Strauss Juliane G, Zimmermann Robert, Hrzenjak Andelko, Zhou Yonggang, Kratky Dagmar, Levak-Frank Sanja, Kostner Gert M, Zechner Rudolf, Frank Sasa
Institute of Molecular Biology, Biochemistry and Microbiology, University of Graz, Harrachgasse 21, A-8010 Graz, Austria.
Biochem J. 2002 Nov 15;368(Pt 1):69-79. doi: 10.1042/BJ20020306.
Endothelial cell-derived lipase (EDL) is a new member of the lipase gene family with high sequence homology with lipoprotein lipase (LPL). EDL is a phospholipase with very little triacylglycerol lipase activity. To investigate the effects of EDL on binding and uptake of high-density lipoprotein (HDL), as well as on the selective uptake of HDL-derived cholesterol esters (CEs), HepG2 cells were infected with adenovirus coding for EDL. For comparison, cells were also infected with LPL and with lacZ as a control. Both HDL binding and particle uptake were increased 1.5-fold and selective HDL-CE uptake was increased 1.8-fold in EDL-infected HepG2 cells compared with controls. The effect of LPL was less pronounced, resulting in 1.1-fold increase in particle uptake and 1.3-fold increase in selective uptake. Inhibition of the enzymic activity with tetrahydrolipstatin (THL) significantly enhanced the effect of EDL, as reflected by a 5.2-fold increase in binding, a 2.6-fold increase in particle uptake and a 1.1-fold increase in CE selective uptake compared with incubations without THL. To elucidate the mechanism responsible for the effects of THL, we analysed the abundance of heparin-releasable EDL protein from infected HepG2 cells upon incubations with THL, HDL and free (non-esterified) fatty acids (FFAs). In the presence of THL, vastly more EDL protein remained bound to the cell surface. Additionally, HDL and FFAs reduced the amount of cell-surface-bound EDL, suggesting that fatty acids that are liberated from phospholipids in HDL release EDL from the cell surface. This was substantiated further by the finding that, in contrast with EDL, the amount of cell-surface-bound enzymically inactive mutant EDL (MUT-EDL) was not reduced in the presence of HDL and foetal calf serum. The increased amount of cell-surface-bound MUT-EDL in the presence of THL suggested that the enzymic inactivity of MUT-EDL, as well as an augmenting effect of THL that is independent of its ability to inactivate the enzyme, are responsible for the increased amount of cell-surface-bound EDL in the presence of THL. Furthermore, in cells expressing MUT-EDL, binding and holoparticle uptake were markedly higher compared with cells expressing the active EDL, and could be increased further in the presence of THL. Despite 1.7-fold higher binding and 1.8-fold higher holoparticle uptake, the selective CE uptake by MUT-EDL-expressing cells was comparable with EDL-expressing cells and was even decreased 1.3-fold with THL. Experiments in CLA-1 (CD-36 and LIMPII analogous 1, the human homologue of scavenger receptor class B type I)-deficient HEK-293 cells demonstrated that EDL alone has the ability to stimulate HDL-CE selective uptake independently of CLA-1. Thus our results demonstrate that EDL mediates both HDL binding and uptake, and the selective uptake of HDL-CE, independently of lipolysis and CLA-1.
内皮细胞源性脂肪酶(EDL)是脂肪酶基因家族的新成员,与脂蛋白脂肪酶(LPL)具有高度的序列同源性。EDL是一种磷脂酶,三酰甘油脂肪酶活性极低。为了研究EDL对高密度脂蛋白(HDL)结合和摄取以及HDL衍生胆固醇酯(CEs)选择性摄取的影响,用编码EDL的腺病毒感染HepG2细胞。作为对照,细胞也用LPL和lacZ感染。与对照相比,在EDL感染的HepG2细胞中,HDL结合和颗粒摄取增加了1.5倍,HDL-CE选择性摄取增加了1.8倍。LPL的作用不太明显,导致颗粒摄取增加1.1倍,选择性摄取增加1.3倍。用四氢脂抑素(THL)抑制酶活性显著增强了EDL的作用,与未添加THL的孵育相比,结合增加了5.2倍,颗粒摄取增加了2.6倍,CE选择性摄取增加了1.1倍。为了阐明THL作用的机制,我们分析了在用THL、HDL和游离(非酯化)脂肪酸(FFA)孵育后,感染的HepG2细胞中肝素可释放的EDL蛋白的丰度。在存在THL的情况下,大量更多的EDL蛋白仍与细胞表面结合。此外,HDL和FFA减少了细胞表面结合的EDL量,表明从HDL中的磷脂释放的脂肪酸将EDL从细胞表面释放出来。这一发现进一步得到证实,即与EDL相反,在存在HDL和胎牛血清的情况下,细胞表面结合的无酶活性突变型EDL(MUT-EDL)的量没有减少。在存在THL的情况下,细胞表面结合的MUT-EDL量增加表明,MUT-EDL的酶无活性以及THL的增强作用(与其使酶失活的能力无关)是存在THL时细胞表面结合的EDL量增加的原因。此外,在表达MUT-EDL的细胞中,与表达活性EDL的细胞相比,结合和全颗粒摄取明显更高,并且在存在THL的情况下可以进一步增加。尽管结合增加了1.7倍,全颗粒摄取增加了1.8倍,但表达MUT-EDL的细胞的CE选择性摄取与表达EDL的细胞相当,并且在添加THL时甚至降低了1.3倍。在CLA-1(CD-36和LIMPII类似物1,清道夫受体B类I型的人类同源物)缺陷的HEK-293细胞中的实验表明,单独的EDL具有独立于CLA-1刺激HDL-CE选择性摄取的能力。因此,我们的结果表明,EDL介导HDL的结合和摄取以及HDL-CE的选择性摄取,与脂解作用和CLA-1无关。