[1]. World Health Organization. 2025. Lung
cancer. WHO Fact Sheet. Retrieved 8 November 2025.
[2]. American Cancer Society. 2025, Lung
Cancer Statistics: How Common Is Lung Cancer? ACS. Retrieved 8 November
2025.
[3]. Boța, M., Vlaia, L., Jîjie, A. R.,
Marcovici, I., Crişan, F., Oancea, C., Dehelean, C. A., Mateescu, T., and
Moacă, E. A., 2024, Exploring synergistic interactions between natural
compounds and conventional chemotherapeutic drugs in preclinical models of lung
cancer. Pharmaceuticals, 17(5), 598.
[4]. Fan, K. Q., Zhang, L., Song, F., Zhang,
Y. H., Chen, T., Cheng, X., Su, N., Zou, Y., Yu, T., Tan, F., Xu, W., and Yan,
Z., 2025, Pharmacological properties and therapeutic potential of berberine: a
comprehensive review. Frontiers
in Pharmacology,
16, 1604071.
[5]. Ni, L., Sun, P., Fan, X., Li, Z., Ren,
H., and Li, J., 2022, Berberine inhibits FOXM1-dependent transcriptional
regulation of POLE2 and interferes with survival of lung adenocarcinoma. Frontiers in Pharmacology, 12, 775514.
[6]. Tamtaji, Z., Sheikhsagha, E., Behnam, M.,
Nabavizadeh, F., Shafiee Ardestani, M., Rahmati-Dehkordi, F., Aschner, M.,
Mirzaei, H., and Tamtaji, O. R., 2025, Berberine and lung cancer: from pure
form to its nanoformulations. Asia-Pacific Journal of Clinical Oncology, 21(5), 455–464.
[7]. Almatroodi, S. A., Alsahli, M. A., and
Rahmani, A. H., 2022, Berberine: an important emphasis on its anticancer
effects through modulation of various cell signaling pathways. Molecules, 27(18).
[8]. Xiong, R. G., Huang, S. Y., Wu, S. X.,
Zhou, D. D., Yang, Z. J., Saimaiti, A., Zhao, C. N., Shang, A., Zhang, Y. J.,
Gan, R. Y., and Li, H. Bin, 2022, Anticancer effects and mechanisms of
berberine from medicinal herbs: an update review. Molecules, 27(14).
[9]. Zhang, C., Sheng, J., Li, G., Zhao, L.,
Wang, Y., Yang, W., Yao, X., Sun, L., Zhang, Z., and Cui, R., 2020, Effects of
berberine and its derivatives on cancer: a systems pharmacology review. Frontiers in Pharmacology, 10, 1461.
[10]. Liu, L., Fan, J., Ai, G., Liu, J., Luo,
N., Li, C., and Cheng, Z., 2019, Berberine in combination with cisplatin
induces necroptosis and apoptosis in ovarian cancer cells. Biological Research, 52(1), 1–14.
[11]. Zhu, M. L., Yin, Y. L., Ping, S., Yu, H.
Y., Wan, G. R., Jian, X., and Li, P., 2017, Berberine promotes ischemia-induced
angiogenesis in mice heart via upregulation of microRNA-29b. Clinical and Experimental Hypertension, 39(7), 672–679.
[12]. Chen, Q. Q., Shi, J. M., Ding, Z., Xia,
Q., Zheng, T. S., Ren, Y. B., Li, M., and Fan, L. H., 2019, Berberine induces
apoptosis in non-small-cell lung cancer cells by upregulating miR-19a targeting
tissue factor. Cancer
Management and Research,
11, 9005–9015.
[13]. Du, W., Zhang, R., Muhammad, B., and Pei,
D., 2022, Targeting the COP9 signalosome for cancer therapy. Cancer Biology & Medicine, 19(5), 573.
[14]. Chen, J., Huang, X., Tao, C., Wang, L.,
Chen, Z., Li, X., Zeng, Q., Ma, M., Zhang, R., and Wu, Z., 2020, Berberine
chloride suppresses non-small cell lung cancer by deregulating Sin3A/TOP2B
pathway in vitro and in vivo. Cancer Chemotherapy and Pharmacology, 86(1), 151–161.
[15]. Gu, W., Zhang, M., Gao, F., Niu, Y., Sun,
L., Xia, H., Li, W., Zhang, Y., Guo, Z., and Du, G., 2022, Berberine regulates
PADI4-related macrophage function to prevent lung cancer. International Immunopharmacology, 110, 108965.
[16]. Wang, Q., Wu, H., Wu, Q., and Zhong, S.,
2023, Berberine targets KIF20A and CCNE2 to inhibit the progression of
non-small cell lung cancer via the PI3K/AKT pathway. Drug Development Research, 84(5), 907–921.
[17]. Liao, W., Zhang, R., Chen, G., Zhu, X.,
Wu, W., Chen, Z., Jiang, C., Lin, Z., Ma, L., and Yu, H., 2024, Berberine
synergises with ferroptosis inducer sensitizing NSCLC to ferroptosis in
p53-dependent SLC7A11-GPX4 pathway. Biomedicine & Pharmacotherapy, 176, 116832.
[18]. Tong, N., Zhang, J., Chen, Y., Li, Z.,
Luo, Y., Zuo, H., and Zhao, X., 2012, Berberine sensitizes multiple human
cancer cells to the anticancer effects of doxorubicin in vitro. Oncology Letters, 3(6), 1263.
[19]. Chen, Z., Vallega, K. A., Chen, H., Zhou,
J., Ramalingam, S. S., and Sun, S. Y., 2022, The natural product berberine
synergizes with osimertinib preferentially against MET-amplified
osimertinib-resistant lung cancer via direct MET inhibition. Pharmacological Research, 175, 105998.
[20]. Zheng, F., Wu, J. J., Tang, Q., Xiao, Q.,
Wu, W. Y., and Hann, S. S., 2018, The enhancement of combination of berberine
and metformin in inhibition of DNMT1 gene expression through interplay of SP1
and PDPK1. Journal
of Cellular and Molecular Medicine, 22(1), 600–612.
[21]. Liu, Y., Liu, X., Zhang, N., Yin, M.,
Dong, J., Zeng, Q., Mao, G., Song, D., Liu, L., and Deng, H., 2020, Berberine
diminishes cancer cell PD-L1 expression and facilitates antitumor immunity via
inhibiting the deubiquitination activity of CSN5. Acta Pharmaceutica Sinica B, 10(12), 2299–2312.
[22]. Zhang, L., Zhou, C., Zhang, S., Chen, X.,
Liu, J., Xu, F., and Liang, W., 2022, Chemotherapy reinforces anti-tumor immune
response and enhances clinical efficacy of immune checkpoint inhibitors. Frontiers in Oncology, 12, 939249.
[23]. Sun, P., Wang, Z., Ma, Y., Liu,Y., Xue,
Y., Li, Y., Gao, X., Wang, Y., and Chu, M., 2025, Advance in identified targets
of berberine. Frontiers
in Pharmacology,
16, 1500511.
[24]. Katiyar, S. K., Meeran, S. M., Katiyar,
N., and Akhtar, S., 2009, p53 cooperates in berberine-induced growth inhibition
and apoptosis of non-small cell human lung cancer cells in vitro and tumor
xenograft growth in vivo. Molecular Carcinogenesis, 48(1), 24–37.
[25]. James, M. A., Fu, H., Liu, Y., Chen, D.
R., and You, M., 2011, Dietary administration of berberine or Phellodendron amurense extract inhibits cell cycle progression
and lung tumorigenesis. Molecular
Carcinogenesis,
50(1), 1–7.
[26]. Nie, P., Zhao, Y., and Xu, H., 2023,
Synthesis, applications, toxicity and toxicity mechanisms of silver
nanoparticles: a review. Ecotoxicology
and Environmental Safety,
253, 114636.
[27]. Qi, H. W., Xin, L. Y., Xu, X., Ji, X. X.,
and Fan, L. H., 2014,
Epithelial-to-mesenchymal transition markers to predict response of berberine
in suppressing lung cancer invasion and metastasis. Journal of Translational Medicine, 12, 22.
[28]. Li, J., Liu, F., Jiang, S., Liu, J.,
Chen, X., Zhang, S., and Zhao, H., 2018, Berberine hydrochloride inhibits cell proliferation
and promotes apoptosis of non-small cell lung cancer via the suppression of the
MMP2 and Bcl-2/Bax signaling pathways. Oncology Letters, 15, 7409–7414.
[29]. Xi, S., Chuang, K., Fang, K., Lee, Y.,
Chung, J., and Chuang, Y., 2014, Effect of berberine on activity and mRNA expression of
N-acetyltransferase in human lung cancer cell line A549. Journal of Traditional Chinese Medicine, 34(3),
302–308.
[30]. Zheng, F., Tang, Q., Wu, J., Zhao, S.,
Liang, Z., Li, L., Wu, W., and Hann, S., 2014, p38α MAPK-mediated induction and
interaction of FOXO3a and p53 contribute to the inhibited growth and induced
apoptosis of human lung adenocarcinoma cells by berberine. Journal of Experimental & Clinical
Cancer Research, 33(1), 36.
[31]. Fu, L., Chen, W., Guo, W., Wang, J.,
Tian, Y., Shi, D., Zhang, X., Qiu, H., Xiao, X., Kang, T., et al., 2013, Berberine targets AP-2/hTERT, NF-κB/COX-2,
HIF-1α/VEGF and cytochrome-c/caspase signaling to suppress human cancer cell
growth. PLoS ONE, 8,
e69240.
[32]. Meng, M., Geng, S., Du, Z., Yao, J.,
Zheng, Y., Li, Z., Zhang, Z., Li, J., Duan, Y., and Du, G., 2017, Berberine and cinnamaldehyde together
prevent lung carcinogenesis. Oncotarget, 8(44),
76385–76397.
[33]. Lu, J.J., Fu, L., Tang, Z., Zhang, C.,
Qin, L., Wang, J., Yu, Z., Shi, D., Xiao, X., Xie, F., Huang, W., and Deng, W.,
2016, Melatonin
inhibits AP-2β/hTERT, NF-κB/COX-2 and Akt/ERK and activates caspase/Cyto C
signaling to enhance the antitumor activity of berberine in lung cancer cells. Oncotarget, 7(3), 2985–3001.
[34]. Shen, F., Zheng, Y. S., Dong, L., Cao,
Z., and Cao, J., 2024, Enhanced
tumor suppression in colorectal cancer via berberine-loaded PEG-PLGA
nanoparticles. Frontiers in
Pharmacology, 15, 1500731.
[35]. Peng, P. L., Hsieh,
Y. S., Wang, C.J., Hsu, J. L., and Chou, F. P.,
2006, Inhibitory
effect of berberine on the invasion of human lung cancer cells via decreased
production of urokinase-plasminogen activator and matrix metalloproteinase-2. Toxicology and Applied Pharmacology, 214(1),
8–15.
[36]. Zhu, T., Li, L. L., Xiao, G. F., Luo, Q. Z.,
Liu, Q.Z ., Yao, K. T., and Xiao, G. H., 2015, Berberine increases doxorubicin sensitivity by
suppressing STAT3 in lung cancer. Journal
of Bioenergetics and Biomembranes, 43(7), 1487–1502.
