Pilzer David, Fishelson Zvi
Department of Cell and Developmental Biology, Sackler School of Medicine, Tel Aviv University, Tel Aviv 69978, Israel.
Int Immunol. 2005 Sep;17(9):1239-48. doi: 10.1093/intimm/dxh300. Epub 2005 Aug 9.
The membrane attack complex (MAC) of the complement system is causing membrane damage and cell death. For protection, cells have adopted several resistance mechanisms, including removal of the membrane-inserted MAC by vesiculation. To identify proteins involved in MAC vesiculation, extracellular proteins released from K562 cells in response to treatment with sub-lytic complement were separated by acrylamide gel electrophoresis and protein bands were extracted, digested into peptides and the peptides were analyzed by mass spectrometry. A 75-kDa protein that was abundant in the supernatant of complement-treated cells was identified as mortalin/GRP75. Analysis by western blotting demonstrated that as early as 5 min after exposure to sub-lytic doses of complement, mortalin was released from K562 cells. Mortalin was released after complete activation of the complement system and formation of C5b-8, and even more so when C5b-9 was formed. Other pore formers, such as streptolysin O and melittin, did not induce release of mortalin. As shown, mortalin can bind to complement C8 and C9 and is shed in vesicles containing C9 and complement MACs. Anti-mortalin antibodies reduced mortalin release from complement-treated cells and elevated the extent of cell death by complement. Inhibitors of protein kinase C and extracellular signal-regulated protein kinase also prevented mortalin release from complement-activated cells. These results suggest that mortalin/GRP75 promotes the shedding of membrane vesicles loaded with complement MAC and protects cells from complement-mediated lysis.
补体系统的膜攻击复合物(MAC)会造成膜损伤和细胞死亡。为了自我保护,细胞采用了多种抵抗机制,包括通过形成囊泡来清除插入膜内的MAC。为了鉴定参与MAC囊泡化的蛋白质,将K562细胞在亚溶细胞剂量补体处理后释放的细胞外蛋白质通过丙烯酰胺凝胶电泳进行分离,提取蛋白条带,消化成肽段,然后通过质谱分析这些肽段。在补体处理细胞的上清液中大量存在的一种75 kDa蛋白质被鉴定为mortalin/GRP75。蛋白质印迹分析表明,早在暴露于亚溶细胞剂量的补体后5分钟,mortalin就从K562细胞中释放出来。在补体系统完全激活并形成C5b-8后,mortalin被释放,而当形成C5b-9时,释放得更多。其他孔形成剂,如链球菌溶血素O和蜂毒素,不会诱导mortalin的释放。如图所示,mortalin可以结合补体C8和C9,并在含有C9和补体MAC的囊泡中脱落。抗mortalin抗体减少了补体处理细胞中mortalin的释放,并增加了补体介导的细胞死亡程度。蛋白激酶C和细胞外信号调节蛋白激酶的抑制剂也阻止了mortalin从补体激活细胞中释放。这些结果表明,mortalin/GRP75促进了装载补体MAC的膜囊泡的脱落,并保护细胞免受补体介导的裂解。