{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,13]],"date-time":"2026-10-13T16:59:19Z","timestamp":1791910759902,"version":"4.3.4"},"reference-count":207,"publisher":"MDPI AG","issue":"3","license":[{"start":{"date-parts":[[2025,1,24]],"date-time":"2025-01-24T00:00:00Z","timestamp":1737676800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"National Institute of Health","award":["R01HL122937"],"award-info":[{"award-number":["R01HL122937"]}]},{"name":"National Institute of Health","award":["R01HL150007"],"award-info":[{"award-number":["R01HL150007"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Cells"],"abstract":"<jats:p>Lysosomes in mammalian cells are recognized as key digestive organelles, containing a variety of hydrolytic enzymes that enable the processing of both endogenous and exogenous substrates. These organelles digest various macromolecules and recycle them through the autophagy\u2013lysosomal system. Recent research has expanded our understanding of lysosomes, identifying them not only as centers of degradation but also as crucial regulators of nutrient sensing, immunity, secretion, and other vital cellular functions. The lysosomal pathway plays a significant role in vascular regulation and is implicated in diseases such as atherosclerosis. During atherosclerotic plaque formation, macrophages initially engulf large quantities of lipoproteins, triggering pathogenic responses that include lysosomal dysfunction, foam cell formation, and subsequent atherosclerosis development. Lysosomal dysfunction, along with the inefficient degradation of apoptotic cells and the accumulation of modified low-density lipoproteins, negatively impacts atherosclerotic lesion progression. Recent studies have highlighted that lysosomal dysfunction contributes critically to atherosclerosis in a cell- and stage-specific manner. In this review, we discuss the mechanisms of lysosomal biogenesis and its regulatory role in atherosclerotic lesions. Based on these lysosomal functions, we propose that targeting lysosomes could offer a novel therapeutic approach for atherosclerosis, shedding light on the connection between lysosomal dysfunction and disease progression while offering new insights into potential anti-atherosclerotic strategies.<\/jats:p>","DOI":"10.3390\/cells14030183","type":"journal-article","created":{"date-parts":[[2025,1,24]],"date-time":"2025-01-24T03:26:31Z","timestamp":1737689191000},"page":"183","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":13,"title":["Lysosome Functions in Atherosclerosis: A Potential Therapeutic Target"],"prefix":"10.3390","volume":"14","author":[{"given":"Zhengchao","family":"Wang","sequence":"first","affiliation":[{"name":"Department of Pharmacological and Pharmaceutical Sciences, College of Pharmacy, University of Houston, Houston, TX 77204, USA"},{"name":"Provincial Key Laboratory for Developmental Biology and Neurosciences, College of Life Sciences, Fujian Normal University, Fuzhou 350007, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0614-9394","authenticated-orcid":false,"given":"Xiang","family":"Li","sequence":"additional","affiliation":[{"name":"Department of Pharmacological and Pharmaceutical Sciences, College of Pharmacy, University of Houston, Houston, TX 77204, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alexandra K.","family":"Moura","sequence":"additional","affiliation":[{"name":"Department of Pharmacological and Pharmaceutical Sciences, College of Pharmacy, University of Houston, Houston, TX 77204, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0003-4408-2001","authenticated-orcid":false,"given":"Jenny Z.","family":"Hu","sequence":"additional","affiliation":[{"name":"Department of Pharmacological and Pharmaceutical Sciences, College of Pharmacy, University of Houston, Houston, TX 77204, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yun-Ting","family":"Wang","sequence":"additional","affiliation":[{"name":"Department of Pharmacological and Pharmaceutical Sciences, College of Pharmacy, University of Houston, Houston, TX 77204, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8338-3013","authenticated-orcid":false,"given":"Yang","family":"Zhang","sequence":"additional","affiliation":[{"name":"Department of Pharmacological and Pharmaceutical Sciences, College of Pharmacy, University of Houston, Houston, TX 77204, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2025,1,24]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"228","DOI":"10.1016\/0014-4800(74)90057-4","article-title":"Lysosomes of the arterial wall. II. Subcellular fractionation of aortic cells from rabbits with experimantal atheroma","volume":"20","author":"Peters","year":"1974","journal-title":"Exp. Mol. Pathol."},{"key":"ref_2","first-page":"1","article-title":"Lysosomes of the arterial wall. IV. Cytochemical localization of acid phosphatase and catalase in smooth muscle cells and foam cells from rabbit atheromatous aorta","volume":"76","author":"Shio","year":"1974","journal-title":"Am. J. Pathol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"223","DOI":"10.1038\/s41580-023-00676-x","article-title":"Lysosomes as coordinators of cellular catabolism, metabolic signalling and organ physiology","volume":"25","author":"Settembre","year":"2024","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"926","DOI":"10.1038\/s41580-024-00757-5","article-title":"Mechanisms of autophagy-lysosome dysfunction in neurodegenerative diseases","volume":"25","author":"Nixon","year":"2024","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"923","DOI":"10.1038\/s41573-019-0036-1","article-title":"Lysosomes as a therapeutic target","volume":"18","author":"Bonam","year":"2019","journal-title":"Nat. Rev. Drug Discov."},{"key":"ref_6","doi-asserted-by":"crossref","unstructured":"Wojtasi\u0144ska, A., Fr\u0105k, W., Lisi\u0144ska, W., Sapeda, N., M\u0142ynarska, E., Rysz, J., and Franczyk, B. (2023). Novel insights into the molecular mechanisms of atherosclerosis. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms241713434"},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Gencer, S., Evans, B.R., van der Vorst, E.P.C., D\u00f6ring, Y., and Weber, C. (2021). Inflammatory chemokines in atherosclerosis. Cells, 10.","DOI":"10.3390\/cells10020226"},{"key":"ref_8","doi-asserted-by":"crossref","unstructured":"Aronova, A., Tosato, F., Naser, N., and Asare, Y. (2023). Innate immune pathways in atherosclerosis-from signaling to long-term epigenetic reprogramming. Cells, 12.","DOI":"10.20944\/preprints202309.0797.v1"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"233","DOI":"10.1038\/35025203","article-title":"Atherosclerosis","volume":"407","author":"Lusis","year":"2000","journal-title":"Nature"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"1685","DOI":"10.1056\/NEJMra043430","article-title":"Inflammation, atherosclerosis, and coronary artery disease","volume":"352","author":"Hansson","year":"2005","journal-title":"N. Engl. J. Med."},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Jebari-Benslaiman, S., Galicia-Garc\u00eda, U., Larrea-Sebal, A., Olaetxea, J.R., Alloza, I., Vandenbroeck, K., Benito-Vicente, A., and Mart\u00edn, C. (2022). Pathophysiology of atherosclerosis. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms23063346"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"762","DOI":"10.1038\/s41569-024-01037-7","article-title":"Efferocytosis in atherosclerosis","volume":"21","author":"Adkar","year":"2024","journal-title":"Nat. Rev. Cardiol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"312","DOI":"10.1038\/s41569-023-00957-0","article-title":"Programmed death of macrophages in atherosclerosis: Mechanisms and therapeutic targets","volume":"21","author":"Zurek","year":"2024","journal-title":"Nat. Rev. Cardiol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"2110","DOI":"10.1161\/CIRCULATIONAHA.120.049922","article-title":"Fate and state of vascular smooth muscle cells in atherosclerosis","volume":"143","author":"Miano","year":"2021","journal-title":"Circulation"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"727","DOI":"10.1038\/s41569-019-0227-9","article-title":"Vascular smooth muscle cells in atherosclerosis","volume":"16","author":"Basatemur","year":"2019","journal-title":"Nat. Rev. Cardiol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"277","DOI":"10.33594\/000000373","article-title":"Lysosome function in cardiovascular diseases","volume":"55","author":"Bhat","year":"2021","journal-title":"Cell Physiol. Biochem."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"223","DOI":"10.12997\/jla.2023.12.3.223","article-title":"Mechanisms of the Wnt pathways as a potential target pathway in atherosclerosis","volume":"12","author":"Poznyak","year":"2023","journal-title":"J. Lipid Atheroscler."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"101","DOI":"10.1038\/s41580-019-0185-4","article-title":"Lysosomes as dynamic regulators of cell and organismal homeostasis","volume":"21","author":"Ballabio","year":"2020","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"329","DOI":"10.1016\/S0092-8674(88)80026-6","article-title":"The mannose 6-phosphate receptor and the biogenesis of lysosomes","volume":"52","author":"Griffiths","year":"1988","journal-title":"Cell"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"636","DOI":"10.1016\/j.bbamcr.2008.12.008","article-title":"Molecular and cellular basis of lysosomal transmembrane protein dysfunction","volume":"1793","author":"Ruivo","year":"2009","journal-title":"Biochim. Biophys. Acta."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"29737","DOI":"10.1074\/jbc.M202369200","article-title":"Human mannose 6-phosphate-uncovering enzyme is synthesized as a proenzyme that is activated by the endoprotease furin","volume":"277","author":"Do","year":"2002","journal-title":"J. Biol. Chem."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"573","DOI":"10.1111\/jnc.14564","article-title":"Lysosome biogenesis in health and disease","volume":"148","author":"Bajaj","year":"2019","journal-title":"J. Neurochem."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"770","DOI":"10.1016\/j.cell.2007.10.018","article-title":"LIMP-2 is a receptor for lysosomal mannose-6-phosphate-independent targeting of beta-glucocerebrosidase","volume":"131","author":"Reczek","year":"2007","journal-title":"Cell"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"283","DOI":"10.1038\/nrm3565","article-title":"Signals from the lysosome: A control centre for cellular clearance and energy metabolism","volume":"14","author":"Settembre","year":"2013","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"512","DOI":"10.1038\/s41419-022-04966-8","article-title":"Novel role for caspase 1 inhibitor VX765 in suppressing NLRP3 inflammasome assembly and atherosclerosis via promoting mitophagy and efferocytosis","volume":"13","author":"Jin","year":"2022","journal-title":"Cell Death. Dis."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1186\/s13020-020-0286-1","article-title":"13-Methylberberine improves endothelial dysfunction by inhibiting NLRP3 inflammasome activation via autophagy induction in human umbilical vein endothelial cells","volume":"15","author":"Peng","year":"2020","journal-title":"Chin. Med."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"964","DOI":"10.1016\/j.jacbts.2021.09.010","article-title":"Rivaroxaban, a direct oral factor Xa inhibitor, attenuates atherosclerosis by alleviating factor Xa-PAR2-mediated autophagy suppression","volume":"6","author":"Ito","year":"2021","journal-title":"JACC Basic. Transl. Sci."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"1065","DOI":"10.1038\/ncb3407","article-title":"Protein kinase C controls lysosome biogenesis independently of mTORC1","volume":"18","author":"Li","year":"2016","journal-title":"Nat. Cell Biol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"1429","DOI":"10.1126\/science.1204592","article-title":"TFEB links autophagy to lysosomal biogenesis","volume":"332","author":"Settembre","year":"2011","journal-title":"Science"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"942","DOI":"10.1038\/s41467-018-03340-7","article-title":"MAP4K3 mediates amino acid-dependent regulation of autophagy via phosphorylation of TFEB","volume":"9","author":"Hsu","year":"2018","journal-title":"Nat. Commun."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"14338","DOI":"10.1038\/ncomms14338","article-title":"mTORC1-independent TFEB activation via Akt inhibition promotes cellular clearance in neurodegenerative storage diseases","volume":"8","author":"Palmieri","year":"2017","journal-title":"Nat. Commun."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"288","DOI":"10.1038\/ncb3114","article-title":"Lysosomal calcium signalling regulates autophagy through calcineurin and TFEB","volume":"17","author":"Medina","year":"2015","journal-title":"Nat. Cell Biol."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"748","DOI":"10.1038\/cr.2017.45","article-title":"Fasting-induced hormonal regulation of lysosomal function","volume":"27","author":"Chen","year":"2017","journal-title":"Cell Res."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"3852","DOI":"10.1093\/hmg\/ddr306","article-title":"Characterization of the CLEAR network reveals an integrated control of cellular clearance pathways","volume":"20","author":"Palmieri","year":"2011","journal-title":"Hum. Mol. Genet."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1016\/j.tibs.2020.09.004","article-title":"How lysosomes sense, integrate, and cope with stress","volume":"46","author":"Saftig","year":"2021","journal-title":"Trends Biochem. Sci."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1038\/s41556-018-0244-7","article-title":"The lysosome as a cellular centre for signalling, metabolism and quality control","volume":"21","author":"Lawrence","year":"2019","journal-title":"Nat. Cell Biol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1128\/EC.01.1.11-21.2002","article-title":"Autophagy in the eukaryotic cell","volume":"1","author":"Reggiori","year":"2002","journal-title":"Eukaryot. Cell"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"357","DOI":"10.1016\/j.freeradbiomed.2011.10.485","article-title":"NAD(P)H oxidase-dependent intracellular and extracellular O2\u2212 production in coronary arterial myocytes from CD38 knockout mice","volume":"52","author":"Xu","year":"2012","journal-title":"Free Radic. Biol. Med."},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Li, X., Zhang, Y., Xia, M., Gulbins, E., Boini, K.M., and Li, P.L. (2014). Activation of Nlrp3 inflammasomes enhances macrophage lipid-deposition and migration: Implication of a novel role of inflammasome in atherogenesis. PLoS ONE, 9.","DOI":"10.1371\/journal.pone.0087552"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"73229","DOI":"10.18632\/oncotarget.12302","article-title":"Enhancement of endothelial permeability by free fatty acid through lysosomal cathepsin B-mediated Nlrp3 inflammasome activation","volume":"7","author":"Wang","year":"2016","journal-title":"Oncotarget"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"811","DOI":"10.1080\/15548627.2019.1637200","article-title":"Enzyme replacement therapy with recombinant pro-CTSD (cathepsin D) corrects defective proteolysis and autophagy in neuronal ceroid lipofuscinosis","volume":"16","author":"Marques","year":"2020","journal-title":"Autophagy"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"137","DOI":"10.1016\/S0962-8924(03)00005-9","article-title":"At the acidic edge: Emerging functions for lysosomal membrane proteins","volume":"13","author":"Eskelinen","year":"2003","journal-title":"Trends Cell Biol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"918","DOI":"10.1136\/jclinpath-2012-201302","article-title":"Cholesteryl ester storage disease: A rare and possibly treatable cause of premature vascular disease and cirrhosis","volume":"66","author":"Reynolds","year":"2013","journal-title":"J. Clin. Pathol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1186\/s13045-021-01087-1","article-title":"Role of lysosomes in physiological activities, diseases, and therapy","volume":"14","author":"Zhang","year":"2021","journal-title":"J. Hematol. Oncol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"C473","DOI":"10.1152\/ajpcell.00415.2023","article-title":"Cholesterol trafficking, lysosomal function, and atherosclerosis","volume":"326","author":"Skeyni","year":"2024","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"287","DOI":"10.1161\/CIRCRESAHA.120.315890","article-title":"The molecular basis of predicting atherosclerotic cardiovascular disease risk","volume":"128","author":"Nayor","year":"2021","journal-title":"Circ. Res."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"617","DOI":"10.1038\/s42255-024-01015-w","article-title":"Dysregulated cellular metabolism in atherosclerosis: Mediators and therapeutic opportunities","volume":"6","author":"Stroope","year":"2024","journal-title":"Nat. Metab."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"824","DOI":"10.1038\/s41569-024-01023-z","article-title":"Targeting immune cell recruitment in atherosclerosis","volume":"21","author":"Weber","year":"2024","journal-title":"Nat. Rev. Cardiol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"1630","DOI":"10.1016\/j.cell.2022.04.004","article-title":"Atherosclerosis: Recent developments","volume":"185","author":"Lusis","year":"2022","journal-title":"Cell"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"750","DOI":"10.1038\/nature00804","article-title":"Subendothelial retention of atherogenic lipoproteins in early atherosclerosis","volume":"417","author":"Gustafsson","year":"2002","journal-title":"Nature"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"691","DOI":"10.1038\/s41419-023-06206-z","article-title":"Macrophage polarization and metabolism in atherosclerosis","volume":"14","author":"Hou","year":"2023","journal-title":"Cell Death Dis."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"540","DOI":"10.1093\/cvr\/cvy022","article-title":"Smooth muscle cell fate and plasticity in atherosclerosis","volume":"114","author":"Allahverdian","year":"2018","journal-title":"Cardiovasc. Res."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"868","DOI":"10.1038\/nature01323","article-title":"Inflammation in atherosclerosis","volume":"420","author":"Libby","year":"2002","journal-title":"Nature"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"503","DOI":"10.1016\/S0092-8674(01)00238-0","article-title":"Atherosclerosis. the road ahead","volume":"104","author":"Glass","year":"2001","journal-title":"Cell"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"1371","DOI":"10.1056\/NEJM198705283162204","article-title":"Compensatory enlargement of human atherosclerotic coronary arteries","volume":"316","author":"Glagov","year":"1987","journal-title":"N. Engl. J. Med."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"108152","DOI":"10.1016\/j.pharmthera.2022.108152","article-title":"Endothelial shear stress signal transduction and atherogenesis: From mechanisms to therapeutics","volume":"235","author":"He","year":"2022","journal-title":"Pharmacol. Ther."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"295","DOI":"10.1080\/08923973.2022.2043896","article-title":"MiR-140-5p upregulation suppressed beta-glycerophosphate-induced vascular smooth muscle cell calcification via targeting TLR4","volume":"44","author":"Zhang","year":"2022","journal-title":"Immunopharmacol. Immunotoxicol."},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"Chen, Q., Qi, X., Zhang, W., Zhang, Y., Bi, Y., Meng, Q., Bian, H., and Li, Y. (2021). Catalpol inhibits macrophage polarization and prevents postmenopausal atherosclerosis through regulating estrogen eeceptor alpha. Front. Pharmacol., 12.","DOI":"10.3389\/fphar.2021.655081"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"377","DOI":"10.1016\/j.bbrc.2010.03.026","article-title":"The autophagy-lysosome pathway: A novel mechanism involved in the processing of oxidized LDL in human vascular endothelial cells","volume":"394","author":"Zhang","year":"2010","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"E8675","DOI":"10.1073\/pnas.1702223114","article-title":"Autophagy is required for endothelial cell alignment and atheroprotection under physiological blood flow","volume":"114","author":"Vion","year":"2017","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_61","doi-asserted-by":"crossref","unstructured":"Wu, W., Xu, H., Wang, Z., Mao, Y., Yuan, L., Luo, W., Cui, Z., Cui, T., Wang, X.L., and Shen, Y.H. (2015). PINK1-Parkin-mediated mitophagy protects mitochondrial integrity and prevents metabolic stress-induced endothelial injury. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0132499"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"1281","DOI":"10.1038\/nm.3288","article-title":"Autophagy regulates endothelial cell processing, maturation and secretion of von Willebrand factor","volume":"19","author":"Torisu","year":"2013","journal-title":"Nat. Med."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"1244","DOI":"10.1126\/science.1153124","article-title":"Ceramide triggers budding of exosome vesicles into multivesicular endosomes","volume":"319","author":"Trajkovic","year":"2008","journal-title":"Science"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"495","DOI":"10.1161\/ATVBAHA.116.308455","article-title":"Endosialin promotes atherosclerosis through phenotypic remodeling of vascular smooth muscle cells","volume":"37","author":"Hasanov","year":"2017","journal-title":"Arterioscler. Thromb. Vasc. Biol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"692","DOI":"10.1161\/CIRCRESAHA.115.306361","article-title":"Vascular smooth muscle cells in atherosclerosis","volume":"118","author":"Bennett","year":"2016","journal-title":"Circ. Res."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"772","DOI":"10.1161\/hh2101.098617","article-title":"Extracellular nucleotides induce arterial smooth muscle cell migration via osteopontin","volume":"89","author":"Chaulet","year":"2001","journal-title":"Circ. Res."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"980","DOI":"10.1080\/15548627.2020.1741202","article-title":"The P2RY12 receptor promotes VSMC-derived foam cell formation by inhibiting autophagy in advanced atherosclerosis","volume":"17","author":"Pi","year":"2021","journal-title":"Autophagy"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"831","DOI":"10.1161\/CIRCRESAHA.121.320047","article-title":"Autophagy is differentially regulated in leukocyte and nonleukocyte foam cells during atherosclerosis","volume":"130","author":"Robichaud","year":"2022","journal-title":"Circ. Res."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"622","DOI":"10.1093\/cvr\/cvy007","article-title":"Vascular smooth muscle cell death, autophagy and senescence in atherosclerosis","volume":"114","author":"Grootaert","year":"2018","journal-title":"Cardiovasc. Res."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"698","DOI":"10.1038\/s43587-021-00089-5","article-title":"Senescent cells suppress innate smooth muscle cell repair functions in atherosclerosis","volume":"1","author":"Childs","year":"2021","journal-title":"Nat. Aging."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"1511","DOI":"10.1007\/s00109-021-02109-8","article-title":"Lipid accumulation and novel insight into vascular smooth muscle cells in atherosclerosis","volume":"99","author":"Liu","year":"2021","journal-title":"J. Mol. Med."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"435","DOI":"10.1042\/CS20201293","article-title":"Metabolism in atherosclerotic plaques: Immunoregulatory mechanisms in the arterial wall","volume":"136","author":"Forteza","year":"2022","journal-title":"Clin. Sci."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"154939","DOI":"10.1016\/j.cyto.2019.154939","article-title":"Who\u2019s in charge here? Macrophage colony stimulating factor and granulocyte macrophage colony stimulating factor: Competing factors in macrophage polarization","volume":"127","author":"Trus","year":"2020","journal-title":"Cytokine"},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"251","DOI":"10.1016\/j.freeradbiomed.2022.02.010","article-title":"Macrophage polarization by potential nutraceutical compounds: A strategic approach to counteract inflammation in atherosclerosis","volume":"181","author":"Sottero","year":"2022","journal-title":"Free. Radic. Biol. Med."},{"key":"ref_75","doi-asserted-by":"crossref","unstructured":"Mushenkova, N.V., Nikiforov, N.G., Melnichenko, A.A., Kalmykov, V., Shakhpazyan, N.K., Orekhova, V.A., and Orekhov, A.N. (2022). Functional phenotypes of intraplaque macrophages and their distinct roles in atherosclerosis development and atheroinflammation. Biomedicines, 10.","DOI":"10.3390\/biomedicines10020452"},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"898","DOI":"10.1038\/cr.2013.75","article-title":"ROS play a critical role in the differentiation of alternatively activated macrophages and the occurrence of tumor-associated macrophages","volume":"23","author":"Zhang","year":"2013","journal-title":"Cell Res."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1161\/01.ATV.0000150649.39934.13","article-title":"Oxidative stress and vascular disease","volume":"25","author":"Madamanchi","year":"2005","journal-title":"Arterioscler. Thromb. Vasc. Biol."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"421","DOI":"10.1161\/CIRCRESAHA.114.302153","article-title":"Macrophage mitochondrial oxidative stress promotes atherosclerosis and nuclear factor-kappaB-mediated inflammation in macrophages","volume":"114","author":"Wang","year":"2014","journal-title":"Circ. Res."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"e76","DOI":"10.1161\/CIRCRESAHA.116.308362","article-title":"Ogg1-dependent DNA repair regulates NLRP3 inflammasome and prevents atherosclerosis","volume":"119","author":"Tumurkhuu","year":"2016","journal-title":"Circ. Res."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.jnutbio.2018.12.005","article-title":"Macrophage plasticity, polarization and function in response to curcumin, a diet-derived polyphenol, as an immunomodulatory agent","volume":"66","author":"Mohammadi","year":"2019","journal-title":"J. Nutr. Biochem."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"2751","DOI":"10.1007\/s00018-020-03453-7","article-title":"Infection and atherosclerosis: TLR-dependent pathways","volume":"77","author":"Li","year":"2020","journal-title":"Cell Mol. Life. Sci."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"534","DOI":"10.1016\/j.cmet.2012.02.011","article-title":"Autophagy links inflammasomes to atherosclerotic progression","volume":"15","author":"Razani","year":"2012","journal-title":"Cell Metab."},{"key":"ref_83","doi-asserted-by":"crossref","unstructured":"Jian, X., Liu, Y., Zhao, Z., Zhao, L., Wang, D., and Liu, Q. (2019). The role of traditional Chinese medicine in the treatment of atherosclerosis through the regulation of macrophage activity. Biomed. Pharmacother., 118.","DOI":"10.1016\/j.biopha.2019.109375"},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"1083","DOI":"10.1172\/JCI70391","article-title":"Cyclin-dependent kinase inhibitor 2B regulates efferocytosis and atherosclerosis","volume":"124","author":"Kojima","year":"2014","journal-title":"J. Clin. Investig."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"36","DOI":"10.1038\/nri2675","article-title":"Macrophage death and defective inflammation resolution in atherosclerosis","volume":"10","author":"Tabas","year":"2010","journal-title":"Nat. Rev. Immunol."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"1221","DOI":"10.1161\/01.RES.0000170946.56981.5c","article-title":"New insights into the regulation of HDL metabolism and reverse cholesterol transport","volume":"96","author":"Lewis","year":"2005","journal-title":"Circ. Res."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"2292","DOI":"10.1001\/jama.290.17.2292","article-title":"Effect of recombinant ApoA-I Milano on coronary atherosclerosis in patients with acute coronary syndromes: A randomized controlled trial","volume":"290","author":"Nissen","year":"2003","journal-title":"JAMA"},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"re3","DOI":"10.1126\/scisignal.272re3","article-title":"CD36, a scavenger receptor involved in immunity, metabolism, angiogenesis, and behavior","volume":"2","author":"Silverstein","year":"2009","journal-title":"Sci. Signal."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"348","DOI":"10.1161\/01.CIR.99.3.348","article-title":"Shed membrane microparticles with procoagulant potential in human atherosclerotic plaques: A role for apoptosis in plaque thrombogenicity","volume":"99","author":"Mallat","year":"1999","journal-title":"Circulation"},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"1258","DOI":"10.1172\/JCI19628","article-title":"Abundant progenitor cells in the adventitia contribute to atherosclerosis of vein grafts in ApoE-deficient mice","volume":"113","author":"Hu","year":"2004","journal-title":"J. Clin. Investig."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"9349","DOI":"10.1073\/pnas.0711382105","article-title":"A sonic hedgehog signaling domain in the arterial adventitia supports resident Sca1+ smooth muscle progenitor cells","volume":"105","author":"Passman","year":"2008","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"435","DOI":"10.2217\/rme.09.1","article-title":"Vascular regeneration: Engineering the stem cell microenvironment","volume":"4","author":"Sun","year":"2009","journal-title":"Regen. Med."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"440","DOI":"10.1160\/TH09-11-0794","article-title":"The mechanism of stem cell differentiation into smooth muscle cells","volume":"104","author":"Xiao","year":"2010","journal-title":"Thromb. Haemost."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"C342","DOI":"10.1152\/ajpcell.00341.2006","article-title":"Stem cell-derived Sca-1+ progenitors differentiate into smooth muscle cells, which is mediated by collagen IV-integrin alpha1\/beta1\/alphav and PDGF receptor pathways","volume":"292","author":"Xiao","year":"2007","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"C711","DOI":"10.1152\/ajpcell.00442.2008","article-title":"Embryonic stem cell differentiation into smooth muscle cells is mediated by Nox4-produced H2O2","volume":"296","author":"Xiao","year":"2009","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"870","DOI":"10.1161\/CIRCRESAHA.109.211417","article-title":"Crucial role of nrf3 in smooth muscle cell differentiation from stem cells","volume":"106","author":"Pepe","year":"2010","journal-title":"Circ. Res."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"908","DOI":"10.1161\/ATVBAHA.116.307373","article-title":"Mesenchymal stem cells attenuate NADPH oxidase-dependent high mobility group box 1 production and inhibit abdominal aortic aneurysms","volume":"36","author":"Sharma","year":"2016","journal-title":"Arterioscler. Thromb. Vasc. Biol."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"870","DOI":"10.1038\/nm.3651","article-title":"Stem cell aging: Mechanisms, regulators and therapeutic opportunities","volume":"20","author":"Oh","year":"2014","journal-title":"Nat. Med."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"409","DOI":"10.1080\/15548627.2021.1936358","article-title":"Enhanced autophagy in Becn1(F121A\/F121A) knockin mice counteracts aging-related neural stem cell exhaustion and dysfunction","volume":"18","author":"Wang","year":"2022","journal-title":"Autophagy"},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"1277","DOI":"10.1126\/science.aag3048","article-title":"Lysosome activation clears aggregates and enhances quiescent neural stem cell activation during aging","volume":"359","author":"Leeman","year":"2018","journal-title":"Science"},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"1437","DOI":"10.1038\/nm.2539","article-title":"Stem cells and the vasculature","volume":"17","author":"Bautch","year":"2011","journal-title":"Nat. Med."},{"key":"ref_102","doi-asserted-by":"crossref","unstructured":"Bonacina, F., Zhang, X., Manel, N., Yvan-Charvet, L., Razani, B., and Norata, G.D. (Nat. Rev. Cardiol., 2024). Lysosomes in the immunometabolic reprogramming of immune cells in atherosclerosis, Nat. Rev. Cardiol., Online ahead of print.","DOI":"10.1038\/s41569-024-01072-4"},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"2202","DOI":"10.1096\/fj.13-243105","article-title":"Hypercholesterolemia-induced priming of hematopoietic stem and progenitor cells aggravates atherosclerosis","volume":"28","author":"Seijkens","year":"2014","journal-title":"FASEB J."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"253","DOI":"10.1161\/ATVBAHA.114.305144","article-title":"Lymphocytes in atherosclerosis","volume":"35","author":"Hedrick","year":"2015","journal-title":"Arterioscler. Thromb. Vasc. Biol."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"3946320221092188","DOI":"10.1177\/03946320221092188","article-title":"Immune system and atherosclerosis: Hostile or friendly relationship","volume":"36","author":"Razmkhah","year":"2022","journal-title":"Int. J. Immunopathol. Pharmacol."},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"889","DOI":"10.1161\/CIRCRESAHA.111.263186","article-title":"Lymphocytes and the adventitial immune response in atherosclerosis","volume":"110","author":"Campbell","year":"2012","journal-title":"Circ. Res."},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"522","DOI":"10.1038\/s41569-021-00668-4","article-title":"Therapeutic strategies targeting inflammation and immunity in atherosclerosis: How to proceed?","volume":"19","author":"Engelen","year":"2022","journal-title":"Nat. Rev. Cardiol."},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"546","DOI":"10.1016\/j.jacbts.2021.01.006","article-title":"B cells in atherosclerosis: Mechanisms and potential clinical applications","volume":"6","author":"Pattarabanjird","year":"2021","journal-title":"JACC Basic. Transl. Sci."},{"key":"ref_109","doi-asserted-by":"crossref","unstructured":"Vellasamy, D.M., Lee, S.J., Goh, K.W., Goh, B.H., Tang, Y.Q., Ming, L.C., and Yap, W.H. (2022). Targeting immune senescence in atherosclerosis. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms232113059"},{"key":"ref_110","doi-asserted-by":"crossref","unstructured":"Marques, A.R.A., Ramos, C., Machado-Oliveira, G., and Vieira, O.V. (2021). Lysosome (dys)function in atherosclerosis-A big weight on the shoulders of a small organelle. Front. Cell Dev. Biol., 9.","DOI":"10.3389\/fcell.2021.658995"},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"e57265","DOI":"10.15252\/embr.202357265","article-title":"Lysosomes in senescence and aging","volume":"24","author":"Tan","year":"2023","journal-title":"EMBO Rep."},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"518","DOI":"10.1093\/jleuko\/qiad117","article-title":"Lysosomal control of dendritic cell function","volume":"114","author":"Yang","year":"2023","journal-title":"J. Leukoc. Biol."},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"84","DOI":"10.1002\/1873-3468.14742","article-title":"Structural view on autophagosome formation","volume":"598","author":"Noda","year":"2024","journal-title":"FEBS Lett."},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"103767","DOI":"10.1016\/j.drudis.2023.103767","article-title":"Lysosome and related protein degradation technologies","volume":"28","author":"Zheng","year":"2023","journal-title":"Drug. Discov. Today"},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"e347","DOI":"10.1002\/mco2.347","article-title":"Chaperone-mediated autophagy: Molecular mechanisms, biological functions, and diseases","volume":"4","author":"Yao","year":"2023","journal-title":"MedComm"},{"key":"ref_116","doi-asserted-by":"crossref","first-page":"1535","DOI":"10.1089\/ars.2007.1692","article-title":"Ceramide: A novel player in reactive oxygen species-induced signaling?","volume":"9","author":"Dumitru","year":"2007","journal-title":"Antioxid. Redox. Signal."},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"417","DOI":"10.1016\/j.cmet.2014.06.009","article-title":"Deficient chaperone-mediated autophagy in liver leads to metabolic dysregulation","volume":"20","author":"Schneider","year":"2014","journal-title":"Cell Metab."},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"759","DOI":"10.1038\/ncb3166","article-title":"Degradation of lipid droplet-associated proteins by chaperone-mediated autophagy facilitates lipolysis","volume":"17","author":"Kaushik","year":"2015","journal-title":"Nat. Cell Biol."},{"key":"ref_119","doi-asserted-by":"crossref","unstructured":"Oku, M., and Sakai, Y. (2018). Three Distinct Types of Microautophagy Based on Membrane Dynamics and Molecular Machineries. Bioessays, 40.","DOI":"10.1002\/bies.201800008"},{"key":"ref_120","doi-asserted-by":"crossref","first-page":"2505","DOI":"10.1080\/15548627.2022.2096397","article-title":"Chaperone-mediated autophagy protects against atherosclerosis","volume":"18","author":"Cuervo","year":"2022","journal-title":"Autophagy"},{"key":"ref_121","doi-asserted-by":"crossref","unstructured":"Iacano, A.J., Lewis, H., Hazen, J.E., Andro, H., Smith, J.D., and Gulshan, K. (2019). Miltefosine increases macrophage cholesterol release and inhibits NLRP3-inflammasome assembly and IL-1beta release. Sci. Rep., 9.","DOI":"10.1038\/s41598-019-47610-w"},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"221","DOI":"10.1038\/nature09663","article-title":"A role for mitochondria in NLRP3 inflammasome activation","volume":"469","author":"Zhou","year":"2011","journal-title":"Nature"},{"key":"ref_123","doi-asserted-by":"crossref","first-page":"222","DOI":"10.1038\/ni.1980","article-title":"Autophagy proteins regulate innate immune responses by inhibiting the release of mitochondrial DNA mediated by the NALP3 inflammasome","volume":"12","author":"Nakahira","year":"2011","journal-title":"Nat. Immunol."},{"key":"ref_124","doi-asserted-by":"crossref","first-page":"758","DOI":"10.1007\/s10753-020-01375-x","article-title":"Clematichinenoside AR alleviates foam cell formation and the inflammatory response in Ox-LDL-induced RAW264.7 cells by activating autophagy","volume":"44","author":"Diao","year":"2021","journal-title":"Inflammation"},{"key":"ref_125","doi-asserted-by":"crossref","unstructured":"Peng, S., Xu, L.W., Che, X.Y., Xiao, Q.Q., Pu, J., Shao, Q., and He, B. (2018). Atorvastatin inhibits inflammatory response, attenuates lipid deposition, and improves the stability of vulnerable atherosclerotic plaques by modulating autophagy. Front. Pharmacol., 9.","DOI":"10.3389\/fphar.2018.00438"},{"key":"ref_126","doi-asserted-by":"crossref","first-page":"117","DOI":"10.1038\/nri3800","article-title":"Role of innate and adaptive immune mechanisms in cardiac injury and repair","volume":"15","author":"Epelman","year":"2015","journal-title":"Nat. Rev. Immunol."},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"1722","DOI":"10.1161\/CIRCRESAHA.118.311362","article-title":"NLRP3 Inflammasome and the IL-1 Pathway in Atherosclerosis","volume":"122","author":"Grebe","year":"2018","journal-title":"Circ. Res."},{"key":"ref_128","doi-asserted-by":"crossref","first-page":"1357","DOI":"10.1038\/nature08938","article-title":"NLRP3 inflammasomes are required for atherogenesis and activated by cholesterol crystals","volume":"464","author":"Duewell","year":"2010","journal-title":"Nature"},{"key":"ref_129","doi-asserted-by":"crossref","first-page":"2105","DOI":"10.4049\/jimmunol.1601563","article-title":"Oxidized low-density lipoprotein immune complex priming of the Nlrp3 inflammasome involves TLR and FcgammaR cooperation and is dependent on CARD9","volume":"198","author":"Rhoads","year":"2017","journal-title":"J. Immunol."},{"key":"ref_130","doi-asserted-by":"crossref","first-page":"812","DOI":"10.1038\/ni.2639","article-title":"CD36 coordinates NLRP3 inflammasome activation by facilitating intracellular nucleation of soluble ligands into particulate ligands in sterile inflammation","volume":"14","author":"Sheedy","year":"2013","journal-title":"Nat. Immunol."},{"key":"ref_131","doi-asserted-by":"crossref","first-page":"5398","DOI":"10.1182\/blood-2009-10-250803","article-title":"Human NLRP3 inflammasome activation is Nox1-4 independent","volume":"115","author":"Jansen","year":"2010","journal-title":"Blood"},{"key":"ref_132","doi-asserted-by":"crossref","first-page":"1733","DOI":"10.1111\/bph.13425","article-title":"NADPH oxidases in oxidant production by microglia: Activating receptors, pharmacology and association with disease","volume":"174","author":"McBean","year":"2017","journal-title":"Br. J. Pharmacol."},{"key":"ref_133","doi-asserted-by":"crossref","first-page":"746","DOI":"10.1016\/j.hlc.2013.01.012","article-title":"NLRP3 inflammasomes show high expression in aorta of patients with atherosclerosis","volume":"22","author":"Zheng","year":"2013","journal-title":"Heart Lung Circ."},{"key":"ref_134","doi-asserted-by":"crossref","first-page":"e003031","DOI":"10.1161\/JAHA.115.003031","article-title":"NLRP3 inflammasome expression and activation in human atherosclerosis","volume":"5","author":"Folkersen","year":"2016","journal-title":"J. Am. Heart. Assoc."},{"key":"ref_135","doi-asserted-by":"crossref","first-page":"162","DOI":"10.1016\/j.bbrc.2012.07.058","article-title":"Critical role of caspase-1 in vascular inflammation and development of atherosclerosis in Western diet-fed apolipoprotein E-deficient mice","volume":"425","author":"Usui","year":"2012","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_136","doi-asserted-by":"crossref","first-page":"F857","DOI":"10.1152\/ajprenal.00203.2014","article-title":"Mitochondrial dysfunction confers albumin-induced NLRP3 inflammasome activation and renal tubular injury","volume":"308","author":"Zhuang","year":"2015","journal-title":"Am. J. Physiol. Renal. Physiol."},{"key":"ref_137","doi-asserted-by":"crossref","first-page":"443","DOI":"10.5551\/jat.RV17001","article-title":"Role of NLRP3 inflammasomes in atherosclerosis","volume":"24","author":"Karasawa","year":"2017","journal-title":"J. Atheroscler. Thromb."},{"key":"ref_138","doi-asserted-by":"crossref","first-page":"449","DOI":"10.1097\/MOL.0000000000000219","article-title":"Endothelial and smooth muscle cell transformation in atherosclerosis","volume":"26","author":"Lao","year":"2015","journal-title":"Curr. Opin. Lipidol."},{"key":"ref_139","doi-asserted-by":"crossref","first-page":"388","DOI":"10.1097\/MOL.0000000000000209","article-title":"Cholesterol efflux capacity, macrophage reverse cholesterol transport and cardioprotective HDL","volume":"26","author":"Hutchins","year":"2015","journal-title":"Curr. Opin. Lipidol."},{"key":"ref_140","first-page":"9","article-title":"Catastrophic antiphospholipid syndrome: A clinical review","volume":"3","author":"Nayer","year":"2014","journal-title":"J. Nephropathol."},{"key":"ref_141","doi-asserted-by":"crossref","first-page":"1138","DOI":"10.1172\/JCI115932","article-title":"Lysophosphatidylcholine, a component of atherogenic lipoproteins, induces mononuclear leukocyte adhesion molecules in cultured human and rabbit arterial endothelial cells","volume":"90","author":"Kume","year":"1992","journal-title":"J. Clin. Investig."},{"key":"ref_142","doi-asserted-by":"crossref","first-page":"191","DOI":"10.1016\/S0021-9150(03)00097-2","article-title":"Adhesion molecules and atherosclerosis","volume":"170","author":"Blankenberg","year":"2003","journal-title":"Atherosclerosis"},{"key":"ref_143","doi-asserted-by":"crossref","first-page":"283","DOI":"10.1016\/S1537-1891(02)00253-7","article-title":"Regulation of chemokine expression in atherosclerosis","volume":"38","author":"Brooks","year":"2002","journal-title":"Vascul. Pharmacol."},{"key":"ref_144","doi-asserted-by":"crossref","first-page":"297","DOI":"10.1016\/S0021-9150(97)00223-2","article-title":"Monocytic cell adhesion to endothelial cells stimulated by oxidized low density lipoprotein is mediated by distinct endothelial ligands","volume":"136","author":"Erl","year":"1998","journal-title":"Atherosclerosis"},{"key":"ref_145","first-page":"35","article-title":"Combination of metformin with other antioxidants may increase its renoprotective efficacy","volume":"2","author":"Baradaran","year":"2013","journal-title":"J. Renal. Inj. Prev."},{"key":"ref_146","doi-asserted-by":"crossref","first-page":"3883","DOI":"10.1073\/pnas.81.12.3883","article-title":"Modification of low density lipoprotein by endothelial cells involves lipid peroxidation and degradation of low density lipoprotein phospholipids","volume":"81","author":"Steinbrecher","year":"1984","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_147","doi-asserted-by":"crossref","first-page":"458","DOI":"10.1159\/000177139","article-title":"Non-lipid-related effects of 3-hydroxy-3-methylglutaryl coenzyme A reductase inhibitors","volume":"87","author":"Corsini","year":"1996","journal-title":"Cardiology"},{"key":"ref_148","doi-asserted-by":"crossref","first-page":"163","DOI":"10.1172\/JCI10310","article-title":"Increased atherosclerosis in LDL receptor-null mice lacking ACAT1 in macrophages","volume":"107","author":"Fazio","year":"2001","journal-title":"J. Clin. Investig."},{"key":"ref_149","doi-asserted-by":"crossref","first-page":"136","DOI":"10.1097\/MED.0b013e3283507836","article-title":"Early steps in reverse cholesterol transport: Cholesteryl ester hydrolase and other hydrolases","volume":"19","author":"Ghosh","year":"2012","journal-title":"Curr. Opin. Endocrinol. Diabetes. Obes."},{"key":"ref_150","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1042\/bj20031811","article-title":"Hormone-sensitive lipase--new roles for an old enzyme","volume":"379","author":"Yeaman","year":"2004","journal-title":"Biochem. J."},{"key":"ref_151","doi-asserted-by":"crossref","first-page":"653","DOI":"10.1161\/CIRCRESAHA.115.306256","article-title":"Macrophage phenotype and function in different stages of atherosclerosis","volume":"118","author":"Tabas","year":"2016","journal-title":"Circ. Res."},{"key":"ref_152","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1111\/imr.12218","article-title":"Macrophage phenotypes in atherosclerosis","volume":"262","author":"Colin","year":"2014","journal-title":"Immunol. Rev."},{"key":"ref_153","doi-asserted-by":"crossref","first-page":"14","DOI":"10.1093\/cvr\/cvn099","article-title":"Early atherosclerosis in humans: Role of diffuse intimal thickening and extracellular matrix proteoglycans","volume":"79","author":"Nakashima","year":"2008","journal-title":"Cardiovasc. Res."},{"key":"ref_154","doi-asserted-by":"crossref","first-page":"1832","DOI":"10.1161\/CIRCULATIONAHA.106.676890","article-title":"Subendothelial lipoprotein retention as the initiating process in atherosclerosis: Update and therapeutic implications","volume":"116","author":"Tabas","year":"2007","journal-title":"Circulation"},{"key":"ref_155","doi-asserted-by":"crossref","first-page":"115","DOI":"10.1056\/NEJM199901143400207","article-title":"Atherosclerosis--an inflammatory disease","volume":"340","author":"Ross","year":"1999","journal-title":"N. Engl. J. Med."},{"key":"ref_156","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1046\/j.1365-3083.1999.00559.x","article-title":"Detection of B cells and proinflammatory cytokines in atherosclerotic plaques of hypercholesterolaemic apolipoprotein E knockout mice","volume":"50","author":"Zhou","year":"1999","journal-title":"Scand. J. Immunol."},{"key":"ref_157","doi-asserted-by":"crossref","unstructured":"Chellan, B., Rojas, E., Zhang, C., and Hofmann Bowman, M.A. (2018). Enzyme-modified non-oxidized LDL (ELDL) induces human coronary artery smooth muscle cell transformation to a migratory and osteoblast-like phenotype. Sci. Rep., 8.","DOI":"10.1038\/s41598-018-30073-w"},{"key":"ref_158","doi-asserted-by":"crossref","first-page":"535","DOI":"10.1161\/ATVBAHA.114.304029","article-title":"Cholesterol loading reprograms the microRNA-143\/145-myocardin axis to convert aortic smooth muscle cells to a dysfunctional macrophage-like phenotype","volume":"35","author":"Vengrenyuk","year":"2015","journal-title":"Arterioscler. Thromb. Vasc. Biol."},{"key":"ref_159","doi-asserted-by":"crossref","first-page":"56","DOI":"10.1038\/s41572-019-0106-z","article-title":"Atherosclerosis","volume":"5","author":"Libby","year":"2019","journal-title":"Nat. Rev. Dis. Primers."},{"key":"ref_160","doi-asserted-by":"crossref","first-page":"590","DOI":"10.1093\/cvr\/cvy010","article-title":"Role of smooth muscle cells in vascular calcification: Implications in atherosclerosis and arterial stiffness","volume":"114","author":"Durham","year":"2018","journal-title":"Cardiovasc. Res."},{"key":"ref_161","doi-asserted-by":"crossref","first-page":"328","DOI":"10.1161\/ATVBAHA.116.308507","article-title":"Rac2 modulates atherosclerotic calcification by regulating macrophage interleukin-1beta production","volume":"37","author":"Ceneri","year":"2017","journal-title":"Arterioscler. Thromb. Vasc. Biol."},{"key":"ref_162","doi-asserted-by":"crossref","unstructured":"Zhu, Y., and McFarlane, H.E. (2022). Regulation of cellulose synthesis via exocytosis and endocytosis. Curr. Opin. Plant. Biol., 69.","DOI":"10.1016\/j.pbi.2022.102273"},{"key":"ref_163","doi-asserted-by":"crossref","unstructured":"Mironov, A.A., Mironov, A., Sanavio, B., Krol, S., and Beznoussenko, G.V. (2023). Intracellular membrane transport in vascular endothelial cells. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms24065791"},{"key":"ref_164","doi-asserted-by":"crossref","unstructured":"Szewczyk-Roszczenko, O.K., Roszczenko, P., Shmakova, A., Finiuk, N., Holota, S., Lesyk, R., Bielawska, A., Vassetzky, Y., and Bielawski, K. (2023). The chemical inhibitors of endocytosis: From mechanisms to potential clinical applications. Cells, 12.","DOI":"10.3390\/cells12182312"},{"key":"ref_165","doi-asserted-by":"crossref","unstructured":"Wilfling, F., Kaksonen, M., and Stachowiak, J. (2023). Protein condensates as flexible platforms for membrane traffic. Curr. Opin. Cell Biol., 85.","DOI":"10.1016\/j.ceb.2023.102258"},{"key":"ref_166","doi-asserted-by":"crossref","first-page":"619","DOI":"10.1016\/j.cell.2010.02.014","article-title":"Autoimmunity and the clearance of dead cells","volume":"140","author":"Nagata","year":"2010","journal-title":"Cell"},{"key":"ref_167","doi-asserted-by":"crossref","first-page":"545","DOI":"10.1016\/j.cmet.2012.01.022","article-title":"Macrophage autophagy plays a protective role in advanced atherosclerosis","volume":"15","author":"Liao","year":"2012","journal-title":"Cell Metab."},{"key":"ref_168","doi-asserted-by":"crossref","first-page":"139","DOI":"10.1016\/j.cell.2016.02.034","article-title":"Lysosomal disorders drive susceptibility to tuberculosis by compromising macrophage migration","volume":"165","author":"Berg","year":"2016","journal-title":"Cell"},{"key":"ref_169","doi-asserted-by":"crossref","first-page":"166","DOI":"10.1038\/nri3607","article-title":"Apoptotic cell clearance: Basic biology and therapeutic potential","volume":"14","author":"Poon","year":"2014","journal-title":"Nat. Rev. Immunol."},{"key":"ref_170","doi-asserted-by":"crossref","first-page":"398","DOI":"10.1038\/s41580-020-0232-1","article-title":"The clearance of dead cells by efferocytosis","volume":"21","author":"Martinez","year":"2020","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_171","doi-asserted-by":"crossref","unstructured":"Chang, H.F., Schirra, C., Pattu, V., Krause, E., and Becherer, U. (2023). Lytic granule exocytosis at immune synapses: Lessons from neuronal synapses. Front. Immunol., 14.","DOI":"10.3389\/fimmu.2023.1177670"},{"key":"ref_172","doi-asserted-by":"crossref","unstructured":"Buratta, S., Tancini, B., Sagini, K., Delo, F., Chiaradia, E., Urbanelli, L., and Emiliani, C. (2020). Lysosomal exocytosis, exosome release and secretory autophagy: The autophagic- and endo-lysosomal systems go extracellular. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21072576"},{"key":"ref_173","doi-asserted-by":"crossref","unstructured":"Gibson, M.S., Domingues, N., and Vieira, O.V. (2018). Lipid and non-lipid factors affecting macrophage dysfunction and inflammation in atherosclerosis. Front. Physiol., 9.","DOI":"10.3389\/fphys.2018.00654"},{"key":"ref_174","doi-asserted-by":"crossref","first-page":"978","DOI":"10.1161\/HYPERTENSIONAHA.111.180935","article-title":"Cysteine protease cathepsins in atherosclerosis-based vascular disease and its complications","volume":"58","author":"Cheng","year":"2011","journal-title":"Hypertension"},{"key":"ref_175","first-page":"163","article-title":"The function of cathepsins B, D, and X in atherosclerosis","volume":"6","author":"Zhao","year":"2016","journal-title":"Am. J. Cardiovasc. Dis."},{"key":"ref_176","doi-asserted-by":"crossref","first-page":"188","DOI":"10.1161\/ATVBAHA.108.181578","article-title":"Leukocyte cathepsin S is a potent regulator of both cell and matrix turnover in advanced atherosclerosis","volume":"29","author":"Bot","year":"2009","journal-title":"Arterioscler. Thromb. Vasc. Biol."},{"key":"ref_177","doi-asserted-by":"crossref","first-page":"3Q","DOI":"10.1016\/j.amjcard.2006.09.020","article-title":"Atherosclerosis: Disease biology affecting the coronary vasculature","volume":"98","author":"Libby","year":"2006","journal-title":"Am. J. Cardiol."},{"key":"ref_178","doi-asserted-by":"crossref","first-page":"2581","DOI":"10.1194\/jlr.M005702","article-title":"Autophagy plays a protective role in free cholesterol overload-induced death of smooth muscle cells","volume":"51","author":"Xu","year":"2010","journal-title":"J. Lipid. Res."},{"key":"ref_179","doi-asserted-by":"crossref","first-page":"655","DOI":"10.1016\/j.cmet.2011.03.023","article-title":"Autophagy regulates cholesterol efflux from macrophage foam cells via lysosomal acid lipase","volume":"13","author":"Ouimet","year":"2011","journal-title":"Cell Metab."},{"key":"ref_180","doi-asserted-by":"crossref","first-page":"403","DOI":"10.1161\/CIRCGENETICS.110.958728","article-title":"A genome-wide association study identifies LIPA as a susceptibility gene for coronary artery disease","volume":"4","author":"Wild","year":"2011","journal-title":"Circ. Cardiovasc. Genet."},{"key":"ref_181","doi-asserted-by":"crossref","first-page":"1055","DOI":"10.1161\/01.RES.87.11.1055","article-title":"Apoptosis regulates human vascular calcification in vitro: Evidence for initiation of vascular calcification by apoptotic bodies","volume":"87","author":"Proudfoot","year":"2000","journal-title":"Circ. Res."},{"key":"ref_182","doi-asserted-by":"crossref","first-page":"728","DOI":"10.1016\/j.cell.2011.10.026","article-title":"Autophagy: Renovation of cells and tissues","volume":"147","author":"Mizushima","year":"2011","journal-title":"Cell"},{"key":"ref_183","doi-asserted-by":"crossref","unstructured":"Henderson, J.M., Weber, C., and Santovito, D. (2021). Beyond self-recycling: Cell-specific role of autophagy in atherosclerosis. Cells, 10.","DOI":"10.3390\/cells10030625"},{"key":"ref_184","doi-asserted-by":"crossref","first-page":"148","DOI":"10.1038\/nm985","article-title":"Trehalose alleviates polyglutamine-mediated pathology in a mouse model of Huntington disease","volume":"10","author":"Tanaka","year":"2004","journal-title":"Nat. Med."},{"key":"ref_185","doi-asserted-by":"crossref","first-page":"1308","DOI":"10.4161\/auto.25188","article-title":"Trehalose delays the progression of amyotrophic lateral sclerosis by enhancing autophagy in motoneurons","volume":"9","author":"Castillo","year":"2013","journal-title":"Autophagy"},{"key":"ref_186","doi-asserted-by":"crossref","first-page":"3311","DOI":"10.1172\/JCI69625","article-title":"Amyloidogenic peptide oligomer accumulation in autophagy-deficient beta cells induces diabetes","volume":"124","author":"Kim","year":"2014","journal-title":"J. Clin. Investig."},{"key":"ref_187","doi-asserted-by":"crossref","first-page":"1942","DOI":"10.1161\/ATVBAHA.114.303342","article-title":"Induction of lysosomal biogenesis in atherosclerotic macrophages can rescue lipid-induced lysosomal dysfunction and downstream sequelae","volume":"34","author":"Emanuel","year":"2014","journal-title":"Arterioscler. Thromb. Vasc. Biol."},{"key":"ref_188","doi-asserted-by":"crossref","first-page":"1369","DOI":"10.1161\/CIRCRESAHA.117.312333","article-title":"Lysosomal cholesterol hydrolysis couples efferocytosis to anti-inflammatory oxysterol production","volume":"122","author":"Viaud","year":"2018","journal-title":"Circ. Res."},{"key":"ref_189","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1161\/01.ATV.0000107030.22053.1e","article-title":"Reduction of atherosclerotic plaques by lysosomal acid lipase supplementation","volume":"24","author":"Du","year":"2004","journal-title":"Arterioscler. Thromb. Vasc. Biol."},{"key":"ref_190","doi-asserted-by":"crossref","unstructured":"Dubland, J.A., and Francis, G.A. (2015). Lysosomal acid lipase: At the crossroads of normal and atherogenic cholesterol metabolism. Front. Cell Dev. Biol., 3.","DOI":"10.3389\/fcell.2015.00003"},{"key":"ref_191","doi-asserted-by":"crossref","first-page":"jcs221739","DOI":"10.1242\/jcs.221739","article-title":"Lysosomal storage disorders\u2014Challenges, concepts and avenues for therapy: Beyond rare diseases","volume":"132","author":"Marques","year":"2019","journal-title":"J. Cell Sci."},{"key":"ref_192","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1016\/S0022-2275(20)30162-0","article-title":"Increased glycosphingolipid levels in serum and aortae of apolipoprotein E gene knockout mice","volume":"43","author":"Garner","year":"2002","journal-title":"J. Lipid. Res."},{"key":"ref_193","doi-asserted-by":"crossref","first-page":"2403","DOI":"10.1161\/CIRCULATIONAHA.113.007559","article-title":"Inhibition of glycosphingolipid synthesis ameliorates atherosclerosis and arterial stiffness in apolipoprotein E-\/- mice and rabbits fed a high-fat and -cholesterol diet","volume":"129","author":"Chatterjee","year":"2014","journal-title":"Circulation"},{"key":"ref_194","doi-asserted-by":"crossref","first-page":"351","DOI":"10.1038\/s41569-018-0002-3","article-title":"Cysteine protease cathepsins in cardiovascular disease: From basic research to clinical trials","volume":"15","author":"Liu","year":"2018","journal-title":"Nat. Rev. Cardiol."},{"key":"ref_195","doi-asserted-by":"crossref","first-page":"897","DOI":"10.1172\/JCI200314915","article-title":"Deficiency of cathepsin S reduces atherosclerosis in LDL receptor-deficient mice","volume":"111","author":"Sukhova","year":"2003","journal-title":"J. Clin. Investig."},{"key":"ref_196","doi-asserted-by":"crossref","first-page":"98","DOI":"10.1161\/CIRCULATIONAHA.105.561449","article-title":"Disruption of the cathepsin K gene reduces atherosclerosis progression and induces plaque fibrosis but accelerates macrophage foam cell formation","volume":"113","author":"Lutgens","year":"2006","journal-title":"Circulation"},{"key":"ref_197","doi-asserted-by":"crossref","first-page":"2065","DOI":"10.1161\/CIRCULATIONAHA.107.688523","article-title":"Cathepsin L deficiency reduces diet-induced atherosclerosis in low-density lipoprotein receptor-knockout mice","volume":"115","author":"Kitamoto","year":"2007","journal-title":"Circulation"},{"key":"ref_198","doi-asserted-by":"crossref","first-page":"3359","DOI":"10.1111\/bph.13626","article-title":"Continuous administration of the mTORC1 inhibitor everolimus induces tolerance and decreases autophagy in mice","volume":"173","author":"Kurdi","year":"2016","journal-title":"Br. J. Pharmacol."},{"key":"ref_199","doi-asserted-by":"crossref","unstructured":"Zhang, K.S., Schecker, J., Krull, A., Riechert, E., J\u00fcrgensen, L., Kamuf-Schenk, V., Burghaus, J., Kiper, L., Cao Ho, T., and W\u00f6ltje, K. (2019). PRAS40 suppresses atherogenesis through inhibition of mTORC1-dependent pro-inflammatory signaling in endothelial cells. Sci. Rep., 9.","DOI":"10.1038\/s41598-019-53098-1"},{"key":"ref_200","doi-asserted-by":"crossref","first-page":"902","DOI":"10.1038\/35022595","article-title":"Accumulation of autophagic vacuoles and cardiomyopathy in LAMP-2-deficient mice","volume":"406","author":"Tanaka","year":"2000","journal-title":"Nature"},{"key":"ref_201","doi-asserted-by":"crossref","first-page":"15750","DOI":"10.1038\/ncomms15750","article-title":"Exploiting macrophage autophagy-lysosomal biogenesis as a therapy for atherosclerosis","volume":"8","author":"Sergin","year":"2017","journal-title":"Nat. Commun."},{"key":"ref_202","doi-asserted-by":"crossref","first-page":"ra9","DOI":"10.1126\/scisignal.2004754","article-title":"The nutrient-responsive transcription factor TFE3 promotes autophagy, lysosomal biogenesis, and clearance of cellular debris","volume":"7","author":"Martina","year":"2014","journal-title":"Sci. Signal."},{"key":"ref_203","doi-asserted-by":"crossref","first-page":"C1034","DOI":"10.1152\/ajpcell.00294.2019","article-title":"Contribution of transcription factor EB to adipoRon-induced inhibition of arterial smooth muscle cell proliferation and migration","volume":"317","author":"Wang","year":"2019","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"ref_204","doi-asserted-by":"crossref","first-page":"676","DOI":"10.1038\/s41419-019-1931-4","article-title":"Activation of TFEB ameliorates dedifferentiation of arterial smooth muscle cells and neointima formation in mice with high-fat diet","volume":"10","author":"Wang","year":"2019","journal-title":"Cell Death. Dis."},{"key":"ref_205","doi-asserted-by":"crossref","first-page":"225","DOI":"10.1016\/j.tem.2014.03.010","article-title":"Self-eating in the plaque: What macrophage autophagy reveals about atherosclerosis","volume":"25","author":"Sergin","year":"2014","journal-title":"Trends. Endocrinol. Metab."},{"key":"ref_206","doi-asserted-by":"crossref","first-page":"333ra50","DOI":"10.1126\/scitranslmed.aad6100","article-title":"Cyclodextrin promotes atherosclerosis regression via macrophage reprogramming","volume":"8","author":"Zimmer","year":"2016","journal-title":"Sci. Transl. Med."},{"key":"ref_207","doi-asserted-by":"crossref","unstructured":"Coisne, C., Hallier-Vanuxeem, D., Boucau, M.C., Hachani, J., Tilloy, S., Bricout, H., Monflier, E., Wils, D., Serpelloni, M., and Parissaux, X. (2016). Beta-cyclodextrins decrease cholesterol release and ABC-associated transporter expression in smooth muscle cells and aortic endothelial cells. Front. Physiol., 7.","DOI":"10.3389\/fphys.2016.00185"}],"container-title":["Cells"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2073-4409\/14\/3\/183\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,8]],"date-time":"2025-10-08T10:35:09Z","timestamp":1759919709000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2073-4409\/14\/3\/183"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,1,24]]},"references-count":207,"journal-issue":{"issue":"3","published-online":{"date-parts":[[2025,2]]}},"alternative-id":["cells14030183"],"URL":"https:\/\/doi.org\/10.3390\/cells14030183","relation":{},"ISSN":["2073-4409"],"issn-type":[{"value":"2073-4409","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,1,24]]}}}