Archaeal and bacterial diversity in an arsenic-rich shallow-Sea hydrothermal system undergoing phase separation

Frontiers in Microbiology

Volumn 4, Issue JUL, 2013, Pages

  Price, Roy E ^a   Lesniewski, Ryan ^a   Nitzsche, Katja S ^b,c   Meyerdierks, Anke ^b   Saltikov, Chad ^d   Pichler, Thomas ^e   Amend, Jan P ^a  

^a UNIVERSITY OF SOUTHERN CALIFORNIA   (United States)

^b MAX PLANCK INSTITUTE FOR MARINE MICROBIOLOGY   (Germany)

^c UNIVERSITY OF TÜBINGEN   (Germany)

^d UNIVERSITY OF CALIFORNIA   (United States)

^e UNIVERSITY OF BREMEN   (Germany)

Author keywords

Arsenic; Gradients; Hydrothermal; Microbial diversity; Milos; Phase separation

Indexed keywords

ARSENIC; RNA 16S;

ARCHAEOGLOBALES; ARCHAEON; ARTICLE; BACTERIAL METABOLISM; BACTEROIDETES; CONTROLLED STUDY; DESULFUROCOCCALES; EPSILONPROTEOBACTERIA; EURYARCHAEOTA; FIRMICUTES; GENE LIBRARY; GEOCHEMISTRY; GEOGRAPHIC DISTRIBUTION; HYDROTHERMAL VENT; ISLAND (GEOLOGICAL); MICROBIAL DIVERSITY; NONHUMAN; NUCLEOTIDE SEQUENCE; PH; PHYLOGENETIC TREE; POLYMERASE CHAIN REACTION; SALINITY; SEA; SEQUENCE HOMOLOGY; SPECIES DIVERSITY; TEMPERATURE; THERMOCOCCALES; THERMOPLASMATALES; THERMOPROTEALES;

ARCHAEA; ARCHAEOGLOBALES; ARCOBACTER SP. SOLAR LAKE; BACILLUS SP.; BACTERIA (MICROORGANISMS); BACTEROIDETES; BETAPROTEOBACTERIA; DESULFUROCOCCALES; EPSILONPROTEOBACTERIA; EURYARCHAEOTA; FIRMICUTES; FLAVOBACTERIA; PROKARYOTA; STAPHYLOTHERMUS; THERMOCOCCALES; THERMOPLASMATALES; THERMOPROTEALES;

EID: 84885346780     PISSN: None     EISSN: 1664302X     Source Type: Journal    
DOI: 10.3389/fmicb.2013.00158     Document Type: Article

References (76)

1. Akerman, N. (2009). Microbial diversity and geochemical energy sources of Tutum Bay, Ambitle Island, Papua New Guinea, an arsenic-rich shallow-sea hydrothermal system. Ph. D. dissertation, Washington University in St. Louis.
2. Akerman, N., Price, R., Pichler, T., and Amend, J. P. (2011). Energy sources for chemlithotrophs in an arsenic- and iron-rich shallow-sea hydrothermal system. Geobiology 9, 436-445.
3. Alonso, C., Warnecke, F., Amann, R., and Pernthaler, J. (2007). High local and global diversity of Flavobacteria in marine plankton. Environ. Microbiol. 9, 1253-1266. doi: 10.1111/j.1462-2920.2007.01244.x
4. Amend, J. P., Rogers, K. L., Shock, E. L., Gurrieri, S., and Inguaggiato, S. (2003). Energetics of chemolithoautotrophy in the hydrothermal system of Vulcano Island, southern Italy. Geobiology 1, 37-58. doi: 10.1046/j.1472-4669.2003.00006.x
5. Arab, H., Voelker, H., and Thomm, M. (2000). Thermococcus aegaeicus sp. nov. and Staphylothermus hellenicus sp. nov., two novel hyperthermophilic archaea isolated from geothermally heated vents offPalaeochori Bay, Milos, Greece. Int. J. Syst. Evol. Microbiol. 50, 2101-2108. doi: 10.1099/00207713-50-6-2101
6. Bayraktarov, E., Price, R. E., Ferdelman, T. G., and Finster, K. (2013). The pH and pCO2 dependence of sulfate reduction in shallow-sea hydrothermal CO2-venting sediments (Milos Island, Greece). Front. Microbiol. 4, 1-10. doi: 10.3389/fmicb.2013.00111
7. Brinkhoff, T., Sievert, S., Kuever, J., and Muyzer, G. (1999). Distribution and diversity of sulfur-oxidizing Thiomicrospira spp. at a shallow-water hydrothermal vent in the Aegean Sea (Milos, Greece). Appl. Environ. Microbiol. 65, 3843-3849.
8. Burggraf, S., Larsen, N., Woese, C. R., and Stetter, K. (1993). An intron within the 16S ribosomal RNA gene of the archaeon Pyrobaculum aerophilum. Proc. Natl. Acad. Sci. U.S.A. 90, 2547-2550. doi: 10.1073/pnas.90.6.2547
9. Cavalca, L., Corsini, A., Zaccheo, P., Andreoni, V., and Muyzer, G. (2013). Microbial transformations of arsenic: perspectives for biological removal of arsenic from water. Future Microbiol. 8, 753-768. doi: 10.2217/fmb.13.38
10. Cox, M. E., Launay, J., and Paris, J. P. (1982). "Geochemistry of low temperature geothermal systems in New Caledonia," in Pacific Geothermal Conference, (Auckland: University of Auckland), 453-459.
11. Dando, P. R., Thomm, M., Arab, H., Brehmer, M., Hooper, L. E., Jochimsen, B., et al. (1998). Microbiology of shallow hydrothermal sites offPalaeochori Bay, Milos (Hellenic Volcanic Arc). Cah. Biol. Mar. 39, 369-372.
12. Dando, P. R., Hughes, J. A., Leahy, Y., Niven, S. J., Taylor, L. J., and Smith, C. (1995). Gas venting rates from the submarine hydrothermal areas around the island of Milos, Hellenic volcanic arc. Cont. Shelf Res. 15, 913-929. doi: 10.1016/0278-4343(95)80002-U
13. Dando, P. R., Stüben, D., and Varnavas, S. P. (1999). Hydrothermalism in the Mediterranean Sea. Prog. Oceanogr. 44, 333-367. doi: 10.1016/S0079-6611(99)00032-4
14. Dick, G. J., Lee, Y. E., and Tebo, B. M. (2006). Manganese(II)-oxidizing Bacillus spores in Guaymas Basin hydrothermal sediments and plumes. Appl. Environ. Microbiol. 72, 3184-3190. doi: 10.1128/AEM.72.5.3184-3190.2006
15. Donachie, S. P., Bowman, J. P., On, S. L. W., and Alam, M. (2005). Arcobacter halophilus sp. nov., the first obligate halophile in the genus Arcobacter. Int. J. Syst. Evol. Microbiol. 55, 1271-1277. doi: 10.1099/ijs.0.63581-0
16. Flores, G. E., Campbell, J. H., Kirshtein, J. D., Meneghin, J., Podar, M., Steinberg, J. I., et al. (2011). Microbial community structure of hydrothermal deposits from geochemically different vent fields along the Mid-Atlantic Ridge. Environ. Microbiol. 13, 2158-2171. doi: 10.1111/j.1462-2920.2011.02463.x
17. Flores, G. E., Wagner, I. D., Liu, Y., and Reysenbach, A. L. (2012). Distribution, abundance, and diversity patterns of the thermoacidophilic "deep-sea hydrothermal vent euryarchaeota 2". Front. Microbiol. 3:47. doi: 10.3389/fmicb.2012.00047
18. Forrest, M. J., Ledesma-Vazquez, J., Ussler, I, W., Kulongoski, J. T., Hilton, D. R., and Greene, H. G. (2005). Gas geochemistry of a shallow submarine hydrothermal vent associated with the El Requeson fault zone, Bahia Concepcion, Baja California Sur, Mexico. Chem. Geol. 224, 82-95. doi: 10.1016/j.chemgeo.2005.07.015
19. German, C. R., and Von Damm, K. L. (2003). "Hydrothermal processes," in Treatise on Geochemistry, eds H. D. Holland and K. K. Turekian (Amsterdam: Elsevier), 145-180.
20. Graumann, P. (2012). Bacillus: Cellular and Molecular Biology. (Norfolk: Caister Academic Press).
21. Guindon, S., and Gascuel, O. (2003). A simple, fast, and accurate algorithm to estimate large phylogenies by maximum likelihood. Syst. Biol. 52, 696-704. doi: 10.1080/10635150390235520
22. Handley, K. M., Hery, M., and Lloyd, J. R. (2009). Redox cycling of arsenic by the hydrothermal marine bacterium Marinobacter santoriniensis. Environ. Microbiol. 11, 1601-1611. doi: 10.1111/j.1462-2920.2009.01890.x
23. Huber, T., Faulkner, G., and Hugenholtz, P. (2004). Bellerophon; a program to detect chimeric sequences in multiple sequence alignments. Bioinformatics 20, 2317-2319. doi: 10.1093/bioinformatics/bth226
24. Italiano, F., Bonfanti, P., Ditta, M., Petrini, R., and Slejko, F. (2009). Helium and carbon isotopes in the dissolved gases of Friuli Region (NE Italy): geochemical evidence of CO2 production and degassing over a seismically active area. Chem. Geol. 266, 76-85. doi: 10.1016/j.chemgeo.2009.05.022
25. Italiano, F., and Nuccio, F. (1991). Geochemical investigations of submarine volcanic exhalations to the east of Panarea, Aeolian Islands, Italy. J. Volcanol. Geother. Res. 46, 125-141. doi: 10.1016/0377-0273(91)90079-F
26. Itoh, T., Nomura, N., and Sako, Y. (2003). Distribution of 16S rRNA introns among the family Thermoproteaceae and their evolutionary implications. Extremophiles 7, 229-233.
27. Itoh, T., Yoshikawa, N., and Takashina, T. (2007). Thermogymnomonas acidicola gen. nov., sp. nov., a novel thermoacidophilic, cell wall-less archaeon in the order Thermoplasmatales, isolated from a solfataric soil in Hakone, Japan. Int. J. Syst. Evol. Microbiol. 57, 2557-2561. doi: 10.1099/ijs.0.65203-0
28. Jochimsen, B., Peinemann-Simon, S., Volker, H., Stuben, D., Botz, R., Stoffers, P., et al. (1997). Stetteria hydrogenophila gen. nov. and sp. nov., a novel mixotrophic sulfur-dependent crenarchaeote isolated from Milos, Greece. Extremophiles 1, 67-73. doi: 10.1007/s007920050016
29. Kirchman, D. L. (2002). The ecology of Cytophata-Flavobacteria in aquatic environments. FEMS Microbiol. Ecol. 39, 91-100. doi: 10.1016/S0168-6496(01)00206-9
30. Kormas, K. A., Tivey, M. K., Von Damm, K. L., and Teske, A. (2006). Bacterial and archaeal phylotypes associated with distinct mineralogical layers of a white smoker spire from a deep-sea hydrothermal vent site (9°N, East Pacific Rise). Environ. Microbiol. 8, 909-920. doi: 10.1111/j.1462-2920.2005.00978.x
31. Koski, R. A., Pichler, T., and Foster, A. L. (2001). "The fate of arsenic in submarine hydrothermal environments: a summary of recent data," in USGS Workshop on Arsenic in the Environment (Denver, CO).
32. Lane, D. J. (1991). "16S/23S rRNA sequencing," in Nucleic Acid Techniques in Bacterial Systematics, eds E. Stackebrandt and M. Goodfellow (Chichester: John Wiley and Sons), 115-175.
33. Li, S.-G., Tang, Y.-Q., Nie, Y., Cai, M., and Wu, X.-L. (2011). Complete genome sequence of Polymorphum gilvum SL003B-26A1T, a crude oil-degrading bacterium from oil polluted saline soil. J. Bacteriol. 193, 2894-2895. doi: 10.1128/JB.00333-11
34. Maugeri, T. L., Gugliandolo, C., Caccamo, D., Panico, A., Lama, L., Gambacorta, A., et al. (2002). A halophilic thermotolerant Bacillus isolated from a marine hot spring able to produce a new exopolysaccharide. Biotechnol. Lett. 24, 515-519. doi: 10.1023/A:1014891431233
35. McCarthy, K. T., Pichler, T., and Price, R. E. (2005). Geochemistry of Champagne Hot Springs shallow hydrothermal vent field and associated sediments, Dominica, Lesser Antilles. Chem. Geol. 224, 55-68. doi: 10.1016/j.chemgeo.2005.07.014
36. Meyer-Dombard, D., Amend, J. P., and Osburn, M. R. (2013). Microbial diversity and potential for arsenic and iron biogeochemical cycling at an arsenic-rich, shallow-sea hydrothermal vent (Tutum Bay, Papua New Guinea). Chem. Geol. 348, 37-47. doi: 10.1016/j.chemgeo.2012.02.024
37. Meyer-Dombard, D. R., Price, R. E., Pichler, T., and Amend, J. P. (2011). Prokaryotic populations in arsenic-rich shallow-sea hydrothermal sediments of Ambitle Island, Papua New Guinea. Geomicrobiol. J. 29, 1-17.
38. Morinaga, Y., Nomura, N., and Sako, Y. (2002). Population dynamics of archaeal mobile introns in natural environments: a shrewd invasion strategy of the latent parasitic DNA. Microbes Environ. 17, 153-163. doi: 10.1264/jsme2.17.153
39. Nakagawa, S., Takai, K., Inagaki, F., Chiba, H., Ishibashi, J.-I., Kataoka, S., et al. (2005). Variability in microbial community and venting chemistry in a sediment-hosted backarc hydrothermal system: impacts of subseafloor phase-separation. FEMS Microbiol. Ecol. 54, 141-155. doi: 10.1016/j.femsec.2005.03.007
40. Nakano, M. M., and Zuber, P. (1998). Anaerobic growth of a "strict aerobe" (Bacillus subtilis). Annu. Rev. Microbiol. 52, 165-190. doi: 10.1146/annurev.micro.52.1.165
41. Nitzsche, K. (2010). Microbial Diversity of Hydrothermally Influenced Arsenic-Rich Sediments offthe Coast of Milos Island, Greece. Diploma thesis, TU Bergakademie, Freiberg, Germany.
42. Nunoura, T., Oida, H., Miyazaki, M., and Suzuki, Y. (2008). Thermosulfidibacter takaii gen. nov., sp. nov., a thermophilic, hydrogen-oxidizing, sulfur-reducing chemolithoautotroph isolated from a deep-sea hydrothermal field in the Southern Okinawa Trough. Int. J. Syst. Evol. Microbiol. 58, 659-665. doi: 10.1099/ijs.0.64615-0
43. Nunoura, T., and Takai, K. (2009). Comparison of microbial communities associated with phase-separation-induced hydrothermal fluids at the Yonaguni Knoll IV hydrothermal field, the Southern Okinawa Trough. FEMS Microbiol. Ecol. 67, 351-370. doi: 10.1111/j.1574-6941.2008.00636.x
44. Oremland, R. S., and Stolz, J. F. (2003). The ecology of arsenic. Science 300, 939-944. doi: 10.1126/science.1081903
45. Pati, A., Gronow, S., Lapidus, A., Copeland, A., Glavina Del Rio, T., Nolan, M., et al. (2010). Complete genome sequence of Arcobacter nitrofigilis type strain (CI). Stand. Genomic. Sci. 2, 300-308. doi: 10.4056/sigs.912121
46. Pichler, T. (2005). Stable and radiogenic isotopes as tracers for the origin, mixing and subsurface history of fluids in submarine shallow-water hydrothermal systems. J. Volcanol. Geother. Res. 139, 211-226. doi: 10.1016/j.jvolgeores.2004.08.007
47. Pichler, T., Amend, J. P., Garey, J., Hallock, P., Hsia, N. P., Karlen, D. J., et al. (2006). A natural laboratory to study arsenic geobiocomplexity. EOS 87, 221-225. doi: 10.1029/2006EO230002
48. Pichler, T., Giggenbach, W. F., McInnes, B. I. A., and Duck, B. (1999). Fe-sulfide formation due to seawater-gas-sediment interaction in a shallow-water hydrothermal system, Lihir Island, Papua New Guinea. Econ. Geol. 94, 281-288. doi: 10.2113/gsecongeo.94.2.281
49. Pokrovski, G. S., Zakirov, I. V., Roux, J., Testemale, D., Hazemann, J. L., Bychkov, A., et al. (2002). Experimental study of arsenic speciation in vapor phase to 500°C: implications for As transport and fractionation in low-density crustal fluids and volcanic gases. Geochim. Cosmochim. Acta 66, 3453-3480. doi: 10.1016/S0016-7037(02)00946-8
50. Polz, M. F., and Cavanaugh, C. M. (1995). Dominance of one bacterial phylotype at a Mid-Atlantic ridge hydrothermal vent site. Proc. Natl. Acad. Sci. U.S.A. 92, 7232-7236. doi: 10.1073/pnas.92.16.7232
51. Price, R. E., Amend, J. P., and Pichler, T. (2007). Enhanced geochemical gradients in a marine shallow-water hydrothermal system: unusual arsenic speciation in horizontal and vertical pore water profiles. Appl. Geochem. 22, 2595-2605. doi: 10.1016/j.apgeochem.2007.06.010
52. Price, R. E., Savov, I., Planer-Friedrich, B., Bühring, S., Amend, J. P., and Pichler, T. (2012). Processes influencing extreme As enrichment in shallow-sea hydrothermal fluids of Milos Island, Greece. Chem. Geol. 348, 15-26. doi: 10.1016/j.chemgeo.2012.06.007
53. Reysenbach, A. L., Liu, Y., Banta, A. B., Beveridge, T. J., Kirshtein, J. D., Schouten, S., et al. (2006). A ubiquitous thermoacidophilic archaeon from deep-sea hydrothermal vents. Nat. Lett. 442, 444-447. doi: 10.1038/nature04921
54. Rogers, K. L., and Amend, J. P. (2005). Archaeal diversity and geochemical energy yields in a geothermal well on Vulcano Island, Italy. Geobiology 3, 319-332. doi: 10.1111/j.1472-4669.2006.00064.x
55. Schlesner, H., Lawson, P. A., Collins, M. D., Weiss, N., Wehmeyer, U., Voelker, H., et al. (2001). Filobacillus milensis gen. nov., sp. nov., a new halophilic spore-forming bacterium with Orn-D-Glu-type peptidoglycan. Int. J. Syst. Evol. Microbiol. 51, 425-431.
56. Schloss, P. D., Westcott, S. L., Ryabin, T., Hall, J. R., Hartmann, M., Hollister, E. B., et al. (2009). Introducing mothur: open-source, platform-independent, community-supported software for describing and comparing microbial communities. Appl. Environ. Microbiol. 75, 7537-7541. doi: 10.1128/AEM.01541-09
57. Schrenk, M. O., Kelley, D. S., Bolton, S. A., and Baross, J. A. (2004). Low archaeal diversity linked to subseafloor geochemical processes at the Lost City hydrothermal field, Mid-Atlantic Ridge. Environ. Microbiol. 6, 1086-1095. doi: 10.1111/j.1462-2920.2004.00650.x
58. Seeberg-Elverfeldt, J., Schlüter, M., Feseker, T., and Kölling, M. (2005). Rhizon sampling of pore waters near the sediment/water interface of aquatic systems. Limnol. Oceanogr. Methods 3, 361-371. doi: 10.4319/lom.2005.3.361
59. Sievert, S. M., Brinkhoff, T., Muyzer, G., Ziebis, W., and Kuever, J. (1999). Spatial heterogeneity of bacterial population along an environmental gradient at a shallow submarine hydrothermal vent near Milos Island, Greece. Appl. Environ. Microbiol. 65, 3834-3842.
60. Sievert, S. M., Heidorn, T., and Kuever, J. (2000a). Halothiobacillus kellyi sp. nov., a mesophilic obligately chemolithoautotrophic sulfur-oxidizing bacterium isolated from a shallow-water hydrothermal vent in the Aegean Sea and emended description of the genus Halothiobacillus. Int. J. Syst. Evol. Microbiol. 50, 1229-1237. doi: 10.1099/00207713-50-3-1229
61. Sievert, S. M., Kuever, J., and Muyzer, G. (2000b). Identification of 16S ribosomal DNA-defined bacterial populations at a shallow submarine hydrothermal vent near Milos Island (Greece). Appl. Environ. Microbiol. 66, 3102-3109. doi: 10.1128/AEM.66.7.3102-3109.2000
62. Sievert, S. M., and Kuever, J. (2000). Desulfacinum hydrothermale sp. nov., a thermophilic sulfate-reducing bacterium from geothermally heated sediments near Milos island, Greece. Int. J. Syst. Evol. Microbiol. 50, 1239-1246. doi: 10.1099/00207713-50-3-1239
63. Spano, A., Gugliandolo, C., Lentini, V., Maugeri, T. L., Anzelmo, G., Annarita, P., et al. (2013). A novel EPS-producing strain of Bacillus licheniformis isolated from a shallow vent offPanarea Island (Italy). Curr. Microbiol. 67, 21-29. doi: 10.1007/s00284-013-0327-4
64. Stüben, D., and Glasby, G. P. (1999). Geochemistry of shallow submarine hydrothermal fluids of Paleochori Bay, Milos, Aegean Sea. Explor. Mining Geol. 8, 273-287.
65. Switzer, B. J., Burns, B. A., Buzzelli, J., Stolz, J. F., and Oremland, R. S. (1998). Bacillus arsenicoselenatis sp. nov., and Bacillus selentireducens sp. nov.: two haloalkaliphilies from Mono Lake, California that respire oxyanions of selenium and arsenic. Arch. Microbiol. 171, 19-30. doi: 10.1007/s002030050673
66. Sylvan, J. B., Toner, B. M., and Edwards, K. J. (2012). Life and death of deep-sea vents: bacterial diversity and ecosystem succession on inactive hydrothermal sulfides. mBio 3, 1-10. doi: 10.1128/mBio.00279-11
67. Takai, K., and Horikoshi, K. (1999). Genetic diversity of archaea in deep-sea hydrothermal vent environments. Genetics 152, 1285-1297.
68. Tarasov, V. G., Gebruk, A. V., Mironov, A. N., and Moskalev, L. I. (2005). Deep-sea and shallow-water hydrothermal vent communities: two different phenomena? Chem. Geol. 224, 5-39. doi: 10.1016/j.chemgeo.2005.07.021
69. Tarasov, V. G., Propp, M. V., Propp, L. N., Zhirmunsky, A. V., Namsaraev, B. B., Gorlenko, V. M., et al. (1990). Shallow-water gasohydrothermal vents of Ushishir Volcano and the ecosystem of Kraternaya Bight (The Kurile Islands). Mar. Ecol. 11, 1-23. doi: 10.1111/j.1439-0485.1990.tb00225.x
70. Valsami-Jones, E., Baltatzis, E., Bailey, E. H., Boyce, A. J., Alexander, J. L., Magganas, A., et al. (2005). The geochemistry of fluids from an active shallow submarine hydrothermal system: Milos island, Hellenic Volcanic Arc. J. Volcanol. Geothermal Res. 148, 130-151. doi: 10.1016/j.jvolgeores.2005.03.018
71. Varnavas, S. P., and Cronan, D. S. (1988). Arsenic, antimony and bismuth in sediments and waters from the Santorini hydrothermal field, Greece. Chem. Geol. 67, 295-305. doi: 10.1016/0009-2541(88)90135-0
72. Von Damm, K. L., Lilley, M. D., Shanks, W. C., Brockington, M., Bray, A. M., O'Grady, K. M., et al. (2003). Extraordinary phase separation and segregation in vent fluids from the southern East Pacific Rise. Earth Planet. Sci. Lett. 206, 365-378. doi: 10.1016/S0012-821X(02)01081-6
73. Wang, Q., Garrity, G. M., Tiedje, J. M., and Cole, J. R. (2007). Naive Bayesian classifier for rapid assignment of rRNA sequences into the new bacterial taxonomy. Appl. Environ. Microbiol. 73, 5261-5267. doi: 10.1128/AEM.00062-07
74. Wirsen, C. (2004). Is life thriving deep beneath the seafloor? Oceanus 42, 1-6. Available online at: http://oceanusmag.whoi.edu/v42n2/wirsen.html
75. Zargar, K., Conrad, A., Bernick, D. L., Lowe, T. M., Stolc, V., Hoeft, S., et al. (2012). ArxA, a new clade of arsenite oxidase within the DMSO reductase family of molybdenum oxidoreductases. Environ. Microbiol. 14, 1635-1645. doi: 10.1111/j.1462-2920.2012.02722.x
76. Zargar, K., Hoeft, S. E., Oremland, R. S., and Saltikov, C. (2010). Identification of a novel arsenite oxidase gene, arxA, in the haloalkaliphilic, arsenite-oxidizing bacterium Alkalilimnicola ehrlichii strain MLHE-1. J. Bacteriol. 192, 3755-3762. doi: 10.1128/JB.00244-10