Distribution and abundance of aerobic anoxygenic phototrophic bacteria in the tropical coastal waters of Gunungkidul, Yogyakarta, Indonesia
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Abstract. Aprilyanto V, Sembiring L, Djohan TS. 2020. Distribution and abundance of aerobic anoxygenic phototrophic bacteria in the tropical coastal waters of Gunungkidul, Yogyakarta, Indonesia. Biodiversitas 21: 5506-5513. Aerobic anoxygenic phototroph (AAP) is a community of bacteria capable of performing anoxygenic photosynthesis in the presence of oxygen. AAP abundance in most oceanic regions signals its ecological importance in marine microbial loop. This study was conducted to reveal the distribution and abundance of AAP in Siung coast, Gunungkidul, Yogyakarta, Indonesia. Total bacterioplankton and AAP were enumerated using acridine orange direct count (AODC) and real-time quantitative PCR (qPCR), respectively. Several water physicochemical parameters as well as dissolved nitrate, ammonium, phosphate, and sulfate were also measured. The results showed that total bacterioplankton was distributed thoroughly in the water column with abundance ranging from 2.5×104 to 3.5×104 cells/mL. Within this total bacterioplankton, AAP ranged from 3.83×102 –7.48×102 cells/mL, comprising about 1% to ~2.5%. Inorganic nutrient concentration, mainly nitrate and phosphate were thought to be the regulating factors to the distribution and abundance of both communities. In conclusion, AAP comprises a relatively low portion of the total bacterioplankton community in the tropical coastal water in Gunungkidul, Yogyakarta.
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References
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Béjà O, Suzuki MT, Heidelberg JF, Nelson WC, Preston CM, Hamada T, Eisen JA, Fraser CM, DeLong EF. 2002. Unsuspected diversity among marine aerobic anoxygenic phototrophs. Nature 415: 630–633.
Biebl H, Allgaier M, Tindall BJ, Koblizek M, Lünsdorf H, Pukall R, Wagner-Döbler I. 2005. Dinoroseobacter shibae gen. nov., sp. nov., a new aerobic phototrophic bacterium isolated from dinoflagellates. International Journal of Systematic and Evolutionary Microbiology 55: 1089–1096.
Biebl H, Pukall R, Lünsdorf H, Schulz S, Allgaier M, Tindall BJ, Wagner-Döbler I. 2007. Description of Labrenzia alexandrii gen. nov., sp. nov., a novel alphaproteobacterium containing bacteriochlorophyll a, and a proposal reclassification of Stappia aggregata as Labrenzia aggregata comb. nov., and of Stappia alba as Labrenzia alba comb. nov. International Journal of Systematic and Evolutionary Microbiology 57: 1095–1107.
Boeuf D, Cottrell MT, Kirchman DL, Lebaron P, Jeanthon C. 2013. Summer community structure of aerobic anoxygenic phototrophic bacteria in the western Arctic Ocean. FEMS Microbiology Ecology 85: 417–432.
Brenner DJ, Krieg NR, Staley JT, Garrity M. 2005. 2 Bergey’s manual of systematic bacteriology. 2nd ed. Springer.
Calbet A, Landry MR, Nunnery S. 2001. Bacteria-flagellate interactions in the microbial food web of the oligotrophic subtropical North Pacific. Aquatic Microbial Ecology 23: 283–292.
Church MJ. 2008. Resource control of bacterial dynamics in the sea, pp. 353–354 In Kirchman DL [ed.], Microbial Ecology of the Oceans.
Church MJ, Hutchins DA, Ducklow HW. 2000. Limitation of bacterial growth by dissolved organic matter and iron in the Southern Ocean. Applied and Environmental Microbiology 66: 455–466.
Cottrell MT, Mannino A, Kirchman DL. 2006. Aerobic anoxygenic phototrophic bacteria in the mid-atlantic bight and the north pacific gyre. Applied and Environmental Microbiology 72: 557–564.
Csotonyi JT, Stackebrandt E, Swiderski J, Schumann P, Yurkov V. 2011. Chromocurvus halotolerans gen. nov., sp. nov., a gammaproteobacterial obligately aerobic anoxygenic phototroph, isolated from a Canadian hypersaline spring. Archives of Microbiology 193: 573–582.
Daley RJ, Hobbie JE. 1975. Direct counts of aquatic bacteria by a modified epifluorescence technique. Limnology and Oceanography 20: 875–882.
Du H, Jiao N, Hu Y, Zeng Y. 2006. Real-time PCR for quantification of aerobic anoxygenic phototrophic bacteria based on pufM gene in marine environment. Journal of Experimental Marine Biology and Ecology 329: 113–121.
Epstein SS, Shiaris MP. 1992. Size-selective grazing of coastal bacterioplankton by natural assemblages of pigmented flagellates, colorless flagellates, and ciliates. Microbial Ecology 23: 211–225.
Ferrera I, Gasol JM, Sebastián M, Hojerová E, Kobížek M. 2011. Comparison of growth rates of aerobic anoxygenic phototrophic bacteria and other bacterioplankton groups in coastal mediterranean waters. Applied and Environmental Microbiology 77: 7451–7458.
Ferrera I, Sánchez O, Kolá?ová E, Koblížek M, Gasol JM. 2017. Light enhances the growth rates of natural populations of aerobic anoxygenic phototrophic bacteria. ISME Journal 11: 2391–2393.
Garcia-Chaves MC, Cottrell MT, Kirchman DL, Derry AM, Bogard MJ, Del Giorgio PA. 2015. Major contribution of both zooplankton and protists to the top-down regulation of freshwater aerobic anoxygenic phototrophic bacteria. Aquatic Microbial Ecology 76: 71–83.
Gonzalez JM, Sherr EB, Sherr BF. 1990. Size-selective grazing on bacteria by natural assemblages of estuarine flagellates and ciliates. Applied and Environmental Microbiology 56: 583–589.
Hahn MW, Höfle MG. 2001. Grazing of protozoa and its effect on populations of aquatic bacteria. FEMS Microbiology Ecology 35: 113–121.
Hobbie JE, Daley RJ, Jasper S. 1977. Use of nuclepore filters for counting bacteria by fluorescence microscopy. Applied and Environmental Microbiology 33: 1225–1228.
Hojerová E, Mašín M, Brunet C, Ferrera I, Gasol JM, Koblížek M. 2011. Distribution and Growth of Aerobic Anoxygenic Phototrophs in the Mediterranean Sea. Environmental Microbiology 13: 2717–2725.
Jung Y-T, Park S, Lee J-S, Yoon J-H. 2017. Roseobacter ponti sp. nov., isolated from seawater. International Journal of Systematic and Evolutionary Microbiology 67: 2189–2194.
Kirchman DL, Stegman MR, Nikrad MP, Cottrell MT. 2014. Abundance, size, and activity of aerobic anoxygenic phototrophic bacteria in coastal waters of the West Antarctic Peninsula. Aquatic Microbial Ecology 73: 41–49.
Kolber ZS, Van Dover CL, Niederman RA, Falkowski PG. 2000. Bacterial photosynthesis in surface waters of the open ocean. Nature 407: 177–179.
Kolber ZS, Plumley FG, Lang AS, Beatty JT, Blankenship RE, VanDover CL, Vetriani C, Koblizek M, Rathgeber C, Falkowski PG. 2001. Contribution of aerobic photoheterotrophic bacteria to the carbon cycle in the ocean. Science 292: 2492–2495.
Lami R, Cottrell MT, Ras J, Ulloa O, Obernosterer I, Claustre H, Kirchman DL, Lebaron P. 2007. High abundances of aerobic anoxygenic photosynthetic bacteria in the South Pacific Ocean. Applied and Environmental Microbiology 73: 4198–4205.
Muramatsu S, Kanamuro M, Sato-Takabe Y, Hirose S, Muramatsu Y, Takaichi S, Hanada S. 2020. Roseobacter cerasinus sp. nov., isolated from a fish farm. International Journal of Systematic and Evolutionary Microbiology: ijsem004360.
Okamura K, Takamiya K, Nishimura M. 1985. Photosynthetic electron transfer system is inoperative in anaerobic cells of Erythrobacter species strain OCh 114. Archives of Microbiology 142: 12–17.
Park S, Chen S, Yoon J-H. 2020. Erythrobacter insulae sp. nov., isolated from a tidal flat. International Journal of Systematic and Evolutionary Microbiology 70: 1470–1477.
Park S, Jung Y-T, Choi SJ, Yoon J-H. 2017. Erythrobacter aquimixticola sp. nov., isolated from the junction between the ocean and a freshwater spring. International Journal of Systematic and Evolutionary Microbiology 67: 2964–2969.
Parte AC. 2014. LPSN—list of prokaryotic names with standing in nomenclature. Nucleic Acids Research 42: D613–D616.
Parte AC. 2018. LPSN – List of Prokaryotic names with Standing in Nomenclature (bacterio.net), 20 years on. International Journal of Systematic and Evolutionary Microbiology 68: 1825–1829.
Rathgeber C, Beatty JT, Yurkov V. 2004. Aerobic phototrophic bacteria: New evidence for the diversity, ecological importance and applied potential of this previously overlooked group. Photosynthesis Research 81: 113–128.
Raven JA. 2009. Contributions of anoxygenic and oxygenic phototrophy and chemolithotrophy to carbon and oxygen fluxes in aquatic environments. Aquatic Microbial Ecology 56: 177–192.
Ritchie AE, Johnson ZI. 2012. Abundance and genetic diversity of aerobic anoxygenic phototrophic bacteria of coastal regions of the pacific ocean. Applied and Environmental Microbiology 78: 2858–2866.
Rocke E, Pachiadaki MG, Cobban A, Kujawinski EB, Edgcomb VP. 2015. Protist community grazing on prokaryotic prey in deep ocean water masses. PLoS ONE 10: 1–14.
Ruyitno. 1988. Perhitungan Secara Langsung Bakteri Laut Menggunakan Teknik Mikroskop Epifluoresens. XIII: 28–36.
Schwalbach MS, Fuhrman JA. 2005. Wide-ranging abundances of aerobic anoxygenic phototrophic bacteria in the world ocean revealed by epifluorescence microscopy and quantitative PCR. Limnology and Oceanography 50: 620–628.
Sherr BF, Sherr EB. 1984. Role of heterotrophic protozoa in carbon and energy flow in aquatic ecosystems, pp. 412–423 In Klugg MJ, Reddy CA [eds.], Current Perspectives in Microbial Ecology. ASM Press.
Shiba T, Shioi Y, Takamiya KI, Sutton DC, Wilkinson CR. 1991. Distribution and physiology of aerobic bacteria containing bacteriochlorophyll a on the east and west coasts of Australia. Applied and Environmental Microbiology 57: 295–300.
Shiba T, Simidu U. 1982. Bacterium Which Contains Bac teriochlorophyll a. International Journal of Systematic Bacteriology 32: 211–217.
Sieracki ME, Gilg IC, Thier EC, Poulton NJ, Goericke R. 2006. Distribution of planktonic aerobic anoxygenic photoheterotrophic bacteria in the northwest Atlantic. Limnology and Oceanography 51: 38–46.
Swingley WD, Sadekar S, Mastrian SD, Matthies HJ, Hao J, Ramos H, Acharya CR, Conrad AL, Taylor HL, Dejesa LC, Shah MK, O’Huallachain ME, Lince MT, Blankenship RE, Beatty JT, Touchman JW. 2007. The complete genome sequence of Roseobacter denitrificans reveals a mixotrophic rather than photosynthetic metabolism. Journal of Bacteriology 189: 683–690.
Thingstad TF, Zweifel UL, Rassoulzadegan F. 1998. P limitation of heterotrophic bacteria and phytoplankton in the northwest Mediterranean. Limnology and Oceanography 43: 88–94.
Tomaš AV, Šanti? D, Šoli? M, Ordulj M, Jozi? S, Šestanovi? S, Mati? F, Kušpili? G, Gladan ŽN. 2019. Dynamics of Aerobic Anoxygenic Phototrophs along the trophic gradient in the central Adriatic Sea. Deep-Sea Research Part II: Topical Studies in Oceanography 164: 112–121.
Unrein F, Gasol JM, Not F, Forn I, Massana R. 2014. Mixotrophic haptophytes are key bacterial grazers in oligotrophic coastal waters. ISME Journal 8: 164–176.
Wheeler PA, Kirchman DL. 1986. Utilization of inorganic and organic nitrogen by bacteria in marine systems. Limnology and Oceanography 31: 998–1009.
Xing T, Liu Y, Wang N, Xu B, Liu K, Shen L, Gu Z, Guo B, Zhou Y, Liu H. 2017. Erythrobacter arachoides sp. nov., isolated from ice core. International Journal of Systematic and Evolutionary Microbiology 67: 4235–4239.
Yurkov V V. 2006. Aerobic phototrophic proteobacteria, pp. 2006 In Dworkin M, Falkow S, Rosenberg E, Schleifer K, Stackebrandt E [eds.], The Prokaryotes. Springer.
Yurkov V V., Beatty JT. 1998. Aerobic Anoxygenic Phototrophic Bacteria. Microbiology and Molecular Biology Reviews 62: 695–724.
Zhuang L, Liu Y, Wang L, Wang W, Shao Z. 2015. Erythrobacter atlanticus sp. nov., a bacterium from ocean sediment able to degrade polycyclic aromatic hydrocarbons. International Journal of Systematic and Evolutionary Microbiology 65: 3714–3719.
Béjà O, Suzuki MT, Heidelberg JF, Nelson WC, Preston CM, Hamada T, Eisen JA, Fraser CM, DeLong EF. 2002. Unsuspected diversity among marine aerobic anoxygenic phototrophs. Nature 415: 630–633.
Biebl H, Allgaier M, Tindall BJ, Koblizek M, Lünsdorf H, Pukall R, Wagner-Döbler I. 2005. Dinoroseobacter shibae gen. nov., sp. nov., a new aerobic phototrophic bacterium isolated from dinoflagellates. International Journal of Systematic and Evolutionary Microbiology 55: 1089–1096.
Biebl H, Pukall R, Lünsdorf H, Schulz S, Allgaier M, Tindall BJ, Wagner-Döbler I. 2007. Description of Labrenzia alexandrii gen. nov., sp. nov., a novel alphaproteobacterium containing bacteriochlorophyll a, and a proposal reclassification of Stappia aggregata as Labrenzia aggregata comb. nov., and of Stappia alba as Labrenzia alba comb. nov. International Journal of Systematic and Evolutionary Microbiology 57: 1095–1107.
Boeuf D, Cottrell MT, Kirchman DL, Lebaron P, Jeanthon C. 2013. Summer community structure of aerobic anoxygenic phototrophic bacteria in the western Arctic Ocean. FEMS Microbiology Ecology 85: 417–432.
Brenner DJ, Krieg NR, Staley JT, Garrity M. 2005. 2 Bergey’s manual of systematic bacteriology. 2nd ed. Springer.
Calbet A, Landry MR, Nunnery S. 2001. Bacteria-flagellate interactions in the microbial food web of the oligotrophic subtropical North Pacific. Aquatic Microbial Ecology 23: 283–292.
Church MJ. 2008. Resource control of bacterial dynamics in the sea, pp. 353–354 In Kirchman DL [ed.], Microbial Ecology of the Oceans.
Church MJ, Hutchins DA, Ducklow HW. 2000. Limitation of bacterial growth by dissolved organic matter and iron in the Southern Ocean. Applied and Environmental Microbiology 66: 455–466.
Cottrell MT, Mannino A, Kirchman DL. 2006. Aerobic anoxygenic phototrophic bacteria in the mid-atlantic bight and the north pacific gyre. Applied and Environmental Microbiology 72: 557–564.
Csotonyi JT, Stackebrandt E, Swiderski J, Schumann P, Yurkov V. 2011. Chromocurvus halotolerans gen. nov., sp. nov., a gammaproteobacterial obligately aerobic anoxygenic phototroph, isolated from a Canadian hypersaline spring. Archives of Microbiology 193: 573–582.
Daley RJ, Hobbie JE. 1975. Direct counts of aquatic bacteria by a modified epifluorescence technique. Limnology and Oceanography 20: 875–882.
Du H, Jiao N, Hu Y, Zeng Y. 2006. Real-time PCR for quantification of aerobic anoxygenic phototrophic bacteria based on pufM gene in marine environment. Journal of Experimental Marine Biology and Ecology 329: 113–121.
Epstein SS, Shiaris MP. 1992. Size-selective grazing of coastal bacterioplankton by natural assemblages of pigmented flagellates, colorless flagellates, and ciliates. Microbial Ecology 23: 211–225.
Ferrera I, Gasol JM, Sebastián M, Hojerová E, Kobížek M. 2011. Comparison of growth rates of aerobic anoxygenic phototrophic bacteria and other bacterioplankton groups in coastal mediterranean waters. Applied and Environmental Microbiology 77: 7451–7458.
Ferrera I, Sánchez O, Kolá?ová E, Koblížek M, Gasol JM. 2017. Light enhances the growth rates of natural populations of aerobic anoxygenic phototrophic bacteria. ISME Journal 11: 2391–2393.
Garcia-Chaves MC, Cottrell MT, Kirchman DL, Derry AM, Bogard MJ, Del Giorgio PA. 2015. Major contribution of both zooplankton and protists to the top-down regulation of freshwater aerobic anoxygenic phototrophic bacteria. Aquatic Microbial Ecology 76: 71–83.
Gonzalez JM, Sherr EB, Sherr BF. 1990. Size-selective grazing on bacteria by natural assemblages of estuarine flagellates and ciliates. Applied and Environmental Microbiology 56: 583–589.
Hahn MW, Höfle MG. 2001. Grazing of protozoa and its effect on populations of aquatic bacteria. FEMS Microbiology Ecology 35: 113–121.
Hobbie JE, Daley RJ, Jasper S. 1977. Use of nuclepore filters for counting bacteria by fluorescence microscopy. Applied and Environmental Microbiology 33: 1225–1228.
Hojerová E, Mašín M, Brunet C, Ferrera I, Gasol JM, Koblížek M. 2011. Distribution and Growth of Aerobic Anoxygenic Phototrophs in the Mediterranean Sea. Environmental Microbiology 13: 2717–2725.
Jung Y-T, Park S, Lee J-S, Yoon J-H. 2017. Roseobacter ponti sp. nov., isolated from seawater. International Journal of Systematic and Evolutionary Microbiology 67: 2189–2194.
Kirchman DL, Stegman MR, Nikrad MP, Cottrell MT. 2014. Abundance, size, and activity of aerobic anoxygenic phototrophic bacteria in coastal waters of the West Antarctic Peninsula. Aquatic Microbial Ecology 73: 41–49.
Kolber ZS, Van Dover CL, Niederman RA, Falkowski PG. 2000. Bacterial photosynthesis in surface waters of the open ocean. Nature 407: 177–179.
Kolber ZS, Plumley FG, Lang AS, Beatty JT, Blankenship RE, VanDover CL, Vetriani C, Koblizek M, Rathgeber C, Falkowski PG. 2001. Contribution of aerobic photoheterotrophic bacteria to the carbon cycle in the ocean. Science 292: 2492–2495.
Lami R, Cottrell MT, Ras J, Ulloa O, Obernosterer I, Claustre H, Kirchman DL, Lebaron P. 2007. High abundances of aerobic anoxygenic photosynthetic bacteria in the South Pacific Ocean. Applied and Environmental Microbiology 73: 4198–4205.
Muramatsu S, Kanamuro M, Sato-Takabe Y, Hirose S, Muramatsu Y, Takaichi S, Hanada S. 2020. Roseobacter cerasinus sp. nov., isolated from a fish farm. International Journal of Systematic and Evolutionary Microbiology: ijsem004360.
Okamura K, Takamiya K, Nishimura M. 1985. Photosynthetic electron transfer system is inoperative in anaerobic cells of Erythrobacter species strain OCh 114. Archives of Microbiology 142: 12–17.
Park S, Chen S, Yoon J-H. 2020. Erythrobacter insulae sp. nov., isolated from a tidal flat. International Journal of Systematic and Evolutionary Microbiology 70: 1470–1477.
Park S, Jung Y-T, Choi SJ, Yoon J-H. 2017. Erythrobacter aquimixticola sp. nov., isolated from the junction between the ocean and a freshwater spring. International Journal of Systematic and Evolutionary Microbiology 67: 2964–2969.
Parte AC. 2014. LPSN—list of prokaryotic names with standing in nomenclature. Nucleic Acids Research 42: D613–D616.
Parte AC. 2018. LPSN – List of Prokaryotic names with Standing in Nomenclature (bacterio.net), 20 years on. International Journal of Systematic and Evolutionary Microbiology 68: 1825–1829.
Rathgeber C, Beatty JT, Yurkov V. 2004. Aerobic phototrophic bacteria: New evidence for the diversity, ecological importance and applied potential of this previously overlooked group. Photosynthesis Research 81: 113–128.
Raven JA. 2009. Contributions of anoxygenic and oxygenic phototrophy and chemolithotrophy to carbon and oxygen fluxes in aquatic environments. Aquatic Microbial Ecology 56: 177–192.
Ritchie AE, Johnson ZI. 2012. Abundance and genetic diversity of aerobic anoxygenic phototrophic bacteria of coastal regions of the pacific ocean. Applied and Environmental Microbiology 78: 2858–2866.
Rocke E, Pachiadaki MG, Cobban A, Kujawinski EB, Edgcomb VP. 2015. Protist community grazing on prokaryotic prey in deep ocean water masses. PLoS ONE 10: 1–14.
Ruyitno. 1988. Perhitungan Secara Langsung Bakteri Laut Menggunakan Teknik Mikroskop Epifluoresens. XIII: 28–36.
Schwalbach MS, Fuhrman JA. 2005. Wide-ranging abundances of aerobic anoxygenic phototrophic bacteria in the world ocean revealed by epifluorescence microscopy and quantitative PCR. Limnology and Oceanography 50: 620–628.
Sherr BF, Sherr EB. 1984. Role of heterotrophic protozoa in carbon and energy flow in aquatic ecosystems, pp. 412–423 In Klugg MJ, Reddy CA [eds.], Current Perspectives in Microbial Ecology. ASM Press.
Shiba T, Shioi Y, Takamiya KI, Sutton DC, Wilkinson CR. 1991. Distribution and physiology of aerobic bacteria containing bacteriochlorophyll a on the east and west coasts of Australia. Applied and Environmental Microbiology 57: 295–300.
Shiba T, Simidu U. 1982. Bacterium Which Contains Bac teriochlorophyll a. International Journal of Systematic Bacteriology 32: 211–217.
Sieracki ME, Gilg IC, Thier EC, Poulton NJ, Goericke R. 2006. Distribution of planktonic aerobic anoxygenic photoheterotrophic bacteria in the northwest Atlantic. Limnology and Oceanography 51: 38–46.
Swingley WD, Sadekar S, Mastrian SD, Matthies HJ, Hao J, Ramos H, Acharya CR, Conrad AL, Taylor HL, Dejesa LC, Shah MK, O’Huallachain ME, Lince MT, Blankenship RE, Beatty JT, Touchman JW. 2007. The complete genome sequence of Roseobacter denitrificans reveals a mixotrophic rather than photosynthetic metabolism. Journal of Bacteriology 189: 683–690.
Thingstad TF, Zweifel UL, Rassoulzadegan F. 1998. P limitation of heterotrophic bacteria and phytoplankton in the northwest Mediterranean. Limnology and Oceanography 43: 88–94.
Tomaš AV, Šanti? D, Šoli? M, Ordulj M, Jozi? S, Šestanovi? S, Mati? F, Kušpili? G, Gladan ŽN. 2019. Dynamics of Aerobic Anoxygenic Phototrophs along the trophic gradient in the central Adriatic Sea. Deep-Sea Research Part II: Topical Studies in Oceanography 164: 112–121.
Unrein F, Gasol JM, Not F, Forn I, Massana R. 2014. Mixotrophic haptophytes are key bacterial grazers in oligotrophic coastal waters. ISME Journal 8: 164–176.
Wheeler PA, Kirchman DL. 1986. Utilization of inorganic and organic nitrogen by bacteria in marine systems. Limnology and Oceanography 31: 998–1009.
Xing T, Liu Y, Wang N, Xu B, Liu K, Shen L, Gu Z, Guo B, Zhou Y, Liu H. 2017. Erythrobacter arachoides sp. nov., isolated from ice core. International Journal of Systematic and Evolutionary Microbiology 67: 4235–4239.
Yurkov V V. 2006. Aerobic phototrophic proteobacteria, pp. 2006 In Dworkin M, Falkow S, Rosenberg E, Schleifer K, Stackebrandt E [eds.], The Prokaryotes. Springer.
Yurkov V V., Beatty JT. 1998. Aerobic Anoxygenic Phototrophic Bacteria. Microbiology and Molecular Biology Reviews 62: 695–724.
Zhuang L, Liu Y, Wang L, Wang W, Shao Z. 2015. Erythrobacter atlanticus sp. nov., a bacterium from ocean sediment able to degrade polycyclic aromatic hydrocarbons. International Journal of Systematic and Evolutionary Microbiology 65: 3714–3719.
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