Oxidative modification of LDL is known to elicit an array of pro-atherogenic responses, but it is generally underappreciated that oxidized LDL (OxLDL) exists in multiple forms, characterized by different degrees of oxidation and different mixtures of bioactive components. The variable effects of OxLDL reported in the literature can be attributed in large part to the heterogeneous nature of the preparations employed. In this review, we first describe the various subclasses and molecular composition of OxLDL, including the variety of minimally modified LDL preparations. We then describe multiple receptors that recognize various species of OxLDL and discuss the mechanisms responsible for the recognition by specific receptors. Furthermore, we discuss the contentious issues such as the nature of OxLDL in vivo and the physiological oxidizing agents, whether oxidation of LDL is a prerequisite for atherogenesis, whether OxLDL is the major source of lipids in foam cells, whether in some cases it actually induces cholesterol depletion, and finally the Janus-like nature of OxLDL in having both pro- and anti-inflammatory effects. Lastly, we extend our review to discuss the role of LDL oxidation in diseases other than atherosclerosis, including diabetes mellitus, and several autoimmune diseases, such as lupus erythematosus, anti-phospholipid syndrome, and rheumatoid arthritis. Antioxid. Redox Signal. 13, 39–75.
ActonS, RigottiA, LandschulzKT, XuS, HobbsHH, KriegerM. Identification of scavenger receptor SR-BI as a high density lipoprotein receptor. Science, 271:518–520. 1996.
2.
ActonSL, SchererPE, LodishHF, KriegerM. Expression cloning of SR-BI, a CD36-related class B scavenger receptor. J Biol Chem, 269:21003–21009. 1994.
AkibaS, ChibaM, MukaidaY, SatoT. Involvement of reactive oxygen species and SP-1 in fibronectin production by oxidized LDL. Biochem Biophys Res Commun, 310:491–497. 2003.
AugeN, GarciaV, Maupas-SchwalmF, LevadeT, SalvayreR, Negre-SalvayreA. Oxidized LDL-induced smooth muscle cell proliferation involves the EGF receptor/PI-3 kinase/Akt and the sphingolipid signaling pathways. Arterioscler Thromb Vasc Biol, 22:1990–1995. 2002.
11.
AugeN, Nikolova–KarakashianM, CarpentierS, ParthasarathyS, Negre–SalvayreA, SalvayreR, MerrillAHJr., LevadeT. Role of sphingosine 1-phosphate in the mitogenesis induced by oxidized low density lipoprotein in smooth muscle cells via activation of sphingomyelinase, ceramidase, and sphingosine kinase. J Biol Chem, 274:21533–21538. 1999.
12.
AviramM. Modified forms of low density lipoprotein in atherosclerosis. Atherosclerosis, 98:1–9. 1993.
13.
BabaevVR, GleavesLA, CarterKJ, SuzukiH, KodamaT, FazioS, LintonMF. Reduced atherosclerotic lesions in mice deficient for total or macrophage-specific expression of scavenger receptor-A. Arterioscler Thromb Vasc Biol, 20:2593–2599. 2000.
14.
BabiyAV, GebickiJM. Decomposition of lipid hydroperoxides enhances the uptake of low density lipoprotein by macrophages. Acta Biochim Pol, 46:31–42. 1999.
15.
BaeYS, LeeJH, ChoiSH, KimS, AlmazanF, WitztumJL, MillerYI. Macrophages generate reactive oxygen species in response to minimally oxidized low-density lipoprotein: Toll-like receptor 4- and spleen tyrosine kinase-dependent activation of NADPH oxidase 2. Circ Res, 104:210–218. 2009.
16.
BalagopalakrishnaC, BhuniaAK, RifkindJM, ChatterjeeS. Minimally modified low density lipoproteins induce aortic smooth muscle cell proliferation via the activation of mitogen activated protein kinase. Mol Cell Biochem, 170:85–89. 1997.
17.
BancellsC, BenitezSn, VillegasS, JorbaO, Ordonez–LlanosJ, Sachez–QuesadaJL. Novel phospholipolytic activities associated with electronegative low-density lipoprotein are involved in increased self-aggregation. Biochemistry, 47:8186–8194. 2008.
BasuSK, GoldsteinJL, AndersonRGW, BrownMS. Degradation of cationized low density lipoprotein and regulation of cholesterol metabolism in homozygous familial hypercholesterolemia fibroblasts. Proc Natl Acad Sci USA, 73:3178–3182. 1976.
BoullierA, BirdDA, ChangIK, DennisEA, FriedmanP, Gillotte–TaylorK, HorkkoS, PalinskiW, QuehenbergerO, ShawP, SteinbergD, TerpstraV, WitztumJL. Scavenger receptors, oOxidized LDL, and atherosclerosis. Ann NY Acad Sci, 947:214–223. 2001.
29.
BoullierA, GillotteKL, HorkkoS, GreenSR, FriedmanP, DennisEA, WitztumJL, SteinbergD, QuehenbergerO. The binding of oxidized low density lipoprotein to mouse CD36 is mediated in part by oxidized phospholipids that are associated with both the lipid and protein moieties of the lipoprotein. J Biol Chem, 275:9163–9169. 2000.
30.
BoullierA, LiY, QuehenbergerO, PalinskiW, TabasI, WitztumJL, MillerYI. Minimally oxidized LDL offsets the apoptotic effects of extensively oxidized LDL and free cholesterol in macrophages. Arterioscler Thromb Vasc Biol, 26:1169–1176. 2006.
31.
BoyanovskyB, KarakashianA, KingK, GiltiayN, Nikolova–KarakashianM. Uptake and metabolism of low density lipoproteins with elevated ceramide content by human microvascular endothelial cells. Implications for the regulation of apoptosis. J Biol Chem, 278:26992–26999. 2003.
32.
BrownAJ, DeanRT, JessupW. Free and esterified oxysterol: formation during copper-oxidation of low density lipoprotein and uptake by macrophages. J Lipid Res, 37:320–335. 1996.
33.
BrownAJ, JessupW. Oxysterols and atherosclerosis. Atherosclerosis, 142:1–28. 1999.
34.
BrownAJ, ManderEL, GelissenIC, KritharidesL, DeanRT, JessupW. Cholesterol and oxysterol metabolism and subcellular distribution in macrophage foam cells: Accumulation of oxidized esters in lysosomes. J Lipid Res, 41:226–237. 2000.
35.
BrownMS, GoldsteinJL. Lipoprotein metabolism in the macrophage. Annu Rev Biochem, 52:223–261. 1983.
36.
BrownMS, GoldsteinJL. A receptor-mediated pathway for cholesterol homeostasis. Science, 232:34–47. 1986.
37.
BucciM, GrattonJP, RudicRD, AcevedoL, RoviezzoF, CirinoG, SessaWC. In vivo delivery of the caveolin-1 scaffolding domain inhibits nitric oxide synthesis and reduces inflammation. Nat Med, 6:1362–1367. 2000.
38.
BurkittMJ. A critical overview of the chemistry of copper-dependent low density lipoprotein oxidation: Roles of lipid hydroperoxides, [alpha]-tocopherol, tThiols, and ceruloplasmin. Arch Biochem Biophys, 394:117–135. 2001.
39.
ByfieldFJ, TikkuS, RothblatGH, GoochKJ, LevitanI. OxLDL increases endothelial stiffness, force generation and network formation. J Lipid Res, 47:715–723. 2006.
40.
CaoG, GarciaCK, WyneKL, SchultzRA, ParkerKL, HobbsHH. Structure and localization of the human gene encoding SR-BI/CLA-1. Evidence for transcriptional control by steroidogenic factor 1. J Biol Chem, 272:33068–33076. 1997.
41.
ChangCL, HsuHY, LinHY, ChiangW, LeeH. Lysophosphatidic acid-induced oxidized low-density lipoprotein uptake is class A scavenger receptor-dependent in macrophages. Prostagland Other Lipid Med, 87:20–25. 2008.
42.
ChenM, MasakiT, SawamuraT. LOX-1, the receptor for oxidized low-density lipoprotein identified from endothelial cells: Implications in endothelial dysfunction and atherosclerosis. Pharmacol Therapeut, 95:89–100. 2002.
43.
ChoiHK, HernánMA, SeegerJD, RobinsJM, WolfeF. Methotrexate and mortality in patients with rheumatoid arthritis: A prospective study. Lancet, 359:1173–1177. 2002.
Collot–TeixeiraS, MartinJ, McDermott–RoeC, PostonR, McGregorJL. CD36 and macrophages in atherosclerosis. Cardiovasc Res, 75:468–477. 2007.
46.
CominaciniL, PasiniAF, GarbinU, DavoliA, TosettiML, CampagnolaM, RigoniA, PastorinoAM, Lo CascioV, SawamuraT. Oxidized low fensity lipoprotein (ox-LDL) binding to ox-LDL receptor-1 in endothelial cells induces the activation of NF-kappa B through an increased production of intracellular reactive oxygen species. J Biol Chem, 275:12633–12638. 2000.
47.
CracowskiJL, DurandT, BessardG. Isoprostanes as a biomarker of lipid peroxidation in humans: Physiology, pharmacology and clinical implications. Trends Pharmacol Sci, 23:360–366. 2002.
48.
CuiMZ, LaagE, SunL, TanM, ZhaoG, XuX. Lysophosphatidic acid induces early growth response gene 1 expression in vascular smooth muscle cells: CRE and SRE mediate the transcription. Arterioscler Thromb Vasc Biol, 26:1029–1035. 2006.
49.
CyrusT, PraticoD, ZhaoL, WitztumJL, RaderDJ, RokachJ, FitzGeraldGA, FunkCD. Absence of 12/15-lipoxygenase expression decreases lipid peroxidation and atherogenesis in apolipoprotein E-deficient mice. Circulation, 103:2277–2282. 2001.
DavidGM. The iron hypothesis. Does iron cause atherosclerosis?Clin Cardiol, 19:925–929. 1996.
52.
de BeerMC, ZhaoZ, WebbNR, van der WesthuyzenDR, de VilliersWJS. Lack of a direct role for macrosialin in oxidized LDL metabolism. J Lipid Res, 44:674–685. 2003.
53.
de WintherMPJ, GijbelsMJJ, van DijkKW, van GorpPJJ, SuzukiH, KodamaT, FrantsRR, HavekesLM, HofkerMH. Scavenger receptor deficiency leads to more complex atherosclerotic lesions in APOE3Leiden transgenic mice. Atherosclerosis, 144:315–321. 1999.
54.
de WintherMPJ, van DijkKW, HavekesLM, HofkerMH. Macrophage scavenger receptor Class A: A multifunctional receptor in aAtherosclerosis. Arterioscler Thromb Vasc Biol, 20:290–297. 2000.
55.
DejagerS, Mietus–SynderM, PitasRE. Oxidized low density lipoproteins bind to the scavenger receptor expressed by rabbit smooth muscle cells and acrophages. Arterioscler Thromb, 13:371–378. 1993.
56.
DixonWG, SymmonsDPM. What effects might anti-TNF[alpha] treatment be expected to have on cardiovascular morbidity and mortality in rheumatoid arthritis? A review of the role of TNF[alpha] in cardiovascular pathophysiology. Annals Rheum Dis, 66:1132–1136. 2007.
57.
DrabM, VerkadeP, ElgerM, KasperM, LohnM, LauterbachB, MenneJ, LindschauC, MendeF, LuftFC, SchedlA, HallerH, KurzchaliaTV. Loss of caveolae, vascular dysfunction, and pulmonary defects in caveolin-1 gene-disrupted mice. Science, 293:2449–2452. 2001.
58.
DuenasAI, AcevesM, Fernandez–PisoneroI, GomezC, OrdunaA, CrespoMS, Garcia–RodriguezC. Selective attenuation of Toll-like receptor 2 signalling may explain the atheroprotective effect of sphingosine 1-phosphate. Cardiovasc Res, 79:537–544. 2008.
59.
ElomaaO, KangasM, SahlbergC, TuukkanenJ, SormunenR, LiakkaA, ThesleffI, KraalG, TryggvasonK. Cloning of a novel bacteria-binding receptor structurally related to scavenger receptors and expressed in a subset of acrophages. Cell, 80:603–609. 1995.
60.
ElshourbagyNA, LiX, TerrettJ, VanHornS, GrossMS, AdamouJE, AndersonKM, WebbCL, LyskoPG. Molecular characterization of a human scavenger receptor, human MARCO. Eur J Biochem, 267:919–926. 2000.
61.
EndemannG, StantonLW, MaddenKS, BryantCM, WhiteRT, ProtterAA. CD36 is a receptor for oxidized low density lipoprotein. J Biol Chem, 268:11811–11816. 1993.
62.
FebbraioM, GuyE, SilversteinRL. Stem cell transplantation reveals that absence of macrophage CD36 is protective against atherosclerosis. Arterioscler Thromb Vasc Biol, 24:2333–2338. 2004.
63.
FebbraioM, PodrezEA, SmithJD, HajjarDP, HazenSL, HoffHF, SharmaK, SilversteinRL. Targeted disruption of the class B scavenger receptor CD36 protects against atherosclerotic lesion development in mice. J Clin Invest, 105:1049–1056. 2000.
FrankPG, LeeH, ParkDS, TandonNN, SchererPE, LisantiMP. Genetic ablation of caveolin-1 confers protection against atherosclerosis. Arterioscler Thromb Vasc Biol, 24:98–105. 2004.
66.
FreemanM, AshkenasJ, ReesDJG, KingsleyDM, CopelandNG, JenkinsNA, KriegerM. An ancient, highly conserved family of cysteine-rich protein domains revealed by cloning type I and type II murine macrophage scavenger receptors. Proc Natl Acad Sci U A, 87:8810–8814. 1990.
67.
FreemanM, EkkelY, RohrerL, PenmanM, FreedmanNJ, ChisolmGM, KriegerM. Expression of type I and type II bovine scavenger receptors in Chinese hamster ovary cells: Lipid droplet accumulation and nonreciprocal cross competition by acetylated and oxidized low density lipoprotein. Proc Natl Acad Sci USA, 88:4931–4935. 1991.
68.
FrostegårdJ, SvenungssonE, WuR, GunnarssonI, LundbergIE, KlareskogL, HörkköS, WitztumJL. Lipid peroxidation is enhanced in patients with systemic lupus erythematosus and is associated with arterial and renal disease manifestations. Arthritis Rheum, 52:192–200. 2005.
69.
FunkCD, CyrusT. 12/15-Lipoxygenase, oxidative modification of LDL and atherogenesis. Trends Cardiovasc Med, 11:116–124. 2004.
70.
GargiuloS, GambaP, SotteroB, BiasiF, ChiarpottoE, ServiddioG, VendemialeG, Poli, LeonarduzziG. The core-aldehyde 9-oxononanoyl cholesterol increases the level of transforming growth factor beta1-specific receptors on promonocytic U937 cell membranes. Aging Cell, 8:77–87. 2009.
71.
Garrido–SánchezL, CardonaF, García–FuentesE, Rojo–MartínezG, Gómez- -ZumaqueroJM, PicónMJ, SoriguerFJ, TinahonesFJ. Anti-oxidized low-density lipoprotein antibody levels are associated with the development of type 2 diabetes mellitus. Eur J Clin Invest, 38:615–621. 2008.
72.
GaubatzJW, GillardBK, MasseyJB, HoogeveenRC, HuangM, LloydEE, RayaJL, YangCY, PownallHJ. Dynamics of dense electronegative low density lipoproteins and their preferential association with lipoprotein phospholipase A2. JLipid Res, 48:348–357. 2007.
73.
GesquiereL, ChoW, SubbaiahPV. Role of group IIa and group V secretory phospholipases A 2 in the metabolism of lipoproteins. Substrate specificities of the enzymes and the regulation of their activities by sphingomyelin. Biochemistry, 41:4911–4920. 2002.
GoldsteinJL, HoYK, BasuSK, BrownMS. Binding site on macrophages that mediates uptake and degradation of acetylated low density lipoprotein, producing massive cholesterol deposition. Proc Natl Acad Sci USA, 76:333–337. 1979.
76.
Gomez–ZumaqueroJM, TinahonesFJ, De RamonE, CampsM, GarridoL, SoriguerFJ. Association of biological markers of activity of systemic lupus erythematosus with levels of anti-oxidized low-density lipoprotein antibodies. Rheumatology, 43:510–513. 2004.
77.
GonzalezMA, SelwynAP. Endothelial function, inflammation, and prognosis in cardiovascular disease. Am J Med, 115:99S–106S. 2003.
78.
GoughPJ, GreavesDR, GordonS. A naturally occurring isoform of the human macrophage scavenger receptor (SR-A) gene generated by alternative splicing blocks modified LDL uptake. J Lipid Res, 39:531–543. 1998.
79.
GoughPJ, GreavesDR, SuzukiH, HakkinenT, HiltunenMO, TurunenM, HerttualaSY, KodamaT, GordonS. Analysis of macrophage scavenger receptor (SR-A) expression in human aortic atherosclerotic lesions. Arterioscler Thromb Vasc Biol, 19:461–471. 1999.
80.
GraierWF, KostnerGM. Glycated low-density lipoprotein and atherogenesis: the missing link between diabetes mellitus and hypercholesterolaemia?Eur J Clin Invest, 27:457–459. 1997.
81.
GrandlM, BaredSM, LiebischG, WernerT, BarlageS, SchmitzG. E-LDL and Ox-LDL differentially regulate ceramide and cholesterol raft microdomains in human Macrophages. Cytometry A, 69:189–191. 2006.
82.
GreenspanP, YuH, MaoF, GutmanRL. Cholesterol deposition in macrophages: foam cell formation mediated by cholesterol-enriched oxidized low density lipoprotein. J Lipid Res, 38:101–109. 1997.
83.
GustinC, Van SteenbruggeM, RaesM. LPA modulates monocyte migration directly and via LPA-stimulated endothelial cells. Am J Physiol Cell Physiol, 295:C905–C914. 2008.
84.
GuyE, KuchibhotlaS, SilversteinR, FebbraioM. Continued inhibition of atherosclerotic lesion development in long-term Western diet fed CD36°/apoE° mice. Atherosclerosis, 192:123–130. 2007.
85.
HaberlandME, OlchCL, FolgelmanAM. Role of lysines in mediating interaction of modified low density lipoproteins with the scavenger receptor of human monocyte macrophages. J Biol Chem, 259:11305–11311. 1984.
86.
HaberlandME, SteinbrecherUP. Modified low-density lipoproteins: Diversity and biological relevance in atherogenesis. Monogr Hum Genet, 14:35. 1992.
87.
HalliwellB, WhitemanM. Measuring reactive species and oxidative damage in vivo and in cell culture: How should you do it and what do the results mean?Br J Pharmacol, 142:231–255. 2004.
88.
HammadSM, TahaTA, NareikaA, JohnsonKR, Lopes–VirellaMF, ObeidLM. Oxidized LDL immune complexes induce release of sphingosine kinase in human U937 monocytic cells. Prostagland Other Lipid Med, 79:126–140. 2006.
89.
HammadSM, TwalWO, BarthJL, SmithKJ, SaadAF, VirellaG, ArgravesWS, Lopes–VirellaMF. Oxidized LDL immune complexes and oxidized LDL differentially affect the expression of genes involved with inflammation and survival in human U937 monocytic cells. Atherosclerosis, 202:394–404. 2009.
HaraY, KusumiY, MitsumataM, LiXK, FujinoM. Lysophosphatidylcholine upregulates LOX-1, chemokine receptors, and activation-related transcription factors in human T-cell line Jurkat. J Thromb Thrombolysis, 26:113–118. 2008.
92.
HaratsD, ShaishA, GeorgeJ, MulkinsM, KuriharaH, LevkovitzH, SigalE. Overexpression of 15-lipoxygenase in vascular endothelium accelerates early atherosclerosis in LDL receptor-deficient mice. Arterioscler Thromb Vasc Biol, 20:2100–2105. 2000.
93.
HarkewiczR, HartvigsenK, AlmazanF, DennisEA, WitztumJL, MillerYI. Cholesteryl ester hydroperoxides are biologically active components of minimally oxidized low density lipoprotein. J Biol Chem, 283:10241–10251. 2008.
94.
HazenSL. Oxidized phospholipids as endogenous pattern recognition ligands in Innate Immunity. J Biol Chem, 283:15527–15531. 2008.
95.
HeineckeJW. Lipoprotein oxidation in cardiovascular disease: Chief culprit or innocent bystander?J Exp Med, 203:813–816. 2006.
96.
HeineckeJW. Mechanisms of oxidative damage of low density lipoprotein in human atherosclerosis. Curr Opin Lipidol, 8:268–274. 1997.
97.
HeineckeJW. Oxidized amino acids: Culprits in human atherosclerosis and indicators of oxidative stress. Free Radical Biol Med, 32:1090–1101. 2002.
98.
HenriksenT, MahoneyEM, SteinbergD. Enhanced macrophage degradation of biologically modified low density lipoprotein. Arteriosclerosis, 149–159. 1983.
99.
HenriksenT, MahoneyEM, SteinbergD. Enhanced macrophage degradation of low density lipoprotein previously incubated with cultured endothelial cells: recognition by receptors for acetylated low density lipoproteins. Proc Natl Acad Sci USA, 78:6499–6503. 1981.
100.
HenryPD. Hyperlipidemic endothelial injury and angiogenesis. Basic Res Cardiol, 89:107–114. 1994.
HiltunenTP, LuomaJS, NikkariT, Yla–HerttualaS. Expression of LDL receptor, VLDL receptor, LDL receptor–related protein, and scavenger receptor in rabbit atherosclerotic lesions: Marked induction of scavenger receptor and VLDL receptor expression during lesion development. Circulation, 97:1079–1086. 1998.
103.
HolopainenJM, MedinaOP, MetsoAJ, KinnunenPKJ. Sphingomyelinase activity associated with human plasma low density lipoprotein. Possible functional implications. J Biol Chem, 275:16484–16489. 2000.
104.
HolvoetP, DonckJ, LandeloosM, BrouwersE, LuijtensK, ArnoutJ, LesaffreE, VanrenterghemY, CollenD. Correlation between oxidized low density lipoproteins and von Willebrand factor in chronic renal failure. Thrombosis Haemost, 76:663–669. 1996.
105.
HolvoetP, PerezG, ZhaoZ, BrouwersE, BernarH, CollenD. Malondialdehyde-modified low density lipoproteins in patients with atherosclerotic disease. J Clin Invest, 95:2611–2619. 1995.
106.
HörkköS, BirdDA, MillerE, ItabeH, LeitingerN, SubbanagounderG, BerlinerJA, FriedmanP, DennisEA, CurtissLK, PalinskiW, WitztumJL. Monoclonal autoantibodies specific for oxidized phospholipids or oxidized phospholipid-protein adducts inhibit macrophage uptake of oxidized low-density lipoproteins. J Clin Invest, 103:117–128. 1999.
107.
HsiehCC, YenMH, YenCH, LauYT. Oxidized low density lipoprotein induces apoptosis via generation of reactive oxygen species in vascular smooth muscle cells. Cardiovasc Res, 49:135–145. 2001.
108.
HuangJT, WelchJS, RicoteM, BinderCJ, WillsonTM, KellyC, WitztumJL, FunkCD, ConradD, GlassCK. Interleukin-4-dependent production of PPAR-[gamma] ligands in macrophages by 12/15-lipoxygenase. Nature, 400:378–382. 1999.
109.
HuangZH, BatesEJ, FerranteJV, HiiCST, PoulosA, RobinsonBS, FerranteA. Inhibition of stimulus-induced endothelial cell intercellular adhesion molecule-1, E-selectin, and vascular cellular adhesion molecule-1 expression by arachidonic acid and its hydroxy and hydroperoxy derivatives. Circ Res, 80:149–158. 1997.
110.
HuberJ, BoechzeltH, KartenB, SurboeckM, BochkovVN, BinderBR, SattlerW, LeitingerN. Oxidized cholesteryl linoleates stimulate endothelial cells to bind monocytes via the extracellular signal-regulated kinase ½ pathway. Arterioscler Thromb Vasc Biol, 22:581–586. 2002.
111.
HughesDA, FraserIP, GordonS. Murine macrophage scavenger receptor: In vivo expression and function as receptor for macrophage adhesion in lymphoid and non-lymphoid organs. Eur J Immunol, 25:466–473. 1995.
112.
HultheJ. Antibodies to oxidized LDL in atherosclerosis development. Clinical and animal studies. Clinica Chimica Acta, 348:1–8. 2004.
113.
HuoY, ZhaoL, HymanMC, ShashkinP, HarryBL, BurcinT, ForlowSB, StarkMA, SmithDF, ClarkeS, SrinivasanS, HedrickCC, PraticoD, WitztumJL, NadlerJL, FunkCD, LeyK. Critical role of macrophage 12/15-lipoxygenase for atherosclerosis in apolipoprotein E-deficient mice. Circulation, 110:2024–2031. 2004.
114.
HuszarD, VarbanML, RinningerF, FeeleyR, AraiT, Fairchild–HuntressV, DonovanMJ, TallAR. Increased LDL cholesterol and atherosclerosis in LDL receptor-deficient mice with attenuated expression of scavenger receptor B1. Arterioscler Thromb Vasc Biol, 20:1068–1073. 2000.
115.
IchiI, NakaharaK, MiyashitaY, HidakaA, KutsukakeS, InoueK, MaruyamaT, MiwaY, HaradaM, TsushimaM, KojoS. Kisei Cohort Study G. Association of ceramides in human plasma with risk factors of atherosclerosis. Lipids, 41:859. 2006.
IshikawaK, NavabM, LeitingerN, FogelmanAM, LusisAJ. Induction of heme oxygenase-1 inhibits the monocyte transmigration induced by mildly oxidized LDL. J Clin Invest, 100:1209–1216. 1997.
118.
ItabeH. Oxidative Modification of LDL: Its Pathological Role in Atherosclerosis. Clin Rev Allergy Immunol, 37:4–11. 2009.
119.
ItabeH, MoriM, FujimotoY, HigashiY, TakanoT. Minimally modified LDL is an oxidized LDL enriched with oxidized phosphatidylcholines. J Biochem (Tokyo), 134:459–465. 2003.
120.
ItabeH, YamamotoH, ImanakaT, ShimamuraK, UchiyamaH, KimuraJ, SanakaT, HataY, TakanoT. Sensitive detection of oxidatively modified low density lipoprotein using a monoclonal antibody. J Lipid Res, 37:45–53. 1996.
121.
JamesMJ, van ReykD, RyeKA, DeanRT, ClelandLG, BarterPJ, JessupW. Low density lipoprotein of synovial fluid in inflammatory joint disease is mildly oxidized. Lipids, 33:1115–1121. 1998.
JeromeW. Advanced atherosclerotic foam cell formation has features of an acquired lysosomal storage disorder. Rejuvenation Res, 9:245–255. 2006.
124.
JessupW, KrithairidesL, StockerR. Lipid oxidation in atherogenesis: An overview. Biochem Soc Trans, 32:134–138. 2004.
125.
JessupW, KritharidesL. Metabolism of oxidized LDL by macrophages. Curr Opin Lipidol, 11:473–481. 2000.
126.
JessupW, ManderEL, DeanRT. The intracellular storage and turnover of apolipoprotein B of oxidized LDL in macrophages. Biochim Biophys Acta, 1126:167–177. 1992.
127.
JialalI, ChaitA. Differences in the metabolism of oxidatively modified low density lipoprotein and acetylated low density lipoprotein by human endothelial cells: Inhibition of cholesterol esterification by oxidatively modified low density lipoprotein. J Lipid Res, 30:1561–1568. 1989.
128.
JiangY, OliverP, DaviesKE, PlattN. Identification and characterization of murine SCARA5, a novel class A scavenger receptor that is expressed by populations of epithelial cells. J Biol Chem, 281:11834–11845. 2006.
129.
KakutaniM, MasakiT, SawamuraT. A platelet–endothelium interaction mediated by lectin-like oxidized low-density lipoprotein receptor-1. Proc Natl Acad Sci USA, 97:360–364. 2000.
130.
KakutaniM, UedaM, NarukoT, MasakiT, SawamuraT. Accumulation of LOX-1 ligand in plasma and atherosclerotic lesions of Watanabe heritable hyperlipidemic rabbits: Identification by a novel enzyme immunoassay. Biochem Biophys Res Commun, 282:180–185. 2001.
131.
KanayamaM, YamaguchiS, ShibataT, ShibataN, KobayashiM, NagaiR, AraiH, TakahashiK, UchidaK. Identification of a serum component that regulates cyclooxygenase-2 gene expression in cooperation with 4-hydroxy-2-nonenal. J Biol Chem, 282:24166–24174. 2007.
132.
KarvonenJ, PaivansaloM, KesaniemiYA, HorkkoS. Immunoglobulin M type of autoantibodies to oxidized low-density lipoprotein has an inverse relation to carotid artery atherosclerosis. Circulation, 108:2107–2112. 2003.
133.
KataokaH, KumeN, MiyamotoS, MinamiM, MoriwakiH, MuraseT, SawamuraT, MasakiT, HashimotoN, KitaT. Expression of lectin-like oxidized low-density lipoprotein receptor-1 in human atherosclerotic lesions. Circulation, 99:3110–3117. 1999.
134.
KatoR, MoriC, KitazatoK, ArataS, ObamaT, MoriM, TakahashiK, AiuchiT, TakanoT, ItabeH. Transient increase in plasma oxidized LDL during the progression of atherosclerosis in apolipoprotein E knockout mice. Arterioscler Thromb Vasc Biol, 29:33–39. 2009.
135.
KawaiY, SaitoA, ShibataN, KobayashiM, YamadaS, OsawaT, UchidaK. Covalent binding of oxidized cholesteryl esters to protein: Implications for oxidative modification of low density lipoprotein and atherosclerosis. J Biol Chem, 278:21040–21049. 2003.
136.
KernH, VolkT, KnauerschieferS, MiethT, RustowB, KoxWJ, SchlameM. Stimulation of monocytes and platelets by short-chain phosphatidylcholines with and without terminal carboxyl group. Biochim Biophys Acta, 1394:33–42. 1998.
KinnunenPKJ, HolopainenJM. Sphingomyelinase activity of LDL: A link between atherosclerosis, ceramide, and apoptosis?Trend Cardiovasc Med, 12:37–42. 2002.
139.
KobayashiK, KishiM, AtsumiT, BertolacciniML, MakinoH, SakairiN, YamamotoI, YasudaT, KhamashtaMA, HughesGRV, KoikeT, VoelkerDR, MatsuuraE. Circulating oxidized LDL forms complexes with {beta}2-glycoprotein I: Implication as an atherogenic autoantigen. J Lipid Res, 44:716–726. 2003.
140.
KodamaT, FreemanM, RohrerL, ZabreckyJ, MatsudairaP, KriegerM. Type I macrophage scavenger receptor contains α-helical and collagen-like coiled coils. Nature, 343:531–535. 1990.
141.
KodamaT, ReddyP, KishimotoC, KriegerM. Purification and characterization of a bovine acetyl low density lipoprotein receptor. Proc Natl Acad Sci USA, 85:9238–9242. 1988.
142.
KonesR. The Jupiter study, CRP screening, and aggressive statin therapy. Implications for the primary prevention of cardiovascular disease. Ther Adv Cardiovasc Dis, 3:309–315. 2009.
143.
KotaniK, SatohN, KatoY, ArakiR, KoyamaK, OkajimaT, TanabeM, OishiM, YamakageH, YamadaK, HattoriM, ShimatsuA. A novel oxidized low-density lipoprotein marker, serum amyloid A-LDL, is associated with obesity and the metabolic syndrome. Atherosclerosis, 204:526–531. 2009.
144.
KowalskyGB, ByfieldFJ, LevitanI. OxLDL facilitates flow-induced realignment of aortic endothelial cells. Am J Physiol Cell Physiol, 295:C332–C340. 2008.
145.
KriegerM, HerzJ. Structures and functions: macrophage scavenger receptors and LDL receptor-related protein (LRP)Annu Rev Biochem, 63:601–637. 1994.
146.
Kris–EthertonPM, LichtensteinAH, HowardBV, SteinbergD, WitztumJL. for the Nutrition Committee of the American Heart Association Council on Nutrition PAaM. Antioxidant vitamin supplements and cardiovascular disease. Circulation, 110:637–641. 2004.
147.
KruthHS. Lipoprotein cholesterol and atherosclerosis. Curr Mol Med, 1:633–653. 2001.
148.
KruthHS, HuangW, IshiiI, ZhangWY. Macrophage foam cell formation with native low density lipoprotein. J Biol Chem, 277:34573–34580. 2002.
149.
KuIA, ImbodenJB, HsuePY, GanzP. Rheumatoid arthritis: A model of systemic inflammation driving atherosclerosis. Circul J, 73:977–985. 2009.
150.
KuchibhotlaS, VanegasD, KennedyDJ, GuyE, NimakoG, MortonRE, FebbraioM. Absence of CD36 protects against atherosclerosis in ApoE knock-out mice with no additional protection provided by absence of scavenger receptor A I/II. Cardiovasc Res, 78:185–196. 2008.
151.
KuhnH, ChaitidisP, RoffeisJ, WaltherM. Arachidonic acid metabolites in the cardiovascular system: The role of lipoxygenase isoforms in atherogenesis with particular emphasis on vascular remodeling. J Cardiovasc Pharmacol, 50:609–620. 2007.
KunjathoorVV, FebbraioM, PodrezEA, MooreKJ, AnderssonL, KoehnS, RheeJS, SilversteinR, HoffHF, FreemanMW. Scavenger receptors Class A-I/II and CD36 are the principal receptors responsible for the uptake of modified low density lipoprotein leading to lipid loading in macrophages. J Biol Chem, 277:49982–49988. 2002.
LandschulzKT, PathakRK, RigottiA, KriegerM, HobbsHH. Regulation of scavenger receptor, class B, type I, a high density lipoprotein receptor, in liver and steroidogenic tissues of the rat. J Clin Invest, 98:984–995. 1996.
156.
LeeHS, SongCY. Oxidized low-density lipoprotein and oxidative stress in the development of glomerulosclerosis. Am J Nephrol, 29:62–70. 2009.
157.
LeeJY, JungGY, HeoHJ, YunMR, ParkJY, BaeSS, HongKW, LeeWS, KimCD. 4-Hydroxynonenal induces vascular smooth muscle cell apoptosis through mitochondrial generation of reactive oxygen species. Toxicol Lett, 166:212–221. 2006.
158.
LeitingerN, WatsonAD, HamaSY, IvandicB, QiaoJH, HuberJ, FaullKF, GrassDS, NavabM, FogelmanAM, de BeerFC, LusisAj, BerlinerJA. Role of group II secretory phospholipase A 2 in atherosclerosis: 2. Potential involvement of biologically active oxidized phospholipids. Arterioscler Thromb Vasc Biol, 19:1291–1298. 1999.
159.
Lemaire–EwingSp, BerthierA, RoyerM, LogetteE, CorcosL, BouchotA, MonierS, PrunetC, RaveneauM, RebeC, DesrumauxC, LizardG, NeelD. 7b- Hydroxycholesterol and 25-hydroxycholesterol-induced interleukin-8 secretion involves a calcium-dependent activation of c-fos via the ERK1/2 signaling pathway in THP-1 cells. Cell Biol Toxicol, 25:127–139. 2009.
160.
LeonarduzziG, ChiarpottoE, BiasiF, PoliG. 4-Hydroxynonenal and cholesterol oxidation products in atherosclerosis. Mol Nut Food Res, 49:1044–1049. 2005.
161.
LiD, MehtaJL. Intracellular signaling of LOX-1 in endothelial cell apoptosis. Circ Res, 104:566–568. 2009.
162.
LiH, FreemanMW, LibbyP. Regulation of smooth muscle cell scavenger receptor expression in vivo by atherogenic diets and in vitro by cytokines. J Clin Invest, 95:122–133. 1995.
163.
LiR, MouillesseauxKP, MontoyaD, CruzD, GharaviN, DunM, KoroniakL, BerlinerJA. Identification of prostaglandin E2 receptor subtype 2 as a receptor activated by OxPAPC. Circ Res, 98:642–650. 2006.
164.
LibbyP. Inflammation in atherosclerosis. Nature, 420:868–874. 2002.
165.
LoidlA, ClausR, IngolicE, DeignerHP, HermetterA. Role of ceramide in activation of stress-associated MAP kinases by minimally modified LDL in vascular smooth muscle cells. Biochim Biophys Acta Mol Basis Dis, 1690:150–158. 2004.
166.
Lopes–VirellaMF, BinzafarN, RackleyS, AkiraT, LaVia M, VirellaG. The uptake of LDL-IC by human macrophages: Predominant involvement of the Fc[gamma]RI receptor. Atherosclerosis, 135:161. 1997.
167.
LopezD, Garcia–ValladaresI, Palafox–SanchezCA, De La TorreIG, KobayashiK, MatsuuraE, LopezLR. Oxidized low-density lipoprotein/β2-glycoprotein I complexes and autoantibodies to oxLig-1/β2-glycoprotein I in patients with systemic lupus erythematosus and antiphospholipid syndrome. Am J Clin Pathol, 121:426–436. 2004.
168.
LopezD, KobayashiK, MerrillJT, MatsuuraE, LopezLR. IgG autoantibodies against β2-glycoprotein I complexed with a lipid ligand derived from oxidized low-density lipoprotein are associated with arterial thrombosis in antiphospholipid syndrome. Clin Devel Immunol, 10:203–211. 2003.
169.
LopezLR, BucknerTR, HurkeyBL, KobayashiBL, MatsuuraE. Determination of oxidized low-density lipoproteins (ox-LDL) versus ox-LDL/β2GPI complexes for the assessment of autoimmune-mediated atherosclerosis. Ann NY Acad Sci, 1109:303–310. 2007.
170.
LordanS, MackrillJJ, O'BrienNM. Oxysterols and mechanisms of apoptotic signaling: Implications in the pathology of degenerative diseases. J Nut Biochem, 20:321–336. 2009.
171.
LougheedM, LumCM, LingW, SuzukiH, KodamaT, SteinbrecherU. High affinity saturable uptake of oxidized low density lipoprotein by macrophages from mice lacking the scavenger receptor class A Type I/II. J Biol Chem, 272:12938–12944. 1997.
172.
LougheedM, ZhangHF, SteinbrecherUP. Oxidized low density lipoprotein is resistant to cathepsins and accumulates within macrophages. J Biol Chem, 266:14519–14525. 1991.
173.
LouridaE, GeorgiadisA, PapavasiliouE, PapathanasiouA, DrososA, TselepisA. Patients with early rheumatoid arthritis exhibit elevated autoantibody titers against mildly oxidized low-density lipoprotein and exhibit decreased activity of the lipoprotein-associated phospholipase A2. Arthritis Res Ther, 9:R19. 2007.
LuJ, YangJH, BurnsAR, ChenHH, TangD, WalterscheidJP, SuzukiSS, YangCY, SawamuraT, ChenCH. Mediation of electronegative low-density lipoprotein signaling by LOX-1: A possible mechanism of endothelial apoptosis. Circ Res, 104:619–627. 2009.
176.
ManderEL, DeanRT, StanleyKK, JessupW. Apolipoprotein B of oxidized LDL accumulates in the lysosomes of macrophages. Biochim Biophys Acta (BBA)–Lipids Lipid Metab, 1212:80–92. 1994.
177.
Manning–TobinJJ, MooreKJ, SeimonTA, BellSA, SharukM, Varez–LeiteJI, de WintherMPJ, TabasI, FreemanMW. Loss of SR-A and CD36 activity reduces atherosclerotic lesion complexity without abrogating foam cell formation in hyperlipidemic mice. Arterioscler Thromb Vasc Biol, 29:19–26. 2009.
178.
MaorI, AviramM. Oxidized low density lipoprotein leads to macrophage accumulation of unesterified cholesterol as a result of lysosomal trapping of the lipoprotein hydrolyzed cholesteryl ester. J Lipid Res, 35:803–819. 1994.
MerryP, GrootveldM, LunecJ, BlakeD. Oxidative damage to lipids within the inflamed human joint provides evidence of radical-mediated hypoxic-reperfusion injury. Am J Clin Nutr, 53:362S–369S. 1991.
188.
MillerYI, ViriyakosolS, BinderCJ, FeramiscoJR, KirklandTN, WitztumJL. Minimally modified LDL binds to CD14, induces macrophage spreading via TLR4/MD-2, and inhibits phagocytosis of apoptotic cells. J Biol Chem, 278:1561–1568. 2003.
189.
MizutaniT, SonodaY, MinegishiT, WakabayashiK, MiyamotoK. Cloning, characterization, and cellular distribution of rat scavenger receptor class B Type I (SRBI) in the ovary. Biochem Biophys Res Commun, 234:499–505. 1997.
190.
MontuschiP, BarnesPJ, RobertsLJ. Isoprostanes: Markers and mediators of oxidative stress. FASEB J, 18:1791–1800. 2004.
MooreKJ, KunjathoorVV, KoehnSL, ManningJJ, TsengAA, SilverJM, McKeeM, FreemanMW. Loss of receptor-mediated lipid uptake via scavenger receptor A or CD36 pathways does not ameliorate atherosclerosis in hyperlipidemic mice. J Clin Invest, 115:2192–2201. 2005.
193.
MoriwakiH, KumeN, SawamuraT, AoyamaT, HoshikawaH, OchiH, NishiE, MasakiT, KitaT. Ligand specificity of LOX-1, a novel receptor for oxidized low-density lipoprotein. Arterioscler Thromb Vasc Biol, 18:1541–1547. 1998.
194.
MorrowJD. The Isoprostanes—Unique products of arachidonate peroxidation: Their role as mediators of oxidant stress. Curr Pharm Des, 12:895–902. 2006.
195.
MurakamiM, KudoI. New phospholipase A2 isozymes with a potential role in atherosclerosis. Curr Opin Lipidol, 14:431–436. 2003.
MurphyJE, TedburyPR, Homer–VanniasinkamS, WalkerJH, PonnambalamS. Biochemistry and cell biology of mammalian scavenger receptors. Atherosclerosis, 182:1. 2005.
198.
NagarajanS. Anti-OxLDL IgG blocks OxLDL interaction with CD36, but promotes Fc[gamma]R, CD32A-dependent inflammatory cell adhesion. Immunol Lett, 108:52–61. 2007.
199.
NagyL, TontonozP, AlvarezJGA, ChenH, EvansRM. Oxidized LDL regulates macrophage gene expression through ligand activation of PPAR gamma. Cell, 93:229–240. 1998.
200.
NaitoM, SuzukiH, MoriT, MatsumotoA, KodamaT, TakahashiK. Coexpression of type I and type II human macrophage scavenger receptors in macrophages of various organs and foam cells in atherosclerotic lesions. Am J Pathol, 141:591–599. 1992.
NakamuraK, FunakoshiH, MiyamotoK, TokunagaF, NakamuraT. Molecular cloning and functional characterization of a human scavenger receptor with C-type lectin (SRCL), a novel member of a scavenger receptor family. Biochem Biophys Res Commun, 280:1028–1035. 2001.
203.
NakamuraK, FunakoshiH, TokunagaF, NakamuraT. Molecular cloning of a mouse scavenger receptor with C-type lectin (SRCL)(1), a novel member of the scavenger receptor family. Biochim Biophys Acta, 1522:53–58. 2001.
204.
NakhjavaniM, EsteghamatiA, AsgaraniF, KhalilzadehO, NikzamirA, SafariR. Association of oxidized low-density lipoprotein and transforming growth factor-beta in type 2 diabetic patients: A cross-sectional study. Trans Res, 153:86–90. 2009.
205.
NavabM, HamaSY, AnantharamaiahGM, HassanK, HoughGP, WatsonAD, ReddyST, SevanianA, FonarowGC, FogelmanAM. Normal high density lipoprotein inhibits three steps in the formation of mildly oxidized low density lipoprotein: Steps 2 and 3. J Lipid Res, 41:1495–1508. 2000.
NicholsonAC. Expression of CD36 in macrophages and atherosclerosis. The role of lipid regulation of PPAR[gamma] signaling. Trends Cardiovasc Med, 14:8–12. 2004.
208.
NicholsonAC, FriedaS, PearceA, SilversteinRL. Oxidized LDL binds to CD36 on human monocyte-derived macrophages and transfected cell lines: Evidence implicating the lipid moiety of the lipoprotein as the binding site. Arterioscler Thromb Vasc Biol, 15:269–275. 1995.
209.
NilssonJ, HanssonGK. Autoimmunity in atherosclerosis: A protective response losing control?J Intern Med, 263:464–478. 2008.
210.
ObinataH, HattoriT, NakaneS, TateiK, IzumiT. Identification of 9- hydroxyoctadecadienoic acid and other oxidized free fatty acids as ligands of the G protein-coupled receptor G2A. J Biol Chem, 280:40676–40683. 2005.
211.
OhtaniK, SuzukiY, EdaS, KawaiT, KaseT, KeshiH, SakaiY, FukuohA, SakamotoT, ItabeH, SuzutaniT, OgasawaraM, YoshidaI, WakamiyaN. The membrane-type collectin CL-P1 is a scavenger eeceptor on vascular endothelial cells. J Biol Chem, 276:44222–44228. 2001.
212.
OkajimaF. Plasma lipoproteins behave as carriers of extracellular sphingosine 1- phosphate: Is this an atherogenic mediator or an anti-atherogenic mediator?Biochim Biophys Acta, 1582:132–137. 2002.
213.
PalinskiW, HorkkoS, MillerE, SteinbrecherUP, PowellHC, CurtissLK, WitztumJL. Cloning of monoclonal autoantibodies to epitopes of oxidized lipoproteins from apolipoprotein e-deficient mice. Demonstration of epitopes of oxidized low density lipoprotein in human plasma. J Clin Invest, 98:800–814. 1996.
214.
ParkYM, FebbraioM, SilversteinRL. CD36 modulates migration of mouse and human macrophages in response to oxidized LDL and may contribute to macrophage trapping in the arterial intima. J Clin Invest, 119:136–145. 2009.
215.
ParsonsMS, BarrettL, LittleC, GrantMD. Harnessing CD36 to rein in inflammation, Endocr Metab Immune. Disord Drug Targets, 8:184–191. 2008.
216.
ParthasarathyS, FongLG, OteroD, SteinbergD. Recognition of solubilized apoproteins from delipidated, oxidized low density lipoprotein (LDL) by the acetyl-LDL receptor. Proc Natl Acad Sci USA, 84:537–540. 1987.
217.
ParthasarathyS, LitvinovD, SelvarajanK, GarelnabiM. Lipid peroxidation and decomposition. Conflicting roles in plaque vulnerability and stability. Biochim Biophys Acta, 1781:221–231. 2008.
218.
PearsonA, LuxA, KriegerM. Expression cloning of dSR-CI, a class C macrophage-specific scavenger receptor from Drosophila melanogaster. Proc Natl Acad Sci USA, 92:4056–4060. 1995.
219.
PengoV, BisonE, RuffattiA, IlicetoS. Antibodies to oxidized LDL/[beta]2- glycoprotein I in antiphospholipid syndrome patients with venous and arterial thromboembolism. Thromb Res, 122:556–559. 2008.
220.
PetersMJL, van HalmVP, NurmohamedMT, DamoiseauxJ, TervaertJWC, TwiskJWR, DijkmansBAC, VoskuylAE. Relations between autoantibodies against oxidized low-density lipoprotein, inflammation, subclinical atherosclerosis, and cardiovascular disease in rheumatoid arthritis. J Rheumatol, 35:1495–1499. 2008.
221.
PitasRE. Expression of the acetyl low density lipoprotein receptor by rabbit fibroblasts and smooth muscle cells. Up-regulation by phorbol esters. J Biol Chem, 265:12722–12727. 1990.
222.
PitasRE, BoylesJ, MahleyRW, BissellDM. Uptake of chemically modified low density lipoproteins in vivo is mediated by specific endothelial cells. J Cell Biol, 100:103–117. 1985.
223.
PlüddemannA, MukhopadhyayS, GordonS. The interaction of macrophage receptors with bacterial ligands. Expert Rev Mol Med, 22:1–25. 2006.
PodrezEA, FebbraioM, SheibaniN, SchmittD, SilversteinRL, HajjarDP, CohenPA, FrazierWA, HoffHF, HazenSL. Macrophage scavenger receptor CD36 is the major receptor for LDL modified by monocyte-generated reactive nitrogen species. J Clin Invest, 105:1095–1108. 2000.
226.
PodrezEA, HoppeG, O'NeilJ, HoffHF. Phospholipids in oxidized LDL not adducted to apoB are recognized by the CD36 scavenger receptor. Free Rad Biol Med, 34:356–364. 2003.
227.
PodrezEA, HoppeG, O'NeilJ, SayreLM, SheibaniN, HoffHF. Macrophage receptors responsible for distinct recognition of low density lipoprotein containing pyrrole or pyridinium adducts: Models of oxidized low density lipoprotein. J Lipid Res, 41:1455–1463. 2000.
228.
PodrezEA, PoliakovE, ShenZ, ZhangR, DengY, SunM, FintonPJ, ShanL, GugiuB, FoxPL, HoffHF, SalomonRG, HazenSL. Identification of a novel fFamily of oxidized phospholipids that serve as ligands for the macrophage scavenger receptor CD36. J Biol Chem, 277:38503–38516. 2002.
229.
QuinnMT, ParthasarathyS, SteinbergD. Lysophosphatidylcholine: A chemotactic factor for human monocytes and its potential role in atherogenesis. Proc NatlAcad Sci USA, 85:2805–2809. 1988.
230.
RaghavamenonA, GarelnabiM, BabuS, AldrichA, LitvinovD, ParthasarathyS. Tocopherol is ineffective in preventing the decomposition of preformed lipid peroxides and may promote the accumulation of toxic aldehydes: A potential explanation for the failure of antioxidants to affect human atherosclerosis. Antioxid Redox Signal, 11:1237–1248. 2009.
231.
RavetchJV, KinetJP. Fc Receptors. Annu Rev Immunol, 9:457–492. 1991.
232.
RigottiA, ActonSL, KriegerM. The Class B scavenger receptors SR-BI and CD36 are receptors for anionic phospholipids. J Biol Chem, 270:16221–16224. 1995.
233.
RigottiA, TrigattiBL, PenmanM, RayburnH, HerzJ, KriegerM. A targeted mutation in the murine gene encoding the high density lipoprotein (HDL) receptor scavenger receptor class B type I reveals its key role in HDL metabolism. Proc Natl Acad Sci USA, 94:12610–12615. 1997.
234.
RiosFJ, JancarS, MeloIB, KetelhuthDF, GidlundM. Role of PPAR-gamma in the modulation of CD36 and FcgammaRII induced by LDL with low and high degrees of oxidation during the differentiation of the monocytic THP-1 cell line. Cell Physiol Biochem, 22:549–556. 2008.
235.
RobbesynF, SalvayreR, Negre–SalvayreA. Dual role of oxidized LDL on the NF-KappaB signaling pathway. Free Rad Res, 38:541–551. 2004.
236.
RohrerL, FreemanM, KodamaT, PenmanM, KriegerM. Coiled-coil fibrous domains mediate ligand binding by macrophage scavenger receptor type II. Nature, 343:570–572. 1990.
RomaP, CatapanoAL, BertulliSM, VaresiL, FumagalliR, BerniniF. Oxidized LDL increase free cholesterol and fail to stimulate cholesterol esterification in murine macrophages. Biochem Biophys Res Commun, 171:123–131. 1990.
239.
RosenfeldME, PalinskiW, Yla–HerttualaS, ButlerS, WitztumJL. Distribution of oxidation specific lipid-protein adducts and apolipoprotein B in atherosclerotic lesions of varying severity from WHHL rabbits. Arteriosclerosis, 10:336–349. 1990.
240.
RossR. Atherosclerosis—An inflammatory disease. N Engl J Med, 340:115–126. 1999.
241.
SaadAF, VirellaG, ChassereauC, BoackleRJ, Lopes–VirellaMF. OxLDL immune complexes activate complement and induce cytokine production by MonoMac 6 cells and human macrophages. J Lipid Res, 47:1975–1983. 2006.
242.
SakaguchiH, TakeyaM, SuzukiH, HakamataH, KodamaT, HoriuchiS, GordonS, van der LaanLJ, KraalG, IshibashiS, KitamuraN, TakahashiK. Role of macrophage scavenger receptors in diet-induced atherosclerosis in mice. Lab Invest, 78:423–434. 1998.
243.
SalonenJT, KorpelaH, SalonenR, NyyssonenK, Yla–HerttualaS, YamamotoR, ButlerS, PalinskiW, WitztumJL. Autoantibody against oxidised LDL and progression of carotid atherosclerosis. The Lancet, 339:883–887. 1992.
244.
SambranoGR, TerpstraV, SteinbergD. Independent mechanisms for macrophage binding and macrophage phagocytosis of damaged erythrocytes: Evidence of receptor cooperativity. Arterioscler Thromb Vasc Biol, 17:3442–3448. 1997.
SantanamN, ParthasarathyS. Paradoxical actions of antioxidants in the oxidation of low density lipoprotein by peroxidases. J Clin Invest, 95:2594–2600. 1995.
SchisselSL, Tweedie–HardmanJ, RappJH, GrahamG, WilliamsKJ, TabasI. Rabbit aorta and human atherosclerotic lesions hydrolyze the sphingomyelin of retained low-density lipoprotein. Proposed role for arterial wall sphingomyelinase in subendothelial retention and aggregation of atherogenic lipoproteins. J Clin Invest, 98:1455–1464. 1996.
250.
SelmanL, SkjodtK, NielsenO, FloridonC, HolmskovU, HansenS. Expression and tissue localization of collectin placenta 1 (CL-P1, SRCL) in human tissues. Mol Immunol, 45:3278. 2008.
251.
SevanianA, BittolobonG, CazzolatoG, HodisH, HwangJ, ZamburliniA, MaiorinoM, UrsiniF. Ldl(-) is a lipid hydroperoxide-enriched circulating lipoprotein. J Lipid Res, 38:419–428. 1997.
252.
SevanianA, HodisHN, HwangJ, McLeodLL, PetersonH. Characterization of endothelial cell injury by cholesterol oxidation products found in oxidized LDL. J Lipid Res, 36:1971–1986. 1995.
253.
ShaulPW. Endothelial nitric oxide synthase, caveolae, and the development of atherosclerosis. J Physiol, 15:21–33. 2003.
254.
ShawPX, GoodyearCS, ChangMK, WitztumJL, SilvermanGJ. The autoreactivity of anti-phosphorylcholine antibodies for atherosclerosis-associated neo-antigens and apoptotic cells. J Immunol, 15:6151–6157. 2003.
255.
ShihHH, ZhangS, CaoW, HahnA, WangJ, PaulsenJE, HarnishDC. CRP is a novel ligand for the oxidized LDL receptor LOX-1. Am J Physiol Heart Circ Physiol, 296:H1643–H1650. 2009.
SiessW. Platelet interaction with bioactive lipids formed by mild oxidation of low- density lipoprotein. Pathophysiol Haemost Thromb, 35:292–304. 2006.
258.
SiessW, TigyiG. Thrombogenic and atherogenic activities of lysophosphatidic acid. J Cell Biochem, 92:1086–1094. 2004.
259.
SiessW, ZanglKJ, EsslerM, BauerM, BrandlR, CorrinthC, BittmanR, TigyiG, AepfelbacherM. Lysophosphatidic acid mediates the rapid activation of platelets and endothelial cells by mildly oxidized low density lipoprotein and accumulates in human atherosclerotic lesions. Proc Natl Acad Sci USA, 96:6931–6936. 1999.
SinghDK, SubbaiahPV. Modulation of the activity and arachidonic acid selectivity of group X secretory phospholipase A 2 by sphingolipids. J Lipid Res, 48:683–692. 2007.
262.
StantonLW, WhiteRT, BryantCM, ProtterAA, EndemannG. A macrophage Fc receptor for IgG is also a receptor for oxidized low density lipoprotein. J Biol Chem, 267:22446–22451. 1992.
263.
StaryHC, ChandlerAB, GlagovS, GuytonJR, InsullWJ, E.RM, SchafferSA, J.SC, WagnerWD, WisslerRW. A report from the Committee on Vascular Lesions of the Council on Arteriosclerosis, American Heart Association. Circulation, 89:2462–2478. 1994.
264.
SteinbergD, ParthasarathyS, CarewTE, KhooJC, WitztumJL. Beyond cholesterol. Modifications of low-density lipoprotein that increase its atherogenicity. N Engl J Med, 320:915–924. 1989.
265.
SteinbrecherUP. Oxidation of human low density lipoprotein results in derivatization of lysine residues of apolipoprotein B by lipid peroxide decomposition products. J Biol Chem, 262:3603–3608. 1987.
266.
SteinbrecherUP. Receptors for oxidized low density lipoprotein. Biochim Biophys Acta, 1436:279–298. 1999.
267.
SteinbrecherUP, WitztumJL, ParthasarathyS, SteinbergD. Decrease in reactive amino groups during oxidation or endothelial cell modification of LDL. Correlation with changes in receptor-mediated catabolism. Arteriosclerosis, 7:135–143. 1987.
268.
StockerR, KeaneyKJJr. New insights on oxidative stress in the artery wall. J Throm Haem, 3:1825–1834. 2005.
269.
StockerR, KeaneyJFJr. Role of oxidative modifications in atherosclerosis. Physiol Rev, 84:1381–1478. 2004.
270.
SuJ, GeorgiadesA, WuR, ThulinT, de FaireU, FrostegårdJ. Antibodies of IgM subclass to phosphorylcholine and oxidized LDL are protective factors for atherosclerosis in patients with hypertension. Atherosclerosis, 188:160–166. 2006.
271.
SubbaiahPV, SubramanianVS, WangK. Novel physiological function of sphingomyelin in plasma. Inhibition of lipid peroxidation in low density lipoproteins. J Biol Chem, 274:36409–36414. 1999.
272.
SubbanagounderG, WatsonAD, BerlinerJA. Bioactive products of phospholipid oxidation: Isolation, identification, measurement and activities. Free Radical Biol Med, 28:1751–1761. 2000.
273.
SubbanagounderG, WongJW, LeeH, FaullKF, MillerE, WitztumJL, BerlinerJA. Epoxyisoprostane and epoxycyclopentenone phospholipids regulate monocyte chemotactic protein-1 and interleukin-8 synthesis. Formation of these oxidized phospholipids in response to interleukin-1 beta. J Biol Chem, 277:7271–7281. 2002.
SuzukiH, KuriharaY, TakeyaM, KamadaN, KataokaM, JishageK, UedaO, SakaguchiH, HigashiT, SuzukiT, TakashimaY, KawabeY, CynshiO, WadaY, HondaM, KuriharaH, AburataniH, DoiT, MatsumotoA, AzumaS, NodaT, ToyodaY, ItakuraH, YazakiY, KodamaT. A role for macrophage scavenger receptors in atherosclerosis and susceptibility to infection. Nature, 386:292–296. 1997.
276.
SzantoA, NagyL. The many faces of PPAR[gamma]: Anti-inflammatory by any means?Immunobiology, 213:789–803. 2008.
277.
TabuchiM, InoueK, Usui–KataokaH, KobayashiK, TeramotoM, TakasugiK, ShikataK, YamamuraM, AndoK, NishidaK, KasaharaJ, KumeN, LopezLR, MitsudoK, NobuyoshiM, YasudaT, KitaT, MakinoH, MatsuuraE. The association of C-reactive protein with an oxidative metabolite of LDL and its implication in atherosclerosis. J Lipid Res, 48:768–781. 2007.
278.
TerpstraV, BirdDA, SteinbergD. Evidence that the lipid moiety of oxidized low density lipoprotein plays a role in its interaction with macrophage receptors. Proc Natl Acad Sci USA, 95:1806–1811. 1998.
279.
TertovVV, SobeninIA, OrekhovAN. Similarity between naturally occurring modified desialylated, electronegative and aortic low density lipoprotein. 25:313–319. 1996.
280.
ThomsonMJ, PuntmannV, KaskiJC. Atherosclerosis and oxidant stress: The end of the road for antioxidant vitamin treatment?Cardiovasc Drugs Ther, 21:195–210. 2007.
281.
TietgeUJF, PraticoD, DingT, FunkCD, HildebrandRB, Van BerkelT, Van EckM. Macrophage-specific expression of group IIA sPLA2 results in accelerated atherogenesis by increasing oxidative stress. J Lipid Res, 46:1604–1614. 2005.
282.
TolozaSMA, UribeAG, McGwinGJ, AlarcónGS, FesslerBJ, BastianHM, ViláLM, WuR, ShoenfeldY, RosemanJM, ReveilleJD. Systemic lupus erythematosus in a multiethnic US cohort (LUMINA): XXIII. Baseline predictors of vascular events. Arthritis Rheum, 50:3947–3957. 2004.
283.
TrigattiB, RigottiA. Scavenger receptor class B type I (SR-BI) and high-density lipoprotein metabolism: Recent lessons from genetically manipulated mice. Int J Tissue React, 22:29–37. 2000.
284.
TsimikasS. Oxidative biomarkers in the diagnosis and prognosis of cardiovascular disease. Am J Cardiol, 98:9P–17P. 2006.
285.
UittenbogaardA, ShaulPW, YuhannaIS, BlairA, SmartEJ. High density lipoprotein prevents oxidized low density lipoprotein-induced inhibition of endothelial nitric-oxide synthase localization and activation in caveolae. J Biol Chem, 275:11278–11283. 2000.
286.
UpstonJM, NiuX, BrownAJ, MashimaR, WangH, SenthilmohanR, KettleAJ, DeanRT, StockerR. Disease stage-dependent accumulation of lipid and protein oxidation products in human atherosclerosis. Am J Pathol, 160:701–710. 2002.
van den EijndenMMED, van NoortJT, HollaarL, van der LaarseA, BertinaRM. Cholesterol or triglyceride loading of human monocyte-derived macrophages by incubation with modified lipoproteins does not induce tissue factor expression. Arterioscler Thromb Vasc Biol, 19:384–392. 1999.
289.
van der VeldeAE, GroenAK. Shifting gears: Liver SR-BI drives reverse cholesterol transport in macrophages. J Clin Invest, 115:2699–2701. 2005.
290.
van MeeterenLA, MoolenaarWH. Regulation and biological activities of the autotaxin-LPA axis. Prog Lipid Res, 46:145–160. 2007.
291.
VarbanML, RinningerF, WangN, Fairchild–HuntressV, DunmoreJH, FangQ, GosselinML, DixonKL, DeedsJD, ActonSL, TallAR, HuszarD. Targeted mutation reveals a central role for SR-BI in hepatic selective uptake of high density lipoprotein cholesterol. Proc Natl Acad Sci USA, 95:4619–4624. 1998.
292.
VerhoyeE, LangloisMR. Circulating oxidized low-density lipoprotein: A biomarker of atherosclerosis and cardiovascular risk?Clin Chem Lab Med, 47:128. 2009.
293.
VirellaG, Lopes–VirellaMF. Atherogenesis and the humoral immune response to modified lipoproteins. Atherosclerosis, 200:239–246. 2008.
294.
VirellaG, ThorpeSR, AldersonNL, StephanEM, AtchleyD, WagnerF, Lopes- -VirellaMF. Autoimmune response to advanced glycosylation end-products of human LDL. J Lipid Res, 44:487–493. 2003.
295.
WadaY, KurodaT, MurasawaA, TanabeN, NakanoM, GejyoF. Autoantibodies against oxidized low-density lipoprotein (LDL) and carotid atherosclerosis in patients with rheumatoid arthritis. Clin Exp Rheumatol, 23:482–486. 2005.
296.
WaltonKA, ColeAL, YehM, SubbanagounderG, KrutzikSR, ModlinRL, LucasRM, NakaiJ, SmartEJ, VoraDK, BerlinerJA. Specific phospholipid oxidation products inhibit ligand activation of toll-like receptors 4 and 2. Arterioscler Thromb Vasc Biol, 23:1197–1203. 2003.
297.
WatsonAD, SubbanagounderG, WelsbieDS, FaullKF, NavabM, JungME, FogelmanAM, BerlinerJA. Structural identification of a novel pro-inflammatory epoxyisoprostane phospholipid in mildly oxidized low density lipoprotein. J Biol Chem, 274:24787–24798. 1999.
298.
WebbNR, de VilliersWJ, ConnellPM, de BeerFC, van der WesthuyzenDR. Alternative forms of the scavenger receptor BI (SR-BI)J Lipid Res, 38:1490–1495. 1997.
299.
WenY, LeakeDS. Low density lipoprotein undergoes oxidation within lysosomes in cells. Circ Res, 100:1337–1343. 2007.
300.
WinyardPGTF, EsterbauerH, KusML, BlakeDR, MorrisCJ. Presence of foam cells containing oxidised low density lipoprotein in the synovial membrane from patients with rheumatoid arthritis. Ann Rheum Dis, 52:677–680. 1993.
301.
WittwerJ, HersbergerM. The two faces of the 15-lipoxygenase in atherosclerosis. Prostagland Leukotrienes Essent Fatty Acids, 77:67–77. 2007.
302.
WitztumJL, SteinbergD. The oxidative modification hypothesis of atherosclerosis: Does it hold for humans?Trends Cardiovasc Med, 11:93. 2001.
303.
WitztumJL, SteinbergD. Role of oxidized low density lipoprotein in atherogenesis. J Clin Invest, 88:1785–1792. 1991.
304.
WoenckhausC, KaufmannA, BufeldD, GemsaD, SprengerH, GröneHJ. Hypochlorite-modified LDL: Chemotactic potential and chemokine induction in human monocytes. Clin Immunol Immunopathol, 86:27–33. 1998.
305.
WongM–L, XieB, BeatiniN, PhuP, MaratheS, JohnsA, GoldPW, HirschE, WilliamsKJ, LicinioJ, TabasI. Acute systemic inflammation up-regulates secretory sphingomyelinase in vivo: A possible link between inflammatory cytokines and atherogenesis. Proc Natl Acad Sci USA, 97:8681–8686. 2000.
306.
WuBJ, KathirK, WittingPK, BeckK, ChoyK, LiC, CroftKD, MoriTA, TanousD, AdamsMR, LauAK, StockerR. Antioxidants protect from atherosclerosis by a heme oxygenase-1 pathway that is independent of free radical scavenging. J Exp Med, 203:1117–1127. 2006.
307.
YamamotoN, ToyodaM, AbeM, KobayashiT, KobayashiK, KatoM, MiyauchiM, KimuraM, UmezonoT, SuzukiD. Lectin-like oxidized LDL receptor-1 (LOX-1) expression in the tubulointerstitial area likely plays an important role in human diabetic nephropathy. Int Med, 48:189–194. 2009.
308.
YanceyPG, JeromeWG. Lysosomal sequestration of free and esterified cholesterol from oxidized low density lipoprotein in macrophages of different species. J Lipid Res, 39:1349–1361. 1998.
309.
YehM, ColeAL, ChoiJ, LiuY, TulchinskyD, QiaoJ–H, FishbeinMC, DooleyAN, HovnanianT, MouilleseauxK, VoraDK, YangW–P, GargalovicP, KirchgessnerT, ShyyJYJ, BerlinerJA. Role for sterol regulatory element-binding protein in activation of endothelial cells by phospholipid oxidation products. Circ Res, 95:780–788. 2004.
310.
Yla–HerttualaS. Is oxidized low-density lipoprotein present in vivo?Curr Opin Lipidol, 9:337–344. 1998.
311.
ZhangH, YangY, SteinbrecherUP. Structural requirements for the binding of modified proteins to the scavenger receptor of macrophages. J Biol Chem, 268:5535–5542. 1993.
312.
ZhangY, Da SilvaJR, ReillyM, BillheimerJT, RothblatGH, RaderDJ. Hepatic expression of scavenger receptor class B type I (SR-BI) is a positive regulator of macrophage reverse cholesterol transport in vivo. J Clin Invest, 115:2870–2874. 2005.
313.
ZhaoZ, de BeerMC, CaiL, AsmisR, de BeerFC, de VilliersWJS, van der WesthuyzenDR. Low-density lipoprotein from apolipoprotein E-deficient mice induces macrophage lipid accumulation in a CD36 and scavenger receptor class A-dependent manner. Arterioscler Thromb Vasc Biol, 25:168–173. 2005.
