REFERENCES
1. Tacke F, Horn P, Wai-Sun Wong V, et al. EASL-EASD-EASO Clinical Practice Guidelines on the management of metabolic dysfunction-associated steatotic liver disease (MASLD). Obesity Facts. 2024;17:374-443.
2. Rinella ME, Lazarus JV, Ratziu V, et al.; NAFLD Nomenclature consensus group. A multisociety Delphi consensus statement on new fatty liver disease nomenclature. J Hepatol. 2023;79:1542-56.
3. Targher G, Byrne CD, Lonardo A, Zoppini G, Barbui C. Non-alcoholic fatty liver disease and risk of incident cardiovascular disease: a meta-analysis. J Hepatol. 2016;65:589-600.
4. Chalasani N, Younossi Z, Lavine JE, et al. The diagnosis and management of nonalcoholic fatty liver disease: Practice guidance from the American Association for the Study of Liver Diseases. Hepatology. 2018;67:328-57.
6. Soto-Gutierrez A, Gough A, Vernetti LA, Taylor DL, Monga SP. Pre-clinical and clinical investigations of metabolic zonation in liver diseases: the potential of microphysiology systems. Exp Biol Med. 2017;242:1605-16.
7. Li Z, Luo G, Gan C, et al. Spatially resolved multi-omics of human metabolic dysfunction-associated steatotic liver disease. Nat Genet. 2025;57:3112-25.
8. Vandereyken K, Sifrim A, Thienpont B, Voet T. Methods and applications for single-cell and spatial multi-omics. Nat Rev Genet. 2023;24:494-515.
9. Ding J, Adiconis X, Simmons SK, et al. Systematic comparison of single-cell and single-nucleus RNA-sequencing methods. Nat Biotechnol. 2020;38:737-46.
10. Cunningham RP, Porat-Shliom N. Liver zonation - revisiting old questions with new technologies. Front Physiol. 2021;12:732929.
11. Yang Z, Zhao J, Xie K, Tang C, Gan C, Gao J. MASLD development: from molecular pathogenesis toward therapeutic strategies. Chin Med J. 2025;138:1807-24.
12. Christofides A, Konstantinidou E, Jani C, Boussiotis VA. The role of peroxisome proliferator-activated receptors (PPAR) in immune responses. Metabolism. 2021;114:154338.
13. Grygiel-Górniak B. Peroxisome proliferator-activated receptors and their ligands: nutritional and clinical implications--a review. Nutr J. 2014;13:17.
14. Hu K, Liu S, Sheng S, et al. Heterogeneity of hepatic macrophages in MASLD/MASH: Lipid-associated programs, molecular mechanisms, spatial niches, and therapeutic implications. ILIVER. 2026;5:100234.
15. Shi S, Zhou Y, Zhang H, Zhang J. TREM2 in MASH: integrating lipid metabolism and immune response. Front Immunol. 2025;16:1604837.
16. Xu Y, Hillman H, Chang M, et al. Identification of conserved and tissue-restricted transcriptional profiles for lipid associated macrophages. Commun Biol. 2025;8:953.
17. Govaere O, Petersen SK, Martinez-Lopez N, et al. Macrophage scavenger receptor 1 mediates lipid-induced inflammation in non-alcoholic fatty liver disease. J Hepatol. 2022;76:1001-12.
18. Jaitin DA, Adlung L, Thaiss CA, et al. Lipid-associated macrophages control metabolic homeostasis in a trem2-dependent manner. Cell. 2019;178:686-98.e14.
19. Zhang M, Haughey M, Wang NY, et al. Targeting the Wnt signaling pathway through R-spondin 3 identifies an anti-fibrosis treatment strategy for multiple organs. PLoS One. 2020;15:e0229445.
20. Candels LS, Becker S, Trautwein C. PLA2G7: a new player in shaping energy metabolism and lifespan. Signal Transduct Target Ther. 2022;7:195.
21. Ye J, Song R, Gong X, Li X, Shao C, Zhong B. Fibrosis severity in MASLD determines the predictive value of Lp-PLA2 for carotid atherosclerosis in type 2 diabetes: a cross-sectional study. Biomedicines. 2025;13:2431.
22. Sawada K, Chung H, Softic S, Moreno-Fernandez ME, Divanovic S. The bidirectional immune crosstalk in metabolic dysfunction-associated steatotic liver disease. Cell Metab. 2023;35:1852-71.
23. Ma C, Wang S, Dong B, Tian Y. Metabolic reprogramming of immune cells in MASH. Hepatology. 2025;Epub ahead of print.
24. Lefere S, Tacke F. Macrophages in obesity and non-alcoholic fatty liver disease: Crosstalk with metabolism. JHEP Rep. 2019;1:30-43.
25. Liu W, Li M, Guo H, et al. Single-cell transcriptome analysis of liver immune microenvironment changes induced by microplastics in mice with non-alcoholic fatty liver. Sci Total Environ. 2024;912:168308.
26. De Ponti FF, Liu Z, Scott CL. Understanding the complex macrophage landscape in MASLD. JHEP Rep. 2024;6:101196.
27. Vonderlin J, Chavakis T, Sieweke M, Tacke F. The multifaceted roles of macrophages in NAFLD pathogenesis. Cell Mol Gastroenterol Hepatol. 2023;15:1311-24.
28. Schleicher J, Dahmen U, Guthke R, Schuster S. Zonation of hepatic fat accumulation: insights from mathematical modelling of nutrient gradients and fatty acid uptake. J R Soc Interface. 2017;14:20170443.
29. van der Graaff D, Kwanten WJ, Francque SM. The potential role of vascular alterations and subsequent impaired liver blood flow and hepatic hypoxia in the pathophysiology of non-alcoholic steatohepatitis. Med Hypotheses. 2019;122:188-97.
30. Xu R, Vujić N, Bianco V, et al. Lipid-associated macrophages between aggravation and alleviation of metabolic diseases. Trends Endocrinol Metab. 2024;35:981-95.
31. Chen T, Zhang H, Shan W, Zhou J, You Y. Liver sinusoidal endothelial cells in hepatic fibrosis: opportunities for future strategies. Biochem Biophys Res Commun. 2025;766:151881.
32. Yin X, Yi H, Wang L, Wu W, Wu X, Yu L. RSPOs facilitated HSC activation and promoted hepatic fibrogenesis. Oncotarget. 2016;7:63767-78.
33. Sugimoto A, Saito Y, Wang G, et al. Hepatic stellate cells control liver zonation, size and functions via R-spondin 3. Nature. 2025;640:752-61.
34. Singh A, Akhtar A, Shukla P. Exploring hepatic stellate cell-driven fibrosis: therapeutic advances and future perspectives. ADMET DMPK. 2025;13:2874.
35. Makri ES, Mouskeftara T, Gika H, et al. Serum and liver lipidome following empagliflozin administration for six months in a fast food diet mouse model. Int J Mol Sci. 2025;26:9273.
36. Bertran L, Capellades J, Abelló S, Richart C. Untargeted lipidomic analysis of metabolic dysfunction-associated steatohepatitis in women with morbid obesity. PLoS One. 2025;20:e0318557.
37. Heymann CJF, Mak AL, Holleboom AG, et al. The plasma lipidome varies with the severity of metabolic dysfunction-associated steatotic liver disease. Lipids Health Dis. 2024;23:402.
38. Pebriana RB, Chen T, Derks RJE, et al. Non-invasive identification of steatohepatitis in patients with MASLD using a sterol and lipidomic signature. J Lipid Res. 2025;66:100845.
39. Meyer J, Teixeira AM, Richter S, et al. Sex differences in diet-induced MASLD - are female mice naturally protected? Front Endocrinol. 2025;16:1567573.
40. Makri ES, Xanthopoulos K, Mavrommatis Parasidis P, et al. Partial validation of a six-month high-fat diet and fructose-glucose drink combination as a mouse model of nonalcoholic fatty liver disease. Endocrine. 2024;85:704-16.
41. Ballestri S, Nascimbeni F, Baldelli E, Marrazzo A, Romagnoli D, Lonardo A. NAFLD as a sexual dimorphic disease: role of gender and reproductive status in the development and progression of nonalcoholic fatty liver disease and inherent cardiovascular risk. Adv Ther. 2017;34:1291-326.
42. Cherubini A, Della Torre S, Pelusi S, Valenti L. Sexual dimorphism of metabolic dysfunction-associated steatotic liver disease. Trends Mol Med. 2024;30:1126-36.
43. Geng A, Cui C, Luo Z, et al. Computational methods for spatial multi-omics integration. Biotechnol Adv. 2026;87:108807.






