Stafilokokal enterotoksinler

Abstract

Stafilokokal gıda zehirlenmeleri başta Staphylococcus aureus olmak üzere enterotoksijenik stafilokoklar tarafından gıdalarda oluşturulan enterotoksinlerin alınması sonucu şekillenen ve tüm dünyada yaygın olarak görülen en önemli intoksikasyonlardan biridir. Enterotoksijenik stafilokoklar en fazla kırmızı et, kanatlı eti, balık eti ve ürünleri ile süt ve ürünleri gibi proteince zengin hayvansal gıdalarda bulunur. Enterotoksijenik stafilokoklar genotipik ve fenotipik özellikleri temel alınarak karakterize edilmişlerdir. Stafilokokal enterotoksinler (SE) yaklaşık 26-31 kDa’luk tekli polipeptidlerdir. SE’ler antijenik özellikleri temel alınarak 5 büyük serolojik tipe (SEA, SEB, SEC, SED, SEE) ayrılmıştır. Son yıllarda SE’lerin yeni tiplerinin de var olduğu (SEG, SEH, SEI, SEJ, SEK, SEL, SEM, SEN ve SEO) bildirilmiş ancak bu enterotoksinlerin gıda zehirlenmeleri ile ilişkileri henüz tam olarak açıklığa kavuşturulamamıştır. SE’lerin spesifik olmayan T hücre proliferasyonunu uyaran süperantijen fonksiyonlarının yanı sıra güçlü gastrointestinal toksin olma özellikleri de vardır. SE’lerin ısıya ve tripsin, pepsin gibi gastrointestinal enzimlere dirençli olmaları en önemli özelliklerinden biridir. SEA stafilokokal gıda zehirlenmelerinde en sık rastlanan enterotoksindir ve bunu SED izlemektedir. İntoksikasyonun oluşabilmesi için 94-184 ng enterotoksinin alınması yeterlidir. SEA hücrenin logaritmik fazında üretilmekte ve σ70 benzeri bir faktörden etkilenmekte bunun yanısıra SEB, SEC ve SED çoğunlukla eksponensiyal fazdan durgunluk fazına geçiş aşamasında üretilip accessory gene regulator (agr) tarafından regüle edilmektedir. SE’lerin enterotoksijenik ve süperantijenik aktivitelerinin açıklığa kavuşturulması için daha çok çalışmaya gereksinim duyulmaktadır

Keywords

• Enterotoksijenik stafilokok,enterotoksin,genotipik özellik,gıda

Staphylococcal enterotoxins

Abstract

Staphylococcal food intoxication is one of the most common types of foodborne disease around the world. Staphylococcal food poisoning results from the ingestion of food containing preformed staphylococcal enterotoxins produced by the enterotoxigenic staphylococci mainly Staphylococcus aureus. The bacterium is most prevalent in protein rich foods of animal origin such as meat, poultry, fish and milk products. Enterotoxigenic strains of staphylococci have been extensively characterized based on genotypic and phenotypic characteristics. SEs are single polypeptides of approximately 26 to 31 kDa. SEs have been divided into 5 major serological types (SEA through SEE) on the basis of their antigenic properties. In recent years new types of SEs (SEG, SEH, SEI, SEJ, SEK, SEL, SEM, SEN, and SEO) have been also identified. However, little is known about relationship between the new SEs and foodborne intoxication at present. Staphylococcal enterotoxins (SEs) function both as potent gastrointestinal toxins as well as superantigens that stimulate non-specific T-cell proliferation. The intoxication generally is not lethal and the elderly are more susceptible than younger individuals. SEA is the enterotoxin most frequently associated with staphylococcal foodborne outbreaks followed by SED. The amount of enterotoxin necessary to cause intoxication is very small about 94-184 ng. SEs are resistant to inactivation by gastrointestinal proteolytic enzymes, such as trypsin and pepsin. The enterotoxins are quite heat resistant and the heat stability is very important property of SEs in terms of food poisoning. The SEA is produced throughout the log phase, while SEB, SEC, and SED are produced primarily during the transition from the exponential to the stationary phases of growth. Expression of SEB, SEC, and SED is affected by accessory gene regulator (agr), while SEA expressed simultaneously with the σ70-like factor. Current methods for detection of the identified enterotoxins are based on the reaction between the enterotoxins and specific antibodies. More studies are needed to answer the questions regarding the activities of SEs both as enterotoxigenic and as superantigens

Keywords

• Enterotoxigenic staphylococci,enterotoxin,genotypic property,food

References

1. Abe J, Ito Y, Onimaru M, Kohsaka T, Takeda T (2000): Characterization and distribution of a new enterotoxin- related superantigen produced by Staphylococcus aureus. Microbiol Immunol, 44, 79-88.
2. Atanassova V, Meindl A, Ring C (2001): Prevalence of Staphylococcus aureus and staphylococcal enterotoxins in raw pork and uncooked smoked ham a comparison of classical culturing detection and RFLP- PCR. J Food Prot, 63, 1144-1153.
3. Balaban N, Rasooly A (2000): Staphylococcal enteroto- xins. Int J Food Microbiol, 61,1-10.
4. Bayles KW, Iandolo JJ (1989): Genetic and molecular analyses of the gene encoding staphylococcal enterotoxin D. J Bacteriol, 171, 4799-4806.
5. Belay N, Rasooly A (2002): Staphylococcus aureus growth and enterotoxin A production in anaerobic environment. J Food Prot, 65, 199-204.
6. Bergdoll MS (1989): Staphylococcus aureus. 463-523. In: MP Doyle (Ed), Foodborne Bacterial Pathogens. Marcel Dekker, New York.
7. Betley MJ, Mekalanos JJ (1985): Staphylococcal enterotoxin A is encoded by phage. Science, 229, 185-187.
8. Betley MJ, Mekalanos JJ (1988): Nucleotide sequence of the type A staphylococcal enterotoxin gene. J Bacteriol, 170, 34-41.

9. Bohach GA, Schlievert PM (1987): Nucleotide sequence of the staphylococcal enterotoxin C1 gene and relatedness to other pyrogenic toxins. Mol Gen Genet, 209, 15-20.
10. Bone FJ, Bogie D, Morgan-Jones SC (1989): Staphylococcal food poisoning from sheep milk cheese. Epidemiol Infect, 103, 449-458.
11. Borst DW, Betley MJ (1994): Phage-associated differences in staphylococcal enterotoxin A gene (sea) expression correlate with sea allele class. Infect Immun
12. Buzby JC, Roberts T (1997): Economic costs and trade impacts of microbial foodborne illness. World Health Stat Q 50, 57-66.
13. Carpenter DF, Silverman, GJ (1974): Staphylococcal enterotoxin B and nuclease production under controlled dissolved oxygen conditions. Appl Microbiol, 28, 628-637.
14. Ciborowski P, Jeljaszewlcz (1985): Staphylococcal enzymes and virulence. 146-183. In: IA Holder (Ed), Bacterial Enzymes and Virulence. CRC Press, Florida.
15. Couch JL, Betley MJ (1989): Nucleotide sequence of the type C3 staphylococcal enterotoxin gene suggests that intergenic recombination causes antigenic variation. J Bacteriol, 171, 4507-4510.
16. Couch JL, Soltis MT, Betley MJ (1988): Cloning and nucleotide sequence of the type E staphylococcal enterotoxin gene. J Bacteriol, 170, 2954-2960.
17. Dahiya RS, Speck ML (1968): Hydrogen peroxide formation by lactobacilli and its effect on Staphylococcus aureus. J Dairy Sci, 10, 1568-1572.
18. Dinges MM, Orwin PM, Schlievert PM (2000). Exotoxins of Staphylococcus aureus. Clin Microbiol Reviews, 13, 16-34.
19. Domenech A, Hernandez FJ, Orden JA, Goyache J, Lopez B, Suarez G, Gomez-Lucia E (1992): Effect of six organic acids on staphylococcal growth and enterotoxin production. Z Lebensm Unters Forsch, 194, 124-128.
20. Erol İ (2003): Besin Hijyeni ve Mikrobiyolojisi. Ders Notu. Ankara Üniv Vet Fak Besin Hijyeni ve Tek Anabi- lim Dalı. Basılmamış.
21. Erol İ, Hildebrandt G (1992): Einfluss von Starterkulturen auf das Wachstum pathogener Keime in türkischer Rohwurst. Fleischwirtsch, 72, 90-97.
22. Erol İ, Usca A (1996): Donmuş piliç karkaslarında izole edilen koagulaz pozitif stafilokokların enterotoksin oluş- turma yeteneklerinin SET-RPLA testi ile belirlenmesi. An- kara Üniv Vet Fak Derg, 43, 443-448.
23. Erol İ, Mutluer B, Vatansever L (1993): A tipi enterotoksin oluşturan Staphylococcus aureus’un çiğ köf- tede üreme ve toksin oluşturma yeteneğinin belirlenmesi. Gıda, 18, 315-318.
24. Evenson ML, Hinds MW, Bernstein RS, Bergdool MS (1988): Estimation of human dose of staphylococcal enterotoxin A from a large outbreak of staphylococcal food poisoning involving chocolate milk. Int J Food Microbiol, 7, 311-316.
25. Fields BA, Malchiodi EL, Li H, Ysern X, Staufffacher CV, Schlievert PM (1996): Crystal structure of a T- cell receptor beta- chain complexed with a superantigen. Nature, 384, 188-192.
26. Fink PJ, Matis LA, Mcelligott DL, Bookman M, Hedrick SM (1986): Correlations between T- cell specificity and the structure of the antigen receptor. Nature, 321, 219-226.
27. Genigeorgis CA (1989): Present state of knowledge on staphylococcal intoxication. Int J Food Microbiol, 9, 327- 360.
28. Gonzalez-Fandos E, Otero A, Sierra M, Garcia-Lopez ML, Prieto M (1994): Effect of three commercial starters on growth of Staphylococcus aureus and enterotoxins (A- D) and thermonuclease production in broth. Int J Food Microbiol, 24, 321-327.
29. Haines WC, Harmon LG (1973): Effect of variations in conditions of incubation upon inhibition of Staphylococcus aureus by Pediococcus cerevisiae and Streptococcus lactis. Appl Microbiol, 25, 169-172.
30. Halpin-Dohnalek MI, Marth EH (1989): Staphylococcus aureus: Production of extracellular compounds and behavior in foods- A review. J Food Prot, 52, 267-282.
31. Harris TO, Grossman D, Kappler JW, Marrack P, Rich RR (1993): Lack of complete correlation between emetic and T-cell- stimulatory activities of staphylococcal enterotoxins. Infect Immun, 61, 3175-3183.
32. Harvey J, Patterson JT, Gibbs PA (1982)
33. Enterotoxigenicity of Staphylococcus aureus strains isolated from poultry: Raw poultry carcasses as a potential food- poisoning hazard. J Appl Bacteriol, 52, 251-258.
34. Holmberg SD, Blake PA (1984): Staphylococcal food poisoning in the United States. New facts and old misconceptions. J Am Med Assoc, 251, 487-489.
35. Hovde CD, Hackett SP, Bohach GA (1990): Nucleotide sequence of the staphylococcal enterotoxin C3 gene: sequence comparison of all three type C staphylococcal enterotoxins. Mol Gen Genet, 220, 329-333.
36. Jablonski LM, Bohach GA (1997): Staphylococcus aureus. 353-375. In: Food Microbiology Fundamentals and Frontiers. MP Doyle, LR Beuchat, TJ Montville (Eds), ASM Press, Washington DC.
37. Jay JM (1996): Staphylococcal gastroenteritis. 429-450. In: Modern Food Microbiology. 5th ed., NewYork.
38. Johns MB, Khan SA (1988): Staphylococcal enterotoxin B gene is associated with a discrete genetic element. J Bacteriol, 170, 4033-4039.
39. Kısa Ö, Albay A, Erol İ, Sırıken B, Esin N, Gün H, Yurtyeri A (1996): Kremalı pastalardan izole edilen koagulaz pozitif stafilokokların enterotoksin oluşturma ö- zelliklerinin VIDAS yöntemiyle belirlenmesi. Ankara Üniv Vet Fak Derg, 43,405-411.
40. Krakauer T (1999): Immune response to staphylococcal superantigens. Immunol Res, 20, 163-173.
41. Morita TN, Patterson JE, Woodburn MJ (1979): Magnesium and iron addition to casein hydrolysate medium for production of staphylococcal enterotoxins A, B, and C. Appl Environ Microbiol, 38, 39-42.
42. Munson SH, Tremaine MT, Betley MJ, Welch RA (1998): Identification and characterization of staphylo- coccal enterotoxin types G and I from Staphylococcus aureus. Infect Immun, 66, 3337-3348.
43. Mutluer B, Erol İ, Kaymaz Ş, Akgün S (1993): Enterotoksijenik Staphylococcus aureus suşlarının beyaz peynirde üretim ve olgunlaşma sırasındaki üreme ve enterotoksin oluşturma yetenekleri. Ankara Üniv Vet Fak Derg, 40, 413-426.
44. Omoe K, Ishikawa M, Shimoda Y, Hu, D-L, Ueda S, Shinagawa K (2002): Detection of seg, seh, and sei genes in Staphylococcus aureus isolates and determination of the enterotoxin productivities of S. aureus isolates harboring seg, seh , or sei genes. J Clin Microbiol, 40, 857-862.
45. Orwin, PM, Leung DYM, Donahue HL, Novick RP, Schlievert PM (2001): Biochemical and biological properties of staphylococcal enterotoxin K. Infect Immun
46. Pettersson H, Forsberg G (2002): Staphylococcal enterotoxin H contrasts closely related enterotoxins in species reactivity. Immunology, 106, 71-79.
47. Qi Y, Miller KJ (2000): Effect of low water activity on staphylococcal enterotoxin A and B biosynthesis. J Food Prot, 63, 473-478.
48. Rao L, Karls RK, Betley MJ (1995): In vitro transcription of pathogenesis-related genes by purified RNA polymerase from Staphylococcus aureus. J Bacteriol, 177, 2609-2614.
49. Rose SA, Modi NK, Tranter HS, Bailey NE, Stringer MF, Hambleton P (1988): Studies on the irradiation of Clostridium botulinum and Staphylococcus aureus. J Appl Bacteriol, 65, 223-229.
50. Sheldrake RF, Hoare RJT, Hutchinson JE (1980): Post- milking iodine teat skin disinfectants. I. Bactericidal efficacy. J Dairy Res, 47, 19-26.
51. Sokari T (1991): Distribution of enterotoxigenic Staphylococcus aureus in ready-to-eat foods in eastern Nigeria. Int J Food Microbiol, 12, 275-280.
52. Su YC, Wong AC (1993). Optimal condition for the production of unidentified staphylococcal enterotoxins. J Food Prot, 56, 313-316.
53. Su YC, Wong AC (1995): Identification and purification of a new staphylococcal enterotoxin H. Appl Environ Microbiol, 61, 1438- 1443.
54. Sundstrom M, Abrahmsen L, Antonsson P, Mehindate K, Mourad W, Dohlsten M (1996): The crystal structure of staphylococcal enterotoxin type dreveals Zn2+- mediated homodimerization. EMBO J, 15, 6832-6840.
55. Sutherland J, Varnam A (2002): Enterotoxin-producing Staphylococcus, Shigella, Yersinia, Vibrio, Aeromonas and Plesiomonas. 384-415. In: CW Blackburn, PJ McClure (Eds), Foodborne Pathogens. CRC press, Washington, DC.
56. Tibana A, Rayman K, Akhtar M, Szabo R (1987): Thermal stability of enterotoxins A, B, and C in a buffered system. J Food Prot, 50, 239-242.
57. Tremaine MT, Brockman DK, Betley MJ (1993): Staphylococcal enterotoxin A gene (sea) expression is not affected by the accessory gene regulator (agr). Infect Immun, 61, 356-359.
58. Todd ECD (1989): Preliminary estimates of costs of foodborne disease in the United States. J Food Prot, 52, 595-601.
59. Tompkin RB, Ambrosino JM, Stozek SK (1973): Effect of pH, sodium chloride and sodium nitrate on enterotoxin A production. Appl Microbiol, 26, 833-837.
60. Van Den Bussche RA, Lyon JD, Bohach G.A (1993): Molecular evolution of the staphylococcal and streptococcal pyrogenic toxin gene family. Mol Phylogenet Evol, 2, 281- 292.
61. Wieneke AA, Roberts D, Gilbert RT (1993): Staphylococcal food poisoning in the United Kingdom, 1969-1990. Epidemiol Infect, 110, 519-531.
62. Wisell, KT (2000): Regulation of virulence gene expression in Staphylococcus aureus. Karolinska University Press, Sweden.
63. Zhang S, Iandolo JJ, Stewart GC (1998): The enterotoxin D plasmid of Staphylococcus aureus encodes a second enterotoxin determinant (sej). FEMS Microbiol Lett,168, 227-233.