Li Yanhua, Treffers Emmely E, Napthine Sawsan, Tas Ali, Zhu Longchao, Sun Zhi, Bell Susanne, Mark Brian L, van Veelen Peter A, van Hemert Martijn J, Firth Andrew E, Brierley Ian, Snijder Eric J, Fang Ying
Department of Veterinary and Biomedical Sciences and Department of Biology/Microbiology, South Dakota State University, Brookings, SD 57007;Department of Diagnostic Medicine/Pathobiology, College of Veterinary Medicine, Kansas State University, Manhattan, KS 66506;
Molecular Virology Laboratory, Department of Medical Microbiology, Leiden University Medical Center, 2333 ZA, Leiden, The Netherlands;Department of Immunohematology and Blood Transfusion, Leiden University Medical Center, 2333 ZA, Leiden, The Netherlands;
Proc Natl Acad Sci U S A. 2014 May 27;111(21):E2172-81. doi: 10.1073/pnas.1321930111. Epub 2014 May 13.
Programmed -1 ribosomal frameshifting (-1 PRF) is a widely used translational mechanism facilitating the expression of two polypeptides from a single mRNA. Commonly, the ribosome interacts with an mRNA secondary structure that promotes -1 frameshifting on a homopolymeric slippery sequence. Recently, we described an unusual -2 frameshifting (-2 PRF) signal directing efficient expression of a transframe protein [nonstructural protein 2TF (nsp2TF)] of porcine reproductive and respiratory syndrome virus (PRRSV) from an alternative reading frame overlapping the viral replicase gene. Unusually, this arterivirus PRF signal lacks an obvious stimulatory RNA secondary structure, but as confirmed here, can also direct the occurrence of -1 PRF, yielding a third, truncated nsp2 variant named "nsp2N." Remarkably, we now show that both -2 and -1 PRF are transactivated by a protein factor, specifically a PRRSV replicase subunit (nsp1β). Embedded in nsp1β's papain-like autoproteinase domain, we identified a highly conserved, putative RNA-binding motif that is critical for PRF transactivation. The minimal RNA sequence required for PRF was mapped within a 34-nt region that includes the slippery sequence and a downstream conserved CCCANCUCC motif. Interaction of nsp1β with the PRF signal was demonstrated in pull-down assays. These studies demonstrate for the first time, to our knowledge, that a protein can function as a transactivator of ribosomal frameshifting. The newly identified frameshifting determinants provide potential antiviral targets for arterivirus disease control and prevention. Moreover, protein-induced transactivation of frameshifting may be a widely used mechanism, potentially including previously undiscovered viral strategies to regulate viral gene expression and/or modulate host cell translation upon infection.
程序性-1核糖体移码(-1 PRF)是一种广泛使用的翻译机制,可促进从单个mRNA表达两种多肽。通常,核糖体与mRNA二级结构相互作用,该二级结构促进在同聚物滑序列上的-1移码。最近,我们描述了一种不寻常的-2移码(-2 PRF)信号,该信号指导猪繁殖与呼吸综合征病毒(PRRSV)的转框蛋白[非结构蛋白2TF(nsp2TF)]从与病毒复制酶基因重叠的另一个阅读框高效表达。异常的是,这种动脉炎病毒PRF信号缺乏明显的刺激性RNA二级结构,但正如在此证实的那样,它也可以指导-1 PRF的发生,产生第三种截短的nsp2变体,称为“nsp2N”。值得注意的是,我们现在表明,-2和-1 PRF均被一种蛋白质因子反式激活,特别是PRRSV复制酶亚基(nsp1β)。嵌入nsp1β的木瓜蛋白酶样自蛋白酶结构域中,我们鉴定出一个高度保守的推定RNA结合基序,该基序对于PRF反式激活至关重要。PRF所需的最小RNA序列定位在一个34个核苷酸的区域内,该区域包括滑序列和下游保守的CCCANCUCC基序。在下拉试验中证明了nsp1β与PRF信号的相互作用。据我们所知,这些研究首次证明一种蛋白质可以作为核糖体移码的反式激活因子。新发现的移码决定因素为动脉炎病毒疾病的控制和预防提供了潜在的抗病毒靶点。此外,蛋白质诱导的移码反式激活可能是一种广泛使用的机制,可能包括以前未发现的病毒策略,以在感染时调节病毒基因表达和/或调节宿主细胞翻译。