van Hoogstraten LMC, Vrieling A, van der Heijden AG, Kogevinas M, Richters A, Kiemeney LA. Global trends in the epidemiology of bladder cancer: challenges for public health and clinical practice. Nat Rev Clin Oncol. 2023;20:287–304.
Google Scholar
Xu W, Liang T, Fang H, Fu L, Deng D, Tan X, et al. Single-cell RNA sequencing identifies MMP11(+) cancer-associated fibroblasts as drivers of angiogenesis and bladder cancer progression. Adv Sci. 2025;12:e02774.
Google Scholar
Li F, Zheng Z, Chen W, Li D, Zhang H, Zhu Y, et al. Regulation of cisplatin resistance in bladder cancer by epigenetic mechanisms. Drug Resist Updat. 2023;68:100938.
Google Scholar
Zhao H, Lin N, Ho VWS, Liu K, Chen X, Wu H, et al. Patient-derived bladder cancer organoids as a valuable tool for understanding tumor biology and developing personalized treatment. Adv Sci. 2025;12:e2414558.
Google Scholar
Zhao Y, Xing Z, Zhao Y, Xu H, Liu R, Yang T, et al. Lactylation prognostic signature identifies DHCR7 as a modulator of chemoresistance and immunotherapy efficacy in bladder cancer. Front Immunol. 2025;16:1585727.
Google Scholar
Deng M, Zhou Z, Chen J, Li X, Liu Z, Ye J, et al. Enhanced oxidative phosphorylation driven by TACO1 mitochondrial translocation promotes stemness and cisplatin resistance in bladder cancer. Adv Sci. 2025;12:e2408599.
Google Scholar
Lei Y, Tang R, Xu J, Wang W, Zhang B, Liu J, et al. Applications of single-cell sequencing in cancer research: progress and perspectives. J Hematol Oncol. 2021;14:91.
Google Scholar
Ward PS, Thompson CB. Metabolic reprogramming: a cancer hallmark even warburg did not anticipate. Cancer Cell. 2012;21:297–308.
Google Scholar
Pavlova NN, Zhu J, Thompson CB. The hallmarks of cancer metabolism: still emerging. Cell Metab. 2022;34:355–77.
Google Scholar
Zhao X, Guo B, Sun W, Yu J, Cui L. Targeting squalene epoxidase confers metabolic vulnerability and overcomes chemoresistance in HNSCC. Adv Sci. 2023;10:e2206878.
Google Scholar
Cerqueira NM, Oliveira EF, Gesto DS, Santos-Martins D, Moreira C, Moorthy HN, et al. Cholesterol biosynthesis: a mechanistic overview. Biochemistry. 2016;55:5483–506.
Google Scholar
Liu X, Bao X, Hu M, Chang H, Jiao M, Cheng J, et al. Inhibition of PCSK9 potentiates immune checkpoint therapy for cancer. Nature. 2020;588:693–8.
Google Scholar
Zeng Y, Luo Y, Zhao K, Liu S, Wu K, Wu Y, et al. m6A-mediated induction of 7-dehydrocholesterol reductase stimulates cholesterol synthesis and cAMP signaling to promote bladder cancer metastasis. Cancer Res. 2024;84:3402–18.
Google Scholar
Tan L, Zhou H, Zhang B, Kwong DLW, Jia Y, Huang J, et al. Multiomics identifies a cholesterol-TFEB-PLD3-TLR9 axis driving immunosuppressive tumor-associated macrophage polarization in esophageal squamous cell carcinoma. Proc Natl Acad Sci USA. 2026;123:e2520427123.
Google Scholar
Hu C, Qiao W, Li X, Ning ZK, Liu J, Dalangood S, et al. Tumor-secreted FGF21 acts as an immune suppressor by rewiring cholesterol metabolism of CD8( + )T cells. Cell Metab. 2024;36:630–47.e638.
Google Scholar
Tang L, Wei R, Chen R, Fan G, Zhou J, Qi Z, et al. Establishment and validation of a cholesterol metabolism-related prognostic signature for hepatocellular carcinoma. Comput Struct Biotechnol J. 2022;20:4402–14.
Google Scholar
Sun R, Wang X, Wang Z, Li C, Shao Q, Liu X, et al. Transcriptomic insights into Alzheimer’s disease: differentially expressed genes and cholesterol metabolism. CNS Neurosci Ther. 2026;32:e70833.
Google Scholar
Xu M, Hu J, Pu L, Liu J, Yang Y, Li Q, et al. Multi-omics integration identifies the cholesterol metabolic enzyme DHCR24 as a key driver in breast cancer. Biology (Basel). 2025;15:40.
Google Scholar
Wang Z, Mao J, Zhang Y, Yang W, Sun D, Lu Z, et al. Pan-cancer analysis reveals the potential role of DHCR24 in bladder cancer via interactions with HRAS to facilitate cholesterol synthesis. Oncol Lett. 2025;30:385.
Google Scholar
Sezgin E, Levental I, Mayor S, Eggeling C. The mystery of membrane organization: composition, regulation and roles of lipid rafts. Nat Rev Mol Cell Biol. 2017;18:361–74.
Google Scholar
Pan Z, Wang K, Wang X, Jia Z, Yang Y, Duan Y, et al. Cholesterol promotes EGFR-TKIs resistance in NSCLC by inducing EGFR/Src/Erk/SP1 signaling-mediated ERRalpha re-expression. Mol Cancer. 2022;21:77.
Google Scholar
Nakajima R, Deguchi R, Komori H, Zhao L, Zhou Y, Shirasawa M, et al. The TFDP1 gene coding for DP1, the heterodimeric partner of the transcription factor E2F, is a target of deregulated E2F. Biochem Biophys Res Commun. 2023;663:154–62.
Google Scholar
Iaquinta PJ, Lees JA. Life and death decisions by the E2F transcription factors. Curr Opin Cell Biol. 2007;19:649–57.
Google Scholar
Trimarchi JM, Lees JA. Sibling rivalry in the E2F family. Nat Rev Mol Cell Biol. 2002;3:11–20.
Google Scholar
Ju G, Lin Q, Lu L, Lin Z, Huang D, Lin Y, et al. TFDP1 drives triple-negative breast cancer development through senescence suppression and serves as a therapeutic target for topotecan. Int J Biol Macromol. 2025;310:143543.
Google Scholar
Comperat E, Amin MB, Cathomas R, Choudhury A, De Santis M, Kamat A, et al. Current best practice for bladder cancer: a narrative review of diagnostics and treatments. Lancet. 2022;400:1712–21.
Google Scholar
Lin P, Xing Z, Hong Y, Xu H, Shao S, Luo L, et al. Integrin beta4 drives immune evasion and therapeutic resistance to PD-1 blockade in bladder cancer via MEK/ERK signaling. J Biol Chem. 2026;302:110941.
Google Scholar
Wang Q, Cao Y, Shen L, Xiao T, Cao R, Wei S, et al. Regulation of PD-L1 through direct binding of cholesterol to CRAC motifs. Sci Adv. 2022;8:eabq4722.
Google Scholar
Du A, Zhou Y, Deng X, Yuan D, Li K, Luo Y, et al. DCLK1 drives malignant progression and chemoresistance of bladder cancer by deubiquitinating HDAC6. Mol Cancer. 2026;25:46.
Google Scholar
Xing Z, Xu H, Lin P, Hong Y, Shao S, Yang T, et al. ITGB4 up-regulated by STAT3 reduces the sensitivity of bladder cancer to cisplatin by suppressing p53. Br J Cancer. 2026;134:1704–15.
Google Scholar
Dianatinasab M, Wesselius A, Salehi-Abargouei A, Yu EYW, Fararouei M, Brinkman M, et al. Dietary fats and their sources in association with the risk of bladder cancer: a pooled analysis of 11 prospective cohort studies. Int J Cancer. 2022;151:44–55.
Google Scholar
Shih HJ, Lin KH, Wen YC, Fan YC, Tsai PS, Huang CJ. Increased risk of bladder cancer in young adult men with hyperlipidemia: a population-based cohort study. Medicine. 2021;100:e28125.
Google Scholar
Li X, Sun Z, Zeng R, Liu R, Lin P, Xing Z, et al. Diet-metabolism-transcription axis modulates the sensitivity to CDK4/6 inhibitors through RB1 in prostate cancer. Cell Rep. 2025;44:116530.
Google Scholar
Xing Z, Yang T, Li X, Xu H, Hong Y, Shao S, et al. High-glucose-associated YTHDC1 lactylation reduces the sensitivity of bladder cancer to enfortumab vedotin therapy. Cell Rep. 2025;44:115545.
Google Scholar
Hanahan D, Weinberg RA. Hallmarks of cancer: the next generation. Cell. 2011;144:646–74.
Google Scholar
Pavlova NN, Thompson CB. The emerging hallmarks of cancer metabolism. Cell Metab. 2016;23:27–47.
Google Scholar
Lu J, Chen S, Bai X, Liao M, Qiu Y, Zheng LL, et al. Targeting cholesterol metabolism in cancer: from molecular mechanisms to therapeutic implications. Biochem Pharmacol. 2023;218:115907.
Google Scholar
Wang X, Lee D, Xu H, Sui Y, Meisenhelder J, Hunter T. PIN1 prolyl isomerase promotes initiation and progression of bladder cancer through the SREBP2-mediated cholesterol biosynthesis pathway. Cancer Discov. 2025;15:633–55.
Google Scholar
Li M, Liu T, Shi J, Zhou F, Deng Z, Luo Y, et al. USP43 promotes gemcitabine resistance by regulating cholesterol homeostasis through E2F1 stabilization in bladder cancer. J Exp Clin Cancer Res. 2025;45:23.
Google Scholar
Wu M, Zhou X, Zhou X, Wang G, Zeng Y, Li J, et al. ZDHHC3-mediated SCAP S-acylation promotes cholesterol biosynthesis and tumor immune escape in hepatocellular carcinoma. Cell Rep. 2024;43:114962.
Google Scholar
Jiang W, Hu JW, He XR, Jin WL, He XY. Statins: a repurposed drug to fight cancer. J Exp Clin Cancer Res. 2021;40:241.
Google Scholar
Symvoulidis P, Tsioutis C, Zamboglou C, Agouridis AP. The effect of statins on the incidence and prognosis of bladder cancer: a systematic review and meta-analysis. Curr Oncol. 2023;30:6648–65.
Google Scholar
Luu W, Zerenturk EJ, Kristiana I, Bucknall MP, Sharpe LJ, Brown AJ. Signaling regulates activity of DHCR24, the final enzyme in cholesterol synthesis. J Lipid Res. 2014;55:410–20.
Google Scholar
Chan SL, Chan AW. Development of serum DHCR24 antibody as a marker for hepatocellular carcinoma: the end of the beginning. EBioMedicine. 2015;2:497–8.
Google Scholar
Romanuik TL, Ueda T, Le N, Haile S, Yong TM, Thomson T, et al. Novel biomarkers for prostate cancer including noncoding transcripts. Am J Pathol. 2009;175:2264–76.
Google Scholar
Qiu T, Cao J, Chen W, Wang J, Wang Y, Zhao L, et al. 24-Dehydrocholesterol reductase promotes the growth of breast cancer stem-like cells through the Hedgehog pathway. Cancer Sci. 2020;111:3653–64.
Google Scholar
Lee GT, Ha YS, Jung YS, Moon SK, Kang HW, Lee OJ, et al. DHCR24 is an independent predictor of progression in patients with non-muscle-invasive urothelial carcinoma, and its functional role is involved in the aggressive properties of urothelial carcinoma cells. Ann Surg Oncol. 2014;21:S538–545.
Google Scholar
Liu XP, Yin XH, Meng XY, Yan XH, Cao Y, Zeng XT, et al. DHCR24 predicts poor clinicopathological features of patients with bladder cancer: a STROBE-compliant study. Medicine. 2018;97:e11830.
Google Scholar
Fu X, Wang Z. DHCR24 in tumor diagnosis and treatment: a comprehensive review. Technol Cancer Res Treat. 2024;23:15330338241259780.
Google Scholar
Greenlee JD, Subramanian T, Liu K, King MR. Rafting down the metastatic cascade: the role of lipid rafts in cancer metastasis, cell death, and clinical outcomes. Cancer Res. 2021;81:5–17.
Google Scholar
Ramseger R, White R, Kroger S. Transmembrane form agrin-induced process formation requires lipid rafts and the activation of Fyn and MAPK. J Biol Chem. 2009;284:7697–705.
Google Scholar
Xu R, Song J, Ruze R, Chen Y, Yin X, Wang C, et al. SQLE promotes pancreatic cancer growth by attenuating ER stress and activating lipid rafts-regulated Src/PI3K/Akt signaling pathway. Cell Death Dis. 2023;14:497.
Google Scholar
Yu S, Wang L, Che D, Zhang M, Li M, Naito M, et al. Targeting CRABP-II overcomes pancreatic cancer drug resistance by reversing lipid raft cholesterol accumulation and AKT survival signaling. J Exp Clin Cancer Res. 2022;41:88.
Google Scholar
Raji L, Tetteh A, Amin A. Role of c-Src in carcinogenesis and drug resistance. Cancers. 2023;16:32.
Google Scholar
Roskoski R Jr. Src protein-tyrosine kinase structure, mechanism, and small molecule inhibitors. Pharmacol Res. 2015;94:9–25.
Google Scholar
Tang Y, Deng X, Wang Y, Zhou Q, Zhang C, Liu Z, et al. NSUN2-mediated m5C modification of SOCS3 mRNA modulates macrophage polarization in bladder cancer. Cell Death Dis. 2025;17:75.
Google Scholar
Kao WH, Liao LZ, Chen YA, Lo UG, Pong RC, Hernandez E, et al. SPHK1 promotes bladder cancer metastasis via PD-L2/c-Src/FAK signaling cascade. Cell Death Dis. 2024;15:678.
Google Scholar
Gong Y, Gao D, Shi Y, Fan G, Yu X, Yang E, et al. SRC enhanced cisplatin resistance in bladder cancer by reprogramming glycolysis and pentose phosphate pathway. Commun Biol. 2025;8:36.
Google Scholar
Zhang Y, Wang J, Zhang G, Cai H. TFDP1 is a potential diagnostic, immunological and prognostic biomarker in pan-cancer. Asian J Surg. 2024;47:2481–3.
Google Scholar
Morimoto Y, Mizushima T, Wu X, Okuzaki D, Yokoyama Y, Inoue A, et al. miR-4711-5p regulates cancer stemness and cell cycle progression via KLF5, MDM2 and TFDP1 in colon cancer cells. Br J Cancer. 2020;122:1037–49.
Google Scholar
Kent LN, Leone G. The broken cycle: E2F dysfunction in cancer. Nat Rev Cancer. 2019;19:326–38.
Google Scholar
Denechaud PD, Lopez-Mejia IC, Giralt A, Lai Q, Blanchet E, Delacuisine B, et al. E2F1 mediates sustained lipogenesis and contributes to hepatic steatosis. J Clin Invest. 2016;126:137–50.
Google Scholar
Shen D, Gao Y, Huang Q, Xuan Y, Yao Y, Gu L, et al. E2F1 promotes proliferation and metastasis of clear cell renal cell carcinoma via activation of SREBP1-dependent fatty acid biosynthesis. Cancer Lett. 2021;514:48–62.
Google Scholar

