Berberine

Last Updated: September 28 2022

Berberine is a plant alkaloid with a long history of use in Ayurvedic and traditional Chinese medicine for the management of various health conditions. Its seemingly potent ability to reduce blood sugar rivals that of some anti-diabetic drugs.

Berberine is most often used for

What is berberine?

Berberine is an alkaloid found in the barks, leaves, twigs, rhizomes, roots, and/or stems of various plants, such as the barberry, Oregon grape, and tree turmeric. Traditionally, berberine has been used as an antimicrobial, antiprotozoal, and antidiarrheal agent in Ayurvedic medicine and traditional Chinese medicine. Most research in humans has examined berberine’s effects on markers of glycemic control, blood lipids, markers of liver function, and anthropometric parameters in people with metabolic disorders.

What are berberine’s main benefits?

Evidence from clinical trials conducted in people with type 2 diabetes suggests that berberine is able to reduce blood sugar to a similar extent as some anti-diabetic drugs. In people with metabolic disorders, limited research suggests that berberine may improve blood lipids and liver enzymes, and reduce body weight and fat mass. However, most of the available trials examining the effects of berberine on the aforementioned outcomes are of poor methodological quality.

What are berberine’s main drawbacks?

Although berberine appears to be generally safe in normal doses, more long-term research on its safety is needed. In some people, supplementation with berberine has been reported to cause gastrointestinal side effects, including diarrhea, constipation, flatulence, and stomach pain. Due to its ability to reduce blood sugar, berberine may increase the risk for hypoglycemia in high doses. Berberine has a high potential of interacting with a number of drugs, with some of these interactions increasing the chances of serious complications.

How does berberine work?

The main mechanism through which berberine exerts its effects appears to be the stimulation of the enzyme adenosine monophosphate-activated protein kinase (AMPK), a master regulator of cellular energy homeostasis that is involved in a wide range of biological processes, including the regulation of glucose, lipid, and protein metabolism.

What else is Berberine known as?
Berberine should not be confused with:
  • Piperine (Black Pepper extract)
  • Berberol (Brand name)
  • Berberrubine (Metabolite)
Dosage information

The standard dose of berberine is 900-2,000mg a day, divided into three to four doses.

Berberine should be taken with a meal, or shortly after, to take advantage of the blood glucose and lipid spike associated with eating.

Too much berberine at once can result in stomach upset, cramping, and diarrhea.

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      Research Breakdown

      References
      1.^Vuddanda PR, Chakraborty S, Singh SBerberine: a potential phytochemical with multispectrum therapeutic activitiesExpert Opin Investig Drugs.(2010 Oct)
      2.^Cernáková M, Kostálová DAntimicrobial activity of berberine--a constituent of Mahonia aquifoliumFolia Microbiol (Praha).(2002)
      5.^Ghosh AK, Bhattacharyya FK, Ghosh DKLeishmania donovani: amastigote inhibition and mode of action of berberineExp Parasitol.(1985 Dec)
      7.^Grycová L, Dostál J, Marek RQuaternary protoberberine alkaloidsPhytochemistry.(2007 Jan)
      8.^Singh IP, Mahajan SBerberine and its derivatives: a patent review (2009 - 2012)Expert Opin Ther Pat.(2012 Dec 12)
      9.^Krivogorsky B, Pernat JA, Douglas KA, Czerniecki NJ, Grundt PStructure-activity studies of some berberine analogs as inhibitors of Toxoplasma gondiiBioorg Med Chem Lett.(2012 Feb 22)
      12.^Tillhon M, Guamán Ortiz LM, Lombardi P, Scovassi AIBerberine: new perspectives for old remediesBiochem Pharmacol.(2012 Nov 15)
      13.^Wang JR, Tanaka T, Zhang H, Kouno I, Jiang ZHFormation and conformation of baicalin-berberine and wogonoside-berberine complexesChem Pharm Bull (Tokyo).(2012)
      15.^Cheng Z, Chen AF, Wu F, Sheng L, Zhang HK, Gu M, Li YY, Zhang LN, Hu LH, Li JY, Li J8,8-Dimethyldihydroberberine with improved bioavailability and oral efficacy on obese and diabetic mouse modelsBioorg Med Chem.(2010 Aug 15)
      16.^Subeki, Matsuura H, Takahashi K, Yamasaki M, Yamato O, Maede Y, Katakura K, Suzuki M, Trimurningsih, Chairul, Yoshihara TAntibabesial activity of protoberberine alkaloids and 20-hydroxyecdysone from Arcangelisia flava against Babesia gibsoni in cultureJ Vet Med Sci.(2005 Feb)
      17.^Schramm A, Baburin I, Hering S, Hamburger MHERG channel inhibitors in extracts of Coptidis rhizomaPlanta Med.(2011 May)
      20.^Pan GY, Wang GJ, Sun JG, Huang ZJ, Zhao XC, Gu Y, Liu XDInhibitory action of berberine on glucose absorptionYao Xue Xue Bao.(2003 Dec)
      21.^Maeng HJ, Yoo HJ, Kim IW, Song IS, Chung SJ, Shim CKP-glycoprotein-mediated transport of berberine across Caco-2 cell monolayersJ Pharm Sci.(2002 Dec)
      22.^Chen W, Miao YQ, Fan DJ, Yang SS, Lin X, Meng LK, Tang XBioavailability study of berberine and the enhancing effects of TPGS on intestinal absorption in ratsAAPS PharmSciTech.(2011 Jun)
      23.^Pan GY, Wang GJ, Liu XD, Fawcett JP, Xie YYThe involvement of P-glycoprotein in berberine absorptionPharmacol Toxicol.(2002 Oct)
      24.^Chae HW, Kim IW, Jin HE, Kim DD, Chung SJ, Shim CKEffect of ion-pair formation with bile salts on the in vitro cellular transport of berberineArch Pharm Res.(2008 Jan)
      26.^Qiu W, Jiang XH, Liu CX, Ju Y, Jin JXEffect of berberine on the pharmacokinetics of substrates of CYP3A and P-gpPhytother Res.(2009 Nov)
      28.^Hwang YH, Cho WK, Jang D, Ha JH, Jung K, Yun HI, Ma JYEffects of berberine and hwangryunhaedok-tang on oral bioavailability and pharmacokinetics of ciprofloxacin in ratsEvid Based Complement Alternat Med.(2012)
      29.^Shan YQ, Ren G, Wang YX, Pang J, Zhao ZY, Yao J, You XF, Si SY, Song DQ, Kong WJ, Jiang JDBerberine analogue IMB-Y53 improves glucose-lowering efficacy by averting cellular efflux especially P-glycoprotein effluxMetabolism.(2012 Oct 15)
      30.^Di Pierro F, Villanova N, Agostini F, Marzocchi R, Soverini V, Marchesini GPilot study on the additive effects of berberine and oral type 2 diabetes agents for patients with suboptimal glycemic controlDiabetes Metab Syndr Obes.(2012)
      33.^Zhang M, Lv X, Li J, Meng Z, Wang Q, Chang W, Li W, Chen L, Liu YSodium caprate augments the hypoglycemic effect of berberine via AMPK in inhibiting hepatic gluconeogenesisMol Cell Endocrinol.(2012 Nov 5)
      36.^Zhang Y, Li X, Zou D, Liu W, Yang J, Zhu N, Huo L, Wang M, Hong J, Wu P, Ren G, Ning GTreatment of type 2 diabetes and dyslipidemia with the natural plant alkaloid berberineJ Clin Endocrinol Metab.(2008 Jul)
      37.^Khin-Maung-U, Myo-Khin, Nyunt-Nyunt-Wai, Aye-Kyaw, Tin-UClinical trial of berberine in acute watery diarrhoeaBr Med J (Clin Res Ed).(1985 Dec 7)
      40.^Wang YX, Kong WJ, Li YH, Tang S, Li Z, Li YB, Shan YQ, Bi CW, Jiang JD, Song DQSynthesis and structure-activity relationship of berberine analogues in LDLR up-regulation and AMPK activationBioorg Med Chem.(2012 Nov 15)
      41.^Li Y, Ren G, Wang YX, Kong WJ, Yang P, Wang YM, Li YH, Yi H, Li ZR, Song DQ, Jiang JDBioactivities of berberine metabolites after transformation through CYP450 isoenzymesJ Transl Med.(2011 May 15)
      43.^Qiu F, Zhu Z, Kang N, Piao S, Qin G, Yao XIsolation and identification of urinary metabolites of berberine in rats and humansDrug Metab Dispos.(2008 Nov)
      44.^Pan JF, Yu C, Zhu DY, Zhang H, Zeng JF, Jiang SH, Ren JYIdentification of three sulfate-conjugated metabolites of berberine chloride in healthy volunteers' urine after oral administrationActa Pharmacol Sin.(2002 Jan)
      47.^Guo Y, Chen Y, Tan ZR, Klaassen CD, Zhou HHRepeated administration of berberine inhibits cytochromes P450 in humansEur J Clin Pharmacol.(2012 Feb)
      48.^Xin HW, Wu XC, Li Q, Yu AR, Zhong MY, Liu YYThe effects of berberine on the pharmacokinetics of cyclosporin A in healthy volunteersMethods Find Exp Clin Pharmacol.(2006 Jan-Feb)
      53.^Chatuphonprasert W, Nemoto N, Sakuma T, Jarukamjorn KModulations of cytochrome P450 expression in diabetic mice by berberineChem Biol Interact.(2012 Mar 5)
      54.^Cheng Q, Aleksunes LM, Manautou JE, Cherrington NJ, Scheffer GL, Yamasaki H, Slitt ALDrug-metabolizing enzyme and transporter expression in a mouse model of diabetes and obesityMol Pharm.(2008 Jan-Feb)
      55.^Viollet B, Guigas B, Sanz Garcia N, Leclerc J, Foretz M, Andreelli FCellular and molecular mechanisms of metformin: an overviewClin Sci (Lond).(2012 Mar)
      56.^Han Y, Wang Q, Song P, Zhu Y, Zou MHRedox regulation of the AMP-activated protein kinasePLoS One.(2010 Nov 5)
      57.^Cheng Z, Pang T, Gu M, Gao AH, Xie CM, Li JY, Nan FJ, Li JBerberine-stimulated glucose uptake in L6 myotubes involves both AMPK and p38 MAPKBiochim Biophys Acta.(2006 Nov)
      60.^Kim WS, Lee YS, Cha SH, Jeong HW, Choe SS, Lee MR, Oh GT, Park HS, Lee KU, Lane MD, Kim JBBerberine improves lipid dysregulation in obesity by controlling central and peripheral AMPK activityAm J Physiol Endocrinol Metab.(2009 Apr)
      61.^Brusq JM, Ancellin N, Grondin P, Guillard R, Martin S, Saintillan Y, Issandou MInhibition of lipid synthesis through activation of AMP kinase: an additional mechanism for the hypolipidemic effects of berberineJ Lipid Res.(2006 Jun)
      62.^Kong W, Wei J, Abidi P, Lin M, Inaba S, Li C, Wang Y, Wang Z, Si S, Pan H, Wang S, Wu J, Wang Y, Li Z, Liu J, Jiang JDBerberine is a novel cholesterol-lowering drug working through a unique mechanism distinct from statinsNat Med.(2004 Dec)
      64.^Turnley AM, Stapleton D, Mann RJ, Witters LA, Kemp BE, Bartlett PFCellular distribution and developmental expression of AMP-activated protein kinase isoforms in mouse central nervous systemJ Neurochem.(1999 Apr)
      66.^Ou TM, Lu YJ, Tan JH, Huang ZS, Wong KY, Gu LQG-quadruplexes: targets in anticancer drug designChemMedChem.(2008 May)
      67.^Monchaud D, Teulade-Fichou MPA hitchhiker's guide to G-quadruplex ligandsOrg Biomol Chem.(2008 Feb 21)
      68.^Arora A, Balasubramanian C, Kumar N, Agrawal S, Ojha RP, Maiti SBinding of berberine to human telomeric quadruplex - spectroscopic, calorimetric and molecular modeling studiesFEBS J.(2008 Aug)
      69.^Zhang WJ, Ou TM, Lu YJ, Huang YY, Wu WB, Huang ZS, Zhou JL, Wong KY, Gu LQ9-Substituted berberine derivatives as G-quadruplex stabilizing ligands in telomeric DNABioorg Med Chem.(2007 Aug 15)
      76.^Hui KK, Yu JL, Chan WF, Tse EInteraction of berberine with human platelet alpha 2 adrenoceptorsLife Sci.(1991)
      77.^Vizi ES, Toth I, Somogyi GT, Szabo L, Harsing LG Jr, Szantay CBerbanes: a new class of selective alpha 2-adrenoceptor antagonistsJ Med Chem.(1987 Aug)
      79.^Kulkarni SK, Dhir AOn the mechanism of antidepressant-like action of berberine chlorideEur J Pharmacol.(2008 Jul 28)
      81.^Kong LD, Cheng CH, Tan RXMonoamine oxidase inhibitors from rhizoma of Coptis chinensisPlanta Med.(2001 Feb)
      82.^Castillo J, Hung J, Rodriguez M, Bastidas E, Laboren I, Jaimes ALED fluorescence spectroscopy for direct determination of monoamine oxidase B inactivationAnal Biochem.(2005 Aug 15)
      84.^Kalalian-Moghaddam H, Baluchnejadmojarad T, Roghani M, Goshadrou F, Ronaghi AHippocampal synaptic plasticity restoration and anti-apoptotic effect underlie berberine improvement of learning and memory in streptozotocin-diabetic ratsEur J Pharmacol.(2013 Jan 5)
      85.^Reisi P, Babri S, Alaei H, Sharifi MR, Mohaddes G, Noorbakhsh SM, Lashgari RTreadmill running improves long-term potentiation (LTP) defects in streptozotocin-induced diabetes at dentate gyrus in ratsPathophysiology.(2010 Feb)
      88.^Bhutada P, Mundhada Y, Bansod K, Tawari S, Patil S, Dixit P, Umathe S, Mundhada DProtection of cholinergic and antioxidant system contributes to the effect of berberine ameliorating memory dysfunction in rat model of streptozotocin-induced diabetesBehav Brain Res.(2011 Jun 20)
      90.^Kulkarni SK, Dhir Asigma-1 receptors in major depression and anxietyExpert Rev Neurother.(2009 Jul)
      94.^Fontanilla D, Johannessen M, Hajipour AR, Cozzi NV, Jackson MB, Ruoho AEThe hallucinogen N,N-dimethyltryptamine (DMT) is an endogenous sigma-1 receptor regulatorScience.(2009 Feb 13)
      96.^Jung HA, Min BS, Yokozawa T, Lee JH, Kim YS, Choi JSAnti-Alzheimer and antioxidant activities of Coptidis Rhizoma alkaloidsBiol Pharm Bull.(2009 Aug)
      97.^Ingkaninan K, Phengpa P, Yuenyongsawad S, Khorana NAcetylcholinesterase inhibitors from Stephania venosa tuberJ Pharm Pharmacol.(2006 May)
      98.^Chen XW, Di YM, Zhang J, Zhou ZW, Li CG, Zhou SFInteraction of herbal compounds with biological targets: a case study with berberineScientificWorldJournal.(2012)
      99.^Asai M, Iwata N, Yoshikawa A, Aizaki Y, Ishiura S, Saido TC, Maruyama KBerberine alters the processing of Alzheimer's amyloid precursor protein to decrease Abeta secretionBiochem Biophys Res Commun.(2007 Jan 12)
      103.^Hu J, Chai Y, Wang Y, Kheir MM, Li H, Yuan Z, Wan H, Xing D, Lei F, Du LPI3K p55γ promoter activity enhancement is involved in the anti-apoptotic effect of berberine against cerebral ischemia-reperfusionEur J Pharmacol.(2012 Jan 15)
      104.^Kwon IH, Choi HS, Shin KS, Lee BK, Lee CK, Hwang BY, Lim SC, Lee MKEffects of berberine on 6-hydroxydopamine-induced neurotoxicity in PC12 cells and a rat model of Parkinson's diseaseNeurosci Lett.(2010 Dec 3)
      105.^Segura Aguilar J, Kostrzewa RMNeurotoxins and neurotoxic species implicated in neurodegenerationNeurotox Res.(2004)
      109.^Wang LH, Li XL, Li Q, Fu Y, Yu HJ, Sun YQ, Zhang L, Shan HLBerberine alleviates ischemic arrhythmias via recovering depressed I(to) and I(Ca) currents in diabetic ratsPhytomedicine.(2012 Feb 15)
      111.^Kim YM, Ha YM, Jin YC, Shi LY, Lee YS, Kim HJ, Seo HG, Choi JS, Kim YS, Kang SS, Lee JH, Chang KCPalmatine from Coptidis rhizoma reduces ischemia-reperfusion-mediated acute myocardial injury in the ratFood Chem Toxicol.(2009 Aug)
      112.^Soloff MS, Gal S, Hoare S, Peters CA, Hunzicker-Dunn M, Anderson GD, Wood TGCloning, characterization, and expression of the rat relaxin geneGene.(2003 Dec 24)
      113.^Gu HP, Lin S, Xu M, Yu HY, Du XJ, Zhang YY, Yuan G, Gao WUp-regulating relaxin expression by G-quadruplex interactive ligand to achieve antifibrotic actionEndocrinology.(2012 Aug)
      114.^Du XJ, Bathgate RA, Samuel CS, Dart AM, Summers RJCardiovascular effects of relaxin: from basic science to clinical therapyNat Rev Cardiol.(2010 Jan)
      118.^Zheng L, Zhou Z, Tao D, Lan TProtective effect of berberine on cardiac myocyte injured by ischemia-reperfusionSichuan Da Xue Xue Bao Yi Xue Ban.(2003 Jul)
      119.^Xiao HB, Sun ZL, Zhang HB, Zhang DSBerberine inhibits dyslipidemia in C57BL/6 mice with lipopolysaccharide induced inflammationPharmacol Rep.(2012 Jul)
      120.^Lee S, Lim HJ, Park JH, Lee KS, Jang Y, Park HYBerberine-induced LDLR up-regulation involves JNK pathwayBiochem Biophys Res Commun.(2007 Nov 3)
      122.^Duff CJ, Hooper NMPCSK9: an emerging target for treatment of hypercholesterolemiaExpert Opin Ther Targets.(2011 Feb)
      123.^Feingold KR, Moser AH, Shigenaga JK, Patzek SM, Grunfeld CInflammation stimulates the expression of PCSK9Biochem Biophys Res Commun.(2008 Sep 19)
      126.^Hu Y, Ehli EA, Kittelsrud J, Ronan PJ, Munger K, Downey T, Bohlen K, Callahan L, Munson V, Jahnke M, Marshall LL, Nelson K, Huizenga P, Hansen R, Soundy TJ, Davies GELipid-lowering effect of berberine in human subjects and ratsPhytomedicine.(2012 Jul 15)
      127.^Derosa G, D'Angelo A, Bonaventura A, Bianchi L, Romano D, Maffioli PEffects of berberine on lipid profile in subjects with low cardiovascular riskExpert Opin Biol Ther.(2013 Apr)
      128.^Dong H, Wang N, Zhao L, Lu FBerberine in the treatment of type 2 diabetes mellitus: a systemic review and meta-analysisEvid Based Complement Alternat Med.(2012)
      129.^Fatehi-Hassanabad Z, Jafarzadeh M, Tarhini A, Fatehi MThe antihypertensive and vasodilator effects of aqueous extract from Berberis vulgaris fruit on hypertensive ratsPhytother Res.(2005 Mar)
      130.^Chun YT, Yip TT, Lau KL, Kong YC, Sankawa UA biochemical study on the hypotensive effect of berberine in ratsGen Pharmacol.(1979)
      131.^Lau CW, Yao XQ, Chen ZY, Ko WH, Huang YCardiovascular actions of berberineCardiovasc Drug Rev.(2001 Fall)
      135.^Bova S, Padrini R, Goldman WF, Berman DM, Cargnelli GOn the mechanism of vasodilating action of berberine: possible role of inositol lipid signaling systemJ Pharmacol Exp Ther.(1992 Apr)
      136.^Huang CG, Chu ZL, Wei SJ, Jiang H, Jiao BHEffect of berberine on arachidonic acid metabolism in rabbit platelets and endothelial cellsThromb Res.(2002 May 15)
      138.^Cao JW, Luo HS, Yu BP, Huang XD, Sheng ZX, Yu JPEffects of berberine on intracellular free calcium in smooth muscle cells of Guinea pig colonDigestion.(2001)
      143.^Huang C, Zhang Y, Gong Z, Sheng X, Li Z, Zhang W, Qin YBerberine inhibits 3T3-L1 adipocyte differentiation through the PPARgamma pathwayBiochem Biophys Res Commun.(2006 Sep 22)
      146.^Pham TP, Kwon J, Shin JBerberine exerts anti-adipogenic activity through up-regulation of C/EBP inhibitors, CHOP and DEC2Biochem Biophys Res Commun.(2011 Sep 23)
      149.^Zhou L, Wang X, Yang Y, Wu L, Li F, Zhang R, Yuan G, Wang N, Chen M, Ning GBerberine attenuates cAMP-induced lipolysis via reducing the inhibition of phosphodiesterase in 3T3-L1 adipocytesBiochim Biophys Acta.(2011 Apr)
      150.^Zhou L, Wang X, Shao L, Yang Y, Shang W, Yuan G, Jiang B, Li F, Tang J, Jing H, Chen MBerberine acutely inhibits insulin secretion from beta-cells through 3',5'-cyclic adenosine 5'-monophosphate signaling pathwayEndocrinology.(2008 Sep)
      151.^Kadowaki T, Yamauchi TAdiponectin and adiponectin receptorsEndocr Rev.(2005 May)
      154.^Li Y, Wang P, Zhuang Y, Lin H, Li Y, Liu L, Meng Q, Cui T, Liu J, Li ZActivation of AMPK by berberine promotes adiponectin multimerization in 3T3-L1 adipocytesFEBS Lett.(2011 Jun 23)
      158.^Chen C, Zhang Y, Huang CBerberine inhibits PTP1B activity and mimics insulin actionBiochem Biophys Res Commun.(2010 Jul 2)
      159.^Cok A, Plaisier C, Salie MJ, Oram DS, Chenge J, Louters LLBerberine acutely activates the glucose transport activity of GLUT1Biochimie.(2011 Jul)
      160.^Kim SH, Shin EJ, Kim ED, Bayaraa T, Frost SC, Hyun CKBerberine activates GLUT1-mediated glucose uptake in 3T3-L1 adipocytesBiol Pharm Bull.(2007 Nov)
      162.^Yi P, Lu FE, Xu LJ, Chen G, Dong H, Wang KFBerberine reverses free-fatty-acid-induced insulin resistance in 3T3-L1 adipocytes through targeting IKKbetaWorld J Gastroenterol.(2008 Feb 14)
      163.^Choi BH, Ahn IS, Kim YH, Park JW, Lee SY, Hyun CK, Do MSBerberine reduces the expression of adipogenic enzymes and inflammatory molecules of 3T3-L1 adipocyteExp Mol Med.(2006 Dec 31)
      165.^Yan HM, Xia MF, Wang Y, Chang XX, Yao XZ, Rao SX, Zeng MS, Tu YF, Feng R, Jia WP, Liu J, Deng W, Jiang JD, Gao XEfficacy of Berberine in Patients with Non-Alcoholic Fatty Liver DiseasePLoS One.(2015 Aug 7)
      166.^Pan GY, Huang ZJ, Wang GJ, Fawcett JP, Liu XD, Zhao XC, Sun JG, Xie YYThe antihyperglycaemic activity of berberine arises from a decrease of glucose absorptionPlanta Med.(2003 Jul)
      168.^Liu L, Yu YL, Yang JS, Li Y, Liu YW, Liang Y, Liu XD, Xie L, Wang GJBerberine suppresses intestinal disaccharidases with beneficial metabolic effects in diabetic states, evidences from in vivo and in vitro studyNaunyn Schmiedebergs Arch Pharmacol.(2010 Apr)
      169.^Liu L, Deng Y, Yu S, Lu S, Xie L, Liu XBerberine attenuates intestinal disaccharidases in streptozotocin-induced diabetic ratsPharmazie.(2008 May)
      173.^Zhang H, Wei J, Xue R, Wu JD, Zhao W, Wang ZZ, Wang SK, Zhou ZX, Song DQ, Wang YM, Pan HN, Kong WJ, Jiang JDBerberine lowers blood glucose in type 2 diabetes mellitus patients through increasing insulin receptor expressionMetabolism.(2010 Feb)
      174.^Pérez-Rubio KG, González-Ortiz M, Martínez-Abundis E, Robles-Cervantes JA, Espinel-Bermúdez MCEffect of berberine administration on metabolic syndrome, insulin sensitivity, and insulin secretionMetab Syndr Relat Disord.(2013 Oct)
      177.^Ma X, Egawa T, Kimura H, Karaike K, Masuda S, Iwanaka N, Hayashi TBerberine-induced activation of 5'-adenosine monophosphate-activated protein kinase and glucose transport in rat skeletal musclesMetabolism.(2010 Nov)
      178.^Gomes AP, Duarte FV, Nunes P, Hubbard BP, Teodoro JS, Varela AT, Jones JG, Sinclair DA, Palmeira CM, Rolo APBerberine protects against high fat diet-induced dysfunction in muscle mitochondria by inducing SIRT1-dependent mitochondrial biogenesisBiochim Biophys Acta.(2012 Feb)
      179.^Liu LZ, Cheung SC, Lan LL, Ho SK, Xu HX, Chan JC, Tong PCBerberine modulates insulin signaling transduction in insulin-resistant cellsMol Cell Endocrinol.(2010 Apr 12)
      180.^Kong WJ, Zhang H, Song DQ, Xue R, Zhao W, Wei J, Wang YM, Shan N, Zhou ZX, Yang P, You XF, Li ZR, Si SY, Zhao LX, Pan HN, Jiang JDBerberine reduces insulin resistance through protein kinase C-dependent up-regulation of insulin receptor expressionMetabolism.(2009 Jan)
      181.^Krawiec BJ, Nystrom GJ, Frost RA, Jefferson LS, Lang CHAMP-activated protein kinase agonists increase mRNA content of the muscle-specific ubiquitin ligases MAFbx and MuRF1 in C2C12 cellsAm J Physiol Endocrinol Metab.(2007 Jun)
      182.^Wang XH, Jiang SM, Sun QWEffects of berberine on human rheumatoid arthritis fibroblast-like synoviocytesExp Biol Med (Maywood).(2011 Jul 1)
      184.^Hu PF, Chen WP, Tang JL, Bao JP, Wu LDProtective effects of berberine in an experimental rat osteoarthritis modelPhytother Res.(2011 Jun)
      185.^Ivanovska N, Philipov S, Hristova MInfluence of berberine on T-cell mediated immunityImmunopharmacol Immunotoxicol.(1999 Nov)
      186.^Yan F, Wang L, Shi Y, Cao H, Liu L, Washington MK, Chaturvedi R, Israel DA, Cao H, Wang B, Peek RM Jr, Wilson KT, Polk DBBerberine promotes recovery of colitis and inhibits inflammatory responses in colonic macrophages and epithelial cells in DSS-treated miceAm J Physiol Gastrointest Liver Physiol.(2012 Mar 1)
      188.^Kuo CL, Chi CW, Liu TYThe anti-inflammatory potential of berberine in vitro and in vivoCancer Lett.(2004 Jan 20)
      189.^Park JE, Cuong TD, Hung TM, Lee I, Na M, Kim JC, Ryoo S, Lee JH, Choi JS, Woo MH, Min BSAlkaloids from Chelidonium majus and their inhibitory effects on LPS-induced NO production in RAW264.7 cellsBioorg Med Chem Lett.(2011 Dec 1)
      192.^Kim BH, Kim M, Yin CH, Jee JG, Sandoval C, Lee H, Bach EA, Hahm DH, Baeg GHInhibition of the signalling kinase JAK3 alleviates inflammation in monoarthritic ratsBr J Pharmacol.(2011 Sep)
      193.^Papageorgiou AC, Wikman LEIs JAK3 a new drug target for immunomodulation-based therapiesTrends Pharmacol Sci.(2004 Nov)
      194.^Malaviya R, Zhu D, Dibirdik I, Uckun FMTargeting Janus kinase 3 in mast cells prevents immediate hypersensitivity reactions and anaphylaxisJ Biol Chem.(1999 Sep 17)
      196.^Schröder M, Kaufman RJThe mammalian unfolded protein responseAnnu Rev Biochem.(2005)
      197.^Chen TC, Lai KC, Yang JS, Liao CL, Hsia TC, Chen GW, Lin JJ, Lin HJ, Chiu TH, Tang YJ, Chung JGInvolvement of reactive oxygen species and caspase-dependent pathway in berberine-induced cell cycle arrest and apoptosis in C6 rat glioma cellsInt J Oncol.(2009 Jun)
      199.^Lin JP, Yang JS, Chang NW, Chiu TH, Su CC, Lu KW, Ho YT, Yeh CC, Mei-Dueyang, Lin HJ, Chung JGGADD153 mediates berberine-induced apoptosis in human cervical cancer Ca ski cellsAnticancer Res.(2007 Sep-Oct)
      200.^Lin CC, Yang JS, Chen JT, Fan S, Yu FS, Yang JL, Lu CC, Kao MC, Huang AC, Lu HF, Chung JGBerberine induces apoptosis in human HSC-3 oral cancer cells via simultaneous activation of the death receptor-mediated and mitochondrial pathwayAnticancer Res.(2007 Sep-Oct)
      204.^Zha W, Liang G, Xiao J, Studer EJ, Hylemon PB, Pandak WM Jr, Wang G, Li X, Zhou HBerberine inhibits HIV protease inhibitor-induced inflammatory response by modulating ER stress signaling pathways in murine macrophagesPLoS One.(2010 Feb 9)
      205.^Zhou J, Zhou S, Tang J, Zhang K, Guang L, Huang Y, Xu Y, Ying Y, Zhang L, Li DProtective effect of berberine on beta cells in streptozotocin- and high-carbohydrate/high-fat diet-induced diabetic ratsEur J Pharmacol.(2009 Mar 15)
      206.^Liu J, He C, Zhou K, Wang J, Kang JXCoptis extracts enhance the anticancer effect of estrogen receptor antagonists on human breast cancer cellsBiochem Biophys Res Commun.(2009 Jan 9)
      207.^Doyle ME, Egan JMMechanisms of action of glucagon-like peptide 1 in the pancreasPharmacol Ther.(2007 Mar)
      208.^Meier JJ, Nauck MAGlucagon-like peptide 1(GLP-1) in biology and pathologyDiabetes Metab Res Rev.(2005 Mar-Apr)
      210.^Shou-Si Lu, Yun-Li Yu, Hao-Jie Zhu1, Xiao-Dong Liu, Li Liu, Yao-Wu Liu, Ping Wang, Lin Xie and Guang-Ji WangJ Endocrinol.(2008)
      211.^Li J1, Gu L, Zhang H, Liu T, Tian D, Zhou M, Zhou SBerberine represses DAXX gene transcription and induces cancer cell apoptosisLab Invest.(2013 Mar)
      212.^Marverti G1, Ligabue A, Lombardi P, Ferrari S, Monti MG, Frassineti C, Costi MPModulation of the expression of folate cycle enzymes and polyamine metabolism by berberine in cisplatin-sensitive and -resistant human ovarian cancer cellsInt J Oncol.(2013 Oct)
      213.^Tsang CM, Cheung YC, Lui VW, Yip YL, Zhang G, Lin VW, Cheung KC, Feng Y1, Tsao SWBerberine suppresses tumorigenicity and growth of nasopharyngeal carcinoma cells by inhibiting STAT3 activation induced by tumor associated fibroblastsBMC Cancer.(2013 Dec 31)
      215.^Fu L1, Chen W, Guo W, Wang J, Tian Y, Shi D, Zhang X, Qiu H, Xiao X, Kang T, Huang W, Wang S, Deng WBerberine Targets AP-2/hTERT, NF-κB/COX-2, HIF-1α/VEGF and Cytochrome-c/Caspase Signaling to Suppress Human Cancer Cell GrowthPLoS One.(2013 Jul 15)
      216.^Ho YT1, Lu CC, Yang JS, Chiang JH, Li TC, Ip SW, Hsia TC, Liao CL, Lin JG, Wood WG, Chung JGBerberine induced apoptosis via promoting the expression of caspase-8, -9 and -3, apoptosis-inducing factor and endonuclease G in SCC-4 human tongue squamous carcinoma cancer cellsAnticancer Res.(2009 Oct)
      219.^Eisenberg-Lerner A1, Kimchi AThe paradox of autophagy and its implication in cancer etiology and therapyApoptosis.(2009 Apr)
      220.^Wang N1, Feng Y, Zhu M, Tsang CM, Man K, Tong Y, Tsao SWBerberine induces autophagic cell death and mitochondrial apoptosis in liver cancer cells: the cellular mechanismJ Cell Biochem.(2010 Dec 15)
      223.^Tong N, Zhang J, Chen Y, Li Z, Luo Y, Zuo H, Zhao XBerberine sensitizes mutliple human cancer cells to the anticancer effects of doxorubicin in vitroOncol Lett.(2012 Jun)
      224.^Zhao X, Zhang J, Tong N, Chen Y, Luo YProtective effects of berberine on doxorubicin-induced hepatotoxicity in miceBiol Pharm Bull.(2012)
      225.^Choi MS1, Yuk DY, Oh JH, Jung HY, Han SB, Moon DC, Hong JTBerberine inhibits human neuroblastoma cell growth through induction of p53-dependent apoptosisAnticancer Res.(2008 Nov-Dec)
      226.^Eom KS1, Hong JM, Youn MJ, So HS, Park R, Kim JM, Kim TYBerberine induces G1 arrest and apoptosis in human glioblastoma T98G cells through mitochondrial/caspases pathwayBiol Pharm Bull.(2008 Apr)
      227.^Patil JB1, Kim J, Jayaprakasha GKBerberine induces apoptosis in breast cancer cells (MCF-7) through mitochondrial-dependent pathwayEur J Pharmacol.(2010 Oct 25)
      230.^Park KS, Kim JB, Bae J, Park SY, Jee HG, Lee KE, Youn YKBerberine inhibited the growth of thyroid cancer cell lines 8505C and TPC1Yonsei Med J.(2012 Mar)
      231.^Wang L, Liu L, Shi Y, Cao H, Chaturvedi R, Calcutt MW, Hu T, Ren X, Wilson KT, Polk DB, Yan FBerberine induces caspase-independent cell death in colon tumor cells through activation of apoptosis-inducing factorPLoS One.(2012)
      232.^Choi MS1, Oh JH, Kim SM, Jung HY, Yoo HS, Lee YM, Moon DC, Han SB, Hong JTBerberine inhibits p53-dependent cell growth through induction of apoptosis of prostate cancer cellsInt J Oncol.(2009 May)
      234.^Letasiová S1, Jantová S, Cipák L, Múcková MBerberine-antiproliferative activity in vitro and induction of apoptosis/necrosis of the U937 and B16 cellsCancer Lett.(2006 Aug 8)
      236.^Xia X, Yan J, Shen Y, Tang K, Yin J, Zhang Y, Yang D, Liang H, Ye J, Weng JBerberine improves glucose metabolism in diabetic rats by inhibition of hepatic gluconeogenesisPLoS One.(2011 Feb 3)
      238.^Cameron J, Ranheim T, Kulseth MA, Leren TP, Berge KEBerberine decreases PCSK9 expression in HepG2 cellsAtherosclerosis.(2008 Dec)
      239.^Zhang BJ, Xu D, Guo Y, Ping J, Chen LB, Wang HProtection by and anti-oxidant mechanism of berberine against rat liver fibrosis induced by multiple hepatotoxic factorsClin Exp Pharmacol Physiol.(2008 Mar)
      240.^Sun X, Zhang X, Hu H, Lu Y, Chen J, Yasuda K, Wang HBerberine inhibits hepatic stellate cell proliferation and prevents experimental liver fibrosisBiol Pharm Bull.(2009 Sep)
      241.^Wang N, Feng Y, Cheung F, Chow OY, Wang X, Su W, Tong YA comparative study on the hepatoprotective action of bear bile and coptidis rhizoma aqueous extract on experimental liver fibrosis in ratsBMC Complement Altern Med.(2012 Nov 29)
      242.^Chang X, Yan H, Fei J, Jiang M, Zhu H, Lu D, Gao XBerberine reduces methylation of the MTTP promoter and alleviates fatty liver induced by a high-fat diet in ratsJ Lipid Res.(2010 Sep)
      243.^Wetterau JR, Lin MC, Jamil HMicrosomal triglyceride transfer proteinBiochim Biophys Acta.(1997 Apr 1)
      244.^Lu SS, Yu YL, Zhu HJ, Liu XD, Liu L, Liu YW, Wang P, Xie L, Wang GJBerberine promotes glucagon-like peptide-1 (7-36) amide secretion in streptozotocin-induced diabetic ratsJ Endocrinol.(2009 Feb)
      246.^Yuan J, Shen XZ, Zhu XSEffect of berberine on transit time of human small intestineZhongguo Zhong Xi Yi Jie He Za Zhi.(1994 Dec)
      247.^Strober W, Fuss I, Mannon PThe fundamental basis of inflammatory bowel diseaseJ Clin Invest.(2007 Mar)
      248.^Xavier RJ, Podolsky DKUnravelling the pathogenesis of inflammatory bowel diseaseNature.(2007 Jul 26)
      252.^Hong T, Yang Z, Lv CF, Zhang YSuppressive effect of berberine on experimental dextran sulfate sodium-induced colitisImmunopharmacol Immunotoxicol.(2012 Jun)
      253.^Zhang X, Zhao Y, Zhang M, Pang X, Xu J, Kang C, Li M, Zhang C, Zhang Z, Zhang Y, Li X, Ning G, Zhao LStructural changes of gut microbiota during berberine-mediated prevention of obesity and insulin resistance in high-fat diet-fed ratsPLoS One.(2012)
      254.^Jiang XW, Zhang Y, Zhu YL, Zhang H, Lu K, Li FF, Peng HYEffects of berberine gelatin on recurrent aphthous stomatitis: a randomized, placebo-controlled, double-blind trial in a Chinese cohortOral Surg Oral Med Oral Pathol Oral Radiol.(2012 Dec 12)
      255.^Gao N, Zhao TY, Li XJThe protective effect of berberine on β-cell lipoapoptosisJ Endocrinol Invest.(2011 Feb)
      258.^Ko BS, Choi SB, Park SK, Jang JS, Kim YE, Park SInsulin sensitizing and insulinotropic action of berberine from Cortidis rhizomaBiol Pharm Bull.(2005 Aug)
      259.^Yin J, Hu R, Chen M, Tang J, Li F, Yang Y, Chen JEffects of berberine on glucose metabolism in vitroMetabolism.(2002 Nov)
      262.^Tsuboi T, da Silva Xavier G, Holz GG, Jouaville LS, Thomas AP, Rutter GAGlucagon-like peptide-1 mobilizes intracellular Ca2+ and stimulates mitochondrial ATP synthesis in pancreatic MIN6 beta-cellsBiochem J.(2003 Jan 15)
      263.^Rayasam GV, Tulasi VK, Sundaram S, Singh W, Kant R, Davis JA, Saini KS, Ray AIdentification of berberine as a novel agonist of fatty acid receptor GPR40Phytother Res.(2010 Aug)
      264.^Abdel-Haq H, Cometa MF, Palmery M, Leone MG, Silvestrini B, Saso LRelaxant effects of Hydrastis canadensis L. and its major alkaloids on guinea pig isolated tracheaPharmacol Toxicol.(2000 Nov)
      266.^Tang LQ, Wang FL, Zhu LN, Lv F, Liu S, Zhang STBerberine ameliorates renal injury by regulating G proteins-AC- cAMP signaling in diabetic rats with nephropathyMol Biol Rep.(2012 Dec 25)
      267.^Wu D, Wen W, Qi CL, Zhao RX, Lü JH, Zhong CY, Chen YYAmeliorative effect of berberine on renal damage in rats with diabetes induced by high-fat diet and streptozotocinPhytomedicine.(2012 Jun 15)
      268.^Liu W, Tang F, Deng Y, Li X, Lan T, Zhang X, Huang H, Liu PBerberine reduces fibronectin and collagen accumulation in rat glomerular mesangial cells cultured under high glucose conditionMol Cell Biochem.(2009 May)
      269.^Jiang Q, Liu P, Wu X, Liu W, Shen X, Lan T, Xu S, Peng J, Xie X, Huang HBerberine attenuates lipopolysaccharide-induced extracelluar matrix accumulation and inflammation in rat mesangial cells: involvement of NF-κB signaling pathwayMol Cell Endocrinol.(2011 Jan 1)
      270.^Liu W, Zhang X, Liu P, Shen X, Lan T, Li W, Jiang Q, Xie X, Huang HEffects of berberine on matrix accumulation and NF-kappa B signal pathway in alloxan-induced diabetic mice with renal injuryEur J Pharmacol.(2010 Jul 25)
      272.^Mogyorósi A, Kapoor A, Isono M, Kapoor S, Sharma K, Ziyadeh FNUtility of serum and urinary transforming growth factor-beta levels as markers of diabetic nephropathyNephron.(2000 Oct)
      273.^Lan T, Liu W, Xie X, Huang K, Peng J, Huang J, Shen X, Liu P, Yang H, Huang HBerberine suppresses high glucose-induced TGF-β1 and fibronectin synthesis in mesangial cells through inhibition of sphingosine kinase 1/AP-1 pathwayEur J Pharmacol.(2012 Dec 15)
      274.^Taha TA, Hannun YA, Obeid LMSphingosine kinase: biochemical and cellular regulation and role in diseaseJ Biochem Mol Biol.(2006 Mar 31)
      275.^Lan T, Liu W, Xie X, Xu S, Huang K, Peng J, Shen X, Liu P, Wang L, Xia P, Huang HSphingosine kinase-1 pathway mediates high glucose-induced fibronectin expression in glomerular mesangial cellsMol Endocrinol.(2011 Dec)
      276.^Lan T, Shen X, Liu P, Liu W, Xu S, Xie X, Jiang Q, Li W, Huang HBerberine ameliorates renal injury in diabetic C57BL/6 mice: Involvement of suppression of SphK-S1P signaling pathwayArch Biochem Biophys.(2010 Oct 15)
      277.^Liu W, Liu P, Tao S, Deng Y, Li X, Lan T, Zhang X, Guo F, Huang W, Chen F, Huang H, Zhou SFBerberine inhibits aldose reductase and oxidative stress in rat mesangial cells cultured under high glucoseArch Biochem Biophys.(2008 Jul 15)
      278.^Liu WH, Hei ZQ, Nie H, Tang FT, Huang HQ, Li XJ, Deng YH, Chen SR, Guo FF, Huang WG, Chen FY, Liu PQBerberine ameliorates renal injury in streptozotocin-induced diabetic rats by suppression of both oxidative stress and aldose reductaseChin Med J (Engl).(2008 Apr 20)
      279.^Bashir S, Gilani AHAntiurolithic effect of berberine is mediated through multiple pathwaysEur J Pharmacol.(2011 Jan 25)
      281.^Bashir S, Gilani AH, Siddiqui AA, Pervez S, Khan SR, Sarfaraz NJ, Shah AJBerberis vulgaris root bark extract prevents hyperoxaluria induced urolithiasis in ratsPhytother Res.(2010 Aug)
      283.^Namba T, Sekiya K, Toshinal A, Kadota S, Hatanaka T, Katayama K, Koizumi TStudy on baths with crude drug. II.: the effects of coptidis rhizoma extracts as skin permeation enhancerYakugaku Zasshi.(1995 Aug)
      284.^Inbaraj JJ, Kukielczak BM, Bilski P, Sandvik SL, Chignell CFPhotochemistry and photocytotoxicity of alkaloids from Goldenseal (Hydrastis canadensis L.) 1. BerberineChem Res Toxicol.(2001 Nov)
      286.^Tan Y, Ming Z, Tang Q, Jiang Z, Hu B, Xiang JEffect of berberine on the mRNA expression of nitric oxide synthase (NOS) in rat corpus cavernosumJ Huazhong Univ Sci Technolog Med Sci.(2005)
      289.^Tan Y, Tang Q, Hu BR, Xiang JZEffects of berberine on cyclic GMP and cyclic AMP levels in rabbit corpus cavernosum in vitroZhonghua Nan Ke Xue.(2005 Jun)
      290.^Drewes SE, George J, Khan FRecent findings on natural products with erectile-dysfunction activityPhytochemistry.(2003 Apr)
      293.^Wei W, Zhao H, Wang A, Sui M, Liang K, Deng H, Ma Y, Zhang Y, Zhang H, Guan YA clinical study on the short-term effect of berberine in comparison to metformin on the metabolic characteristics of women with polycystic ovary syndromeEur J Endocrinol.(2012 Jan)
      294.^Zhao W, Xue R, Zhou ZX, Kong WJ, Jiang JDReduction of blood lipid by berberine in hyperlipidemic patients with chronic hepatitis or liver cirrhosisBiomed Pharmacother.(2008 Dec)
      297.^Vogel RAPCSK9 inhibition: the next statinJ Am Coll Cardiol.(2012 Jun 19)
      298.^Rashid S, Curtis DE, Garuti R, Anderson NN, Bashmakov Y, Ho YK, Hammer RE, Moon YA, Horton JDDecreased plasma cholesterol and hypersensitivity to statins in mice lacking Pcsk9Proc Natl Acad Sci U S A.(2005 Apr 12)
      299.^Welder G, Zineh I, Pacanowski MA, Troutt JS, Cao G, Konrad RJHigh-dose atorvastatin causes a rapid sustained increase in human serum PCSK9 and disrupts its correlation with LDL cholesterolJ Lipid Res.(2010 Sep)
      300.^Dubuc G, Chamberland A, Wassef H, Davignon J, Seidah NG, Bernier L, Prat AStatins upregulate PCSK9, the gene encoding the proprotein convertase neural apoptosis-regulated convertase-1 implicated in familial hypercholesterolemiaArterioscler Thromb Vasc Biol.(2004 Aug)
      301.^Kong WJ, Wei J, Zuo ZY, Wang YM, Song DQ, You XF, Zhao LX, Pan HN, Jiang JDCombination of simvastatin with berberine improves the lipid-lowering efficacyMetabolism.(2008 Aug)
      302.^Carlomagno G, Pirozzi C, Mercurio V, Ruvolo A, Fazio SEffects of a nutraceutical combination on left ventricular remodeling and vasoreactivity in subjects with the metabolic syndromeNutr Metab Cardiovasc Dis.(2012 May)
      304.^Marazzi G, Cacciotti L, Pelliccia F, Iaia L, Volterrani M, Caminiti G, Sposato B, Massaro R, Grieco F, Rosano GLong-term effects of nutraceuticals (berberine, red yeast rice, policosanol) in elderly hypercholesterolemic patientsAdv Ther.(2011 Dec)
      306.^Liu J, Zhang J, Shi Y, Grimsgaard S, Alraek T, Fønnebø VChinese red yeast rice (Monascus purpureus) for primary hyperlipidemia: a meta-analysis of randomized controlled trialsChin Med.(2006 Nov 23)
      307.^Zhou S, Lim LY, Chowbay BHerbal modulation of P-glycoproteinDrug Metab Rev.(2004 Feb)
      308.^Xue L, Jiao L, Wang Y, Nie Y, Han T, Jiang Y, Rahman K, Zhang Q, Qin LEffects and interaction of icariin, curculigoside, and berberine in er-xian decoction, a traditional chinese medicinal formula, on osteoclastic bone resorptionEvid Based Complement Alternat Med.(2012)
      309.^Vrzal R, Zdarilová A, Ulrichová J, Bláha L, Giesy JP, Dvorák ZActivation of the aryl hydrocarbon receptor by berberine in HepG2 and H4IIE cells: Biphasic effect on CYP1A1Biochem Pharmacol.(2005 Sep 15)
      310.^Zhi D, Feng PF, Sun JL, Guo F, Zhang R, Zhao X, Li BXThe enhancement of cardiac toxicity by concomitant administration of Berberine and macrolidesEur J Pharm Sci.(2015 Aug 30)