{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,23]],"date-time":"2026-10-23T20:59:47Z","timestamp":1792789187848,"version":"4.5.0"},"reference-count":61,"publisher":"MDPI AG","issue":"19","license":[{"start":{"date-parts":[[2026,9,24]],"date-time":"2026-09-24T00:00:00Z","timestamp":1790208000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Science Committee of the Ministry of Science and Higher Education of the Republic of Kazakhstan","award":["AP26194471"],"award-info":[{"award-number":["AP26194471"]}],"id":[{"id":"https:\/\/ror.org\/05tyne317","id-type":"ROR","asserted-by":"crossref"}]},{"award":["AP26194471"],"award-info":[{"award-number":["AP26194471"]}],"id":[{"id":"https:\/\/ror.org\/05tyne317","id-type":"ROR","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Biology"],"abstract":"<jats:p>Geothermal-associated microbial communities remain poorly characterized in Central Asia. Here, shotgun metagenomics was used to compare microbial communities associated with thermally influenced substrates from the Chundzha and Rakhman geothermal systems in Kazakhstan. Six metagenomes representing three spatial replicates per site yielded approximately 6.5\u20137.3 Gb of clean sequence data per sample. Species-level richness showed a directional trend toward higher values in Chundzha, whereas Shannon diversity was similar between sites. Genus-level Bray\u2013Curtis ordination showed apparent site-associated separation, with sampling site explaining 87.3% of the variation, although the effect was not statistically significant (PERMANOVA, R2 = 0.873, p = 0.10). Chundzha showed greater relative representation of several halophilic and halotolerant taxa, whereas Rakhman showed greater representation of soil-associated lineages. Broad COG and KEGG functional profiles were largely shared between sites. Several hydrogenase-, nitrogenase-, and RuBisCO-associated genes showed higher relative representation in Rakhman, whereas selected fermentation- and photosynthesis-associated functions were relatively higher in Chundzha; however, none of the targeted KO comparisons remained significant after FDR correction. These results indicate candidate taxonomic and functional patterns that should be regarded as exploratory because of the limited replication. The dataset provides a baseline for future genome-resolved and metatranscriptomic studies of geothermal-associated microbial communities in Kazakhstan.<\/jats:p>","DOI":"10.3390\/biology15191704","type":"journal-article","created":{"date-parts":[[2026,9,24]],"date-time":"2026-09-24T13:26:11Z","timestamp":1790256371000},"page":"1704","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Comparative Shotgun Metagenomics of Microbial Communities Associated with Two Geothermal Systems in Kazakhstan"],"prefix":"10.3390","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-5612-1002","authenticated-orcid":false,"given":"Ardak B.","family":"Kakimova","sequence":"first","affiliation":[{"name":"Department of Chemical and Biological Technologies, K. Turysov Institute of Geology and Oil and Gas Engineering, Satbayev University, 22 Satpaev Street, Almaty 050013, Kazakhstan"},{"name":"Department of Biotechnology, Faculty of Biology and Biotechnology, Al-Farabi Kazakh National University, Al-Farabi 71, Almaty 050038, Kazakhstan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yelena G.","family":"Zaparina","sequence":"additional","affiliation":[{"name":"Department of Botany and Agroecology, Faculty of Biology and Biotechnology, Al-Farabi Kazakh National University, Al-Farabi 71, Almaty 050038, Kazakhstan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Bolatkhan K.","family":"Zayadan","sequence":"additional","affiliation":[{"name":"Department of Biotechnology, Faculty of Biology and Biotechnology, Al-Farabi Kazakh National University, Al-Farabi 71, Almaty 050038, Kazakhstan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4340-8749","authenticated-orcid":false,"given":"Sandugash K.","family":"Sandybayeva","sequence":"additional","affiliation":[{"name":"Department of Chemical and Biological Technologies, K. Turysov Institute of Geology and Oil and Gas Engineering, Satbayev University, 22 Satpaev Street, Almaty 050013, Kazakhstan"},{"name":"Department of Biotechnology, Faculty of Biology and Biotechnology, Al-Farabi Kazakh National University, Al-Farabi 71, Almaty 050038, Kazakhstan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0000-0764-8756","authenticated-orcid":false,"given":"Dinara","family":"Kelden","sequence":"additional","affiliation":[{"name":"Department of Chemical and Biological Technologies, K. Turysov Institute of Geology and Oil and Gas Engineering, Satbayev University, 22 Satpaev Street, Almaty 050013, Kazakhstan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Aliyam A.","family":"Dauletova","sequence":"additional","affiliation":[{"name":"Department of Biotechnology, Faculty of Biology and Biotechnology, Al-Farabi Kazakh National University, Al-Farabi 71, Almaty 050038, Kazakhstan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Silvano","family":"Piazza","sequence":"additional","affiliation":[{"name":"Computational Biology, International Centre for Genetic Engineering and Biotechnology, ICGEB, 34149 Trieste, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,9,24]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"1505","DOI":"10.1089\/ast.2018.2016","article-title":"The intersection of geology, geochemistry, and microbiology in continental hydrothermal systems","volume":"19","author":"Colman","year":"2019","journal-title":"Astrobiology"},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Inskeep, W.P., Rusch, D.B., Jay, Z.J., Herrgard, M.J., Kozubal, M.A., Richardson, T.H., Macur, R.E., Hamamura, N., Jennings, R.M., and Fouke, B.W. (2010). Metagenomes from high-temperature chemotrophic systems reveal geochemical controls on microbial community structure and function. PLoS ONE, 5.","DOI":"10.1371\/journal.pone.0009773"},{"key":"ref_3","first-page":"100160","article-title":"Taxonomic and functional metagenomics profiling of Tuwa and Unnai hot springs microbial communities","volume":"26","author":"Vora","year":"2023","journal-title":"Ecol. Genet. Genom."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"2876","DOI":"10.1038\/s41467-018-05020-y","article-title":"Microbial biogeography of 925 geothermal springs in New Zealand","volume":"9","author":"Power","year":"2018","journal-title":"Nat. Commun."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"1166","DOI":"10.1038\/ismej.2013.237","article-title":"Humboldt\u2019s spa: Microbial diversity is controlled by temperature in geothermal environments","volume":"8","author":"Sharp","year":"2014","journal-title":"ISME J."},{"key":"ref_6","doi-asserted-by":"crossref","unstructured":"Wang, S., Hou, W., Dong, H., Jiang, H., Huang, L., Wu, G., Zhang, C., Song, Z., Zhang, Y., and Ren, H. (2013). Control of temperature on microbial community structure in hot springs of the Tibetan Plateau. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0062901"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"718","DOI":"10.1038\/ismej.2012.157","article-title":"Sediment microbial communities in Great Boiling Spring are controlled by temperature and distinct from water communities","volume":"7","author":"Cole","year":"2013","journal-title":"ISME J."},{"key":"ref_8","doi-asserted-by":"crossref","unstructured":"Bowen De Le\u00f3n, K., Gerlach, R., Peyton, B.M., and Fields, M.W. (2013). Archaeal and bacterial communities in three alkaline hot springs in Heart Lake Geyser Basin, Yellowstone National Park. Front. Microbiol., 4.","DOI":"10.3389\/fmicb.2013.00330"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"5917","DOI":"10.1038\/s41598-020-62797-z","article-title":"Microbiome and ecology of a hot spring-microbialite system on the Trans-Himalayan Plateau","volume":"10","author":"Roy","year":"2020","journal-title":"Sci. Rep."},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"DeCastro, M.-E., Escuder-Rodr\u00edguez, J.-J., Becerra, M., Rodr\u00edguez-Belmonte, E., and Gonz\u00e1lez-Siso, M.-I. (2021). Comparative metagenomic analysis of two hot springs from Ourense (Northwestern Spain) and others worldwide. Front. Microbiol., 12.","DOI":"10.3389\/fmicb.2021.769065"},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Klatt, C.G., Inskeep, W.P., Herrgard, M.J., Jay, Z.J., Rusch, D.B., Tringe, S.G., Parenteau, M.N., Ward, D.M., Boomer, S.M., and Bryant, D.A. (2013). Community structure and function of high-temperature chlorophototrophic microbial mats inhabiting diverse geothermal environments. Front. Microbiol., 4.","DOI":"10.3389\/fmicb.2013.00106"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1038\/s41597-025-04493-z","article-title":"Recovery of nearly 3,000 archaeal genomes from 152 terrestrial geothermal spring metagenomes","volume":"12","author":"Qi","year":"2025","journal-title":"Sci. Data"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"782","DOI":"10.1038\/s41597-026-07139-w","article-title":"Genome-resolved metagenomic survey of 500 samples from 56 hot springs across the Western US","volume":"13","author":"Korchagina","year":"2026","journal-title":"Sci. Data"},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Chen, F., Cheng, M., Rong, D., Wang, Y., Liang, R., Irfan, M., Kang, Y., and Cao, Y. (2025). Metagenomic insights into the microbial communities and functional traits of hot springs in Guizhou Province, China. Front. Microbiol., 16.","DOI":"10.3389\/fmicb.2025.1615879"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"e70004","DOI":"10.1002\/mbo3.70004","article-title":"The metagenomic landscape of a high-altitude geothermal spring in Tajikistan reveals a novel Desulfurococcaceae member, Zestomicrobium tamdykulense gen. nov., sp. nov","volume":"13","author":"Dzhuraeva","year":"2024","journal-title":"MicrobiologyOpen"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"2204","DOI":"10.1111\/j.1462-2920.2011.02475.x","article-title":"Biological nitrogen fixation in acidic high-temperature geothermal springs in Yellowstone National Park, Wyoming","volume":"13","author":"Hamilton","year":"2011","journal-title":"Environ. Microbiol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"126644","DOI":"10.1016\/j.syapm.2025.126644","article-title":"Nitrogen fixation in Pampinifervens, a new species-rich genus of Aquificaceae that inhabits a wide pH range in terrestrial hot springs","volume":"48","author":"Palmer","year":"2025","journal-title":"Syst. Appl. Microbiol."},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"Luo, S., Chen, X., Guo, S., Hu, S., Dong, Z., and Geng, J. (2026). Temperature-driven metabolic adaptation in thermophilic microbial communities of Western Sichuan hot springs. BMC Microbiol., 26.","DOI":"10.1186\/s12866-026-04921-z"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"fiae062","DOI":"10.1093\/femsec\/fiae062","article-title":"Quantifying genome-specific carbon fixation in a 750-meter deep subsurface hydrothermal microbial community","volume":"100","author":"Coskun","year":"2024","journal-title":"FEMS Microbiol. Ecol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"56","DOI":"10.1186\/s40793-026-00875-x","article-title":"Exploring the biotechnological potential of terrestrial hot spring microbiomes for CO2 utilisation","volume":"21","author":"Stead","year":"2026","journal-title":"Environ. Microbiome"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"2926","DOI":"10.1007\/s00284-021-02545-2","article-title":"Metagenomics and culture-based diversity analysis of the bacterial community in the Zharkent geothermal spring in Kazakhstan","volume":"78","author":"Mashzhan","year":"2021","journal-title":"Curr. Microbiol."},{"key":"ref_22","first-page":"112","article-title":"Isolation and study of morphological and cultural properties of cyanobacterial community from hot springs in Almaty region","volume":"75","author":"Sandybayeva","year":"2023","journal-title":"Eurasian J. Ecol."},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Kozhagulova, A., Yapiyev, V., Karabayanova, L., Dillinger, A., Zavaley, V., Kalitova, A., Bayramov, E., Holbrook, J., Grasby, S.E., and Fustic, M. (2023). Geological controls on the geothermal system and hydrogeochemistry of the deep low-salinity Upper Cretaceous aquifers in the Zharkent (eastern Ily) Basin, south-eastern Kazakhstan. Front. Earth Sci., 11.","DOI":"10.3389\/feart.2023.1212064"},{"key":"ref_24","first-page":"1560","article-title":"Prospects for the development of ecotourism in the territory of the Katon-Karagai State National Nature Park of the East Kazakhstan Region","volume":"45","author":"Turyspekova","year":"2022","journal-title":"Geoj. Tour. Geosites"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"i884","DOI":"10.1093\/bioinformatics\/bty560","article-title":"Fastp: An ultra-fast all-in-one FASTQ preprocessor","volume":"34","author":"Chen","year":"2018","journal-title":"Bioinformatics"},{"key":"ref_26","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_27","doi-asserted-by":"crossref","unstructured":"Hyatt, D., Chen, G.-L., 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_28","doi-asserted-by":"crossref","first-page":"3150","DOI":"10.1093\/bioinformatics\/bts565","article-title":"CD-HIT: Accelerated for clustering the next-generation sequencing data","volume":"28","author":"Fu","year":"2012","journal-title":"Bioinformatics"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"366","DOI":"10.1038\/s41592-021-01101-x","article-title":"Sensitive protein alignments at tree-of-life scale using DIAMOND","volume":"18","author":"Buchfink","year":"2021","journal-title":"Nat. Methods"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"R60","DOI":"10.1186\/gb-2011-12-6-r60","article-title":"Metagenomic biomarker discovery and explanation","volume":"12","author":"Segata","year":"2011","journal-title":"Genome Biol."},{"key":"ref_31","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_32","doi-asserted-by":"crossref","first-page":"D274","DOI":"10.1093\/nar\/gkaa1018","article-title":"COG database update: Focus on microbial diversity, model organisms, and widespread pathogens","volume":"49","author":"Galperin","year":"2021","journal-title":"Nucleic Acids Res."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"117","DOI":"10.1111\/j.1442-9993.1993.tb00438.x","article-title":"Non-parametric multivariate analyses of changes in community structure","volume":"18","author":"Clarke","year":"1993","journal-title":"Aust. J. Ecol."},{"key":"ref_34","first-page":"32","article-title":"A new method for non-parametric multivariate analysis of variance","volume":"26","author":"Anderson","year":"2001","journal-title":"Austral Ecol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"245","DOI":"10.1111\/j.1541-0420.2005.00440.x","article-title":"Distance-based tests for homogeneity of multivariate dispersions","volume":"62","author":"Anderson","year":"2006","journal-title":"Biometrics"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1111\/j.2517-6161.1995.tb02031.x","article-title":"Controlling the false discovery rate: A practical and powerful approach to multiple testing","volume":"57","author":"Benjamini","year":"1995","journal-title":"J. R. Stat. Soc. Ser. B Stat. Methodol."},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Massello, F.L., Chan, C.S., Chan, K.-G., Goh, K.M., Donati, E., and Urbieta, M.S. (2020). Meta-analysis of microbial communities in hot springs: Recurrent taxa and complex shaping factors beyond pH and temperature. Microorganisms, 8.","DOI":"10.3390\/microorganisms8060906"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"7506","DOI":"10.1038\/s41467-024-51841-5","article-title":"Covariation of hot spring geochemistry with microbial genomic diversity, function, and evolution","volume":"15","author":"Colman","year":"2024","journal-title":"Nat. Commun."},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Podar, P.T., Yang, Z., Bj\u00f6rnsd\u00f3ttir, S.H., and Podar, M. (2020). Comparative analysis of microbial diversity across temperature gradients in hot springs from Yellowstone and Iceland. Front. Microbiol., 11.","DOI":"10.3389\/fmicb.2020.01625"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"137574","DOI":"10.1016\/j.scitotenv.2020.137574","article-title":"Temperature governs the distribution of hot spring microbial community in three hydrothermal fields, Eastern Tibetan Plateau Geothermal Belt, Western China","volume":"720","author":"Guo","year":"2020","journal-title":"Sci. Total Environ."},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Rowe, L., Dowd, S.E., Davidson, K., Kovarik, C., VanAken, M., Jarabek, A., and Taylor, C. (2024). Comparing microbial populations from diverse hydrothermal features in Yellowstone National Park: Hot springs and mud volcanoes. Front. Microbiol., 15.","DOI":"10.3389\/fmicb.2024.1409664"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"681","DOI":"10.1038\/s41467-019-08499-1","article-title":"Mixing of meteoric and geothermal fluids supports hyperdiverse chemosynthetic hydrothermal communities","volume":"10","author":"Colman","year":"2019","journal-title":"Nat. Commun."},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Sharma, N., Kumar, J., Abedin, M.M., Sahoo, D., Pandey, A., Rai, A.K., and Singh, S.P. (2020). Metagenomics revealing molecular profiling of community structure and metabolic pathways in natural hot springs of the Sikkim Himalaya. BMC Microbiol., 20.","DOI":"10.1186\/s12866-020-01923-3"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"fiaa246","DOI":"10.1093\/femsec\/fiaa246","article-title":"An emerging view of the diversity, ecology and function of Archaea in alkaline hydrothermal environments","volume":"97","author":"Mueller","year":"2021","journal-title":"FEMS Microbiol. Ecol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"4066","DOI":"10.1038\/s41467-024-48498-5","article-title":"Analysis of nearly 3000 archaeal genomes from terrestrial geothermal springs sheds light on interconnected biogeochemical processes","volume":"15","author":"Qi","year":"2024","journal-title":"Nat. Commun."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"135","DOI":"10.1007\/s00792-016-0889-8","article-title":"Microbial community analysis of pH 4 thermal springs in Yellowstone National Park","volume":"21","author":"Jiang","year":"2017","journal-title":"Extremophiles"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"22243","DOI":"10.1038\/s41598-022-26299-4","article-title":"A metagenomic insight into the microbiomes of geothermal springs in the Subantarctic Kerguelen Islands","volume":"12","author":"Allioux","year":"2022","journal-title":"Sci. Rep."},{"key":"ref_48","doi-asserted-by":"crossref","unstructured":"Saxena, R., Dhakan, D.B., Mittal, P., Waiker, P., Chowdhury, A., Ghatak, A., and Sharma, V.K. (2017). Metagenomic analysis of hot springs in Central India reveals hydrocarbon degrading thermophiles and pathways essential for survival in extreme environments. Front. Microbiol., 7.","DOI":"10.3389\/fmicb.2016.02123"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"117144","DOI":"10.1016\/j.envres.2023.117144","article-title":"Metagenome-assembled genomes reveal carbohydrate degradation and element metabolism of microorganisms inhabiting Tengchong hot springs, China","volume":"238","author":"Zhang","year":"2023","journal-title":"Environ. Res."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"1099","DOI":"10.1007\/s12088-024-01248-z","article-title":"NGS-Based Metagenomics Depicting Taxonomic and Functional Insights into North-Western Himalayan Hot Springs","volume":"64","author":"Rangra","year":"2024","journal-title":"Indian J. Microbiol."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1186\/s40793-024-00572-7","article-title":"Integrating multi-platform assembly to recover MAGs from hot spring biofilms: Insights into microbial diversity, biofilm formation, and carbohydrate degradation","volume":"19","author":"Liew","year":"2024","journal-title":"Environ. Microbiome"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"936","DOI":"10.1038\/s41559-018-0519-1","article-title":"Function and functional redundancy in microbial systems","volume":"2","author":"Louca","year":"2018","journal-title":"Nat. Ecol. Evol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"e00991-21","DOI":"10.1128\/msystems.00991-21","article-title":"Microbial functional diversity correlates with species diversity along a temperature gradient","volume":"7","author":"Ruhl","year":"2022","journal-title":"mSystems"},{"key":"ref_54","doi-asserted-by":"crossref","unstructured":"Hunt, K.A., Jennings, R.M., Inskeep, W.P., and Carlson, R.P. (2018). Multiscale analysis of autotroph-heterotroph interactions in a high-temperature microbial community. PLoS Comput. Biol., 14.","DOI":"10.1371\/journal.pcbi.1006431"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"108391","DOI":"10.1016\/j.fbio.2026.108391","article-title":"Metagenomics-driven discovery of next-generation fermentation biocatalysts: From enzyme mining to synthetic biology applications","volume":"77","author":"Wani","year":"2026","journal-title":"Food Biosci."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"1689","DOI":"10.1021\/acs.biochem.5b01328","article-title":"[NiFe]-Hydrogenase maturation","volume":"55","author":"Lacasse","year":"2016","journal-title":"Biochemistry"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"875","DOI":"10.1021\/acs.accounts.0c00022","article-title":"Structural insight into [NiFe] hydrogenase maturation by transient complexes between Hyp proteins","volume":"53","author":"Miki","year":"2020","journal-title":"Acc. Chem. Res."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"761","DOI":"10.1038\/ismej.2015.153","article-title":"Genomic and metagenomic surveys of hydrogenase distribution indicate H2 is a widely utilised energy source for microbial growth and survival","volume":"10","author":"Greening","year":"2016","journal-title":"ISME J."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1264\/jsme2.ME17134","article-title":"Nitrogen fixation in thermophilic chemosynthetic microbial communities depending on hydrogen, sulfate, and carbon dioxide","volume":"33","author":"Nishihara","year":"2018","journal-title":"Microbes Environ."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"261","DOI":"10.1146\/annurev-marine-120709-142712","article-title":"Beyond the Calvin cycle: Autotrophic carbon fixation in the ocean","volume":"3","author":"Sievert","year":"2011","journal-title":"Annu. Rev. Mar. Sci."},{"key":"ref_61","doi-asserted-by":"crossref","unstructured":"Xie, Y.-G., Cao, X.-R., Qi, Y.-L., Chen, L., Mao, Y.-H., Li, Y., Wang, C.-J., Li, Z.-W., Qu, Y.-N., and Li, W.-J. (2026). Genome-resolved discovery of Candidatus Vitaminotrophota reveals carbon fixation and multi-vitamin biosynthetic potential in hot springs. NPJ Biofilms Microbiomes.","DOI":"10.1038\/s41522-026-01093-6"}],"container-title":["Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2079-7737\/15\/19\/1704\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,24]],"date-time":"2026-09-24T13:38:32Z","timestamp":1790257112000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2079-7737\/15\/19\/1704"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,24]]},"references-count":61,"journal-issue":{"issue":"19","published-online":{"date-parts":[[2026,10]]}},"alternative-id":["biology15191704"],"URL":"https:\/\/doi.org\/10.3390\/biology15191704","relation":{},"ISSN":["2079-7737"],"issn-type":[{"value":"2079-7737","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,24]]}}}