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Yet, chronic treatments with anti-inflammatory agents, such as methotrexate (MTX), failed to modulate disease progression, possibly for the limited drug bioavailability and plaque deposition. Here, MTX\u2013lipid conjugates, based on 1,2-distearoyl-sn-glycero-3-phosphoethanolamine (DSPE), were integrated in the structure of spherical polymeric nanoparticles (MTX-SPNs) or intercalated in the lipid bilayer of liposomes (MTX-LIP). Although, both nanoparticles were colloidally stable with an average diameter of \u223c200 nm, MTX-LIP exhibited a higher encapsulation efficiency (&amp;gt;70%) and slower release rate (\u223c50% at 10 h) compared to MTX-SPN. In primary bone marrow derived macrophages (BMDMs), MTX-LIP modulated the transcellular transport of oxLDL more efficiently than free MTX mostly by inducing a 2-fold overexpression of ABCA1 (regulating oxLDL efflux), while the effect on CD36 and SRA-1 (regulating oxLDL influx) was minimal. Furthermore, in BMDMs, MTX-LIP showed a stronger anti-inflammatory activity than free MTX, reducing the expression of IL-1\u03b2 by 3-fold, IL-6 by 2-fold, and also moderately of TNF-\u03b1. In 28 days high-fat-diet-fed apoE\u2013\/\u2013 mice, MTX-LIP reduced the mean plaque area by 2-fold and the hematic amounts of RANTES by half as compared to free MTX. These results would suggest that the nanoenhanced delivery to vascular plaques of the anti-inflammatory DSPE-MTX conjugate could effectively modulate the disease progression by halting monocytes\u2019 maturation and recruitment already at the onset of atherosclerosis.<\/jats:p>","DOI":"10.1021\/acsami.0c12202","type":"journal-article","created":{"date-parts":[[2020,7,28]],"date-time":"2020-07-28T21:21:31Z","timestamp":1595971291000},"page":"37943-37956","source":"Crossref","is-referenced-by-count":32,"title":["Modulating\nLipoprotein Transcellular Transport and\nAtherosclerotic Plaque Formation in ApoE\u2013\/\u2013 Mice via Nanoformulated Lipid\u2013Methotrexate Conjugates"],"prefix":"10.1021","volume":"12","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-4216-724X","authenticated-orcid":true,"given":"Valentina","family":"Di Francesco","sequence":"first","affiliation":[{"name":"Fondazione Istituto Italiano di Tecnologia , , Via Morego 30 , ,","place":["Genoa, Italy, 16163"]},{"name":"University of Genoa , , Via Opera Pia, 13 , ,","place":["Genoa, Italy, 16145"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7736-1187","authenticated-orcid":true,"given":"Danila","family":"Gurgone","sequence":"additional","affiliation":[{"name":"University of Glasgow , , ,","place":["Glasgow, United Kingdom, G12 8TA"]},{"name":"University of Naples Federico II , , ,","place":["Naples, Italy, 80131"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9715-3876","authenticated-orcid":true,"given":"Roberto","family":"Palomba","sequence":"additional","affiliation":[{"name":"Fondazione Istituto Italiano di Tecnologia , , Via Morego 30 , ,","place":["Genoa, Italy, 16163"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1560-3571","authenticated-orcid":true,"given":"Miguel Filipe Moreira Marques","family":"Ferreira","sequence":"additional","affiliation":[{"name":"Fondazione Istituto Italiano di Tecnologia , , Via Morego 30 , ,","place":["Genoa, Italy, 16163"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3790-5749","authenticated-orcid":true,"given":"Tiziano","family":"Catelani","sequence":"additional","affiliation":[{"name":"Istituto Italiano di Tecnologia , , via Morego 30 , ,","place":["Genova, Italy, 16163"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3087-2363","authenticated-orcid":true,"given":"Antonio","family":"Cervadoro","sequence":"additional","affiliation":[{"name":"Fondazione Istituto Italiano di Tecnologia , , Via Morego 30 , ,","place":["Genoa, Italy, 16163"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3926-4225","authenticated-orcid":true,"given":"Pasquale","family":"Maffia","sequence":"additional","affiliation":[{"name":"University of Glasgow , , ,","place":["Glasgow, United Kingdom, G12 8TA"]},{"name":"University of Naples Federico II , , ,","place":["Naples, Italy, 80131"]},{"name":"University of Glasgow , , ,","place":["Glasgow, United Kingdom, G12 8TA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6050-4188","authenticated-orcid":true,"given":"Paolo","family":"Decuzzi","sequence":"additional","affiliation":[{"name":"Fondazione Istituto Italiano di Tecnologia , , Via Morego 30 , ,","place":["Genoa, Italy, 16163"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2020,7,28]]},"reference":[{"key":"2026090112145667700_cit1","doi-asserted-by":"crossref","DOI":"10.1016\/j.jacc.2017.10.081","article-title":"Taming Immune\nand Inflammatory Responses to Treat Atherosclerosis","author":"Libby","year":"2018","journal-title":"Journal of the American College of Cardiology"},{"key":"2026090112145667700_cit2","doi-asserted-by":"publisher","first-page":"1078","DOI":"10.1073\/pnas.1322725111","article-title":"Probing Nanoparticle\nTranslocation across the Permeable Endothelium in Experimental Atherosclerosis","volume":"111","author":"Kim","year":"2014","journal-title":"Proc. Natl. Acad. Sci. U. S. A."},{"key":"2026090112145667700_cit3","doi-asserted-by":"publisher","first-page":"341","DOI":"10.1016\/j.cell.2011.04.005","article-title":"Macrophages in the\nPathogenesis of Atherosclerosis","volume":"145","author":"Moore","year":"2011","journal-title":"Cell"},{"key":"2026090112145667700_cit4","doi-asserted-by":"publisher","first-page":"3898","DOI":"10.1111\/bph.13818","article-title":"Targeting\nInflammation to Reduce Cardiovascular Disease Risk: A Realistic Clinical\nProspect?","volume":"174","author":"Welsh","year":"2017","journal-title":"British journal of pharmacology"},{"key":"2026090112145667700_cit5","doi-asserted-by":"publisher","first-page":"1119","DOI":"10.1056\/NEJMoa1707914","article-title":"Antiinflammatory Therapy\nwith Canakinumab for Atherosclerotic Disease","volume":"377","author":"Ridker","year":"2017","journal-title":"N. Engl. J. 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J. Nanomed."},{"key":"2026090112145667700_cit14","doi-asserted-by":"publisher","first-page":"3767","DOI":"10.2147\/IJN.S129324","article-title":"Methotrexate Carried in Lipid Core\nNanoparticles Reduces Myocardial Infarction Size and Improves Cardiac\nFunction in Rats","volume":"12","author":"Maranhao","year":"2017","journal-title":"Int. J. Nanomed."},{"key":"2026090112145667700_cit15","doi-asserted-by":"publisher","first-page":"531","DOI":"10.1007\/s10557-013-6488-3","article-title":"Anti-Atherogenic\nEffects of Methotrexate Carried by a Lipid Nanoemulsion That Binds\nto Ldl Receptors in Cholesterol-Fed Rabbits","volume":"27","author":"Bulgarelli","year":"2013","journal-title":"Cardiovasc. 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Med."},{"key":"2026090112145667700_cit18","doi-asserted-by":"publisher","first-page":"5785","DOI":"10.1021\/acsnano.7b01385","article-title":"Hyaluronan\nNanoparticles\nSelectively Target Plaque-Associated Macrophages and Improve Plaque\nStability in Atherosclerosis","volume":"11","author":"Beldman","year":"2017","journal-title":"ACS Nano"},{"key":"2026090112145667700_cit19","doi-asserted-by":"publisher","first-page":"13888","DOI":"10.1021\/acsami.8b17928","article-title":"Promoting the Delivery\nof Nanoparticles to Atherosclerotic\nPlaques by DNA Coating","volume":"11","author":"Zhang","year":"2019","journal-title":"ACS Appl. Mater. Interfaces"},{"key":"2026090112145667700_cit20","doi-asserted-by":"publisher","first-page":"16","DOI":"10.1038\/ncpcardio1397","article-title":"Hemodynamic\nShear Stress and the Endothelium in Cardiovascular Pathophysiology","volume":"6","author":"Davies","year":"2009","journal-title":"Nat. Clin. Pract. Cardiovasc. Med."},{"key":"2026090112145667700_cit21","doi-asserted-by":"publisher","first-page":"975","DOI":"10.1114\/1.140","article-title":"Finite Element Modeling of Three-Dimensional\nPulsatile Flow in the Abdominal Aorta: Relevance to Atherosclerosis","volume":"26","author":"Taylor","year":"1998","journal-title":"Ann. Biomed. Eng."},{"key":"2026090112145667700_cit22","doi-asserted-by":"publisher","first-page":"585","DOI":"10.1007\/s10237-013-0520-1","article-title":"Vascular Deposition Patterns for\nNanoparticles in an Inflamed Patient-Specific Arterial Tree","volume":"13","author":"Hossain","year":"2014","journal-title":"Biomech. Model. Mechanobiol."},{"key":"2026090112145667700_cit23","doi-asserted-by":"publisher","first-page":"364","DOI":"10.1016\/j.atherosclerosis.2011.04.016","article-title":"Targeting\nTherapeutics to the Vascular Wall in Atherosclerosis\u2013Carrier\nSize Matters","volume":"217","author":"Charoenphol","year":"2011","journal-title":"Atherosclerosis"},{"key":"2026090112145667700_cit24","doi-asserted-by":"publisher","first-page":"13759","DOI":"10.1021\/acsnano.8b08875","article-title":"Nanoparticle-Aided\nCharacterization of Arterial Endothelial Architecture During Atherosclerosis\nProgression and Metabolic Therapy","volume":"13","author":"Beldman","year":"2019","journal-title":"ACS Nano"},{"key":"2026090112145667700_cit25","doi-asserted-by":"publisher","first-page":"109","DOI":"10.1021\/acsnano.7b07720","article-title":"Nanoparticle Functionalization with\nPlatelet Membrane Enables Multifactored\nBiological Targeting and Detection of Atherosclerosis","volume":"12","author":"Wei","year":"2018","journal-title":"ACS Nano"},{"key":"2026090112145667700_cit26","doi-asserted-by":"publisher","first-page":"8885","DOI":"10.1021\/acsnano.5b02611","article-title":"Multifunctional\nNanoparticles Facilitate Molecular Targeting and Mirna Delivery to\nInhibit Atherosclerosis in Apoe\u2013\/\u2013Mice","volume":"9","author":"Kheirolomoom","year":"2015","journal-title":"ACS Nano"},{"key":"2026090112145667700_cit27","doi-asserted-by":"publisher","first-page":"9702","DOI":"10.1021\/acsami.8b18190","article-title":"Sr-a-Targeted Phase-Transition\nNanoparticles for the Detection and Treatment of Atherosclerotic Vulnerable\nPlaques","volume":"11","author":"Ye","year":"2019","journal-title":"ACS Appl. 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Liq."},{"key":"2026090112145667700_cit32","doi-asserted-by":"publisher","first-page":"422","DOI":"10.1093\/rheumatology\/28.5.422","article-title":"Synovial and Serum Levels of Methotrexate\nDuring Methotrexate Therapy\nof Rheumatoid Arthritis","volume":"28","author":"Tishler","year":"1989","journal-title":"Rheumatology"},{"key":"2026090112145667700_cit33","doi-asserted-by":"publisher","first-page":"2139","DOI":"10.1016\/j.nano.2016.05.012","article-title":"Spherical\nPolymeric Nanoconstructs for Combined Chemotherapeutic\nand Anti-Inflammatory Therapies","volume":"12","author":"Lee","year":"2016","journal-title":"Nanomedicine"},{"key":"2026090112145667700_cit34","doi-asserted-by":"publisher","DOI":"10.3791\/58154","article-title":"Correlative\nLight Electron Microscopy (Clem) for Tracking and Imaging Viral Protein\nAssociated Structures in Cryo-Immobilized Cells","author":"Santarella-Mellwig","year":"2018","journal-title":"J. Visualized Exp."},{"key":"2026090112145667700_cit35","doi-asserted-by":"publisher","first-page":"472","DOI":"10.4049\/jimmunol.1601445","article-title":"Complement\nProtein C1q Enhances Macrophage\nFoam Cell Survival and Efferocytosis","volume":"198","author":"Pulanco","year":"2017","journal-title":"J. Immunol."},{"key":"2026090112145667700_cit36","doi-asserted-by":"publisher","first-page":"720","DOI":"10.1172\/JCI113377","article-title":"Stimulated Arachidonate\nMetabolism During Foam Cell Transformation\nof Mouse Peritoneal Macrophages with Oxidized Low Density Lipoprotein","volume":"81","author":"Yokode","year":"1988","journal-title":"J. Clin. Invest."},{"key":"2026090112145667700_cit37","doi-asserted-by":"publisher","first-page":"591","DOI":"10.1111\/tra.12576","article-title":"7-Ketocholesterol Impairs Phagocytosis and Efferocytosis\nVia Dysregulation of Phosphatidylinositol 4, 5-Bisphosphate","volume":"19","author":"Lu","year":"2018","journal-title":"Traffic"},{"key":"2026090112145667700_cit38","doi-asserted-by":"publisher","first-page":"3675","DOI":"10.1002\/art.24040","article-title":"Atheroprotective Effects of Methotrexate on Reverse\nCholesterol Transport Proteins and Foam Cell Transformation in Human\nThp-1 Monocyte\/Macrophages","volume":"58","author":"Reiss","year":"2008","journal-title":"Arthritis Rheum."},{"key":"2026090112145667700_cit39","doi-asserted-by":"publisher","first-page":"139","DOI":"10.1016\/j.vph.2004.08.003","article-title":"Cd36, Oxidized Ldl\nand Ppar Gamma: Pathological Interactions\nin Macrophages and Atherosclerosis","volume":"41","author":"Nicholson","year":"2004","journal-title":"Vasc. Pharmacol."},{"key":"2026090112145667700_cit40","doi-asserted-by":"publisher","first-page":"460","DOI":"10.1016\/j.jconrel.2013.06.012","article-title":"Rosiglitazone-Loaded Nanospheres for Modulating Macrophage-Specific\nInflammation in Obesity","volume":"170","author":"Di Mascolo","year":"2013","journal-title":"J. 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Controlled Release"},{"key":"2026090112145667700_cit43","doi-asserted-by":"publisher","first-page":"5785","DOI":"10.1021\/acsnano.7b01385","article-title":"Hyaluronan\nNanoparticles Selectively Target Plaque-Associated Macrophages and\nImprove Plaque Stability in Atherosclerosis","volume":"11","author":"Beldman","year":"2017","journal-title":"ACS Nano"},{"key":"2026090112145667700_cit44","doi-asserted-by":"publisher","first-page":"E6731","DOI":"10.1073\/pnas.1609629113","article-title":"Immune Cell Screening of a Nanoparticle Library Improves\nAtherosclerosis Therapy","volume":"113","author":"Tang","year":"2016","journal-title":"Proc. Natl. Acad. Sci.\nU. S. A."},{"key":"2026090112145667700_cit45","doi-asserted-by":"publisher","first-page":"21654","DOI":"10.1074\/jbc.272.34.21654","article-title":"Native\nand Modified Low Density Lipoproteins Increase the Functional Expression\nof the Macrophage Class B Scavenger Receptor, Cd36","volume":"272","author":"Han","year":"1997","journal-title":"J. Biol. 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