{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,7]],"date-time":"2026-10-07T23:49:36Z","timestamp":1791416976582,"version":"4.3.3"},"reference-count":97,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,10,8]],"date-time":"2026-10-08T00:00:00Z","timestamp":1791417600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2026,10,8]],"date-time":"2026-10-08T00:00:00Z","timestamp":1791417600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Microbiome"],"DOI":"10.1186\/s40168-026-02536-0","type":"journal-article","created":{"date-parts":[[2026,10,7]],"date-time":"2026-10-07T23:02:34Z","timestamp":1791414154000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Novel Antarctic chemolithotroph drives iron biomineralization"],"prefix":"10.1186","volume":"14","author":[{"given":"Jaekyung","family":"Yoon","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Boyoung","family":"Lee","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kyu-Cheul","family":"Yoo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Min-Jung","family":"Kwak","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hae Jung","family":"Song","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chung Yeon","family":"Hwang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yusook","family":"Chung","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kitae","family":"Kim","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Soon-Kyeong","family":"Kwon","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ju Yeon","family":"Song","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hwan Su","family":"Yoon","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7715-6992","authenticated-orcid":false,"given":"Jihyun F.","family":"Kim","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,10,8]]},"reference":[{"key":"2536_CR1","unstructured":"Madigan MT, Aiyer J, Buckley DH, Sattley WM, Stahl DA. Brock biology of microorganisms. eBook. 16th ed. Hoboken, NJ: Pearson Higher Ed; 2021."},{"key":"2536_CR2","doi-asserted-by":"publisher","first-page":"123","DOI":"10.1038\/s43017-020-00116-w","volume":"2","author":"NJ Planavsky","year":"2021","unstructured":"Planavsky NJ, Crowe SA, Fakhraee M, Beaty B, Reinhard CT, Mills BJW, et al. Evolution of the structure and impact of Earth\u2019s biosphere. Nat Rev Earth Environ. 2021;2:123\u201339. https:\/\/doi.org\/10.1038\/s43017-020-00116-w.","journal-title":"Nat Rev Earth Environ"},{"key":"2536_CR3","doi-asserted-by":"publisher","first-page":"2857","DOI":"10.1098\/rstb.2011.0129","volume":"366","author":"NH Sleep","year":"2011","unstructured":"Sleep NH, Bird DK, Pope EC. Serpentinite and the dawn of life. Philos Trans R Soc Lond B Biol Sci. 2011;366:2857\u201369.","journal-title":"Philos Trans R Soc Lond B Biol Sci"},{"key":"2536_CR4","doi-asserted-by":"publisher","first-page":"572","DOI":"10.1038\/s41579-024-01044-y","volume":"22","author":"TW Lyons","year":"2024","unstructured":"Lyons TW, Tino CJ, Fournier GP, Anderson RE, Leavitt WD, Konhauser KO, et al. Co-evolution of early Earth environments and microbial life. Nat Rev Microbiol. 2024;22:572\u201386.","journal-title":"Nat Rev Microbiol"},{"key":"2536_CR5","doi-asserted-by":"publisher","first-page":"307","DOI":"10.1038\/nature13068","volume":"506","author":"TW Lyons","year":"2014","unstructured":"Lyons TW, Reinhard CT, Planavsky NJ. The rise of oxygen in Earth\u2019s early ocean and atmosphere. Nature. 2014;506:307\u201315.","journal-title":"Nature"},{"key":"2536_CR6","doi-asserted-by":"publisher","first-page":"140","DOI":"10.1016\/j.earscirev.2017.06.012","volume":"172","author":"KO Konhauser","year":"2017","unstructured":"Konhauser KO, Planavsky NJ, Hardisty DS, Robbins LJ, Warchola TJ, Haugaard R, et al. Iron formations: a global record of Neoarchaean to Palaeoproterozoic environmental history. Earth Sci Rev. 2017;172:140\u201377.","journal-title":"Earth Sci Rev"},{"key":"2536_CR7","doi-asserted-by":"publisher","first-page":"138","DOI":"10.1016\/j.freeradbiomed.2019.05.005","volume":"140","author":"NJ Tosca","year":"2019","unstructured":"Tosca NJ, Jiang CZ, Rasmussen B, Muhling J. Products of the iron cycle on the early Earth. Free Radic Biol Med. 2019;140:138\u201353.","journal-title":"Free Radic Biol Med"},{"key":"2536_CR8","doi-asserted-by":"publisher","first-page":"467","DOI":"10.2113\/gsecongeo.105.3.467","volume":"105","author":"A Bekker","year":"2010","unstructured":"Bekker A, Slack JF, Planavsky N, Krapez B, Hofmann A, Konhauser KO, et al. Iron formation: the sedimentary product of a complex interplay among mantle, tectonic, oceanic, and biospheric processes. Econ Geol. 2010;105:467\u2013508.","journal-title":"Econ Geol"},{"key":"2536_CR9","doi-asserted-by":"publisher","first-page":"1135","DOI":"10.2113\/gsecongeo.68.7.1135","volume":"68","author":"P Cloud","year":"1973","unstructured":"Cloud P. Paleoecological significance of the banded iron-formation. Econ Geol. 1973;68:1135\u201343.","journal-title":"Econ Geol"},{"key":"2536_CR10","doi-asserted-by":"publisher","first-page":"509","DOI":"10.1111\/gbi.12192","volume":"14","author":"CS Chan","year":"2016","unstructured":"Chan CS, Emerson D, Luther GW III. The role of microaerophilic Fe-oxidizing micro-organisms in producing banded iron formations. Geobiology. 2016;14:509\u201328.","journal-title":"Geobiology"},{"key":"2536_CR11","doi-asserted-by":"publisher","first-page":"558","DOI":"10.1038\/s41561-019-0372-0","volume":"12","author":"LJ Robbins","year":"2019","unstructured":"Robbins LJ, Funk SP, Flynn SL, Warchola TJ, Li ZQ, Lalonde SV, et al. Hydrogeological constraints on the formation of Palaeoproterozoic banded iron formations. Nat Geosci. 2019;12:558\u201363.","journal-title":"Nat Geosci"},{"key":"2536_CR12","doi-asserted-by":"publisher","first-page":"593","DOI":"10.1007\/s12542-021-00598-z","volume":"95","author":"CL Dreher","year":"2021","unstructured":"Dreher CL, Schad M, Robbins LJ, Konhauser KO, Kappler A, Joshi P. Microbial processes during deposition and diagenesis of Banded Iron Formations. Palaontol Z. 2021;95:593\u2013610.","journal-title":"Palaontol Z"},{"key":"2536_CR13","doi-asserted-by":"publisher","first-page":"1198","DOI":"10.1126\/science.1228026","volume":"339","author":"SA Marcott","year":"2013","unstructured":"Marcott SA, Shakun JD, Clark PU, Mix AC. A reconstruction of regional and global temperature for the past 11,300 years. Science. 2013;339:1198\u2013201.","journal-title":"Science"},{"key":"2536_CR14","doi-asserted-by":"publisher","first-page":"39","DOI":"10.1038\/nature25152","volume":"553","author":"B Bereiter","year":"2018","unstructured":"Bereiter B, Shackleton S, Baggenstos D, Kawamura K, Severinghaus J. Mean global ocean temperatures during the last glacial transition. Nature. 2018;553:39\u201344.","journal-title":"Nature"},{"key":"2536_CR15","doi-asserted-by":"publisher","first-page":"548","DOI":"10.1038\/s41586-020-03155-x","volume":"589","author":"S Bova","year":"2021","unstructured":"Bova S, Rosenthal Y, Liu ZY, Godad SP, Yan M. Seasonal origin of the thermal maxima at the Holocene and the last interglacial. Nature. 2021;589:548\u201353.","journal-title":"Nature"},{"key":"2536_CR16","doi-asserted-by":"publisher","first-page":"356","DOI":"10.1038\/nature25454","volume":"554","author":"K Rehfeld","year":"2018","unstructured":"Rehfeld K, M\u00fcnch T, Ho SL, Laepple T. Global patterns of declining temperature variability from the Last Glacial Maximum to the Holocene. Nature. 2018;554:356\u20139.","journal-title":"Nature"},{"key":"2536_CR17","doi-asserted-by":"publisher","first-page":"171","DOI":"10.1038\/nature14258","volume":"519","author":"SL Lewis","year":"2015","unstructured":"Lewis SL, Maslin MA. Defining the Anthropocene. Nature. 2015;519:171\u201380.","journal-title":"Nature"},{"key":"2536_CR18","doi-asserted-by":"publisher","DOI":"10.1126\/science.aad2622","volume":"351","author":"CN Waters","year":"2016","unstructured":"Waters CN, Zalasiewicz J, Summerhayes C, Barnosky AD, Poirier C, Ga\u0142uszka A, et al. The Anthropocene is functionally and stratigraphically distinct from the Holocene. Science. 2016;351:aad2622.","journal-title":"Science"},{"key":"2536_CR19","doi-asserted-by":"publisher","first-page":"1321","DOI":"10.1126\/science.aaz5489","volume":"367","author":"RE Bell","year":"2020","unstructured":"Bell RE, Seroussi H. History, mass loss, structure, and dynamic behavior of the Antarctic Ice Sheet. Science. 2020;367:1321\u20135.","journal-title":"Science"},{"key":"2536_CR20","doi-asserted-by":"publisher","first-page":"5635","DOI":"10.1038\/s41467-019-13496-5","volume":"10","author":"JA Smith","year":"2019","unstructured":"Smith JA, Graham AGC, Post AL, Hillenbrand CD, Bart PJ, Powell RD. The marine geological imprint of Antarctic ice shelves. Nat Commun. 2019;10:5635.","journal-title":"Nat Commun"},{"key":"2536_CR21","doi-asserted-by":"publisher","first-page":"978","DOI":"10.1130\/G48503.1","volume":"49","author":"JA Smith","year":"2021","unstructured":"Smith JA, Hillenbrand C-D, Subt C, Rosenheim BE, Frederichs T, Ehrmann W, et al. History of the Larsen C Ice Shelf reconstructed from sub\u2013ice shelf and offshore sediments. Geology. 2021;49:978\u201382.","journal-title":"Geology"},{"key":"2536_CR22","doi-asserted-by":"publisher","first-page":"223","DOI":"10.1038\/s41586-018-0171-6","volume":"558","author":"A Shepherd","year":"2018","unstructured":"Shepherd A, Fricker HA, Farrell SL. Trends and connections across the Antarctic cryosphere. Nature. 2018;558:223\u201332.","journal-title":"Nature"},{"key":"2536_CR23","doi-asserted-by":"publisher","first-page":"5786","DOI":"10.1038\/s41467-019-13741-x","volume":"10","author":"J Jung","year":"2019","unstructured":"Jung J, Yoo KC, Rosenheim BE, Conway TM, Lee JI, Yoon HI, et al. Microbial Fe(III) reduction as a potential iron source from Holocene sediments beneath Larsen Ice Shelf. Nat Commun. 2019;10:5786.","journal-title":"Nat Commun"},{"key":"2536_CR24","unstructured":"Yoo KC, Lee JI, Lee MK, Kim S, Subt C, Rosenheim B, et al. Snowball Earth (the Cryogenian) reincarnated in the Holocene sediments beneath the Larsen C Ice Shelf (Antarctica); Submitted."},{"key":"2536_CR25","doi-asserted-by":"publisher","first-page":"1404","DOI":"10.1002\/2016GC006578","volume":"18","author":"C Subt","year":"2017","unstructured":"Subt C, Yoon HI, Yoo KC, Lee JI, Leventer A, Domack EW, et al. Sub-ice shelf sediment geochronology utilizing novel radiocarbon methodology for highly detrital sediments. Geochem Geophys Geosyst. 2017;18:1404\u201318.","journal-title":"Geochem Geophys Geosyst"},{"key":"2536_CR26","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0105585","volume":"9","author":"CFA de Bourcy","year":"2014","unstructured":"de Bourcy CFA, De Vlaminck I, Kanbar JN, Wang J, Gawad C, Quake SR. A quantitative comparison of single-cell whole genome amplification methods. PLoS ONE. 2014;9:e105585.","journal-title":"PLoS ONE"},{"key":"2536_CR27","doi-asserted-by":"publisher","first-page":"852","DOI":"10.1038\/s41587-019-0209-9","volume":"37","author":"E Bolyen","year":"2019","unstructured":"Bolyen E, Rideout JR, Dillon MR, Bokulich NA, Abnet CC, Al-Ghalith GA, et al. Reproducible, interactive, scalable and extensible microbiome data science using QIIME 2. Nat Biotechnol. 2019;37:852\u20137.","journal-title":"Nat Biotechnol"},{"key":"2536_CR28","doi-asserted-by":"publisher","DOI":"10.7717\/peerj.2584","volume":"4","author":"T Rognes","year":"2016","unstructured":"Rognes T, Flouri T, Nichols B, Quince C, Mah\u00e9 F. VSEARCH: a versatile open source tool for metagenomics. PeerJ. 2016;4:e2584.","journal-title":"PeerJ"},{"key":"2536_CR29","doi-asserted-by":"publisher","first-page":"90","DOI":"10.1186\/s40168-018-0470-z","volume":"6","author":"NA Bokulich","year":"2018","unstructured":"Bokulich NA, Kaehler BD, Rideout JR, Dillon M, Bolyen E, Knight R, et al. Optimizing taxonomic classification of marker-gene amplicon sequences with QIIME 2\u2019s q2-feature-classifier plugin. Microbiome. 2018;6:90.","journal-title":"Microbiome"},{"key":"2536_CR30","doi-asserted-by":"publisher","first-page":"D643","DOI":"10.1093\/nar\/gkt1209","volume":"42","author":"P Yilmaz","year":"2014","unstructured":"Yilmaz P, Parfrey LW, Yarza P, Gerken J, Pruesse E, Quast C, et al. The SILVA and \u201cAll-species Living Tree Project (LTP)\u201d taxonomic frameworks. Nucleic Acids Res. 2014;42:D643\u20138.","journal-title":"Nucleic Acids Res"},{"key":"2536_CR31","doi-asserted-by":"publisher","first-page":"1823","DOI":"10.1093\/bioinformatics\/bts252","volume":"28","author":"E Pruesse","year":"2012","unstructured":"Pruesse E, Peplies J, Gl\u00f6ckner FO. SINA: accurate high-throughput multiple sequence alignment of ribosomal RNA genes. Bioinformatics. 2012;28:1823\u20139.","journal-title":"Bioinformatics"},{"key":"2536_CR32","doi-asserted-by":"publisher","first-page":"268","DOI":"10.1093\/molbev\/msu300","volume":"32","author":"LT Nguyen","year":"2015","unstructured":"Nguyen LT, Schmidt HA, von Haeseler A, Minh BQ. IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Mol Biol Evol. 2015;32:268\u201374.","journal-title":"Mol Biol Evol"},{"key":"2536_CR33","doi-asserted-by":"publisher","first-page":"W293","DOI":"10.1093\/nar\/gkab301","volume":"49","author":"I Letunic","year":"2021","unstructured":"Letunic I, Bork P. Interactive Tree Of Life (iTOL) v5: an online tool for phylogenetic tree display and annotation. Nucleic Acids Res. 2021;49:W293\u20136.","journal-title":"Nucleic Acids Res"},{"key":"2536_CR34","doi-asserted-by":"publisher","first-page":"R60","DOI":"10.1186\/gb-2011-12-6-r60","volume":"12","author":"N Segata","year":"2011","unstructured":"Segata N, Izard J, Waldron L, Gevers D, Miropolsky L, Garrett WS, et al. Metagenomic biomarker discovery and explanation. Genome Biol. 2011;12:R60.","journal-title":"Genome Biol"},{"key":"2536_CR35","doi-asserted-by":"publisher","first-page":"312","DOI":"10.1111\/1755-0998.13704","volume":"23","author":"M Shaffer","year":"2023","unstructured":"Shaffer M, Thurimella K, Sterrett JD, Lozupone CA. SCNIC: sparse correlation network investigation for compositional data. Mol Ecol Resour. 2023;23:312\u201325.","journal-title":"Mol Ecol Resour"},{"key":"2536_CR36","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pcbi.1002687","volume":"8","author":"J Friedman","year":"2012","unstructured":"Friedman J, Alm EJ. Inferring correlation networks from genomic survey data. PLOS Comput Biol. 2012;8:e1002687.","journal-title":"PLOS Comput Biol"},{"key":"2536_CR37","doi-asserted-by":"publisher","first-page":"2498","DOI":"10.1101\/gr.1239303","volume":"13","author":"P Shannon","year":"2003","unstructured":"Shannon P, Markiel A, Ozier O, Baliga NS, Wang JT, Ramage D, et al. Cytoscape: a software environment for integrated models of biomolecular interaction networks. Genome Res. 2003;13:2498\u2013504.","journal-title":"Genome Res"},{"key":"2536_CR38","doi-asserted-by":"publisher","first-page":"158","DOI":"10.1186\/s40168-018-0541-1","volume":"6","author":"GV Uritskiy","year":"2018","unstructured":"Uritskiy GV, DiRuggiero J, Taylor J. MetaWRAP-a flexible pipeline for genome-resolved metagenomic data analysis. Microbiome. 2018;6:158.","journal-title":"Microbiome"},{"key":"2536_CR39","doi-asserted-by":"publisher","first-page":"1674","DOI":"10.1093\/bioinformatics\/btv033","volume":"31","author":"D Li","year":"2015","unstructured":"Li D, Liu CM, Luo R, Sadakane K, Lam TW. MEGAHIT: an ultra-fast single-node solution for large and complex metagenomics assembly via succinct de Bruijn graph. Bioinformatics. 2015;31:1674\u20136.","journal-title":"Bioinformatics"},{"key":"2536_CR40","doi-asserted-by":"publisher","first-page":"119","DOI":"10.1186\/1471-2105-11-119","volume":"11","author":"D Hyatt","year":"2010","unstructured":"Hyatt D, Chen GL, Locascio PF, Land ML, Larimer FW, Hauser LJ. Prodigal: prokaryotic gene recognition and translation initiation site identification. BMC Bioinformatics. 2010;11:119.","journal-title":"BMC Bioinformatics"},{"key":"2536_CR41","doi-asserted-by":"publisher","first-page":"2251","DOI":"10.1093\/bioinformatics\/btz859","volume":"36","author":"T Aramaki","year":"2020","unstructured":"Aramaki T, Blanc-Mathieu R, Endo H, Ohkubo K, Kanehisa M, Goto S, et al. KofamKOALA: KEGG Ortholog assignment based on profile HMM and adaptive score threshold. Bioinformatics. 2020;36:2251\u20132.","journal-title":"Bioinformatics"},{"key":"2536_CR42","doi-asserted-by":"publisher","first-page":"28","DOI":"10.1002\/pro.3711","volume":"29","author":"M Kanehisa","year":"2020","unstructured":"Kanehisa M, Sato Y. KEGG Mapper for inferring cellular functions from protein sequences. Protein Sci. 2020;29:28\u201335.","journal-title":"Protein Sci"},{"key":"2536_CR43","doi-asserted-by":"publisher","first-page":"37","DOI":"10.3389\/fmicb.2020.00037","volume":"11","author":"AI Garber","year":"2020","unstructured":"Garber AI, Nealson KH, Okamoto A, McAllister SM, Chan CS, Barco RA, et al. FeGenie: a comprehensive tool for the identification of iron genes and iron gene neighborhoods in genome and metagenome assemblies. Front Microbiol. 2020;11:37.","journal-title":"Front Microbiol"},{"key":"2536_CR44","volume":"7","author":"S Neukirchen","year":"2021","unstructured":"Neukirchen S, Sousa FL. DiSCo: a sequence-based type-specific predictor of Dsr-dependent dissimilatory sulphur metabolism in microbial data. Microb Genom. 2021;7:000603.","journal-title":"Microb Genom"},{"key":"2536_CR45","doi-asserted-by":"publisher","first-page":"1040","DOI":"10.1093\/bioinformatics\/bty741","volume":"35","author":"Q Tu","year":"2019","unstructured":"Tu Q, Lin L, Cheng L, Deng Y, He Z. NCycDB: a curated integrative database for fast and accurate metagenomic profiling of nitrogen cycling genes. Bioinformatics. 2019;35:1040\u20138.","journal-title":"Bioinformatics"},{"key":"2536_CR46","doi-asserted-by":"publisher","first-page":"1144","DOI":"10.1038\/nmeth.3103","volume":"11","author":"J Alneberg","year":"2014","unstructured":"Alneberg J, Bjarnason BS, de Bruijn I, Schirmer M, Quick J, Ijaz UZ, et al. Binning metagenomic contigs by coverage and composition. Nat Methods. 2014;11:1144\u20136.","journal-title":"Nat Methods"},{"key":"2536_CR47","doi-asserted-by":"publisher","first-page":"605","DOI":"10.1093\/bioinformatics\/btv638","volume":"32","author":"YW Wu","year":"2016","unstructured":"Wu YW, Simmons BA, Singer SWMB. MaxBin 2.0: an automated binning algorithm to recover genomes from multiple metagenomic datasets. Bioinformatics. 2016;32:605\u20137.","journal-title":"Bioinformatics"},{"key":"2536_CR48","doi-asserted-by":"publisher","DOI":"10.7717\/peerj.7359","volume":"7","author":"DD Kang","year":"2019","unstructured":"Kang DD, Li F, Kirton E, Thomas A, Egan R, An H, et al. MetaBAT 2: an adaptive binning algorithm for robust and efficient genome reconstruction from metagenome assemblies. PeerJ. 2019;7:e7359.","journal-title":"PeerJ"},{"key":"2536_CR49","doi-asserted-by":"publisher","first-page":"1043","DOI":"10.1101\/gr.186072.114","volume":"25","author":"DH Parks","year":"2015","unstructured":"Parks DH, Imelfort M, Skennerton CT, Hugenholtz P, Tyson GW. CheckM: assessing the quality of microbial genomes recovered from isolates, single cells, and metagenomes. Genome Res. 2015;25:1043\u201355.","journal-title":"Genome Res"},{"key":"2536_CR50","doi-asserted-by":"publisher","first-page":"357","DOI":"10.1038\/nmeth.1923","volume":"9","author":"B Langmead","year":"2012","unstructured":"Langmead B, Salzberg SL. Fast gapped-read alignment with Bowtie 2. Nat Methods. 2012;9:357\u20139.","journal-title":"Nat Methods"},{"key":"2536_CR51","doi-asserted-by":"publisher","first-page":"455","DOI":"10.1089\/cmb.2012.0021","volume":"19","author":"A Bankevich","year":"2012","unstructured":"Bankevich A, Nurk S, Antipov D, Gurevich AA, Dvorkin M, Kulikov AS, et al. SPAdes: a new genome assembly algorithm and its applications to single-cell sequencing. J Comput Biol. 2012;19:455\u201377.","journal-title":"J Comput Biol"},{"key":"2536_CR52","doi-asserted-by":"publisher","first-page":"1925","DOI":"10.1093\/bioinformatics\/btz848","volume":"36","author":"PA Chaumeil","year":"2019","unstructured":"Chaumeil PA, Mussig AJ, Hugenholtz P, Parks DH. GTDB-Tk: a toolkit to classify genomes with the Genome Taxonomy Database. Bioinformatics. 2019;36:1925\u20137.","journal-title":"Bioinformatics"},{"key":"2536_CR53","doi-asserted-by":"publisher","first-page":"2068","DOI":"10.1093\/bioinformatics\/btu153","volume":"30","author":"T Seemann","year":"2014","unstructured":"Seemann T. Prokka: rapid prokaryotic genome annotation. Bioinformatics. 2014;30:2068\u20139.","journal-title":"Bioinformatics"},{"key":"2536_CR54","doi-asserted-by":"publisher","first-page":"5825","DOI":"10.1093\/molbev\/msab293","volume":"38","author":"CP Cantalapiedra","year":"2021","unstructured":"Cantalapiedra CP, Hern\u00e1ndez-Plaza A, Letunic I, Bork P, Huerta-Cepas J. eggNOG-mapper v2: functional annotation, orthology assignments, and domain prediction at the metagenomic scale. Mol Biol Evol. 2021;38:5825\u20139.","journal-title":"Mol Biol Evol"},{"key":"2536_CR55","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1038\/s41564-020-00834-3","volume":"6","author":"AM Eren","year":"2021","unstructured":"Eren AM, Kiefl E, Shaiber A, Veseli I, Miller SE, Schechter MS, et al. Community-led, integrated, reproducible multi-omics with anvi\u2019o. Nat Microbiol. 2021;6:3\u20136.","journal-title":"Nat Microbiol"},{"key":"2536_CR56","doi-asserted-by":"publisher","first-page":"967","DOI":"10.1099\/ijsem.0.002593","volume":"68","author":"WB Whitman","year":"2018","unstructured":"Whitman WB, Oren A, Chuvochina M, da Costa MS, Garrity GM, Rainey FA, et al. Proposal of the suffix -ota to denote phyla. Addendum to \u2018Proposal to include the rank of phylum in the International Code of Nomenclature of Prokaryotes.\u2019 Int J Syst Evol Microbiol. 2018;68:967\u20139.","journal-title":"Int J Syst Evol Microbiol"},{"key":"2536_CR57","doi-asserted-by":"publisher","first-page":"754","DOI":"10.1093\/bioinformatics\/17.8.754","volume":"17","author":"JP Huelsenbeck","year":"2001","unstructured":"Huelsenbeck JP, Ronquist F. MrBayes: Bayesian inference of phylogenetic trees. Bioinformatics. 2001;17:754\u20135.","journal-title":"Bioinformatics"},{"key":"2536_CR58","doi-asserted-by":"publisher","first-page":"451","DOI":"10.1016\/j.mimet.2007.02.012","volume":"69","author":"J Bertaux","year":"2007","unstructured":"Bertaux J, Gloger U, Schmid M, Hartmann A, Scheu S. Routine fluorescence in situ hybridization in soil. J Microbiol Methods. 2007;69:451\u201360.","journal-title":"J Microbiol Methods"},{"key":"2536_CR59","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1264\/jsme2.ME12107","volume":"28","author":"K Kubota","year":"2013","unstructured":"Kubota K. CARD-FISH for environmental microorganisms: technical advancement and future applications. Microbes Environ. 2013;28:3\u201312.","journal-title":"Microbes Environ"},{"key":"2536_CR60","doi-asserted-by":"publisher","DOI":"10.3389\/fmicb.2019.01383","volume":"10","author":"M Lukumbuzya","year":"2019","unstructured":"Lukumbuzya M, Schmid M, Pjevac P, Daims H. A multicolor fluorescence in situ hybridization approach using an extended set of fluorophores to visualize microorganisms. Front Microbiol. 2019;10:1383.","journal-title":"Front Microbiol"},{"key":"2536_CR61","doi-asserted-by":"publisher","first-page":"420","DOI":"10.1038\/s41587-019-0036-z","volume":"37","author":"JJA Almagro Armenteros","year":"2019","unstructured":"Almagro Armenteros JJA, Tsirigos KD, S\u00f8nderby CK, Petersen TN, Winther O, Brunak S, et al. SignalP 5.0 improves signal peptide predictions using deep neural networks. Nat Biotechnol. 2019;37:420\u20133.","journal-title":"Nat Biotechnol"},{"key":"2536_CR62","doi-asserted-by":"publisher","first-page":"W402","DOI":"10.1093\/nar\/gkz297","volume":"47","author":"DWA Buchan","year":"2019","unstructured":"Buchan DWA, Jones DT. The PSIPRED protein analysis workbench: 20 years on. Nucleic Acids Res. 2019;47:W402-7.","journal-title":"Nucleic Acids Res"},{"key":"2536_CR63","doi-asserted-by":"publisher","first-page":"845","DOI":"10.1038\/nprot.2015.053","volume":"10","author":"LA Kelley","year":"2015","unstructured":"Kelley LA, Mezulis S, Yates CM, Wass MN, Sternberg MJE. The Phyre2 web portal for protein modeling, prediction and analysis. Nat Protoc. 2015;10:845\u201358.","journal-title":"Nat Protoc"},{"key":"2536_CR64","doi-asserted-by":"publisher","first-page":"679","DOI":"10.1038\/s41592-022-01488-1","volume":"19","author":"M Mirdita","year":"2022","unstructured":"Mirdita M, Sch\u00fctze K, Moriwaki Y, Heo L, Ovchinnikov S, Steinegger M. ColabFold: making protein folding accessible to all. Nat Methods. 2022;19:679\u201382.","journal-title":"Nat Methods"},{"key":"2536_CR65","doi-asserted-by":"publisher","first-page":"583","DOI":"10.1038\/s41586-021-03819-2","volume":"596","author":"J Jumper","year":"2021","unstructured":"Jumper J, Evans R, Pritzel A, Green T, Figurnov M, Ronneberger O, et al. Highly accurate protein structure prediction with AlphaFold. Nature. 2021;596:583\u20139.","journal-title":"Nature"},{"key":"2536_CR66","doi-asserted-by":"publisher","first-page":"W400","DOI":"10.1093\/nar\/gkh417","volume":"32","author":"PG Bagos","year":"2004","unstructured":"Bagos PG, Liakopoulos TD, Spyropoulos IC, Hamodrakas SJ. PRED-TMBB: a web server for predicting the topology of beta-barrel outer membrane proteins. Nucleic Acids Res. 2004;32:W400\u20134.","journal-title":"Nucleic Acids Res"},{"key":"2536_CR67","doi-asserted-by":"publisher","first-page":"e0107421","DOI":"10.1128\/mBio.01074-21","volume":"12","author":"JL Keffer","year":"2021","unstructured":"Keffer JL, McAllister SM, Garber AI, Hallahan BJ, Sutherland MC, Rozovsky S, et al. Iron oxidation by a fused cytochrome-porin common to diverse iron-oxidizing bacteria. MBio. 2021;12:e0107421.","journal-title":"MBio"},{"key":"2536_CR68","doi-asserted-by":"publisher","first-page":"4673","DOI":"10.1093\/nar\/22.22.4673","volume":"22","author":"JD Thompson","year":"1994","unstructured":"Thompson JD, Higgins DG, Gibson TJ. CLUSTAL W: improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position-specific gap penalties and weight matrix choice. Nucleic Acids Res. 1994;22:4673\u201380.","journal-title":"Nucleic Acids Res"},{"key":"2536_CR69","doi-asserted-by":"publisher","first-page":"744","DOI":"10.1006\/bbrc.1998.9549","volume":"251","author":"E Arslan","year":"1998","unstructured":"Arslan E, Schulz H, Zufferey R, K\u00fcnzler P, Th\u00f6ny-Meyer L. Overproduction of the Bradyrhizobium japonicum c-type cytochrome subunits of the cbb3 oxidase in Escherichia coli. Biochem Biophys Res Commun. 1998;251:744\u20137.","journal-title":"Biochem Biophys Res Commun"},{"key":"2536_CR70","doi-asserted-by":"publisher","first-page":"1100","DOI":"10.1038\/nbt.4232","volume":"36","author":"MJ Kwak","year":"2018","unstructured":"Kwak MJ, Kong HG, Choi K, Kwon SK, Song JY, Lee J, et al. Rhizosphere microbiome structure alters to enable wilt resistance in tomato. Nat Biotechnol. 2018;36:1100\u20139.","journal-title":"Nat Biotechnol"},{"key":"2536_CR71","doi-asserted-by":"publisher","first-page":"725","DOI":"10.1038\/nbt.3893","volume":"35","author":"RM Bowers","year":"2017","unstructured":"Bowers RM, Kyrpides NC, Stepanauskas R, Harmon-Smith M, Doud D, Reddy TBK, et al. Minimum information about a single amplified genome (MISAG) and a metagenome-assembled genome (MIMAG) of bacteria and archaea. Nat Biotechnol. 2017;35:725\u201331.","journal-title":"Nat Biotechnol"},{"key":"2536_CR72","doi-asserted-by":"publisher","first-page":"E1166","DOI":"10.1073\/pnas.1716667115","volume":"115","author":"PS Adam","year":"2018","unstructured":"Adam PS, Borrel G, Gribaldo S. Evolutionary history of carbon monoxide dehydrogenase\/acetyl-CoA synthase, one of the oldest enzymatic complexes. Proc Natl Acad Sci U S A. 2018;115:E1166-73.","journal-title":"Proc Natl Acad Sci U S A"},{"key":"2536_CR73","doi-asserted-by":"publisher","first-page":"360","DOI":"10.1038\/s41579-020-00502-7","volume":"19","author":"A Kappler","year":"2021","unstructured":"Kappler A, Bryce C, Mansor M, Lueder U, Byrne JM, Swanner ED. An evolving view on biogeochemical cycling of iron. Nat Rev Microbiol. 2021;19:360\u201374.","journal-title":"Nat Rev Microbiol"},{"key":"2536_CR74","doi-asserted-by":"publisher","DOI":"10.1128\/msystems.00553-19","volume":"5","author":"SM McAllister","year":"2020","unstructured":"McAllister SM, Polson SW, Butterfield DA, Glazer BT, Sylvan JB, Chan CS. Validating the Cyc2 neutrophilic iron oxidation pathway using meta-omics of Zetaproteobacteria iron mats at marine hydrothermal vents. mSystems. 2020;5:e00553\u201319.","journal-title":"mSystems"},{"key":"2536_CR75","doi-asserted-by":"publisher","DOI":"10.1128\/AEM.00496-21","volume":"87","author":"Y-M Huang","year":"2021","unstructured":"Huang Y-M, Straub D, Blackwell N, Kappler A, Kleindienst S. Meta-omics reveal Gallionellaceae and Rhodanobacter species as interdependent key players for Fe(II) oxidation and nitrate reduction in the autotrophic enrichment culture KS. Appl Environ Microbiol. 2021;87:e0049621.","journal-title":"Appl Environ Microbiol"},{"key":"2536_CR76","doi-asserted-by":"publisher","first-page":"1079","DOI":"10.1038\/s41587-020-0501-8","volume":"38","author":"DH Parks","year":"2020","unstructured":"Parks DH, Chuvochina M, Chaumeil PA, Rinke C, Mussig AJ, Hugenholtz P. A complete domain-to-species taxonomy for Bacteria and archaea. Nat Biotechnol. 2020;38:1079\u201386.","journal-title":"Nat Biotechnol"},{"key":"2536_CR77","doi-asserted-by":"publisher","first-page":"726","DOI":"10.1111\/1462-2920.14865","volume":"22","author":"P Geesink","year":"2020","unstructured":"Geesink P, Wegner CE, Probst AJ, Herrmann M, Dam HT, Kaster AK, et al. Genome-inferred spatio-temporal resolution of an uncultivated Roizmanbacterium reveals its ecological preferences in groundwater. Environ Microbiol. 2020;22:726\u201337.","journal-title":"Environ Microbiol"},{"key":"2536_CR78","doi-asserted-by":"publisher","DOI":"10.1016\/j.jhazmat.2024.135955","volume":"480","author":"Z Li","year":"2024","unstructured":"Li Z, He Y, Zhang H, Qian H, Wang Y. Biotransformations of arsenic in marine sediments across marginal slope to hadal zone. J Hazard Mater. 2024;480:135955. https:\/\/doi.org\/10.1016\/j.jhazmat.2024.135955.","journal-title":"J Hazard Mater"},{"key":"2536_CR79","doi-asserted-by":"publisher","first-page":"16076","DOI":"10.1038\/srep16076","volume":"5","author":"SK Kwon","year":"2015","unstructured":"Kwon SK, Kim SK, Lee DH, Kim JF. Comparative genomics and experimental evolution of Escherichia coli BL21(DE3) strains reveal the landscape of toxicity escape from membrane protein overproduction. Sci Rep. 2015;5:16076.","journal-title":"Sci Rep"},{"key":"2536_CR80","doi-asserted-by":"publisher","first-page":"573","DOI":"10.5194\/se-9-573-2018","volume":"9","author":"EC Chi Fru","year":"2018","unstructured":"Chi Fru EC, Kilias S, Ivarsson M, Rattray JE, Gkika K, McDonald I, et al. Sedimentary mechanisms of a modern banded iron formation on Milos Island. Greece Solid Earth. 2018;9:573\u201398.","journal-title":"Greece Solid Earth"},{"key":"2536_CR81","doi-asserted-by":"publisher","first-page":"451","DOI":"10.1038\/s41586-019-1436-4","volume":"572","author":"EJ Javaux","year":"2019","unstructured":"Javaux EJ. Challenges in evidencing the earliest traces of life. Nature. 2019;572:451\u201360.","journal-title":"Nature"},{"key":"2536_CR82","doi-asserted-by":"publisher","first-page":"R567","DOI":"10.1016\/j.cub.2009.05.054","volume":"19","author":"AL Sessions","year":"2009","unstructured":"Sessions AL, Doughty DM, Welander PV, Summons RE, Newman DK. The continuing puzzle of the great oxidation event. Curr Biol. 2009;19:R567\u201374.","journal-title":"Curr Biol"},{"key":"2536_CR83","doi-asserted-by":"publisher","first-page":"3462","DOI":"10.1111\/1462-2920.14328","volume":"20","author":"C Bryce","year":"2018","unstructured":"Bryce C, Blackwell N, Schmidt C, Otte J, Huang YM, Kleindienst S, et al. Microbial anaerobic Fe(II) oxidation \u2013 ecology, mechanisms and environmental implications. Environ Microbiol. 2018;20:3462\u201383.","journal-title":"Environ Microbiol"},{"key":"2536_CR84","doi-asserted-by":"publisher","first-page":"752","DOI":"10.1038\/nrmicro1490","volume":"4","author":"KA Weber","year":"2006","unstructured":"Weber KA, Achenbach LA, Coates JD. Microorganisms pumping iron: anaerobic microbial iron oxidation and reduction. Nat Rev Microbiol. 2006;4:752\u201364.","journal-title":"Nat Rev Microbiol"},{"key":"2536_CR85","doi-asserted-by":"publisher","first-page":"180","DOI":"10.1111\/gbi.12019","volume":"11","author":"N Klueglein","year":"2013","unstructured":"Klueglein N, Kappler A. Abiotic oxidation of Fe(II) by reactive nitrogen species in cultures of the nitrate-reducing Fe(II) oxidizer Acidovorax sp. BoFeN1\u2014questioning the existence of enzymatic Fe(II) oxidation. Geobiology. 2013;11:180\u201390.","journal-title":"Geobiology"},{"key":"2536_CR86","doi-asserted-by":"publisher","first-page":"1051","DOI":"10.1128\/AEM.03277-13","volume":"80","author":"N Klueglein","year":"2014","unstructured":"Klueglein N, Zeitvogel F, Stierhof YD, Floetenmeyer M, Konhauser KO, Kappler A, et al. Potential role of nitrite for abiotic Fe(II) oxidation and cell encrustation during nitrate reduction by denitrifying bacteria. Appl Environ Microbiol. 2014;80:1051\u201361.","journal-title":"Appl Environ Microbiol"},{"key":"2536_CR87","doi-asserted-by":"publisher","DOI":"10.1093\/ismeco\/ycae008","volume":"4","author":"F Corbera-Rubio","year":"2024","unstructured":"Corbera-Rubio F, Stouten GR, Bruins J, Dost SF, Merkel AY, M\u00fcller S, et al. \u201cCandidatus Siderophilus nitratireducens\u201d: a putative nap-dependent nitrate-reducing iron oxidizer within the new order Siderophiliales. ISME Commun. 2024;4:ycae008.","journal-title":"ISME Commun"},{"key":"2536_CR88","doi-asserted-by":"publisher","first-page":"2656","DOI":"10.1128\/AEM.03493-15","volume":"82","author":"S He","year":"2016","unstructured":"He S, Tominski C, Kappler A, Behrens S, Roden EE. Metagenomic analyses of the autotrophic Fe(II)-oxidizing, nitrate-reducing enrichment culture KS. Appl Environ Microbiol. 2016;82:2656\u201368.","journal-title":"Appl Environ Microbiol"},{"key":"2536_CR89","volume":"6","author":"D Emerson","year":"2015","unstructured":"Emerson D. The irony of iron \u2013 biogenic iron oxides as an iron source to the ocean. Front Microbiol. 2015;6:1502.","journal-title":"Front Microbiol"},{"key":"2536_CR90","doi-asserted-by":"publisher","first-page":"8193","DOI":"10.1073\/pnas.1505515112","volume":"112","author":"WQ Li","year":"2015","unstructured":"Li WQ, Beard BL, Johnson CM. Biologically recycled continental iron is a major component in banded iron formations. Proc Natl Acad Sci U S A. 2015;112:8193\u20138.","journal-title":"Proc Natl Acad Sci U S A"},{"key":"2536_CR91","doi-asserted-by":"publisher","first-page":"269","DOI":"10.1029\/2005EO290001","volume":"86","author":"E Domack","year":"2005","unstructured":"Domack E, Ishman S, Leventer A, Sylva S, Willmott V, Huber B. A chemotrophic ecosystem found beneath Antarctic ice shelf. Eos Trans Am Geophys Union. 2005;86:269\u201372.","journal-title":"Eos Trans Am Geophys Union"},{"key":"2536_CR92","doi-asserted-by":"publisher","first-page":"671","DOI":"10.1038\/s41579-018-0046-8","volume":"16","author":"WD Orsi","year":"2018","unstructured":"Orsi WD. Ecology and evolution of seafloor and subseafloor microbial communities. Nat Rev Microbiol. 2018;16:671\u201383.","journal-title":"Nat Rev Microbiol"},{"key":"2536_CR93","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.1600983","volume":"3","author":"PF Hoffman","year":"2017","unstructured":"Hoffman PF, Abbot DS, Ashkenazy Y, Benn DI, Brocks JJ, Cohen PA, et al. Snowball Earth climate dynamics and Cryogenian geology-geobiology. Sci Adv. 2017;3:e1600983.","journal-title":"Sci Adv"},{"key":"2536_CR94","doi-asserted-by":"publisher","first-page":"535","DOI":"10.1038\/nature19355","volume":"537","author":"AP Nutman","year":"2016","unstructured":"Nutman AP, Bennett VC, Friend CRL, Van Kranendonk MJ, Chivas AR. Rapid emergence of life shown by discovery of 3,700-million-year-old microbial structures. Nature. 2016;537:535\u20138.","journal-title":"Nature"},{"key":"2536_CR95","doi-asserted-by":"publisher","first-page":"60","DOI":"10.1038\/nature21377","volume":"543","author":"MS Dodd","year":"2017","unstructured":"Dodd MS, Papineau D, Grenne T, Slack JF, Rittner M, Pirajno F, et al. Evidence for early life in Earth\u2019s oldest hydrothermal vent precipitates. Nature. 2017;543:60\u20134.","journal-title":"Nature"},{"key":"2536_CR96","doi-asserted-by":"publisher","first-page":"161","DOI":"10.1016\/j.bbabio.2012.10.001","volume":"1827","author":"M Ilbert","year":"2013","unstructured":"Ilbert M, Bonnefoy V. Insight into the evolution of the iron oxidation pathways. Biochim Biophys Acta. 2013;1827:161\u201375.","journal-title":"Biochim Biophys Acta"},{"key":"2536_CR97","doi-asserted-by":"publisher","DOI":"10.3389\/fmicb.2018.00513","volume":"9","author":"A Price","year":"2018","unstructured":"Price A, Pearson VK, Schwenzer SP, Miot J, Olsson-Francis K. Nitrate-dependent iron oxidation: a potential Mars metabolism. Front Microbiol. 2018;9:513.","journal-title":"Front Microbiol"}],"container-title":["Microbiome"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s40168-026-02536-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s40168-026-02536-0","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s40168-026-02536-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,7]],"date-time":"2026-10-07T23:02:36Z","timestamp":1791414156000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1186\/s40168-026-02536-0"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,10,8]]},"references-count":97,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["2536"],"URL":"https:\/\/doi.org\/10.1186\/s40168-026-02536-0","relation":{},"ISSN":["2049-2618"],"issn-type":[{"value":"2049-2618","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,10,8]]},"assertion":[{"value":"16 April 2026","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"25 August 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"8 October 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"Not applicable.","order":1,"name":"Ethics","label":"Ethics approval and consent to participate","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"Not applicable.","order":2,"name":"Ethics","label":"Consent for publication","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors declare that they have no competing interests.","order":3,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Declarations"}}],"article-number":"225"}}