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The strains recommended for use in the bacterial reverse mutation test (OECD guideline 471) can be certified as non-genetically modified organisms

The strains recommended for use in the bacterial reverse mutation test (OECD guideline 471) can... The bacterial reverse mutation test, commonly called Ames test, is used worldwide. In Japan, the genetically modified organisms (GMOs) are regulated under the Cartagena Domestic Law, and organisms obtained by self- cloning and/or natural occurrence would be exempted from the law case by case. The strains of Salmonella typhimurium and Escherichia coli recommended for use in the bacterial reverse mutation test (OECD guideline 471), have been considered as non-GMOs because they can be constructed by self-cloning or naturally occurring bacterial strains, or do not disturb the biological diversity. The present article explains the reasons why these tester strains should be classified as non-GMOs. Keywords: Bacterial reverse mutation test, Genetically modified organisms, Biodiversity, Natural occurrence, Self- cloning, pKM101, pAQ1 Definition of genetically modified organisms Gene mutations of the Ames tester strains recommended Genetically modified organisms (GMOs) are defined as in OECD guideline 471 an organism “in which the genetic material (DNA) has The strains which are used in the bacterial reverse been altered in a way that does not occur naturally by mutation test (OECD guideline 471) [4] are derivatives mating and/or natural recombination, without using of S. enterica serovar Typhimurium (S. typhimurium) modern recombinant DNA technology” [1]. Accordingly, LT2 or E. coli B strain [5–7]. All the Ames tester strains organisms are considered to be a non-GMO if they are recommended for use in the bacterial reverse mutation made by the transfer of genetic material through bacter- test are listed in Table 1. The Salmonella tester strains ial conjugation between same/different species. For ex- harbor different mutations (hisD3052, hisG46, hisC3076, ample, the transfer of the antibiotic resistance genes hisG428, hisD6610 and hisO1242) in the genes of the naturally occurs by bacterial conjugation in a broad host histidine operon of S. typhimurium.The Salmonella range [2]. It is also known that bacteria gain the property strains originated from S. typhimurium LT2 are histidine of antibiotic resistance through the genetic mutations auxotrophs which are the result from treatment with and horizontal transfer of the antibiotic resistance genes mutagens or radiation [5, 6, 8–13]. In addition, the all under selective pressures [3]. Therefore, non-GMOs are Salmonella tester strains carry an rfa (deep rough) mu- not considered to disturb the biological diversity. tation for permeation of test chemicals, and the strains except for TA102 have a deletion mutation of uvrB gene to keep adducts generated with test chemicals as well as gal, chl, and bio genes [5, 6, 14]. The two tester strains of E. coli carry a terminating ochre mutation in the trpE gene as well as a uvrA mutation [15, 16]. Thus, the * Correspondence: a-hakura@hhc.eisai.co.jp Tsukuba Drug Safety, Eisai Co., Ltd., 5-1-3 Tokodai, Tsukuba-shi, Ibaraki genetic background changes (mutations) in the Ames 300-2635, Japan Full list of author information is available at the end of the article © 2016 Sugiyama et al. Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The Creative Commons Public Domain Dedication waiver (http://creativecommons.org/publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated. Sugiyama et al. Genes and Environment (2016) 38:2 Page 2 of 3 Table 1 Strains recommended for use in the bacterial reverse mutation test (OECD guideline 471) Strain Orignal Genotype Plasmid S. typhimurium TA98 S. typhimurium LT2 hisD3052 rfa Δ(gal chl bio uvrB) pKM101 S. typhimurium TA100 S. typhimurium LT2 hisG46 rfa Δ(gal chl bio uvrB) pKM101 S. typhimurium TA1535 S. typhimurium LT2 hisG46 rfa Δ(gal chl bio uvrB) None S. typhimurium TA1537 S. typhimurium LT2 hisC3076 rfa Δ(gal chl bio uvrB) None S. typhimurium TA102 S. typhimurium LT2 hisG428 rfa galE hisΔ(G)8476 pKM101, pAQ1 S. typhimurium TA97/TA97a S. typhimurium LT2 hisD6610 hisO1242 rfa Δ(gal chl bio uvrB) pKM101 E. coli WP2uvrA E. coli B trpE uvrA None E. coli WP2uvrA/pKM101 E. coli B trpE uvrA pKM101 tester strains can naturally occur without using modern introduced into the genome of S. typhimurium. The recombinant DNA technology. hisG428 gene is inserted into the ampicillin resistance gene. It was also reported that S. typhimurium LT2 pKM101 plasmid can naturally occur and self- strains are inherently non-colicinogenic, but the strain transmissible was shown to have an ability to receive colicin plasmids As shown in Table 1, the five strains of S. typhimurium from E. coli through conjugation [17, 25, 26]. Thus S. (TA98, TA100, TA102, TA97 and TA97a) and one strain typhimurium has a possibility to have the plasmid pMB1 of E. coli (WP2uvrA/pKM101) harbor plasmid pKM101. as well as toxin colicin naturally in their cells even with- Plasmid pKM101 carries an ampicillin resistance gene out introducing pAQ1. Therefore, the introduction of and mucAB genes encoding analogs of UmuD/C pro- pAQ1 into the Ames tester strains does not disturb the teins of E. coli, which are involved in error-prone DNA biological diversity of S. typhimurium, and pAQ1 plas- repair [6, 17]. pKM101 (35.4 kb) is derived from its clin- mid can be generated via self-cloning technology and ically isolated parent R46 plasmid by an in vivo 14-kb transferred to S. typhimurium LT2. deletion [18]. R46 plasmid contains four drug-resistance genes, while pKM101 dose not contain the other three Conclusion drug-resistance genes with the exception of the ampicil- In Japan, the Cartagena Domestic Law regulates living lin resistance gene [19, 20]. In addition, since R plasmids organisms resulting from modern biotechnology in- have a self-transmissible nature, pKM101 is normally cluding recombinant DNA technology, and probable present in the members of the family Enterobacteriaceae exemptions for microorganisms obtained by self- including the genera Salmonella and Escherichia [6, 17]. cloning and/or “natural occurrence” are assessed and Taken together, plasmid pKM101 is considered to be a decided case by case (for each produced organism) derivative of a naturally occurring plasmid, and self- [27, 28]. Based on the following stated reasons, we transmittable. conclude that all the Ames tester strains recommended for use in the bacterial mutation test [4] can be certi- pAQ1 plasmid in the Ames tester strains does not disturb fied as non-GMOs; the biological diversity The S. typhimurium TA102 strain harbors plasmids 1) Genetic backgrounds of the nine strains pAQ1 in addition to pKM101. The pAQ1 is a derivative recommended for use in the bacterial mutation test of pBR322 and carries the target DNA sequence for re- [4] can be generated spontaneously, or by radiation version, hisG428, a part of the histidine biosynthetic op- or chemicals. eron originated from S. typhimurium. Thus, hisG428 is a 2) pKM101 harbored in the tester strains TA97, TA97a, self-cloned gene. The vector pBR322 consists of the fol- TA98, TA100, TA102, and WP2uvrA/pKM101 is a lowing DNA segments assembled in vitro; the tetracyc- naturally occurring plasmid and self-transmittable. line resistance gene, ampicillin resistance gene, and the 3) pAQ1 plasmid which the strain TA102 carries, can replicator regions derived from colicin plasmid, pMB1 be generated via self-cloning technology and trans- [21, 22]. The two drug resistance genes are derived from ferred to S. typhimurium LT2 by conjugation. transposons, Tn10 and Tn3, respectively [22, 23]. Trans- posons are known to be transferred between related bac- Competing interests teria [24]. So, the drug resistance genes can be naturally The authors declare that they have no competing interests. Sugiyama et al. Genes and Environment (2016) 38:2 Page 3 of 3 Authors’ contributions 22. Bolivar F, Rodriguez RL, Greene PJ, Betlach MC, Heyneker HL, Boyer HW, KS drafted the manuscript. MY, AH, and TA critically reviewed on the et al. Construction and characterization of new cloning vehicles. II. A manuscript. All authors read and approved the final manuscript. multipurpose cloning system. Gene. 1977;2:95–113. 23. Bukhari AI, Shapiro JA, Adhya SL. DNA insertion elements, plasmids and episomes. Cold Spring Harbor, N.Y.: Cold Spring Harbor Laboratory; Acknowledgements 1977. p. 639–40. The authors are grateful to the members of JEMS (The Japanese 24. de la Cruz F, Davies J. Horizontal gene transfer and the origin of species: Environmental Mutagen Society)/BMS (The Bacterial Mutagenicity Study lessons from bacteria. Trends Microbiol. 2000;8:128–33. Group) for their valuable comments and helpful discussion on these 25. Levin DE, Hollstein M, Christman MF, Schwiers EA, Ames BN. A new contents. Salmonella tester strain (TA102) with A� T base pairs at the site of mutation detects oxidative mutagens. Proc Natl Acad Sci U S A. 1982;79:7445–9. Author details 1 26. Sutcliffe J. Complete nucleotide sequence of the Escherichia coli plasmid Division of Genetics and Mutagenesis, National Institute of Health Sciences, 2 pBR322, Cold Spring Harb Symp Quant Biol. 1979;43 Pt 1: 77-90. 1-18-1 Kamiyoga, Setagaya-ku, Tokyo 158-8501, Japan. Drug Safety Research 27. The Cartagena Protocol on Biosafety to the Convention on Biological Center, Tokushima Research Institute, Otsuka Pharmaceutical Co., Ltd., 463-10 3 Diversity, Montreal, 2000. URL: https://www.cbd.int/doc/legal/cartagena- Kagasuno, Kawauchi-cho, Tokushima 771-0192, Japan. Tsukuba Drug Safety, protocol-en.pdf . Accessed 11 Jan 2016. Eisai Co., Ltd., 5-1-3 Tokodai, Tsukuba-shi, Ibaraki 300-2635, Japan. 28. The Conservation and Sustainable Use of Biological Diversity through Regulations on the Use of Living Modified Organisms (Act No 97 of 2003). Received: 17 October 2015 Accepted: 4 December 2015 References 1. European Commission. Directive 2001/18/EC of the European Parliament and the Council of 12 March 2001 on the deliberate release into the environment of genetically modified organisms and repealing Council Directive 90/220/EEC, L106. Official J Eur Communities. 2001: 1-38. 2. Courvalin P. Transfer of antibiotic resistance genes between gram-positive and gram-negative bacteria. Antimicrob Agents Chemother. 1994;38:1447. 3. Mazel D, Davies J. Antibiotic resistance in microbes. Cell Mol Life Sci. 1999; 56:742–54. 4. OECD guideline for the testing of chemicals 471. Bacterial Reverse Mutation Test. 1997. 5. Ames BN, Lee FD, Durstonv WE. An improved bacterial test system for the detection and classification of mutagens and carcinogens. Proc Natl Acad Sci U S A. 1973;70:782–6. 6. Maron DM, Ames BN. Revised methods for the Salmonella mutagenicity test. Mutat Res. 1983;113:173–215. 7. Mortelmans K, Riccio ES. The bacterial tryptophan reverse mutation assay with Escherichia coli WP2. Mutat Res. 2000;455:61–9. 8. Ames BN, Durston WE, Yamasaki E, Lee FD. Carcinogens are mutagens: a simple test system combining liver homogenates for activation and bacteria for detection. Proc Natl Acad Sci U S A. 1973;70:2281–5. 9. Gee P, Maron DM, Ames BN. Detection and classification of mutagens: a set of base-specific Salmonella tester strains. Proc Natl Acad Sci U S A. 1994;24:11606–10. 10. Mortelmans K, Zeiger E. The Ames Salmonella/microsome mutagenicity assay. Mutat Res. 2000;455:29–60. 11. Tejs S. The Ames test: a methodological short review. Environ Biotechnol. 2008;4:7–14. 12. Hartman PE, Hartman Z, Stahl RC, Ames BN. Classification and mapping of spontaneous and induced mutations in the histidine operon of Salmonella. Adv Genet. 1971;16:1–34. 13. Whitfield HJ, Martin RG, Ames BN. Classification of aminotransferase (C gene) mutants in the histidine operon. J Mol Biol. 1966;21:335–55. 14. O'Donovan M. The comparative responses of Salmonella typhimurium TA1537 and TA97a to a range of reference mutagens and novel compounds. Mutagenesis. 1990;5:267–74. 15. Green M, Muriel W. Mutagen testing using Trp reversion in Escherichia coli. Submit your next manuscript to BioMed Central Mutat Res. 1976;38:3–32. 16. Parry JM, Parry EM. Genetic toxicology: principles and methods. and we will help you at every step: Methods Mol Biol. 2012;817:21–34. • We accept pre-submission inquiries 17. Mortelmans K. Isolation of plasmid pKM101 in the Stocker laboratory. Mutat Res. 2006;612:151–64. � Our selector tool helps you to find the most relevant journal 18. Winans SC, Walker GC. Conjugal transfer system of the IncN plasmid � We provide round the clock customer support pKM101. J Bacteriol. 1985;161:402–10. � Convenient online submission 19. Langer PJ, Walker GC. Restriction endonuclease cleavage map of pKM101: relationship to parental plasmid R46. Mol Gen Genet. 1981;182:268–72. � Thorough peer review 20. Langer PJ, Shanabruch WG, Walker GC. Functional organization of plasmid � Inclusion in PubMed and all major indexing services pKM101. J Bacteriol. 1981;145:1310–6. � Maximum visibility for your research 21. Betlach MC, Hershfield V, Chow L, Brown W, Goodman HM, Boyer HW. A restriction endonuclease analysis of the bacterial plasmid controlling the Submit your manuscript at EcoRI restricition and modification of DNA. Fed Proc. 1976;35:2035. www.biomedcentral.com/submit http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.png Genes and Environment Springer Journals

The strains recommended for use in the bacterial reverse mutation test (OECD guideline 471) can be certified as non-genetically modified organisms

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Springer Journals
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Copyright © 2016 by The Author(s)
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Biomedicine; Human Genetics
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Abstract

The bacterial reverse mutation test, commonly called Ames test, is used worldwide. In Japan, the genetically modified organisms (GMOs) are regulated under the Cartagena Domestic Law, and organisms obtained by self- cloning and/or natural occurrence would be exempted from the law case by case. The strains of Salmonella typhimurium and Escherichia coli recommended for use in the bacterial reverse mutation test (OECD guideline 471), have been considered as non-GMOs because they can be constructed by self-cloning or naturally occurring bacterial strains, or do not disturb the biological diversity. The present article explains the reasons why these tester strains should be classified as non-GMOs. Keywords: Bacterial reverse mutation test, Genetically modified organisms, Biodiversity, Natural occurrence, Self- cloning, pKM101, pAQ1 Definition of genetically modified organisms Gene mutations of the Ames tester strains recommended Genetically modified organisms (GMOs) are defined as in OECD guideline 471 an organism “in which the genetic material (DNA) has The strains which are used in the bacterial reverse been altered in a way that does not occur naturally by mutation test (OECD guideline 471) [4] are derivatives mating and/or natural recombination, without using of S. enterica serovar Typhimurium (S. typhimurium) modern recombinant DNA technology” [1]. Accordingly, LT2 or E. coli B strain [5–7]. All the Ames tester strains organisms are considered to be a non-GMO if they are recommended for use in the bacterial reverse mutation made by the transfer of genetic material through bacter- test are listed in Table 1. The Salmonella tester strains ial conjugation between same/different species. For ex- harbor different mutations (hisD3052, hisG46, hisC3076, ample, the transfer of the antibiotic resistance genes hisG428, hisD6610 and hisO1242) in the genes of the naturally occurs by bacterial conjugation in a broad host histidine operon of S. typhimurium.The Salmonella range [2]. It is also known that bacteria gain the property strains originated from S. typhimurium LT2 are histidine of antibiotic resistance through the genetic mutations auxotrophs which are the result from treatment with and horizontal transfer of the antibiotic resistance genes mutagens or radiation [5, 6, 8–13]. In addition, the all under selective pressures [3]. Therefore, non-GMOs are Salmonella tester strains carry an rfa (deep rough) mu- not considered to disturb the biological diversity. tation for permeation of test chemicals, and the strains except for TA102 have a deletion mutation of uvrB gene to keep adducts generated with test chemicals as well as gal, chl, and bio genes [5, 6, 14]. The two tester strains of E. coli carry a terminating ochre mutation in the trpE gene as well as a uvrA mutation [15, 16]. Thus, the * Correspondence: a-hakura@hhc.eisai.co.jp Tsukuba Drug Safety, Eisai Co., Ltd., 5-1-3 Tokodai, Tsukuba-shi, Ibaraki genetic background changes (mutations) in the Ames 300-2635, Japan Full list of author information is available at the end of the article © 2016 Sugiyama et al. Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The Creative Commons Public Domain Dedication waiver (http://creativecommons.org/publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated. Sugiyama et al. Genes and Environment (2016) 38:2 Page 2 of 3 Table 1 Strains recommended for use in the bacterial reverse mutation test (OECD guideline 471) Strain Orignal Genotype Plasmid S. typhimurium TA98 S. typhimurium LT2 hisD3052 rfa Δ(gal chl bio uvrB) pKM101 S. typhimurium TA100 S. typhimurium LT2 hisG46 rfa Δ(gal chl bio uvrB) pKM101 S. typhimurium TA1535 S. typhimurium LT2 hisG46 rfa Δ(gal chl bio uvrB) None S. typhimurium TA1537 S. typhimurium LT2 hisC3076 rfa Δ(gal chl bio uvrB) None S. typhimurium TA102 S. typhimurium LT2 hisG428 rfa galE hisΔ(G)8476 pKM101, pAQ1 S. typhimurium TA97/TA97a S. typhimurium LT2 hisD6610 hisO1242 rfa Δ(gal chl bio uvrB) pKM101 E. coli WP2uvrA E. coli B trpE uvrA None E. coli WP2uvrA/pKM101 E. coli B trpE uvrA pKM101 tester strains can naturally occur without using modern introduced into the genome of S. typhimurium. The recombinant DNA technology. hisG428 gene is inserted into the ampicillin resistance gene. It was also reported that S. typhimurium LT2 pKM101 plasmid can naturally occur and self- strains are inherently non-colicinogenic, but the strain transmissible was shown to have an ability to receive colicin plasmids As shown in Table 1, the five strains of S. typhimurium from E. coli through conjugation [17, 25, 26]. Thus S. (TA98, TA100, TA102, TA97 and TA97a) and one strain typhimurium has a possibility to have the plasmid pMB1 of E. coli (WP2uvrA/pKM101) harbor plasmid pKM101. as well as toxin colicin naturally in their cells even with- Plasmid pKM101 carries an ampicillin resistance gene out introducing pAQ1. Therefore, the introduction of and mucAB genes encoding analogs of UmuD/C pro- pAQ1 into the Ames tester strains does not disturb the teins of E. coli, which are involved in error-prone DNA biological diversity of S. typhimurium, and pAQ1 plas- repair [6, 17]. pKM101 (35.4 kb) is derived from its clin- mid can be generated via self-cloning technology and ically isolated parent R46 plasmid by an in vivo 14-kb transferred to S. typhimurium LT2. deletion [18]. R46 plasmid contains four drug-resistance genes, while pKM101 dose not contain the other three Conclusion drug-resistance genes with the exception of the ampicil- In Japan, the Cartagena Domestic Law regulates living lin resistance gene [19, 20]. In addition, since R plasmids organisms resulting from modern biotechnology in- have a self-transmissible nature, pKM101 is normally cluding recombinant DNA technology, and probable present in the members of the family Enterobacteriaceae exemptions for microorganisms obtained by self- including the genera Salmonella and Escherichia [6, 17]. cloning and/or “natural occurrence” are assessed and Taken together, plasmid pKM101 is considered to be a decided case by case (for each produced organism) derivative of a naturally occurring plasmid, and self- [27, 28]. Based on the following stated reasons, we transmittable. conclude that all the Ames tester strains recommended for use in the bacterial mutation test [4] can be certi- pAQ1 plasmid in the Ames tester strains does not disturb fied as non-GMOs; the biological diversity The S. typhimurium TA102 strain harbors plasmids 1) Genetic backgrounds of the nine strains pAQ1 in addition to pKM101. The pAQ1 is a derivative recommended for use in the bacterial mutation test of pBR322 and carries the target DNA sequence for re- [4] can be generated spontaneously, or by radiation version, hisG428, a part of the histidine biosynthetic op- or chemicals. eron originated from S. typhimurium. Thus, hisG428 is a 2) pKM101 harbored in the tester strains TA97, TA97a, self-cloned gene. The vector pBR322 consists of the fol- TA98, TA100, TA102, and WP2uvrA/pKM101 is a lowing DNA segments assembled in vitro; the tetracyc- naturally occurring plasmid and self-transmittable. line resistance gene, ampicillin resistance gene, and the 3) pAQ1 plasmid which the strain TA102 carries, can replicator regions derived from colicin plasmid, pMB1 be generated via self-cloning technology and trans- [21, 22]. The two drug resistance genes are derived from ferred to S. typhimurium LT2 by conjugation. transposons, Tn10 and Tn3, respectively [22, 23]. Trans- posons are known to be transferred between related bac- Competing interests teria [24]. So, the drug resistance genes can be naturally The authors declare that they have no competing interests. Sugiyama et al. Genes and Environment (2016) 38:2 Page 3 of 3 Authors’ contributions 22. Bolivar F, Rodriguez RL, Greene PJ, Betlach MC, Heyneker HL, Boyer HW, KS drafted the manuscript. MY, AH, and TA critically reviewed on the et al. Construction and characterization of new cloning vehicles. II. A manuscript. All authors read and approved the final manuscript. multipurpose cloning system. Gene. 1977;2:95–113. 23. Bukhari AI, Shapiro JA, Adhya SL. DNA insertion elements, plasmids and episomes. Cold Spring Harbor, N.Y.: Cold Spring Harbor Laboratory; Acknowledgements 1977. p. 639–40. The authors are grateful to the members of JEMS (The Japanese 24. de la Cruz F, Davies J. Horizontal gene transfer and the origin of species: Environmental Mutagen Society)/BMS (The Bacterial Mutagenicity Study lessons from bacteria. Trends Microbiol. 2000;8:128–33. Group) for their valuable comments and helpful discussion on these 25. Levin DE, Hollstein M, Christman MF, Schwiers EA, Ames BN. A new contents. Salmonella tester strain (TA102) with A� T base pairs at the site of mutation detects oxidative mutagens. Proc Natl Acad Sci U S A. 1982;79:7445–9. Author details 1 26. Sutcliffe J. Complete nucleotide sequence of the Escherichia coli plasmid Division of Genetics and Mutagenesis, National Institute of Health Sciences, 2 pBR322, Cold Spring Harb Symp Quant Biol. 1979;43 Pt 1: 77-90. 1-18-1 Kamiyoga, Setagaya-ku, Tokyo 158-8501, Japan. Drug Safety Research 27. The Cartagena Protocol on Biosafety to the Convention on Biological Center, Tokushima Research Institute, Otsuka Pharmaceutical Co., Ltd., 463-10 3 Diversity, Montreal, 2000. URL: https://www.cbd.int/doc/legal/cartagena- Kagasuno, Kawauchi-cho, Tokushima 771-0192, Japan. Tsukuba Drug Safety, protocol-en.pdf . Accessed 11 Jan 2016. Eisai Co., Ltd., 5-1-3 Tokodai, Tsukuba-shi, Ibaraki 300-2635, Japan. 28. The Conservation and Sustainable Use of Biological Diversity through Regulations on the Use of Living Modified Organisms (Act No 97 of 2003). Received: 17 October 2015 Accepted: 4 December 2015 References 1. European Commission. Directive 2001/18/EC of the European Parliament and the Council of 12 March 2001 on the deliberate release into the environment of genetically modified organisms and repealing Council Directive 90/220/EEC, L106. Official J Eur Communities. 2001: 1-38. 2. Courvalin P. Transfer of antibiotic resistance genes between gram-positive and gram-negative bacteria. Antimicrob Agents Chemother. 1994;38:1447. 3. Mazel D, Davies J. Antibiotic resistance in microbes. Cell Mol Life Sci. 1999; 56:742–54. 4. OECD guideline for the testing of chemicals 471. Bacterial Reverse Mutation Test. 1997. 5. Ames BN, Lee FD, Durstonv WE. An improved bacterial test system for the detection and classification of mutagens and carcinogens. Proc Natl Acad Sci U S A. 1973;70:782–6. 6. Maron DM, Ames BN. Revised methods for the Salmonella mutagenicity test. Mutat Res. 1983;113:173–215. 7. Mortelmans K, Riccio ES. The bacterial tryptophan reverse mutation assay with Escherichia coli WP2. Mutat Res. 2000;455:61–9. 8. Ames BN, Durston WE, Yamasaki E, Lee FD. Carcinogens are mutagens: a simple test system combining liver homogenates for activation and bacteria for detection. Proc Natl Acad Sci U S A. 1973;70:2281–5. 9. Gee P, Maron DM, Ames BN. Detection and classification of mutagens: a set of base-specific Salmonella tester strains. Proc Natl Acad Sci U S A. 1994;24:11606–10. 10. Mortelmans K, Zeiger E. The Ames Salmonella/microsome mutagenicity assay. Mutat Res. 2000;455:29–60. 11. Tejs S. The Ames test: a methodological short review. Environ Biotechnol. 2008;4:7–14. 12. Hartman PE, Hartman Z, Stahl RC, Ames BN. Classification and mapping of spontaneous and induced mutations in the histidine operon of Salmonella. Adv Genet. 1971;16:1–34. 13. Whitfield HJ, Martin RG, Ames BN. Classification of aminotransferase (C gene) mutants in the histidine operon. J Mol Biol. 1966;21:335–55. 14. O'Donovan M. The comparative responses of Salmonella typhimurium TA1537 and TA97a to a range of reference mutagens and novel compounds. Mutagenesis. 1990;5:267–74. 15. Green M, Muriel W. Mutagen testing using Trp reversion in Escherichia coli. Submit your next manuscript to BioMed Central Mutat Res. 1976;38:3–32. 16. Parry JM, Parry EM. Genetic toxicology: principles and methods. and we will help you at every step: Methods Mol Biol. 2012;817:21–34. • We accept pre-submission inquiries 17. Mortelmans K. Isolation of plasmid pKM101 in the Stocker laboratory. Mutat Res. 2006;612:151–64. � Our selector tool helps you to find the most relevant journal 18. Winans SC, Walker GC. Conjugal transfer system of the IncN plasmid � We provide round the clock customer support pKM101. J Bacteriol. 1985;161:402–10. � Convenient online submission 19. Langer PJ, Walker GC. Restriction endonuclease cleavage map of pKM101: relationship to parental plasmid R46. Mol Gen Genet. 1981;182:268–72. � Thorough peer review 20. Langer PJ, Shanabruch WG, Walker GC. Functional organization of plasmid � Inclusion in PubMed and all major indexing services pKM101. J Bacteriol. 1981;145:1310–6. � Maximum visibility for your research 21. Betlach MC, Hershfield V, Chow L, Brown W, Goodman HM, Boyer HW. A restriction endonuclease analysis of the bacterial plasmid controlling the Submit your manuscript at EcoRI restricition and modification of DNA. Fed Proc. 1976;35:2035. www.biomedcentral.com/submit

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Genes and Environment – Springer Journals

Published: Jan 22, 2016

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