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1 Medical Microbiology Unit, University of Louvain, Avenue Hippocrate 54, B-1200 Brussels, Belgium
2 Department of Clinical Chemistry, Microbiology and Immunology, Blok A, Ghent University Hospital, De Pintelaan 185, B-9000 Ghent, Belgium
3 Laboratorium voor Microbiologie, Fac. Wetenschappen, Vakgroep WE10V, Ledeganckstraat 35, B-9000 Ghent, Belgium
4 Culture Collection of the University of Göteborg, Department of Clinical Bacteriology, Guldhedsgatan 10, 6tr, S-413 46 Göteborg, Sweden
Correspondence
Mario Vaneechoutte
Mario.Vaneechoutte{at}rug.ac.be
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Tables containing details of the strains used and 16S rDNA sequence similarity values are available as supplementary data in IJSEM Online.
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reported that three DNA groups could be recognized within C. terrigena; these three groups could also be recognized by immunotyping and whole-cell protein patterns, but could not be differentiated phenotypically. They were therefore not named as separate species, but maintained as genospecies within C. terrigena. Vaneechoutte et al. (1992) showed that these three DNA groups gave clearly different rDNA restriction digestion patterns, based on amplified 16S, spacer and the 5' part of 23S rDNA.
In a comparative study of SDS-PAGE whole-cell protein patterns of Comamonas strains (Willems et al., 1991), Comamonas testosteroni and Comamonas acidovorans (now Delftia acidovorans) were found to have distinct and characteristic protein patterns. For 46 C. terrigena strains, including E. Falsen (EF) group 10 strains and the monotype strain of Aquaspirillum aquaticum, more variable protein patterns were observed. By numerical analysis, they were grouped into four electropherovars, 1a and 1b, 2 and 3, which correspond to C. terrigena DNA groups 1, 2 and 3, respectively.
In immunodiffusion analysis with antisera against one or two representative strains of each of the DNA groups of C. terrigena, three groups were again easily distinguishable. Strains reacted strongly with the antiserum of their own DNA group, and not at all, or weakly to moderately, with antisera of other groups (Willems et al., 1991).
Data obtained in the present study enable us to describe these three DNA groups as separate species. We propose to rename C. terrigena DNA group 2 as Comamonas aquatica comb. nov., and C. terrigena DNA group 3 as Comamonas kerstersii sp. nov. The strains used in this study and the characteristics tested for each strain are available as supplementary data in IJSEM Online.
Sequencing of the 16S rRNA gene and DNA similarity calculations and cluster analysis of the sequences obtained together with sequences from GenBank were carried out as described previously (Vaneechoutte et al., 2000; Nemec et al., 2001). Fig. 1 presents the 16S rDNA similarity tree, with Delftia acidovorans IAM 12409T as the outgroup. The distance matrix is available as supplementary data in IJSEM Online. From these data, it is clear that the three DNA groups cluster separately and cannot be confused with C. testosteroni or the recently described species Comamonas denitrificans (Gumaelius et al., 2001), Comamonas nitrativorans (Etchebehere et al., 2001) or Comamonas koreensis (Chang et al., 2002).
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; the API20NE and API ID 32 GN systems (bioMérieux) and susceptibility testing to desferrioxamine (Desferal) (Lindsay & Riley, 1991; Laffineur et al., 2002) were also used. It was possible to differentiate the three groups from C. testosteroni and from each other using these methods (Table 1).
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) and C. nitrativorans (Etchebehere et al., 2001) and the other Comamonas species is that both of the former species reduce nitrate to nitrogen. C. koreensis (Chang et al., 2002) is the only non-motile Comamonas species described so far.
All three species described in the present study are Gram-negative rods that are motile by multitrichous polar flagella, often with bipolar tufts. They are aerobic and non-fermentative, do not acidify or assimilate sugars, are urease-negative, positive for nitrate reduction and negative for nitrite reduction. They are negative for the production of indole and H2S, do not hydrolyse gelatin and produce no lysine decarboxylase, ornithine decarboxylase or arginine dihydrolase. Aesculin is not hydrolysed. Phosphatase, trypsin and acidification of ethylene glycol (Laffineur et al., 2002) give negative reactions. Citrate is not assimilated at 30 °C when tested with the ID 32 GN system (AUX medium; bioMérieux), but assimilation–alkalinization is variable on Simmons' citrate agar.
The major cellular fatty acids of the three species are C16 : 1, C16 : 0 and C18 : 1
9c, but C10 : 0 3-OH, C12 : 0 and C14 : 0 are also present, as well as small amounts of C15 : 0 and C17 : 0 cyclo. In contrast to C. testosteroni, C16 : 0 2-OH is either not detected or is only present in trace amounts.
Amplification of tRNA intergenic spacers (Welsh & McClelland, 1991) and separation of the amplified DNA fragments by capillary electrophoresis (Vaneechoutte et al., 1998; Baele et al., 2000, 2001) also makes it possible to differentiate between isolates of these three species and C. testosteroni and Delftia acidovorans (Table 2). A full database with all established tRNA PCR fingerprints is freely available upon request.
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), and given the clear biochemical and genotypic (16S rDNA sequence and tRNA-PCR profile) differences between the three DNA groups as established in this study, we propose that C. terrigena actually comprises three separate species and that C. terrigena DNA group 2 should be renamed as C. aquatica, and C. terrigena DNA group 3 as C. kerstersii.
Description of Comamonas aquatica (Hylemon et al. 1973) comb. nov.
Comamonas aquatica (a.qua'ti.ca. L. fem. adj. aquatica living in water).
Basonym: Aquaspirillum aquaticum Hylemon et al. 1973.
Gram-negative rods, motile by multitrichous polar flagella, often with bipolar tufts. Grows aerobically on tryptic soy agar and broth at 30 and 35 °C. Moderate and delayed growth in broth may be observed at 40 °C; there is no growth at 42 °C. After 24 h incubation at 30 °C, colonies reach a diameter of 1–1·5 mm on blood agar. Oxidase and catalase activities are positive. Carbohydrates are neither acidified nor assimilated. Ethylene glycol is not acidified. Urease, indole and H2S are not produced; negative for lysine decarboxylase, ornithine decarboxylase and arginine dihydrolase. Nitrates are reduced, but nitrites are not. Tyrosine and gelatin are not hydrolysed. Simmons' citrate agar test is positive. On Simmons' minimal agar base, L-alanine and glycine are neither assimilated nor alkalinized. Using ID 32 GN strips, propionate, valerate, 3-hydroxybutyrate, L-proline, itaconate, suberate, acetate and lactate are assimilated. Utilization of caprate and 4-hydroxybenzoate is variable. 3-Hydroxybenzoate and L-alanine are not utilized. Alkaline phosphatase, trypsin and pyrrolidone arylamidase are negative. Strains are susceptible to desferrioxamine and colistin. The main cellular fatty acids are C16 : 1, C16 : 0, C18 : 19c, C10 : 0 3-OH, C12 : 0 and C14 : 0. The DNA G+C content is 62·8–66·7 mol%.
The type strain is LMG 2370T (=CCUG 15845T=ATCC 11330T). Its DNA G+C content is 64·0 mol%. The sequence of its 16S rDNA is available under EMBL/GenBank accession number AJ430344.
Description of Comamonas kerstersii sp. nov.
Comamonas kerstersii (ker.ster'si.i. N.L. masc. gen. n. kerstersii of Kersters, in honour of Karel Kersters, a Belgian microbiologist, for his contribution to bacterial taxonomy).
Gram-negative rods, motile by multitrichous polar flagella, often with bipolar tufts. Grows aerobically on tryptic soy agar and broth at 30, 35, 42 and 44 °C. Optimal growth occurs at 35–40 °C. After 24 h incubation at 30 °C, colonies reach a diameter of 1·5 mm on blood agar, and a diameter of more than 2 mm may be reached at 40 °C. Oxidase and catalase activities are positive. Carbohydrates are neither acidified nor assimilated. Ethylene glycol is not acidified. Urease, indole and H2S are not produced; negative for lysine decarboxylase, ornithine decarboxylase and arginine dihydrolase. Nitrates are reduced but nitrites are not. Simmons' citrate agar test may be positive, but irregular and delayed. Tyrosine hydrolysis is positive but gelatin hydrolysis is negative. On Simmons' minimal agar base, L-alanine and glycine are neither assimilated nor alkalinized. Using ID 32 GN strips, valerate, 3-hydroxybutyrate, L-proline, itaconate, suberate, acetate, lactate and 4-hydroxybenzoate are assimilated. Utilization of caprate is variable. L-Alanine and 3-hydroxybenzoate are negative. Alkaline phosphatase, benzylarginine arylamidase and pyrrolidone arylamidase are negative. Strains are susceptible to desferrioxamine and colistin. The main cellular fatty acids are C16 : 1, C16 : 0, C18 : 19c, C10 : 0 3-OH, C12 : 0 and C14 : 0. DNA G+C content is 59·7–63·3 mol%.
The type strain is LMG 3475T (=CCUG 15333T=AF61T). Its DNA G+C content is 61·0 mol%. The sequence of its 16S rDNA is available under EMBL/GenBank accession number AJ430347.
Emended description of Comamonas terrigena sensu stricto (ex Hugh 1962) De Vos et al. 1985
Gram-negative rods, motile by multitrichous polar flagella, often with bipolar tufts. Grows aerobically on tryptic soy agar and broth at 30 and 35 °C, but not at 40 °C. After 24 h incubation at 30 °C, colonies reach a diameter of 1 mm on blood agar. Oxidase and catalase activities are positive. Carbohydrates are neither acidified nor assimilated. Ethylene glycol is not acidified. Urease, indole and H2S are not produced; negative for lysine decarboxylase, ornithine decarboxylase and arginine dihydrolase. Nitrates are reduced but nitrites are not. Simmons' citrate agar test is positive, but may be delayed. Tyrosine hydrolysis is positive but gelatin hydrolysis is negative. On Simmons' minimal agar base, L-alanine is assimilated and alkalinized but glycine is not. Using ID 32 GN strips, propionate, valerate, 3-hydroxybutyrate, L-proline, suberate, acetate, lactate and L-alanine are assimilated. Utilization of itaconate and caprate is variable. 3-Hydroxybenzoate and 4-hydroxybenzoate are always negative. Alkaline phosphatase and benzylarginine arylamidase activities are negative but pyrrolidone arylamidase is positive. Strains are susceptible to desferrioxamine and colistin. The main cellular fatty acids are C16 : 1, C16 : 0, C18 : 19c, C10 : 0 3-OH, C12 : 0 and C14 : 0. DNA G+C content is 64·0–66·6 mol%.
The type strain is LMG 1253T (=CCUG 15327T=NCIB 8193T). Its DNA G+C content is 64·0 mol%. The sequence of its 16S rDNA is available under EMBL/GenBank accession number AJ430342.
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ACKNOWLEDGEMENTS |
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REFERENCES |
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Baele, M., Storms, V., Haesebrouck, F., Devriese, L. A., Gillis, M., Verschraegen, G., de Baere, T. & Vaneechoutte, M. (2001). Application and evaluation of the interlaboratory reproducibility of tRNA intergenic length polymorphism analysis (tDNA-PCR) for identification of Streptococcus species. J Clin Microbiol 39, 1436–1442.
Chang, Y.-H., Han, J., Chun, J., Lee, K. C., Rhee, M.-S., Kim, Y.-B. & Bae, K. S. (2002). Comamonas koreensis sp. nov., a non-motile species from wetland in Woopo, Korea. Int J Syst Evol Microbiol 52, 377–381.[Abstract]
De Vos, P., Kersters, K., Falsen, E., Pot, B., Gillis, M., Segers, P. & De Ley, J. (1985). Comamonas Davis and Park 1962 gen. nov., nom. rev. emend., and Comamonas terrigena Hugh 1962 sp. nov., nom. rev. Int J Syst Bacteriol 35, 443–453.
Etchebehere, C., Errazquin, M. I., Dabert, P., Moletta, R. & Muxí, L. (2001). Comamonas nitrativorans sp. nov., a novel denitrifier isolated from a denitrifying reactor treating landfill leachate. Int J Syst Evol Microbiol 51, 977–983.[Abstract]
Gilligan, P. H. & Whittier, S. (1999). Burkholderia, Xenotrophomonas, Ralstonia, Brevundimonas, Comamonas and Acidovorax. In Manual of Clinical Microbiology, 7th edn, pp. 526–538. Edited by P. R. Murray, E. J. Baron, M. A. Pfaller, F. C. Tenover & R. H. Yolken. Washington, DC: American Society for Microbiology.
Gumaelius, L., Magnusson, G., Pettersson, B. & Dalhammar, G. (2001). Comamonas denitrificans sp. nov., an efficient denitrifying bacterium isolated from activated sludge. Int J Syst Evol Microbiol 51, 999–1006.[Abstract]
Laffineur, K., Janssens, M., Charlier, J., Avesani, V., Wauters, G., & Delmée, M. (2002). Biochemical and susceptibility tests useful for identification of nonfermenting gram-negative rods. J Clin Microbiol 40, 1085–1087.
Lindsay, J. A. & Riley, T. V. (1991). Susceptibility to desferrioxamine: a new test for the identification of Staphylococcus epidermidis. J Med Microbiol 35, 45–48.[Abstract]
Nemec, A., De Baere, T., Tjernberg, I., Vaneechoutte, M., van der Reijden, T. J. K. & Dijkshoorn, L. (2001). Acinetobacter ursingii sp. nov. and Acinetobacter schindleri sp. nov., isolated from human clinical specimens. Int J Syst Evol Microbiol 51, 1891–1899.[Abstract]
Vaneechoutte, M., Rossau, R., De Vos, P. & 7 other authors (1992). Rapid identification of bacteria of the Comamonadaceae with amplified ribosomal DNA-restriction analysis (ARDRA). FEMS Microbiol Lett 93, 227–233.[CrossRef]
Vaneechoutte, M., Boerlin, P., Tichy, H.-V., Bannerman, E., Jäger, B. & Bille, J. (1998). Comparison of PCR-based DNA fingerprinting techniques for the identification of Listeria species and their use for atypical Listeria isolates. Int J Syst Bacteriol 48, 127–139.
Vaneechoutte, M., Claeys, G., Steyaert, S., De Baere, T., Peleman, R. & Verschraegen, G. (2000). Isolation of Moraxella canis from an ulcerated metastatic lymph node. J Clin Microbiol 38, 3870–3871.
Welsh, J. & McClelland, M. (1991). Genomic fingerprints produced by PCR with consensus tRNA gene primers. Nucleic Acids Res 19, 861–866.
Willems, A., Pot, B., Falsen, E., Vandamme, P., Gillis, M., Kersters, K. & De Ley, J. (1991). Polyphasic taxonomic study of the emended genus Comamonas: relationship to Aquaspirillum aquaticum, E. Falsen group 10, and other clinical isolates. Int J Syst Bacteriol 41, 427–444.
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