Cain K, Brown D G, Langlais C, Cohen G M
Medical Research Council Toxicology Unit, Hodgkin Building, University of Leicester, P. O. Box 138, Lancaster Road, Leicester LE1 9HN, United Kingdom.
J Biol Chem. 1999 Aug 6;274(32):22686-92. doi: 10.1074/jbc.274.32.22686.
In mammals, apoptotic protease-activating factor 1 (Apaf-1), cytochrome c, and dATP activate caspase-9, which initiates the postmitochondrial-mediated caspase cascade by proteolytic cleavage/activation of effector caspases to form active approximately 60-kDa heterotetramers. We now demonstrate that activation of caspases either in apoptotic cells or following dATP activation of cell lysates results in the formation of two large but different sized protein complexes, the "aposome" and the "microaposome". Surprisingly, most of the DEVDase activity in the lysate was present in the aposome and microaposome complexes with only small amounts of active caspase-3 present as its free approximately 60-kDa heterotetramer. The larger aposome complex (M(r) = approximately 700,000) contained Apaf-1 and processed caspase-9, -3, and -7. The smaller microaposome complex (M(r) = approximately 200,000-300,000) contained active caspase-3 and -7 but little if any Apaf-1 or active caspase-9. Lysates isolated from control THP.1 cells, prior to caspase activation, showed striking differences in the distribution of key apoptotic proteins. Apaf-1 and procaspase-7 may be functionally complexed as they eluted as an approximately 200-300-kDa complex, which did not have caspase cleavage (DEVDase) activity. Procaspase-3 and -9 were present as separate and smaller 60-90-kDa (dimer) complexes. During caspase activation, Apaf-1, caspase-9, and the effector caspases redistributed and formed the aposome. This resulted in the processing of the effector caspases, which were then released, possibly bound to other proteins, to form the microaposome complex.
在哺乳动物中,凋亡蛋白酶激活因子1(Apaf-1)、细胞色素c和dATP可激活半胱天冬酶-9,后者通过对效应半胱天冬酶进行蛋白水解切割/激活,启动线粒体后介导的半胱天冬酶级联反应,形成活性约为60 kDa的异源四聚体。我们现在证明,在凋亡细胞中或细胞裂解物经dATP激活后,半胱天冬酶的激活会导致形成两种大小不同的大蛋白复合物,即“凋亡小体”和“微凋亡小体”。令人惊讶的是,裂解物中的大多数DEVDase活性存在于凋亡小体和微凋亡小体复合物中,只有少量活性半胱天冬酶-3以其游离的约60 kDa异源四聚体形式存在。较大的凋亡小体复合物(相对分子质量约为700,000)包含Apaf-1以及经过加工的半胱天冬酶-9、-3和-7。较小的微凋亡小体复合物(相对分子质量约为200,000 - 300,000)包含活性半胱天冬酶-3和-7,但几乎不含Apaf-1或活性半胱天冬酶-9。在半胱天冬酶激活之前,从对照THP.1细胞中分离的裂解物在关键凋亡蛋白的分布上显示出显著差异。Apaf-1和半胱天冬酶原-7可能在功能上形成复合物,因为它们以约200 - 300 kDa的复合物形式洗脱,该复合物没有半胱天冬酶切割(DEVDase)活性。半胱天冬酶原-3和-9以单独的较小的60 - 90 kDa(二聚体)复合物形式存在。在半胱天冬酶激活过程中,Apaf-1、半胱天冬酶-9和效应半胱天冬酶重新分布并形成凋亡小体。这导致效应半胱天冬酶的加工,然后它们被释放,可能与其他蛋白质结合,形成微凋亡小体复合物。