ESAT-6 proteins: protective antigens and virulence factors?

Horwitz, 1995, Protective immunity against tuberculosis induced by vaccination with major extracellular proteins of Mycobacterium tuberculosis, Proc. Natl. Acad. Sci. U. S. A., 92, 1530, 10.1073/pnas.92.5.1530

Orme, 1993, T cell response to Mycobacterium tuberculosis, J. Infect. Dis., 167, 1481, 10.1093/infdis/167.6.1481

Andersen, 1991, Proteins released from Mycobacterium tuberculosis during growth, Infect. Immun., 59, 1905, 10.1128/IAI.59.6.1905-1910.1991

Andersen, 1995, Recall of long-lived immunity to Mycobacterium tuberculosis infection in mice, J. Immunol., 154, 3359, 10.4049/jimmunol.154.7.3359

Sørensen, 1995, Purification and characterization of a low-molecular-mass T-cell antigen secreted by Mycobacterium tuberculosis, Infect. Immun., 63, 1710, 10.1128/IAI.63.5.1710-1717.1995

Berthet, 1998, A Mycobacterium tuberculosis operon encoding ESAT-6 and a novel low- molecular-mass culture filtrate protein (CFP-10), Microbiology, 144, 3195, 10.1099/00221287-144-11-3195

Gordon, 1999, Identification of variable regions in the genomes of tubercle bacilli using bacterial artificial chromosome arrays, Mol. Microbiol., 32, 643, 10.1046/j.1365-2958.1999.01383.x

Behr, 1999, Comparative genomics of BCG vaccines by whole-genome DNA microarrays, Science, 284, 1520, 10.1126/science.284.5419.1520

Cole, 1998, Deciphering the biology of Mycobacterium tuberculosis from the complete genome sequence, Nature, 393, 537, 10.1038/31159

Tekaia, 1999, Analysis of the proteome of Mycobacterium tuberculosis in silico, Tuber. Lung Dis., 79, 329, 10.1054/tuld.1999.0220

Brown, 2000, The mycosins of Mycobacterium tuberculosis H37Rv: a family of subtilisin-like serine proteases, Gene, 254, 147, 10.1016/S0378-1119(00)00277-8

Renshaw, 2002, Conclusive evidence that the major T-cell antigens of the M. tuberculosis complex ESAT-6 and CFP-10 form a tight, 1:1 complex and characterisation of the structural properties of ESAT-6, CFP-10 and the ESAT-6-CFP-10 complex: implications for pathogenesis and virulence, J. Biol. Chem., 277, 21598, 10.1074/jbc.M201625200

Gey Van Pittius, N.C. et al. (2001) The ESAT-6 gene cluster of Mycobacterium tuberculosis and other high G+C Gram-positive bacteria. Genome Biol. 2, RESEARCH0044 (http://genomebiology.com/)

Marmiesse, 2004, Macro-array and bioinformatic analyses reveal mycobacterial ‘core’ genes, variation in the ESAT-6 gene family and new phylogenetic markers for the Mycobacterium tuberculosis complex, Microbiology, 150, 483, 10.1099/mic.0.26662-0

Sonnenberg, 1997, Definition of Mycobacterium tuberculosis culture filtrate proteins by two-dimensional polyacrylamide gel electrophoresis, N- terminal amino acid sequencing, and electrospray mass spectrometry, Infect. Immun., 65, 4515, 10.1128/IAI.65.11.4515-4524.1997

Rosenkrands, 2000, Mapping and identification of Mycobacterium tuberculosis proteins by two-dimensional gel electrophoresis, microsequencing and immunodetection, Electrophoresis, 21, 935, 10.1002/(SICI)1522-2683(20000301)21:5<935::AID-ELPS935>3.0.CO;2-P

Okkels, L.M. et al. CFP10 discriminates between non-acylated and acetylated ESAT-6 of Mycobacterium tuberculosis by differential interaction. Proteomics (in press)

Polevoda, B. and Sherman, F. (2002) The diversity of acetylated proteins. Genome Biol. 3, reviews0006 (http://genomebiology.com/)

Brandt, 1996, Key epitopes on the ESAT-6 antigen recognized in mice during the recall of protective immunity to Mycobacterium tuberculosis, J. Immunol., 157, 3527, 10.4049/jimmunol.157.8.3527

Pollock, 1997, Predominant recognition of the ESAT-6 protein in the first phase of infection with Mycobacterium bovis in cattle, Infect. Immun., 65, 2587, 10.1128/IAI.65.7.2587-2592.1997

Ravn, 1999, Human T cell responses to the ESAT-6 antigen from Mycobacterium tuberculosis, J. Infect. Dis., 179, 637, 10.1086/314640

Elhay, 1998, Delayed-type hypersensitivity responses to ESAT-6 and MPT64 from Mycobacterium tuberculosis in the guinea pig, Infect. Immun., 66, 3454, 10.1128/IAI.66.7.3454-3456.1998

Mori, 2004, Specific detection of tuberculosis infection with an interferon-gamma based assay using new antigens, Am. J. Respir. Crit. Care Med., 170, 59, 10.1164/rccm.200402-179OC

Lalvani, 2001, Rapid detection of Mycobacterium tuberculosis infection by enumeration of antigen-specific T cells, Am. J. Respir. Crit. Care Med., 163, 824, 10.1164/ajrccm.163.4.2009100

Arend, 2000, Detection of active tuberculosis infection by T cell responses to early-secreted antigenic target 6-kDa protein and culture filtrate protein 10, J. Infect. Dis., 181, 1850, 10.1086/315448

van Pinxteren, 2000, Diagnosis of tuberculosis based on the two specific antigens ESAT-6 and CFP10, Clin. Diagn. Lab. Immunol., 7, 155, 10.1128/CDLI.7.2.155-160.2000

Mustafa, 1998, Comparison of antigen-specific T-cell responses of tuberculosis patients using complex or single antigens of Mycobacterium tuberculosis, Scand. J. Immunol., 48, 535, 10.1046/j.1365-3083.1998.00419.x

Lyashchenko, 1998, Heterogeneous antibody responses in tuberculosis, Infect. Immun., 66, 3936, 10.1128/IAI.66.8.3936-3940.1998

Brusasca, 2003, Antigen recognition by serum antibodies in non-human primates experimentally infected with Mycobacterium tuberculosis, Comp. Med., 53, 165

Lyashchenko, 1998, Diversity of antigen recognition by serum antibodies in experimental bovine tuberculosis, Infect. Immun., 66, 5344, 10.1128/IAI.66.11.5344-5349.1998

Mustafa, 2000, Multiple epitopes from the Mycobacterium tuberculosis ESAT-6 antigen are recognized by antigen-specific human T cell lines, Clin. Infect. Dis., 30, S201, 10.1086/313862

Mustafa, 2003, Human Th1 cell lines recognize the Mycobacterium tuberculosis ESAT-6 antigen and its peptides in association with frequently expressed HLA class II molecules, Scand. J. Immunol., 57, 125, 10.1046/j.1365-3083.2003.01204.x

Dillon, 2000, Molecular and immunological characterization of Mycobacterium tuberculosis CFP-10, an immunodiagnostic antigen missing in Mycobacterium bovis BCG, J. Clin. Microbiol., 38, 3285, 10.1128/JCM.38.9.3285-3290.2000

Skjøt, 2000, Comparative evaluation of low-molecular-mass proteins from Mycobacterium tuberculosis identifies members of the ESAT-6 family as immunodominant T-cell antigens, Infect. Immun., 68, 214, 10.1128/IAI.68.1.214-220.2000

Colangeli, 2000, MTSA-10, the product of the Rv3874 gene of Mycobacterium tuberculosis, elicits tuberculosis-specific, delayed-type hypersensitivity in guinea pigs, Infect. Immun., 68, 990, 10.1128/IAI.68.2.990-993.2000

Okkels, 2003, PPE protein (Rv3873) from DNA segment RD1 of Mycobacterium tuberculosis: Strong recognition of both specific T-cell epitopes and epitopes conserved within the PPE family, Infect. Immun., 71, 6116, 10.1128/IAI.71.11.6116-6123.2003

Agger, 2003, Human T-cell responses to the RD1-encoded protein TB27.4 (Rv3878) from Mycobacterium tuberculosis, Immunology, 110, 507, 10.1111/j.1365-2567.2003.01763.x

Brusasca, 2001, Immunological characterization of antigens encoded by the RD1 region of the Mycobacterium tuberculosis genome, Scand. J. Immunol., 54, 448, 10.1046/j.1365-3083.2001.00975.x

Demangel, 2004, Cell envelope protein PPE68 contributes to Mycobacterium tuberculosis RD1 immunogenicity independently of a 10-kilodalton culture filtrate protein and ESAT-6, Infect. Immun., 72, 2170, 10.1128/IAI.72.4.2170-2176.2004

Liu, 2004, Evaluation of T-cell responses to novel RD1- and RD2-encoded Mycobacterium tuberculosis gene products for specific detection of human tuberculosis infection, Infect. Immun., 72, 2574, 10.1128/IAI.72.5.2574-2581.2004

Cockle, 2002, Identification of novel Mycobacterium tuberculosis antigens with potential as diagnostic reagents or subunit vaccine candidates by comparative genomics, Infect. Immun., 70, 6996, 10.1128/IAI.70.12.6996-7003.2002

Dillon, 1999, Molecular characterization and human T-cell responses to a member of a novel Mycobacterium tuberculosis mtb39 gene family, Infect. Immun., 67, 2941, 10.1128/IAI.67.6.2941-2950.1999

Alderson, 2000, Expression cloning of an immunodominant family of Mycobacterium tuberculosis antigens using human CD4(+) T cells, J. Exp. Med., 191, 551, 10.1084/jem.191.3.551

Skjøt, 2002, Epitope mapping of the immunodominant antigen TB10.4 and the two homologous proteins TB10.3 and TB12.9, which constitute a subfamily of the esat-6 gene family, Infect. Immun., 70, 5446, 10.1128/IAI.70.10.5446-5453.2002

Pathan, 2000, High frequencies of circulating IFN-gamma-secreting CD8 cytotoxic T cells specific for a novel MHC class I-restricted Mycobacterium tuberculosis epitope in M. tuberculosis-infected subjects without disease, Eur. J. Immunol., 30, 2713, 10.1002/1521-4141(200009)30:9<2713::AID-IMMU2713>3.0.CO;2-4

Smith, 2000, Human CD8(+) T cells specific for Mycobacterium tuberculosis secreted antigens in tuberculosis patients and healthy BCG-vaccinated controls in The Gambia, Infect. Immun., 68, 7144, 10.1128/IAI.68.12.7144-7148.2000

Majlessi, 2003, CD8+-T-cell responses of Mycobacterium-infected mice to a newly identified major histocompatibility complex class I-restricted epitope shared by proteins of the ESAT-6 family, Infect. Immun., 71, 7173, 10.1128/IAI.71.12.7173-7177.2003

Hsu, 2003, The primary mechanism of attenuation of bacillus Calmette-Guerin is a loss of secreted lytic function required for invasion of lung interstitial tissue, Proc. Natl. Acad. Sci. U. S. A., 100, 12420, 10.1073/pnas.1635213100

Weber, 2002, Antigenic drift, antigenic shift and interferon antagonists: how bunyaviruses counteract the immune system, Virus Res., 88, 129, 10.1016/S0168-1702(02)00125-9

Bai, 2003, Antigenic drift as a mechanism for tumor evasion of destruction by cytolytic T lymphocytes, J. Clin. Invest., 111, 1487, 10.1172/JCI17656

Haile, 2002, Expression of the mceA, esat-6 and hspX genes in Mycobacterium tuberculosis and their responses to aerobic conditions and to restricted oxygen supply, Microbiology, 148, 3881, 10.1099/00221287-148-12-3881

Shi, 2004, Effect of growth state on transcription levels of genes encoding major secreted antigens of Mycobacterium tuberculosis in the mouse lung, Infect. Immun., 72, 2420, 10.1128/IAI.72.4.2420-2424.2004

Mahairas, 1996, Molecular analysis of genetic differences between Mycobacterium bovis BCG and virulent M. bovis, J. Bacteriol., 178, 1274, 10.1128/jb.178.5.1274-1282.1996

Kamath, 1999, Differential protective efficacy of DNA vaccines expressing secreted proteins of Mycobacterium tuberculosis, Infect. Immun., 67, 1702, 10.1128/IAI.67.4.1702-1707.1999

Li, 1999, Immunogenicity of DNA vaccines expressing tuberculosis proteins fused to tissue plasminogen activator signal sequences, Infect. Immun., 67, 4780, 10.1128/IAI.67.9.4780-4786.1999

Huygen, 1996, Immunogenicity and protective efficacy of a tuberculosis DNA vaccine, Nat. Med., 2, 893, 10.1038/nm0896-893

Brandt, 2000, ESAT-6 subunit vaccination against Mycobacterium tuberculosis, Infect. Immun., 68, 791, 10.1128/IAI.68.2.791-795.2000

Olsen, 2000, Efficient protection against Mycobacterium tuberculosis by vaccination with a single subdominant epitope from the ESAT-6 antigen, Eur. J. Immunol., 30, 1724, 10.1002/1521-4141(200006)30:6<1724::AID-IMMU1724>3.0.CO;2-A

Brodin, 2004, Enhanced protection against extrapulmonary tuberculosis with an attenuated live Mycobacterium microti vaccine inducing T-cell immunity against RD1 antigens, J. Infect. Dis., 190, 115, 10.1086/421468

Pym, 2003, Recombinant BCG exporting ESAT-6 confers enhanced protection against tuberculosis, Nat. Med., 9, 533, 10.1038/nm859

Olsen, 2001, Protection of mice with a tuberculosis subunit vaccine based on a fusion protein of antigen 85B and ESAT-6, Infect. Immun., 69, 2773, 10.1128/IAI.69.5.2773-2778.2001

Olsen, A.W. et al. Protective effect of a tuberculosis subunit vaccine based on a fusion of Ag85B and ESAT-6 in the aerosol guinea pig model. Infect. Immun. (in press)

Andersen, 2000, Specific immune-based diagnosis of tuberculosis, Lancet, 356, 1099, 10.1016/S0140-6736(00)02742-2

Doherty, 2002, Immune responses to the Mycobacterium tuberculosis-specific antigen ESAT-6 signal subclinical infection among contacts of tuberculosis patients, J. Clin. Microbiol., 40, 704, 10.1128/JCM.40.2.704-706.2002

Lalvani, 2001, Enhanced contact tracing and spatial tracking of Mycobacterium tuberculosis infection by enumeration of antigen-specific T cells, Lancet, 357, 2017, 10.1016/S0140-6736(00)05115-1

Vekemans, 2001, Tuberculosis contacts but not patients have higher gamma interferon responses to ESAT-6 than do community controls in The Gambia, Infect. Immun., 69, 6554, 10.1128/IAI.69.10.6554-6557.2001

Pollock, 2003, Specific delayed-type hypersensitivity responses to ESAT-6 identify tuberculosis-infected cattle, J. Clin. Microbiol., 41, 1856, 10.1128/JCM.41.5.1856-1860.2003

Hart, 1977, BCG and vole bacillus vaccines in the prevention of tuberculosis in adolescence and early adult life, BMJ, 2, 293, 10.1136/bmj.2.6082.293

Brodin, 2002, Bacterial artificial chromosome-based comparative genomic analysis identifies Mycobacterium microti as a natural ESAT-6 deletion mutant, Infect. Immun., 70, 5568, 10.1128/IAI.70.10.5568-5578.2002

Pym, 2002, Loss of RD1 contributed to the attenuation of the live tuberculosis vaccines Mycobacterium bovis BCG and Mycobacterium microti, Mol. Microbiol., 46, 709, 10.1046/j.1365-2958.2002.03237.x

Lewis, 2003, Deletion of RD1 from Mycobacterium tuberculosis mimics bacille Calmette-Guerin attenuation, J. Infect. Dis., 187, 117, 10.1086/345862

Stanley, 2003, Acute infection and macrophage subversion by Mycobacterium tuberculosis require a specialized secretion system, Proc. Natl. Acad. Sci. U. S. A., 100, 13001, 10.1073/pnas.2235593100

Wards, 2000, An esat6 knockout mutant of Mycobacterium bovis produced by homologous recombination will contribute to the development of a live tuberculosis vaccine, Tuber. Lung Dis., 80, 185, 10.1054/tuld.2000.0244

Guinn, 2004, Individual RD1-region genes are required for export of ESAT-6/CFP-10 and for virulence of Mycobacterium tuberculosis, Mol. Microbiol., 51, 359, 10.1046/j.1365-2958.2003.03844.x

Bao, 2003, Virulence, immunogenicity, and protective efficacy of two recombinant Mycobacterium bovis bacillus Calmette-Guerin strains expressing the antigen ESAT-6 from Mycobacterium tuberculosis, Infect. Immun., 71, 1656, 10.1128/IAI.71.4.1656-1661.2003

Okkels, 2004, Protein–protein interactions of proteins from the ESAT-6 family of Mycobacterium tuberculosis, J. Bacteriol., 186, 2487, 10.1128/JB.186.8.2487-2491.2004