. 2025 Feb 7;13:e18924. doi: 10.7717/peerj.18924

Table 3. Studies investigating the effects of berberine on browning.

	Dose	Duration	Suggested Pathway	Results	References
3T3-L1 preadipocytes	0.5, 1, 5, 10 µM	7 days	cAMP/PKA	Adipogenic genes (C/EBP-α, PPARγ) ↓, CREB activity↓	Zhang et al. (2015)
HepG2 cells	5, 10, 15 µM	N/A	AMPK	AMPK phosphorylation↑, ACC↑, fatty acid oxidation↑	Brusq et al. (2006)
Male Syrian golden hamsters	100 mg/kg/day	10 days	AMPK	AMPK phosphorylation↑, ACC↑, fatty acid oxidation↑	Brusq et al. (2006)
Male C57BL/6J mice	50 and 100 mg/kg/day	14 days	GDF15	Serum GDF15↑, GFRAL↑, appetite↓, UCP1↑	Li et al. (2023)
Obese C57BLKS/J-Lepr^db/Lepr^db male mice and wild-type mice	5 mg/kg/day	4 weeks	AMPK- PGC1α	Energy expenditure↑, weight gain↓, BAT activity↑, UCP1↑, PGC-1α↑, CIDEA↑	Zhang et al. (2014c)
Male C57BL/6J mice	100 mg/kg/day	10 weeks	AMPK	AMPK↑, complex I↓, AMP/ATP↑, ADP/ATP↑	Turner et al. (2008)
Male Wistar rats	100 mg/kg/day	4 weeks	AMPK	AMPK↑, complex I↓, AMP/ATP↑, ADP/ATP↑	Turner et al. (2008)
Female Sprague-Dawley rats	380 mg/kg/day	2 weeks	N/A	Olanzapine-induce BAT loss↓, weight gain↓, adiposity↓, AMPK↑, UCP1↑, PGC1α↑. Food intake did not change.	Hu et al. (2014)
Male C57BL/6J mice	25 and 100 mg/kg/day	12 weeks	AMPK-SIRT1	Distribution of BAT↑, thermogenesis↑, body weight↓, AMPK/SIRT1 activation↑, PPAR↑ deacetylation↑, UCP1 expression↑	Xu et al. (2021)
Obese C57BLKS/J-Lepr^db/Lepr^db male mice	5 mg/kg/day	26 days	AMPK	Lipogenesis (FAS, PPARγ)↓, expression of browning markers (PGC1α)↑, AMPK activation↑, body weight↓. Food intake did not change.	Lee et al. (2006)
Wistar rats	380 mg/kg/day	2 weeks	AMPK		Lee et al. (2006)
Female Sprague-Dawley rats	380 mg/kg/day	2 weeks	N/A	Blood lipid levels↓, weight loss↑	Hu et al. (2012)
Obese humans	1.5 g/day	12 weeks	N/A	Blood lipid levels↓, weight loss↑	Hu et al. (2012)
Male C57BL/6J mice	1.5 mg/kg/day	6 weeks	AMPK–PRDM16	Both in mice and humans: Brown adipocyte differentiation↑, PRDM16 transcription↑In mice: thermogenesis↑, energy expenditure↑AMPK is essential for the browning effect of berberine.	Wu et al. (2019)
NAFLD patients	1.5 g/day	1 month	AMPK–PRDM16		Wu et al. (2019)

Note:

ACC, acetyl-CoA carboxylase; ADP, adenosine diphosphate; AMP, adenosine monophosphate; AMPK, AMP-activated protein kinase; ATP, adenosine triphosphate; BAT, brown adipose tissue; cAMP, cyclic adenosine monophosphate; C/EBP-α, CCAAT/enhancer-binding protein alpha; CIDEA, cell death-inducing DNA fragmentation factor-like effector A; CREB, cAMP response element-binding protein; FAS, fatty acid synthetase; GDF15, Growth differentiation factor 15; GFRAL, glial cell-derived neurotrophic factor family receptor alpha-like; PGC1α, PPARγ-co-activator-1α; PKA, protein kinase A; PPARγ, Peroxisome proliferator-activated receptor-γ; PRDM16, PR domain containing 16; SIRT1, Sirtuin 1; UCP1, uncoupling protein 1.