[1]

Boada-Romero E, Martinez J, Heckmann BL, Green DR. 2020. The clearance of dead cells by efferocytosis. Nature Reviews Molecular Cell Biology 21:398−414

doi: 10.1038/s41580-020-0232-1
[2]

Ma Y, Jiang T, Zhu X, Xu Y, Wan K, et al. 2024. Efferocytosis in dendritic cells: an overlooked immunoregulatory process. Frontiers in Immunology 15:1415573

doi: 10.3389/fimmu.2024.1415573
[3]

de Cathelineau AM, Henson PM. 2003. The final step in programmed cell death: phagocytes carry apoptotic cells to the grave. Essays in Biochemistry 39:105−117

doi: 10.1042/bse0390105
[4]

Novak ML, Thorp EB. 2013. Shedding light on impaired efferocytosis and nonresolving inflammation. Circulation Research 113:9−12

doi: 10.1161/CIRCRESAHA.113.301583
[5]

Tajbakhsh A, Gheibi hayat SM, Movahedpour A, Savardashtaki A, Loveless R, et al. 2021. The complex roles of efferocytosis in cancer development, metastasis, and treatment. Biomedicine & Pharmacotherapy 140:111776

doi: 10.1016/j.biopha.2021.111776
[6]

Schilperoort M, Ngai D, Sukka SR, Avrampou K, Shi H, et al. 2023. The role of efferocytosis-fueled macrophage metabolism in the resolution of inflammation. Immunological Reviews 319:65−80

doi: 10.1111/imr.13214
[7]

Boucher DM, Robichaud S, Lorant V, Leon JS, Suliman I, et al. 2025. Age-related impairments in immune cell efferocytosis and autophagy hinder atherosclerosis regression. Arteriosclerosis, Thrombosis, and Vascular Biology 45:481−495

doi: 10.1161/ATVBAHA.124.321662
[8]

Chen Q, Liu C, Wang Q, Zhang X, Zheng Y, et al. 2026. The role of macrophage metabolic reprogramming in efferocytosis: a dual-edged sword in atherosclerosis and tumor progression. Molecular Immunology 194:63−72

doi: 10.1016/j.molimm.2026.04.006
[9]

Bäck M, Yurdagul A, Tabas I, Öörni K, Kovanen PT. 2019. Inflammation and its resolution in atherosclerosis: mediators and therapeutic opportunities. Nature Reviews Cardiology 16:389−406

doi: 10.1038/s41569-019-0169-2
[10]

Chen Y, Kou Y, Ni Y, Yang H, Xu C, et al. 2025. Microglia efferocytosis: an emerging mechanism for the resolution of neuroinflammation in Alzheimer's disease. Journal of Neuroinflammation 22:96

doi: 10.1186/s12974-025-03428-0
[11]

Xing J, Wang K, Xu YC, Pei ZJ, Yu QX, et al. 2024. Efferocytosis: unveiling its potential in autoimmune disease and treatment strategies. Autoimmunity Reviews 23:103578

doi: 10.1016/j.autrev.2024.103578
[12]

Astuti Y, Raymant M, Quaranta V, Clarke K, Abudula M, et al. 2024. Efferocytosis reprograms the tumor microenvironment to promote pancreatic cancer liver metastasis. Nature Cancer 5:774−790

doi: 10.1038/s43018-024-00731-2
[13]

Myers KV, Amend SR, Pienta KJ. 2019. Targeting Tyro3, Axl and MerTK (TAM receptors): implications for macrophages in the tumor microenvironment. Molecular Cancer 18:94

doi: 10.1186/s12943-019-1022-2
[14]

Razi S, Yaghmoorian Khojini J, Kargarijam F, Panahi S, Tahershamsi Z, et al. 2023. Macrophage efferocytosis in health and disease. Cell Biochemistry and Function 41:152−165

doi: 10.1002/cbf.3780
[15]

Zhang W, Zeng Y, Xiao Q, Wu Y, Liu J, et al. 2024. An in-situ peptide-antibody self-assembly to block CD47 and CD24 signaling enhances macrophage-mediated phagocytosis and anti-tumor immune responses. Nature Communications 15:5670

doi: 10.1038/s41467-024-49825-6
[16]

Hayat SMG, Bianconi V, Pirro M, Jaafari MR, Hatamipour M, et al. 2020. CD47: role in the immune system and application to cancer therapy. Cellular Oncology 43:19−30

doi: 10.1007/s13402-019-00469-5
[17]

Harris R, Harman DJ, Card TR, Aithal GP, Guha IN. 2017. Prevalence of clinically significant liver disease within the general population, as defined by non-invasive markers of liver fibrosis: a systematic review. The Lancet Gastroenterology & Hepatology 2:288−297

doi: 10.1016/S2468-1253(16)30205-9
[18]

Zamani M, Alizadeh-Tabari S, Ajmera V, Singh S, Murad MH, et al. 2025. Global prevalence of advanced liver fibrosis and cirrhosis in the general population: a systematic review and meta-analysis. Clinical Gastroenterology and Hepatology 23:1123−1134

doi: 10.1016/j.cgh.2024.08.020
[19]

Kim HY, Yu JH, Chon YE, Kim SU, Kim MN, et al. 2024. Prevalence of clinically significant liver fibrosis in the general population: a systematic review and meta-analysis. Clinical and Molecular Hepatology 30:S199−S213

doi: 10.3350/cmh.2024.0351
[20]

Friedman SL, Pinzani M. 2022. Hepatic fibrosis 2022: Unmet needs and a blueprint for the future. Hepatology 75:473−488

doi: 10.1002/hep.32285
[21]

Harrison SA, Bedossa P, Guy CD, Schattenberg JM, Loomba R, et al. 2024. A phase 3, randomized, controlled trial of resmetirom in NASH with liver fibrosis. New England Journal of Medicine 390:497−509

doi: 10.1056/NEJMoa2309000
[22]

Friedman SL, Roll FJ, Boyles J, Bissell DM. 1985. Hepatic lipocytes: the principal collagen-producing cells of normal rat liver. Proceedings of the National Academy of Sciences of the United States of America 82:8681−8685

doi: 10.1073/pnas.82.24.8681
[23]

Yao L, Hu X, Dai K, Yuan M, Liu P, et al. 2022. Mesenchymal stromal cells: promising treatment for liver cirrhosis. Stem Cell Research & Therapy 13:308

doi: 10.1186/s13287-022-03001-z
[24]

Ramachandran P, Matchett KP, Dobie R, Wilson-Kanamori JR, Henderson NC. 2020. Single-cell technologies in hepatology: new insights into liver biology and disease pathogenesis. Nature Reviews Gastroenterology & Hepatology 17:457−472

doi: 10.1038/s41575-020-0304-x
[25]

Zhong Z, Cui XL, Tan KJ, Wu XY, Zhu XJ, et al. 2024. Apoptotic vesicles (apoVs) derived from fibroblast-converted hepatocyte-like cells effectively ameliorate liver fibrosis. Journal of Nanobiotechnology 22:541

doi: 10.1186/s12951-024-02824-7
[26]

Mohammed S, Thadathil N, Selvarani R, Nicklas EH, Wang D, et al. 2021. Necroptosis contributes to chronic inflammation and fibrosis in aging liver. Aging Cell 20:e13512

doi: 10.1111/acel.13512
[27]

Kim KH, Cheng N, Lau LF. 2022. Cellular communication network factor 1-stimulated liver macrophage efferocytosis drives hepatic stellate cell activation and liver fibrosis. Hepatology Communications 6:2798−2811

doi: 10.1002/hep4.2057
[28]

Lu JL, Yu CX, Song LJ. 2023. Programmed cell death in hepatic fibrosis: current and perspectives. Cell Death Discovery 9:449

doi: 10.1038/s41420-023-01749-8
[29]

Mu YP, Ogawa T, Kwada N. 2012. Mechanism of hepatocyte apoptosis in rats with liver fibrosis induced by lipogenic methionine-choline-deficient diet. Zhonghua Bing Li Xue Za Zhi 41:112−118 (in Chinese)

doi: 10.3760/cma.j.issn.0529-5807.2012.02.009
[30]

Wang Z, Du K, Jin N, Tang B, Zhang W. 2023. Macrophage in liver fibrosis: identities and mechanisms. International Immunopharmacology 120:110357

doi: 10.1016/j.intimp.2023.110357
[31]

Morioka S, Maueröder C, Ravichandran KS. 2019. Living on the edge: efferocytosis at the interface of homeostasis and pathology. Immunity 50:1149−1162

doi: 10.1016/j.immuni.2019.04.018
[32]

Hochreiter-Hufford A, Ravichandran KS. 2013. Clearing the dead: apoptotic cell sensing, recognition, engulfment, and digestion. Cold Spring Harbor Perspectives in Biology 5:a008748

doi: 10.1101/cshperspect.a008748
[33]

Elliott MR, Chekeni FB, Trampont PC, Lazarowski ER, Kadl A, et al. 2009. Nucleotides released by apoptotic cells act as a find-me signal to promote phagocytic clearance. Nature 461:282−286

doi: 10.1038/nature08296
[34]

Peter C, Waibel M, Radu CG, Yang LV, Witte ON, et al. 2008. Migration to apoptotic "find-me" signals is mediated via the phagocyte receptor G2A. Journal of Biological Chemistry 283:5296−5305

doi: 10.1074/jbc.M706586200
[35]

Gude DR, Alvarez SE, Paugh SW, Mitra P, Yu J, et al. 2008. Apoptosis induces expression of sphingosine kinase 1 to release sphingosine-1-phosphate as a "come-and-get-me" signal. The FASEB Journal 22:2629−2638

doi: 10.1096/fj.08-107169
[36]

Truman LA, Ford CA, Pasikowska M, Pound JD, Wilkinson SJ, et al. 2008. CX3CL1/fractalkine is released from apoptotic lymphocytes to stimulate macrophage chemotaxis. Blood 112:5026−5036

doi: 10.1182/blood-2008-06-162404
[37]

Tajbakhsh A, Yousefi F, Abedi SM, Rezaee M, Savardashtaki A, et al. 2022. The cross-talk between soluble "Find me" and "Keep out" signals as an initial step in regulating efferocytosis. Journal of Cellular Physiology 237:3113−3126

doi: 10.1002/jcp.30770
[38]

Knies UE, Behrensdorf HA, Mitchell CA, Deutsch U, Risau W, et al. 1998. Regulation of endothelial monocyte-activating polypeptide II release by apoptosis. Proceedings of the National Academy of Sciences of the United States of America 95:12322−12327

doi: 10.1073/pnas.95.21.12322
[39]

Neniskyte U, Kuliesiute U, Vadisiute A, Jevdokimenko K, Coletta L, et al. 2023. Phospholipid scramblase Xkr8 is required for developmental axon pruning via phosphatidylserine exposure. The EMBO Journal 42:EMBJ2022111790

doi: 10.15252/embj.2022111790
[40]

Morioka S, Kajioka D, Yamaoka Y, Ellison RM, Tufan T, et al. 2022. Chimeric efferocytic receptors improve apoptotic cell clearance and alleviate inflammation. Cell 185:4887−4903.e17

doi: 10.1016/j.cell.2022.11.029
[41]

Lemke G, Burstyn-Cohen T. 2010. TAM receptors and the clearance of apoptotic cells. Annals of the New York Academy of Sciences 1209:23−29

doi: 10.1111/j.1749-6632.2010.05744.x
[42]

van der Meer JHM, van der Poll T, van 't Veer C. 2014. TAM receptors, Gas6, and protein S: roles in inflammation and hemostasis. Blood 123:2460−2469

doi: 10.1182/blood-2013-09-528752
[43]

Chen J, Zhong MC, Guo H, Davidson D, Mishel S, et al. 2017. SLAMF7 is critical for phagocytosis of haematopoietic tumour cells via Mac-1 integrin. Nature 544:493−497

doi: 10.1038/nature22076
[44]

Matozaki T, Murata Y, Okazawa H, Ohnishi H. 2009. Functions and molecular mechanisms of the CD47-SIRPα signalling pathway. Trends in Cell Biology 19:72−80

doi: 10.1016/j.tcb.2008.12.001
[45]

Yurdagul A Jr, Subramanian M, Wang X, Crown SB, Ilkayeva OR, et al. 2020. Macrophage metabolism of apoptotic cell-derived arginine promotes continual efferocytosis and resolution of injury. Cell Metabolism 31:518−533.e10

doi: 10.1016/j.cmet.2020.01.001
[46]

Liu Z, Deng Y, Song X, Cai X, Xiong H, et al. 2025. CXCL12/CXCR4 modulates macrophage efferocytosis to induce glomerular crescent formation and fibrosis via ELMO1/DOCK180/RAC1 signaling in ANCA-associated glomerulonephritis. Cellular and Molecular Life Sciences 82:280

doi: 10.1007/s00018-025-05750-5
[47]

Wang Y, Wang J, Zhang J, Wang Y, Wang Y, et al. 2024. Stiffness sensing via Piezo1 enhances macrophage efferocytosis and promotes the resolution of liver fibrosis. Science Advances 10:eadj3289

doi: 10.1126/sciadv.adj3289
[48]

Davidson AJ, Wood W. 2020. Macrophages use distinct actin regulators to switch engulfment strategies and ensure phagocytic plasticity in vivo. Cell Reports 31:107692

doi: 10.1016/j.celrep.2020.107692
[49]

Sukka SR, Ampomah PB, Darville LNF, Ngai D, Wang X, et al. 2024. Efferocytosis drives a tryptophan metabolism pathway in macrophages to promote tissue resolution. Nature Metabolism 6:1736−1755

doi: 10.1038/s42255-024-01115-7
[50]

Wang Y, Subramanian M, Yurdagul A, Jr. , Barbosa-Lorenzi VC, Cai B, et al. 2017. Mitochondrial fission promotes the continued clearance of apoptotic cells by macrophages. Cell 171:331−345.e22

doi: 10.1016/j.cell.2017.08.041
[51]

Kinchen JM, Doukoumetzidis K, Almendinger J, Stergiou L, Tosello-Trampont A, et al. 2008. A pathway for phagosome maturation during engulfment of apoptotic cells. Nature Cell Biology 10:556−566

doi: 10.1038/ncb1718
[52]

Zhao YG, Codogno P, Zhang H. 2021. Machinery, regulation and pathophysiological implications of autophagosome maturation. Nature Reviews Molecular Cell Biology 22:733−750

doi: 10.1038/s41580-021-00392-4
[53]

Wang Y, Liu XY, Wang Y, Zhao WX, Li FD, et al. 2023. NOX2 inhibition stabilizes vulnerable plaques by enhancing macrophage efferocytosis via MertK/PI3K/AKT pathway. Redox Biology 64:102763

doi: 10.1016/j.redox.2023.102763
[54]

Jiang S, Park DW, Stigler WS, Creighton J, Ravi S, et al. 2013. Mitochondria and AMP-activated protein kinase-dependent mechanism of efferocytosis. Journal of Biological Chemistry 288:26013−26026

doi: 10.1074/jbc.M113.489468
[55]

Garcia-Macia M, Santos-Ledo A, Leslie J, Paish HL, Collins AL, et al. 2021. A mammalian target of rapamycin-perilipin 3 (mTORC1-Plin3) pathway is essential to activate lipophagy and protects against hepatosteatosis. Hepatology 74:3441−3459

doi: 10.1002/hep.32048
[56]

Thorp E, Vaisar T, Subramanian M, Mautner L, Blobel C, et al. 2011. Shedding of the Mer tyrosine kinase receptor is mediated by ADAM17 protein through a pathway involving reactive oxygen species, protein kinase Cδ, and p38 mitogen-activated protein kinase (MAPK). Journal of Biological Chemistry 286:33335−33344

doi: 10.1074/jbc.M111.263020
[57]

Wei S, Guan G, Luan X, Yu C, Miao L, et al. 2025. NLRP3 inflammasome constrains liver regeneration through impairing MerTK-mediated macrophage efferocytosis. Science Advances 11:eadq5786

doi: 10.1126/sciadv.adq5786
[58]

Babuta M, Morel C, de Carvalho Ribeiro M, Calenda C, Ortega-Ribera M, et al. 2024. Neutrophil extracellular traps activate hepatic stellate cells and monocytes via NLRP3 sensing in alcohol-induced acceleration of MASH fibrosis. Gut 73:1854−1869

doi: 10.1136/gutjnl-2023-331447
[59]

Mridha AR, Wree A, Robertson AAB, Yeh MM, Johnson CD, et al. 2017. NLRP3 inflammasome blockade reduces liver inflammation and fibrosis in experimental NASH in mice. Journal of Hepatology 66:1037−1046

doi: 10.1016/j.jhep.2017.01.022
[60]

Ramachandran P, Pellicoro A, Vernon MA, Boulter L, Aucott RL, et al. 2012. Differential Ly-6C expression identifies the recruited macrophage phenotype, which orchestrates the regression of murine liver fibrosis. Proceedings of the National Academy of Sciences of the United States of America 109:E3186−E3195

doi: 10.1073/pnas.1119964109
[61]

Huo S, Li B, Du J, Zhang X, Zhang J, et al. 2023. Dibutyl phthalate induces liver fibrosis via p38MAPK/NF-κB/NLRP3-mediated pyroptosis. Science of the Total Environment 897:165500

doi: 10.1016/j.scitotenv.2023.165500
[62]

Liebold I, Meyer S, Heine M, Kuhl A, Witt J, et al. 2023. TREM2 regulates the removal of apoptotic cells and inflammatory processes during the progression of NAFLD. Cells 12:341

doi: 10.3390/cells12030341
[63]

De Ponti FF, Bujko A, Liu Z, Collins PJ, Schuermans S, et al. 2025. Spatially restricted and ontogenically distinct hepatic macrophages are required for tissue repair. Immunity 58:362−380.e10

doi: 10.1016/j.immuni.2025.01.002
[64]

Xiong X, Kuang H, Ansari S, Liu T, Gong J, et al. 2019. Landscape of intercellular crosstalk in healthy and NASH liver revealed by single-cell secretome gene analysis. Molecular Cell 75:644−660.e5

doi: 10.1016/j.molcel.2019.07.028
[65]

Junior, Lai YS, Nguyen HT, Salmanida FP, Chang KT. 2021. MERTK+/hi M2c macrophages induced by baicalin alleviate non-alcoholic fatty liver disease. International Journal of Molecular Sciences 22:10604

doi: 10.3390/ijms221910604
[66]

An P, Wei LL, Zhao S, Sverdlov DY, Vaid KA, et al. 2020. Hepatocyte mitochondria-derived danger signals directly activate hepatic stellate cells and drive progression of liver fibrosis. Nature Communications 11:2362

doi: 10.1038/s41467-020-16092-0
[67]

Ganguly S, Rosenthal SB, Ishizuka K, Troutman TD, Rohm TV, et al. 2024. Lipid-associated macrophages' promotion of fibrosis resolution during MASH regression requires TREM2. Proceedings of the National Academy of Sciences of the United States of America 121:e2405746121

doi: 10.1073/pnas.2405746121
[68]

Hendrikx T, Porsch F, Kiss MG, Rajcic D, Papac-Miličević N, et al. 2022. Soluble TREM2 levels reflect the recruitment and expansion of TREM2+ macrophages that localize to fibrotic areas and limit NASH. Journal of Hepatology 77:1373−1385

doi: 10.1016/j.jhep.2022.06.004
[69]

Indira Chandran V, Wernberg CW, Lauridsen MM, Skytthe MK, Bendixen SM, et al. 2023. Circulating TREM2 as a noninvasive diagnostic biomarker for NASH in patients with elevated liver stiffness. Hepatology 77:558−572

doi: 10.1002/hep.32620
[70]

Shi H, Wang X, Li F, Gerlach BD, Yurdagul A Jr, et al. 2022. CD47-SIRPα axis blockade in NASH promotes necroptotic hepatocyte clearance by liver macrophages and decreases hepatic fibrosis. Science Translational Medicine 14:eabp8309

doi: 10.1126/scitranslmed.abp8309
[71]

Cai B, Dongiovanni P, Corey KE, Wang X, Shmarakov IO, et al. 2020. Macrophage MerTK promotes liver fibrosis in nonalcoholic steatohepatitis. Cell Metabolism 31:406−421.e7

doi: 10.1016/j.cmet.2019.11.013
[72]

Popov Y, Sverdlov DY, Bhaskar KR, Sharma AK, Millonig G, et al. 2010. Macrophage-mediated phagocytosis of apoptotic cholangiocytes contributes to reversal of experimental biliary fibrosis. American Journal of Physiology-Gastrointestinal and Liver Physiology 298:G323−G334

doi: 10.1152/ajpgi.00394.2009
[73]

Shi H, Wang X, Sloas C, Gerlach B, Yurdagul A Jr, et al. 2025. Impaired TIM4-mediated efferocytosis by liver macrophages contributes to fibrosis in metabolic dysfunction–associated steatohepatitis. Science Translational Medicine 17:eadv2106

doi: 10.1126/scitranslmed.adv2106
[74]

Wu H, Chen G, Wang J, Deng M, Yuan F, et al. 2020. TIM-4 interference in Kupffer cells against CCL4-induced liver fibrosis by mediating Akt1/Mitophagy signalling pathway. Cell Proliferation 53:e12731

doi: 10.1111/cpr.12731
[75]

Kuang S, Feng C, Yuan Y, Li M, Mo C, et al. 2026. Baoganning decoction mitigates liver fibrosis via regulating IDO1-mediated macrophage efferocytosis. Phytomedicine 151:157798

doi: 10.1016/j.phymed.2026.157798
[76]

Eom JA, Park IG, Hyun JY, Lee NY, Kwon GH, et al. 2026. Pharmabiotics, Phocaeicola dorei, ameliorates cholestatic liver fibrosis by alleviating macrophage efferocytosis of neutrophils. Nature Communications 17:6516

doi: 10.1038/s41467-026-73166-1
[77]

Lee IH, Im E, Lee HJ, Sim DY, Lee JH, et al. 2021. Apoptotic and antihepatofibrotic effect of honokiol via activation of GSK3β and suppression of Wnt/β-catenin pathway in hepatic stellate cells. Phytotherapy Research 35:452−462

doi: 10.1002/ptr.6824
[78]

Jindal A, Bruzzì S, Sutti S, Locatelli I, Bozzola C, et al. 2015. Fat-laden macrophages modulate lobular inflammation in nonalcoholic steatohepatitis (NASH). Experimental and Molecular Pathology 99:155−162

doi: 10.1016/j.yexmp.2015.06.015
[79]

Wang W, Gao Y, Chen Y, Cheng M, Sang Y, et al. 2025. TGF-β inhibitors: the future for prevention and treatment of liver fibrosis? Frontiers in Immunology 16:1583616

doi: 10.3389/fimmu.2025.1583616
[80]

Peng Y, Lei H, Zhao J, Wang H, Luo Z, et al. 2025. GDF10 attenuates MASH progression by restoring quiescent hepatic stellate cells via competitive inhibition of TGF-β/SMAD2 signaling. International Journal of Biological Sciences 21:6997−7012

doi: 10.7150/ijbs.123784
[81]

Luedde T, Kaplowitz N, Schwabe RF. 2014. Cell death and cell death responses in liver disease: mechanisms and clinical relevance. Gastroenterology 147:765−783.e4

doi: 10.1053/j.gastro.2014.07.018
[82]

Pastore M, Caligiuri A, Raggi C, Navari N, Piombanti B, et al. 2022. Macrophage MerTK promotes profibrogenic cross-talk with hepatic stellate cells via soluble mediators. JHEP Reports 4:100444

doi: 10.1016/j.jhepr.2022.100444
[83]

Petta S, Valenti L, Marra F, Grimaudo S, Tripodo C, et al. 2016. MERTK rs4374383 polymorphism affects the severity of fibrosis in non-alcoholic fatty liver disease. Journal of Hepatology 64:682−690

doi: 10.1016/j.jhep.2015.10.016
[84]

Remmerie A, Martens L, Thoné T, Castoldi A, Seurinck R, et al. 2020. Osteopontin expression identifies a subset of recruited macrophages distinct from kupffer cells in the fatty liver. Immunity 53:641−657.e14

doi: 10.1016/j.immuni.2020.08.004
[85]

Fabre T, Barron AMS, Christensen SM, Asano S, Bound K, et al. 2023. Identification of a broadly fibrogenic macrophage subset induced by type 3 inflammation. Science Immunology 8:eadd8945

doi: 10.1126/sciimmunol.add8945
[86]

Ramachandran P, Dobie R, Wilson-Kanamori JR, Dora EF, Henderson BEP, et al. 2019. Resolving the fibrotic niche of human liver cirrhosis at single-cell level. Nature 575:512−518

doi: 10.1038/s41586-019-1631-3
[87]

Lee HJ, Jeng YM, Chen YL, Chung L, Yuan RH. 2014. Gas6/Axl pathway promotes tumor invasion through the transcriptional activation of Slug in hepatocellular carcinoma. Carcinogenesis 35:769−775

doi: 10.1093/carcin/bgt372
[88]

Reichl P, Dengler M, van Zijl F, Huber H, Fuhrlinger G, et al. 2015. Axl activates autocrine transforming growth factor-β signaling in hepatocellular carcinoma. Hepatology 61:930−941

doi: 10.1002/hep.27492
[89]

Fourcot A, Couchie D, Chobert MN, Zafrani ES, Mavier P, et al. 2011. Gas6 deficiency prevents liver inflammation, steatohepatitis, and fibrosis in mice. American Journal of Physiology-Gastrointestinal and Liver Physiology 300:G1043−G1053

doi: 10.1152/ajpgi.00311.2010
[90]

Mederacke I, Filliol A, Affo S, Nair A, Hernandez C, et al. 2022. The purinergic P2Y14 receptor links hepatocyte death to hepatic stellate cell activation and fibrogenesis in the liver. Science Translational Medicine 14:eabe5795

doi: 10.1126/scitranslmed.abe5795
[91]

Zhan SS, Jiang JX, Wu J, Halsted C, Friedman SL, et al. 2006. Phagocytosis of apoptotic bodies by hepatic stellate cells induces NADPH oxidase and is associated with liver fibrosis in vivo. Hepatology 43:435−443

doi: 10.1002/hep.21093
[92]

Canbay A, Feldstein AE, Higuchi H, Werneburg N, Grambihler A, et al. 2003. Kupffer cell engulfment of apoptotic bodies stimulates death ligand and cytokine expression. Hepatology 38:1188−1198

doi: 10.1053/jhep.2003.50472
[93]

Jiang JX, Chen X, Hsu DK, Baghy K, Serizawa N, et al. 2012. Galectin-3 modulates phagocytosis-induced stellate cell activation and liver fibrosis in vivo. American Journal of Physiology-Gastrointestinal and Liver Physiology 302:G439−G446

doi: 10.1152/ajpgi.00257.2011
[94]

Miyazaki H, Sawada T, Kiyohira M, Yu Z, Nakamura K, et al. 2014. Fatty acid binding protein 7 regulates phagocytosis and cytokine production in Kupffer cells during liver injury. The American Journal of Pathology 184:2505−2515

doi: 10.1016/j.ajpath.2014.05.015
[95]

Wang X, He Q, Zhou C, Xu Y, Liu D, et al. 2023. Prolonged hypernutrition impairs TREM2-dependent efferocytosis to license chronic liver inflammation and NASH development. Immunity 56:58−77.e11

doi: 10.1016/j.immuni.2022.11.013
[96]

Muendlein HI, Connolly WM, Leiriao J, Nolan MA, Judge J, et al. 2025. TNF switches homeostatic efferocytosis to lytic caspase-8–dependent pyroptosis and IL-1β maturation. Science Immunology 10:eadq0043

doi: 10.1126/sciimmunol.adq0043
[97]

Shan S, Chao S, Liu Z, Wang S, Liu Z, et al. 2024. TREM2 protects against inflammation by regulating the release of mito-DAMPs from hepatocytes during liver fibrosis. Free Radical Biology and Medicine 220:154−165

doi: 10.1016/j.freeradbiomed.2024.05.004
[98]

Zhou L, Lu Y, Qiu X, Chen Z, Tang Y, et al. 2025. Lipid droplet efferocytosis attenuates proinflammatory signaling in macrophages via TREM2- and MS4A7-dependent mechanisms. Cell Reports 44:115310

doi: 10.1016/j.celrep.2025.115310
[99]

Yang F, Yang W, Zhang Y, Ye W, Zhao J, et al. 2026. H4K12 lactylation drives TREM2high macrophages differentiation in liver fibrosis. Hepatology 00:Ahead of Print

doi: 10.1097/HEP.0000000000001808
[100]

Li B, Hu J, Xu H. 2024. Integrated single-cell and bulk RNA sequencing reveals immune-related SPP1+ macrophages as a potential strategy for predicting the prognosis and treatment of liver fibrosis and hepatocellular carcinoma. Frontiers in Immunology 15:1455383

doi: 10.3389/fimmu.2024.1455383
[101]

Nusse Y, Kubes P. 2025. Liver macrophages: development, dynamics, and functions. Cellular & Molecular Immunology 22:1178−1189

doi: 10.1038/s41423-025-01298-3
[102]

Mitsui Y, Satoh T. 2025. Functional diversity of disorder-specific macrophages involved in various diseases. Inflammation and Regeneration 45:29

doi: 10.1186/s41232-025-00390-5
[103]

Wang Y, Guan Y, Feng D, Maccioni L, Parra MA, et al. 2025. Infiltrating macrophages replace Kupffer cells and play diverse roles in severe alcohol-associated hepatitis. Cellular & Molecular Immunology 22:1262−1275

doi: 10.1038/s41423-025-01343-1
[104]

Patseas D, El-Masry A, Liu Z, Ramachandran P, Triantafyllou E. 2025. Myeloid cells in chronic liver inflammation. Cellular & Molecular Immunology 22:1237−1261

doi: 10.1038/s41423-025-01324-4
[105]

Lee J, Kim CM, Cha JH, Park JY, Yu YS, et al. 2022. Multiplexed digital spatial protein profiling reveals distinct phenotypes of mononuclear phagocytes in livers with advanced fibrosis. Cells 11:3387

doi: 10.3390/cells11213387
[106]

Huang D, Xu M, Wang H, Zhao Y, Zhang Z, et al. 2025. SIRPα blockade therapy potentiates immunotherapy by inhibiting PD-L1+ myeloid cells in hepatocellular carcinoma. Cell Death & Disease 16:451

doi: 10.1038/s41419-025-07779-7
[107]

Hardesty JE, Warner JB, Song YL, Rouchka EC, McClain CJ, et al. 2023. Resolvin D1 attenuated liver injury caused by chronic ethanol and acute LPS challenge in mice. The FASEB Journal 37:e22705

doi: 10.1096/fj.202200778R
[108]

Kang JW, Choi HS, Shin JK, Lee SM. 2019. Resolvin D1 activates the sphingosine-1-phosphate signaling pathway in murine livers with ischemia/reperfusion injury. Biochemical and Biophysical Research Communications 514:1058−1065

doi: 10.1016/j.bbrc.2019.05.041
[109]

Bae SH, Kim JH, Park TH, Lee K, Lee BI, et al. 2022. BMS794833 inhibits macrophage efferocytosis by directly binding to MERTK and inhibiting its activity. Experimental & Molecular Medicine 54:1450−1460

doi: 10.1038/s12276-022-00840-x
[110]

Bárcena C, Stefanovic M, Tutusaus A, Joannas L, Menéndez A, et al. 2015. Gas6/Axl pathway is activated in chronic liver disease and its targeting reduces fibrosis via hepatic stellate cell inactivation. Journal of Hepatology 63:670−678

doi: 10.1016/j.jhep.2015.04.013
[111]

Chen CC, Chen CY, Yeh CT, Liu YT, Leu YL, et al. 2023. Corylin Attenuates CCl4-induced liver fibrosis in mice by regulating the GAS6/AXL signaling pathway in hepatic stellate cells. International Journal of Molecular Sciences 24:16936

doi: 10.3390/ijms242316936
[112]

Kariolis MS, Miao YR, Diep A, Nash SE, Olcina MM, et al. 2017. Inhibition of the GAS6/AXL pathway augments the efficacy of chemotherapies. Journal of Clinical Investigation 127:183−198

doi: 10.1172/jci85610
[113]

Pan Z, El Sharkway R, Bayoumi A, Metwally M, Gloss BS, et al. 2024. Inhibition of MERTK reduces organ fibrosis in mouse models of fibrotic disease. Science Translational Medicine 16:eadj0133

doi: 10.1126/scitranslmed.adj0133
[114]

Bernsmeier C, Pop OT, Singanayagam A, Triantafyllou E, Patel VC, et al. 2015. Patients with acute-on-chronic liver failure have increased numbers of regulatory immune cells expressing the receptor tyrosine kinase MERTK. Gastroenterology 148:603−615.e14

doi: 10.1053/j.gastro.2014.11.045
[115]

Kim DH, Sung M, Park MS, Sun EG, Yoon S, et al. 2024. Galectin 3-binding protein (LGALS3BP) depletion attenuates hepatic fibrosis by reducing transforming growth factor-β1 (TGF-β1) availability and inhibits hepatocarcinogenesis. Cancer Communications 44:1106−1129

doi: 10.1002/cac2.12600
[116]

Hu K, Gong X, Ai Q, Lin L, Dai J, et al. 2017. Endogenous AMPK acts as a detrimental factor in fulminant hepatitis via potentiating JNK-dependent hepatocyte apoptosis. Cell Death & Disease 8:e2637

doi: 10.1038/cddis.2017.62
[117]

Jang YJ, An SY, Kim JH. 2017. Identification of MFGE8 in mesenchymal stem cell secretome as an anti-fibrotic factor in liver fibrosis. BMB Reports 50:58−59

doi: 10.5483/bmbrep.2017.50.2.012
[118]

Grøndal SM, Tutusaus A, Boix L, Reig M, Blø M, et al. 2024. Dynamic changes in immune cell populations by AXL kinase targeting diminish liver inflammation and fibrosis in experimental MASH. Frontiers in Immunology 15:1400553

doi: 10.3389/fimmu.2024.1400553
[119]

Li HY, Ju D, Zhang DW, Li H, Kong LM, et al. 2015. Activation of TGF-β1-CD147 positive feedback loop in hepatic stellate cells promotes liver fibrosis. Scientific Reports 5:16552

doi: 10.1038/srep16552
[120]

Pan Z, Eslam M. 2024. MERTK and fibrosis: a new target for therapy. DNA and Cell Biology 43:311−314

doi: 10.1089/dna.2024.0099
[121]

Staufer K, Huber H, Zessner-Spitzenberg J, Stauber R, Finkenstedt A, et al. 2023. Gas6 in chronic liver disease — a novel blood-based biomarker for liver fibrosis. Cell Death Discovery 9:282

doi: 10.1038/s41420-023-01551-6
[122]

Matsuda T, Kaji K, Nishimura N, Asada S, Koizumi A, et al. 2024. Cabozantinib prevents the progression of metabolic dysfunction-associated steatohepatitis by inhibiting the activation of hepatic stellate cell and macrophage and attenuating angiogenic activity. Heliyon 10:e38647

doi: 10.1016/j.heliyon.2024.e38647
[123]

Lai YS, Putra RBDS, Aui SP, Chang KT. 2018. M2C polarization by baicalin enhances efferocytosis via upregulation of MERTK receptor. The American Journal of Chinese Medicine 46:1899−1914

doi: 10.1142/S0192415X18500957
[124]

Liu S, Wu J, Stolarz A, Zhang H, Boerma M, et al. 2023. PCSK9 attenuates efferocytosis in endothelial cells and promotes vascular aging. Theranostics 13:2914−2929

doi: 10.7150/thno.83914
[125]

Gao J, Yang Z, Song Y, Shao L, Dong X, et al. 2026. TRIM13 in situ engineering boosts anti-inflammatory capacity of CAR-Ms for liver fibrosis therapy. Nature Communications 17:2077

doi: 10.1038/s41467-026-69858-3
[126]

Laurindo LF, de Carvalho GM, de Oliveira Zanuso B, Figueira ME, Direito R, et al. 2023. Curcumin-based nanomedicines in the treatment of inflammatory and immunomodulated diseases: an evidence-based comprehensive review. Pharmaceutics 15:229

doi: 10.3390/pharmaceutics15010229
[127]

Uz M, Kalaga M, Pothuraju R, Ju J, Junker WM, et al. 2019. Dual delivery nanoscale device for miR-345 and gemcitabine co-delivery to treat pancreatic cancer. Journal of Controlled Release 294:237−246

doi: 10.1016/j.jconrel.2018.12.031
[128]

Li HM, Zhang JY, Wang XQ, Ye LT, Ren B, et al. 2025. Therapeutic potential of PDA@CeO2 in suppressing hepatic stellate cell activation and preventing liver fibrosis. International Journal of Nanomedicine 20:9073−9091

doi: 10.2147/IJN.S521372
[129]

Li F, Cheng Z, Sun J, Cheng X, Li C, et al. 2023. The combination of sinusoidal perfusion enhancement and apoptosis inhibition by riociguat plus a galactose-PEGylated bilirubin multiplexing nanomedicine ameliorates liver fibrosis progression. Nano Letters 23:4126−4135

doi: 10.1021/acs.nanolett.2c04726
[130]

Li Y, Li P, Song J, Zhang X, Xiao H, et al. 2026. Targeting efferocytosis for tissue regeneration: from microenvironment reprogramming to clinical translation. Theranostics 16:3697−3734

doi: 10.7150/thno.126081
[131]

Gong D, Shi W, Yi SJ, Chen H, Groffen J, et al. 2012. TGFβ signaling plays a critical role in promoting alternative macrophage activation. BMC Immunology 13:31

doi: 10.1186/1471-2172-13-31