Chapter 7 : The Reemergence of Severe Group A Streptococcal Disease: an Evolutionary Perspective

MyBook is a cheap paperback edition of the original book and will be sold at uniform, low price.

Ebook: Choose a downloadable PDF or ePub file. Chapter is a downloadable PDF file. File must be downloaded within 48 hours of purchase

Buy this Chapter
Digital (?) $15.00

Preview this chapter:
Zoom in

The Reemergence of Severe Group A Streptococcal Disease: an Evolutionary Perspective, Page 1 of 2

| /docserver/preview/fulltext/10.1128/9781555816940/9781555811211_Chap07-1.gif /docserver/preview/fulltext/10.1128/9781555816940/9781555811211_Chap07-2.gif


This chapter reviews changes in the epidemiology of group A streptococcal infections and describes factors associated with the fitness and virulence of the pathogen. It also highlights the genetic diversity of group A streptococci, which, acted on by host factors, may account for periodic changes in disease severity. Childbed fever was one of the most frequent causes of death among postpartum women. A number of characteristics of group A streptococci may contribute to their fitness. These determinants can be categorized into three functional classes: adherence and colonization, invasion and replication, and avoidance of host defenses. Activation of the alternate complement pathway produces C5a, which is one of the primary mediators of chemotaxis in human tissue, attracting neutrophils to sites of infection. Horizontal gene transfer has resulted in -like genes and regulons with mosaic structures. Such an ability to recombine, in conjunction with strong selective pressures, can accelerate the evolution of functional diversity. The current increase in severe disease, particularly streptococcal toxic shock syndrome (STSS) and necrotizing fasciitis, is most likely related to changes in serotype distribution, production of toxins, and/or other factors. As immunity to these virulence factors increases, virulence will be lost. The author believes that this resurgence of more severe group A streptococcal disease does not represent the natural selection of a more virulent clone that will predominate but rather that as population immunity increases one will once again return to periods of waxing and waning of group A streptococcal disease severity.

Citation: Low D, McGeer A, Schwartz B. 1998. The Reemergence of Severe Group A Streptococcal Disease: an Evolutionary Perspective, p 93-123. In Scheld W, Armstrong D, Hughes J (ed), Emerging Infections 1. ASM Press, Washington, DC. doi: 10.1128/9781555816940.ch7

Key Concept Ranking

Tumor Necrosis Factor alpha
Restriction Fragment Length Polymorphism
Toxic Shock Syndrome
Highlighted Text: Show | Hide
Loading full text...

Full text loading...


Image of Figure 1.
Figure 1.

The •like genes of group A streptococci are located adjacent to each other at a chromosomal locus called the regulon, where they are flanked by a regulatory gene called and the gene. which encodes a C5a peptidase. (Reprinted from [123] with permission from M. A. Kehoe and Elsevier Science.)

Citation: Low D, McGeer A, Schwartz B. 1998. The Reemergence of Severe Group A Streptococcal Disease: an Evolutionary Perspective, p 93-123. In Scheld W, Armstrong D, Hughes J (ed), Emerging Infections 1. ASM Press, Washington, DC. doi: 10.1128/9781555816940.ch7
Permissions and Reprints Request Permissions
Download as Powerpoint
Image of Figure 2.
Figure 2.

Bridging of T cells and antigen-presenting cell s. A schematic model of a superantigen (SAg) interaction with a TCR and an MHC class II molecule is shown. APC. antigen-presenting cell; Ag. antigen. (Reprinted from Clinical Microbiology Reviews 125) with permission from M. Kotb and the American Society for Microbiology.)

Citation: Low D, McGeer A, Schwartz B. 1998. The Reemergence of Severe Group A Streptococcal Disease: an Evolutionary Perspective, p 93-123. In Scheld W, Armstrong D, Hughes J (ed), Emerging Infections 1. ASM Press, Washington, DC. doi: 10.1128/9781555816940.ch7
Permissions and Reprints Request Permissions
Download as Powerpoint
Image of Figure 3.
Figure 3.

Interplay between T-cell• and antigen-presenting cell (APC)-derived cytokines and induction of an inflammatory cytokine cascade by superantigens. IFN, interferon; Sag, Superantigen; TNF, tumor necrosis factor. (Reprinted (from ( ) with permission from M. Kotb and the American Society), for Microbiology.)

Citation: Low D, McGeer A, Schwartz B. 1998. The Reemergence of Severe Group A Streptococcal Disease: an Evolutionary Perspective, p 93-123. In Scheld W, Armstrong D, Hughes J (ed), Emerging Infections 1. ASM Press, Washington, DC. doi: 10.1128/9781555816940.ch7
Permissions and Reprints Request Permissions
Download as Powerpoint
Image of Figure 4.
Figure 4.

Reported cases of scarlet fever per 100.000 persons and scarlet fever case fatality rates in Oslo, Norway, from 1863 to 1878. (Reprinted from the [ ] with permission from R. M. Krause and the University of Chicago Press.)

Citation: Low D, McGeer A, Schwartz B. 1998. The Reemergence of Severe Group A Streptococcal Disease: an Evolutionary Perspective, p 93-123. In Scheld W, Armstrong D, Hughes J (ed), Emerging Infections 1. ASM Press, Washington, DC. doi: 10.1128/9781555816940.ch7
Permissions and Reprints Request Permissions
Download as Powerpoint
Image of Figure 5.
Figure 5.

Monthly incidences of group A streptococcal bacteremia in Sweden. 1987 10 1989. (Reprinted from the [ ] with permission from A. Strömberg and the University of Chicago Press.)

Citation: Low D, McGeer A, Schwartz B. 1998. The Reemergence of Severe Group A Streptococcal Disease: an Evolutionary Perspective, p 93-123. In Scheld W, Armstrong D, Hughes J (ed), Emerging Infections 1. ASM Press, Washington, DC. doi: 10.1128/9781555816940.ch7
Permissions and Reprints Request Permissions
Download as Powerpoint
Image of Figure 6.
Figure 6.

(A) Temporal distribution of New Zealand M type 1 and 12 isolates as percentages or all M typeable isolates by year. (B) Years in which each RFLP type was found among New Zealand isolates. (Reprinted from the [346] with permission from D. R. Martin and the University of Chicago Press.,

Citation: Low D, McGeer A, Schwartz B. 1998. The Reemergence of Severe Group A Streptococcal Disease: an Evolutionary Perspective, p 93-123. In Scheld W, Armstrong D, Hughes J (ed), Emerging Infections 1. ASM Press, Washington, DC. doi: 10.1128/9781555816940.ch7
Permissions and Reprints Request Permissions
Download as Powerpoint


1. Accardo, P.,, P. Sánchez-Corral,, O. Criado,, E. García,, and S. Rodriguez de Córdoba. 1996. Binding of human complement component C4b-binding protein (C4BP) to Streptococcus pyogenes involves the C4b-binding site. J. Immunol. 157:49354939.
2. Adams, E. M.,, S. Gudmundsson,, D. E. Yocum,, R. C. Haselby,, W. A. Craig,, and W. R. Suodstrom. 1985. Streptococcal myositis. Arch. Intern. Med. 145:10201023.
3. Åkesson, P.,, K. H. Schmidt,, J. Cooney,, and L. Björck. 1994. M1 protein and protein H: IgGFc- and albumin-binding streptococcal surface protein and encoded by adjacent gene. Biochem. J. 300:877886.
4. Åkesson, P.,, A. G. Sjoholm,, and L. Björck. 1996. Protein SIC, a novel extracellular protein of Streptococcus pyogenes interfering with complement function. J. Biol. Chem. 271:10811088.
5. Alkan, M.,, I. Ofek,, and E. H. Beachey. 1977. Adherence of pharyngeal and skin strains of group A streptococci to human skin and oral epithelial cells. Infect. Immun. 18:555557.
6. Anderson, R. M.,, and R. M. May. 1982. Coevolution of host and parasite. Parasitology 85:411426.
7. Anthony, B. F.,, E. L. Kaplan,, L. W. Wannamaker,, and S. S. Chapman. 1976. The dynamics of streptococcal infections in a defined population of children: serotypes associated with skin and respiratory infections. Am. J. Epidemiol. 104:652666.
8. Barrett, A. M.,, and G. A. Gresham. 1958. Acute streptococcal myositis. Lancet i:347351.
9. Barry, W.,, L. Hudgins,, S. T. Donta,, and E. L. Pesanti. 1992. Intravenous immunoglobulin therapy for toxic shock syndrome. JAMA 267:33153316.
10. Beachey, E. H.,, and H. S. Courtney. 1987. Bacterial adherence: the attachment of group a streptococci to mucosal surfaces. Rev. Infect. Dis. 9(Suppl.5):S475S481.
11. Beathard, A.,, and J. C. Guckian. 1967. Necrotizing fasciitis due to group A β-hemolytic streptococci. Arch. Intern. Med. 120:6367.
12. Ben Nasr, A.,, H. Herwald,, W. Muller-Ester),, and L. Björck. 1995. Human kininogens interact with M protein. a bacterial surface protein and virulence determinant. Biochem. J. 305:173180.
13. Berge, A.,, and L. Björck. 1995. Streptococcal cysteine proteinase releases biologically active fragments of streptococcal surface proteins. J. Biol. Chem. 270:98629867.
14. Berge, A.,, and U. Sjobring. 1993. PAM. a novel plasminogen-binding protein from Streptococcus pyogenes. J Biol. Chem. 268:2541725424.
15. Berkelman, R. L.,, D. Martin,, D. R. Graham,, J. Mowry,, R. Freisem,, J. A. Weber,, J. L. Ho,, and J. R. Allen. 1982. Streptococcal wound infections caused by a vaginal carrier. JAMA 247:26802682.
16. Bessen, D.,, and V. A. Fischetti. 1990. A human IgG receptor of group A streptococci is associated with tissue site of infection and streptococcal disease. J. Infect. Dis. 161:747754.
17. Bessen, D. E.,, and V. A. Fischetti. 1990. Differentiation between two biologically distinct classes of group A streptococci by limited substitutions of amino acids within the shared region of M protein-like molecules. J. Exp. Med. 172:17571764.
18. Bessen, D. E.,, and V. A. Fischetti. 1992. Nucleotide sequences of two adjacent M or M-like protein genes of group A streptococci: different RNA transcript levels and identification of a unique immunoglobulin A-binding protein. Infect. Immun. 60:124135.
19. Bessen, D. E.,, and S. K. Hollingshead. 1994. Allelic polymorphism of emm loci provides evidence for horizontal gene spread in group A streptococci. Proc. Natl. Acad. Sci. USA 91:32803284.
20. Bessen, D. E.,, C. M. Solir,, T. L. Readdy,, and S. K. Hollingshead. 1996. Genetic correlates of throat and skin isolates of group A streptococci. J. Infect. Dis. 173:896900.
21. Bhakdi, S.,, H. Bayley,, A. Valeva,, J. Walev,, B. Walker,, U. Weller,, M. Kehoe,, and M. Palmer. 1996. Staphylococcal alpha-toxin, streptolysin-O. and Escherichia coli hemolysin: prototypes of pore-forming bacterial cytolysins. Arch. Microbiol. 165:7379.
22. Bibler, M. R.,, and G. W. Rouan. 1986. Cryptogenic group A streptococcal bacteremia: experience at an urban general hospital and review of the literature. Rev. Infect. Dis. 8:941951.
23. Bisno, A. L.,, and D. L. Stevens. 1996. Streptococcal infections of skin and soft tissues. N. Engl. J. Med. 334:240245.
24. Brundage, J. F.,, J. D. Gunzenhauser,, J. N. Longfield,, M. V. Rubertone,, S. L. Ludwig,, F. A. Rubin, and Eo L. Kaplan. 1996. Epidemiology and control of acute respiratory disease with emphasis on group A beta-hemolytic streptococcus: a decade of U.S. army experience. Pediatrics 97(Suppl.):964970.
25. Bryant, A. E.,, M. A. Kehoe,, and D. L. Stevens. 1992. Streptococcal pyrogenic exotoxin A and streptolysin O enhance polymorphonuclear leukocyte binding to gelatin matrixes. J. Infect. Dis. 166:165169.
26. Caparon, M. G.,, and J. R. Scott. 1987. Identification of a gene that regulates expression of M protein, the major virulence determinant of group A streptococci. Proc. Natl. Acad. Sci. USA 84:86778681.
27. Caparon, M. G.,, D. L. Stevens,, A. Olsén,, and J. R. Scott. 1991. Role of M protein in adherence of group A streptococci. Infect. Immun. 59:18111817.
28. Carapctis, J.,, R. Robins•Browne,, D. Martin,, T. Shelby•James,, and G. Hogg. 1995. Increasing severity of invasive group A streptococcal disease in Australia: clinical and molecular epidemiological features and identification of a new virulent M-nontypeable clone. Clin. Infect. Dis. 21:12201227.
29. Cartwright, K.,, M. Logan,, C. McNulty,, S. Harrison,, R. George,, A. Efstratiou,, M. McEvoy,, and N. Begg. 1995. A cluster of cases of streptococcal necrotizing fasciitis in Gloucestershire. Epidemiol. Infect. 115:387397.
30.Centers for Disease Control. 1990. Group A beta-hemolytic streptococcal bacteremia-Colorado. Morbid. Mortal. Weekly Rep. 39:311.
31. Charles, D., and 8. Larsen. 1986. Streptococcal puerperal sepsis and obstetric infections: a historical perspective. Rev. Infect. Dis. 8:411422.
32. Chaussee, M. S.,, J. Liu,, D. L. Stevens,, and J. J. Ferretti. 1996. Genetic and phenotypic diversity among isolates of Streptococcus pyogenes from invasive infections. J. Infect. Dis. 173:901908.
33.Chelsom. J., A. Halstensen, T. Haga. and E. A. Hoiby. 1994. Necrotising fasciitis due to group A streptococci in western Norway: incidence and clinical features. Lancet 344:11111115.
34. Chen, C.,, N. Bormann. and P. P. Cleary. 1993. VirR and Mry are homologous trans-acting regulators of M protein and C5a peptidase expression in group A streptococci. Mol. Cell. Genet.241:685693.
35. Chen, C.,, and P. P. Cleary. 1990. Complete nucleotide sequence of the streptococcal C5a peptidase gene of Streptococcus pyogenes. J. Biol. Chem. 265:31613167.
36. Chmouryguina, I.,, A. Suvorov,, P. Ferrieri,, and P. P. Cleary. 1996. Conservation of the C5a peptidase genes in group A and B streptococci. Infect. Immun. 64:23872390.
37. Choi, Y.,, J . A. Lafferty,, and J. R. Clements. 1990. Selective expansion of T cells expressing Vβ2 in toxic shock syndrome. J. Exp. Med. 172:981984.
38. Claesson, B. E. B.,, and U. L.-E. Claesson. 1985. An outbreak of endometritis in a maternity unit caused by spread of group A streptococci from a showerhead. J. Hosp. Infect. 6:304311.
39. Claesson, B. E. B.,, N. G. Svensson,, L. Gotthardsson,, L. Gotthardsson,, and B. Garden. 1992. A foodborne outbreak of group A streptococcal disease at a birthday party. Scand. J. Infect. Dis. 24:577586.
40.. Cleary, P. P.,, E. L. Kaplan,, J. P. Handley,, A. Wlazlo,, M. H. Kim,, A. R. Hauser,, and P. M. Schlievert. 1992. Clonal basis for resurgence of serious Streptococcus pyogenes disease in the 1980s. Lancet 339:518521.
41. Cleary, P. P.,, D. LaPenta,, D. Heath,, E. J. Haanes,, and C. Chen,. 1991. A virulence regulon in Streptococcus pyogenes. p. 147151. In G. M. Dunny,. P. P. Cleary,. and L. L. McKay (ed.). Genetics and Molecular Biology of Streptococci, Lactococci. and Enterococci. American Society for Microbiology, Washington. D.C..
42. Cleary, P. P.,, J. Peterson,, C. Chen,, and C. Nelson. 1991. Virulent human strains of group G streptococci express a C5a peptidase enzyme similar to that produced by group A streptococci. Infect. Immun. 59:23052310.
43. Cleary, P. P.,, U. Prahbu,, J. B. Dale,, D. E. Wexler,, and J. Handley. 1992. Streptococcal C5a peptidase is a highly specific endopeptidase. Infect. Immun. 60:52195223.
44. Cockerill, F. R., III, K. L. MacDonald, R. L. Thompson, F. Roberson, P. C. Kohner, J. Besser• Wiek, J. M. Manahan, J. M. Musser, P. M. Schlievert, J. Talbot, B. Frankfort, J. M. Steckelberg, W. R. Wilson, M. T. Osterholm, and the Investigation Team. 1997. An outbreak of invasive group A streptococcal disease associated with high carriage rates of the invasive clone among school-aged children. JAMA 277:3843.
45. Colman, G.,, A. Tanna,, A. Efstratiou,, and E. Gaworzewska. 1993. The serotypes of Streptococcus pyogenes present in Britain during 1980-1990 and their association with disease. J. Med. Microbiol. 39: 165178.
46. Cone, L. A.,, D. R. Woodward,, P. M. Schlievert,, and G. S. Tomory. 1987. Clinical and bacteriologic observations of a toxic shock-like syndrome due to Streptococcus pyogenes. N. Engl. J. Med. 317:146149.
47. Connolly, T. J .,, D. J. Pavelka,, E. F. Lanspa,, and T. L. Connolly. 1989. Toxic shock-like syndrome associated with necrotizing Streptococcus pyogenes infection. Henry Ford Hasp. Med. J. 37:6972.
48. Cooper, D. A.,, and R. A. Joske. 1954. Acute streptococcal gangrene of the skin. Aust. N. Z. J. Surg. 23:268272.
49. Courtney, H. S.,, M. S. Bronze,, J. B. Dale,, and D. L. Hasty. 1994. Analysis of the role of M24 protein in group A streptococcal adhesion and colonization by use of ω-interposon mutagenesis. Infect. Immun. 62:48684873.
50. Courtney, H. S.,, J. B. Dale,, and D. L. Hasty. 1996. Differential effects of the streptococcal fibronectin-binding protein. FBP54, on adhesion of group A streptococci to human buccal cells and HEp•2 tissue culture cells. Infect. Immun. 64:24152419.
51. Courtney, H. S.,, D. L. Hasty,, J. B. Dale,, and T. P. Poirier. 1992. A 28-kilodalton fibronectin• binding protein of group A streptococci. Curr. Microbial. 25:245250.
52. Courtney, H. S.,, Y. Li,, J. B. Dale,, and D. L. Hasty. 1994. Cloning. sequencing, and expression of a fibronectin / fibrinogen-binding protein from group A streptococci. Infect. Immun. 62:39373946.
53. Cruickshank, J. G.,, R. J. C. Hart,, M. George,, and T. G. Feest. 1981. Fatal streptococcal septicaemia. Br. Med. J . 282:19441945.
54. Dale, J. B.,, R. G. Washburn,, M. B. Marques,, and M. R. Wessels. 1996. Hyaluronate capsule and surface M protein in resistance to opsonization of group A streptococci. Infect. Immun. 64:14951501.
55. Dan, M.,, S. Maximova,, Y. Siegman, Igra, R. Gulman, and H. H. Rolmensch. 1990. Varied presentations of sporadic group A streptococcal bacteremia: c1inical experience and attempt at classification. Rev. Infect. Dis. 12:537542.
56. Davies, H. D.,, A. McGeer. B. Schwartz, K. Green, D. Cann, A. E. Simor, D. E. Low, and the Ontario Group A Streptococcal Study Group. 1996. Invasive group A streptococcal infections in Ontario, Canada. N. Engl. J. Med. 335:547554.
57. Demers, B.,, B. Schwartz,, D. E. Low,, P. Wilson,, K. Fergoson, The Ontario Group A Streptococcal Study Project, and A. McGeer. 1993. Clinical features of streptococcal toxic shock syndrome in Ontario. Canada: a prospective study, abstr. 1396, p. 374. In Program and Abstracts of the 33rd Interscience Conference on Antimicrobial Agents and Chemotherapy. American Society for Microbiology, Washington, D.C..
58. Descamps, V.,, J. Aitken,, and M. G. Lee. 1994. Hippocrates on necrotising fasciitis. Lancet 344:556.
59. Dillon, H. C.,, C. W. Derrick,, and M. S. Dillon. 1974. M-antigens common to pyodenna and acute glomerulonephritis. J. Infect. Dis. 103:257267.
60. DiPersio, J. R.,, T. M. File, Jr.,, D. L. Stevens,, W. G. Gardner,, G. Petropoulos,, and K. Dinsa. 1996. Spread of serious disease-producing M3 clones of group A streptococcus among family members and health care workers. Clin. Infect. Dis. 22:490495.
61. Doebbeling, B. N.,, and R. P. Wenzel. 1989. Spontaneous streptococcal gangrenous myositis: survival with early debridement. South. Med. J. 32:900902.
62. Donaldson, P. M. W.,, B. Naylor,, J. W. Lowe,, and D. R. Gouldesbrough. 1993. Rapidly fatal necroti sing fasciitis caused by Streptococcus pyogenes. J. Clin. Pathol. 46:617620.
63. Duma, R. J.,, A. N. Weinberg,, T. F. Medrek,, and L. J . Kunz. 1969. Streptococcal infections: a bacteriologic and clinical study of streptococcal bacteremia. Medicine 48:87127.
64. Eichenbaum, Z.,, E. Muller,, S. A. Morse,, and J. R. Scott. 1996. Acquisition of iron from host proteins by the group A streptococcus. Infect. Immun. 64:54285429.
65. Einarsson, O. J .,, and M. Pers. 1986. Streptococcal gangrene of the eyelids. Scand. J. Plast. Reconstr. Surg. 20:331335.
66. Ellen, R. P.,, and R. J. Gibbons. 1972. M protein-associated adherence of Streptococcus pyogenes to epithelial surfaces: prerequisite for virulence. Infect. Immun. 5:826830.
67. Ewald, P. W. 1996. Guarding against the most dangerous emerging pathogens: insights from evolutionary biology. Emerg. Infect. Dis. 2:245256.
68. Falck, G.,, and J . Kjellander. 1992. Outbreak of group A streptococcal infection in a day-care center. Pediatr. Infect. Dis. J. 11:914919.
69. Farley, T. A.,, S. A. Wilson,, F. Mahoney,, K. Y. Kelso,, D. R. Johnson,, and E. L. Kaplan. 1993. Direct inoculation of food as the cause of an outbreak of group A streptococcal pharyngitis. J. Infect. Dis. 167:12321235.
70. Fedden, W. F. 1909. Six cases of acute infective gangrene of the extremities. Proc. R. Soc. Med. 2(Pt.1 ):213.
71. Ferrieri, P.,, A. S. Dajani,, L. W. Wannamaker,, and S. S. Chapman. 1972. Natural history of impetigo. I. Site sequence of acquisition and familial patterns of spread of cutaneous streptococci. J. Clin. Invest. 51:28512862.
72. Fischetti, V. A. 1989. Streptococcal M protein: design and biological behavior. Clin. Microbial. Rev. 2:285314.
73. Fischetti, V. A. 1991. Streptococcal M protein. Sci. Am. 264:5865.
74. Fischetti, V. A.,, W. M. Hndges,, and D. E. Hurby. 1990. Protection against streptococcal pharyngeal colonization with a vaccinia:M protein recombinant. Science 244:14871490.
75. Forni, A. L.,, E. Kaplan,, P. M. Schlievert,, and R. S. Roberts. 1995. Clinical and microbiological characteristics of severe group A streptococcus infection and streptococcal toxic shock syndrome. Clin. Infect. Dis. 21:333340.
76. Francis, J.,, and R. E. Warren. 1988. Streptococcus pyogenes bacteraemia in Cambridge-a review of 67 episodes. Q. J. Med. 68:603613.
77. Frick, I. M.,, P. Akesson,, J. Cooney,, U. Sjobring,, K. H. Schmidt,, H. Gomi,, S. Haltori,, C. Tagawa,, F. Kishimoto,, and L. Björck. 1994. Protein H—a surface protein of Streptococcus pyagenes with separate binding sites for IgG and albumin. Mol. Microbial. 12:143151.
78. Gastanaduy, A. S.,, E. L. Kaplan,, B. B. Huwe,, C. McKay,, and L. W. Wannamaker. 1980. Failure of penicillin to eradicate group A streptococci during an outbreak of pharyngitis. Lancet ii:498502.
79. Gaworzewska, E.,, and G. Colman. 1988. Changes in the pattern of infection caused by Streptococcus pyogenes. Epidemial. Infect. 100:257269.
80. Gerlach, D.,, H. Knoll,, W. Kohler,, J. H. Ozegowski,, and V. Hribalova. 1983. Isolation and characterization of erythrogenic toxins. V. Identity of erythrogenic toxin type B and streptococcal proteinase precursor. Zentralbl. Bakteriol. Mikrabiol. Hyg. Abt. I Orig. A 255:221233.
81. Gibson, C. M.,, and M. G. Caparon. 1996. Insertional inactivation of Streptococcus pyogenes sod suggests that prtF is regulated in response to a superoxide signal. J. Bacteriol. 178:46884695.
82. Giuliano, A.,, F. Lewis, Jr., K. Hadley, and F. W. Blaisdell. 1977. Bacteriology of necrotizing fasciitis. Am. J. Surg. 134:5257.
83. Goepel, J. R.,, D. G. Richards,, D. M. Harris,, and L. Henry. 1980. Fulminant Streptococcus pyogenes infection. Br. Med. J. 281:1412.
84. Gonzales-Ruiz, A.,, G. L. Ridgway,, S. L. Cohen,, C. P. Hunt,, G. McGrouther,, and M. Adiseshiah. 1995. Varicella gangrenosa with toxic shock-like syndrome due to group A streptococcus infection in an adult: case repon . Clin. Infect. Dis. 20:10581060.
85. Gordon, G.,, B. A. S. Dale,, and D. Lochhead. 1994. An outbreak of group A haemolytic streptococcal puerperal sepsis spread by the communal use of bidets. Br. J. Obstet. Gynaecol. 101:447448.
86. Gordon, J.E.,, and G. F. Badger. 1934. The isolation time of scarlet fever. Am. J. Public Health 24:438.
87. Gray, B. M., 1991. Streptococcal infections, p. 639673. In A. S. Evans, and P. S. Brachman (ed.), Bacteria/Infections of Humans: Epidemiology and Control. Plenum Medical Book Company, New York, N.Y..
88. Gray, C. G.,, J. Escamilla,, K. C. Hyams,, J. P. Struewing,, E. L. Kaplan,, and A. K. Tupponce. 1991. Hyperendemic Streptococcus pyogenes infection despite prophylaxis with penicillin G benzathine. N. Engl. J. Med. 325:9297.
89. Haanes, E. J.,, and P. P. Cleary. 1989. Identification of a divergent M protein gene and an M protein-related gene family in Streptococcus pyogenes serotype 49. J. Bacteriol. 171:63976408.
90. Haanes, E. J.,, D. G. Heath,, and P. P. Cleary. 1992. Architecture of the vir regulons of group A streptococci parallels opacity factor phenotype and M protein class. J. Bacteriol. 174:49674976 .
91. Hable, K. A.,, C. Horstbeirer,, A. D. Wold,, and J. A. Washington. 1973. Group A β-hemolytic streptococcemia: bacteriologic and clinical study of 44 cases. Mayo Clin. Proc. 48:336339.
92. Hanski, E.,, and M. G. Caparon. 1992. Protein F. a fibronectin binding protein is an adhesin of group A streptococci. Proc. Natl. Acad. Sci. USA 89:61726176.
93. Hanski, E.,, J. Jaffe,, and V. Ozeri. 1996. Proteins F1 and F2 of Streptococcus pyogenes. Properties of fibronectin binding. Adv. Exp. Med. Biol. 408:141150.
94. Harbaugh, M. P.,, A. Podbielski,, S. Hugl,, and P. P. Cleary. 1993. Nucleotide substitutions and small-scale insertion produce size and antigenic variation in group A streptococcal M1 protein. Mol. Microbiol. 8:981991.
95. Hasty, D. L.,, I. Ofek,, H. S. Courtney,, and R. J. Doyle. 1992. Multiple adhesins of streptococci. Infect. Immun. 60:21472152.
96. Heggie, A. D.,, M. R. Jacobs,, P. E. Linz,, D. P. Han,, E. L. Kaplan,, and B. Boxerbaum. 1992. Prevalence and characteristics of pharyngeal group A β-hemolytic streplococci in US navy recruits receiving benzathine penicillin prophylaxis. J. Infect. Dis. 166:10061013.
97. Herwald, H.,, M. Collin,, W. Muller-Esterl, arid L. Björck. 1996. Streptococcal cysteine proteinase releases kinins: a novel virulence mechanism. J. Exp. Med. 184:665673.
98. Hoge, C. W.,, B. Schwartz,, D. F. Talkington,, R. F. Breiman,, E. M. MacNeill,, and S. J. Englender. 1993. The changing epidemiology of invasive group A streptococcal infections and the emergence of streptococcal toxic shock-like syndrome. JAMA 269:585589.
99. Hollingshead, S. K.,, T. L. Readdy,, D. L. Yung,, and D. E. Bessen. 1993. Structural heterogeneity of the emm gene cluster in group A streptococci. Mol. Microbiol. 8:707717.
100. Hollingshead, S. K.,, J. W. Simecka,, and S. M. Michalek. 1993. Role of M protein in pharyngeal colonization by group A streptococci in rats. Infect. Immun. 6:22772283.
101. Holm, S. E.,, A. Norrby,, A. M. Bergholm,, and M. Norgren. 1992. Aspects of pathogenesis of serious group A streptococcal infections in Sweden. J. Infect. Dis. 166:3137.
102. Horstmann, R. D.,, H. J. Sievertsen,, J. Knobloch,, and V. A. Fischetti. 1988. Antiphagocytic activity of streptococcal M protein: selective binding of complement control protein factor H. Proc. Natl. Acad. Sci. USA 85:16571661.
103. Hribalova, V. 1988. Streptococcus pyogenes and the toxic shock syndrome. Ann. Intern. Med. 108:772. (Letter).
104. Husmann, L. K.,, D.-L. Yung,, S. K. Hollingshead,, and J. R. Scott. 1997. Role of putative virulence factors of Streptococcus pyogenes in mouse models of long-term throat colonization and pneumonia. Infect. Immun. 65:14221430.
105. Hynes, R. 0., and K. M. Yamada. 1982. Fibronectins: multifunctional modular glycoproteins. J. Cell Biol. 95:369377.
106. Ichiyama, S.,, K. Nakashima,, K. Shimokata,, M. Ohta,, Y. Shimizu,, K. Ooe,, H. Igarashi,, and T. Murai. 1997. Transmission of Sireptococcus pyogenes causing toxic shock-like syndrome among family members and confirmation by DNA macrorestriction analysis. J. Infect. Dis. 175:723726.
107. Ispahani, P.,, F. E. Donald,, and A. J. D. Aveline. 1988. Streptococcus pyogenes bacteraemia: an old enemy subdued. but not defeated. J. Infect. 16:3746.
108. Jaffe, J.,, S. Natanson-Yaron,, M. G. Caparon,, and E. Hanski. 1996. Protein F2. a novel fibro nectin-binding protein from Streptococcus pyogenes, possesses two binding domains. Mol. Microbiol. 21:373384.
109. James, W. E. S.,, G. F. Badger,, and J. H. Dingle. 1960. A study of illness in a group of Cleveland families. XIX. The epidemiology of the acquisition of group A streptococci and of associated illnesses. N. Engl. J. Med. 262:687694.
110. Ji, Y.,, L. MeLandsborough,, A. Kondagunta,, and P. P. Cleary. 1996. C5a peptidase alters clearance and trafficking of group A streptococci by infected mice. Infect. Immun. 64:503510.
111. . Johnson, D. R.,, D. L. Stevens,, and E. L. Kaplan. 1992. Epidemiologic analysis of group A streptococcal serotypes associated with severe systemic infections, rheumatic fever, or uncomplicated pharyngitis. J. Infect. Dis. 166:374382.
112. Kaplan, E. 1993. Global assessment of rheumatic fever and rheumatic heart disease at the close of the century. The influences and dynamics of population and pathogens: a failure to realize prevention? (The T. Duckett Jones Memorial Lecture.) Circulation 88: 19641972.
113. Kaplan, E. L. 1996. Recent epidemiology of group A streptococcal infections in North America and abroad: an overview. Pediatrics 97:945948.
114. Kaplan, E. L.,, D. R. Johnson,, and P. P. Cleary. 1989. Group A streptococcal serotypes isolated from patients and sibling contacts during the resurgence of rheumatic fever in the United States in the mid-1980s. J. Infect. Dis. 159:101103.
115. Kappler, J.,, B. Kotzin,, L. Herron,, E. W. Gelfand,, R. D. Bigler,, A. Boylston,, S. Carrel,, D. N. Posnelt,, Y. Choi, aud P. Marraek. 1989. VII-specific stimulation of human T cells by staphylococcal toxins. Science 244:811813.
116. Kapur, V.,, S. Kanjilal,, M. R. Hamrick,, L.-L. Li,, T. S. Whittam,, S. A. Sawyer,, and J. M. Musser. 1995. Molecular population genetic analysis of the streptokinase gene of Streptococcus pyogenes: mosaic alleles generated by recombination. Mol. Microbiol. 16:509519.
117. Kapur, V.,, J. T. Maffei,, R. S. Greer,, L. L. Li,, G. J. Adams,, and J. M. Musser. 1994. Vaccination with streptococcal extracellular cysteine protease (interleukin-1β convertase) protects mice against challenge with heterologous group A streptococci. Microb. Pathog. 16:443450.
118. Kapur, V.,, M. W. Majesky,, L. L. Li,, R. A. Black,, and J. M. Musser. 1993. C1eavagc of interleukin I J.1 (IL-1βprecursor to produce active IL-1β by a conserved extracellular cysteine protease from Streptococcus pyogenes. Proc. Natl. Acad. Sci. USA 90:76767680.
119. Kapur, V.,, S. Topouzis,, M. W. Majesky,, L. L. Li,, M. R. Hamrick,, R. J. Hamill,, J. M. Patti,, and J. M. Musser. 1993. A conserved Streptococcus pyogenes extracellular cysteine protease cleaves human fibronectin and degrades vitronectin. Microb. Pathog. 15:327346.
120. Katz, A. R.t and D. M. Morens. 1992. Severe streptococcal infections in historical perspective. Clin. Infect. Dis. 14:298307.
121. Kaul, R.,, A. McGeer,, D. E. Low,, K. Green,, B. Schwartz, Ontario Group A Streptococcal Study, and A. E. Simor. Population-based surveillance for group A streptococcal necrotizing fasciitis: clinical feature, prognostic indicators and microbiologic analysis of 77 cases. Am. J. Med , in press.
122. Keefer, C. S.,, F. J. Inglefinger,, and W. W. Spink. 1937. Significance of hemolytic streptococci bacteraemia: a study of two hundred and forty six patients. Arch. Intern. Med. 60:10841097.
123. Kehoe, M. A.,, V. Kapur,, A. M. Whatmore,, and J. M. Musser. 1996. Horizontal gene transfer among group A streptococci: implications for pathogenesis and epidemiology. Trends Microbiol. 4:436443.
124. Kihlberg, B. M.,, J . Cooney,, M. G. Cap. ron, A. Olsen, and L. Björck. 1995. Biological properties of a Streptococcus pyogenes mutant generated by Tn916 insertion in mga. Microb. Pathog. 19:299315.
125. Kotb, M. 1995. Bacterial pyrogenic exotoxins as superantigens. Clin. Microbiol. Rev. 8:411426.
126. Krause, R. M. 1993. Dynamic of emergence. J. Infect. Dis. 170:265271.
127. Kreikemeyer, B.,, S. R. Talay,, and G. S. Chhatwal. 1995. Characterization of a novel fibronectin-binding sulface protein in group A streptococci. Microb. Pathog. 19:299315.
128. Kuttner, A. G.,, and E. Krumwiede. 1994. Observations on the epidemiology of streptococcal pharyngitis and the relation of streptococcal carriers to the occurrence of outbreaks. J. Clin. Invest. 23:139.
129. Kuusela, P.,, M. Ullberg,, O. Saksela,, and G. Kronvall. 1992. Tissue-type plasminogen activator-mediated activation of plasminogen on the surface of group A, C. and G streptococci. Infect. Immun. 60:196201.
130. Lamothe, F.,, P. D'Amico,, P. Ghosn,, C. Tremblay,, J. Braldy,, and J. V. Patenaude. 1995. Clinical usefulness of intravenous human immunoglobulins in invasive group A streptococcal infections: case repon and review. Clin. Infect. Dis. 21:14691470.
131. Lancefield, R. C. 1962. Current knowledge of the type specific M antigens of group A streptococci. J. Immunol. 89:307313.
132. Lannigan, R.,, Z. Hussain,, and T. W. Austin. 1985. Streptococcus pyogenes as a cause of nosocomial infection in a critical care unit. Diagn. Microbial. Infect. Dis. 3:337341.
133. LaPenta, D.,, C. Rubens,, E. Chi,, and P. P. Cleary. 1994. Group A streptococci efficiently invade human respiratory epithelial cells. Proc. Natl. Acad. Sci. USA 91:1211512119.
134. Lee, P. K.,, and P. M. Schlievert. 1991 . Molecular genetics of pyrogenic exotoxin "superantigens" of group A streptococci and Staphylococcus aureus. Curr. Top. Microbiol. Immunol. 174:119.
135. Lenski, R. E.,, and R. M. May. 1994. The evolution of virulence in parasites and pathogens: reconciliation between two competing hypotheses. J. Theor. Biol. 169:253265.
136. Levin, B. R. 1996. The evolution and maintenance of virulence in microparasites. Emerg. Infect. Dis. 2:93102.
137. Levin, B. R.,, and C. S. Eden. 1990. Selection and evolution of virulence in bacteria: an ecumenical excursion and modest suggestion. Parasitology l00:S103S115.
138. Levin, S.,, and D. Pimentel. 1981. Selection of intermediate rates of increase in parasite host systems. Am. Nat. 117:308315.
139. Llpsltch, M.,, and E. R. Moxon. 1997. Virulence and transmissibility of pathogens: what is the relationship? Trends Microbiol. 5:3137.
140. Lossos, I. S.,, I. Felsenstein,, R. Breuer,, and D. Engelhard. 1992. Food-borne outbreak of group A β-hemolytic streptococcal pharyngitis. Arch. Intern. Med. 152:853855.
141. Lottenberg, R. 1996. Contact activation proteins and the bacterial surface. Trends Microbiol. 4:413415.
142. Lottenberg, R.,, L. E. Desjardin,, H. Wang,, and M. D. P. Boyle. 1992. Streptokinase-producing streptococci grown in human plasma acquire unregulated cell-associated plasmin activity. J. Infect. Dis. 166:436440.
143. Lottenberg, R.,, D. Minning-Wenz,, and M. D. P. Boyle. 1994. Capturing host plasmin(ogen): a common mechanism for invasive pathogens? Trends Microbial. 2:2024.
144. Loudon, I. 1994. Necrotising fasciitis. hospital gangrene. and phagedena. Lancet 344:1416.
145. Maddox, J. S.,, J. C. Ware,, and H. C. Dillon, Jr. 1985. The natural history of streptococcal skin infection: prevention with topical antibiotics. J. Am. Acad. Dermatol. 13:207212.
146. Martin, D. R.,, and L. A. Single. 1993. Molecular epidemiology of group A streptococcus M type 1 infections. J. Infect. Dis. 167:11121117.
147. Martin, P. R.,, and E. A. Høiby. 1990. Streptococcal serogroup A epidemic in Norway 1987-1988. Scand. J. Infect. Dis. 22:421429.
148. Mastro, T. D.,, T. A. Farley,, J. A. Elliott,, R. R. Facklam,, J. R. Perks,, J. L. Hadler,, R. C. Gond,, and J. S. Spika. 1990. An outbreak of surgical-wound infections due to group A streptococcus canied on the scalp. N. Engl. J. Med. 323:968972.
149. Mattingly, P. C.,, and A. G. Mowat. 1981. Streptococcal pyomyositis following a sore throat. Rheumatol. Rehabil. 20:151152.
150. Mcintyre, D. M. 1968. An epidemic of Streptococcus pyogenes puerperal and post-operative sepsis with an unusual carrier site: the anus. Am. J. Obstet. Gynecol. 101:308314.
151. McIver, K.,, A. S. Heath,, and J. R. Scott. 1995. Regulation of virulence by environmental signals in group A streptococci: influence of osmolarity, temperature, gas exchange. and iron limitation on emm transcription. Infect. Immun. 63:45404542.
152. McKee, W. M.,, J. M. DiCaprio,, C. E. Roberts, Jr, and J. C. Sherris. 1966. Anal carriage as the probable source of a streptococcal epidemic. Lancet ii:l0071009.
153. McLandsborough, L. A.,, and P. P. Cleary. 1995. Insertional inactivation of virR in Streptococcus pyogenes M49 desmonstrates that VirR functions as a positive regulator of ScpA, FcRA. OF, and M protein. FEMS Microbiol. Lett. 128:4552.
154. McLean, T. R.,, D. M. Musher,, and R. J. Hamill. 1991. Streptococcal myonecrosis due to Streptococcus pyogenes. Ther. Challenges 2:1722.
155. Meleney, F. L. 1924. Hemolytic Streptococcus gangrene. Arch. Surg. 9:317364.
156. Meleney, F. L. 1933. A differential diagnosis between certain types of infectious gangrene of the skin: with particular reference to haemolytic Streptococcus gangrene and bacterial synergistic gangrene. Surg. Gynecol. Obstet. 56:847867.
157. Memlsh, Z. A.,, D. Gravel•Tropper,, C. Oxley,, B. Toye,, and G. E. Garber. 1994. Group A streptococcal endometritis: report of an outbreak. and review of the literature. Can. J. Infect. Dis. 5:276281.
158. Miethke, T.,, K. Duschek,, C. Wahl,, K. Heeg,, and H. Wagner. 1993. Pathogenesis of the toxic shock syndrome: T cell mediated lethal shock caused by the superantigen TSST-1. Eur. J. Immunol. 23:14941500.
159. Miles, L. T.,, J. B. Jacobs,, P. D. Gittelman,, and A. S. Lebowitz. 1992. Streptococcal gangrene of the head and neck: a case report and review of the literature. Head Neck 14:143147.
160. Miller, C. W.,, J. F. Prescott,, K. A. Mathews,, S. D. Betschel,, J. A. Yager,, V. Guru,, and D. E. Low. 1996. Streptococcal toxic shock syndrome in dogs. J. Am. Vet. Med. Assoc. 209:14211426.
161. Molinari, G.,, S. R. Talay,, P. Valentin-Weigand,, M. Rohde,, and G. S. Chhatwal. 1997. The fibronectin-binding protein of Streptococcus pyogenes, SfbI. is involved in the internalization of group A streptococci by epithelial cells. Infect. Immun. 65:13571363.
162. Mollick, J. A.,, G. G. Miller,, J. M. Musser,, R. G. Cook,, D. Grossman,, and R. R. Rich. 1993. A novel superantigen isolated from pathogenic strains of Streptococcus pyogenes with aminoterminal homology to staphylococcal enterotoxins B and C. J. Clin. Invest. 92:710719.
163. Moore, D. L.,, G. Delage,, H. Labelle,, and M. Gauthier. 1986. Preacute streptococcal pyomyositis: report of two cases and review of the literature. J. Pediatr. Orthop. 6:232235.
164. Morse, S. S. 1990. Regulating viral traffic. Issues Sci. Technol. 7:8184.
165. Morse, S. S. 1991. Emerging viruses: defining the rules for viral traffic. Perspect. Biol. Med. 34:387409.
166. Morse, S. S., 1993. Examining the origins of emerging viruses. p. 10-28. In S. S. Morse (ed.). Emerging Viruses. Oxford University Press, New York, N.Y..
167. Morse, S. S.,, and A. Schluederberg. 1990. Emerging viruses: the evolution of viruses and viral diseases. J. Infect. Dis. 162:17.
168. Moses, A. E.,, A. Ziv,, M. Harari,, G. Rahav,, M. Shapiro,, and D. Engelhard. 1995. Increased incidence and severity of Streptococcus pyogenes bacteremia in young children. Pediatr. Infect. Dis. J. 14:767770.
169. Müller-Alouf, H.,, J. E. Aloof,, D. Gerlach,, J . •H. Ozegowski, C. Fitting, and J.•M. Cavaillon. 1996. Human pro- and anti-inflammatory cytokine patterns induced by Streptococcus pyogenes erythrogenic (pyrogenic) exotoxin A and C superantigens. Infect. Immun. 64:14501453.
170. 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 88:26682672.
171. Musser, J. M.,, V. Kapur,, J. Szeto,, X. Pan,, D. S. Swanson,, and D. R. Martin. 1995. Genetic diversity and relationships among Streptococcus pyogenes strains expressing serotype M1 protein: recent intercontinental spread of a subclone causing episodes of invasive disease. Infect. Immun. 63:9941003.
172. Nadal, D.,, R. P. Lauener,, and C. P. Braegger. 1993. T cell activation and cytokine release in streptococcal toxic shock• like syndrome. J. Pediatr. 122:727729.
173. Navarro, V. J.,, P. I. Axelrod,, W. Pinover,, H. S. Hockfield,, and J. R. Kostman. 1993. A com• parison of Streptococcus pyogenes (group A streptococcal) bacteremia at an urban and a suburban hospital. Arch. Intern. Med. 153:26792684.
174. Nemoto, E.,, H. Rikiishi,, S. Sugawara,, S. Okamoto,, K. Tamura,, Y. Maruyama,, and K. Kumagai. 1996. Isolation of a new superantigen with potent mitogenic activity to murine T cells from Streptococcus pyogenes. FEMS Immunol. Med. Microbiol. 15:8191.
175. Norgren, M. 1992. Genetic diversity in TIMI group A streptococci in relation to clinical outcome of in fect ion. J. Infect. Dis. 166:10141020.
176. Norrby-Teglund, A.,, R. Kaul,, and D. E. Low. 1996. Plasma from patients with severe invasive group A streptococcal infections treated with nonnal polyspecific IgG inhibits streptococcal-superantigen-induced T cell prolife ration and cytokine production. J. Immunol. 156:30573064.
177. Norrby-Teglund, A.,, R. Kaul,, D. E. Low,, A. McGeer,, J. Andersson,, U. Andersson,, and M. Kolb. 1996. Evidence for the presence of streptococcal-superantigen-neutralizing antibodies in normal polyspecific immunoglobulin G. Infect. Immun. 64:53955398.
178. Norrby-Teglund, A.,, D. Newlon,, M. Kolb,, S. E. Holm,, and M. Norgren. 1994. Superantigenic propenies of the group A streptococcal exotoxin SpeF (MF). Infect. Immun. 62:52275233.
179. Norrby-Teglund, A.,, M. Norgren,, S. E. Holm,, U. Andersson,, and J. Andersson. 1994. Similar cytokine induction profiles of a novel streptococcal mitogenic factor, MF, and pyrogenic exotoxins A and B. Infect. Immun. 62:37313738.
180. Okada, N.,, R. T. Geist,, and M. G. Caparon. 1993. Positive transcriptional control of mry regulates virulence in the group A streptococcus. Mol. Microbiol. 7:893903.
181. Okada, N.,, M. Liszewski,, J. Atkinson,, and M. Caparon. 1995. Membrane cofactor protein (CD46) is a keratinocyte receptor for the M protein of the group A streptococcus. Proc. Natl. Acad. Sci. USA 92:24892493.
182. Okada, N.,, A. P. Pentland,, P. Falk,, and M. G. Caparon. 1994. M protein and protein F act as important determinants of cell-specific tropism of Streptotoccus pyogenes in skin tissue. J. Clin. Invest. 94:965977.
183. O'Toole, P. 1992. Two major classes in [he M protein family in group A streptococci. Proc. Natl. Acad. Sci. USA 89:86618665.
184. Overholt, E. M.,, P. W. Flint,, E. L. Overholt,, and C. S. Murakami. 1992. Necrotizing fasciitis of the eyelids. Otolaryngol. Head Neck Surg. 106:339343.
185. Paine, T. F., Jr.,, R. P. Novick,, and W. H. Hall, Jr. 1963. Fatal gangrene caused by Streptococcus pyogenes. Arch. Intern. Med. 112:936939.
186. Pancholi, V.,, and V. A. Fischetti. 1992. A major surface protein on group A streptococci is a glyceraldehyde-3-phosphate-dehydrogenase with multiple binding act ivity. J. Exp. Med. 176:415426.
187. Perez, C. M.,, B. M. Kubak,, H. G. Cryer,, S. Salehmugodam,, P. Vespa,, and D. Farmer. 1997. Adjunctive treatment of streptococcal toxic shock syndrome using intravenous immunoglobulin: case report and review. Am. J. Med. 102:111112.
188. Perez-Casal, J.,, M. G. Caparon,, and J. R. Scott. 1991. Mry. a trans-acting positive regulator of the M protein gene of Streptococcus pyogenes with similarity to the receptor proteins of two-component regulatory sytems. J. Bacteriol. 173:26172624.
189. Perez-Casal, J.,, N. Okada,, M. G. Caparon,, and J. R. Scott. 1995. Role of the conserved C-repeat region of the M protein of Streptococcus pyogenes. Mol. Microbiol. 15:907916.
190. Podbielski, A. 1993. Three different types of organization of the vir regulon in group A streptococci. Mol. Gen. Genet. 237:287300.
191. Podbielski, A.,, A. Flosdorff,, and J. Weber-Heynemann. 1995. The group A streptococcal virR49 gene controls expression of four structural vir regulon genes. Infect. Immun. 63:920.
192. Podbielski, A.,, J. Hawlitzky,, T. D. Pack,, A. Flosdorff,, and M. D. Boyle. 1994. A group A streptococcal Enn protein potentially resulting from intergenomic recombination exhibits atypical immunoglobulin-binding characteristics. Mol. Microbiol. 12:725736.
193. Podbielski, A.,, J. A. Peterson,, and P. Cleary. 1992. Surface protein-CAT reporter fusions demonstrate differential gene express ion in the vir regulon of Streptococcus pyogenes. Mol. Microbial. 6:22532265.
194. Podbielski, A.,, N. Schnitzler,, P. Beyhs,, and M. D. Boyle. 1996. M-related protein (MrP) contributes to group A streptococcal resistance to phagocytosis by human granulocytes. Mol. Microbiol. 19:429441.
195. Podbielski, A.,, M. Woischnik,, B. Pohl,, and K. H. Schmidt. 1996. What is the size of the group A streptococcal vir regulon? The Mga regulator affects expression of secreted and surface virulence factors. Med. Microbial. Immun. 185:171181.
196. Poirier, T. P.,, M. A. Kehoe,, E. Whitnack,, M. E. Dockter,, and E. H. Beachey. 1989. Fibrinogen binding and resistance to phagocytosis of Streptococcus sanguis expressing cloned M protein of Streptococcus pyogenes. Infect. Immun. 57:2935.
197. Quinn, R. W. 1982. Epidemiology of group A streptococcal infections-their changing frequency and severity. Yale J. Biol. Med. 55:265270.
198. Quinn, R. W., 1982. Streptococcal infections. p. 525552. In A. S. Evans, and H. A. Feldman (ed.), Bacterial Infections of Humans. Plenum Medical. New York, N.V..
199. Quinn, R. W. 1989. Comprehensive review of morbidity and mortality trends for rheumatic fever, streptococcal disease, and scarlet fever: the decline of rheumatic fever. Rev. Infect. Dis. 11:928953.
200. Raeder, R.,, and M. D. Boyle. 1993. Association of type II immunoglobulin G-binding protein expression and survival of group A streptococci in human blood. Infect. Immun. 61:36963702.
201. Raeder, R.,, and M. D. Boyle. 1993. Association between expression of immunoglobulin G-binding proteins by group A streptococci and virulence in a mouse skin infection model. Infect. Immun. 61:13781384.
202. Rakonjac, J. V.,, J. C. Robbins,, and V. A. Fischetti. 1995. DNA sequence of the serum opacity factor of group A streptococci: identification of a fibronectin-binding repeat domain. Infect. Immun. 63:622631.
203. Rammelkamp, C. H., Jr.,, E. A. Mortimer, Jr.,, and E. Wolinsky. 1964. Transmission of streptococcal and staphylococcal infections. Ann. Intern. Med. 60:753758.
204. Read, A. F. 1994. The evolution of virulence. Trends Microbiol. 2:7376.
205. Reitmeyer, J. c.,, A. Ewert,, M. A. Crawford,, G. R. Reitmeyer,, and L. Mock. 1993. Survival of group A streptococci in dried human blood. J. Med. Microbiol. 38:6163.
206. Richman, D. D.,, S. J. Breton,, and D. A. Goldmann. 1977. Scarlet fever and group A streptococcal surgical wound infection traced to an anal carrier. J. Pediatr. 90:387390.
207. Rye, M., K H. Beachey, and E. Whitnaek. 1989. Ultrastructural localization of the fibrinogen-binding domain of streptococcal M protein. Infect. Immun. 57:23972404.
208. Schaffner, W.,, L. B. Lefkowitz, Jr.,, J. S. Goodman,, and M. G. Koenig. 1969. Hospital outbreak of infections with group A streptococci traced to an asymptomatic anal carrier. N. Engl. J. Med. 280:12241225.
209. Schlievert, P. M. 1995. The role of superantigens in human disease. Curr. Opin. Infect. Dis. 8:170174.
210. Schlievert, P. M.,, A. P. Assimacopoulos,, and P. P. Cleary. 1996. Severe invasive group A streptococcal disease: clinical description and mechanisms of pathogenesis. J. Lab. Clin. Med. 127:1322.
211. Schmidt, K. H.,, K. Mann,, J. Cooney,, and W. Kohler. 1993. Multiple binding of type 3 streptococcal M protein to human fibrinogen, albumin and fibronectin. FEMS Immunol. Med. Microbiol. 7:135144.
212. Schnitzler, N.,, A. Podbielski,, G. Baumgarten,, M. Mignon,, and A. Kaufhold. 1995. M or M-like protein gene polymorphisms in human group G streptococci. J. Clin. Microbiol. 33:356363.
213. Schwartz, B.,, J. A. Elliott,, J . C. Butler,, P. A. Simon,, B. L. Jameson,, G. E. Welch,, and R. R. Facklam. 1992. Clusters of invasive group A streptococcal infections in family, hospital. and nursing home settings. Clin. Infect. Dis. 15:277284.
214. Schwartz, B.,, R. R. Facklam,, and R. F. Breiman. 1990. Changing epidemiology of group A streptococcal infection in the USA. Lancet 336:11671171.
215. Sela, S.,, A. Aviv,, A. Tovi,, I. Burstein,, M. G. Caparon,, and E. Hanski. 1993. An adhesin of Streptococcus pyogenes binds fibronectin via two distinct domains. Mol. Microbiol. 10:10491055.
216. Shanley, T. P.,, D. Schrier,, V. Kapur,, M. Kehoe,, J. M. Musser,, and P. A. Ward. 1996. Streptococcal cysteine protease augments lung injury induced by products of group A streptococci. Infect. Immun. 64:870877.
217. Sharma, A. K.,, and M. K. Pangburn. 1997. Localization by site-directed mutagenesis of the site in human complement factor H that binds to Streptococcus pyogenes M protein. Infect. Immun. 65:484487.
218. Simpson, W. J.,, J. M. Musser,, and P. P. Cleary. 1992. Evidence consistent with horizontal transfer of the gene (emm12) encoding serotype M12 protein between group A and group G pathogenic streptococci. Infect. Immun. 60:18901893.
219. Single, L. A.,, and D. R. Marlin. 1992. Clonal differences within M-types of the group A streptococcus revealed by pulsed field gel electrophoresis. FEMS Microbiol. Lett. 91:8590.
220. Sirinavian, S.,, and G. H. McCracken, Jr. 1979. Primary suppurative myositis in children. Am. J. Dis. Child. 133:263265.
221. Soares, S.,, K. G. Kristinsson,, J. M. Musser,, and A. Tomasz. 1993. Evidence for the introduction of a multiresistant clone of serotype 6B Streptococcus pneumoniae from Spain to Iceland in the Late 1980s. J. Infect. Dis. 168:158163.
222. Stamm, W. E.,, J. C. Feeley,, and R. R. Facklam. 1978. Wound infections due to group A streptococcus traced to a vaginal carrier. J. Infect. Dis. 138:287292.
223. Slegmayr, B.,, S. Björck,, S. Holm,, J. Nisell,, A. Rydvall,, and B. Settergren. 1992. Septic shock induced by group A streptococcal infection: clinical and therapeutic aspects. Scand. J. Infect. Dis. 24:589597.
224. Slevens, D. L.,, M. H. Tanner,, J. Winship,, R. Swarts,, K. M. Ries,, P. M. Schlievert,, and E. Kaplan. 1989. Severe group A streptococcal infections associated with a toxic shock-like syndrome and scarlet fever toxin A. N. Engl. J. Med. 321:17.
225. Strasberg, S. M.,, and M. S. Silver. 1968. Hemolytic streptococcus gangrene: an uncommon but frequently fatal infection in the antibiotic era. Am. J. Surg. 115:6368.
226. Slromberg, A.,, V. Romanus,, and L. G. Burman. 1991. Outbreaks of group A streptococcal bacteremia in Sweden: an epidemiologic and clinical study. J. Infect. Dis. 164:595598.
227. Talay, S. R.,, P. Valentin-Weigand,, P. G. Jerlstrom,, K. N. Timmis,, and G. S. Chhatwal. 1992. Fibronectin-binding protein of Streptococcus pyogenes: sequence of the binding domain involved in adherence of streptococci to epithelial cells. Infect. Immun. 60:38373844.
228. Talay, S. R.,, P. Valentin-Weigand,, K. N. Timml., and G. S. Chhatwal. 1994. Domain structure and conserved epitopes of Sfb protein, the fibronectin-binding adhesin of Streptococcus pyogenes. Mol. Microbiol. 13:531539.
229. Talkington, D. F.,, B. Schwartz,, C. M. Black,, J. K. Todd,, J. Elliott,, R. F. Breiman,, and R. R. Facklam. 1993. Association of phenotypic and genotypic characteristics of invasive Streptococcus pyogenes isolates with clinical components of streptococcal toxic shock syndrome. Infect. Immun. 61:33693374.
230. Them, A.,, L. Slenberg,, B. Dahlback,, and G. Lindahl. 1995. Ig-binding surface proteins of Streptococcus pyogenes also bind human C4b-binding protein (C4BP), a regulatory component of the complement system. J. Immunol. 154:375386.
231. Trence, D. L.,, M. Y. Khan,, and D. N. Gerding. 1981. Beta-hemolytic streptococcal bacteremia in adults: association with cold weather in Minnesota. Minn. Med. 64:675679.
232.Tylew.ka, S. K., V. A. Fischetti, and R. J. Gibbons. 1988. Binding selectivity of Streptococcus pyogenes and M protein to epithelial cells differs from that of lipoteichoic acid. Curr. Microbiol. 16:209266.
233. VanHeyningen, T.,, G. Fogg,, D. Yates,, E. Hanski,, and M. Caparon. 1993. Adherence and fibronectio binding are environmentally regulated in the group A streptococci. Mol. Microbial. 9:12131222.
234. Wagner, D. P.,, P. M. Schlieverl,, A. P. Assimacopoulos,, A. Stoehr,, P. J. Carson,, and K. Komadina. 1996. Acute group G streptococcal myositis associated with streptococcal toxic shock syndrome: case report and review. Clin. Infect. Dis. 23:11591161.
235. Wang, H.,, R. Lottenberg,, and M. D. P. Boyle. 1995. Analysis of the interaction of group A streptococci with fibrinogen. streptokinase and plasminogen. Microb. Pathog. 18:153166.
236. Wang, H.,, R. Lottenberg,, and M. D. P. Boyle. 1995. Role for fibrinogen in the streptokinase-dependent acquisition of plasmin(ogen) by group A streptococci. J. Infect. Dis. 171:8592.
237. Wang, J.•R., and M. W. Stinson. 1994. M protein mediates streptococcal adhesion to HEp-2 cells. Infect. Immun. 62:442448.
238. Wannamaker, L. W. 1970. Differences between streptococcal infections of the throat and of the skin. N. Engl. J. Med. 282:2331.
239. Watanabe•Ohnishi, R.,, D. E. Low,, A. McGeer,, D. L. Stevens,, P. M. Schlievert,, D. Newton,, B. Schwartz,, B. Kreiswirth, and Ontario Group A Streptococcal Study. 1995. Selective depletion of Vβ•bearing T cells in patients with severe invasive group A streptococcal infections and streptococcal toxic shock syndrome. J. Infect. Dis. 171:7484.
240. Webb, H. E.,, N. W. Hoover,, D. R. Nichols,, and L. A. Weed. 1962. Streptococcal gangrene. Arch. Surg. 85:969973.
241. Weinbren, M. J.,, and R. M. Perinpanayagam. 1992. Streptococcal necrotising fasciitis. J. Infect. 25:299302.
242. Wessels, M. R.,, and M. S. Bronze. 1994. Critical role of the group A streptococcal capsule in pharyngeal colonization and infection in mice. Proc. Natl. Acad. Sci. USA 91:1223812242.
243. Wessels, M. R.,, J. B. Goldberg,, A. E. Moses,, and T. J. DiCesare. 1994. Effects on virulence of mutations in a locus essential for hyaluronic acid expression in group A streptococci. Infect. Immun. 62:433441.
244. Wessels, M. R.,, A. E. Moses,, J. B. Goldberg,, and T. J. DiCesare. 1991. Hyaluronic acid capsule is a virulence factor for mucoid group A streptococci. Proc. Natl. Acad. Sci. USA 88:83178321.
245. Whatmore, A. M.,, V. Kapur,, J. M. Musser,, and M. A. Kehoe. 1995. Molecular population genetic analysis of the emm subdivision of group A streptococcal emm-like genes: horizontal gene transfer and restricted variation among enn genes. Mol. Microbial. 15:10391048.
246.. Whatmore, A. M.,, V. Kapur,, D. J. Sullivan,, J . M. Musser,, and M. A. Kehoe. 1994. Noncongruent relationships between variation in emm gene sequences and the population generic structure of group A streptococci. Mol. Microbial. 14:619631.
247. Whatmore, A. M.,, and M. A. Kehoe. 1994. Horizontal gene transfer in the evolution of group A streptococcal emm-like genes: gene mosaics and variation in Vir regulons. Mol. Microbiol. 11:363374.
248. White, W. L. 1953. Hemolytic Streptococcus gangrene: a report of seven cases. Plast. Reconstr. Surg. 2:114.
249. Whitnack, E., and E- H. Beachey. 1985. Biochemical and biological properties of the binding of human fibrinogen to M protein in group A streptococci. J. Bacteriol. 164:350358.
250. Wilson, B. 1952. Necrotizing fasciitis. Am. Surg. 18:416431.
251. Wolf, B. B.,, C. M. Gibson,, V. Kapur,, I. M. Hussaini,, J. M. Musser,, and S. L. Gonias. 1994. Proteolytically active streptococcal pyrogenic exotoxin B cleaves monocytic cell urokinase receptor and releases an active fragment of the receptor from the cell surface. J. Biol. Chem. 269: 3068230687.
252. Wood, T. F.,, M. A. Potter,, and O. Jonasson. 1993. Streptococcal toxic shock-like syndrome: the importance of surgical intervention. Ann. Surg. 217:109114.
253.The Working Group on Severe Streptococcal Infections. 1993. Defining the group A streptococcal toxic shock syndrome. Rationale and consensus definition. JAMA 269:384, 390391.
254.World Health Organization. 1984. Rheumatic Fever and Rheumatic Heart Disease. World Health Organization. Geneva, Switzerland.
255. Xu, S.,, and C. M. Collins. 1996. Temperature regulation of the streptococcal pyrogenic exotoxin A-encoding gene (specA). Infect. Immun. 64:53995402.
256. Yoder, E. L.,, J. Mendez,, and R. Khatib. 1987. Spontaneous gangrenous myositis induced by Streptococcus pyogenes: case report and review of the literature. Rev. Infect. Dis. 9:382385.
257. Yutsudo, T.,, K. Okumura,, M. Iwasaki,, A. Hara,. S. Kamitani,, W. Minamide,, H. Igarashi,, and Y. Hinuma. 1994. The gene encoding a new mitogenic factor in a Streptococcus pyogenes strain is distributed on1y in group A streptococci. Infect. Immun. 62:40004004.
258. Zhao, Y.•X., A. Abdelnour, T. Kalland, and A. Tarkowski. 1995. Overexpression of the T-cell receptor Vβ3 in transgenic mice increases mortality during infection by enterotoxin A-producing Staphylococcus aureus. Infect. Immun. 63:44634469.


Generic image for table
Table 1.

Possible virulence and transmissibility determinants of group A streptococci: adherence and colonization

Citation: Low D, McGeer A, Schwartz B. 1998. The Reemergence of Severe Group A Streptococcal Disease: an Evolutionary Perspective, p 93-123. In Scheld W, Armstrong D, Hughes J (ed), Emerging Infections 1. ASM Press, Washington, DC. doi: 10.1128/9781555816940.ch7