{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,2,6]],"date-time":"2026-02-06T02:06:24Z","timestamp":1770343584876,"version":"3.49.0"},"reference-count":36,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2025,12,9]],"date-time":"2025-12-09T00:00:00Z","timestamp":1765238400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2025,12,10]],"date-time":"2025-12-10T00:00:00Z","timestamp":1765324800000},"content-version":"vor","delay-in-days":1,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100002347","name":"Bundesministerium f\u00fcr Bildung und Forschung","doi-asserted-by":"publisher","award":["Grant No. 031B1056"],"award-info":[{"award-number":["Grant No. 031B1056"]}],"id":[{"id":"10.13039\/501100002347","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Karlsruher Institut f\u00fcr Technologie (KIT)"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Appl Microbiol Biotechnol"],"abstract":"\n Abstract<\/jats:title>\n \n Parageobacillus thermoglucosidasius<\/jats:italic>\n is a carboxydotrophic microorganism that produces H\n 2<\/jats:sub>\n through the water-gas shift (WGS) reaction, using carbon monoxide (CO) as the main substrate. CO is a common constituent of syngas, alongside CO\n 2<\/jats:sub>\n , H\n 2<\/jats:sub>\n , O\n 2<\/jats:sub>\n , and other gases. The facultatively anaerobic nature of\n P. thermoglucosidasius<\/jats:italic>\n is particularly pertinent for hydrogenogenesis from O\n 2<\/jats:sub>\n -containing syngas. Here, we evaluated the effects of different syngas compositions (5, 12, and 20% of H\n 2<\/jats:sub>\n gas, with constant CO and CO\n 2<\/jats:sub>\n ; 10, 30, and 50% CO gas with constant CO\n 2<\/jats:sub>\n and H\n 2<\/jats:sub>\n ) on hydrogenogenesis at the bioreactor scale. Electron balance analysis showed that 88\u201391% of electrons coming from CO were converted into H\n 2<\/jats:sub>\n , regardless of the gas composition. The presence of H\n 2<\/jats:sub>\n in different compositions had no inhibitory effect on hydrogen production rate (HPR), and the maximum HPR corresponded to 13.65 L H\n 2<\/jats:sub>\n L\u207b\n 1<\/jats:sup>\n day\u207b\n 1<\/jats:sup>\n in\u00a0fermentations containing 30% CO. A carbon source, other than CO, is needed for biomass formation of\n P. thermoglucosidasius<\/jats:italic>\n . Acetate was shown to be the primary intermediate metabolite of glucose metabolism, but could also be used as an initial carbon source for biomass generation. When this carbon source was used, most electrons from CO were converted to H\n 2<\/jats:sub>\n , demonstrating that this organic acid can be used as an effective alternative to glucose for H\n 2<\/jats:sub>\n production with\n P. thermoglucosidasius<\/jats:italic>\n .\n <\/jats:p>\n <\/jats:sec>\n \n Key points<\/jats:title>\n \n \u2022\n Evaluation of lab-defined syngas at different compositions for H2 production with P. thermoglucosidasius at the bioreactor scale<\/jats:italic>\n .\n <\/jats:p>\n \n \u2022\n Hydrogen presence in the headspace was not inhibiting for subsequent H2 production<\/jats:italic>\n .\n <\/jats:p>\n \n \u2022\n Acetate can replace glucose to generate biomass when growing P. thermoglucosidasius<\/jats:italic>\n .\n <\/jats:p>\n <\/jats:sec>","DOI":"10.1007\/s00253-025-13659-z","type":"journal-article","created":{"date-parts":[[2025,12,9]],"date-time":"2025-12-09T20:33:47Z","timestamp":1765312427000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Evaluation of lab-defined syngas and acetate as substrates for H2 production with Parageobacillus thermoglucosidasius DSM 6285"],"prefix":"10.1007","volume":"109","author":[{"given":"Magda S.","family":"Ardila","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Habibu","family":"Aliyu","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Pieter","family":"de Maayer","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Anke","family":"Neumann","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"297","published-online":{"date-parts":[[2025,12,9]]},"reference":[{"key":"13659_CR1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-023-41105-z","author":"P Achakulwisut","year":"2023","unstructured":"Achakulwisut P, Erickson P, Guivarch C, Schaeffer R, Brutschin E, Pye S (2023) Global fossil fuel reduction pathways under different climate mitigation strategies and ambitions. Nat Commun. https:\/\/doi.org\/10.1038\/s41467-023-41105-z","journal-title":"Nat Commun"},{"key":"13659_CR2","doi-asserted-by":"publisher","DOI":"10.3390\/microorganisms9051090","author":"H Aliyu","year":"2021","unstructured":"Aliyu H, Kastner R, De Maayer P, Neumann A (2021) Carbon monoxide induced metabolic shift in the carboxydotrophic Parageobacillus thermoglucosidasius DSM 6285. Microorganisms. https:\/\/doi.org\/10.3390\/microorganisms9051090","journal-title":"Microorganisms"},{"key":"13659_CR3","doi-asserted-by":"publisher","DOI":"10.3389\/fmicb.2020.00058","author":"AL Arantes","year":"2020","unstructured":"Arantes AL, Moreira JPC, Diender M, Parshina SN, Stams AJM, Alves MM, Alves JI, Sousa DZ (2020) Enrichment of Anaerobic Syngas-Converting Communities and Isolation of a Novel Acetobacterium wieringae Strain JM. Front Microbiol. https:\/\/doi.org\/10.3389\/fmicb.2020.00058","journal-title":"Front Microbiol"},{"key":"13659_CR4","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1186\/s13068-024-02597-z","volume":"18","author":"MS Ardila","year":"2025","unstructured":"Ardila MS, Aliyu H, de Maayer P, Neumann A (2025) Batch and semi-continuous fermentation with Parageobacillus thermoglucosidasius DSM 6285 for H2 production. Biotechnol Biofuels Bioprod 18:3. https:\/\/doi.org\/10.1186\/s13068-024-02597-z","journal-title":"Biotechnol Biofuels Bioprod"},{"key":"13659_CR5","doi-asserted-by":"publisher","DOI":"10.3390\/fermentation10110592","author":"MS Ardila","year":"2024","unstructured":"Ardila MS, Aliyu H, de Maayer P, Neumann A (2024) Effect of different partial pressures on H2 production with Parageobacillus thermoglucosidasius DSM 6285. Fermentation. https:\/\/doi.org\/10.3390\/fermentation10110592","journal-title":"Fermentation"},{"key":"13659_CR6","doi-asserted-by":"publisher","first-page":"5202","DOI":"10.1016\/j.ijhydene.2019.09.006","volume":"45","author":"G Balachandar","year":"2020","unstructured":"Balachandar G, Varanasi JL, Singh V, Singh H, Das D (2020) Biological hydrogen production via dark fermentation: a holistic approach from lab-scale to pilot-scale. Int J Hydrogen Energy 45:5202\u20135215. https:\/\/doi.org\/10.1016\/j.ijhydene.2019.09.006","journal-title":"Int J Hydrogen Energy"},{"key":"13659_CR7","doi-asserted-by":"publisher","DOI":"10.1007\/BF01121069","author":"VI Baranenko","year":"1989","unstructured":"Baranenko VI, Kirov VS (1989) Solubility of hydrogen in water in a broad temperature and pressure range. Sov Atomic Energy. https:\/\/doi.org\/10.1007\/BF01121069","journal-title":"Sov Atomic Energy"},{"key":"13659_CR8","doi-asserted-by":"publisher","first-page":"391","DOI":"10.3390\/reactions2040025","volume":"2","author":"C Benevenuti","year":"2021","unstructured":"Benevenuti C, Amaral P, Ferreira T, Seidl P (2021) Impacts of syngas composition on anaerobic fermentation. Reactions 2:391\u2013407. https:\/\/doi.org\/10.3390\/reactions2040025","journal-title":"Reactions"},{"key":"13659_CR9","doi-asserted-by":"publisher","DOI":"10.1016\/j.envpol.2021.116843","author":"YH Chan","year":"2021","unstructured":"Chan YH, Syed Abdul Rahman SNF, Lahuri HM, Khalid A (2021) Recent progress on CO-rich syngas production via CO2 gasification of various wastes: a critical review on efficiency, challenges and outlook. Environ Pollut. https:\/\/doi.org\/10.1016\/j.envpol.2021.116843","journal-title":"Environ Pollut"},{"key":"13659_CR10","doi-asserted-by":"publisher","first-page":"715","DOI":"10.1016\/j.cej.2019.04.191","volume":"372","author":"B Chezeau","year":"2019","unstructured":"Chezeau B, Fontaine JP, Vial Ch (2019) Analysis of liquid-to-gas mass transfer, mixing and hydrogen production in dark fermentation process. Chem Eng J 372:715\u2013727. https:\/\/doi.org\/10.1016\/j.cej.2019.04.191","journal-title":"Chem Eng J"},{"key":"13659_CR11","doi-asserted-by":"publisher","DOI":"10.3389\/fenrg.2021.642162","author":"J Collis","year":"2021","unstructured":"Collis J, Strunge T, Steubing B, Zimmermann A, Schom\u00e4cker R (2021) Deriving economic potential and GHG emissions of steel mill gas for chemical industry. Front Energy Res. https:\/\/doi.org\/10.3389\/fenrg.2021.642162","journal-title":"Front Energy Res"},{"key":"13659_CR12","doi-asserted-by":"publisher","first-page":"242","DOI":"10.1021\/je60062a007","volume":"19","author":"TE Crozier","year":"1974","unstructured":"Crozier TE, Yamamoto S (1974) Solubility of hydrogen in water, seawater, and NaCl solutions. J Chem Eng Data 19:242\u2013244","journal-title":"J Chem Eng Data"},{"key":"13659_CR13","doi-asserted-by":"publisher","first-page":"279","DOI":"10.1002\/bit.21226","volume":"97","author":"YS Do","year":"2007","unstructured":"Do YS, Smeenk J, Broer KM, Kisting CJ, Brown R, Heindel TJ, Bobik TA, DiSpirito AA (2007) Growth of Rhodospirillum rubrum on synthesis gas: conversion of CO to H2 and poly-\u03b2-hydroxyalkanoate. Biotechnol Bioeng 97:279\u2013286. https:\/\/doi.org\/10.1002\/bit.21226","journal-title":"Biotechnol Bioeng"},{"key":"13659_CR14","doi-asserted-by":"publisher","first-page":"220","DOI":"10.1186\/s13068-017-0910-1","volume":"10","author":"S Esquivel-Elizondo","year":"2017","unstructured":"Esquivel-Elizondo S, Delgado AG, Rittmann BE, Krajmalnik-Brown R (2017) The effects of CO2 and H2 on CO metabolism by pure and mixed microbial cultures. Biotechnol Biofuels 10:220. https:\/\/doi.org\/10.1186\/s13068-017-0910-1","journal-title":"Biotechnol Biofuels"},{"key":"13659_CR15","doi-asserted-by":"publisher","first-page":"761","DOI":"10.1038\/ismej.2015.153","volume":"10","author":"C Greening","year":"2016","unstructured":"Greening C, Biswas A, Carere CR, Jackson CJ, Taylor MC, Stott MB, Cook GM, Morales SE (2016) Genomic and metagenomic surveys of hydrogenase distribution indicate H2 is a widely utilised energy source for microbial growth and survival. ISME J 10:761\u2013777. https:\/\/doi.org\/10.1038\/ismej.2015.153","journal-title":"ISME J"},{"key":"13659_CR16","doi-asserted-by":"publisher","first-page":"2543","DOI":"10.1016\/j.ijhydene.2013.12.022","volume":"39","author":"M Haddad","year":"2014","unstructured":"Haddad M, Cimpoia R, Guiot SR (2014) Performance of Carboxydothermus hydrogenoformans in a gas-lift reactor for syngas upgrading into hydrogen. Int J Hydrogen Energy 39:2543\u20132548. https:\/\/doi.org\/10.1016\/j.ijhydene.2013.12.022","journal-title":"Int J Hydrogen Energy"},{"key":"13659_CR17","doi-asserted-by":"publisher","DOI":"10.3390\/microorganisms11040995","author":"BM Harahap","year":"2023","unstructured":"Harahap BM, Ahring BK (2023) Acetate production from syngas produced from lignocellulosic biomass materials along with gaseous fermentation of the syngas: a review. Microorganisms. https:\/\/doi.org\/10.3390\/microorganisms11040995","journal-title":"Microorganisms"},{"key":"13659_CR18","doi-asserted-by":"publisher","first-page":"118","DOI":"10.1264\/jsme2.23.118","volume":"23","author":"S Hattori","year":"2008","unstructured":"Hattori S (2008) Syntrophic acetate-oxidizing microbes in methanogenic environments. Microbes Environ 23:118\u2013127. https:\/\/doi.org\/10.1264\/jsme2.23.118","journal-title":"Microbes Environ"},{"key":"13659_CR19","doi-asserted-by":"publisher","DOI":"10.1016\/j.biortech.2024.130972","author":"N Hern\u00e1ndez-Herreros","year":"2024","unstructured":"Hern\u00e1ndez-Herreros N, Rodr\u00edguez A, Gal\u00e1n B, Auxiliadora Prieto M (2024) Boosting hydrogen production in Rhodospirillum rubrum by syngas-driven photoheterotrophic adaptive evolution. Bioresour Technol. https:\/\/doi.org\/10.1016\/j.biortech.2024.130972","journal-title":"Bioresour Technol"},{"key":"13659_CR20","doi-asserted-by":"publisher","first-page":"1907","DOI":"10.1038\/s41396-021-00893-2","volume":"15","author":"S Joshi","year":"2021","unstructured":"Joshi S, Robles A, Aguiar S, Delgado AG (2021) The occurrence and ecology of microbial chain elongation of carboxylates in soils. ISME J 15:1907\u20131918. https:\/\/doi.org\/10.1038\/s41396-021-00893-2","journal-title":"ISME J"},{"key":"13659_CR21","doi-asserted-by":"publisher","DOI":"10.3390\/fermentation5030075","author":"K Liu","year":"2019","unstructured":"Liu K, Phillips JR, Sun X, Mohammad S, Huhnke RL, Atiyeh HK (2019) Investigation and modeling of gas-liquid mass transfer in a sparged and non-sparged continuous stirred tank reactor with potential application in syngas fermentation. Fermentation. https:\/\/doi.org\/10.3390\/fermentation5030075","journal-title":"Fermentation"},{"key":"13659_CR22","first-page":"247","volume-title":"Moura JJG","author":"M Martins","year":"2021","unstructured":"Martins M, Pereira IAC, Pita M, De Lacey AL (2021) Biological production of hydrogen BT - enzymes for solving humankind\u2019s problems: natural and artificial systems in health, agriculture, environment and energy. In: Moura I, Maia LB (eds) Moura JJG. Springer International Publishing, Cham, pp 247\u2013273"},{"key":"13659_CR23","doi-asserted-by":"publisher","DOI":"10.1186\/s13568-019-0931-1","volume":"9","author":"T Mohr","year":"2019","unstructured":"Mohr T, Aliyu H, Biebinger L, G\u00f6dert R, Hornberger A, Cowan D, De Maayer P, Neumann A (2019) Effects of different operating parameters on hydrogen production by Parageobacillus thermoglucosidasius DSM 6285. AMB Express 9:207. https:\/\/doi.org\/10.1186\/s13568-019-0931-1","journal-title":"AMB Express"},{"key":"13659_CR24","doi-asserted-by":"publisher","unstructured":"Mohr T, Aliyu H, K\u00fcchlin R, Polliack S, Zwick M, Neumann A, Cowan D, De Maayer P (2018) CO dependent hydrogen production by the facultative anaerobe Parageobacillus thermoglucosidasius. Microb Cell Fact 1\u201312. https:\/\/doi.org\/10.1186\/s12934-018-0954-3","DOI":"10.1186\/s12934-018-0954-3"},{"key":"13659_CR25","doi-asserted-by":"publisher","first-page":"125","DOI":"10.1186\/s12934-024-02391-4","volume":"23","author":"M Mol","year":"2024","unstructured":"Mol M, Ardila MS, Mol BA, Aliyu H, Neumann A, de Maayer P (2024) The effects of synthesis gas feedstocks and oxygen perturbation on hydrogen production by Parageobacillus thermoglucosidasius. Microb Cell Fact 23:125. https:\/\/doi.org\/10.1186\/s12934-024-02391-4","journal-title":"Microb Cell Fact"},{"key":"13659_CR26","doi-asserted-by":"publisher","first-page":"123","DOI":"10.1016\/j.bej.2004.06.001","volume":"21","author":"G Najafpour","year":"2004","unstructured":"Najafpour G, Younesi H, Mohamed AR (2004) Effect of organic substrate on hydrogen production from synthesis gas using Rhodospirillum rubrum, in batch culture. Biochem Eng J 21:123\u2013130. https:\/\/doi.org\/10.1016\/j.bej.2004.06.001","journal-title":"Biochem Eng J"},{"key":"13659_CR27","doi-asserted-by":"publisher","first-page":"597","DOI":"10.1016\/j.rser.2016.09.044","volume":"67","author":"P Nikolaidis","year":"2017","unstructured":"Nikolaidis P, Poullikkas A (2017) A comparative overview of hydrogen production processes. Renew Sustain Energy Rev 67:597\u2013611. https:\/\/doi.org\/10.1016\/j.rser.2016.09.044","journal-title":"Renew Sustain Energy Rev"},{"key":"13659_CR28","doi-asserted-by":"publisher","DOI":"10.1016\/j.biortech.2023.129894","author":"L Perret","year":"2024","unstructured":"Perret L, Boukis N, Sauer J (2024) Synthesis gas fermentation at high cell density: how pH and hydrogen partial pressure affect productivity and product ratio in continuous fermentation. Bioresour Technol. https:\/\/doi.org\/10.1016\/j.biortech.2023.129894","journal-title":"Bioresour Technol"},{"key":"13659_CR29","doi-asserted-by":"publisher","first-page":"147","DOI":"10.1128\/JB.00147-19","volume":"201","author":"S Pinhal","year":"2019","unstructured":"Pinhal S, Ropers D, Geiselmann J, De Jong H, Metcalf WW (2019) Acetate metabolism and the inhibition of bacterial growth by acetate. J Bacteriol 201:147\u2013166. https:\/\/doi.org\/10.1128\/JB.00147-19","journal-title":"J Bacteriol"},{"key":"13659_CR30","doi-asserted-by":"publisher","first-page":"188","DOI":"10.3934\/bioeng.2016.2.188","volume":"3","author":"K Ramachandriya","year":"2016","unstructured":"Ramachandriya K, Kundiyana D, Sharma A, Kumar A, Atiyeh H, Huhnke R, Wilkins M (2016) Critical factors affecting the integration of biomass gasification and syngas fermentation technology. AIMS Bioeng 3:188\u2013210. https:\/\/doi.org\/10.3934\/bioeng.2016.2.188","journal-title":"AIMS Bioeng"},{"key":"13659_CR31","doi-asserted-by":"publisher","first-page":"21","DOI":"10.1016\/j.indcrop.2016.11.015","volume":"106","author":"S Redl","year":"2017","unstructured":"Redl S, Diender M, Jensen T\u00d8, Sousa DZ, Nielsen AT (2017) Exploiting the potential of gas fermentation. Ind Crops Prod 106:21\u201330. https:\/\/doi.org\/10.1016\/j.indcrop.2016.11.015","journal-title":"Ind Crops Prod"},{"key":"13659_CR32","doi-asserted-by":"publisher","DOI":"10.1111\/1751-7915.70063","author":"A Robazza","year":"2024","unstructured":"Robazza A, Raya i Garcia A, Baleeiro FCF, Kleinsteuber S, Neumann A (2024) Acetate shock loads enhance CO uptake rates of anaerobic microbiomes. Microb Biotechnol. https:\/\/doi.org\/10.1111\/1751-7915.70063","journal-title":"Microb Biotechnol"},{"key":"13659_CR33","doi-asserted-by":"publisher","DOI":"10.1186\/s13068-021-02017-6","author":"A Rodr\u00edguez","year":"2021","unstructured":"Rodr\u00edguez A, Hern\u00e1ndez-Herreros N, Garc\u00eda JL, Auxiliadora Prieto M (2021) Enhancement of biohydrogen production rate in Rhodospirillum rubrum by a dynamic CO-feeding strategy using dark fermentation. Biotechnol Biofuels. https:\/\/doi.org\/10.1186\/s13068-021-02017-6","journal-title":"Biotechnol Biofuels"},{"key":"13659_CR34","doi-asserted-by":"publisher","first-page":"10901","DOI":"10.5194\/acp-23-10901-2023","volume":"23","author":"R Sander","year":"2023","unstructured":"Sander R (2023) Compilation of Henry\u2019s law constants (version 5.0.0) for water as solvent. Atmos Chem Phys 23:10901\u201312440. https:\/\/doi.org\/10.5194\/acp-23-10901-2023","journal-title":"Atmos Chem Phys"},{"key":"13659_CR35","doi-asserted-by":"publisher","first-page":"6215","DOI":"10.1007\/s00253-015-6762-3","volume":"99","author":"J Tr\u010dek","year":"2015","unstructured":"Tr\u010dek J, Mira NP, Jarboe LR (2015) Adaptation and tolerance of bacteria against acetic acid. Appl Microbiol Biotechnol 99:6215\u20136229. https:\/\/doi.org\/10.1007\/s00253-015-6762-3","journal-title":"Appl Microbiol Biotechnol"},{"key":"13659_CR36","doi-asserted-by":"publisher","first-page":"2612","DOI":"10.1016\/j.biortech.2007.04.059","volume":"99","author":"H Younesi","year":"2008","unstructured":"Younesi H, Najafpour G, Ku Ismail KS, Mohamed AR, Kamaruddin AH (2008) Biohydrogen production in a continuous stirred tank bioreactor from synthesis gas by anaerobic photosynthetic bacterium: Rhodopirillum rubrum. Bioresour Technol 99:2612\u20132619. https:\/\/doi.org\/10.1016\/j.biortech.2007.04.059","journal-title":"Bioresour Technol"}],"container-title":["Applied Microbiology and Biotechnology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00253-025-13659-z.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s00253-025-13659-z","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00253-025-13659-z.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,2,5]],"date-time":"2026-02-05T12:32:01Z","timestamp":1770294721000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s00253-025-13659-z"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,12,9]]},"references-count":36,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2025,12]]}},"alternative-id":["13659"],"URL":"https:\/\/doi.org\/10.1007\/s00253-025-13659-z","relation":{},"ISSN":["1432-0614"],"issn-type":[{"value":"1432-0614","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,12,9]]},"assertion":[{"value":"9 May 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"14 October 2025","order":2,"name":"revised","label":"Revised","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"24 November 2025","order":3,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"9 December 2025","order":4,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"This article does not contain any studies with human participants or animals performed by any of the authors.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethical approval"}},{"value":"The authors declare no competing interests.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"261"}}