{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,8]],"date-time":"2026-11-08T11:28:48Z","timestamp":1794137328106,"version":"4.8.0"},"reference-count":60,"publisher":"National Academy of Sciences","issue":"14","license":[{"start":{"date-parts":[[2019,3,18]],"date-time":"2019-03-18T00:00:00Z","timestamp":1552867200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["www.pnas.org"],"crossmark-restriction":true},"short-container-title":["Proc. Natl. Acad. Sci. U.S.A."],"published-print":{"date-parts":[[2019,4,2]]},"abstract":"<jats:p>\n                    The exergonic reaction of FeS with H\n                    <jats:sub>2<\/jats:sub>\n                    S to form FeS\n                    <jats:sub>2<\/jats:sub>\n                    (pyrite) and H\n                    <jats:sub>2<\/jats:sub>\n                    was postulated to have operated as an early form of energy metabolism on primordial Earth. Since the Archean, sedimentary pyrite formation has played a major role in the global iron and sulfur cycles, with direct impact on the redox chemistry of the atmosphere. However, the mechanism of sedimentary pyrite formation is still being debated. We present microbial enrichment cultures which grew with FeS, H\n                    <jats:sub>2<\/jats:sub>\n                    S, and CO\n                    <jats:sub>2<\/jats:sub>\n                    as their sole substrates to produce FeS\n                    <jats:sub>2<\/jats:sub>\n                    and CH\n                    <jats:sub>4<\/jats:sub>\n                    . Cultures grew over periods of 3 to 8 mo to cell densities of up to 2 to 9 \u00d7 10\n                    <jats:sup>6<\/jats:sup>\n                    cells per mL\n                    <jats:sup>\u22121<\/jats:sup>\n                    . Transformation of FeS with H\n                    <jats:sub>2<\/jats:sub>\n                    S to FeS\n                    <jats:sub>2<\/jats:sub>\n                    was followed by\n                    <jats:sup>57<\/jats:sup>\n                    Fe M\u00f6ssbauer spectroscopy and showed a clear biological temperature profile with maximum activity at 28 \u00b0C and decreasing activities toward 4 \u00b0C and 60 \u00b0C. CH\n                    <jats:sub>4<\/jats:sub>\n                    was formed concomitantly with FeS\n                    <jats:sub>2<\/jats:sub>\n                    and exhibited the same temperature dependence. Addition of either penicillin or 2-bromoethanesulfonate inhibited both FeS\n                    <jats:sub>2<\/jats:sub>\n                    and CH\n                    <jats:sub>4<\/jats:sub>\n                    production, indicating a coupling of overall pyrite formation to methanogenesis. This hypothesis was supported by a 16S rRNA gene-based phylogenetic analysis, which identified at least one archaeal and five bacterial species. The archaeon was closely related to the hydrogenotrophic methanogen\n                    <jats:italic>Methanospirillum stamsii<\/jats:italic>\n                    , while the bacteria were most closely related to sulfate-reducing Deltaproteobacteria, as well as uncultured Firmicutes and Actinobacteria. Our results show that pyrite formation can be mediated at ambient temperature through a microbially catalyzed redox process, which may serve as a model for a postulated primordial iron\u2212sulfur world.\n                  <\/jats:p>","DOI":"10.1073\/pnas.1814412116","type":"journal-article","created":{"date-parts":[[2019,3,18]],"date-time":"2019-03-18T21:00:03Z","timestamp":1552942803000},"page":"6897-6902","update-policy":"https:\/\/doi.org\/10.1073\/pnas.cm10313","source":"Crossref","is-referenced-by-count":188,"title":["Pyrite formation from FeS and H\n                    <sub>2<\/sub>\n                    S is mediated through microbial redox activity"],"prefix":"10.1073","volume":"116","author":[{"given":"Joana","family":"Thiel","sequence":"first","affiliation":[{"name":"Department of Biology, University of Konstanz, 78464 Konstanz, Germany;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"James M.","family":"Byrne","sequence":"additional","affiliation":[{"name":"Geomicrobiology, Center for Applied Geoscience, Eberhardt Karls University of T\u00fcbingen, 72074 T\u00fcbingen, Germany;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Andreas","family":"Kappler","sequence":"additional","affiliation":[{"name":"Geomicrobiology, Center for Applied Geoscience, Eberhardt Karls University of T\u00fcbingen, 72074 T\u00fcbingen, Germany;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Bernhard","family":"Schink","sequence":"additional","affiliation":[{"name":"Department of Biology, University of Konstanz, 78464 Konstanz, Germany;"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6296-4145","authenticated-orcid":false,"given":"Michael","family":"Pester","sequence":"additional","affiliation":[{"name":"Department of Biology, University of Konstanz, 78464 Konstanz, Germany;"},{"name":"Department of Microorganisms, Leibniz Institute DSMZ\u2013German Culture Collection for Microorganisms and Cell Cultures, 38124 Braunschweig, Germany;"},{"name":"Institute for Microbiology, Technical University of Braunschweig, 38106 Braunschweig, Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"341","published-online":{"date-parts":[[2019,3,18]]},"reference":[{"key":"e_1_3_4_1_2","doi-asserted-by":"crossref","first-page":"514","DOI":"10.1021\/cr0503658","article-title":"Chemistry of iron sulfides","volume":"107","author":"Rickard D","year":"2007","unstructured":"D Rickard, 3rd GW Luther, Chemistry of iron sulfides. 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Geochim Cosmochim Acta 61, 135\u2013147 (1997).","journal-title":"Geochim Cosmochim Acta"},{"key":"e_1_3_4_4_2","doi-asserted-by":"crossref","first-page":"85","DOI":"10.1016\/S0012-821X(01)00352-1","article-title":"A novel iron sulphide mineral switch and its implications for Earth and planetary science","volume":"189","author":"Rickard D","year":"2001","unstructured":"D Rickard, IB Butler, A Oldroyd, A novel iron sulphide mineral switch and its implications for Earth and planetary science. Earth Planet Sci Lett 189, 85\u201391 (2001).","journal-title":"Earth Planet Sci Lett"},{"key":"e_1_3_4_5_2","doi-asserted-by":"crossref","first-page":"1","DOI":"10.2475\/ajs.268.1.1","article-title":"Sedimentary pyrite formation","volume":"268","author":"Berner RA","year":"1970","unstructured":"RA Berner, Sedimentary pyrite formation. Am J Sci 268, 1\u201323 (1970).","journal-title":"Am J Sci"},{"key":"e_1_3_4_6_2","doi-asserted-by":"crossref","first-page":"367","DOI":"10.1016\/j.gca.2017.10.006","article-title":"Sulfate-reducing bacteria influence the nucleation and growth of mackinawite and greigite","volume":"220","author":"Picard A","year":"2018","unstructured":"A Picard, A Gartman, DR Clarke, PR Girguis, Sulfate-reducing bacteria influence the nucleation and growth of mackinawite and greigite. Geochim Cosmochim Acta 220, 367\u2013384 (2018).","journal-title":"Geochim Cosmochim Acta"},{"key":"e_1_3_4_7_2","doi-asserted-by":"crossref","first-page":"4167","DOI":"10.1016\/S0016-7037(97)81466-4","article-title":"Pyrite formation by reactions of iron monosulfides with dissolved inorganic and organic sulfur species","volume":"60","author":"Wilkin RT","year":"1996","unstructured":"RT Wilkin, HL Barnes, Pyrite formation by reactions of iron monosulfides with dissolved inorganic and organic sulfur species. Geochim Cosmochim Acta 60, 4167\u20134179 (1996).","journal-title":"Geochim Cosmochim Acta"},{"key":"e_1_3_4_8_2","doi-asserted-by":"crossref","first-page":"1495","DOI":"10.1016\/0016-7037(91)90122-L","article-title":"Reactions forming pyrite and marcasite from solution. 1. Nucleation of FeS2 below 100\u00b0C","volume":"55","author":"Schoonen MAA","year":"1991","unstructured":"MAA Schoonen, HL Barnes, Reactions forming pyrite and marcasite from solution. 1. Nucleation of FeS2 below 100\u00b0C. Geochim Cosmochim Acta 55, 1495\u20131504 (1991).","journal-title":"Geochim Cosmochim Acta"},{"key":"e_1_3_4_9_2","first-page":"49","article-title":"The microbiological formation of iron sulphides","volume":"20","author":"Rickard D","year":"1969","unstructured":"D Rickard, The microbiological formation of iron sulphides. Stockholm Contrib Geol 20, 49\u201366 (1969).","journal-title":"Stockholm Contrib Geol"},{"key":"e_1_3_4_10_2","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1016\/S0009-2541(99)00198-9","article-title":"Reaction pathways in the Fe-S system below 100\u00b0C","volume":"167","author":"Benning LG","year":"2000","unstructured":"LG Benning, RT Wilkin, HL Barnes, Reaction pathways in the Fe-S system below 100\u00b0C. Chem Geol 167, 25\u201351 (2000).","journal-title":"Chem Geol"},{"key":"e_1_3_4_11_2","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1180\/minmag.1998.62A.1.80","article-title":"In situ determination of the stability of iron monosulphides and kinetics of pyrite formation","volume":"62","author":"Benning L","year":"1998","unstructured":"L Benning, H Barnes, In situ determination of the stability of iron monosulphides and kinetics of pyrite formation. Mineral Mag 62, 151\u2013152 (1998).","journal-title":"Mineral Mag"},{"key":"e_1_3_4_12_2","doi-asserted-by":"crossref","first-page":"605","DOI":"10.1016\/0016-7037(84)90089-9","article-title":"Sedimentary pyrite formation\u2014An update","volume":"48","author":"Berner RA","year":"1984","unstructured":"RA Berner, Sedimentary pyrite formation\u2014An update. Geochim Cosmochim Acta 48, 605\u2013615 (1984).","journal-title":"Geochim Cosmochim Acta"},{"key":"e_1_3_4_13_2","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1128\/AEM.64.1.119-125.1998","article-title":"Elemental sulfur and thiosulfate disproportionation by Desulfocapsa sulfoexigens sp. nov., a new anaerobic bacterium isolated from marine surface sediment","volume":"64","author":"Finster K","year":"1998","unstructured":"K Finster, W Liesack, B Thamdrup, Elemental sulfur and thiosulfate disproportionation by Desulfocapsa sulfoexigens sp. nov., a new anaerobic bacterium isolated from marine surface sediment. 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Limnol Oceanogr 43, 253\u2013264 (1998).","journal-title":"Limnol Oceanogr"},{"key":"e_1_3_4_15_2","doi-asserted-by":"crossref","first-page":"101","DOI":"10.1128\/aem.59.1.101-108.1993","article-title":"Bacterial disproportionation of elemental sulfur coupled to chemical reduction of iron or manganese","volume":"59","author":"Thamdrup B","year":"1993","unstructured":"B Thamdrup, K Finster, JW Hansen, F Bak, Bacterial disproportionation of elemental sulfur coupled to chemical reduction of iron or manganese. Appl Environ Microbiol 59, 101\u2013108 (1993).","journal-title":"Appl Environ Microbiol"},{"key":"e_1_3_4_16_2","doi-asserted-by":"crossref","first-page":"2019","DOI":"10.1016\/S0016-7037(99)00140-4","article-title":"Low temperature anaerobic bacterial diagenesis of ferrous monosulfide to pyrite","volume":"63","author":"Donald R","year":"1999","unstructured":"R Donald, G Southam, Low temperature anaerobic bacterial diagenesis of ferrous monosulfide to pyrite. Geochim Cosmochim Acta 63, 2019\u20132023 (1999).","journal-title":"Geochim Cosmochim Acta"},{"key":"e_1_3_4_17_2","doi-asserted-by":"crossref","first-page":"3581","DOI":"10.1016\/j.gca.2011.03.033","article-title":"A cryptic sulfur cycle driven by iron in the methane zone of marine sediment (Aarhus Bay, Denmark)","volume":"75","author":"Holmkvist L","year":"2011","unstructured":"L Holmkvist, TG Ferdelman, BB J\u00f8rgensen, A cryptic sulfur cycle driven by iron in the methane zone of marine sediment (Aarhus Bay, Denmark). Geochim Cosmochim Acta 75, 3581\u20133599 (2011).","journal-title":"Geochim Cosmochim Acta"},{"key":"e_1_3_4_18_2","doi-asserted-by":"crossref","first-page":"452","DOI":"10.1128\/mr.52.4.452-484.1988","article-title":"Before enzymes and templates: Theory of surface metabolism","volume":"52","author":"W\u00e4chtersh\u00e4user G","year":"1988","unstructured":"G W\u00e4chtersh\u00e4user, Before enzymes and templates: Theory of surface metabolism. Microbiol Rev 52, 452\u2013484 (1988).","journal-title":"Microbiol Rev"},{"key":"e_1_3_4_19_2","doi-asserted-by":"crossref","first-page":"584","DOI":"10.1002\/cbdv.200790052","article-title":"On the chemistry and evolution of the pioneer organism","volume":"4","author":"W\u00e4chtersh\u00e4user G","year":"2007","unstructured":"G W\u00e4chtersh\u00e4user, On the chemistry and evolution of the pioneer organism. Chem Biodivers 4, 584\u2013602 (2007).","journal-title":"Chem Biodivers"},{"key":"e_1_3_4_20_2","doi-asserted-by":"crossref","first-page":"59","DOI":"10.1098\/rstb.2002.1183","article-title":"On the origins of cells: A hypothesis for the evolutionary transitions from abiotic geochemistry to chemoautotrophic prokaryotes, and from prokaryotes to nucleated cells","volume":"358","author":"Martin W","year":"2003","unstructured":"W Martin, MJ Russell, On the origins of cells: A hypothesis for the evolutionary transitions from abiotic geochemistry to chemoautotrophic prokaryotes, and from prokaryotes to nucleated cells. Philos Trans R Soc Lond B Biol Sci 358, 59\u201383, and discussion (2003) 358:83\u201385 (2003).","journal-title":"Philos Trans R Soc Lond B Biol Sci"},{"key":"e_1_3_4_21_2","doi-asserted-by":"crossref","first-page":"334","DOI":"10.1016\/j.gca.2017.08.036","article-title":"Fe(III): S(-II) concentration ratio controls the pathway and the kinetics of pyrite formation during sulfidation of ferric hydroxides","volume":"217","author":"Wan M","year":"2017","unstructured":"M Wan, C Schroder, S Peiffer, Fe(III): S(-II) concentration ratio controls the pathway and the kinetics of pyrite formation during sulfidation of ferric hydroxides. Geochim Cosmochim Acta 217, 334\u2013348 (2017).","journal-title":"Geochim Cosmochim Acta"},{"key":"e_1_3_4_22_2","doi-asserted-by":"crossref","first-page":"319","DOI":"10.1007\/BF02396500","article-title":"M\u00f6ssbauer effect study of natural greigite","volume":"68","author":"Vandenberghe RE","year":"1992","unstructured":"RE Vandenberghe, E de Grave, PMA de Bakker, M Krs, JJ Hus, M\u00f6ssbauer effect study of natural greigite. Hyperfine Interact 68, 319\u2013322 (1992).","journal-title":"Hyperfine Interact"},{"key":"e_1_3_4_23_2","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1186\/1467-4866-8-1","article-title":"Greigite: A true intermediate on the polysulfide pathway to pyrite","volume":"8","author":"Hunger S","year":"2007","unstructured":"S Hunger, LG Benning, Greigite: A true intermediate on the polysulfide pathway to pyrite. Geochem Trans 8, 1 (2007).","journal-title":"Geochem Trans"},{"key":"e_1_3_4_24_2","doi-asserted-by":"crossref","first-page":"496","DOI":"10.1002\/jrs.2729","article-title":"The transformation of mackinawite into greigite studied by Raman spectroscopy","volume":"42","author":"Bourdoiseau JA","year":"2011","unstructured":"JA Bourdoiseau, M Jeannin, C Remazeilles, R Sabota, P Refait, The transformation of mackinawite into greigite studied by Raman spectroscopy. J Raman Spectrosc 42, 496\u2013504 (2011).","journal-title":"J Raman Spectrosc"},{"key":"e_1_3_4_25_2","doi-asserted-by":"crossref","first-page":"285","DOI":"10.1007\/978-1-4684-5409-3_8","article-title":"Use of specific inhibitors in biogeochemistry and microbial ecology","volume":"10","author":"Oremland RS","year":"1988","unstructured":"RS Oremland, DG Capone, Use of specific inhibitors in biogeochemistry and microbial ecology. Adv Microb Ecol 10, 285\u2013383 (1988).","journal-title":"Adv Microb Ecol"},{"key":"e_1_3_4_26_2","doi-asserted-by":"crossref","first-page":"309","DOI":"10.1007\/0-387-30742-7_11","volume-title":"The Prokaryotes","author":"Schink B","year":"2006","unstructured":"B Schink, AJM Stams, Syntrophism among prokaryotes. The Prokaryotes, eds M Dworkin, S Falkow, E Rosenberg, K-H Schleifer, E Stackebrandt (Springer, 3rd Ed, Singapore) Vol 2, 309\u2013335 (2006).","edition":"3"},{"key":"e_1_3_4_27_2","doi-asserted-by":"crossref","first-page":"755","DOI":"10.1128\/aem.48.4.755-757.1984","article-title":"Bacterial dry matter content and biomass estimations","volume":"48","author":"Bratbak G","year":"1984","unstructured":"G Bratbak, I Dundas, Bacterial dry matter content and biomass estimations. Appl Environ Microbiol 48, 755\u2013757 (1984).","journal-title":"Appl Environ Microbiol"},{"key":"e_1_3_4_28_2","first-page":"262","article-title":"Energetics of syntrophic cooperation in methanogenic degradation","volume":"61","author":"Schink B","year":"1997","unstructured":"B Schink, Energetics of syntrophic cooperation in methanogenic degradation. Microbiol Mol Biol Rev 61, 262\u2013280 (1997).","journal-title":"Microbiol Mol Biol Rev"},{"key":"e_1_3_4_29_2","first-page":"457","article-title":"The growth of micro-organisms in relation to their energy supply","volume":"23","author":"Bauchop T","year":"1960","unstructured":"T Bauchop, SR Elsden, The growth of micro-organisms in relation to their energy supply. J Gen Microbiol 23, 457\u2013469 (1960).","journal-title":"J Gen Microbiol"},{"key":"e_1_3_4_30_2","doi-asserted-by":"crossref","first-page":"300","DOI":"10.1007\/BF00521294","article-title":"Maintenance energy: A general model for energy-limited and energy-sufficient growth","volume":"133","author":"Pirt SJ","year":"1982","unstructured":"SJ Pirt, Maintenance energy: A general model for energy-limited and energy-sufficient growth. Arch Microbiol 133, 300\u2013302 (1982).","journal-title":"Arch Microbiol"},{"key":"e_1_3_4_31_2","doi-asserted-by":"crossref","first-page":"180","DOI":"10.1099\/ijs.0.056218-0","article-title":"Methanospirillum stamsii sp. nov., a psychrotolerant, hydrogenotrophic, methanogenic archaeon isolated from an anaerobic expanded granular sludge bed bioreactor operated at low temperature","volume":"64","author":"Parshina SN","year":"2014","unstructured":"SN Parshina, AV Ermakova, M Bomberg, EN Detkova, Methanospirillum stamsii sp. nov., a psychrotolerant, hydrogenotrophic, methanogenic archaeon isolated from an anaerobic expanded granular sludge bed bioreactor operated at low temperature. Int J Syst Evol Microbiol 64, 180\u2013186 (2014).","journal-title":"Int J Syst Evol Microbiol"},{"key":"e_1_3_4_32_2","first-page":"825","article-title":"A mesophilic, sulfate-reducing, rod-shaped, non-spore-forming bacterium","volume":"45","author":"Rozanova EP","year":"1976","unstructured":"EP Rozanova, TN Nazina, A mesophilic, sulfate-reducing, rod-shaped, non-spore-forming bacterium. Microbiologiya (USSR) 45, 825\u2013830 (1976).","journal-title":"Microbiologiya (USSR)"},{"key":"e_1_3_4_33_2","first-page":"634","article-title":"Isolation of a new genus of sulfate-reducing bacteria and description of a new species of this genus, Desulfomicrobium apsheronum gen. nov., sp. nov","volume":"57","author":"Rozanova EP","year":"1988","unstructured":"EP Rozanova, TN Nazina, AS Galushko, Isolation of a new genus of sulfate-reducing bacteria and description of a new species of this genus, Desulfomicrobium apsheronum gen. nov., sp. nov. Microbiologiya (USSR) 57, 634\u2013641 (1988).","journal-title":"Microbiologiya (USSR)"},{"key":"e_1_3_4_34_2","doi-asserted-by":"crossref","first-page":"184","DOI":"10.1007\/BF00415282","article-title":"Chemolithotrophic growth of Desulfovibrio sulfodismutans sp. nov. by disproportionation of inorganic sulfur-compounds","volume":"147","author":"Bak F","year":"1987","unstructured":"F Bak, N Pfennig, Chemolithotrophic growth of Desulfovibrio sulfodismutans sp. nov. by disproportionation of inorganic sulfur-compounds. Arch Microbiol 147, 184\u2013189 (1987).","journal-title":"Arch Microbiol"},{"key":"e_1_3_4_35_2","doi-asserted-by":"crossref","first-page":"545","DOI":"10.1099\/00207713-49-2-545","article-title":"Characterization of the anaerobic propionate-degrading syntrophs Smithella propionica gen. nov., sp. nov. and Syntrophobacter wolinii","volume":"49","author":"Liu Y","year":"1999","unstructured":"Y Liu, DL Balkwill, HC Aldrich, GR Drake, DR Boone, Characterization of the anaerobic propionate-degrading syntrophs Smithella propionica gen. nov., sp. nov. and Syntrophobacter wolinii. Int J Syst Bacteriol 49, 545\u2013556 (1999).","journal-title":"Int J Syst Bacteriol"},{"key":"e_1_3_4_36_2","doi-asserted-by":"crossref","first-page":"353","DOI":"10.1111\/j.1574-6976.1998.tb00376.x","article-title":"Anaerobic respiration with elemental sulfur and with disulfides","volume":"22","author":"Hedderich R","year":"1998","unstructured":"R Hedderich, , Anaerobic respiration with elemental sulfur and with disulfides. FEMS Microbiol Rev 22, 353\u2013381 (1998).","journal-title":"FEMS Microbiol Rev"},{"key":"e_1_3_4_37_2","doi-asserted-by":"crossref","first-page":"292","DOI":"10.1016\/j.mib.2011.03.003","article-title":"Anaerobic oxidation of methane with sulfate: On the reversibility of the reactions that are catalyzed by enzymes also involved in methanogenesis from CO2","volume":"14","author":"Thauer RK","year":"2011","unstructured":"RK Thauer, Anaerobic oxidation of methane with sulfate: On the reversibility of the reactions that are catalyzed by enzymes also involved in methanogenesis from CO2. Curr Opin Microbiol 14, 292\u2013299 (2011).","journal-title":"Curr Opin Microbiol"},{"key":"e_1_3_4_38_2","doi-asserted-by":"crossref","first-page":"688","DOI":"10.1093\/femsre\/fuv020","article-title":"Life under extreme energy limitation: A synthesis of laboratory- and field-based investigations","volume":"39","author":"Lever MA","year":"2015","unstructured":"MA Lever, , Life under extreme energy limitation: A synthesis of laboratory- and field-based investigations. FEMS Microbiol Rev 39, 688\u2013728 (2015).","journal-title":"FEMS Microbiol Rev"},{"key":"e_1_3_4_39_2","doi-asserted-by":"crossref","first-page":"629","DOI":"10.1139\/cjm-2017-0545","article-title":"The effect of gram-positive (Desulfosporosinus orientis) and gram-negative (Desulfovibrio desulfuricans) sulfate-reducing bacteria on iron sulfide mineral precipitation","volume":"64","author":"Stanley W","year":"2018","unstructured":"W Stanley, G Southam, The effect of gram-positive (Desulfosporosinus orientis) and gram-negative (Desulfovibrio desulfuricans) sulfate-reducing bacteria on iron sulfide mineral precipitation. Can J Microbiol 64, 629\u2013637 (2018).","journal-title":"Can J Microbiol"},{"key":"e_1_3_4_40_2","doi-asserted-by":"crossref","first-page":"311","DOI":"10.1146\/annurev-marine-010814-015535","article-title":"Slow microbial life in the seabed","volume":"8","author":"J\u00f8rgensen BB","year":"2016","unstructured":"BB J\u00f8rgensen, IPG Marshall, Slow microbial life in the seabed. Annu Rev Mar Sci 8, 311\u2013332 (2016).","journal-title":"Annu Rev Mar Sci"},{"key":"e_1_3_4_41_2","doi-asserted-by":"crossref","first-page":"932","DOI":"10.1126\/science.1133796","article-title":"Ecology. A starving majority deep beneath the seafloor","volume":"314","author":"J\u00f8rgensen BB","year":"2006","unstructured":"BB J\u00f8rgensen, S D\u2019Hondt, Ecology. A starving majority deep beneath the seafloor. Science 314, 932\u2013934 (2006).","journal-title":"Science"},{"key":"e_1_3_4_42_2","doi-asserted-by":"crossref","first-page":"893","DOI":"10.1016\/j.gca.2004.07.032","article-title":"The yield and isotopic composition of radiolytic H2, a potential energy source for the deep subsurface biosphere","volume":"69","author":"Lin L-H","year":"2005","unstructured":"L-H Lin, G Slater, B Lollar, G Lacrampe-Couloume, TC Onstott, The yield and isotopic composition of radiolytic H2, a potential energy source for the deep subsurface biosphere. 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Characterization of the filamentous gliding Desulfonema limicola gen. nov sp. nov, and Desulfonema magnum sp. nov. Arch Microbiol 134, 286\u2013294 (1983).","journal-title":"Arch Microbiol"},{"key":"e_1_3_4_48_2","doi-asserted-by":"crossref","first-page":"100","DOI":"10.1007\/BF00692721","article-title":"Convenient method to prepare neutral sulfide solution for cultivation of phototrophic sulfur bacteria","volume":"139","author":"Siefert E","year":"1984","unstructured":"E Siefert, N Pfennig, Convenient method to prepare neutral sulfide solution for cultivation of phototrophic sulfur bacteria. 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Nucleic Acids Res 41, D590\u2013D596 (2013).","journal-title":"Nucleic Acids Res"},{"key":"e_1_3_4_56_2","doi-asserted-by":"crossref","first-page":"1363","DOI":"10.1093\/nar\/gkh293","article-title":"ARB: A software environment for sequence data","volume":"32","author":"Ludwig W","year":"2004","unstructured":"W Ludwig, , ARB: A software environment for sequence data. Nucleic Acids Res 32, 1363\u20131371 (2004).","journal-title":"Nucleic Acids Res"},{"key":"e_1_3_4_57_2","doi-asserted-by":"crossref","first-page":"7537","DOI":"10.1128\/AEM.01541-09","article-title":"Introducing mothur: Open-source, platform-independent, community-supported software for describing and comparing microbial communities","volume":"75","author":"Schloss PD","year":"2009","unstructured":"PD Schloss, , Introducing mothur: Open-source, platform-independent, community-supported software for describing and comparing microbial communities. Appl Environ Microbiol 75, 7537\u20137541 (2009).","journal-title":"Appl Environ Microbiol"},{"key":"e_1_3_4_58_2","first-page":"152","article-title":"Taxonomic parameters revisited: Tarnished gold standards","volume":"33","author":"Stackebrandt E","year":"2006","unstructured":"E Stackebrandt, J Ebers, Taxonomic parameters revisited: Tarnished gold standards. Microbiol Today 33, 152\u2013154 (2006).","journal-title":"Microbiol Today"},{"key":"e_1_3_4_59_2","doi-asserted-by":"crossref","first-page":"1312","DOI":"10.1093\/bioinformatics\/btu033","article-title":"RAxML version 8: A tool for phylogenetic analysis and post-analysis of large phylogenies","volume":"30","author":"Stamatakis A","year":"2014","unstructured":"A Stamatakis, RAxML version 8: A tool for phylogenetic analysis and post-analysis of large phylogenies. 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