Fueller Florian, Schmidt Gudula
Institute for Experimental and Clinical Pharmacology and Toxicology, University of Freiburg, 79104 Freiburg, Germany.
Protein Sci. 2008 Aug;17(8):1456-62. doi: 10.1110/ps.035386.108. Epub 2008 Jun 26.
Pathogenic Yersinia strains evade the innate immune responses of the host by producing effector proteins ( Yersinia outer proteins [Yops]), which are directly injected into mammalian cells by a type III secretion system (TTSS). One of these effector proteins (YopT) disrupts the actin cytoskeleton of the host cell resulting in cell rounding. YopT is a cysteine protease that cleaves Rho proteins directly upstream of the post-translationally modified cysteine. Thereby, it releases the GTPases from the membrane leading to inactivation. Small GTPases are modified by isoprenylation of the cysteine of the CAAX box, cleavage of the -AAX tripeptide, and methylation of the cysteine. We have shown that isoprenylation and the endoproteolytic cleavage of the tripeptide of Rho GTPases are essential for YopT-induced cleavage, whereas carboxyl methylation is not required. In the present study, we post-translationally modified RhoA, Rac, Cdc42, and several mutants in vitro and characterized the YopT-induced cleavage with recombinant YopT. We show that farnesylated RhoA is a preferred substrate of YopT compared with the geranylgeranylated GTPase. Geranylgeranylated RhoA, however, is the preferred substrate for YopT-catalyzed cleavage with a threefold faster turnover rate over Rac and Cdc42. Moreover, our data indicate that the composition of the polybasic region of the GTPases defines the specificity and efficiency of the YopT-induced cleavage, and that a space between the polybasic stretch of amino acids at the C terminus and the CAAX box enhances the turnover rate of YopT-catalyzed cleavage.
致病性耶尔森氏菌菌株通过产生效应蛋白(耶尔森氏菌外蛋白[Yops])来逃避宿主的先天免疫反应,这些效应蛋白由III型分泌系统(TTSS)直接注入哺乳动物细胞。其中一种效应蛋白(YopT)会破坏宿主细胞的肌动蛋白细胞骨架,导致细胞变圆。YopT是一种半胱氨酸蛋白酶,可在翻译后修饰的半胱氨酸的直接上游切割Rho蛋白。从而,它使GTP酶从膜上释放出来,导致其失活。小GTP酶通过CAAX盒半胱氨酸的异戊二烯化、-AAX三肽的切割以及半胱氨酸的甲基化进行修饰。我们已经表明,Rho GTP酶的异戊二烯化和三肽的内切蛋白水解切割对于YopT诱导的切割至关重要,而羧基甲基化则不是必需的。在本研究中,我们在体外对RhoA、Rac、Cdc42和几个突变体进行了翻译后修饰,并用重组YopT对YopT诱导的切割进行了表征。我们表明,与香叶基香叶基化的GTP酶相比,法尼基化的RhoA是YopT的首选底物。然而,香叶基香叶基化的RhoA是YopT催化切割的首选底物,其周转速度比Rac和Cdc42快三倍。此外,我们的数据表明,GTP酶多碱性区域的组成定义了YopT诱导切割的特异性和效率,并且C末端氨基酸多碱性延伸与CAAX盒之间的空间提高了YopT催化切割的周转速度。