REFERENCES

1. Webster AK, Phillips PC. Epigenetics and individuality: from concepts to causality across timescales. Nat Rev Genet. 2025;26:406-23.

2. Easwaran H, Weeraratna AT. Unravelling the genetics and epigenetics of the ageing tumour microenvironment in cancer. Nat Rev Cancer. 2025;25:828-47.

3. Mehler M. Epigenetic principles and mechanisms underlying nervous system functions in health and disease. Prog Neurobiol. 2008;86:305-41.

4. Marei HE. Epigenetic regulators in cancer therapy and progression. npj Precis Onc. 2025;9:206.

5. Lennartsson A, Ekwall K. Histone modification patterns and epigenetic codes. Biochim Biophys Acta Gen Subj. 2009;1790:863-8.

6. Villar-garea A, Imhof A. The analysis of histone modifications. Biochim Biophys Acta Proteins Proteom. 2006;1764:1932-9.

7. Chen J, Huang Z, Chen Y, et al. Lactate and lactylation in cancer. Sig Transduct Target Ther. 2025;10:38.

8. Yang B, Li L, Shi D, Zhong T, Xiong H. Lactylation and antitumor immunity. Front Immunol. 2025;16:1690068.

9. Zhu W, Fan C, Hou Y, Zhang Y. Lactylation in tumor microenvironment and immunotherapy resistance: new mechanisms and challenges. Cancer Lett. 2025;627:217835.

10. Feng X, Li D, Wang P, Li X, Li G. Lactylation in cancer: unlocking the key to drug resistance and therapeutic breakthroughs. Oncol Res. 2025;33:3327-46.

11. Zhang D, Tang Z, Huang H, et al. Metabolic regulation of gene expression by histone lactylation. Nature. 2019;574:575-80.

12. Xiao S, Zhang S, Sun K, Huang Q, Li Q, Hu C. Lactate and lactylation: molecular insights into histone and non-histone lactylation in tumor progression, tumor immune microenvironment, and therapeutic strategies. Biomark Res. 2025;13:134.

13. Yang Y, Luo N, Gong Z, Zhou W, Ku Y, Chen Y. Lactate and lysine lactylation of histone regulate transcription in cancer. Heliyon. 2024;10:e38426.

14. Peng X, Du J. Histone and non-histone lactylation: molecular mechanisms, biological functions, diseases, and therapeutic targets. Mol Biomed. 2025;6:38.

15. Barcena-varela M, Monga SP, Lujambio A. Precision models in hepatocellular carcinoma. Nat Rev Gastroenterol Hepatol. 2024;22:191-205.

16. Zhao Z, Cui T, Wei F, et al. Wnt/β-Catenin signaling pathway in hepatocellular carcinoma: pathogenic role and therapeutic target. Front Oncol. 2024;14:1367364.

17. Yang H, Li J, Niu Y, et al. Interactions between the metabolic reprogramming of liver cancer and tumor microenvironment. Front Immunol. 2025;16:1494788.

18. Zhang X, Liang C, Wu C, et al. A rising star involved in tumour immunity: lactylation. J Cell Mol Med. 2024;28:e70146.

19. Zong Z, Xie F, Wang S, et al. Alanyl-tRNA synthetase, AARS1, is a lactate sensor and lactyltransferase that lactylates p53 and contributes to tumorigenesis. Cell. 2024;187:2375-2392.e33.

20. Yang Z, Yan C, Ma J, et al. Lactylome analysis suggests lactylation-dependent mechanisms of metabolic adaptation in hepatocellular carcinoma. Nat Metab. 2023;5:61-79.

21. Wei X, Zou L, Huang Y, et al. LDHA-mediated YAP lactylation promotes the tumor progression of hepatocellular carcinoma by inducing YAP dephosphorylation and activation. Biol Direct. 2025;20:64.

22. Zhao L, Cui H, Li Y, Ye Y, Shen Z. Metabolite to modifier: lactate and lactylation in the evolution of tumors. MedComm. 2025;6:e70413.

23. Zhao P, Qiao C, Wang J, Zhou Y, Zhang C. Histone lactylation facilitates hepatocellular carcinoma progression by upregulating endothelial cell‐specific molecule 1 expression. Mol Carcinog. 2024;63:2078-89.

24. Wu S, Li J, Zhan Y. H3K18 lactylation accelerates liver fibrosis progression through facilitating SOX9 transcription. Exp Cell Res. 2024;440:114135.

25. Huang J, Xie H, Li J, et al. Histone lactylation drives liver cancer metastasis by facilitating NSF1-mediated ferroptosis resistance after microwave ablation. Redox Biology. 2025;81:103553.

26. Li L, Dong J, Xu C, Wang S. Lactate drives senescence-resistant lineages in hepatocellular carcinoma via histone H2B lactylation of NDRG1. Cancer Lett. 2025;616:217567.

27. Xu H, Li L, Wang S, et al. Royal jelly acid suppresses hepatocellular carcinoma tumorigenicity by inhibiting H3 histone lactylation at H3K9la and H3K14la sites. Phytomedicine. 2023;118:154940.

28. Xu K, Zhang K, Wang Y, Gu Y. Comprehensive review of histone lactylation: structure, function, and therapeutic targets. Biochem Pharmacol. 2024;225:116331.

29. Li H, Sun L, Gao P, Hu H. Lactylation in cancer: current understanding and challenges. Cancer Cell. 2024;42:1803-7.

30. Jia Z, Lu S, Wang Z, Ge P. From mechanism to targeted therapy: advances in histone lactylationdriven cancer progression (review). Oncol Lett. 2025;31:1-17.

31. Qin Q, Wang D, Qu Y, et al. Enhanced glycolysis-derived lactate promotes microglial activation in Parkinson’s disease via histone lactylation. npj Parkinsons Dis. 2025;11:3.

32. Wang N, Wang W, Wang X, et al. Histone lactylation boosts reparative gene activation post-myocardial infarction. Circ Res. 2022;131:893-908.

33. Zhu R, Ye X, Lu X, et al. ACSS2 acts as a lactyl-CoA synthetase and couples KAT2A to function as a lactyltransferase for histone lactylation and tumor immune evasion. Cell Metab. 2025;37:361-376.e7.

34. Niu Z, Chen C, Wang S, et al. HBO1 catalyzes lysine lactylation and mediates histone H3K9la to regulate gene transcription. Nat Commun. 2024;15:3561.

35. Zou Y, Cao M, Tai M, et al. A feedback loop driven by H4K12 lactylation and HDAC3 in macrophages regulates lactate‐induced collagen synthesis in fibroblasts via the TGF‐β signaling. Adv Sci. 2025;12:2411408.

36. Llovet JM, Kelley RK, Villanueva A, et al. Hepatocellular carcinoma. Nat Rev Dis Primers. 2021;7:6.

37. Luo X, He X, Zhang X, et al. Hepatocellular carcinoma: signaling pathways, targeted therapy, and immunotherapy. MedComm. 2024;5:e474.

38. Wang J, Liu C, Hu R, Wu L, Li C. Statin therapy: a potential adjuvant to immunotherapies in hepatocellular carcinoma. Front Pharmacol. 2024;15:1324140.

39. Zhang J, Hu C, Xie X, Qi L, Li C, Li S. Immune checkpoint inhibitors in HBV-caused hepatocellular carcinoma therapy. Vaccines. 2023;11:614.

40. Cheng K, Cai N, Zhu J, Yang X, Liang H, Zhang W. Tumor‐associated macrophages in liver cancer: from mechanisms to therapy. Cancer Commun. 2022;42:1112-40.

41. Yang Y, Gao Y, Xiong Y, et al. Research progress of warburg effect in hepatocellular carcinoma. Front Biosci. 2024;29:178.

42. Pan L, Feng F, Wu J, et al. Demethylzeylasteral targets lactate by inhibiting histone lactylation to suppress the tumorigenicity of liver cancer stem cells. Pharmacol Res. 2022;181:106270.

43. Liu Z, Han J, Su S, et al. Histone lactylation facilitates MCM7 expression to maintain stemness and radio-resistance in hepatocellular carcinoma. Biochem Pharmacol. 2025;236:116887.

44. Gao Y, Gong Y, Song X, et al. Dihydroartemisinin inhibits histone lactylation through YAP1 to act as a ‘hot’ switch for ‘cold’ tumor in hepatocellular carcinoma. Phytomedicine. 2025;148:157307.

45. Ji Y, Xu Z, Tang L, et al. O-GlcNAcylation of YBX1 drives a glycolysis-histone lactylation feedback loop in hepatocellular carcinoma. Cancer Lett. 2025;631:217957.

46. Fan M, Liu JS, Wei XL, Nie Y, Liu HL. Histone lactylation-driven ubiquitin-specific protease 34 promotes cisplatin resistance in hepatocellular carcinoma. Gastroenterol Res. 2025;18:23-30.

47. Wang H, Xu M, Zhang T, et al. PYCR1 promotes liver cancer cell growth and metastasis by regulating IRS1 expression through lactylation modification. Clin Transl Med.2024;14:e70045.

48. Zeng Y, Jiang H, Chen Z, et al. Histone lactylation promotes multidrug resistance in hepatocellular carcinoma by forming a positive feedback loop with PTEN. Cell Death Dis. 2025;16:59.

49. Dong R, Fei Y, He Y, et al. Lactylation‐driven HECTD2 limits the response of hepatocellular carcinoma to lenvatinib. Adv Sci. 2025;12:2412559.

50. Ye J, Gao X, Huang X, et al. Integrating single-cell and spatial transcriptomics to uncover and elucidate GP73-mediated pro-angiogenic regulatory networks in hepatocellular carcinoma. Research. 2024;7:0387.

51. Hong H, Chen X, Wang H, Gu X, Yuan Y, Zhang Z. Global profiling of protein lysine lactylation and potential target modified protein analysis in hepatocellular carcinoma. Proteomics. 2023;23:2200432.

52. Liu S, Pan Y, Liu W, et al. Lactylation-driven MVP upregulation boosts immunotherapy resistance by inhibiting PD-L1 degradation in hepatocellular carcinoma. J Immunother Cancer. 2025;13:e012230.

53. Cai J, Song L, Zhang F, et al. Targeting SRSF10 might inhibit M2 macrophage polarization and potentiate anti‐PD‐1 therapy in hepatocellular carcinoma. Cancer Commun. 2024;44:1231-60.

54. Li J, Chen X, Song S, et al. Hexokinase 2-mediated metabolic stress and inflammation burden of liver macrophages via histone lactylation in MASLD. Cell Reports. 2025;44:115350.

55. Du W, Tan S, Peng Y, et al. Histone lactylation-driven YTHDC1 promotes hepatocellular carcinoma progression via lipid metabolism remodeling. Cancer Lett. 2025;611:217426.

56. Wang Z, Liu Z, Lv M, Luan Z, Li T, Hu J. Novel histone modifications and liver cancer: emerging frontiers in epigenetic regulation. Clin Epigenet. 2025;17:30.

57. Ding C, Yan F, Xu B, et al. PRMT3 drives PD-L1-mediated immune escape through activating PDHK1-regulated glycolysis in hepatocellular carcinoma. Cell Death Dis. 2025;16:158.

58. Corrigan DT, Tanwar A, Du M, Martin AM, Zang X. The B7-H3 (CD276) pathway: emerging biology and clinical therapeutics. Trends Pharmacol Sci. 2025;46:975-88.

59. Ma Z, Yang J, Jia W, et al. Histone lactylation-driven B7-H3 expression promotes tumor immune evasion. Theranostics. 2025;15:2338-59.

60. Rho H, Terry AR, Chronis C, Hay N. Hexokinase 2-mediated gene expression via histone lactylation is required for hepatic stellate cell activation and liver fibrosis. Cell Metab. 2023;35:1406-1423.e8.

61. Yasukawa K, Shimada S, Akiyama Y, et al. ACVR2A attenuation impacts lactate production and hyperglycolytic conditions attracting regulatory T cells in hepatocellular carcinoma. Cell Rep Med. 2025;6:102038.

Hepatoma Research
ISSN 2454-2520 (Online) 2394-5079 (Print)

Portico

All published articles are preserved here permanently:

https://www.portico.org/publishers/oae/

Portico

All published articles are preserved here permanently:

https://www.portico.org/publishers/oae/