{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,5]],"date-time":"2026-10-05T15:55:43Z","timestamp":1791215743331,"version":"4.1.0"},"reference-count":137,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,7,28]],"date-time":"2026-07-28T00:00:00Z","timestamp":1785196800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Microbiol."],"abstract":"<jats:p>\n                    Biogas, a mixture of biomethane (bio-CH\n                    <jats:sub>4<\/jats:sub>\n                    ) and carbon dioxide (CO\n                    <jats:sub>2<\/jats:sub>\n                    ), is the product of methanogenesis, the final stage of anaerobic digestion (AD). Biohydrogen (bio-H\n                    <jats:sub>2<\/jats:sub>\n                    ), on the other hand, is produced during the acidogenesis stage through dark fermentation (DF) processes and the transformation of intermediates from other fermentation types, such as the conversion of lactate and acetate into butyrate. Bio-H\n                    <jats:sub>2<\/jats:sub>\n                    and CO\n                    <jats:sub>2<\/jats:sub>\n                    together form fermentation gas, whose volume and bio-H\n                    <jats:sub>2<\/jats:sub>\n                    content depend on the dominant type of acid fermentation. Understanding the complexity of metabolic pathways and the interactions between microorganisms at each stage of AD permits process optimization in two-stage systems, facilitating the production of both bio-H\n                    <jats:sub>2<\/jats:sub>\n                    and bio-CH\n                    <jats:sub>4<\/jats:sub>\n                    . Theoretical calculations indicate that separate recovery of bio-H\n                    <jats:sub>2<\/jats:sub>\n                    and bio-CH\n                    <jats:sub>4<\/jats:sub>\n                    through a two-stage AD process yields a higher energy output from the degraded substrate. However, DF is characterized by limited bio-H\n                    <jats:sub>2<\/jats:sub>\n                    yields due to inherent metabolic constraints and competition among microbial pathways. In contrast, acetogenic pathways consistently generate substrates for methanogenesis. Paradoxically, analyses based on simple substrates such as glucose facilitate a deeper understanding of metabolic pathways and chemical reactions in bacterial cells. Such knowledge is essential for the rational design and critical evaluation of integrated multiproduct AD systems, supporting mindful resource management and sustainable development. This review focuses on (i) the biological limitations of bio-H\u2082 generation during acidogenesis; (ii) physiological constraints that keep bio-H\u2082 technologies at the research and development stage, compared with the commercially established biogas technologies; and (iii) prospects for bio-H\n                    <jats:sub>2<\/jats:sub>\n                    recovery within integrated multiproduct AD systems.\n                  <\/jats:p>","DOI":"10.3389\/fmicb.2026.1881665","type":"journal-article","created":{"date-parts":[[2026,7,28]],"date-time":"2026-07-28T05:37:24Z","timestamp":1785217044000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Can bio-H2 compete with bio-CH4\u2013unequal chances of anaerobic digestion metabolic pathways"],"prefix":"10.3389","volume":"17","author":[{"given":"Anna","family":"Sikora","sequence":"first","affiliation":[{"name":"Laboratory of White Biotechnology, Institute of Biochemistry and Biophysics PAS","place":["Warsaw, Poland"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anna","family":"Detman-Ignatowska","sequence":"additional","affiliation":[{"name":"Laboratory of White Biotechnology, Institute of Biochemistry and Biophysics PAS","place":["Warsaw, Poland"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,7,28]]},"reference":[{"key":"ref1","doi-asserted-by":"publisher","first-page":"1149","DOI":"10.1016\/j.ijhydene.2024.10.218","article-title":"Review of over two decades of research on dark and photo fermentation for biohydrogen production \u2013 a combination of traditional, systematic, and bibliometric approaches","volume":"91","author":"Agyekum","year":"2024","journal-title":"Int. J. Hydrog. Energy"},{"key":"ref2","doi-asserted-by":"publisher","first-page":"335","DOI":"10.1016\/j.ijhydene.2023.05.161","article-title":"Biohydrogen production through dark fermentation: recent trends and advances in transition to a circular bioeconomy","volume":"52","author":"Ahmad","year":"2024","journal-title":"Int. J. Hydrog. Energy"},{"key":"ref3","doi-asserted-by":"publisher","first-page":"500","DOI":"10.3390\/methane3030029","article-title":"Biohydrogen produced via dark fermentation: a review","volume":"3","author":"Albuquerque","year":"2024","journal-title":"Methane"},{"key":"ref4","doi-asserted-by":"publisher","first-page":"2113","DOI":"10.3390\/molecules25092113","article-title":"A simple biorefinery concept to produce 2G-lactic acid from sugar beet pulp (SBP): a high-value target approach to valorize a waste stream","volume":"25","author":"Alves De Oliveira","year":"2020","journal-title":"Molecules"},{"key":"ref5","doi-asserted-by":"publisher","first-page":"123981","DOI":"10.1016\/j.biortech.2020.123981","article-title":"Evaluation of dissolved and immobilized redox mediators on dark fermentation: driving to hydrogen or solventogenic pathway","volume":"317","author":"Atilano-Camino","year":"2020","journal-title":"Bioresour. Technol."},{"key":"ref6","doi-asserted-by":"publisher","first-page":"2573","DOI":"10.1016\/j.ijhydene.2009.01.010","article-title":"Effect of extrinsic lactic acid on fermentative hydrogen production","volume":"34","author":"Baghchehsaraee","year":"2009","journal-title":"Int. J. Hydrog. Energy"},{"key":"ref7","doi-asserted-by":"publisher","first-page":"1255","DOI":"10.3390\/en10091255","article-title":"Integrated bioethanol fermentation\/anaerobic digestion for valorization of sugar beet pulp","volume":"10","author":"Berlowska","year":"2017","journal-title":"Energies"},{"key":"ref8","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1155\/2016\/3154929","article-title":"Simultaneous Saccharification and fermentation of sugar beet pulp for efficient bioethanol production","volume":"2016","author":"Ber\u0142owska","year":"2016","journal-title":"Biomed. Res. Int."},{"key":"ref9","doi-asserted-by":"publisher","first-page":"118101","DOI":"10.1016\/j.envres.2024.118101","article-title":"Hydrogen and methane production through two stage anaerobic digestion of straw residues","volume":"247","author":"Bertasini","year":"2024","journal-title":"Environ. Res."},{"key":"ref10","doi-asserted-by":"publisher","first-page":"2913","DOI":"10.1111\/1462-2920.13082","article-title":"A novel route for ethanol oxidation in the acetogenic bacterium Acetobacterium woodii: the acetaldehyde\/ethanol dehydrogenase pathway","volume":"18","author":"Bertsch","year":"2016","journal-title":"Environ. Microbiol."},{"key":"ref11","doi-asserted-by":"publisher","first-page":"810","DOI":"10.1016\/j.fuel.2016.08.030","article-title":"Biohydrogen and biogas \u2013 an overview on feedstocks and enhancement process","volume":"185","author":"Bharathiraja","year":"2016","journal-title":"Fuel"},{"key":"ref12","doi-asserted-by":"publisher","first-page":"125269","DOI":"10.1016\/j.fuel.2022.125269","article-title":"Microbial electrolysis cell (MEC): reactor configurations, recent advances and strategies in biohydrogen production","volume":"328","author":"Bora","year":"2022","journal-title":"Fuel"},{"key":"ref13","doi-asserted-by":"publisher","first-page":"117440","DOI":"10.1016\/j.watres.2021.117440","article-title":"Biochar boosts dark fermentative H2 production from sugarcane bagasse by selective enrichment\/colonization of functional bacteria and enhancing extracellular electron transfer","volume":"202","author":"Bu","year":"2021","journal-title":"Water Res."},{"key":"ref14","doi-asserted-by":"publisher","first-page":"6713","DOI":"10.1016\/j.ijhydene.2016.03.057","article-title":"Inhibition of dark fermentative bio-hydrogen production: a review","volume":"41","author":"Bundhoo","year":"2016","journal-title":"Int. J. Hydrog. Energy"},{"key":"ref15","doi-asserted-by":"publisher","first-page":"107","DOI":"10.1016\/j.ijhydene.2023.06.253","article-title":"Regulation of the dark fermentation products by electro-fermentation in reactors without membrane","volume":"49","author":"Carde\u00f1a","year":"2024","journal-title":"Int. J. Hydrog. Energy"},{"key":"ref16","doi-asserted-by":"publisher","first-page":"109602","DOI":"10.1016\/j.rser.2019.109602","article-title":"Stability problems in the hydrogen production by dark fermentation: possible causes and solutions","volume":"119","author":"Castell\u00f3","year":"2020","journal-title":"Renew. Sust. Energ. Rev."},{"key":"ref17","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/s40793-023-00545-2","article-title":"A unified compendium of prokaryotic and viral genomes from over 300 anaerobic digestion microbiomes","volume":"19","author":"Centurion","year":"2024","journal-title":"Environmental Microbiome"},{"key":"ref18","doi-asserted-by":"publisher","first-page":"e70074","DOI":"10.1002\/biot.70074","article-title":"Biological routes for biohydrogen production: a clean and carbon-free fuel","volume":"20","author":"Cha","year":"2025","journal-title":"Biotechnol. J."},{"key":"ref19","doi-asserted-by":"publisher","first-page":"125394","DOI":"10.1016\/j.cej.2020.125394","article-title":"Improving hydrogen and methane co-generation in cascading dark fermentation and anaerobic digestion: the effect of magnetite nanoparticles on microbial electron transfer and syntrophism","volume":"397","author":"Cheng","year":"2020","journal-title":"Chem. Eng. J."},{"key":"ref20","doi-asserted-by":"publisher","first-page":"10057","DOI":"10.1016\/j.biortech.2011.08.063","article-title":"Comparative analysis of hydrogen-producing bacterial biofilms and granular sludge formed in continuous cultures of fermentative bacteria","volume":"102","author":"Chojnacka","year":"2011","journal-title":"Bioresour. Technol."},{"key":"ref21","doi-asserted-by":"publisher","first-page":"e0128008","DOI":"10.1371\/journal.pone.0128008","article-title":"Noteworthy facts about a methane-producing microbial community processing acidic effluent from sugar beet molasses fermentation","volume":"10","author":"Chojnacka","year":"2015","journal-title":"PLoS One"},{"key":"ref22","doi-asserted-by":"publisher","first-page":"123713","DOI":"10.1016\/j.biortech.2020.123713","article-title":"Dark fermentative hydrogen production from hydrolyzed sugar beet pulp improved by iron addition","volume":"314","author":"Cieciura-W\u0142och","year":"2020","journal-title":"Bioresour. Technol."},{"key":"ref23","doi-asserted-by":"publisher","first-page":"123029","DOI":"10.1016\/j.renene.2025.123029","article-title":"Enhancing biomass energy recovery in Brazil\u2019s agro-industrial sector: innovating biogas, biomethane, and hydrogen production through peach syrup waste management","volume":"247","author":"Corr\u00eaa","year":"2025","journal-title":"Renew. Energy"},{"key":"ref24","doi-asserted-by":"publisher","first-page":"131014","DOI":"10.1016\/j.energy.2024.131014","article-title":"Experimental study of CO2\/H2\/NH3 influence on CH4 flameless combustion process in semi-industrial furnace","volume":"296","author":"Czyzewski","year":"2024","journal-title":"Energy"},{"key":"ref25","doi-asserted-by":"publisher","first-page":"128842","DOI":"10.1016\/j.fuel.2023.128842","article-title":"Biohydrogen production through dark fermentation from waste biomass: current status and future perspectives on biorefinery development","volume":"350","author":"D\u2019 Silva","year":"2023","journal-title":"Fuel"},{"key":"ref26","doi-asserted-by":"publisher","first-page":"6046","DOI":"10.1016\/j.ijhydene.2008.07.098","article-title":"Advances in biological hydrogen production processes","volume":"33","author":"Das","year":"2008","journal-title":"Int. J. Hydrog. Energy"},{"key":"ref27","doi-asserted-by":"publisher","first-page":"1800","DOI":"10.1128\/AEM.67.4.1800-1804.2001","article-title":"Pathway of propionate oxidation by a syntrophic culture of Smithella propionica and Methanospirillum hungatei","volume":"67","author":"De Bok","year":"2001","journal-title":"Appl. Environ. Microbiol."},{"key":"ref28","doi-asserted-by":"publisher","first-page":"125","DOI":"10.1186\/s13068-021-01968-0","article-title":"Evaluation of acidogenesis products\u2019 effect on biogas production performed with metagenomics and isotopic approaches","volume":"14","author":"Detman","year":"","journal-title":"Biotechnol. Biofuels"},{"key":"ref29","doi-asserted-by":"publisher","first-page":"1023","DOI":"10.15244\/pjoes\/68149","article-title":"Biohydrogen and biomethane (biogas) production in the consecutive stages of anaerobic digestion of molasses","volume":"26","author":"Detman","year":"2017","journal-title":"Pol. J. Environ. Stud."},{"key":"ref30","doi-asserted-by":"publisher","first-page":"10967","DOI":"10.1016\/j.ijhydene.2018.05.004","article-title":"Inhibition of hydrogen-yielding dark fermentation by ascomycetous yeasts","volume":"43","author":"Detman","year":"","journal-title":"Int. J. Hydrog. Energy"},{"key":"ref31","doi-asserted-by":"publisher","first-page":"158","DOI":"10.1186\/s40168-021-01105-x","article-title":"Dynamics of dark fermentation microbial communities in the light of lactate and butyrate production","volume":"9","author":"Detman","year":"","journal-title":"Microbiome"},{"key":"ref32","doi-asserted-by":"publisher","first-page":"612344","DOI":"10.3389\/fmicb.2020.612344","article-title":"Dynamics and complexity of dark fermentation microbial communities producing hydrogen from sugar beet molasses in continuously operating packed bed reactors","volume":"11","author":"Detman","year":"","journal-title":"Front. Microbiol."},{"key":"ref33","doi-asserted-by":"publisher","first-page":"36","DOI":"10.1186\/s12934-019-1085-1","article-title":"Cell factories converting lactate and acetate to butyrate: Clostridium butyricum and microbial communities from dark fermentation bioreactors","volume":"18","author":"Detman","year":"2019","journal-title":"Microb. Cell Factories"},{"key":"ref34","doi-asserted-by":"publisher","first-page":"116","DOI":"10.1186\/s13068-018-1106-z","article-title":"Methane-yielding microbial communities processing lactate-rich substrates: a piece of the anaerobic digestion puzzle","volume":"11","author":"Detman","year":"","journal-title":"Biotechnol. Biofuels"},{"key":"ref35","doi-asserted-by":"publisher","DOI":"10.2139\/ssrn.6301403","article-title":"Single-stage vs. two-stage anaerobic digestion of molasses in a circular economy context: energy recovery and digestate fertilizer value","author":"Detman-Ignatowska","year":"2026","journal-title":"SSRN Elec. J."},{"key":"ref36","doi-asserted-by":"publisher","first-page":"152812","DOI":"10.1016\/j.ijhydene.2025.152812","article-title":"Impact of plant-derived biochars on biohydrogen production from sugar beet molasses in a continuous system: insights into the roles of microbial communities and chemical elements","volume":"199","author":"Detman-Ignatowska","year":"","journal-title":"Int. J. Hydrog. Energy"},{"key":"ref37","doi-asserted-by":"publisher","first-page":"105955","DOI":"10.1016\/j.ibiod.2024.105955","article-title":"Feasibility of successive hydrogen and methane production: effects of temperature and organic loads on energy potential and microbial dynamics","volume":"196","author":"Devens","year":"2025","journal-title":"Int. Biodeterior. Biodegrad."},{"key":"ref38","doi-asserted-by":"publisher","first-page":"571","DOI":"10.1007\/s12155-020-10093-z","article-title":"Effect of lactate fermentation type on the biochemical methane potential of tequila vinasse","volume":"13","author":"Diaz-Cruces","year":"2020","journal-title":"Bioenergy Res."},{"key":"ref39","doi-asserted-by":"publisher","first-page":"36","DOI":"10.1007\/BF02568732","article-title":"The role of an NAD-independent lactate dehydrogenase and acetate in the utilization of lactate Byclostridium acetobutylicum strain P262","volume":"164","author":"Diez-Gonzalez","year":"1995","journal-title":"Arch. Microbiol."},{"key":"ref40","doi-asserted-by":"publisher","first-page":"5810","DOI":"10.1128\/AEM.70.10.5810-5817.2004","article-title":"Lactate-utilizing Bacteria, isolated from human feces, that produce butyrate as a major fermentation product","volume":"70","author":"Duncan","year":"2004","journal-title":"Appl. Environ. Microbiol."},{"key":"ref41","doi-asserted-by":"publisher","first-page":"855","DOI":"10.1002\/bbb.2658","article-title":"Improvement of substrate turnover through integrating dark fermentation into existing biogas plants","volume":"18","author":"Eggers","year":"2024","journal-title":"Biofuels Bioprod. Biorefin."},{"key":"ref42","doi-asserted-by":"publisher","first-page":"656","DOI":"10.1016\/j.rser.2017.05.075","article-title":"A critical review on inhibition of dark biohydrogen fermentation","volume":"79","author":"Elbeshbishy","year":"2017","journal-title":"Renew. Sust. Energ. Rev."},{"key":"ref43","doi-asserted-by":"publisher","first-page":"3371","DOI":"10.1007\/s00253-016-7325-y","article-title":"Microbial communities from 20 different hydrogen-producing reactors studied by 454 pyrosequencing","volume":"100","author":"Etchebehere","year":"2016","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref44","doi-asserted-by":"publisher","first-page":"132007","DOI":"10.1016\/j.biortech.2024.132007","article-title":"Functional microorganisms in hydrogen production: mechanisms and applications","volume":"419","author":"Fan","year":"2025","journal-title":"Bioresour. Technol."},{"key":"ref45","doi-asserted-by":"publisher","first-page":"532","DOI":"10.1016\/j.ijhydene.2023.10.092","article-title":"Fermentative biohydrogen production in sugarcane biorefineries: advances, challenges and prospects","volume":"49","author":"Fuess","year":"2024","journal-title":"Int. J. Hydrog. Energy"},{"key":"ref46","doi-asserted-by":"publisher","first-page":"426","DOI":"10.1016\/j.biortech.2017.09.121","article-title":"Temporal dynamics and metabolic correlation between lactate-producing and hydrogen-producing bacteria in sugarcane vinasse dark fermentation: the key role of lactate","volume":"247","author":"Fuess","year":"2018","journal-title":"Bioresour. Technol."},{"key":"ref47","doi-asserted-by":"publisher","first-page":"121379","DOI":"10.1016\/j.biortech.2019.121379","article-title":"Novel insights on the versatility of biohydrogen production from sugarcane vinasse via thermophilic dark fermentation: impacts of pH-driven operating strategies on acidogenesis metabolite profiles","volume":"286","author":"Fuess","year":"2019","journal-title":"Bioresour. Technol."},{"key":"ref48","doi-asserted-by":"publisher","first-page":"124595","DOI":"10.1016\/j.biortech.2020.124595","article-title":"A review on the factors influencing biohydrogen production from lactate: the key to unlocking enhanced dark fermentative processes","volume":"324","author":"Garc\u00eda-Depraect","year":"","journal-title":"Bioresour. Technol."},{"key":"ref49","doi-asserted-by":"publisher","first-page":"5820","DOI":"10.1016\/j.ijhydene.2016.11.204","article-title":"A novel biohydrogen production process: co-digestion of vinasse and Nejayote as complex raw substrates using a robust inoculum","volume":"42","author":"Garc\u00eda-Depraect","year":"2017","journal-title":"Int. J. Hydrog. Energy"},{"key":"ref50","doi-asserted-by":"publisher","first-page":"151","DOI":"10.1016\/j.fuel.2018.06.126","article-title":"Fermentative biohydrogen production from tequila vinasse via the lactate-acetate pathway: operational performance, kinetic analysis and microbial ecology","volume":"234","author":"Garc\u00eda-Depraect","year":"2018","journal-title":"Fuel"},{"key":"ref51","doi-asserted-by":"publisher","first-page":"127358","DOI":"10.1016\/j.biortech.2022.127358","article-title":"Two-stage anaerobic digestion of food waste: enhanced bioenergy production rate by steering lactate-type fermentation during hydrolysis-acidogenesis","volume":"358","author":"Garc\u00eda-Depraect","year":"2022","journal-title":"Bioresour. Technol."},{"key":"ref52","doi-asserted-by":"publisher","first-page":"11284","DOI":"10.1016\/j.ijhydene.2020.08.209","article-title":"Microbial ecology of a lactate-driven dark fermentation process producing hydrogen under carbohydrate-limiting conditions","volume":"46","author":"Garc\u00eda-Depraect","year":"","journal-title":"Int. J. Hydrog. Energy"},{"key":"ref53","doi-asserted-by":"publisher","first-page":"123160","DOI":"10.1016\/j.biortech.2020.123160","article-title":"Three-stage process for tequila vinasse valorization through sequential lactate, biohydrogen and methane production","volume":"307","author":"Garc\u00eda-Depraect","year":"2020","journal-title":"Bioresour. Technol."},{"key":"ref54","doi-asserted-by":"publisher","first-page":"618","DOI":"10.1016\/j.biortech.2018.11.056","article-title":"Effect of process parameters on enhanced biohydrogen production from tequila vinasse via the lactate-acetate pathway","volume":"273","author":"Garc\u00eda-Depraect","year":"","journal-title":"Bioresour. Technol."},{"key":"ref55","doi-asserted-by":"publisher","first-page":"159","DOI":"10.1016\/j.fuel.2019.04.147","article-title":"Enhanced biohydrogen production from the dark co-fermentation of tequila vinasse and nixtamalization wastewater: novel insights into ecological regulation by pH","volume":"253","author":"Garc\u00eda-Depraect","year":"","journal-title":"Fuel"},{"key":"ref56","doi-asserted-by":"publisher","first-page":"236","DOI":"10.1016\/j.biortech.2019.02.100","article-title":"Lactate- and acetate-based biohydrogen production through dark co-fermentation of tequila vinasse and nixtamalization wastewater: metabolic and microbial community dynamics","volume":"282","author":"Garc\u00eda-Depraect","year":"","journal-title":"Bioresour. Technol."},{"key":"ref57","doi-asserted-by":"publisher","first-page":"6350","DOI":"10.3390\/en17246350","article-title":"Insights into biohydrogen production through dark fermentation of food waste: substrate properties, inocula, and pretreatment strategies","volume":"17","author":"Gbiete","year":"2024","journal-title":"Energies"},{"key":"ref58","doi-asserted-by":"publisher","first-page":"151390","DOI":"10.1016\/j.cej.2024.151390","article-title":"Decipher syntrophies within C2-C4 organic acids-degrading anaerobic microbiomes: a multi-omic exploration","volume":"489","author":"Ghiotto","year":"2024","journal-title":"Chem. Eng. J."},{"key":"ref59","doi-asserted-by":"publisher","first-page":"2182","DOI":"10.1007\/s11356-024-35873-4","article-title":"A comparative evaluation of dark fermentative bioreactor configurations for enhanced hydrogen production","volume":"32","author":"Goren","year":"2025","journal-title":"Environ. Sci. Pollut. Res."},{"key":"ref60","doi-asserted-by":"publisher","first-page":"10789","DOI":"10.3390\/app142310789","article-title":"Microbes and parameters influencing dark fermentation for hydrogen production","volume":"14","author":"Gupta","year":"2024","journal-title":"Appl. Sci."},{"key":"ref61","doi-asserted-by":"publisher","first-page":"21","DOI":"10.2166\/wst.2005.0494","article-title":"Fundamentals of the fermentative production of hydrogen","volume":"52","author":"Hallenbeck","year":"2005","journal-title":"Water Sci. Technol."},{"key":"ref62","doi-asserted-by":"publisher","first-page":"7379","DOI":"10.1016\/j.ijhydene.2008.12.080","article-title":"Fermentative hydrogen production: principles, progress, and prognosis","volume":"34","author":"Hallenbeck","year":"2009","journal-title":"Int. J. Hydrog. Energy"},{"key":"ref63","doi-asserted-by":"publisher","first-page":"259","DOI":"10.1016\/j.biortech.2016.02.131","article-title":"Selective fractionation of sugar beet pulp for release of fermentation and chemical feedstocks; optimisation of thermo-chemical pre-treatment","volume":"209","author":"Hamley-Bennett","year":"2016","journal-title":"Bioresour. Technol."},{"key":"ref64","doi-asserted-by":"publisher","first-page":"693","DOI":"10.1007\/s00253-007-0854-7","article-title":"Quantitative analysis of a high-rate hydrogen-producing microbial community in anaerobic agitated granular sludge bed bioreactors using glucose as substrate","volume":"75","author":"Hung","year":"2007","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref65","doi-asserted-by":"publisher","first-page":"2323","DOI":"10.3390\/agriculture15222323","article-title":"Biohydrogen production from agricultural and livestock by-products by dark fermentation: a data mining approach","volume":"15","author":"Illuminati","year":"2025","journal-title":"Agriculture"},{"key":"ref66","doi-asserted-by":"publisher","first-page":"12699","DOI":"10.1007\/s13399-022-03282-7","article-title":"Bio-hydrogen production through dark fermentation: an overview","volume":"14","author":"Jain","year":"2024","journal-title":"Biomass Conv. Bioref."},{"key":"ref67","doi-asserted-by":"publisher","first-page":"6666","DOI":"10.1016\/j.biortech.2007.11.067","article-title":"Biological hydrogen production by immobilized cells of Clostridium tyrobutyricum JM1 isolated from a food waste treatment process","volume":"99","author":"Jo","year":"2008","journal-title":"Bioresour. Technol."},{"key":"ref68","first-page":"359","volume-title":"Advances in Biotechnology for Food Industry","author":"Joanna","year":"2018"},{"key":"ref69","doi-asserted-by":"publisher","first-page":"136","DOI":"10.1016\/j.biortech.2011.09.093","article-title":"Effects of initial lactic acid concentration, HRTs, and OLRs on bio-hydrogen production from lactate-type fermentation","volume":"103","author":"Kim","year":"2012","journal-title":"Bioresour. Technol."},{"key":"ref70","doi-asserted-by":"publisher","first-page":"14","DOI":"10.1007\/s11783-018-1037-8","article-title":"Biogas and its opportunities\u2014a review","volume":"12","author":"Kougias","year":"2018","journal-title":"Front. Environ. Sci. Eng."},{"key":"ref71","doi-asserted-by":"publisher","first-page":"685","DOI":"10.1007\/s10529-006-9299-9","article-title":"Improving the yield from fermentative hydrogen production","volume":"29","author":"Kraemer","year":"2007","journal-title":"Biotechnol. Lett."},{"key":"ref72","doi-asserted-by":"publisher","first-page":"8393","DOI":"10.1016\/j.biortech.2011.03.041","article-title":"Dark fermentation on biohydrogen production: pure culture","volume":"102","author":"Lee","year":"2011","journal-title":"Bioresour. Technol."},{"key":"ref73","doi-asserted-by":"publisher","first-page":"843","DOI":"10.1128\/JB.01417-07","article-title":"Coupled ferredoxin and Crotonyl coenzyme a (CoA) reduction with NADH catalyzed by the Butyryl-CoA dehydrogenase\/Etf complex from Clostridium kluyveri","volume":"190","author":"Li","year":"2008","journal-title":"J. Bacteriol."},{"key":"ref74","doi-asserted-by":"publisher","first-page":"53","DOI":"10.1007\/s42773-025-00453-3","article-title":"Quantitative analysis of the mechanism of biochar in alleviating product inhibition in different fermentative hydrogen production processes","volume":"7","author":"Li","year":"","journal-title":"Biochar"},{"key":"ref75","doi-asserted-by":"publisher","first-page":"53","DOI":"10.1007\/s42773-025-00453-3","article-title":"Quantitative analysis of the mechanism of biochar in alleviating product inhibition in different fermentative hydrogen production processes","volume":"7","author":"Li","year":"","journal-title":"Biochar"},{"key":"ref76","doi-asserted-by":"publisher","first-page":"122504","DOI":"10.1016\/j.biortech.2019.122504","article-title":"Enhancement of bio-hydrogen yield and pH stability in photo fermentation process using dark fermentation effluent as succedaneum","volume":"297","author":"Li","year":"2020","journal-title":"Bioresour. Technol."},{"key":"ref77","doi-asserted-by":"publisher","first-page":"132822","DOI":"10.1016\/j.biortech.2025.132822","article-title":"Biohydrogen production from food waste using a novel rotational drum reactor integrated with milli-magnetite","volume":"434","author":"Liu","year":"2025","journal-title":"Bioresour. Technol."},{"key":"ref78","doi-asserted-by":"publisher","first-page":"171","DOI":"10.1196\/annals.1419.019","article-title":"Metabolic, phylogenetic, and ecological diversity of the methanogenic Archaea","volume":"1125","author":"Liu","year":"2008","journal-title":"Ann. N. Y. Acad. Sci."},{"key":"ref79","doi-asserted-by":"publisher","first-page":"315","DOI":"10.1016\/j.ijhydene.2024.01.039","article-title":"Recent advances in effect of biochar on fermentative hydrogen production: performance and mechanisms","volume":"57","author":"Lou","year":"2024","journal-title":"Int. J. Hydrog. Energy"},{"key":"ref80","doi-asserted-by":"publisher","first-page":"967","DOI":"10.1016\/j.ijhydene.2024.05.473","article-title":"Analysis of the contribution of different electron transfer pathways for hydrogen production in a bioelectrochemically assisted dark fermentation system","volume":"72","author":"Lv","year":"2024","journal-title":"Int. J. Hydrog. Energy"},{"key":"ref81","doi-asserted-by":"publisher","first-page":"540","DOI":"10.1016\/j.rser.2015.02.032","article-title":"Review on research achievements of biogas from anaerobic digestion","volume":"45","author":"Mao","year":"2015","journal-title":"Renew. Sust. Energ. Rev."},{"key":"ref82","doi-asserted-by":"publisher","first-page":"1231","DOI":"10.3390\/pr13041231","article-title":"Optimizing hydrogen production through efficient organic matter oxidation performed by microbial electrolysis cells","volume":"13","author":"Marchetti","year":"2025","journal-title":"PRO"},{"key":"ref83","doi-asserted-by":"publisher","first-page":"5165","DOI":"10.1016\/j.ijhydene.2019.07.054","article-title":"Biohydrogen and biomethane production from cassava wastewater in a two-stage anaerobic sequencing batch biofilm reactor","volume":"45","author":"Mari","year":"2020","journal-title":"Int. J. Hydrog. Energy"},{"key":"ref84","doi-asserted-by":"publisher","first-page":"128070","DOI":"10.1016\/j.biortech.2022.128070","article-title":"Influence of key operational parameters on biohydrogen production from fruit and vegetable waste via lactate-driven dark fermentation","volume":"364","author":"Mart\u00ednez-Mendoza","year":"2022","journal-title":"Bioresour. Technol."},{"key":"ref85","doi-asserted-by":"publisher","first-page":"236","DOI":"10.1263\/jbb.103.236","article-title":"Hydrogen production by fermentation using acetic acid and lactic acid","volume":"103","author":"Matsumoto","year":"2007","journal-title":"J. Biosci. Bioeng."},{"key":"ref86","doi-asserted-by":"publisher","first-page":"7600","DOI":"10.1073\/pnas.0610456104","article-title":"The genome of Syntrophus aciditrophicus: life at the thermodynamic limit of microbial growth","volume":"104","author":"McInerney","year":"2007","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref87","doi-asserted-by":"publisher","first-page":"111","DOI":"10.1186\/s12934-024-02380-7","article-title":"Unlocking the genome of the non-sourdough Kazachstania humilis MAW1: insights into inhibitory factors and phenotypic properties","volume":"23","author":"Mielecki","year":"2024","journal-title":"Microb. Cell Factories"},{"key":"ref88","doi-asserted-by":"publisher","first-page":"2209","DOI":"10.1039\/D5SE01709B","article-title":"Genome-resolved insights into Ni\/Fe2O3 nanocatalyst-enhanced dark fermentative hydrogen production from food waste","volume":"10","author":"Mishra","year":"2026","journal-title":"Sustain. Energy Fuels"},{"key":"ref89","doi-asserted-by":"publisher","first-page":"126327","DOI":"10.1016\/j.biortech.2021.126327","article-title":"Electron bifurcation reactions in dark fermentation: an overview for better understanding and improvement","volume":"344","author":"Mostafa","year":"2022","journal-title":"Bioresour. Technol."},{"key":"ref90","doi-asserted-by":"publisher","first-page":"6167","DOI":"10.1128\/JB.01605-08","article-title":"Involvement of NADH:acceptor oxidoreductase and butyryl coenzyme a dehydrogenase in reversed electron transport during syntrophic butyrate oxidation by Syntrophomonas wolfei","volume":"191","author":"M\u00fcller","year":"2009","journal-title":"J. Bacteriol."},{"key":"ref91","doi-asserted-by":"publisher","first-page":"615","DOI":"10.1111\/j.1574-6941.2011.01085.x","article-title":"Kinetic modelling of lactate utilization and butyrate production by key human colonic bacterial species: modelling butyrate production by colonic bacterial species","volume":"76","author":"Mu\u00f1oz-Tamayo","year":"2011","journal-title":"FEMS Microbiol. Ecol."},{"key":"ref92","doi-asserted-by":"publisher","first-page":"1","DOI":"10.15341\/mast(2375-9402)\/01.01.2015\/001","article-title":"Dry ice and liquefied biomethane \u2014 two products from biogas for an energetic and economical upgrading of biogas plants","volume":"1","author":"Nachtmann","year":"2015","journal-title":"MAST"},{"key":"ref93","first-page":"331","volume-title":"Water and Wastewater Treatment Technologies","author":"Nguyen","year":"2019"},{"key":"ref94","doi-asserted-by":"publisher","first-page":"924","DOI":"10.1016\/j.renene.2019.09.024","article-title":"Enhanced hydrogen production by a sequential dark and photo fermentation process: effects of initial feedstock composition, dilution and microbial population","volume":"147","author":"Ni\u00f1o-Navarro","year":"2020","journal-title":"Renew. Energy"},{"key":"ref95","doi-asserted-by":"publisher","first-page":"21","DOI":"10.1007\/s12649-009-9001-2","article-title":"Biohydrogen production from biomass and wastes via dark fermentation: a review","volume":"1","author":"Ntaikou","year":"2010","journal-title":"Waste Biomass Valoriz."},{"key":"ref96","doi-asserted-by":"publisher","first-page":"20683","DOI":"10.1007\/s13399-025-06620-7","article-title":"Valorization of green market waste for sequential biohydrogen and biomethane production","volume":"15","author":"\u00d6zm\u0131h\u00e7\u0131","year":"2025","journal-title":"Biomass Conv. Bioref."},{"key":"ref97","doi-asserted-by":"publisher","first-page":"2465","DOI":"10.1007\/s00253-018-8737-7","article-title":"Hydrogen metabolic patterns driven by Clostridium-Streptococcus community shifts in a continuous stirred tank reactor","volume":"102","author":"Palomo-Briones","year":"2018","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref98","doi-asserted-by":"publisher","first-page":"11","DOI":"10.1186\/s42834-021-00085-9","article-title":"Physicochemical, microbiological characterization and phytotoxicity of digestates produced on single-stage and two-stage anaerobic digestion of food waste","volume":"31","author":"Parra-Orobio","year":"2021","journal-title":"Sustain. Environ. Res."},{"key":"ref99","doi-asserted-by":"publisher","first-page":"117","DOI":"10.1016\/j.ijhydene.2021.10.028","article-title":"Enhancing photo fermentative hydrogen production using ethanol rich dark fermentation effluents","volume":"47","author":"Policastro","year":"2022","journal-title":"Int. J. Hydrog. Energy"},{"key":"ref100","doi-asserted-by":"publisher","first-page":"579","DOI":"10.1016\/j.ijhydene.2024.12.175","article-title":"Peanut shell activated carbon doped with nickel-iron nanoparticles as material for improving dark fermentative hydrogen production by Enterobacter aerogenes","volume":"99","author":"Ramprakash","year":"2025","journal-title":"Int. J. Hydrog. Energy"},{"key":"ref101","doi-asserted-by":"publisher","first-page":"112773","DOI":"10.1016\/j.rser.2022.112773","article-title":"Current strategies and future perspectives in biological hydrogen production: a review","volume":"168","author":"Ramprakash","year":"2022","journal-title":"Renew. Sust. Energ. Rev."},{"key":"ref102","doi-asserted-by":"publisher","first-page":"127238","DOI":"10.1016\/j.fuel.2022.127238","article-title":"Elucidating the role of pH and total solids content in the co-production of biohydrogen and carboxylic acids from food waste via lactate-driven dark fermentation","volume":"338","author":"Regueira-Marcos","year":"2023","journal-title":"Fuel"},{"key":"ref103","doi-asserted-by":"crossref","first-page":"141","DOI":"10.1016\/B978-0-444-64203-5.00006-X","volume-title":"Biohydrogen","author":"Sa\u011f\u0131r","year":"2019"},{"key":"ref104","doi-asserted-by":"publisher","first-page":"335","DOI":"10.1016\/j.apenergy.2014.03.024","article-title":"Can two-stage instead of one-stage anaerobic digestion really increase energy recovery from biomass?","volume":"124","author":"Schievano","year":"2014","journal-title":"Appl. Energy"},{"key":"ref105","doi-asserted-by":"publisher","first-page":"8502","DOI":"10.1021\/es301376n","article-title":"Two-stage vs single-stage thermophilic anaerobic digestion: comparison of energy production and biodegradation efficiencies","volume":"46","author":"Schievano","year":"2012","journal-title":"Environ. Sci. Technol."},{"key":"ref106","doi-asserted-by":"crossref","first-page":"471","DOI":"10.1007\/978-3-642-30123-0_59","volume-title":"The Prokaryotes","author":"Schink","year":"2013"},{"key":"ref107","doi-asserted-by":"publisher","first-page":"e115902","DOI":"10.1371\/journal.pone.0115902","article-title":"Degradation of acetaldehyde and its precursors by Pelobacter carbinolicus and P. acetylenicus","volume":"9","author":"Schmidt","year":"2014","journal-title":"PLoS One"},{"key":"ref108","doi-asserted-by":"publisher","first-page":"e56905","DOI":"10.1371\/journal.pone.0056905","article-title":"A proteomic view at the biochemistry of syntrophic butyrate oxidation in Syntrophomonas wolfei","volume":"8","author":"Schmidt","year":"2013","journal-title":"PLoS One"},{"key":"ref109","doi-asserted-by":"publisher","first-page":"547","DOI":"10.1016\/j.rser.2015.08.065","article-title":"State of the art and future concept of food waste fermentation to bioenergy","volume":"53","author":"Sen","year":"2016","journal-title":"Renew. Sust. Energ. Rev."},{"key":"ref110","doi-asserted-by":"publisher","first-page":"355","DOI":"10.1007\/s002530050696","article-title":"Biochemical route and control of butyrate synthesis in Butyribacterium methylotrophicum","volume":"45","author":"Shen","year":"1996","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref111","doi-asserted-by":"crossref","first-page":"489","DOI":"10.1002\/9783527843596.ch15","volume-title":"Iron-Sulfur Clusters","author":"Shima","year":"2025"},{"key":"ref112","doi-asserted-by":"publisher","first-page":"429","DOI":"10.1146\/annurev-micro-090110-102844","article-title":"Genomic insights into Syntrophy: the paradigm for anaerobic metabolic cooperation","volume":"66","author":"Sieber","year":"2012","journal-title":"Ann. Rev. Microbiol."},{"key":"ref113","doi-asserted-by":"publisher","first-page":"18","DOI":"10.15199\/17.2022.12.4","article-title":"Mikroorganizmy jako producenci biowodoru: Od bada\u0144 podstawowych do prac rozwojowych z dziedziny biotechnologii","volume":"1","author":"Sikora","year":"2022","journal-title":"GAS"},{"key":"ref114","volume-title":"Lactic Acid Bacteria - R & D for Food, Health and Livestock Purposes","author":"Sikora","year":"2013"},{"key":"ref115","volume-title":"Fermentation Processes","author":"Sikora","year":"2017"},{"key":"ref116","doi-asserted-by":"publisher","first-page":"121721","DOI":"10.1016\/j.jclepro.2020.121721","article-title":"Production of hydrogen and methane from lignocellulose waste by fermentation. A review of chemical pretreatment for enhancing the efficiency of the digestion process","volume":"267","author":"So\u0142owski","year":"2020","journal-title":"J. Clean. Prod."},{"key":"ref117","doi-asserted-by":"publisher","first-page":"7516","DOI":"10.1073\/pnas.1912289117","article-title":"Functional cooperation of the glycine synthase-reductase and wood\u2013Ljungdahl pathways for autotrophic growth of Clostridium drakei","volume":"117","author":"Song","year":"2020","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref118","doi-asserted-by":"publisher","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":"ref119","doi-asserted-by":"publisher","first-page":"151546","DOI":"10.1016\/j.ijhydene.2025.151546","article-title":"Dark fermentation of biomass for enhanced hydrogen production: a review of pretreatment strategies, microbial enhancement, and process regulation","volume":"175","author":"Sun","year":"2025","journal-title":"Int. J. Hydrog. Energy"},{"key":"ref120","doi-asserted-by":"publisher","first-page":"983","DOI":"10.1002\/er.4342","article-title":"Optimization and kinetic modeling of an enhanced bio-hydrogen fermentation with the addition of synergistic biochar and nickel nanoparticle","volume":"43","author":"Sun","year":"2019","journal-title":"Int. J. Energy Res."},{"key":"ref121","doi-asserted-by":"publisher","first-page":"29","DOI":"10.1016\/j.biortech.2016.07.089","article-title":"Effect of biochar addition on hydrogen and methane production in two-phase anaerobic digestion of aqueous carbohydrates food waste","volume":"219","author":"Sunyoto","year":"2016","journal-title":"Bioresour. Technol."},{"key":"ref122","doi-asserted-by":"publisher","first-page":"761","DOI":"10.1007\/s11157-015-9383-5","article-title":"Biohydrogen production by dark fermentation: scaling-up and technologies integration for a sustainable system","volume":"14","author":"Tapia-Venegas","year":"2015","journal-title":"Rev. Environ. Sci. Biotechnol."},{"key":"ref123","doi-asserted-by":"publisher","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":"ref124","doi-asserted-by":"publisher","first-page":"17766","DOI":"10.1021\/acs.est.4c05994","article-title":"Facilitating intracellular Electron bifurcation by mediating Flavin-based extracellular and transmembrane Electron transfer: a novel role of pyrogenic carbon in dark fermentation for hydrogen production","volume":"58","author":"Tian","year":"2024","journal-title":"Environ. Sci. Technol."},{"key":"ref125","doi-asserted-by":"publisher","first-page":"126580","DOI":"10.1016\/j.biortech.2021.126580","article-title":"Valorization of sugar beet pulp to value-added products: a review","volume":"346","author":"Usmani","year":"2022","journal-title":"Bioresour. Technol."},{"key":"ref126","doi-asserted-by":"publisher","first-page":"670","DOI":"10.1111\/1462-2920.12493","article-title":"A novel mode of lactate metabolism in strictly anaerobic bacteria","volume":"17","author":"Weghoff","year":"2015","journal-title":"Environ. Microbiol."},{"key":"ref127","doi-asserted-by":"publisher","first-page":"fuab057","DOI":"10.1093\/femsre\/fuab057","article-title":"Syntrophic propionate-oxidizing bacteria in methanogenic systems","volume":"46","author":"Westerholm","year":"2022","journal-title":"FEMS Microbiol. Rev."},{"key":"ref128","doi-asserted-by":"publisher","first-page":"100","DOI":"10.1111\/j.1574-6968.2010.02023.x","article-title":"Syntrophaceticus schinkii gen. Nov., sp. nov., an anaerobic, syntrophic acetate-oxidizing bacterium isolated from a mesophilic anaerobic filter","volume":"309","author":"Westerholm","year":"2010","journal-title":"FEMS Microbiol. Lett."},{"key":"ref129","doi-asserted-by":"publisher","first-page":"522","DOI":"10.1016\/j.energy.2012.02.048","article-title":"A novel CO2 cryogenic liquefaction and separation system","volume":"42","author":"Xu","year":"2012","journal-title":"Energy"},{"key":"ref130","doi-asserted-by":"publisher","first-page":"117946","DOI":"10.1016\/j.envres.2023.117946","article-title":"Enhanced thermophilic dark fermentation of hydrogen production from food waste by Fe-modified biochar","volume":"244","author":"Yang","year":"2024","journal-title":"Environ. Res."},{"key":"ref131","doi-asserted-by":"publisher","first-page":"130","DOI":"10.1016\/j.rser.2018.07.029","article-title":"Various additives for improving dark fermentative hydrogen production: a review","volume":"95","author":"Yang","year":"2018","journal-title":"Renew. Sust. Energ. Rev."},{"key":"ref132","doi-asserted-by":"publisher","first-page":"4761","DOI":"10.1016\/j.ijhydene.2007.07.038","article-title":"Biohydrogen production from cheese processing wastewater by anaerobic fermentation using mixed microbial communities","volume":"32","author":"Yang","year":"2007","journal-title":"Int. J. Hydrog. Energy"},{"key":"ref133","doi-asserted-by":"publisher","first-page":"333","DOI":"10.1007\/s13205-025-04507-y","article-title":"Valorization of digestates from organic solid waste as fertilizers, soil improvers, and agricultural prebiotics: panorama and perspectives","volume":"15","author":"Zapata-Morales","year":"2025","journal-title":"3 Biotech"},{"key":"ref134","doi-asserted-by":"publisher","first-page":"4539","DOI":"10.1038\/s41467-024-48790-4","article-title":"Biological fermentation pilot-scale systems and evaluation for commercial viability towards sustainable biohydrogen production","volume":"15","author":"Zhang","year":"","journal-title":"Nat. Commun."},{"key":"ref135","doi-asserted-by":"publisher","first-page":"4539","DOI":"10.1038\/s41467-024-48790-4","article-title":"Biological fermentation pilot-scale systems and evaluation for commercial viability towards sustainable biohydrogen production","volume":"15","author":"Zhang","year":"","journal-title":"Nat. Commun."},{"key":"ref136","doi-asserted-by":"publisher","first-page":"148444","DOI":"10.1016\/j.cej.2023.148444","article-title":"Advances in the biomass valorization in dark fermentation systems: a sustainable approach for biohydrogen production","volume":"481","author":"Zhao","year":"2024","journal-title":"Chem. Eng. J."},{"key":"ref137","doi-asserted-by":"publisher","first-page":"100971","DOI":"10.1016\/j.ceja.2025.100971","article-title":"Extremely effective self-sufficient sugar factory wastewater treatment plant and its methanogenic microbial consortium","volume":"25","author":"Zielenkiewicz","year":"2026","journal-title":"Chem. Eng. J. Adv."}],"updated-by":[{"DOI":"10.3389\/fmicb.2026.1963866","type":"correction","label":"Correction","source":"publisher","updated":{"date-parts":[[2026,9,9]],"date-time":"2026-09-09T00:00:00Z","timestamp":1788912000000}},{"DOI":"10.3389\/fmicb.2026.2010146","type":"correction","label":"Correction","source":"publisher","updated":{"date-parts":[[2026,10,5]],"date-time":"2026-10-05T00:00:00Z","timestamp":1791158400000}}],"container-title":["Frontiers in Microbiology"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fmicb.2026.1881665\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,5]],"date-time":"2026-10-05T15:12:38Z","timestamp":1791213158000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fmicb.2026.1881665\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,28]]},"references-count":137,"alternative-id":["10.3389\/fmicb.2026.1881665"],"URL":"https:\/\/doi.org\/10.3389\/fmicb.2026.1881665","relation":{},"ISSN":["1664-302X"],"issn-type":[{"value":"1664-302X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,28]]},"article-number":"1881665"}}