Prophages Associated with other Streptococcal Species

Temperate bacteriophages are associated with a number of the other streptococcal species and may well form part of a common pool of phage genetic material that is available for horizontal transfer of virulence and other genes. For example, S. mitis is an occasional cause of infective endocarditis, and bacterial binding of platelets is an important mechanism in the pathogenesis of this disease. Platelet binding by S. mitis is mediated by the surface proteins PblA and PblB, encoded by a lysogenic bacteriophage [66, 67]. Thus, it may well be that S. mitis shares a number of genetic characteristics with the better known pathogenic streptococci, and the analysis of the genome of this organism may reveal even further insights into potential virulence mechanisms. No similar association between a temperate phage and the transmission of virulence traits has been seen yet in S. sanguis or S.gordonii. Bacteriophages have been isolated from both S.bovis and S.suis [5356]; however, any role that these elements play in the dissemination of virulence genes or horizontal transfer is unknown.

Note added in proof

Since the preparation of this chapter, two additional genomes of Group A Streptococci have been published; i.e.,

Sumby P, Porcella SF, Madrigal AG, Barbian KD, Virtaneva K, Ricklefs SM, Stur-devant DE, Graham MR, Vuopio-Varkila J, Hoe NP, Musser JM. Evolutionary Origin and Emergence of a Highly Successful Clone of Serotype Ml Group A Streptococcus Involved Multiple Horizontal Gene Transfer Events, J. Infect. Dis. (2005) 192:771-782; and

Nicole M. Green, Stephen B. Beres, Edward A. Graviss, James E. Allison, Allison J. McGeer, Jaana Vuopio-Varkila, Rance B. LeFebvre, and James M. Musser, Genetic Diversity among Type emm28 Group A Streptococcus Strains Causing Invasive Infections and Pharyngitis, J. Clin. Microbiol. (2005) Vol. 43:4083-4091.

And six additional Group B Streptococcus genomes - in one publication: Hervé Tettelin, Vega Masignani, Michael J. Cieslewicz, Claudio Donati, Duccio Medini, Naomi L. Ward, Samuel V. Angiuoli, Jonathan Crabtree, Amanda L. Jones, A. Scott Durkin, Robert T. DeBoy, Tanja M. Davidsen, Marirosa Mora, Maria Scarselli, Immaculada Margarit y Ros, Jeremy D. Peterson, Christopher R. Hauser, Jaideep P. Sundaram, William C. Nelson, Ramana Madupu, Lauren M. Brinkac, Robert J. Dodson, Mary J. Rosovitz, Steven A. Sullivan, Sean C. Daugh-erty, Daniel H. Haft, Jeremy Selengut, Michelle L. Gwinn, Liwei Zhou, Nikhat Zafar, Hoda Khouri, Diana Radune, George Dimitrov, Kisha Watkins, Kevin J. B. O'Connor, Shannon Smith, Teresa R. Utterback, Owen White, Craig E. Rubens, Guido Grandi, Lawrence C. Madoff, Dennis L. Kasper, John L. Telford, Michael R. Wessels, Rino Rappuoli, and Claire M. Fraser. Genome analysis of multiple pathogenic isolates of Streptococcus agalactiae: Implications for the microbial "pangenome". PNAS (2005) 102: 13950-13955.

References universal common ancestor. Nat Rev Microbiol 1: 127-136. 2 Lancefield, R. C. 1928. The antigenic

1 Koonin, E. V. 2003. Comparative geno mics, minimal gene-sets and the last

4 Fischetti, V. A., R. P. Novick, J. J. Fer-retti, D. A. Portnoy, and J. I. Rood, editors. 2005. Gram-Positive Pathogens. ASM Press, Washington, DC

complex of Streptococcus haemolyticus.

5 Ferretti, J. J., W. M. McShan, D. Ajdic, D. J. Savic, G. Savic, K. Lyon, C. Pri-

III. Chemical and immunological prop-

erties of the species-specific substance. J Exp Med 47: 481-491. 3 Kawamura, Y., X. G. Hou, F. Sultana, ton, H. Lai, S. Lin, Y. Qian, H. G. Jia, F. Z. Najar, Q. Ren, H. Zhu, L. Song, J. White, X. Yuan, S. W. Clifton, B. A. Roe, and R. McLaughlin. 2001. Com-

H. Miura, andT. Ezaki. 1995. Determination of 16S rRNA sequences of Streptococcus mitis and Streptococcus gordonii and phylogenetic relationships among plete genome sequence of an Ml strain of Streptococcus pyogenes. Proc Natl Acad Sci USA 98: 4658-4663.

members of the genus Streptococcus. Int J Syst Bacteriol 45: 406-408.

6 Beres, S. B., G. L. Sylva, K. D. Barbian, B. Lei, J. S. Hoff, N. D. Mammarella,

M. Y. Liu, J. C. Smoot, S. F. Porcella, L. D. Parkins, D. S. Campbell, T. M. Smith, J. K. McCormick, D. Y. Leung, P. M. Schlievert, and J. M. Musser. 2002. Genome sequence of a serotype M3 strain of group A Streptococcus: phage-encoded toxins, the high-virulence phenotype, and clone emergence. Proc Natl Acad Sci U S A 99: 1007810083.

7 Nakagawa, I., K. Kurokawa, A. Yama-shita, M. Nakata, Y. Tomiyasu, N. Oka-hashi, S. Kawabata, K. Yamazaki,

T. Shiba, T. Yasunaga, H. Hayashi, M. Hattori, and S. Hamada. 2003. Genome sequence of an M3 strain of Streptococcus pyogenes reveals a large-scale genomic rearrangement in invasive strains and new insights into phage evolution. Genome Res 13: 1042-1055.

8 Banks, D. J., S. F. Porcella, K. D. Barbian, S. B. Beres, L. E. Philips, J. M. Voyich, F. R. DeLeo, J. M. Martin, G. A. Somerville, and J. M. Musser. 2004. Progress toward characterization of the group A Streptococcus metagenome: complete genome sequence of a macro-lide-resistant serotype M6 strain. J Infect Dis 190: 727-738.

9 Smoot, J. C., K. D. Barbian, J. J. Van Gompel, L. M. Smoot, M. S. Chaussee, G. L. Sylva, D. E. Sturdevant, S. M. Rick-lefs, S. F. Porcella, L. D. Parkins, S. B. Beres, D. S. Campbell, T. M. Smith,

Q. Zhang, V. Kapur, J. A. Daly, L. G. Veasy, and J. M. Musser. 2002. Genome sequence and comparative microarray analysis of serotype M18 group A Streptococcus strains associated with acute rheumatic fever outbreaks. Proc Natl Acad Sci U S A 99: 4668-4673.

10 Hynes, W. L., L. Hancock, and J. J. Fer-retti. 1995. Analysis of a second bacteriophage hyaluronidase gene from Streptococcus pyogenes: evidence for a third hyaluronidase involved in extracellular enzymatic activity. Infect Immun 63: 3015-3020.

11 Darling, A. C., B. Mau, F. R. Blattner, and N. T. Perna. 2004. Mauve: multiple alignment ofconserved genomic sequence with rearrangements. Genome Res 14: 1394-1403.

12 Beres, S. B., G. L. Sylva, D. E. Sturdevant, C. N. Granville, M. Liu, S. M. Rick-lefs, A. R. Whitney, L. D. Parkins, N. P. Hoe, G. J. Adams, D. E. Low, F. R. DeLeo, A. McGeer, and J. M. Musser. 2004. Genome-wide molecular dissection ofserotype M3 group A Streptococcus strains causing two epidemics of invasive infections. Proc Natl Acad Sci U S A 101: 11833-11838.

13 Reid, S. D., N. P. Hoe, L. M. Smoot, and J. M. Musser. 2001. Group A Streptococcus: allelic variation, population genetics, and host-pathogen interactions.

J Clin Invest 107: 393-399.

14 Stockbauer, K. E., D. Grigsby, X. Pan, Y. X. Fu, L. M. Mejia, A. Cravioto, and J. M. Musser. 1998. Hypervariability generated by natural selection in an extracellular complement-inhibiting protein of serotype M1 strains of group A Streptococcus. Proc Natl Acad Sci U S A 95: 3128-3133.

15 Lukomski, S., K. Nakashima, I. Abdi, V. J. Cipriano, B. J. Shelvin, E. A. Grav-iss, and J. M. Musser. 2001. Identification and characterization of a second extracellular collagen-like protein made by group A Streptococcus: control of production at the level of translation. Infect Immun 69: 1729-1738.

16 Huang, T. T., H. Malke, and J. J. Ferretti. 1989. Heterogeneity of the streptokinase gene in group A streptococci. Infect Immun 57: 502-506.

17 Musser, J. M., A. R. Hauser, M. H. Kim, P. M. Schlievert, K. Nelson, and R. K. Selander. 1991. Streptococcus pyogenes causing toxic-shock-like syndrome and other invasive diseases: clonal diversity and pyrogenic exotoxin expression. Proc Natl Acad Sci USA 88: 2668-2672.

18 Stockbauer, K. E., L. Magoun, M. Liu, E. H. Burns, Jr., S. Gubba, S. Renish,

X. Pan, S. C. Bodary, E. Baker, J. Coburn, J. M. Leong, and J. M. Musser. 1999. A natural variant ofthe cysteine protease virulence factor of group A Streptococcus with an arginine-glycine-aspartic acid (RGD) motif preferentially binds human integrins avß3 and anbß3. Proc Natl Acad Sci U S A 96: 242-247.

19 McGregor, K. F., B. G. Spratt, A. Kalia, A. Bennett, N. Bilek, B. Beall, and D. E.

Bessen. 2004. Multilocus sequence typing of Streptococcus pyogenes representing most known emm types and distinctions among subpopulation genetic structures. J Bacteriol 186: 4285-4294.

20 Hynes, W. 2004. Virulence factors of the group A streptococci and genes that regulate their expression. Front Biosci 9: 3399-3433.

21 Nagiec, M. J., B. Lei, S. K. Parker, M. L. Vasil, M. Matsumoto, R. M. Ireland,

S. B. Beres, N. P. Hoe, and J. M. Musser. 2004. Analysis of a novel prophage-encoded group A Streptococcus extracellular phospholipase A(2). J Biol Chem 279: 45909-45918.

22 Collin, M., and A. Olsen. 2001. EndoS, a novel secreted protein from Streptococcus pyogenes with endoglycosidase activity on human IgG. EMBO J 20: 30463055.

23 Coye, L. H., and C. M. Collins. 2004. Identification of SpyA, a novel ADP-ribosyltransferase of Streptococcus pyo-genes. Mol Microbiol 54: 89-98.

24 Terao, Y., S. Kawabata, E. Kunitomo, I. Nakagawa, and S. Hamada. 2002. Novel laminin-binding protein of Streptococcus pyogenes, Lbp, is involved in adhesion to epithelial cells. Infect Immun 70: 993-997.

25 von Pawel-Rammingen, U., B. P. Johansson, and L. Bjorck. 2002. IdeS, a novel streptococcal cysteine proteinase with unique specificity for immunoglo-bulin G. EMBO J 21: 1607-1615.

26 Beall, B., G. Gherardi, M. Lovgren, R. R. Facklam, B. A. Forwick, and G. J. Tyrrell. 2000. emm and sof gene sequence variation in relation to serological typing of opacity-factor-positive group A streptococci. Microbiology 146( Pt 5): 11951209.

27 Hendrix, R. W., J. G. Lawrence, G. F. Hatfull, and S. Casjens. 2000. The origins and ongoing evolution of viruses. Trends Microbiol 8: 504-508.

28 McCornick, J. K., Peterson, M. L., and Schlievert, A. M. 2005. Toxins and Superantigens of Group A Streptococci. In: Gram-Positive Pathogens. Fischetti, V. A., R. P. Novick, J. J. Ferretti, D. A. Portnoy, and J. I. Rood, editors. ASM Press, Washington, DC

29 Aziz, R. K., S. A. Ismail, H. W. Park, and M. Kotb. 2004. Post-proteomic identification of a novel phage-encoded streptodornase, Sdal, in invasive M1T1 Streptococcus pyogenes. Mol Microbiol 54: 184-197.

30 Malke, H. 1979. Conjugal transfer of plasmids determining resistance to macrolides, lincosomides and strepto-gramin-B type antibiotics among group A, B, D and H streptococci. FEMS Microbiol Lett 5: 335-338.

31 Hidalgo-Grass, C., M. Ravins, M. Dan-Goor, J. Jaffe, A. E. Moses, and

E. Hanski. 2002. A locus ofgroup A Streptococcus involved in invasive disease and DNA transfer. Mol Microbiol 46: 87-99.

32 Berge, A., M. Rasmussen, and L. Bjorck. 1998. Identification of an insertion sequence located in a region encoding virulence factors of Streptococcus pyogenes. Infect Immun 66: 3449-3453.

F. Campanile, M. R. Oggioni, S. Stefani, and G. Pozzi. 2003. The novel conjuga-tive transposon tn1207.3 carries the macrolide efflux gene mef(A) in Streptococcus pyogenes. Microb Drug Resist 9: 243-247.

U. Dobrindt. 2003. Prokaryotic chromosomes and disease. Science 301: 790793.

35 Wren, B. W. 2000. Microbial genome analysis: insights into virulence, host adaptation and evolution. Nat Rev Genet 1: 30-39.

36 Chaffin, D. O., S. B. Beres, H. H. Yim, and C. E. Rubens. 2000. The serotype of type Ia and III group B streptococci is determined by the polymerase gene within the polycistronic capsule operon. J Bacteriol 182: 4466-4477.

37 Glaser, P., C. Rusniok, C. Buchrieser, F. Chevalier, L. Frangeul, T. Msadek, M. Zouine, E. Couve, L. Lalioui,

C. Poyart, P. Trieu-Cuot, and F. Kunst. 2002. Genome sequence ofStreptococcus agalactiae, a pathogen causing invasive neonatal disease. Mol Microbiol 45: 1499-1513.

38 Tettelin, H., V. Masignani, M. J. Ciesle-wicz, J. A. Eisen, S. Peterson, M. R.

Wessels, I. T. Paulsen, K. E. Nelson, I. Margarit, T. D. Read, L. C. Madoff, A. M. Wolf, M. J. Beanan, L. M. Brinkac, S. C. Daugherty, R. T. DeBoy, A. S. Dur-kin, J. F. Kolonay, R. Madupu, M. R. Lewis, D. Radune, N. B. Fedorova, D. Scanlan, H. Khouri, S. Mulligan, H. A. Carty, R. T. Cline, S. E. Van Aken, J. Gill, M. Scarselli, M. Mora, E. T. Iaco-bini, C. Brettoni, G. Galli, M. Mariani,

G. Grandi, and C. M. Fraser. 2002. Complete genome sequence and comparative genomic analysis of an emerging human pathogen, serotype V Streptococcus agalactiae. Proc Natl Acad Sci USA 99: 12391-12396.

G. Bakken. 1981. Serotyping and bacter-iophage typing of human and bovine group-B streptococci. J Med Microbiol 14: 479-482.

40 Russell, H., N. L. Norcross, and D. E. Kahn. 1969. Isolation and characterization of Streptococcus agalactiae bacteriophage. J Gen Virol 5: 315-317.

41 Stringer, J. 1980. The development of a phage-typing system for group-B streptococci. J Med Microbiol 13: 133-143.

42 Boyer, K., L. Vogel, S. Gotoff, C. Gad-zala, J. Stringer, and W. Maxted. 1980. Nosocomial transmission of bacteriophage type 7/11/12 group B streptococci in a special care nursery. Am J Dis Child 134: 964-966.

43 Franken, C., G. Haase, C. Brandt, J. Weber-Heynemann, S. Martin,

C. Lammler, A. Podbielski, R. Lutticken, and B. Spellerberg. 2001. Horizontal gene transfer and host specificity of beta-haemolytic streptococci: the role of a putative composite transposon containing scpB and lmb. Mol Microbiol 41: 925-935.

44 Malke. H. 2005. Genetics and Pathogenicity Factors of Group C and G Streptococci. In: Gram-Positive Pathogens. Fischetti, V. A., R. P. Novick, J. J. Ferretti, D. A. Portnoy, and J. I. Rood, editors. ASM Press, Washington, DC

45 Carmeli, Y., and K. L. Ruoff.1995. Report of cases of and taxonomic considerations for large-colony-forming

Lancefield group C streptococcal bacteremia. J Clin Microbiol 33: 2114-2117.

E. Gunther, J. Rodel, E. Straube, and K. H. Schmidt. 2002. Superantigen-like gene(s) in human pathogenic Streptococcus dysgalactiae, subsp equisimilis: genomic localisation of the gene encoding streptococcal pyrogenic exotoxin G (speG(dys)). FEMS Immunol Med Microbiol 34: 159-167.

47 Igwe, E. I., P. L. Shewmaker, R. R. Fack-lam, M. M. Farley, C. van Beneden, and B. Beall. 2003. Identification of superantigen genes speM, ssa, and smeZ in invasive strains of beta-hemolytic group C and G streptococci recovered from humans. FEMS Microbiol Lett 229: 259-264.

48 Proft, T., P. D. Webb, V. Handley, and J. D. Fraser. 2003. Two novel superantigens found in both group A and group C Streptococcus. Infect Immun 71: 13611369.

49 Oliver, S. P., R. A. Almeida, and L. F. Calvinho. 1998. Virulence factors of Streptococcus uberis isolated from cows with mastitis. Zentralbl Veterinarmed B 45: 461-471.

50 Gase, K., J. J. Ferretti, C. Primeaux, and W. M. McShan. 1999. Identification, cloning, and expression of the CAMP factor gene (cfa) of group A streptococci. Infect Immun 67: 4725-4731.

51 Rosey, E. L., R. A. Lincoln, P. N. Ward, R. J. Yancey, Jr., and J. A. Leigh. 1999. PauA: a novel plasminogen activator from Streptococcus uberis. FEMS Microbiol Lett 178: 27-33.

52 Staats, J. J., B. L. Plattner, G. C. Stewart, and M. M. Changappa. 1999. Presence ofthe Streptococcus suis suilysin gene and expression of MRP and EF correlates with high virulence in Streptococcus suis type 2 isolates. Vet Microbiol 70: 201-211.

H. Dezfulian, S. J. Labrie, R. Brousseau, S. Moineau, and M. Gottschalk. 2003. Identification of an inducible bacteriophage in a virulent strain of Streptococcus suis serotype 2. Infect Immun 71: 6104-6108.

54 Tarakanov, B. V. 1996. [Biology oflyso-genic strains of Streptococcus bovis and virulent mutants of their temperate phages]. Mikrobiologiia 65: 656-662. In Russian.

55 Klieve, A. V., G. L. Heck, M. A. Prance, and Q. Shu. 1999. Genetic homogeneity and phage susceptibility of ruminal strains of Streptococcus bovis isolated in Australia. Lett Appl Microbiol 29: 108112.

56 Styriak, I., P. Pristas, and P. Javorsky. 1998. Lack of surface receptors not restriction-modification system determines F4 phage resistance in Streptococcus bovis II/1. Folia Microbiol (Praha) 43: 35-38.

57 Hoskins, J., W. E. Alborn, Jr., J. Arnold, L. C. Blaszczak, S. Burgett, B. S. DeHoff, S. T. Estrem, L. Fritz, D. J. Fu,

W. Fuller, C. Geringer, R. Gilmour, J. S. Glass, H. Khoja, A. R. Kraft, R. E. Lagace, D. J. LeBlanc, L. N. Lee, E. J. Lefkowitz, J. Lu, P. Matsushima, S. M. McAhren, M. McHenney, K. McLeaster, C. W. Mundy, T. I. Nicas, F. H. Norris, M. O'Gara, R. B. Peery, G. T. Robertson, P. Rockey, P. M. Sun, M. E. Winkler, Y. Yang, M. Young-Bellido, G. Zhao, C. A. Zook, R. H. Baltz, S. R. Jaskunas, P. R. Rosteck, Jr., P. L. Skatrud, and J. I. Glass. 2001. Genome of the bacterium Streptococcus pneumoniae strain R6. J Bacteriol 183: 5709-5717.

58 Tettelin, H., K. E. Nelson, I. T. Paulsen, J. A. Eisen, T. D. Read, S. Peterson,

J. Heidelberg, R. T. DeBoy, D. H. Haft, R. J. Dodson, A. S. Durkin, M. Gwinn, J. F. Kolonay, W. C. Nelson, J. D. Peterson, L. A. Umayam, O. White, S. L. Salzberg, M. R. Lewis, D. Radune, E. Holtzapple, H. Khouri, A. M. Wolf, T. R. Utterback, C. L. Hansen, L. A. McDonald, T. V. Feldblyum, S. Angiuoli, T. Dickinson, E. K. Hickey, I. E. Holt, B. J. Loftus, F. Yang, H. O. Smith, J. C. Venter, B. A. Dougherty, D. A. Morrison, S. K. Hollingshead, and C. M. Fraser. 2001. Complete genome sequence of a virulent isolate of Streptococcus pneumo-niae. Science 293: 498-506.

59 Claverys, J. P., M. Prudhomme, I. Mortier-Barriere, and B. Martin. 2000. Adaptation to the environment: Streptococcus pneumoniae, a paradigm for recombination-mediated genetic plasticity? Mol Microbiol 35: 251-259.

60 Martin-Galiano, A. J., J. M. Wells, and A. G. de la Campa. 2004. Relationship between codon biased genes, microarray expression values and physiological characteristics of Streptococcus pneumoniae. Microbiology 150: 2313-2325.

61 Dowson, C. G., T. J. Coffey, and B. G. Spratt. 1994. Origin and molecular epidemiology of penicillin-binding-protein-mediated resistance to beta-lactam antibiotics. Trends Microbiol 2: 361-366.

62 Hakenbeck, R., N. Balmelle, B. Weber, C. Gardes, W. Keck, and A. de Saizieu. 2001. Mosaic genes and mosaic chromosomes: intra- and interspecies genomic variation of Streptococcus pneumoniae. Infect Immun 69: 2477-2486.

63 Matsushita, K., W. Fujimaki, H. Kato, T. Uchiyama, H. Igarashi, H. Ohkuni, S. Nagaoka, M. Kawagoe, S. Kotani, and H. Takada. 1995. Immunopathological activities of extracellular products of Streptococcus mitis, particularly a super-antigenic fraction. Infect Immun 63: 785-793.

64 Lu, H. Z., X. H. Weng, B. Zhu, H. Li, Y. K. Yin, Y. X. Zhang, D. W. Haas, and Y. W. Tang. 2003. Major outbreak of toxic shock-like syndrome caused by Streptococcus mitis. J Clin Microbiol 41: 3051-3055.

65 Neeleman, C., C. H. Klaassen, D. M. Klomberg, H. A. de Valk, and J. W. Mouton. 2004. Pneumolysin is a key factor in misidentification of macrolide-resistant Streptococcus pneumoniae and is a putative virulence factor of S. mitis and other streptococci. J Clin Microbiol 42: 4355-4357.

66 Siboo, I. R., B. A. Bensing, and P. M. Sullam. 2003. Genomic organization and molecular characterization of SM1, a temperate bacteriophage of Streptococcus mitis. J Bacteriol 185: 6968-6975.

67 Bensing, B. A., I. R. Siboo, and P. M. Sullam. 2001. Proteins PblA and PblB of Streptococcus mitis, which promote binding to human platelets, are encoded within a lysogenic bacteriophage. Infect Immun 69: 6186-6192.

68 Whiley, R. A., and D. Beighton. 1998. Current classification of the oral streptococci. Oral Microbiol Immunol 13: 195216.

69 Whiley, R. A., D. Beighton, T. G. Win-stanley, H. Y. Fraser, and J. M. Hardie. 1992. Streptococcus intermedius, Streptococcus constellatus, and Streptococcus anginosus (the Streptococcus milleri group): association with different body sites and clinical infections. J Clin Microbiol 30: 243-244.

70 Whiley, R. A., L. M. Hall, J. M. Hardie, and D. Beighton. 1999. A study of small-colony, beta-haemolytic, Lance-field group C streptococci within the anginosus group: description of Streptococcus constellatus subsp. pharyngis subsp. nov., associated with the human throat and pharyngitis. Int J Syst Bacter-iol 49 Pt 4: 1443-1449.

71 Nagamune, H., R. A. Whiley, T. Goto, Y. Inai, T. Maeda, J. M. Hardie, and H. Kourai. 2000. Distribution of the intermedilysin gene among the anginosus group streptococci and correlation between intermedilysin production and deep-seated infection with Streptococcus intermedius. J Clin Microbiol 38: 220226.

72 Tapp, J., M. Thollesson, and B. Herrmann. 2003. Phylogenetic relationships and genotyping of the genus Streptococcus by sequence determination of the RNase P RNA gene, rnpB. Int J Syst Evol Microbiol 53: 1861-1871.

73 Bolotin, A., B. Quinquis, P. Renault, A. Sorokin, S. D. Ehrlich, S. Kulakaus-kas, A. Lapidus, E. Goltsman, M. Mazur, G. D. Pusch, M. Fonstein, R. Overbeek, N. Kyprides, B. Purnelle, D. Prozzi, K. Ngui, D. Masuy, F. Hancy, S. Bur-teau, M. Boutry, J. Delcour, A. Goffeau, and P. Hols. 2004. Complete sequence and comparative genome analysis ofthe dairy bacterium Streptococcus thermophi-lus. Nat Biotechnol 22: 1554-1558.

74 Ajdic, D., W. McShan, R. McLaughlin, G. Savic, J. Chang, M. Carson, C. Pri-meaux, R. Tian, S. Kenton, H. Jia, S. Lin, Y. Qian, S. Li, H. Zhu, F. Najar, H. Lai, J. White, B. Roe, and J. Ferretti. 2002. Genome sequence of Streptococcus mutans UA159, a cariogenic dental pathogen. Proc Natl Acad Sci U S A 99: 14434-14439.

75 Schleifer, K. H., and R. Kilpper-Balz. 1984. Transfer of Streptococcus faecalis and Streptococcus faecium to the genus Enterococcus nom. rev. as Enterococcus faecalis comb. nov. and Enterococcus fae-cium comb. nov. Int J Syst Bacteriol 34: 31-34.

76 Steiner, K., and H. Malke. 2000. Life in protein-rich environments: the relA-in-dependent response of Streptococcus pyogenes to amino acid starvation. Mol Microbiol 38: 1004-1016.

77 McShan, W. M. 2005. The Bacteriophages of Group A Streptococci. In: Gram-Positive Pathogens. Fischetti, V. A., R. P. Novick, J. J. Fer-retti, D. A. Portnoy, and J. I. Rood (eds.). ASM Press, Washington, DC

78 Pritchard, D. G., S. Dong, J. R. Baker, and J. A. Engler. 2004. The bifunctional peptidoglycan lysin of Streptococcus aga-lactiae bacteriophage B30. Microbiology 150: 2079-2087.

79 Obregon, V., P. Garcia, R. Lopez, and J. L. Garcia. 2003. VO1, a temperate bacteriophage of the type 19A multire-sistant epidemic 8249 strain of Streptococcus pneumoniae: analysis of variability of lytic and putative C5 methyltransfer-ase genes. Microb Drug Resist 9: 7-15.

R. Lopez, and P. Garcia. 2003. Genome organization and molecular analysis of the temperate bacteriophage MM1 of Streptococcus pneumoniae. J Bacteriol 185: 2362-2368.

81 Boyce, J. D., B. E. Davidson, and A. J. Hillier. 1995. Identification of prophage genes expressed in lysogens of the Lac-tococcus lactis bacteriophage BK5-T. Appl Environ Microbiol 61: 4099-4104.

82 Ventura, M., C. Canchaya, M. Kleerebe-zem, W. M. de Vos, R. J. Siezen, and H. Brussow. 2003. The prophage sequences of Lactobacillus plantarum strain WCFS1. Virology 316: 245-255.

83 Ventura, M., S. Foley, A. Bruttin, S. C. Chennoufi, C. Canchaya, and H. Brus-sow. 2002. Transcription mapping as a tool in phage genomics: the case of the temperate Streptococcus thermophilus phage Sfi21. Virology 296: 62-76.

84 Canchaya, C., F. Desiere, W. McShan, J. Ferretti, J. Parkhill, and H. Brussow. 2002. Genome analysis of an inducible prophage and prophage remnants integrated in the Streptococcus pyogenes strain SF370. Virology 302: 245-258.

85 Desiere, F., W. M. McShan, D. van Sinderen, J. J. Ferretti, and H. Brussow. 2001. Comparative genomics reveals close genetic relationships between phages from dairy bacteria and pathogenic Streptococci: evolutionary implications for prophage-host interactions. Virology 288: 325-341.

T. J. Gibson. 1994. CLUSTALW: improving the sensitivity of progressive multiple sequence alignment through sequence weighting, positions-specific gap penalties and weight matrix choice. Nucleic Acids Res 22: 4673-4680.

87 Proux, C., D. van Sinderen, J. Suarez, P. Garcia, V. Ladero, G. F. Fitzgerald, F. Desiere, and H. Brussow. 2002. The dilemma of phage taxonomy illustrated by comparative genomics of Sfi21-like Siphoviridae in lactic acid bacteria.

J Bacteriol 184: 6026-6036.

88 Lucchini, S., F. Desiere, and H. Brussow. 1999. Comparative genomics of Streptococcus thermophilus phage species supports a modular evolution theory.

J Virol 73: 8647-8656.

89 Stanley, E., G. F. Fitzgerald, C. Le Mar-rec, B. Fayard, and D. van Sinderen. 1997. Sequence analysis and characterization of phi 01205, a temperate bacteriophage infecting Streptococcus thermophilus CNRZ1205. Microbiology 143(Pt 11): 3417-3429.

90 Tremblay, D. M., and S. Moineau. 1999. Complete genomic sequence ofthe lytic bacteriophage DT1 of Streptococcus thermophilus. Virology 255: 63-76.

91 Banks, D. J., S. B. Beres, and J. M. Musser. 2002. The fundamental contribution of phages to GAS evolution, genome diversification and strain emergence. Trends Microbiol 10: 515-521.

92 Campbell, A. M. 1992. Chromosomal insertion sites for phages and plasmids. J Bacteriol 174: 7495-7499.

93 Groth, A. C., and M. P. Calos. 2004. Phage integrases: biology and applications. J Mol Biol 335: 667-678.

94 McShan, W. M., Y.-F. Tang, and J. J. Ferretti. 1997. Bacteriophage T12 of Streptococcus pyogenes integrates into the gene for a serine tRNA. Mol Microbiol 23: 719-728.

95 Campbell, A., S. J. Schneider, and B. Song. 1992. Lambdoid phages as elements ofbacterial genomes. Genetica 86: 259-267.

96 McShan, W. M., and J. J. Ferretti. 1997. Genetic diversity in temperate bacterio-phages of Streptococcus pyogenes: identification of a second attachment site for phages carrying the erythrogenic toxin A gene. J Bacteriol 179: 6509-6511.

97 Yu, C.-E., and J. J. Ferretti. 1991. Molecular characterization of new group A streptococcal bacteriophages containing the gene for streptococcal erythrogenic toxin A (speA). Mol Gen Genet 231: 161-168.

98 Yu, C.-E., and J. J. Ferretti. 1989. Molecular epidemiologic analysis of the type A streptococcal exotoxin (erythrogenic toxin) gene (speA) in clinical Streptococcus pyogenes strains. Infect Immun 57: 3715-3719.

0 0

Post a comment