{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,11]],"date-time":"2026-09-11T05:04:01Z","timestamp":1789103041193,"version":"build-2803163510"},"reference-count":115,"publisher":"MDPI AG","issue":"3","license":[{"start":{"date-parts":[[2021,3,16]],"date-time":"2021-03-16T00:00:00Z","timestamp":1615852800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100012627","name":"NASA Astrobiology Institute","doi-asserted-by":"publisher","award":["NN13AI60G"],"award-info":[{"award-number":["NN13AI60G"]}],"id":[{"id":"10.13039\/100012627","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["OPP-1745341"],"award-info":[{"award-number":["OPP-1745341"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["EAR-1637066"],"award-info":[{"award-number":["EAR-1637066"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]},{"name":"McMurdo Long Term Ecological Research","award":["OPP-115245"],"award-info":[{"award-number":["OPP-115245"]}]},{"name":"McMurdo Long Term Ecological Research","award":["OPP-1637708"],"award-info":[{"award-number":["OPP-1637708"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Genes"],"abstract":"<jats:p>Sulfide inhibits oxygenic photosynthesis by blocking electron transfer between H2O and the oxygen-evolving complex in the D1 protein of Photosystem II. The ability of cyanobacteria to counter this effect has implications for understanding the productivity of benthic microbial mats in sulfidic environments throughout Earth history. In Lake Fryxell, Antarctica, the benthic, filamentous cyanobacterium Phormidium pseudopriestleyi creates a 1\u20132 mm thick layer of 50 \u00b5mol L\u22121 O2 in otherwise sulfidic water, demonstrating that it sustains oxygenic photosynthesis in the presence of sulfide. A metagenome-assembled genome of P. pseudopriestleyi indicates a genetic capacity for oxygenic photosynthesis, including multiple copies of psbA (encoding the D1 protein of Photosystem II), and anoxygenic photosynthesis with a copy of sqr (encoding the sulfide quinone reductase protein that oxidizes sulfide). The genomic content of P. pseudopriestleyi is consistent with sulfide tolerance mechanisms including increasing psbA expression or directly oxidizing sulfide with sulfide quinone reductase. However, the ability of the organism to reduce Photosystem I via sulfide quinone reductase while Photosystem II is sulfide-inhibited, thereby performing anoxygenic photosynthesis in the presence of sulfide, has yet to be demonstrated.<\/jats:p>","DOI":"10.3390\/genes12030426","type":"journal-article","created":{"date-parts":[[2021,3,16]],"date-time":"2021-03-16T12:01:02Z","timestamp":1615896062000},"page":"426","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":27,"title":["Metabolic Capacity of the Antarctic Cyanobacterium Phormidium pseudopriestleyi That Sustains Oxygenic Photosynthesis in the Presence of Hydrogen Sulfide"],"prefix":"10.3390","volume":"12","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-6604-4618","authenticated-orcid":false,"given":"Jessica E.","family":"Lumian","sequence":"first","affiliation":[{"name":"Microbiology Graduate Group, University of California, Davis, CA 95616, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anne D.","family":"Jungblut","sequence":"additional","affiliation":[{"name":"Life Sciences Department, The Natural History Museum, London SW7 5BD, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Megan L.","family":"Dillion","sequence":"additional","affiliation":[{"name":"Genomics and Bioinformatics, Novozymes, Inc., Davis, CA 95616, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2471-6903","authenticated-orcid":false,"given":"Ian","family":"Hawes","sequence":"additional","affiliation":[{"name":"Coastal Marine Field Station, University of Waikato, Tauranga 3110, New Zealand"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Peter T.","family":"Doran","sequence":"additional","affiliation":[{"name":"Geology and Geophysics, Louisiana State University, Baton Rouge, LA 70803, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tyler J.","family":"Mackey","sequence":"additional","affiliation":[{"name":"Department of Earth and Planetary Sciences, University of New Mexico, Albuquerque, NM 87131, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gregory J.","family":"Dick","sequence":"additional","affiliation":[{"name":"Department of Earth and Environmental Sciences, University of Michigan, Ann Arbor, MI 48109, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Christen L.","family":"Grettenberger","sequence":"additional","affiliation":[{"name":"Department of Earth and Planetary Sciences, University of California, Davis, CA 95616, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7343-2061","authenticated-orcid":false,"given":"Dawn Y.","family":"Sumner","sequence":"additional","affiliation":[{"name":"Department of Earth and Planetary Sciences, University of California, Davis, CA 95616, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2021,3,16]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1016\/j.chemgeo.2013.05.039","article-title":"What Caused the Rise of Atmospheric O2?","volume":"362","author":"Kasting","year":"2013","journal-title":"Chem. Geol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"450","DOI":"10.1038\/24839","article-title":"A New Model for Proterozoic Ocean Chemistry","volume":"396","author":"Canfield","year":"1998","journal-title":"Nature"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"850","DOI":"10.1144\/jgs2016-147","article-title":"Microbial Sulphate Reduction during Neoproterozoic Glaciation, Port Askaig Formation, UK","volume":"174","author":"Parnell","year":"2017","journal-title":"J. Geol. Soc."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"264","DOI":"10.1016\/j.epsl.2011.09.016","article-title":"Molybdenum Evidence for Expansive Sulfidic Water Masses in ~750Ma Oceans","volume":"311","author":"Dahl","year":"2011","journal-title":"Earth Planet. Sci. Lett."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Wang, P., Algeo, T.J., Zhou, Q., Yu, W., Du, Y., Qin, Y., Xu, Y., Yuan, L., and Pan, W. (2019). Large Accumulations of 34S-Enriched Pyrite in a Low-Sulfate Marine Basin: The Sturtian Nanhua Basin, South China. Precambrian Res., 335.","DOI":"10.1016\/j.precamres.2019.105504"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"250","DOI":"10.1111\/j.1472-4669.2012.00322.x","article-title":"Cyanobacterial Life at Low O2: Community Genomics and Function Reveal Metabolic Versatility and Extremely Low Diversity in a Great Lakes Sinkhole Mat","volume":"10","author":"Voorhies","year":"2012","journal-title":"Geobiology"},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Whitton, B.A. (2012). Cyanobacteria in the Cryosphere: Snow, Ice and Extreme Cold. Ecology of Cyanobacteria II: Their Diversity in Space and Time, Springer.","DOI":"10.1007\/978-94-007-3855-3"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"921","DOI":"10.1038\/ismej.2015.167","article-title":"Structure and Function of Natural Sulphide-Oxidizing Microbial Mats under Dynamic Input of Light and Chemical Energy","volume":"10","author":"Klatt","year":"2016","journal-title":"ISME J."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"5596","DOI":"10.1128\/AEM.00715-06","article-title":"Dominant Microbial Populations in Limestone-Corroding Stream Biofilms, Frasassi Cave System, Italy","volume":"72","author":"Macalady","year":"2006","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"1251","DOI":"10.1111\/1462-2920.13654","article-title":"Oxygenic and Anoxygenic Photosynthesis in a Microbial Mat from an Anoxic and Sulfidic Spring","volume":"19","author":"Weber","year":"2017","journal-title":"Environ. Microbiol."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"568","DOI":"10.1038\/ismej.2017.193","article-title":"Cyanobacterial Photosynthesis under Sulfidic Conditions: Insights from the Isolate Leptolyngbya Sp. Strain Hensonii","volume":"12","author":"Hamilton","year":"2018","journal-title":"ISME J."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"887","DOI":"10.1130\/G36966.1","article-title":"Antarctic Microbial Mats: A Modern Analog for Archean Lacustrine Oxygen Oases","volume":"43","author":"Sumner","year":"2015","journal-title":"Geology"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"620","DOI":"10.1128\/AEM.02699-15","article-title":"Microbial Mat Communities along an Oxygen Gradient in a Perennially Ice-Covered Antarctic Lake","volume":"82","author":"Jungblut","year":"2016","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"270","DOI":"10.1016\/0005-2728(79)90045-8","article-title":"Sulfide Inhibition of Photosystem II in Cyanobacteria (Blue-Green Algae) and Tobacco Chloroplasts","volume":"546","author":"Oren","year":"1979","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"736","DOI":"10.1128\/AEM.70.2.736-744.2004","article-title":"Variation in Sulfide Tolerance of Photosystem II in Phylogenetically Diverse Cyanobacteria from Sulfidic Habitats","volume":"70","author":"Miller","year":"2004","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"398","DOI":"10.1128\/aem.51.2.398-407.1986","article-title":"Adaptation to Hydrogen Sulfide of Oxygenic and Anoxygenic Photosynthesis among Cyanobacteria","volume":"51","author":"Cohen","year":"1986","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Dick, G.J., Grim, S.L., and Klatt, J.M. (2018). Controls on O2 Production in Cyanobacterial Mats and Implications for Earth\u2019s Oxygenation. Annu. Rev. Earth Planet. Sci., 46.","DOI":"10.1146\/annurev-earth-082517-010035"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"3301","DOI":"10.1111\/1462-2920.12791","article-title":"Hydrogen Sulfide Can Inhibit and Enhance Oxygenic Photosynthesis in a Cyanobacterium from Sulfidic Springs","volume":"17","author":"Klatt","year":"2015","journal-title":"Environ. Microbiol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1093\/nar\/28.1.27","article-title":"KEGG: Kyoto Encyclopedia of Genes and Genomes","volume":"28","author":"Kanehisa","year":"2000","journal-title":"Nucleic Acids Res."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"1947","DOI":"10.1002\/pro.3715","article-title":"Toward Understanding the Origin and Evolution of Cellular Organisms","volume":"28","author":"Kanehisa","year":"2019","journal-title":"Protein Sci. Publ. Protein Soc."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1016\/0006-3002(58)90313-5","article-title":"The Influence of the Herbicide, DCMU, on the Oxygen-Evolving System of Photosynthesis","volume":"27","author":"Bishop","year":"1958","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"3697","DOI":"10.1007\/s00018-009-0103-6","article-title":"Cyanobacterial PsbA Gene Family: Optimization of Oxygenic Photosynthesis","volume":"66","author":"Mulo","year":"2009","journal-title":"Cell. Mol. Life Sci."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"247","DOI":"10.1016\/j.bbabio.2011.04.011","article-title":"Strategies for PsbA Gene Expression in Cyanobacteria, Green Algae and Higher Plants: From Transcription to PSII Repair","volume":"1817","author":"Mulo","year":"2012","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"12939","DOI":"10.1021\/bi8018916","article-title":"Low-Oxygen Induction of Normally Cryptic PsbA Genes in Cyanobacteria","volume":"47","author":"Summerfield","year":"2008","journal-title":"Biochemistry"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"1310","DOI":"10.1093\/molbev\/msv024","article-title":"Origin and Evolution of Water Oxidation before the Last Common Ancestor of the Cyanobacteria","volume":"32","author":"Cardona","year":"2015","journal-title":"Mol. Biol. Evol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"3336","DOI":"10.1128\/JB.182.12.3336-3344.2000","article-title":"Cyanobacterial Sulfide-Quinone Reductase: Cloning and Heterologous Expression","volume":"182","author":"Bronstein","year":"2000","journal-title":"J. Bacteriol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"623","DOI":"10.1128\/jb.129.2.623-629.1977","article-title":"Occurrence of Facultative Anoxygenic Photosynthesis among Filamentous and Unicellular Cyanobacteria","volume":"129","author":"Garlick","year":"1997","journal-title":"J. Bacteriol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"855","DOI":"10.1128\/jb.123.3.855-861.1975","article-title":"Facultative Anoxygenic Photosynthesis in the Cyanobacterium Oscillatoria Limnetica","volume":"123","author":"Cohen","year":"1975","journal-title":"J. Bacteriol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"489","DOI":"10.1038\/257489a0","article-title":"Sulphide-Dependent Anoxygenic Photosynthesis in the Cyanobacterium Oscillatoria Limnetica","volume":"257","author":"Cohen","year":"1975","journal-title":"Nature"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"321","DOI":"10.1007\/s10498-008-9052-1","article-title":"The Saline Lakes of the McMurdo Dry Valleys, Antarctica","volume":"15","author":"Green","year":"2009","journal-title":"Aquat. Geochem."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"985","DOI":"10.2307\/1313732","article-title":"Dry Valley Streams in Antarctica: Ecosystems Waiting for Water","volume":"49","author":"McKnight","year":"1999","journal-title":"BioScience"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"543","DOI":"10.1080\/00288306.1985.10421206","article-title":"Water Column and Sediment Characteristics of Lake Fryxell, Taylor Valley, Antarctica","volume":"28","author":"Lawrence","year":"1985","journal-title":"N. Z. J. Geol. Geophys."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"1215","DOI":"10.2307\/1937286","article-title":"Production Strategies in Antarctic Inland Waters: Phytoplankton Eco-Physiology in a Permanently Ice-Covered Lake","volume":"62","author":"Vincent","year":"1981","journal-title":"Ecology"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"527","DOI":"10.1130\/0091-7613(1999)027<0527:HOMDVL>2.3.CO;2","article-title":"History of McMurdo Dry Valley Lakes, Antarctica, from Stable Chlorine Isotope Data","volume":"27","author":"Lyons","year":"1999","journal-title":"Geology"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"649","DOI":"10.1046\/j.1365-2427.2000.00612.x","article-title":"Stratification and Dynamics of Microbial Loop Communities in Lake Fryxell, Antarctica","volume":"44","author":"Roberts","year":"2000","journal-title":"Freshw. Biol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"54","DOI":"10.1007\/s003000050104","article-title":"The Microbial Plankton of Lake Fryxell, Southern Victoria Land, Antarctica during the Summers of 1992 and 1994","volume":"17","author":"James","year":"1997","journal-title":"Polar Biol."},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Spigel, R.H., and Priscu, J.C. (2013). Physical Limnology of the McMurdo Dry Valleys Lakes. Ecosystem Dynamics in a Polar Desert: The McMurdo Dry Valleys, Antarctica, American Geophysical Union (AGU).","DOI":"10.1029\/AR072p0153"},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Howard-Williams, C., Schwarz, A., Hawes, I., and Priscu, J.C. (2013). Optical Properties of the McMurdo Dry Valley Lakes, Antarctica. Ecosystem Dynamics in a Polar Desert: The McMurdo Dry Valleys, Antarctica, American Geophysical Union (AGU).","DOI":"10.1029\/AR072p0189"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"179","DOI":"10.1111\/j.0022-3646.1992.00179.x","article-title":"Photosynthesis-Irradiance Relationships in Phytoplankton from the Physically Stable Water Column of a Perennially Ice-Covered Lake (Lake Bonney, Antarctica)","volume":"28","author":"Lizotte","year":"1992","journal-title":"J. Phycol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"130","DOI":"10.1002\/j.1551-8833.2006.tb07570.x","article-title":"Standard Methods for the Examination of Water and Wastewater, 21st Edition","volume":"98","author":"Miner","year":"2006","journal-title":"Am. Water Works Assoc. J."},{"key":"ref_41","unstructured":"Hillman, C. (2018, November 28). Structure of Benthic Microbial Mat Assemblages in Lake Fryxell, Antarctica. Available online: https:\/\/ir.canterbury.ac.nz\/handle\/10092\/8737."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1099\/00221287-111-1-1","article-title":"Generic Assignments, Strain Histories and Properties of Pure Cultures of Cyanobacteria","volume":"111","author":"Rippka","year":"1979","journal-title":"Microbiology"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"614","DOI":"10.1093\/bioinformatics\/btt593","article-title":"PEAR: A Fast and Accurate Illumina Paired-End ReAd MergeR","volume":"30","author":"Zhang","year":"2014","journal-title":"Bioinformatics"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"2114","DOI":"10.1093\/bioinformatics\/btu170","article-title":"Trimmomatic: A Flexible Trimmer for Illumina Sequence Data","volume":"30","author":"Bolger","year":"2014","journal-title":"Bioinformatics"},{"key":"ref_45","doi-asserted-by":"crossref","unstructured":"Crusoe, M.R., Alameldin, H.F., Awad, S., Boucher, E., Caldwell, A., Cartwright, R., Charbonneau, A., Constantinides, B., Edvenson, G., and Fay, S. (2015). The Khmer Software Package: Enabling Efficient Nucleotide Sequence Analysis. F1000Research, 4.","DOI":"10.12688\/f1000research.6924.1"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"1674","DOI":"10.1093\/bioinformatics\/btv033","article-title":"MEGAHIT: An Ultra-Fast Single-Node Solution for Large and Complex Metagenomics Assembly via Succinct de Bruijn Graph","volume":"31","author":"Li","year":"2015","journal-title":"Bioinformatics"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"1072","DOI":"10.1093\/bioinformatics\/btt086","article-title":"QUAST: Quality Assessment Tool for Genome Assemblies","volume":"29","author":"Gurevich","year":"2013","journal-title":"Bioinformatics"},{"key":"ref_48","unstructured":"Li, H. (2013). Aligning Sequence Reads, Clone Sequences and Assembly Contigs with BWA-MEM. arXiv, Available online: http:\/\/arxiv.org\/abs\/1303.3997."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"2078","DOI":"10.1093\/bioinformatics\/btp352","article-title":"The Sequence Alignment\/Map Format and SAMtools","volume":"25","author":"Li","year":"2009","journal-title":"Bioinformatics"},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Eren, A.M., Esen, \u00d6.C., Quince, C., Vineis, J.H., Morrison, H.G., Sogin, M.L., and Delmont, T.O. (2015). Anvi\u2019o: An Advanced Analysis and Visualization Platform for \u2018omics Data. PeerJ, 3.","DOI":"10.7717\/peerj.1319"},{"key":"ref_51","doi-asserted-by":"crossref","unstructured":"Delmont, T.O., and Eren, A.M. (2018). Linking Pangenomes and Metagenomes: The Prochlorococcus Metapangenome. PeerJ, 6.","DOI":"10.7717\/peerj.4320"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"1043","DOI":"10.1101\/gr.186072.114","article-title":"CheckM: Assessing the Quality of Microbial Genomes Recovered from Isolates, Single Cells, and Metagenomes","volume":"25","author":"Parks","year":"2015","journal-title":"Genome Res."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"1925","DOI":"10.1093\/bioinformatics\/btz848","article-title":"GTDB-Tk: A Toolkit to Classify Genomes with the Genome Taxonomy Database","volume":"36","author":"Chaumeil","year":"2020","journal-title":"Bioinformatics"},{"key":"ref_54","doi-asserted-by":"crossref","unstructured":"Hyatt, D., Chen, G., Locascio, P.F., Land, M.L., Larimer, F.W., and Hauser, L.J. (2010). Prodigal: Prokaryotic Gene Recognition and Translation Initiation Site Identification. BMC Bioinform., 11.","DOI":"10.1186\/1471-2105-11-119"},{"key":"ref_55","doi-asserted-by":"crossref","unstructured":"Eddy, S.R. (2011). Accelerated Profile HMM Searches. PLoS Comput. Biol., 7.","DOI":"10.1371\/journal.pcbi.1002195"},{"key":"ref_56","doi-asserted-by":"crossref","unstructured":"Matsen, F.A., Kodner, R.B., and Armbrust, E.V. (2010). Pplacer: Linear Time Maximum-Likelihood and Bayesian Phylogenetic Placement of Sequences onto a Fixed Reference Tree. BMC Bioinform., 11.","DOI":"10.1186\/1471-2105-11-538"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"5114","DOI":"10.1038\/s41467-018-07641-9","article-title":"High Throughput ANI Analysis of 90K Prokaryotic Genomes Reveals Clear Species Boundaries","volume":"9","author":"Jain","year":"2018","journal-title":"Nat. Commun."},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"Price, M.N., Dehal, P.S., and Arkin, A.P. (2010). FastTree 2\u2014Approximately Maximum-Likelihood Trees for Large Alignments. PLoS ONE, 5.","DOI":"10.1371\/journal.pone.0009490"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"132","DOI":"10.1186\/s13059-016-0997-x","article-title":"Mash: Fast Genome and Metagenome Distance Estimation Using MinHash","volume":"17","author":"Ondov","year":"2016","journal-title":"Genome Biol."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"1569","DOI":"10.1093\/bioinformatics\/btq228","article-title":"DendroPy: A Python Library for Phylogenetic Computing","volume":"26","author":"Sukumaran","year":"2010","journal-title":"Bioinformatics"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"357","DOI":"10.1038\/s41586-020-2649-2","article-title":"Array Programming with NumPy","volume":"585","author":"Harris","year":"2020","journal-title":"Nature"},{"key":"ref_62","first-page":"726","article-title":"BlastKOALA and GhostKOALA: KEGG Tools for Functional Characterization of Genome and Metagenome Sequences","volume":"428","author":"Kanehisa","year":"2016","journal-title":"J. Mol. Biol. Comput. Resour. Mol. Biol."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"2068","DOI":"10.1093\/bioinformatics\/btu153","article-title":"Prokka: Rapid Prokaryotic Genome Annotation","volume":"30","author":"Seemann","year":"2014","journal-title":"Bioinformatics"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"566","DOI":"10.1038\/nbt.4163","article-title":"KBase: The United States Department of Energy Systems Biology Knowledgebase","volume":"36","author":"Arkin","year":"2018","journal-title":"Nat. Biotechnol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"164","DOI":"10.1186\/s13059-020-02066-4","article-title":"Exploring Neighborhoods in Large Metagenome Assembly Graphs Using Spacegraphcats Reveals Hidden Sequence Diversity","volume":"21","author":"Brown","year":"2020","journal-title":"Genome Biol."},{"key":"ref_66","doi-asserted-by":"crossref","unstructured":"Camacho, C., Coulouris, G., Avagyan, V., Ma, N., Papadopoulos, J., Bealer, K., and Madden, T.L. (2009). BLAST+: Architecture and Applications. BMC Bioinform., 10.","DOI":"10.1186\/1471-2105-10-421"},{"key":"ref_67","doi-asserted-by":"crossref","unstructured":"Miller, M.A., Pfeiffer, W., and Schwartz, T. (2010, January 14). Creating the CIPRES Science Gateway for Inference of Large Phylogenetic Trees. Proceedings of the 2010 Gateway Computing Environments Workshop (GCE), New Orleans, LA, USA.","DOI":"10.1109\/GCE.2010.5676129"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"2947","DOI":"10.1093\/bioinformatics\/btm404","article-title":"Clustal W and Clustal X Version 2.0","volume":"23","author":"Larkin","year":"2007","journal-title":"Bioinformatics"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"291","DOI":"10.1093\/molbev\/msz189","article-title":"ModelTest-NG: A New and Scalable Tool for the Selection of DNA and Protein Evolutionary Models","volume":"37","author":"Darriba","year":"2020","journal-title":"Mol. Biol. Evol."},{"key":"ref_70","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","year":"2014","journal-title":"Bioinformatics"},{"key":"ref_71","doi-asserted-by":"crossref","unstructured":"Shahak, Y., and Hauska, H. (2008). Advances in Photosynthesis and Respiration. Sulfur Metabolism in Phototrophic Organisms, Sulfide Oxidation from Cyanobacteria to Humans: Sulfide\u2013Quinone Oxidoreductase (SQR), Springer.","DOI":"10.1007\/978-1-4020-6863-8_16"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"1972","DOI":"10.1093\/bioinformatics\/btp348","article-title":"TrimAl: A Tool for Automated Alignment Trimming in Large-Scale Phylogenetic Analyses","volume":"25","year":"2009","journal-title":"Bioinformatics"},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"5279","DOI":"10.1111\/mec.14953","article-title":"Photoecology of the Antarctic Cyanobacterium Leptolyngbya sp. BC1307 Brought to Light through Community Analysis, Comparative Genomics and in Vitro Photosphysiology","volume":"27","author":"Chrismas","year":"2018","journal-title":"Mol. Ecol."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"2142","DOI":"10.1038\/s41396-020-0668-5","article-title":"A Phylogenetically Novel Cyanobacterium Most Closely Related to Gloeobacter","volume":"14","author":"Grettenberger","year":"2020","journal-title":"ISME J."},{"key":"ref_75","doi-asserted-by":"crossref","unstructured":"Tang, J., Du, L.-M., Lian, Y.-M., and Daroch, M. (2019). Complete Genome Sequence and Comparative Analysis of Synechococcus sp. CS-601 (SynAce01), a Cold-Adapted Cyanobacterium from an Oligotrophic Antarctic Habitat. Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20010152"},{"key":"ref_76","doi-asserted-by":"crossref","unstructured":"Lara, Y., Durieu, B., Cornet, L., Verlaine, O., Rippka, R., Pessi, I.S., Misztak, A., Joris, B., Javaux, E.J., and Baurain, D. (2017). Draft Genome Sequence of the Axenic Strain Phormidesmis priestleyi ULC007, a Cyanobacterium Isolated from Lake Bruehwiler (Larsemann Hills). Genome Announc., 5.","DOI":"10.1128\/genomeA.01546-16"},{"key":"ref_77","doi-asserted-by":"crossref","unstructured":"Chrismas, N.A.M., Barker, G., Anesio, A.M., and S\u00e1nchez-Baracaldo, P. (2016). Genomic Mechanisms for Cold Tolerance and Production of Exopolysaccharides in the Arctic Cyanobacterium Phormidesmis Priestleyi BC1401. BMC Genom., 17.","DOI":"10.1186\/s12864-016-2846-4"},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"1053","DOI":"10.1073\/pnas.1217107110","article-title":"Improving the Coverage of the Cyanobacterial Phylum Using Diversity-Driven Genome Sequencing","volume":"110","author":"Shih","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_79","doi-asserted-by":"crossref","unstructured":"Rodriguez-R, L.M., and Konstantinidis, K.T. (2016). The Enveomics Collection: A Toolbox for Specialized Analyses of Microbial Genomes and Metagenomes. PeerJ Inc.","DOI":"10.7287\/peerj.preprints.1900"},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"519","DOI":"10.1111\/j.1462-2920.2005.00717.x","article-title":"Diversity within Cyanobacterial Mat Communities in Variable Salinity Meltwater Ponds of McMurdo Ice Shelf, Antarctica","volume":"7","author":"Jungblut","year":"2005","journal-title":"Environ. Microbiol."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"272","DOI":"10.1111\/j.1574-6941.2006.00110.x","article-title":"Biogeographical Distribution and Ecological Ranges of Benthic Cyanobacteria in East Antarctic Lakes","volume":"57","author":"Taton","year":"2006","journal-title":"FEMS Microbiol. Ecol."},{"key":"ref_82","doi-asserted-by":"crossref","unstructured":"Barco, R.A., Garrity, G.M., Scott, J.J., Amend, J.P., Nealson, K.H., and Emerson, D. (2020). A Genus Definition for Bacteria and Archaea Based on a Standard Genome Relatedness Index. MBio, 11.","DOI":"10.1128\/mBio.02475-19"},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"369","DOI":"10.1146\/annurev.pp.26.060175.002101","article-title":"Phycobiliproteins and Complementary Chromatic Adaptation","volume":"26","author":"Bogorad","year":"1975","journal-title":"Annu. Rev. Plant Physiol."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"7229","DOI":"10.1074\/jbc.M311565200","article-title":"The Efficient Functioning of Photosynthesis and Respiration in Synechocystis sp. PCC 6803 Strictly Requires the Presence of Either Cytochrome c6 or Plastocyanin","volume":"279","author":"Navarro","year":"2004","journal-title":"J. Biol. Chem."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"3430","DOI":"10.1128\/JB.01864-14","article-title":"Coregulated Genes Link Sulfide:Quinone Oxidoreductase and Arsenic Metabolism in Synechocystis Sp. Strain PCC6803","volume":"196","author":"Nagy","year":"2014","journal-title":"J. Bacteriol."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"247","DOI":"10.1016\/j.bbabio.2015.10.007","article-title":"Photosynthetic, Respiratory and Extracellular Electron Transport Pathways in Cyanobacteria","volume":"1857","author":"Bombelli","year":"2016","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_87","doi-asserted-by":"crossref","unstructured":"Mullineaux, C.W. (2014). Electron Transport and Light-Harvesting Switches in Cyanobacteria. Front. Plant Sci., 5.","DOI":"10.3389\/fpls.2014.00007"},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"177","DOI":"10.1007\/s11120-011-9714-5","article-title":"Sequence Variation at the Oxygen-Evolving Centre of Photosystem II: A New Class of \u2018Rogue\u2019 Cyanobacterial D1 Proteins","volume":"110","author":"Murray","year":"2012","journal-title":"Photosynth. Res."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"5325","DOI":"10.1021\/bi00474a017","article-title":"Glu-69 of the D2 Protein in Photosystem II Is a Potential Ligand to Mn Involved in Photosynthetic Oxygen Evolution","volume":"29","author":"Vermaas","year":"1990","journal-title":"Biochemistry"},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"1901","DOI":"10.4319\/lo.2010.55.5.1901","article-title":"Metagenomic Profiling of Arctic Microbial Mat Communities as Nutrient Scavenging and Recycling Systems","volume":"55","author":"Varin","year":"2010","journal-title":"Limnol. Oceanogr."},{"key":"ref_91","doi-asserted-by":"crossref","unstructured":"Dillon, M.L., Hawes, I., Jungblut, A.D., Mackey, T.J., Eisen, J.A., Doran, P.T., and Sumner, D.Y. (2020). Energetic and Environmental Constraints on the Community Structure of Benthic Microbial Mats in Lake Fryxell, Antarctica. FEMS Microbiol. Ecol., 96.","DOI":"10.1093\/femsec\/fiz207"},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"2288","DOI":"10.1111\/1462-2920.12689","article-title":"The FlxABCD-HdrABC Proteins Correspond to a Novel NADH Dehydrogenase\/Heterodisulfide Reductase Widespread in Anaerobic Bacteria and Involved in Ethanol Metabolism in Desulfovibrio Vulgaris Hildenborough","volume":"17","author":"Ramos","year":"2015","journal-title":"Environ. Microbiol."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"277","DOI":"10.1042\/BJ20040746","article-title":"Stabilization of Model Membranes during Drying by Compatible Solutes Involved in the Stress Tolerance of Plants and Microorganisms","volume":"383","author":"Hincha","year":"2004","journal-title":"Biochem. J."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"645","DOI":"10.1128\/aem.57.3.645-648.1991","article-title":"Accumulation of Trehalose and Sucrose in Cyanobacteria Exposed to Matric Water Stress","volume":"57","author":"Hershkovitz","year":"1991","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"519","DOI":"10.1016\/S0304-4165(99)00155-5","article-title":"Involvement of the Compatible Solutes Trehalose and Sucrose in the Response to Salt Stress of a Cyanobacterial Scytonema Species Isolated from Desert Soils","volume":"1472","author":"Behm","year":"1999","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"800","DOI":"10.1093\/icb\/45.5.800","article-title":"Desiccation Tolerance of Prokaryotes: Application of Principles to Human Cells","volume":"45","author":"Potts","year":"2005","journal-title":"Integr. Comp. Biol."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"87","DOI":"10.1111\/j.1574-6976.2010.00234.x","article-title":"Molecular Biology of Cyanobacterial Salt Acclimation","volume":"35","author":"Hagemann","year":"2011","journal-title":"FEMS Microbiol. Rev."},{"key":"ref_98","doi-asserted-by":"crossref","unstructured":"Kirsch, F., Kl\u00e4hn, S., and Hagemann, M. (2019). Salt-Regulated Accumulation of the Compatible Solutes Sucrose and Glucosylglycerol in Cyanobacteria and Its Biotechnological Potential. Front. Microbiol., 10.","DOI":"10.3389\/fmicb.2019.02139"},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"329","DOI":"10.1080\/09670269910001736392","article-title":"UV Protection in Cyanobacteria","volume":"34","author":"Scherer","year":"1999","journal-title":"Eur. J. Phycol."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"463","DOI":"10.3390\/metabo3020463","article-title":"Multiple Roles of Photosynthetic and Sunscreen Pigments in Cyanobacteria Focusing on the Oxidative Stress","volume":"3","author":"Wada","year":"2013","journal-title":"Metabolites"},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"4861","DOI":"10.1128\/AEM.02508-08","article-title":"Organization, Evolution, and Expression Analysis of the Biosynthetic Gene Cluster for Scytonemin, a Cyanobacterial UV-Absorbing Pigment","volume":"75","author":"Sorrels","year":"2009","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_102","doi-asserted-by":"crossref","unstructured":"Mills, L.A., McCormick, A.J., and Lea-Smith, D.J. (2020). Current Knowledge and Recent Advances in Understanding Metabolism of the Model Cyanobacterium Synechocystis sp. PCC 6803. Biosci. Rep., 40.","DOI":"10.1042\/BSR20193325"},{"key":"ref_103","unstructured":"Falkowski, P.G., and Raven, J.A. (2013). Aquatic Photosynthesis, Princeton University Press. [2nd ed.]."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"86","DOI":"10.1016\/j.abb.2010.07.007","article-title":"Roles of Xanthophyll Carotenoids in Protection against Photoinhibition and Oxidative Stress in the Cyanobacterium Synechococcus Sp. Strain PCC 7002","volume":"504","author":"Zhu","year":"2010","journal-title":"Arch. Biochem. Biophys."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"2108","DOI":"10.1038\/ismej.2015.14","article-title":"Chlorophyll f -Driven Photosynthesis in a Cavernous Cyanobacterium","volume":"9","author":"Behrendt","year":"2015","journal-title":"ISME J."},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"1677","DOI":"10.1128\/AEM.02000-07","article-title":"Coping with Our Cold Planet","volume":"74","author":"Rodrigues","year":"2008","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_107","doi-asserted-by":"crossref","unstructured":"Watzer, B., and Forchhammer, K. (2018). Cyanophycin Synthesis Optimizes Nitrogen Utilization in the Unicellular Cyanobacterium Synechocystis Sp. Strain PCC 6803. Appl. Environ. Microbiol., 84.","DOI":"10.1128\/AEM.01298-18"},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"2025","DOI":"10.1128\/AEM.03579-14","article-title":"Anoxygenic Photosynthesis Controls Oxygenic Photosynthesis in a Cyanobacterium from a Sulfidic Spring","volume":"81","author":"Klatt","year":"2015","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"125","DOI":"10.1017\/S0954102089000192","article-title":"Microbial Biomass, Photosynthesis and Chlorophyll a Related Pigments in the Ponds of the McMurdo Ice Shelf, Antarctica","volume":"1","author":"Pridmore","year":"1989","journal-title":"Antarct. Sci."},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1111\/j.1365-2427.1995.tb00882.x","article-title":"Phytoplankton Nutrient Deficiency in Lakes of the McMurdo Dry Valleys, Antarctica","volume":"34","author":"Priscu","year":"1995","journal-title":"Freshw. Biol."},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"118","DOI":"10.1046\/j.1529-8817.1998.340118.x","article-title":"Temperature Dependence of UV Radiation Effects on Antarctic Cyanobacteria","volume":"34","author":"Roos","year":"1998","journal-title":"J. Phycol."},{"key":"ref_112","doi-asserted-by":"crossref","unstructured":"Keys, J.R., and Williams, K. (1981). Origin of Crystalline, Cold Desert Salts in the McMurdo Region, Antarctica. Geochim. Cosmochim. Acta, 2299\u20132309.","DOI":"10.1016\/0016-7037(81)90084-3"},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1017\/S0954102094000039","article-title":"The Chemical Composition of Glacial Melt Water Ponds and Streams on the McMurdo Ice Shelf, Antarctica","volume":"6","author":"Whitehead","year":"1994","journal-title":"Antarct. Sci."},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1016\/j.yqres.2005.04.002","article-title":"Late Quaternary Climate-Driven Environmental Change in the Larsemann Hills, East Antarctica, Multi-Proxy Evidence from a Lake Sediment Core","volume":"64","author":"Hodgson","year":"2005","journal-title":"Quat. Res."},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"369","DOI":"10.1111\/gbi.12280","article-title":"The \u2018Dirty Ice\u2019 of the McMurdo Ice Shelf: Analogues for Biological Oases during the Cryogenian","volume":"16","author":"Hawes","year":"2018","journal-title":"Geobiology"}],"container-title":["Genes"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2073-4425\/12\/3\/426\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T05:36:37Z","timestamp":1760160997000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2073-4425\/12\/3\/426"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,3,16]]},"references-count":115,"journal-issue":{"issue":"3","published-online":{"date-parts":[[2021,3]]}},"alternative-id":["genes12030426"],"URL":"https:\/\/doi.org\/10.3390\/genes12030426","relation":{},"ISSN":["2073-4425"],"issn-type":[{"value":"2073-4425","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,3,16]]}}}