{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T02:54:14Z","timestamp":1792032854962,"version":"4.4.0"},"reference-count":55,"publisher":"American Society for Microbiology","issue":"24","license":[{"start":{"date-parts":[[2009,12,15]],"date-time":"2009-12-15T00:00:00Z","timestamp":1260835200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/journals.asm.org\/non-commercial-tdm-license"}],"content-domain":{"domain":["journals.asm.org"],"crossmark-restriction":true},"short-container-title":["J Bacteriol"],"published-print":{"date-parts":[[2009,12,15]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n          <jats:p>\n            The\n            <jats:italic>Bacillus anthracis<\/jats:italic>\n            spore is the causative agent of the disease anthrax. The outermost structure of the\n            <jats:italic>B. anthracis<\/jats:italic>\n            spore, the exosporium, is a shell composed of approximately 20 proteins. The function of the exosporium remains poorly understood and is an area of active investigation. In this study, we analyzed the previously identified but uncharacterized exosporium protein ExsK. We found that, in contrast to other exosporium proteins, ExsK is present in at least two distinct locations, i.e., the spore surface as well as a more interior location underneath the exosporium. In spores that lack the exosporium basal layer protein ExsFA\/BxpB, ExsK fails to encircle the spore and instead is present at only one spore pole, indicating that ExsK assembly to the spore is partially dependent on ExsFA\/BxpB. In spores lacking the exosporium surface protein BclA, ExsK fails to mature into high-molecular-mass species observed in wild-type spores. These data suggest that the assembly and maturation of ExsK within the exosporium are dependent on ExsFA\/BxpB and BclA. We also found that ExsK is not required for virulence in murine and guinea pig models but that it does inhibit germination. Based on these data, we propose a revised model of exosporium maturation and assembly and suggest a novel role for the exosporium in germination.\n          <\/jats:p>","DOI":"10.1128\/jb.01110-09","type":"journal-article","created":{"date-parts":[[2009,10,17]],"date-time":"2009-10-17T00:46:30Z","timestamp":1255740390000},"page":"7587-7596","update-policy":"https:\/\/doi.org\/10.1128\/asmj-crossmark-policy-page","source":"Crossref","is-referenced-by-count":44,"title":["Roles of the\n            <i>Bacillus anthracis<\/i>\n            Spore Protein ExsK in Exosporium Maturation and Germination"],"prefix":"10.1128","volume":"191","author":[{"given":"Kari M.","family":"Severson","sequence":"first","affiliation":[{"name":"Department of Microbiology and Immunology, Loyola University Medical Center, Maywood, Illinois 60153"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Michael","family":"Mallozzi","sequence":"additional","affiliation":[{"name":"Department of Microbiology and Immunology, Loyola University Medical Center, Maywood, Illinois 60153"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Joel","family":"Bozue","sequence":"additional","affiliation":[{"name":"Bacteriology Division, United States Army Medical Research Institute of Infectious Diseases, Fort Detrick, Frederick, Maryland 21702-5011"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Susan L.","family":"Welkos","sequence":"additional","affiliation":[{"name":"Bacteriology Division, United States Army Medical Research Institute of Infectious Diseases, Fort Detrick, Frederick, Maryland 21702-5011"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Christopher K.","family":"Cote","sequence":"additional","affiliation":[{"name":"Bacteriology Division, United States Army Medical Research Institute of Infectious Diseases, Fort Detrick, Frederick, Maryland 21702-5011"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Katherine L.","family":"Knight","sequence":"additional","affiliation":[{"name":"Department of Microbiology and Immunology, Loyola University Medical Center, Maywood, Illinois 60153"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Adam","family":"Driks","sequence":"additional","affiliation":[{"name":"Department of Microbiology and Immunology, Loyola University Medical Center, Maywood, Illinois 60153"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"235","reference":[{"key":"e_1_3_2_2_2","doi-asserted-by":"crossref","first-page":"33","DOI":"10.1016\/j.femsle.2005.02.016","volume":"245","year":"2005","unstructured":"Baillie, L., S. Hibbs, P. Tsai, G. L. Cao, and G. M. Rosen. 2005. Role of superoxide in the germination of Bacillus anthracis endospores. FEMS Microbiol. Lett. 245 : 33-38.","journal-title":"FEMS Microbiol. Lett."},{"key":"e_1_3_2_3_2","doi-asserted-by":"crossref","first-page":"947","DOI":"10.1111\/j.1365-2958.2008.06206.x","volume":"68","year":"2008","unstructured":"Ball, D. A., R. Taylor, S. J. Todd, C. Redmond, E. Couture-Tosi, P. Sylvestre, A. Moir, and P. A. Bullough. 2008. Structure of the exosporium and sublayers of spores of the Bacillus cereus family revealed by electron crystallography. Mol. Microbiol. 68 : 947-958.","journal-title":"Mol. Microbiol."},{"key":"e_1_3_2_4_2","doi-asserted-by":"crossref","first-page":"2089","DOI":"10.1099\/00221287-148-7-2089","volume":"148","year":"2002","unstructured":"Barlass, P. J., C. W. Houston, M. O. Clements, and A. Moir. 2002. Germination of Bacillus cereus spores in response to L-alanine and to inosine: the roles of gerL and gerQ operons. Microbiology 148 : 2089-2095.","journal-title":"Microbiology"},{"key":"e_1_3_2_5_2","doi-asserted-by":"publisher","DOI":"10.1128\/JB.181.22.7043-7051.1999"},{"key":"e_1_3_2_6_2","doi-asserted-by":"publisher","DOI":"10.1128\/IAI.01202-06"},{"key":"e_1_3_2_7_2","doi-asserted-by":"publisher","DOI":"10.1128\/IAI.00434-07"},{"key":"e_1_3_2_8_2","doi-asserted-by":"publisher","DOI":"10.1128\/IAI.00660-07"},{"key":"e_1_3_2_9_2","doi-asserted-by":"publisher","DOI":"10.1128\/JB.01098-08"},{"key":"e_1_3_2_10_2","doi-asserted-by":"crossref","first-page":"619","DOI":"10.1099\/mic.0.2007\/008292-0","volume":"154","year":"2008","unstructured":"Cote, C. K., J. Bozue, K. L. Moody, T. L. DiMezzo, C. E. Chapman, and S. L. Welkos. 2008. Analysis of a novel spore antigen in Bacillus anthracis that contributes to spore opsonization. Microbiology 154 : 619-632.","journal-title":"Microbiology"},{"key":"e_1_3_2_11_2","doi-asserted-by":"publisher","DOI":"10.1128\/MMBR.63.1.1-20.1999"},{"key":"e_1_3_2_12_2","doi-asserted-by":"crossref","first-page":"251","DOI":"10.1016\/S0966-842X(02)02373-9","volume":"10","year":"2002","unstructured":"Driks, A. 2002. Maximum shields: the assembly and function of the bacterial spore coat. Trends Microbiol. 10 : 251-254.","journal-title":"Trends Microbiol."},{"key":"e_1_3_2_13_2","doi-asserted-by":"publisher","DOI":"10.1128\/JB.00921-06"},{"key":"e_1_3_2_14_2","doi-asserted-by":"crossref","first-page":"1133","DOI":"10.1099\/mic.0.023333-0","volume":"155","year":"2009","unstructured":"Giorno, R., M. Mallozzi, J. Bozue, K. S. Moody, A. Slack, D. Qiu, R. Wang, A. Friedlander, S. Welkos, and A. Driks. 2009. Localization and assembly of proteins comprising the outer structures of the Bacillus anthracis spore. Microbiology 155 : 1133-1145.","journal-title":"Microbiology"},{"key":"e_1_3_2_15_2","doi-asserted-by":"publisher","DOI":"10.1128\/jb.92.4.1261-1262.1966"},{"key":"e_1_3_2_16_2","unstructured":"Molecular biological methods for Bacillus. 1990"},{"key":"e_1_3_2_17_2","doi-asserted-by":"publisher","DOI":"10.1128\/jb.177.12.3394-3406.1995"},{"key":"e_1_3_2_18_2","doi-asserted-by":"crossref","first-page":"555","DOI":"10.1146\/annurev.micro.61.080706.093224","volume":"61","year":"2007","unstructured":"Henriques, A. O., and C. P. Moran, Jr. 2007. Structure, assembly, and function of the spore surface layers. Annu. Rev. Microbiol. 61 : 555-588.","journal-title":"Annu. Rev. Microbiol."},{"key":"e_1_3_2_19_2","doi-asserted-by":"publisher","DOI":"10.1128\/JB.184.5.1296-1303.2002"},{"key":"e_1_3_2_20_2","doi-asserted-by":"crossref","first-page":"487","DOI":"10.1111\/j.1365-2958.2005.04968.x","volume":"59","year":"2006","unstructured":"Kim, H., M. Hahn, P. Grabowski, D. C. McPherson, M. M. Otte, R. Wang, C. C. Ferguson, P. Eichenberger, and A. Driks. 2006. The Bacillus subtilis spore coat protein interaction network. Mol. Microbiol. 59 : 487-502.","journal-title":"Mol. Microbiol."},{"key":"e_1_3_2_21_2","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1073\/pnas.0507121102","volume":"103","year":"2006","unstructured":"Klobutcher, L. A., K. Ragkousi, and P. Setlow. 2006. The Bacillus subtilis spore coat provides \u201ceat resistance\u201d during phagocytic predation by the protozoan Tetrahymena thermophila. Proc. Natl. Acad. Sci. USA 103 : 165-170.","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"e_1_3_2_22_2","doi-asserted-by":"crossref","first-page":"1109","DOI":"10.1271\/bbb.62.1109","volume":"62","year":"1998","unstructured":"Kobayashi, K., K. Hashiguchi, K. Yokozeki, and S. Yamanaka. 1998. Molecular cloning of the transglutaminase gene from Bacillus subtilis and its expression in Escherichia coli. Biosci. Biotechnol. Biochem. 62 : 1109-1114.","journal-title":"Biosci. Biotechnol. Biochem."},{"key":"e_1_3_2_23_2","first-page":"157","volume":"144","year":"1996","unstructured":"Kobayashi, K., Y. Kumazawa, K. Miwa, and S. Yamanaka. 1996. \u03b5-(\u03b3-Glutamyl) lysine cross-links of spore coat proteins and transglutaminase activity in Bacillus subtilis. FEMS Microbiol. Lett. 144 : 157-160.","journal-title":"FEMS Microbiol. Lett."},{"key":"e_1_3_2_24_2","doi-asserted-by":"crossref","first-page":"553","DOI":"10.1016\/S1369-5274(00)00140-5","volume":"3","year":"2000","unstructured":"Kroos, L., and Y. T. Yu. 2000. Regulation of sigma factor activity during Bacillus subtilis development. Curr. Opin. Microbiol. 3 : 553-560.","journal-title":"Curr. Opin. Microbiol."},{"key":"e_1_3_2_25_2","doi-asserted-by":"crossref","first-page":"195","DOI":"10.1099\/ijs.0.02747-0","volume":"54","year":"2004","unstructured":"La Duc, M. T., M. Satomi, and K. Venkateswaran. 2004. Bacillus odysseyi sp. nov., a round-spore-forming bacillus isolated from the Mars Odyssey spacecraft. Int. J. Syst. Evol. Microbiol. 54 : 195-201.","journal-title":"Int. J. Syst. Evol. Microbiol."},{"key":"e_1_3_2_26_2","doi-asserted-by":"crossref","first-page":"1333","DOI":"10.1099\/mic.0.2007\/014399-0","volume":"154","year":"2008","unstructured":"Liang, L., X. He, G. Liu, and H. Tan. 2008. The role of a purine-specific nucleoside hydrolase in spore germination of Bacillus thuringiensis. Microbiology 154 : 1333-1340.","journal-title":"Microbiology"},{"key":"e_1_3_2_27_2","doi-asserted-by":"publisher","DOI":"10.1128\/iai.52.2.509-512.1986"},{"key":"e_1_3_2_28_2","doi-asserted-by":"publisher","DOI":"10.1128\/JB.186.1.164-178.2004"},{"key":"e_1_3_2_29_2","doi-asserted-by":"crossref","first-page":"110","DOI":"10.1111\/j.1574-6968.2008.01380.x","volume":"289","year":"2008","unstructured":"Mallozzi, M., J. Bozue, R. Giorno, K. S. Moody, A. Slack, C. Cote, D. Qiu, R. Wang, P. McKenney, E. M. Lai, J. R. Maddock, A. Friedlander, S. Welkos, P. Eichenberger, and A. Driks. 2008. Characterization of a Bacillus anthracis spore coat-surface protein that influences coat-surface morphology. FEMS Microbiol. Lett. 289 : 110-117.","journal-title":"FEMS Microbiol. Lett."},{"key":"e_1_3_2_30_2","doi-asserted-by":"publisher","DOI":"10.1128\/IAI.00727-07"},{"key":"e_1_3_2_31_2","doi-asserted-by":"publisher","DOI":"10.1128\/JB.01116-06"},{"key":"e_1_3_2_32_2","unstructured":"Assembly and localization of a Bacillus anthracis spore coat protein that undergoes processing."},{"key":"e_1_3_2_33_2","unstructured":"Guide for the care and use of laboratory animals. 1996"},{"key":"e_1_3_2_34_2","doi-asserted-by":"publisher","DOI":"10.1128\/MMBR.64.3.548-572.2000"},{"key":"e_1_3_2_35_2","doi-asserted-by":"publisher","DOI":"10.1128\/jb.115.3.1179-1190.1973"},{"key":"e_1_3_2_36_2","doi-asserted-by":"crossref","first-page":"1261","DOI":"10.1073\/pnas.0709321105","volume":"105","year":"2008","unstructured":"Oliva, C. R., M. K. Swiecki, C. E. Griguer, M. W. Lisanby, D. C. Bullard, C. L. Turnbough, Jr., and J. F. Kearney. 2008. The integrin Mac-1 (CR3) mediates internalization and directs Bacillus anthracis spores into professional phagocytes. Proc. Natl. Acad. Sci. USA 105 : 1261-1266.","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"e_1_3_2_37_2","first-page":"483","year":"2002","unstructured":"Piggot, P., and R. Losick. 2002. Sporulation genes and intercompartmental regulation, p. 483-517. In A. L. Sonenshein, J. A. Hoch, and R. Losick (ed.), Bacillus subtilis and its closest relatives: from genes to cells. American Society for Microbiology, Washington, DC.","journal-title":"Bacillus subtilis and its closest relatives: from genes to cells"},{"key":"e_1_3_2_38_2","doi-asserted-by":"crossref","first-page":"355","DOI":"10.1099\/mic.0.26681-0","volume":"150","year":"2004","unstructured":"Redmond, C., L. W. Baillie, S. Hibbs, A. J. Moir, and A. Moir. 2004. Identification of proteins in the exosporium of Bacillus anthracis. Microbiology 150 : 355-363.","journal-title":"Microbiology"},{"key":"e_1_3_2_39_2","unstructured":"Molecular cloning: a laboratory manual 1989"},{"key":"e_1_3_2_40_2","doi-asserted-by":"crossref","first-page":"550","DOI":"10.1016\/j.mib.2003.10.001","volume":"6","year":"2003","unstructured":"Setlow, P. 2003. Spore germination. Curr. Opin. Microbiol. 6 : 550-556.","journal-title":"Curr. Opin. Microbiol."},{"key":"e_1_3_2_41_2","doi-asserted-by":"crossref","first-page":"514","DOI":"10.1111\/j.1365-2672.2005.02736.x","volume":"101","year":"2006","unstructured":"Setlow, P. 2006. Spores of Bacillus subtilis: their resistance to and killing by radiation, heat and chemicals. J. Appl. Microbiol. 101 : 514-525.","journal-title":"J. Appl. Microbiol."},{"key":"e_1_3_2_42_2","doi-asserted-by":"crossref","first-page":"315","DOI":"10.1016\/j.femsle.2005.03.029","volume":"245","year":"2005","unstructured":"Shatalin, K. Y., and A. A. Neyfakh. 2005. Efficient gene inactivation in Bacillus anthracis. FEMS Microbiol. Lett. 245 : 315-319.","journal-title":"FEMS Microbiol. Lett."},{"key":"e_1_3_2_43_2","doi-asserted-by":"publisher","DOI":"10.1128\/JB.185.6.1903-1910.2003"},{"key":"e_1_3_2_44_2","doi-asserted-by":"publisher","DOI":"10.1128\/JB.187.17.5868-5876.2005"},{"key":"e_1_3_2_45_2","doi-asserted-by":"crossref","first-page":"359","DOI":"10.1111\/j.1365-2958.2007.05658.x","volume":"64","year":"2007","unstructured":"Steichen, C. T., J. F. Kearney, and C. L. Turnbough, Jr. 2007. Non-uniform assembly of the Bacillus anthracis exosporium and a bottle cap model for spore germination and outgrowth. Mol. Microbiol. 64 : 359-367.","journal-title":"Mol. Microbiol."},{"key":"e_1_3_2_46_2","doi-asserted-by":"crossref","first-page":"169","DOI":"10.1046\/j.1365-2958.2000.03000.x","volume":"45","year":"2002","unstructured":"Sylvestre, P., E. Couture-Tosi, and M. Mock. 2002. A collagen-like surface glycoprotein is a structural component of the Bacillus anthracis exosporium. Mol. Microbiol. 45 : 169-178.","journal-title":"Mol. Microbiol."},{"key":"e_1_3_2_47_2","doi-asserted-by":"publisher","DOI":"10.1128\/JB.187.15.5122-5128.2005"},{"key":"e_1_3_2_48_2","doi-asserted-by":"crossref","first-page":"269","DOI":"10.1111\/j.1365-2672.1987.tb02408.x","volume":"62","year":"1987","unstructured":"Titball, R. W., and R. J. Manchee. 1987. Factors affecting the germination of spores of Bacillus anthracis. J. Appl. Bacteriol. 62 : 269-273.","journal-title":"J. Appl. Bacteriol."},{"key":"e_1_3_2_49_2","doi-asserted-by":"publisher","DOI":"10.1128\/JB.185.11.3373-3378.2003"},{"key":"e_1_3_2_50_2","doi-asserted-by":"crossref","first-page":"1001","DOI":"10.1099\/13500872-140-5-1001","volume":"140","year":"1994","unstructured":"Vary, P. S. 1994. Prime time for Bacillus megaterium. Microbiology 140 : 1001-1013.","journal-title":"Microbiology"},{"key":"e_1_3_2_51_2","doi-asserted-by":"publisher","DOI":"10.1128\/IAI.00283-07"},{"key":"e_1_3_2_52_2","doi-asserted-by":"publisher","DOI":"10.1128\/JB.185.4.1462-1464.2003"},{"key":"e_1_3_2_53_2","doi-asserted-by":"crossref","first-page":"253","DOI":"10.1016\/j.mimet.2003.10.019","volume":"56","year":"2004","unstructured":"Welkos, S. L., C. K. Cote, K. M. Rea, and P. H. Gibbs. 2004. A microtiter fluorometric assay to detect the germination of Bacillus anthracis spores and the germination inhibitory effects of antibodies. J. Microbiol. Methods 56 : 253-265.","journal-title":"J. Microbiol. Methods"},{"key":"e_1_3_2_54_2","unstructured":"Reference deleted."},{"key":"e_1_3_2_55_2","doi-asserted-by":"publisher","DOI":"10.1128\/JB.187.22.7753-7764.2005"},{"key":"e_1_3_2_56_2","doi-asserted-by":"publisher","DOI":"10.1128\/JB.186.4.1110-1119.2004"}],"container-title":["Journal of Bacteriology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/journals.asm.org\/doi\/pdf\/10.1128\/JB.01110-09","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/journals.asm.org\/doi\/pdf\/10.1128\/JB.01110-09","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,7,29]],"date-time":"2021-07-29T17:10:20Z","timestamp":1627578620000},"score":1,"resource":{"primary":{"URL":"https:\/\/journals.asm.org\/doi\/10.1128\/JB.01110-09"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2009,12,15]]},"references-count":55,"journal-issue":{"issue":"24","published-print":{"date-parts":[[2009,12,15]]}},"alternative-id":["10.1128\/JB.01110-09"],"URL":"https:\/\/doi.org\/10.1128\/jb.01110-09","relation":{},"ISSN":["0021-9193","1098-5530"],"issn-type":[{"value":"0021-9193","type":"print"},{"value":"1098-5530","type":"electronic"}],"subject":[],"published":{"date-parts":[[2009,12,15]]},"assertion":[{"value":"2009-08-20","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2009-10-07","order":1,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2020-12-20","order":2,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}]}}