berberine metabolites stimulate GLP-1 secretion by alleviating oxidative stress and mitochondrial dysfunction","volume":"52","author":"Yang","year":"2024","journal-title":"Am. J. Chin. Med."},{"key":"B63","doi-asserted-by":"publisher","first-page":"1874","DOI":"10.3390\/molecules28041874","article-title":"Berberine ameliorates metabolic-associated fatty liver disease mediated metabolism disorder and redox homeostasis by upregulating clock genes: Clock and Bmal1 expressions","volume":"28","author":"Ye","year":"2023","journal-title":"Molecules"},{"key":"B64","doi-asserted-by":"publisher","first-page":"374","DOI":"10.1038\/s41420-021-00768-7","article-title":"Berberine alleviates liver fibrosis through inducing ferrous redox to activate ROS-mediated hepatic stellate cells ferroptosis","volume":"7","author":"Yi","year":"2021","journal-title":"Cell Death Discov."},{"key":"B65","doi-asserted-by":"publisher","first-page":"712","DOI":"10.1016\/j.metabol.2008.01.013","article-title":"Efficacy of berberine in patients with type 2 diabetes mellitus","volume":"57","author":"Yin","year":"2008","journal-title":"Metabolism"},{"key":"B66","doi-asserted-by":"publisher","first-page":"1693","DOI":"10.7150\/ijbs.54604","article-title":"Berberine alleviates lipid metabolism disorders via inhibition of mitochondrial complex I in gut and liver","volume":"17","author":"Yu","year":"2021","journal-title":"Int. J. Biol. Sci."},{"key":"B67","doi-asserted-by":"publisher","first-page":"8803404","DOI":"10.1155\/2022\/8803404","article-title":"Mitochondrial stress in metabolic inflammation: modest benefits and full losses","volume":"2022","author":"Yuan","year":"2022","journal-title":"Oxid. Med. Cell Longev."},{"key":"B68","doi-asserted-by":"publisher","first-page":"588517","DOI":"10.3389\/fcimb.2020.588517","article-title":"Effects of berberine on the gastrointestinal microbiota","volume":"10","author":"Zhang","year":"2020","journal-title":"Front. Cell Infect. Microbiol."},{"key":"B69","doi-asserted-by":"publisher","first-page":"4170","DOI":"10.3390\/ijms26094170","article-title":"Combination of berberine and evodiamine alleviates obesity by promoting browning in 3T3-L1 cells and high-fat diet-induced mice","volume":"26","author":"Zhang","year":"2025","journal-title":"Int. J. Mol. Sci."},{"key":"B70","doi-asserted-by":"publisher","first-page":"151","DOI":"10.1111\/dom.70171","article-title":"Dihydroberberine normalizes insulin secretion by regulating glucokinase","volume":"28","author":"Zhang","year":"2026","journal-title":"Diabetes Obes. Metab."},{"key":"B71","doi-asserted-by":"publisher","first-page":"113773","DOI":"10.1016\/j.bcp.2019.113773","article-title":"Circadian pharmacological effects of berberine on chronic colitis in mice: role of the clock component Rev-erb\u03b1","volume":"172","author":"Zhou","year":"2020","journal-title":"Biochem. Pharmacol."},{"key":"B72","doi-asserted-by":"publisher","first-page":"157030","DOI":"10.1016\/j.phymed.2025.157030","article-title":"Multiple target and regulatory pathways of berberine","volume":"146","author":"Zhou","year":"2025","journal-title":"Phytomedicine"},{"key":"B73","doi-asserted-by":"publisher","first-page":"192","DOI":"10.1016\/j.freeradbiomed.2019.06.019","article-title":"Berberine attenuates nonalcoholic hepatic steatosis through the AMPK-SREBP-1c-SCD1 pathway","volume":"141","author":"Zhu","year":"2019","journal-title":"Free Radic. Biol. Med."}],"container-title":["Frontiers in Pharmacology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fphar.2026.1935151\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,8]],"date-time":"2026-09-08T08:02:23Z","timestamp":1788854543000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fphar.2026.1935151\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,8]]},"references-count":73,"alternative-id":["10.3389\/fphar.2026.1935151"],"URL":"https:\/\/doi.org\/10.3389\/fphar.2026.1935151","relation":{},"ISSN":["1663-9812"],"issn-type":[{"value":"1663-9812","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,8]]},"article-number":"1935151"}}