Bacterial inactivation by solar ultraviolet radiation compared with sensitivity to 254 nm radiation

Photochemistry and Photobiology

Volumn 85, Issue 5, 2009, Pages 1043-1052

  Coohill, Thomas P ^a   Sagripanti, Jose Luis ^b  

^a SIENA COLLEGE   (United States)

^b EDGEWOOD CHEMICAL BIOLOGICAL CENTER   (United States)

Author keywords

[No Author keywords available]

Indexed keywords

BACILLUS ANTHRACIS; BACILLUS SUBTILIS; BACTERIAL SPORE; PHYSIOLOGY; RADIATION EXPOSURE; REVIEW; ULTRAVIOLET RADIATION;

BACILLUS ANTHRACIS; BACILLUS SUBTILIS; SPORES, BACTERIAL; ULTRAVIOLET RAYS;

BACTERIA (MICROORGANISMS);

EID: 69649098080     PISSN: 00318655     EISSN: None     Source Type: Journal    
DOI: 10.1111/j.1751-1097.2009.00586.x     Document Type: Review

References (121)

1. Calkins, J. T. Thordardottir (1980) The ecological significance of solar UV radiation on aquatic organisms. Nature 283, 563 566.
2. Giese, A. C. (1976) Living with the Sun's Ultraviolet Rays. Plenum Press, New York.
3. Mitchell, D. L. D. Karentz (1993) The induction and repair of DNA photodamage in the environment. In Environmental UV Photobiology (Edited by A. R. Young, J. Moan, L. O. Bjorn W. Nultsch pp. 345 371. Plenum Press, London.
4. Coohill, T. P., D.-P. Häder D. L. Mitchell (1996) Environmental ultraviolet photobiology. Photochem. Photobiol. 64 (3 401 402.
5. Jagger, J. (1985) Solar-UV Actions on Living Cells. Praeger, New York.
6. Nicholson, W. L., A. C. Schuerger P. Setlow (2005) The solar UV environment and bacterial spore UV resistance: considerations for Earth-to-Mars transport by natural processes and human spaceflight. Mut. Res. 517, 249 264.
7. Dudley, J. P. M. H. Woodford (2002) Bioweapons, biodiversity, and ecocide: Potential effects of biological weapons on biological diversity. Bioscience 52, 583 592.
8. Kortepeter, M. G. G. W. Parker (2000) Potential biological weapons threats. Available at: http://www.cdc.gov/ncidod/eid/vol5no4/kortepeter.htm. Accessed date 4 August 2005.
9. Whitney, E. A. S., M. E. Beatty, T. H. Taylor Jr., R. Weyant, J. Sobel, M. J. Arduino D. A. Ashford (2003) Inactivation of Bacillus anthracis spores. Emerg. Infect. Dis. 9, 623 627.
10. Coohill, T. P. (1996) Stratospheric ozone loss, ultraviolet effects and action spectroscopy. Adv. Space Res. 18 (12 27 33.
11. Christiaens, F. J., A. Chardon, A. Fourtanier J. E. Frederick (2005) Standard ultraviolet daylight for nonextreme exposure conditions. Photochem. Photobiol. 81, 874 878.
12. Diffey, B. L. (1977) The calculation of the spectral distribution of natural ultraviolet radiation under clear day conditions. Phys. Med. Biol. 22 (2 309 316.
13. Godar, D. E. (2005) UV doses worldwide. Photochem. Photobiol. 81, 736 749.
14. Ireland, W. R. Sacher (1996) The angular distribution of solar ultraviolet, visible and near-infrared radiation from cloudless skies. Photochem. Photobiol. 63 (4 483 486.
15. Kerr, J. B. C. T. McElroy (1993) Evidence for large upward trends of ultraviolet-B radiation linked to ozone depletion. Science 262, 1032 1034.
16. Micheletti, M. I., R. D. Piacentini S. Madronich (2003) Sensitivity of biologically active UV radiation to stratospheric ozone Changes: Effects of action spectrum shape and wavelength range. Photochem. Photobiol. 78 (5 456 461.
17. Sliney, D. H., R. L., Wood Jr., P. M. Moscato W. J. Marshall (1990) Ultraviolet exposure in the outdoor environment: Measurements of ambient ultraviolet exposure levels at large zenith angels. In Light, Lasers, and Synchrotron Radiation (Edited by M. Grandolfo, A. Rindi D. H. Sliney pp. 169 179. Plenum Press, New York.
18. Coohill, T. P. (1986) Virus-cell interactions as probes for vacuum-ultraviolet radiation damage and repair. Photochem. Photobiol. 44, 359 363.
19. Jagger, J. (1967) Introduction to Research in Ultraviolet Photobiology. Prentice-Hall, Inc., Englewood Cliffs, NJ.
20. Coohill, T. P. J.-L. Sagripanti (2008) Bacterial inactivation by germicidal ultraviolet radiation. Photochem. Photobiol. 84, 1084 1090.
21. UV Products. Bacteria Destruction Chart. San Gabriel, CA.
22. Lytle, C. D. J.-L. Sagripanti (2005) Predicted inactivation of viruses of relevance to biodefense by solar radiation. J. Virol. 79, 1 9.
23. Heisler, G. M., R. H. Grant, W. Gao J. R. Slusser (2004) Solar ultraviolet-B radiation in urban environments: The case of Baltimore, Maryland. Photochem. Photobiol. 80, 422 428.
24. Puskeppeleit, M., L. E. Quintern, S. El Naggar, J.-U. Schott, U. Eschweiler, G. Horneck H. Bucker (1992) Long-term dosimetry of solar UV radiation in Antarctica with spores of Bacillus subtilis. Appl. Environ. Microbiol. 58, 2355 2359.
25. Munakata, N. (1995) Continual increase in biologically effective dose of solar UV radiation determined by spore dosimetry from 1980 to 1993 in Tokyo. J. Photochem. Photobiol. B: Biol. 31, 63 68.
26. Jagger, J. (1981) Near-UV radiation effects on microorganisms. Photochem. Photobiol. 34, 761 768.
27. Tyrrell, R. M. A. Souza-Neto (1981) Lethal effects of natural solar ultraviolet radiation in repair proficient and repair deficient strains of Escherichia coli: Actions and interactions. Photochem. Photobiol. 34, 331 337.
28. Xue, Y. W. L. Nicholson (1996) The two main spore DNA repair pathways, nucleotide excision repair and spore photoproduct lyase, are sufficient for the resistance of Bacillus subtilis spores to artificial UV-C and UV-B but not to solar radiation. Appl. Environ. Microbiol. 62 (7 2221 2227.
29. Webb, R. B. M. S. Brown (1976) Sensitivity of strains of Escherichia coli differing in repair capability to far UV, near UV and visible radiations. Photochem. Photobiol. 24, 425 432.
30. Tyrrell, R. M. (1979) Lethal cellular changes induced by near ultraviolet radiation. Acta Biol. Med. Ger. 38, 1259 1269.
31. Kelner, A. (1951) Action spectra for photoreactivation of ultraviolet-irradiated Escherichia coli and Streptomyces griseus. J. Gen. Physiol. 34, 835 852.
32. Webb, R. B. (1997) Lethal and mutagenic effects of near-ultraviolet radiation. Photochem. Photobiol. Rev. 2, 169 261.
33. Tyrrell, R. M. (1977) Solar dosimetry with repair deficient bacterial spores: Action spectra, photoproduct measurements and a comparison with other biological systems. Photochem. Photobiol. 27, 571 579.
34. Young, A. R., J. Moan, L. O. Bjorn W. Nultsch (1993) Environmental UV Photobiology. Plenum Press, NY.
35. Peak, M. J. J. G. Peak (1989) Solar-ultraviolet-induced damage to DNA. Photodermatology 6, 1 15.
36. USDA UV-B Monitoring Program. Available at: http://UVB.nrel.colostate. edu. Accessed on 10 June 2009.
37. Coohill, T. (1994) Exposure response curves action spectra and amplification factors. NATO ASI Series. In Stratospheric Ozone Depletion-UV-B in the Biosphere, Vol. 118 (Edited by R. H. Biggs M. E. B. Joyner pp. 57 62. Springer-Verlag, New York.
38. Morowitz, H. J. (1950) Absorption effects in volume irradiation of microorganisms. Science 111, 229 230.
39. Nicholson, W. L. B. Galeano (2003) UV Resistance of Bacillus antracis spores revisited: Validation of Bacillus subtilis spores as UV surrogates for spores of B. anthracis sterne. Appl. Environ. Microbiol. 69 (2 1327 1330.
40. Cerf, O. (1977) A review: Tailing of survival curves of bacterial spores. J. Appl. Bacteriol. 42, 1 19.
41. Yousef, A. E. E. H. Marth (1988) Inactivation of Listeria monocytogenes by ultraviolet energy. J. Food Sci. 53 (2 571 573.
42. Aas, P., M. Lyons, R. Pledger, D. L. Mitchell W. H. Jeffrey (1996) Inhibition of bacterial activities by solar radiation in nearshore waters and the Gulf of Mexico. Aquat. Microb. Ecol. 11, 229 238.
43. Harm, W. (1969) Biological determination of the germicidal activity of sunlight. Radiat. Res. 40, 63 69.
44. Harm, W. (1979) Relative effectiveness of the 300-320 nm spectral region of sunlight for the production of primary lethal damage in E. coli cells. Mut. Res. 60, 263 270.
45. Miller, R. V., W. Jeffrey, D. Mitchell M. Elasri (1999) Bacterial responses to ultraviolet light. ASM News 65 (8 535 538.
46. Kubitschek, H. E. M. J. Peak (1979) Action spectrum for growth delay induced by near-ultraviolet light in E. coli B/r K. Photochem. Photobiol. 31, 55 58.
47. Hanawalt, P. C. (1989) Concepts and models for DNA repair: from Escherichia coli to mammalian cells. Environ. Mol. Mutagen. 14 (16 90 98.
48. Friedberg, E. C., G. C. Walker W. Siede (1995) DNA Repair and Mutagenesis. ASM Press, Washington, DC.
49. Douki, T., B. Setlow P. Setlow (2005) Photosensitization of DNA by dipicolinic acid, a major component of spores of Bacillus species. Photochem. Photobiol. Sci. 4, 591 597.
50. Setlow, P. (1994) Mechanisms which contribute to the long-term survival of spores of Bacillus species. J. Appl. Bacteriol. (Symposium Supplement) 76, 49S 60S.
51. Nicholson, W. L. (1995) Photoreactivation in the genus Bacillus. Curr. Microbiol. 31, 361 364.
52. Stuy, J. H. (1955) Photoreactivation of ultraviolet-inactivated bacilli. Biochim. Biophys. Acta 17, 238 240.
53. Harris, G. D., V. D. Adams, D. L. Sorensen M. S. Curtis (1987) Ultraviolet inactivation of selected bacteria and viruses with photoreactivation of the bacteria. Wat. Res. 21 (6 687 692.
54. Kelner, A. (1949) Effect of visible light on the recovery of Streptomyces griseus conidia from ultraviolet irradiation injury. PNAS (USA) 35, 73 79.
55. Hoyer, O. (2000) The status of UV technology in Europe. IUVA News 2, 22 27.
56. Pakker, H., R. T. S. Martins, P. Boelen A. G. J. Buma (2000) Effects of temperature on the photoreactivation of ultraviolet-B-induced DNA damage in Palmaria palmate (Rhodophyta). J. Phycol. 36, 334 341.
57. Mitchell, D. L. R. S. Nairn (1989) The biology of the (6-4) photoproduct. Photochem. Photobiol. 49, 805 819.
58. Slieman, T. A. W. L. Nicholson (2000) Artificial solar UV radiation induces strand breaks and cyclobutane pyrimidine dimers in Bacillus subtilis spore DNA. Appl. Environ. Microbiol. 66, 199 205.
59. Eisenstark, A. (1971) Mutagenic and lethal effects of visible and near-ultraviolet light on bacterial cells. Adv. Genet. 12, 167 198.
60. Hollaender, A. (1943) Effect of long ultraviolet and short visible radiation (3500 to 4900 Å) on Escherichia coli. J. Bacteriol. 46, 531 541.
61. Horneck, G. (1995) Quantification of the biological effectiveness of environmental UV radiation. J. Photochem. Photobiol. B: Biol. 31, 43 49.
62. Kramer, G. F. B. N. Ames (1987) Oxidative mechanisms of toxicity of low-intensity near UV light in Salmonella typhimurium. J. Bacteriol. 169 (5 2259 2266.
63. Matsungaga, T., K. Hieda O. Nikaido (1991) Wavelength dependent formation of thymine dimers and (6-4) photoproducts in DNA by monochromatic ultraviolet light ranging from 150 to 365 nm. Photochem. Photobiol. 54 (3 403 410.
64. Nicholson, W. L., B. Setlow P. Setlow (2002) UV Photochemistry of DNA in vitro and in Bacillus subtilis spores at earth-ambient and low atmospheric pressure: Implications for spore survival on other planets or moons in the solar system. Astrobiology 2, 417 425.
65. Peak, M. J. J. G. Peak (1982) Single-strand breaks induced in Bacillus subtilis DNA by ultraviolet light: Action spectrum and properties. Photochem. Photobiol. 35, 675 680.
66. Peak, M. J., J. G. Peak, M. P. Moehring R. B. Webb (1984) Ultraviolet action spectra for DNA dimer induction, lethality, and mutagenesis in Escherichia coli with emphasis on the UVB region. Photochem. Photobiol. 40 (5 613 620.
67. Peak, J. G., M. J. Peak R. W. Tuveson (1983) Ultraviolet action spectra for aerobic and anaerobic inactivation of Escherichia coli strains specifically sensitive and resistant to near ultraviolet radiations. Photochem. Photobiol. 38 (5 541 543.
68. Sutherland, J. C. (2002) Biological effects of polychromatic light. J. Photochem. Photobiol. 76, 164 170.
69. Turner, M. A. R. B. Webb (1981) Comparative mutagenesis and interaction between near-ultraviolet (313- to 405-nm) and far-ultraviolet (254-nm) radiation in Escherichia coli strains with differing repair capabilities. J. Bacteriol. 147 (2 410S 417S.
70. Webb, R. B. M. S. Brown (1982) Genetic damage in Escherichia coli K12 ab2480 by broad-spectrum near-ultraviolet radiation. Science 215, 991 993.
71. Fajardo-Cavazos, P. W. L. Nicholson (2000) The TRAP-like splA protein is a trans-acting negative regulator of spore photoproduct lyase synthesis during Bacillus subtilis sporulation. J. Bacteriol. 182, 555 560.
72. Moeller, R., T. Douki, J. Cadet, E. Stackebrandt, W. L. Nicholson, P. Rettberg, G. Reitz G. Horneck (2007) UV-radiation-induced formation of DNA bipyrimidine photoproducts in Bacillus subtilis endospores and their repair during germination. Int. Microbiol. 10, 39 46.
73. Peak, H. J. J. G. Peak (1983) Use of action spectra for identifying molecular targets and mechanisms of action of solar ultraviolet light. Physiol. Plant 58, 367 372.
74. Joux, F., W. H. Jeffrey, P. Lebaron D. L. Mitchell (1999) Marine bacterial isolates display diverse responses to UV-B radiation. Appl. Environ. Microbiol. 65 (8 3820 3827.
75. Sagripanti, J.-L. D. Lytle (2004) Inactivation of viruses by solar ultraviolet radiation after release in U.S. cities. Edgewood Chemical Biological Center Document ECBC-TR-411. Aberdeen Proving Ground, U.S. Army, MD.
76. Rettberg, P. G. Horneck (1999) Intrinsic and extrinsic biomarkers for the assessment of risks from environmental UV radiation. J. Epidemiol. 9 (6S S78 S83.
77. Slieman, T. A. W. L. Nicholson (2001) Role of dipicolinic acid in survival of Bacillus subtilis spores exposed to artificial and solar UV radiation. Appl. Environ. Microbiol. 67 (3 1274 1279.
78. Caldwell, M. M. (1971) Solar ultraviolet radiation and the growth and development of higher plants. In Photophysiology (Edited by A. C. Giese 6, pp. 131 177. Academic Press, New York.
79. Setlow, R. B. (1974) The wavelengths in sunlight effective in producing skin cancer: A theoretical analysis. PNAS 71 (9 3363 3366.
80. Jeffrey, W. H., R. J. Pledger, P. Aas, S. Hager, R. B. Coffin, R. Von Haven D. L. Mitchell (1996) Diel and depth profiles of DNA photodamage in bacterioplankton exposed to ambient solar ultraviolet radiation. Mar. Ecol. Prog. Ser. 137, 283 291.
81. Ashwood-Smith, M. J., J. Copeland J. Wilcockson (1967) Sunlight and frozen bacteria. Nature 214, 33 35.
82. Billen, D. M. M. Fletcher (1974) Inactivation of dark-repair-deficient mutants of Escherichia coli by sunlight. Int. J. Radiat. Biol. 26, 73 76.
83. Buchbinder, L., M. Solowey E. B. Phelps (1941) Studies on microorganisms in simulated room environments. III. The survival rates of streptococci in the presence of natural, daylight and sunlight, and artificial illumination. J. Bacteriol. 42, 353 366.
84. Buchbinder, L. (1942) The bacterial effects of daylight and sunlight on chained gram positive cocci in simulated room environment: Theoretical and practical considerations. Aerobiology 17, 267 270.
85. Caldwell, M. M., L. O. Björn, J. F. Bornman, S. D. Fling, G. Kulandaivelu, A. H. Teramura M. Tevini (1998) Effects of increased solar ultraviolet radiation on terrestrial ecosystems. J. Photochem. Photobiol. B: Biol. 46, 40 52.
86. Conroy, R. M., M. E. Meegan, T. Joyce, K. McGuigan J. Barnes (2005) Solar disinfection of drinking water protects against cholera in children under 6 years of age. Arch. Dis. Child. 85, 293 295.
87. Davies, C. M. L. M. Evison (1991) Sunlight and the survival of enteric bacteria in natural waters. J. Appl. Bacteriol. 70, 265 274.
88. Fujioka, R. S. O. T. Narikawa (1982) Effect of sunlight on enumeration of indicator bacteria under field conditions. Appl. Environ. Microbiol. 44 (2 395 401.
89. Gameson, A. L. H. D. J. Gould (1975) The effects of solar radiation on the mortality of some terrestrial bacteria in sea water. Proceedings of the International Symposium on Sea Outfalls, pp. 209 219. Pergamon Press, New York.
90. Grigsby, P. J. Calkins (1979) The inactivation of a natural population of coliform bacteria by sunlight. Photochem. Photobiol. 31, 291 294.
91. Hernandez, E., G. A. Ferreyra W. P. MacCormack (2002) Effect of solar radiation on two Antarctic marine bacterial strains. Polar Biol. 25, 453 459.
92. Hughes, K. A. (2005) Effect of Antarctic solar radiation on sewage bacteria viability. Wat. Res. 39, 2237 2244.
93. Kehoe, S. C., M. R. Barer, L. O. Devlin K. G. McGuigan (2004) Batch process solar disinfection is an efficient means of disinfecting drinking water contaminated with Shigella dysenteriae type I. Appl. Microbiol. 38, 410 414.
94. Lonnen, J., S. Kilvington, S. C. Kehoe, F. Al-Touati K. G. McGuigan (2005) Solar and photocatalytic disinfection of protozoan, fungal and bacterial microbes in drinking water. Wat. Res. 39, 877 883.
95. Munakata, N. (1981) Killing and mutagenic action of sunlight upon Bacillus subtilis spores: A dosimetric system. Mut. Res. 82, 263 268.
96. Resnick, M. A. (1970) Sunlight-induced killing in Saccharomyces cerevisiae. Nature 226, 377 378.
97. Riesenman, P. J. W. L. Nicholson (2000) Role of the spore coat layers in Bacillus subtilis spore resistance to hydrogen peroxide, artificial UV-C, UV-B and solar UV radiation. Appl. Environ. Microbiol. 66 (2 620 626.
98. Sommer, B., A. Mariňo, Y. Solarte, M. L. Salas, C. Dierolf, C. Valiente, D. Mora, R. Rechsteiner, P. Setter, W. Wirojanagud, H. Ajarmeh, A. Al-Hassan M. Wegelin (1997) SODIS - An emerging water treatment process. Aqua 46 (3 127 137.
99. Weinbauer, M. G., S. W. Wilhelm, C. A. Suttle, R. J. Pledger D. L. Mitchell (1999) Sunlight-induced DNA damage and resistance in natural viral communities. Aquat. Microb. Ecol. 17, 111 120.
100. Smith, R. J., S. C. Kehoe, K. G. McGuigan M. R. Barer (2000) Effects of simulated solar disinfection of water on infectivity of Salmonella typhimurium. Appl. Microbiol. 31, 284 288.
101. De Flora, S., A. Camoirano, A. Izzotti C. Bennicelli (1990) Potent genotoxicity of halogen lamps, compared to fluorescent light and sunlight. Carcinogenesis 11 (12 2171 2177.
102. Beebe, J. M. G. W. Pirsch (1958) Response of air-borne species of pasteurella to artificial radiation simulating sunlight under different conditions of relative humidity. Appl. Microbiol. 6, 127 138.
103. Khaengraeng, R. R. H. Reed (2005) Oxygen and photoinactivation of Escherichia coli in UVA and sunlight. J. Appl. Microbiol. 99, 39 50.
104. Knudson, G. B. (1986) Photoreactivation of ultraviolet-irradiated, plasmid-bearing, and plasmid-free strains of Bacillus anthracis. Appl. Environ. Microbiol. 52 (3 444 449.
105. ASTM International (2005) Standard Test Method E 2414-05. Quantitative Sporicidal Three-Step Method to Determine Sporicidal Efficacy of Liquids, Liquid Sprays, and Vapor or Gases on Contaminated Carrier Surfaces. American Society for Testing and Materials, West Conshohocken, PA.
106. Sagripanti, J.-L., M. Carrera, J. Insalaco, M. Ziemski, J. Rogers R. Zandomeni (2007) Virulent spores of Bacillus anthracis and other Bacillus species deposited on solid surfaces have similar sensitivity to chemical decontaminants. J. Appl. Microbiol. 102, 11 21.
107. Munakata, N., S. Kazadzis, A. F. Bais, K. Hieda, G. Ronto, P. Rettberg G. Horneck (2000) Comparisons of spore dosimetry and spectral photometry of solar-UV radiation at four sites in Japan and Europe. Photochem. Photobiol. 72, 739 745.
108. Bérces, A., A. Fekete, S. Gáspár, P. Gróf, P. Rettberg, G. Horneck G. Rontó (1999) Biological UV dosimeters in the assessment of the biological hazard from environmental radiation. J. Photochem. Photobiol. B: Biol. 53, 36 43.
109. Quintern, L. E., M. Puskeppeleit, P. Rainer, S. Weber, S. El Nagger, U. Eschweiler G. Horneck (1994) Continuous dosimetry of the biologically harmful UV-radiation in Antarctica with a biofilm technique. J. Photochem. Photobiol. B: Biol. 22, 59 66.
110. Moeller, R., E. Stackebrant, T. Douki, J. Cadet, P. Rettberg, H.-J. Mollenkopf, G. Reitz G. Horneck (2007) DNA bipyrimidine photoproduct repair and transcriptional response of UV-C irradiated Bacillus subtilis. Arch. Microbiol. 188, 421 431.
111. Lindberg, C. G. Horneck (1991) Action spectra for survival and spore photoproduct formation of Bacillus subtilis irradiated with short-wavelength (200-300 nm) UV at atmospheric pressure and in vacuo. J. Photochem. Photobiol. B: Biol. 11, 69 80.
112. Munakata, N. (1989) Genotoxic action of sunlight upon Bacillus subtilis spores: Monitoring studies in Tokyo, Japan. J. Radiat. Res. 30, 338 351.
113. Setlow, P. (2001) Resistance of spores of Bacillus species to ultraviolet light. Environ. Mol. Mutagen. 38, 97 104.
114. Munakata, N., F. Morohoshi, K. Hieda, K. Suzuzi, Y. Furusawa, H. Shimura T. Ito (1996) Experimental correspondence between spore dosimetry and spectral photometry of solar ultraviolet radiation. Photochem. Photobiol. 63, 74 78.
115. Coohill, T. P. (1991) Photobiology school: Action spectra again? Photochem. Photobiol. 54 (5 859 870.
116. Nicholson, W. L., N. Munakata, G. Horneck, H. J. Melosh P. Setlow (2000) Resistance of Bacillus endospores to extreme terrestrial and extraterrestrial environments. Microbiol. Mol. Biol. Rev. 64 (3 548 572.
117. Moeller, R., G. Horneck, R. Facius E. Stackebrandt (2005) Role of pigmentation in protecting Bacillus sp. endospores against environmental UV radiation. FEMS Microbiol. Ecol. 51, 231 236.
118. Saxena, D., E. Ben-Dov, R. Manasherob, Z. Barak, S. Boussiba A. Zaritsky (2002) A UV tolerant mutant of Bacillus thuringiensis subsp. kurstaki producing melanCurr. Microbiol. 44, 25 30.
119. Tuveson, R. W., R. A. Larson J. Kagan (1988) Role of cloned carotenoid genes expressed in Escherichia coli in protecting against inactivation by near-UV light and specific phototoxic molecules. J. Bacteriol. 170 (10 4675 4680.
120. Sharp, D. G. (1939) The lethal action of short ultraviolet rays on several common pathogenic bacteria. J. Bacteriol. 37, 447 460.
121. Helgason, E., O. A. Økstad, d. A. Caugant, H. A. Johansen, A. Fouet, M. Mock, I. Hegna A. E. Kolstø (2000) Bacillus antracis, Bacillus cereus and Bacillus thuringiensis - One species on the basis of genetic evidence. Am. Soc. Microbiol. 66, 2627 2630.