Gcebe Nomakorinte, Michel Anita, Gey van Pittius Nicolaas C, Rutten Victor
Tuberculosis Laboratory, Agricultural Research Council - Onderstepoort Veterinary InstituteOnderstepoort, South Africa; Department of Veterinary Tropical Diseases, Faculty of Veterinary Science, University of PretoriaOnderstepoort, South Africa.
Department of Veterinary Tropical Diseases, Faculty of Veterinary Science, University of Pretoria Onderstepoort, South Africa.
Front Microbiol. 2016 Jun 7;7:795. doi: 10.3389/fmicb.2016.00795. eCollection 2016.
The Esx and PE/PPE families of proteins are among the most immunodominant mycobacterial antigens and have thus been the focus of research to develop vaccines and immunological tests for diagnosis of bovine and human tuberculosis, mainly caused by Mycobacterium bovis and Mycobacterium tuberculosis, respectively. In non-tuberculous mycobacteria (NTM), multiple copies of genes encoding homologous proteins have mainly been identified in pathogenic Mycobacterium species phylogenically related to Mycobacterium tuberculosis and Mycobacterium bovis. Only ancestral copies of these genes have been identified in nonpathogenic NTM species like Mycobacterium smegmatis, Mycobacterium sp. KMS, Mycobacterium sp. MCS, and Mycobacterium sp. JLS. In this study we elucidated the genomes of four nonpathogenic NTM species, viz Mycobacterium komanii sp. nov., Mycobacterium malmesburii sp. nov., Mycobacterium nonchromogenicum, and Mycobacterium fortuitum ATCC 6841. These genomes were investigated for genes encoding for the Esx and PE/PPE (situated in the esx cluster) family of proteins as well as adjacent genes situated in the ESX-1 to ESX-5 regions. To identify proteins actually expressed, comparative proteomic analyses of purified protein derivatives from three of the NTM as well as Mycobacterium kansasii ATCC 12478 and the commercially available purified protein derivatives from Mycobacterium bovis and Mycobacterium avium was performed. The genomic analysis revealed the occurrence in each of the four NTM, orthologs of the genes encoding for the Esx family, the PE and PPE family proteins in M. bovis and M. tuberculosis. The identification of genes of the ESX-1, ESX-3, and ESX-4 region including esxA, esxB, ppe68, pe5, and pe35 adds to earlier reports of these genes in nonpathogenic NTM like M. smegmatis, Mycobacterium sp. JLS and Mycobacterium KMS. This report is also the first to identify esxN gene situated within the ESX-5 locus in M. nonchromogenicum. Our proteomics analysis identified a total of 609 proteins in the six PPDs and 22 of these were identified as shared between PPD of M.bovis and one or more of the NTM PPDs. Previously characterized M tuberculosis/M. bovis homologous immunogenic proteins detected in one or more of the nonpathogenic NTM in this study included CFP-10 (detected in M. malmesburii sp. nov. PPD), GroES (detected in all NTM PPDs but M. malmesburii sp. nov.), DnaK (detected in all NTM PPDs), and GroEL (detected in all NTM PPDs). This study confirms reports that the ESX-1, ESX-3, and ESX-4 regions are ancestral regions and thus found in the genomes of most mycobacteria. Identification of NTM homologs of immunogenic proteins warrants further investigation of their ability to cause cross-reactive immune responses with MTBC antigens.
Esx和PE/PPE蛋白家族是分枝杆菌中免疫原性最强的抗原之一,因此一直是开发用于诊断牛结核病和人类结核病疫苗及免疫检测的研究重点,牛结核病和人类结核病分别主要由牛分枝杆菌和结核分枝杆菌引起。在非结核分枝杆菌(NTM)中,编码同源蛋白的基因多拷贝主要在与结核分枝杆菌和牛分枝杆菌系统发育相关的致病性分枝杆菌物种中被鉴定出来。在耻垢分枝杆菌、KMS分枝杆菌、MCS分枝杆菌和JLS分枝杆菌等非致病性NTM物种中,仅鉴定出这些基因的原始拷贝。在本研究中,我们阐明了四种非致病性NTM物种的基因组,即新种科曼尼分枝杆菌、新种马尔姆斯伯里分枝杆菌、非产色分枝杆菌和偶然分枝杆菌ATCC 6841。对这些基因组进行了编码Esx和PE/PPE(位于esx簇中)蛋白家族的基因以及位于ESX-1至ESX-5区域的相邻基因的研究。为了鉴定实际表达的蛋白质,对三种NTM以及堪萨斯分枝杆菌ATCC 12478的纯化蛋白衍生物和市售的牛分枝杆菌及鸟分枝杆菌纯化蛋白衍生物进行了比较蛋白质组学分析。基因组分析揭示了在这四种NTM的每一种中,存在与牛分枝杆菌和结核分枝杆菌中编码Esx家族、PE和PPE家族蛋白的基因直系同源的基因。ESX-1、ESX-3和ESX-4区域基因的鉴定,包括esxA、esxB、ppe68、pe5和pe35,补充了早期关于这些基因在耻垢分枝杆菌、JLS分枝杆菌和KMS分枝杆菌等非致病性NTM中的报道。本报告也是首次在非产色分枝杆菌的ESX-5位点内鉴定出esxN基因。我们的蛋白质组学分析在六种纯化蛋白衍生物中总共鉴定出609种蛋白质,其中22种被鉴定为在牛分枝杆菌的纯化蛋白衍生物与一种或多种NTM的纯化蛋白衍生物之间共享。在本研究中,在一种或多种非致病性NTM中检测到先前已鉴定的结核分枝杆菌/牛分枝杆菌同源免疫原性蛋白,包括CFP-10(在新种马尔姆斯伯里分枝杆菌的纯化蛋白衍生物中检测到)、GroES(在除新种马尔姆斯伯里分枝杆菌外的所有NTM的纯化蛋白衍生物中检测到)、DnaK(在所有NTM的纯化蛋白衍生物中检测到)和GroEL(在所有NTM的纯化蛋白衍生物中检测到)。本研究证实了关于ESX-1、ESX-3和ESX-4区域是祖先区域,因此存在于大多数分枝杆菌基因组中的报道。免疫原性蛋白的NTM同源物的鉴定值得进一步研究它们引起与结核分枝杆菌复合群抗原交叉反应性免疫应答的能力。