{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,9]],"date-time":"2026-10-09T23:25:27Z","timestamp":1791588327514,"version":"4.3.4"},"reference-count":57,"publisher":"MDPI AG","issue":"9","license":[{"start":{"date-parts":[[2025,4,28]],"date-time":"2025-04-28T00:00:00Z","timestamp":1745798400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["31860011"],"award-info":[{"award-number":["31860011"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["51864037"],"award-info":[{"award-number":["51864037"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["2020GG0018"],"award-info":[{"award-number":["2020GG0018"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["2022YFHH0026"],"award-info":[{"award-number":["2022YFHH0026"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["2022ZY0213"],"award-info":[{"award-number":["2022ZY0213"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["2021MS05066"],"award-info":[{"award-number":["2021MS05066"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["2023QNJS154"],"award-info":[{"award-number":["2023QNJS154"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Key Technology Research Program Project of Inner Mongolia Autonomous Region","award":["31860011"],"award-info":[{"award-number":["31860011"]}]},{"name":"Key Technology Research Program Project of Inner Mongolia Autonomous Region","award":["51864037"],"award-info":[{"award-number":["51864037"]}]},{"name":"Key Technology Research Program Project of Inner Mongolia Autonomous Region","award":["2020GG0018"],"award-info":[{"award-number":["2020GG0018"]}]},{"name":"Key Technology Research Program Project of Inner Mongolia Autonomous Region","award":["2022YFHH0026"],"award-info":[{"award-number":["2022YFHH0026"]}]},{"name":"Key Technology Research Program Project of Inner Mongolia Autonomous Region","award":["2022ZY0213"],"award-info":[{"award-number":["2022ZY0213"]}]},{"name":"Key Technology Research Program Project of Inner Mongolia Autonomous Region","award":["2021MS05066"],"award-info":[{"award-number":["2021MS05066"]}]},{"name":"Key Technology Research Program Project of Inner Mongolia Autonomous Region","award":["2023QNJS154"],"award-info":[{"award-number":["2023QNJS154"]}]},{"name":"Key R&amp;D and Achievement Transformation Plan Project of Inner Mongolia Autonomous Region","award":["31860011"],"award-info":[{"award-number":["31860011"]}]},{"name":"Key R&amp;D and Achievement Transformation Plan Project of Inner Mongolia Autonomous Region","award":["51864037"],"award-info":[{"award-number":["51864037"]}]},{"name":"Key R&amp;D and Achievement Transformation Plan Project of Inner Mongolia Autonomous Region","award":["2020GG0018"],"award-info":[{"award-number":["2020GG0018"]}]},{"name":"Key R&amp;D and Achievement Transformation Plan Project of Inner Mongolia Autonomous Region","award":["2022YFHH0026"],"award-info":[{"award-number":["2022YFHH0026"]}]},{"name":"Key R&amp;D and Achievement Transformation Plan Project of Inner Mongolia Autonomous Region","award":["2022ZY0213"],"award-info":[{"award-number":["2022ZY0213"]}]},{"name":"Key R&amp;D and Achievement Transformation Plan Project of Inner Mongolia Autonomous Region","award":["2021MS05066"],"award-info":[{"award-number":["2021MS05066"]}]},{"name":"Key R&amp;D and Achievement Transformation Plan Project of Inner Mongolia Autonomous Region","award":["2023QNJS154"],"award-info":[{"award-number":["2023QNJS154"]}]},{"name":"foundation of central governmental guidance for local scientific and technological development","award":["31860011"],"award-info":[{"award-number":["31860011"]}]},{"name":"foundation of central governmental guidance for local scientific and technological development","award":["51864037"],"award-info":[{"award-number":["51864037"]}]},{"name":"foundation of central governmental guidance for local scientific and technological development","award":["2020GG0018"],"award-info":[{"award-number":["2020GG0018"]}]},{"name":"foundation of central governmental guidance for local scientific and technological development","award":["2022YFHH0026"],"award-info":[{"award-number":["2022YFHH0026"]}]},{"name":"foundation of central governmental guidance for local scientific and technological development","award":["2022ZY0213"],"award-info":[{"award-number":["2022ZY0213"]}]},{"name":"foundation of central governmental guidance for local scientific and technological development","award":["2021MS05066"],"award-info":[{"award-number":["2021MS05066"]}]},{"name":"foundation of central governmental guidance for local scientific and technological development","award":["2023QNJS154"],"award-info":[{"award-number":["2023QNJS154"]}]},{"name":"National Natural Science Foundation of Inner Mongolia","award":["31860011"],"award-info":[{"award-number":["31860011"]}]},{"name":"National Natural Science Foundation of Inner Mongolia","award":["51864037"],"award-info":[{"award-number":["51864037"]}]},{"name":"National Natural Science Foundation of Inner Mongolia","award":["2020GG0018"],"award-info":[{"award-number":["2020GG0018"]}]},{"name":"National Natural Science Foundation of Inner Mongolia","award":["2022YFHH0026"],"award-info":[{"award-number":["2022YFHH0026"]}]},{"name":"National Natural Science Foundation of Inner Mongolia","award":["2022ZY0213"],"award-info":[{"award-number":["2022ZY0213"]}]},{"name":"National Natural Science Foundation of Inner Mongolia","award":["2021MS05066"],"award-info":[{"award-number":["2021MS05066"]}]},{"name":"National Natural Science Foundation of Inner Mongolia","award":["2023QNJS154"],"award-info":[{"award-number":["2023QNJS154"]}]},{"name":"fundamental research funds for the directly affiliated universities in Inner Mongolia","award":["31860011"],"award-info":[{"award-number":["31860011"]}]},{"name":"fundamental research funds for the directly affiliated universities in Inner Mongolia","award":["51864037"],"award-info":[{"award-number":["51864037"]}]},{"name":"fundamental research funds for the directly affiliated universities in Inner Mongolia","award":["2020GG0018"],"award-info":[{"award-number":["2020GG0018"]}]},{"name":"fundamental research funds for the directly affiliated universities in Inner Mongolia","award":["2022YFHH0026"],"award-info":[{"award-number":["2022YFHH0026"]}]},{"name":"fundamental research funds for the directly affiliated universities in Inner Mongolia","award":["2022ZY0213"],"award-info":[{"award-number":["2022ZY0213"]}]},{"name":"fundamental research funds for the directly affiliated universities in Inner Mongolia","award":["2021MS05066"],"award-info":[{"award-number":["2021MS05066"]}]},{"name":"fundamental research funds for the directly affiliated universities in Inner Mongolia","award":["2023QNJS154"],"award-info":[{"award-number":["2023QNJS154"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>Direct interspecies electron transfer (DIET) is a syntrophic metabolism wherein free electrons are directly transferred between microorganisms without the mediation of intermediates such as molecular hydrogen or formate. Previous research has demonstrated that Methanothrix harundinacea 6Ac is capable of reducing carbon dioxide through DIET. However, the mechanisms underlying electron uptake in M. harundinacea 6Ac during DIET remain poorly understood. This study aims to elucidate the electron and proton flux in M. harundinacea 6Ac during DIET and to propose a model for electron uptake in this organism, primarily based on the analysis of gene transcript levels, genomic characteristics of M. harundinacea 6Ac, and the pathways generating fully reduced ferridoxin (Fdred2\u2212), reduced coenzyme F420 (F420H2), coenzyme M (CoM-SH), and coenzyme B (CoB-SH) during DIET. The findings suggest that membrane-bound heterodisulfide reductase (HdrED), F420H2-dehydrogenase lacking subunit F (Fpo\u2212), and cytoplasmic heterodisulfide reductase (HdrABC)-subunit B of F420-reducing hydrogenase (FrhB) complex play critical roles in electron uptake in M. harundinacea 6Ac during DIET. Specifically, Fpo\u2212 is responsible for generating Fdred2\u2212 with reduced methanophenazine (MPH2), driven by a proton motive force, while HdrED facilitates the reduction of heterodisulfide of coenzyme M and coenzyme B (CoM-S-S-CoB) to CoM-SH and CoB-SH using MPH2. Additionally, cytoplasmic heterodisulfide reductase HdrABC and subunit B of coenzyme F420-hydrogenase complex (HdrABC-FrhB complex) catalyzes the reduction of oxidized coenzyme F420 (F420) to F420H2, utilizing CoM-SH, CoB-SH, and Fdred2\u2212. This study represents the first genetics-based functional characterization of electron and proton flux in M. harundinacea 6Ac during DIET, providing a model for further investigation of electron uptake in Methanosaeta species. Furthermore, it deepens our understanding of the mechanisms underlying electron uptake in methanogens during DIET.<\/jats:p>","DOI":"10.3390\/ijms26094195","type":"journal-article","created":{"date-parts":[[2025,4,29]],"date-time":"2025-04-29T07:49:36Z","timestamp":1745912976000},"page":"4195","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":5,"title":["Analysis of Mechanisms for Electron Uptake by Methanothrix\u00a0harundinacea 6Ac During Direct Interspecies Electron Transfer"],"prefix":"10.3390","volume":"26","author":[{"given":"Lei","family":"Wang","sequence":"first","affiliation":[{"name":"School of Life Science and Technology, Inner Mongolia University of Science and Technology, Baotou 014010, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaoman","family":"Shan","sequence":"additional","affiliation":[{"name":"School of Life Science and Technology, Inner Mongolia University of Science and Technology, Baotou 014010, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yanhui","family":"Xu","sequence":"additional","affiliation":[{"name":"School of Life Science and Technology, Inner Mongolia University of Science and Technology, Baotou 014010, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Quan","family":"Xi","sequence":"additional","affiliation":[{"name":"School of Mining and Coal, Inner Mongolia University of Science and Technology, Baotou 014010, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Haiming","family":"Jiang","sequence":"additional","affiliation":[{"name":"School of Life Science and Technology, Inner Mongolia University of Science and Technology, Baotou 014010, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xia","family":"Li","sequence":"additional","affiliation":[{"name":"School of Mining and Coal, Inner Mongolia University of Science and Technology, Baotou 014010, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2025,4,28]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"20","DOI":"10.1007\/BF00406313","article-title":"Methanobacillus omelianskii, a symbiotic association of two species of bacteria","volume":"59","author":"Bryant","year":"1967","journal-title":"Arch. Mikrobiol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"4247","DOI":"10.1128\/AEM.68.9.4247-4252.2002","article-title":"Biochemical evidence for formate transfer in syntrophic propionate-oxidizing cocultures of Syntrophobacter fumaroxidans and Methanospirillum hungatei","volume":"68","author":"Luijten","year":"2002","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"568","DOI":"10.1038\/nrmicro2166","article-title":"Electron transfer in syntrophic communities of anaerobic bacteria and archaea","volume":"7","author":"Stams","year":"2009","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"Rosenberg, E., DeLong, E.F., Lory, S., Stackebrandt, E., and Thompson, F. (2013). Syntrophism among prokaryotes. The Prokaryotes, Springer.","DOI":"10.1007\/978-3-642-30144-5"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"58","DOI":"10.1196\/annals.1419.005","article-title":"Physiology, ecology, phylogeny, and genomics of microorganisms capable of syntrophic metabolism","volume":"1125","author":"Mcinerney","year":"2008","journal-title":"Ann. N. Y. Acad. Sci."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"441","DOI":"10.1038\/nrmicro1892","article-title":"The ecology and biotechnology of sulphate reducing bacteria","volume":"6","author":"Muyzer","year":"2008","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"1949","DOI":"10.1128\/aem.60.6.1949-1955.1994","article-title":"Autotrophic, hydrogen-oxidizing, denitrifying bacteria in groundwater, potential agents for bioremediation of nitrate contamination","volume":"60","author":"Smith","year":"1998","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1420","DOI":"10.1128\/AEM.02508-06","article-title":"Growth yields in bacterial denitrification and nitrate ammonification","volume":"73","author":"Strohm","year":"2007","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"196","DOI":"10.1021\/es001360p","article-title":"In situ stimulation of groundwater denitrification with formate to remediate nitrate contamination","volume":"35","author":"Smith","year":"2001","journal-title":"Environ. Sci. Technol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"8683","DOI":"10.1128\/AEM.71.12.8683-8691.2005","article-title":"Investigation of an acetate-fed denitrifying microbial community by stable isotope probing, full-cycle rRNA analysis, and fluorescent in situ hybridization-microautoradiography","volume":"71","author":"Ginige","year":"2005","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"346","DOI":"10.1128\/aem.41.2.346-354.1981","article-title":"Anaerobic degradation of lactate by syntrophic associations of Methanosarcina barkeri and Desulfovibrio species and effect of H2 on acetate degradation","volume":"41","author":"McInerney","year":"1981","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Doloman, A., Boeren, S., Miller, C.D., and Sousa, D.Z. (2022). Stimulating effect of trichococcus flocculiformis on a coculture of Syntrophomonas wolfei and Methanospirillum hungatei. Appl. Environ. Microbiol., 88.","DOI":"10.1128\/aem.00391-22"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"307","DOI":"10.1016\/j.jbiosc.2016.02.002","article-title":"Denitrification by Pseudomonas stutzeri coupled with CO2 reduction by Sporomusa ovata with hydrogen as an electron donor assisted by solid-phase humin","volume":"122","author":"Xiao","year":"2016","journal-title":"J. Biosci. Bioeng."},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Ozuolmez, D., Na, H., Lever, M.A., Kjeldsen, K.U., J\u00f8rgensen, B.B., and Plugge, C.M. (2015). Methanogenic archaea and sulfate reducing bacteria co-cultured on acetate: Teamwork or coexistence?. Front. Microbiol., 6.","DOI":"10.3389\/fmicb.2015.00492"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1111\/j.1574-6968.1986.tb01939.x","article-title":"Interspecies hydrogen transfer in co-cultures of methanol-utilizing acidogens and sulfate-reducing or methanogenic bacteria","volume":"38","author":"Heijthuijsen","year":"1986","journal-title":"FEMS Microbiol. Lett."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1413","DOI":"10.1126\/science.1196526","article-title":"Direct exchange of electrons within aggregates of an evolved syntrophic coculture of anaerobic bacteria","volume":"330","author":"Summers","year":"2010","journal-title":"Science"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"10042","DOI":"10.1073\/pnas.1117592109","article-title":"Microbial interspecies electron transfer via electric currents through conductive minerals","volume":"109","author":"Kato","year":"2012","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"4599","DOI":"10.1128\/AEM.00895-14","article-title":"Direct interspecies electron transfer between Geobacter metallireducens and Methanosarcina barkeri","volume":"80","author":"Rotaru","year":"2014","journal-title":"Appl. Environ. Microbiol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"408","DOI":"10.1039\/C3EE42189A","article-title":"A new model for electron flow during anaerobic digestion: Direct interspecies electron transfer to Methanosaeta for the reduction of carbon dioxide to methane","volume":"7","author":"Rotaru","year":"2014","journal-title":"Energy Environ. Sci."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"587","DOI":"10.1038\/nature15733","article-title":"Intercellular wiring enables electron transfer between methanotrophic archaea and bacteria","volume":"526","author":"Wegener","year":"2015","journal-title":"Nature"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"13924","DOI":"10.1038\/ncomms13924","article-title":"Syntrophic anaerobic photosynthesis via direct interspecies electron transfer","volume":"8","author":"Ha","year":"2017","journal-title":"Nat. Commun."},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Yee, M.O., and Rotaru, A.E. (2020). Extracellular electron uptake in Methanosarcinales is independent of multiheme c-type cytochromes. Sci. Rep., 10.","DOI":"10.1038\/s41598-019-57206-z"},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Zhou, J.J., Smith, J.A., Li, M., and Holmes, D.E. (2023). Methane production by Methanothrix thermoacetophila via direct interspecies electron transfer with Geobacter metallireducens. mBio, 14.","DOI":"10.1128\/mbio.00360-23"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"1774","DOI":"10.3390\/microbiolres14040122","article-title":"Geobacter grbiciae\u2014A new electron donor in the formation of co-cultures via direct interspecies electron transfer","volume":"14","author":"Deng","year":"2023","journal-title":"Microbiol. Res."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"e202409192","DOI":"10.1002\/anie.202409192","article-title":"Solar-Driven Methanogenesis through Microbial Ecosystem Engineering on Carbon Nitride","volume":"63","author":"Kalathil","year":"2024","journal-title":"Angew. Chem. Int. Ed."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"4896","DOI":"10.1039\/c1ee02229f","article-title":"Live wires: Direct extracellular electron exchange for bioenergy and the bioremediation of energy-related contamination","volume":"4","author":"Lovley","year":"2011","journal-title":"Energy Environ. Sci."},{"key":"ref_27","doi-asserted-by":"crossref","unstructured":"Rotaru, A.E., Woodard, T.L., Nevin, K.P., and Lovley, D.R. (2015). Link between capacity for current production and syntrophic growth in Geobacter species. Front. Microbiol., 6.","DOI":"10.3389\/fmicb.2015.00744"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"370","DOI":"10.1038\/s41396-021-01078-7","article-title":"Light-driven carbon dioxide reduction to methane by Methanosarcina barkeri in an electric syntrophic coculture","volume":"16","author":"Huang","year":"2022","journal-title":"ISME J."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"837","DOI":"10.1038\/s41396-019-0575-9","article-title":"Syntrophus conductive pili demonstrate that common hydrogen-donating syntrophs can have a direct electron transfer option","volume":"14","author":"Walker","year":"2020","journal-title":"ISME J."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"e202402318","DOI":"10.1002\/anie.202402318","article-title":"Unnatural Direct Interspecies Electron Transfer Enabled by Living Cell-Cell Click Chemistry","volume":"63","author":"Zhao","year":"2024","journal-title":"Angew. Chem. Int. Ed."},{"key":"ref_31","doi-asserted-by":"crossref","unstructured":"Holmes, D.E., Zhou, J.J., Ueki, T., Woodard, T., and Lovley, D.R. (2021). Mechanisms for Electron Uptake by Methanosarcina acetivorans during Direct Interspecies Electron Transfer. mBio, 12.","DOI":"10.1128\/mBio.02344-21"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"eabh1852","DOI":"10.1126\/sciadv.abh1852","article-title":"Syntrophic interspecies electron transfer drives carbon fixation and growth by Rhodopseudomonas palustris under dark, anoxic conditions","volume":"7","author":"Liu","year":"2021","journal-title":"Sci. Adv."},{"key":"ref_33","doi-asserted-by":"crossref","unstructured":"Yee, M.O., Snoeyenbos-West, O.L., Thamdrup, B., Ottosen, L.D.M., and Rotaru, A.E. (2019). Extracellular Electron Uptake by Two Methanosarcina Species. Front. Microbiol., 7.","DOI":"10.3389\/fenrg.2019.00029"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"531","DOI":"10.1038\/nature15512","article-title":"Single cell activity reveals direct electron transfer in methanotrophic consortia","volume":"526","author":"McGlynn","year":"2015","journal-title":"Nature"},{"key":"ref_35","doi-asserted-by":"crossref","unstructured":"Holmes, D.E., Rotaru, A.E., Ueki, T., Shrestha, P.M., Ferry, J.G., and Lovley, D.R. (2018). Electron and Proton Flux for Carbon Dioxide Reduction in Methanosarcina barkeri During Direct Interspecies Electron Transfer. Front. Microbiol., 9.","DOI":"10.3389\/fmicb.2018.03109"},{"key":"ref_36","unstructured":"Wang, L. (2020). Research on Stoichiometrical Production of Methane by Microbes via Direct Interspecies Electron Transfer. [Master\u2019s Thesis, Inner Mongolia University of Science and Technology]."},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Wang, L., Nevin, K.P., Woodard, T.L., Mu, B.Z., and Lovley, D.R. (2016). Expanding the Diet for DIET: Electron Donors Supporting Direct Interspecies Electron Transfer (DIET) in Defined Co-Cultures. Front. Microbiol., 7.","DOI":"10.3389\/fmicb.2016.00236"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"1277","DOI":"10.1111\/j.1742-4658.2011.08048.x","article-title":"Re-evaluation of the function of the F420 dehydrogenase in electron transport of Methanosarcina mazei","volume":"278","author":"Welte","year":"2011","journal-title":"FEBS J."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"579","DOI":"10.1038\/nrmicro1931","article-title":"Methanogenic archaea: Ecologically relevant differences in energy conservation","volume":"6","author":"Thauer","year":"2008","journal-title":"Nat. Rev. Microbiol."},{"key":"ref_40","doi-asserted-by":"crossref","unstructured":"Arnulf, K., Thomas, H., Jennifer, F., Laura, V.N., and Reinhard, R. (2015). Cytochromes c in Archaea-distribution, maturation, cell architecture, and the special case of Ignicoccus hospitalis. Front. Microbiol., 6.","DOI":"10.3389\/fmicb.2015.00439"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"3921","DOI":"10.1128\/jb.174.12.3921-3927.1992","article-title":"Characterization of Cytochromes from Methanosarcina Strain G\u00f6l and Their Involvement in Electron Transport during Growth on Methanol","volume":"174","author":"Kamlage","year":"1992","journal-title":"J. Bacteriol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"333","DOI":"10.1002\/cbic.200390053","article-title":"Redox Potentials of Methanophenazine and CoB-S-S-CoM, Factors Involved in Electron Transport in Methanogenic Archaea","volume":"4","author":"Tietze","year":"2003","journal-title":"Chembiochem"},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Zhu, J.X., Zheng, H.J., Ai, G.M., Zhang, G.S., Liu, D., Liu, X.L., and Dong, X.Z. (2012). The Genome Characteristics and Predicted Function of Methyl-Group Oxidation Pathway in the Obligate Aceticlastic Methanogens, Methanosaeta spp.. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0036756"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"2868","DOI":"10.1128\/JB.00162-11","article-title":"Membrane-bound electron transport in Methanosaeta thermophila","volume":"193","author":"Welte","year":"2011","journal-title":"J. Bacteriol."},{"key":"ref_45","first-page":"1130","article-title":"Bioenergetics and anaerobic respiratory chains of aceticlastic methanogens","volume":"1837","author":"Welte","year":"2014","journal-title":"BBA"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"674","DOI":"10.1128\/JB.01307-09","article-title":"Function of Ech Hydrogenase in Ferredoxin-Dependent, Membrane-Bound Electron Transport in Methanosarcina mazei","volume":"192","author":"Welte","year":"2010","journal-title":"J. Bacteriol."},{"key":"ref_47","doi-asserted-by":"crossref","unstructured":"Yan, Z., Wang, M., and Ferry, J.G. (2017). A Ferredoxin- and F420H2-dependent, electron-bifurcating, heterodisulfide reductase with homologs in the domains Bacteria and Archaea. mBio, 8.","DOI":"10.1128\/mBio.02285-16"},{"key":"ref_48","doi-asserted-by":"crossref","unstructured":"Yan, Z., and Ferry, J.G. (2018). Electron Bifurcation and Confurcation in Methanogenesis and Reverse Methanogenesis. Front. Microbiol., 9.","DOI":"10.3389\/fmicb.2018.01322"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"507","DOI":"10.1146\/annurev.biochem.030508.152103","article-title":"Hydrogenases from Methanogenic Archaea, Nickel, a Novel Cofactor, and H2 Storage","volume":"79","author":"Thauer","year":"2010","journal-title":"Annu. Rev. Biochem."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"146931","DOI":"10.1016\/j.cej.2023.146931","article-title":"Harnessing Carbonaceous materials\u2019 multifaceted roles for enhanced anaerobic digestion performance","volume":"477","author":"Zhang","year":"2023","journal-title":"Chem. Eng. J."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"1513","DOI":"10.1007\/s00018-002-8526-3","article-title":"Redox-driven proton translocation in methanogenic Archaea","volume":"59","author":"Deppenmeier","year":"2002","journal-title":"Cell. Mol. Life Sci."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"843","DOI":"10.1111\/j.1365-2958.2009.06990.x","article-title":"Methanogenesis by Methanosarcina acetivorans involves two structurally and functionally distinct classes of heterodisulfide reductase","volume":"75","author":"Buan","year":"2010","journal-title":"Mol. Microbiol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1099\/ijs.0.63887-0","article-title":"Methanosaeta harundinacea sp. nov., a novel acetate-scavenging methanogen isolated from a UASB reactor","volume":"56","author":"Ma","year":"2006","journal-title":"Int. J. Syst. Evol. Microbiol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"15473","DOI":"10.1073\/pnas.1208779109","article-title":"Essential anaplerotic role for the energy-converting hydrogenase Eha in hydrogenotrophic methanogenesis","volume":"109","author":"Lie","year":"2012","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_55","doi-asserted-by":"crossref","unstructured":"Arshad, A., Speth, D.R., Graaf RMd Camp, H.J.O., Jetten, M.S.M., and Welte, C.U. (2015). A metagenomics-based metabolic model of nitrate-dependent anaerobic oxidation of methane by Methanoperedens-like archaea. Front. Microbiol., 6.","DOI":"10.3389\/fmicb.2015.01423"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"1929","DOI":"10.1038\/s41396-018-0109-x","article-title":"A methanotrophic archaeon couples anaerobic oxidation of methane to Fe(III) reduction","volume":"12","author":"Cai","year":"2018","journal-title":"ISME J."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"1030","DOI":"10.1038\/s41396-020-0590-x","article-title":"Anaerobic methane oxidation coupled to manganese reduction by members of the Methanoperedenaceae","volume":"14","author":"Leu","year":"2020","journal-title":"ISME J."}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/26\/9\/4195\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,9]],"date-time":"2025-10-09T17:23:32Z","timestamp":1760030612000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/26\/9\/4195"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,4,28]]},"references-count":57,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2025,5]]}},"alternative-id":["ijms26094195"],"URL":"https:\/\/doi.org\/10.3390\/ijms26094195","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,4,28]]}}}