{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,17]],"date-time":"2026-08-17T11:59:35Z","timestamp":1786967975604,"version":"build-2736575974"},"reference-count":34,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2025,10,8]],"date-time":"2025-10-08T00:00:00Z","timestamp":1759881600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2025,10,8]],"date-time":"2025-10-08T00:00:00Z","timestamp":1759881600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100004527","name":"Energimyndigheten","doi-asserted-by":"publisher","award":["P47516-1"],"award-info":[{"award-number":["P47516-1"]}],"id":[{"id":"10.13039\/501100004527","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004359","name":"Vetenskapsr\u00e5det","doi-asserted-by":"publisher","award":["2020-05318"],"award-info":[{"award-number":["2020-05318"]}],"id":[{"id":"10.13039\/501100004359","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Bio4Energy"},{"name":"Umea University"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Biotechnol. Biofuels Bioprod."],"abstract":"<jats:title>Abstract<\/jats:title>\n          <jats:sec>\n            <jats:title>Background<\/jats:title>\n            <jats:p>Aeration plays a critical role in the bioconversion of pretreated lignocellulose by enhancing lytic-polysaccharide-monooxygenase(LPMO)-supported enzymatic saccharification. However, its broader impact, particularly on fermentation inhibitors, remains insufficiently understood. The hypothesis that aeration not only promotes LPMO activity, which has been shown clearly in previous studies, but also affects fermentation inhibitors was investigated in experiments with softwood pretreatment liquids. The effects of aeration were explored through chemical analysis of fermentation inhibitors and through subsequent fermentations with the xylose-utilizing <jats:italic>Saccharomyces cerevisiae<\/jats:italic> yeast CelluX\u21224 to test the fermentability. Controls in which N<jats:sub>2<\/jats:sub> rather than air was supplied to the pretreatment liquids were used to distinguish between evaporation effects and effects caused by oxidation due to O<jats:sub>2<\/jats:sub> in air. In separate experiments, two redox-dependent detoxification methods, treatments with sulfite and laccase, were further investigated.<\/jats:p>\n          <\/jats:sec>\n          <jats:sec>\n            <jats:title>Results<\/jats:title>\n            <jats:p>While aeration had no negative effects on the subsequent fermentation of a sugar control, it compromised the fermentability of the pretreatment liquids. Compared to the N<jats:sub>2<\/jats:sub> control, subsequent fermentation of aerated samples showed reduced consumption of fermentable sugar (glucose, mannose, xylose) at 0.61 compared to 0.76\u00a0g\u00a0L<jats:sup>\u22121<\/jats:sup>\u00a0h<jats:sup>\u22121<\/jats:sup>, and lower ethanol productivity (0.23 vs. 0.30\u00a0g\u00a0L<jats:sup>\u22121<\/jats:sup>\u00a0h<jats:sup>\u22121<\/jats:sup>). Apart from more commonly studied pretreatment by-products (such as aliphatic carboxylic acids, furan aldehydes, and phenolics), methanol (~\u20091\u00a0g\u00a0L<jats:sup>\u22121<\/jats:sup>) was detected in both pretreatment liquids. The methanol concentration decreased during gas addition, which was attributed to evaporation. Compared to the initial pretreatment liquid, aerated reaction mixtures exhibited slightly elevated levels of formaldehyde, but lower levels of furfural and vanillin. Sulfite detoxification was successful under both aeration and N<jats:sub>2<\/jats:sub> conditions. Treatment with laccase was found to have variable effects on the fermentability depending on the conditions applied.<\/jats:p>\n          <\/jats:sec>\n          <jats:sec>\n            <jats:title>Conclusions<\/jats:title>\n            <jats:p>The results underscore the dual role of aeration in softwood bioconversion, positive for promoting LPMO activity but potentially negative with respect to subsequent fermentability, and highlight the need to carefully tailor aeration strategies for the design of efficient biochemical processing of lignocellulosic feedstocks. Treatment with reducing agents, such as sulfite, emerges as a possibility to alleviate negative side-effects on the fermentability when aeration is used to promote LPMO activity.<\/jats:p>\n          <\/jats:sec>","DOI":"10.1186\/s13068-025-02708-4","type":"journal-article","created":{"date-parts":[[2025,10,8]],"date-time":"2025-10-08T07:34:49Z","timestamp":1759908889000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":2,"title":["Effects of aeration of softwood pretreatment liquid on inhibitors and fermentability using Saccharomyces cerevisiae yeast"],"prefix":"10.1186","volume":"18","author":[{"given":"Chaojun","family":"Tang","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Carlos","family":"Mart\u00edn","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Leif J.","family":"J\u00f6nsson","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2025,10,8]]},"reference":[{"key":"2708_CR1","doi-asserted-by":"publisher","first-page":"484","DOI":"10.1126\/science.1114736","volume":"311","author":"AJ Ragauskas","year":"2006","unstructured":"Ragauskas AJ, Williams CK, Davison BH, Britovsek G, Cairney J, Eckert CA, et al. The path forward for biofuels and biomaterials. Science. 2006;311:484\u20139.","journal-title":"Science"},{"key":"2708_CR2","doi-asserted-by":"publisher","first-page":"1210","DOI":"10.1111\/gcbb.12844","volume":"13","author":"AL Cowie","year":"2021","unstructured":"Cowie AL, Berndes G, Bentsen NS, Brand\u00e3o M, Cherubini F, Egnell G, et al. Applying a science-based systems perspective to dispel misconceptions about climate effects of forest bioenergy. GCB Bioenergy. 2021;13:1210\u201331.","journal-title":"GCB Bioenergy"},{"key":"2708_CR3","volume":"19","author":"P B\u00f6rjesson","year":"2023","unstructured":"B\u00f6rjesson P, Bj\u00f6rnsson L, Ericsson K, Lantz M. Systems perspectives on combined production of advanced biojet fuel and biofuels in existing industrial infrastructure in Sweden. Energy Convers Manage. 2023;19:100404.","journal-title":"Energy Convers Manage"},{"key":"2708_CR4","doi-asserted-by":"publisher","first-page":"294","DOI":"10.1186\/s13068-019-1634-1","volume":"12","author":"M Galbe","year":"2019","unstructured":"Galbe M, Wallberg O. Pretreatment for biorefineries: a review of common methods for efficient utilisation of lignocellulosic materials. Biotechnol Biofuels. 2019;12:294.","journal-title":"Biotechnol Biofuels"},{"key":"2708_CR5","doi-asserted-by":"publisher","first-page":"587","DOI":"10.1016\/j.rser.2019.02.024","volume":"107","author":"JC Solarte-Toro","year":"2019","unstructured":"Solarte-Toro JC, Romero-Garc\u00eda JM, Mart\u00ednez-Pati\u00f1o JC, Ruiz-Ramos E, Castro-Galiano E, Cardona-Alzate CA. Acid pretreatment of lignocellulosic biomass for energy vectors production: a review focused on operational conditions and techno-economic assessment for bioethanol production. Renew Sustain Energy Rev. 2019;107:587\u2013601.","journal-title":"Renew Sustain Energy Rev"},{"key":"2708_CR6","doi-asserted-by":"publisher","DOI":"10.3389\/fbioe.2022.846592","volume":"10","author":"C Mart\u00edn","year":"2022","unstructured":"Mart\u00edn C, Dixit P, Momayez F, J\u00f6nsson LJ. Hydrothermal pretreatment of lignocellulosic feedstocks to facilitate biochemical conversion. Front Bioeng Biotechnol. 2022;10:846592.","journal-title":"Front Bioeng Biotechnol"},{"key":"2708_CR7","doi-asserted-by":"publisher","DOI":"10.1016\/j.biortech.2022.128469","volume":"369","author":"HA Ruiz","year":"2023","unstructured":"Ruiz HA, Sganzerla WG, Larnaudie V, Veersma RJ, van Erven G, Shiva, et al. Advances in process design, techno-economic assessment and environmental aspects for hydrothermal pretreatment in the fractionation of biomass under biorefinery concept. Bioresour Technol. 2023;369:128469.","journal-title":"Bioresour Technol"},{"key":"2708_CR8","doi-asserted-by":"publisher","first-page":"200","DOI":"10.1016\/j.biortech.2017.09.081","volume":"247","author":"Z Wang","year":"2018","unstructured":"Wang Z, Wu G, J\u00f6nsson LJ. Effects of impregnation of softwood with sulfuric acid and sulfur dioxide on chemical and physical characteristics, enzymatic digestibility, and fermentability. Bioresour Technol. 2018;247:200\u20138.","journal-title":"Bioresour Technol"},{"key":"2708_CR9","doi-asserted-by":"publisher","first-page":"430","DOI":"10.1002\/bit.22829","volume":"107","author":"B Du","year":"2010","unstructured":"Du B, Sharma LN, Becker C, Chen S-F, Mowery RA, van Walsum GP, et al. Effect of varying feedstock-pretreatment chemistry combinations on the formation and accumulation of potentially inhibitory degradation products in biomass hydrolysates. Biotechnol Bioeng. 2010;107:430\u201340.","journal-title":"Biotechnol Bioeng"},{"key":"2708_CR10","doi-asserted-by":"publisher","first-page":"103","DOI":"10.1016\/j.biortech.2015.10.009","volume":"199","author":"LJ J\u00f6nsson","year":"2016","unstructured":"J\u00f6nsson LJ, Mart\u00edn C. Pretreatment of lignocellulose: formation of inhibitory by-products and strategies for minimizing their effects. Bioresour Technol. 2016;199:103\u201312.","journal-title":"Bioresour Technol"},{"key":"2708_CR11","doi-asserted-by":"publisher","first-page":"16","DOI":"10.1186\/1754-6834-6-16","volume":"6","author":"LJ J\u00f6nsson","year":"2013","unstructured":"J\u00f6nsson LJ, Alriksson B, Nilvebrant N-O. Bioconversion of lignocellulose: inhibitors and detoxification. Biotechnol Biofuels. 2013;6:16.","journal-title":"Biotechnol Biofuels"},{"key":"2708_CR12","doi-asserted-by":"publisher","first-page":"691","DOI":"10.1007\/s002530051233","volume":"49","author":"LJ J\u00f6nsson","year":"1998","unstructured":"J\u00f6nsson LJ, Palmqvist E, Nilvebrant N-O, Hahn-H\u00e4gerdal B. Detoxification of wood hydrolysates with laccase and peroxidase from the white-rot fungus Trametes versicolor. Appl Microbiol Biotechnol. 1998;49:691\u20137.","journal-title":"Appl Microbiol Biotechnol"},{"key":"2708_CR13","doi-asserted-by":"publisher","first-page":"298","DOI":"10.1016\/j.biortech.2011.11.080","volume":"104","author":"M Kolb","year":"2012","unstructured":"Kolb M, Sieber V, Amann M, Faulstich M, Schieder D. Removal of monomer delignification products by laccase from Trametes versicolor. Bioresour Technol. 2012;104:298\u2013304.","journal-title":"Bioresour Technol"},{"key":"2708_CR14","doi-asserted-by":"publisher","first-page":"214","DOI":"10.3390\/fermentation9030214","volume":"9","author":"D Ibarra","year":"2023","unstructured":"Ibarra D, Eugenio ME, Alvira P, Ballesteros I, Ballesteros M, Negro MJ. Effect of laccase detoxification on bioethanol production from liquid fraction of steam-pretreated olive tree pruning. Fermentation. 2023;9:214.","journal-title":"Fermentation"},{"key":"2708_CR15","doi-asserted-by":"publisher","first-page":"1254","DOI":"10.1016\/j.biortech.2010.08.037","volume":"102","author":"B Alriksson","year":"2011","unstructured":"Alriksson B, Cavka A, J\u00f6nsson LJ. Improving the fermentability of enzymatic hydrolysates of lignocellulose through chemical in-situ detoxification with reducing agents. Bioresour Technol. 2011;102:1254\u201363.","journal-title":"Bioresour Technol"},{"key":"2708_CR16","doi-asserted-by":"publisher","first-page":"1458","DOI":"10.1016\/j.biotechadv.2012.03.002","volume":"30","author":"JS Van Dyk","year":"2012","unstructured":"Van Dyk JS, Pletschke BI. A review of lignocellulose bioconversion using enzymatic hydrolysis and synergistic cooperation between enzymes-factors affecting enzymes, conversion and synergy. Biotechnol Adv. 2012;30:1458\u201380.","journal-title":"Biotechnol Adv"},{"key":"2708_CR17","doi-asserted-by":"publisher","first-page":"747","DOI":"10.1016\/j.tibtech.2015.09.006","volume":"33","author":"GR Hemsworth","year":"2015","unstructured":"Hemsworth GR, Johnston EM, Davies GJ, Walton PH. Lytic polysaccharide monooxygenases in biomass conversion. Trends Biotechnol. 2015;33:747\u201361.","journal-title":"Trends Biotechnol"},{"key":"2708_CR18","doi-asserted-by":"publisher","first-page":"551","DOI":"10.1042\/EBC20220156","volume":"67","author":"M S\u00f8rlie","year":"2023","unstructured":"S\u00f8rlie M, Keller MB, Westh P. The interplay between lytic polysaccharide monooxygenases and glycoside hydrolases. Essays Biochem. 2023;67:551\u20139.","journal-title":"Essays Biochem"},{"key":"2708_CR19","doi-asserted-by":"publisher","first-page":"15079","DOI":"10.1073\/pnas.1105776108","volume":"108","author":"RJ Quinlan","year":"2011","unstructured":"Quinlan RJ, Sweeney MD, Leggio LL, Otten H, Poulsen JCN, et al. Insights into the oxidative degradation of cellulose by a copper metalloenzyme that exploits biomass components. Proc Natl Acad Sci U S A. 2011;108:15079\u201384.","journal-title":"Proc Natl Acad Sci U S A"},{"key":"2708_CR20","doi-asserted-by":"publisher","first-page":"480","DOI":"10.1016\/j.biortech.2012.01.116","volume":"110","author":"M Dimarogona","year":"2012","unstructured":"Dimarogona M, Topakas E, Olsson L, Christakopoulos P. Lignin boosts the cellulase performance of a GH-61 enzyme from Sporotrichum thermophile. Bioresour Technol. 2012;110:480\u20137.","journal-title":"Bioresour Technol"},{"key":"2708_CR21","doi-asserted-by":"publisher","DOI":"10.3389\/fbioe.2022.1071159","volume":"10","author":"C Tang","year":"2022","unstructured":"Tang C, Gandla ML, J\u00f6nsson LJ. Comparison of solid and liquid fractions of pretreated Norway spruce as reductants in LPMO-supported saccharification of cellulose. Front Bioeng Biotechnol. 2022;10:1071159.","journal-title":"Front Bioeng Biotechnol"},{"key":"2708_CR22","doi-asserted-by":"publisher","first-page":"800","DOI":"10.1080\/07388551.2019.1619157","volume":"39","author":"L Favaro","year":"2019","unstructured":"Favaro L, Jansen T, van Zyl WH. Exploring industrial and natural Saccharomyces cerevisiae strains for the bio-based economy from biomass: the case of bioethanol. Crit Rev Biotechnol. 2019;39:800\u201316.","journal-title":"Crit Rev Biotechnol"},{"key":"2708_CR23","doi-asserted-by":"publisher","first-page":"149","DOI":"10.1099\/00221287-44-2-149","volume":"44","author":"RH De Deken","year":"1966","unstructured":"De Deken RH. The crabtree effect: a regulatory system in yeast. J Gen Microbiol. 1966;44:149\u201356.","journal-title":"J Gen Microbiol"},{"key":"2708_CR24","doi-asserted-by":"publisher","first-page":"59","DOI":"10.1002\/bit.25098","volume":"111","author":"D Cannella","year":"2014","unstructured":"Cannella D, J\u00f8rgensen H. Do new cellulolytic enzyme preparations affect the industrial strategies for high solids lignocellulosic ethanol production? Biotechnol Bioeng. 2014;111:59\u201368.","journal-title":"Biotechnol Bioeng"},{"key":"2708_CR25","doi-asserted-by":"publisher","first-page":"5192","DOI":"10.1021\/acssuschemeng.8b06092","volume":"7","author":"N Zhong","year":"2019","unstructured":"Zhong N, Chandra R, Saddler JN. Sulfite post-treatment to simultaneously detoxify and improve the enzymatic hydrolysis and fermentation of a steam-pretreated softwood lodgepole pine whole slurry. ACS Sustain Chem Eng. 2019;7:5192\u20139.","journal-title":"ACS Sustain Chem Eng"},{"issue":"12","key":"2708_CR26","doi-asserted-by":"publisher","first-page":"1419723","DOI":"10.3389\/fbioe.2024.1419723","volume":"2024","author":"C Tang","year":"2024","unstructured":"Tang C, Cavka A, Bui M, J\u00f6nsson LJ. Comparison of simultaneous saccharification and fermentation with LPMO-supported hybrid hydrolysis and fermentation. Front Bioeng Biotechnol. 2024;2024(12):1419723.","journal-title":"Front Bioeng Biotechnol"},{"key":"2708_CR27","doi-asserted-by":"publisher","DOI":"10.1186\/s13068-023-02407-y","volume":"16","author":"C Tang","year":"2023","unstructured":"Tang C, Gandla ML, J\u00f6nsson LJ. LPMO-supported saccharification of biomass: effects of continuous aeration of reaction mixtures with variable fractions of water-insoluble solids and cellulolytic enzymes. Biotechnol Biofuels Bioprod. 2023;16:156.","journal-title":"Biotechnol Biofuels Bioprod"},{"key":"2708_CR28","doi-asserted-by":"publisher","first-page":"242","DOI":"10.1016\/j.biortech.2018.04.074","volume":"262","author":"C Mart\u00edn","year":"2018","unstructured":"Mart\u00edn C, Wu G, Wang Z, Stagge S, J\u00f6nsson LJ. Formation of microbial inhibitors in steam-explosion pretreatment of softwood impregnated with sulfuric acid and sulfur dioxide. Bioresour Technol. 2018;262:242\u201350.","journal-title":"Bioresour Technol"},{"key":"2708_CR29","doi-asserted-by":"publisher","first-page":"9747","DOI":"10.1021\/acs.jafc.5b04803","volume":"63","author":"A Cavka","year":"2015","unstructured":"Cavka A, Stagge S, J\u00f6nsson LJ. Identification of small aliphatic aldehydes in pretreated lignocellulosic feedstocks and evaluation of their inhibitory effects on yeast. J Agric Food Chem. 2015;63:9747\u201354.","journal-title":"J Agric Food Chem"},{"key":"2708_CR30","doi-asserted-by":"publisher","first-page":"10001","DOI":"10.1021\/acssuschemeng.3c01320","volume":"11","author":"M Chen","year":"2023","unstructured":"Chen M, Li Y, Lu F, Luterbacher JS, Ralph J. Lignin hydrogenolysis: phenolic monomers from lignin and associated phenolates across plant clades. ACS Sustain Chem Eng. 2023;11:10001\u201317.","journal-title":"ACS Sustain Chem Eng"},{"key":"2708_CR31","doi-asserted-by":"publisher","first-page":"654","DOI":"10.1007\/s12155-013-9392-6","volume":"7","author":"VD Mitchell","year":"2014","unstructured":"Mitchell VD, Taylor CM, Bauer S. Comprehensive analysis of monomeric phenolics in dilute acid plant hydrolysates. Bioenerg Res. 2014;7:654\u201369.","journal-title":"Bioenerg Res"},{"key":"2708_CR32","doi-asserted-by":"publisher","DOI":"10.1186\/s13568-015-0149-9","volume":"5","author":"S Stagge","year":"2015","unstructured":"Stagge S, Cavka A, J\u00f6nsson LJ. Identification of benzoquinones in pretreated lignocellulosic feedstocks and inhibitory effects on yeast. AMB Expr. 2015;5:62.","journal-title":"AMB Expr"},{"key":"2708_CR33","doi-asserted-by":"publisher","first-page":"617","DOI":"10.1385\/ABAB:84-86:1-9:617","volume":"84\u201386","author":"S Larsson","year":"2000","unstructured":"Larsson S, Quintana-S\u00e1inz A, Reimann A, Nilvebrant N-O, J\u00f6nsson LJ. The influence of lignocellulose-derived aromatic compounds on oxygen-limited growth and ethanolic fermentation by Saccharomyces cerevisiae. Appl Biochem Biotechnol. 2000;84\u201386:617\u201332.","journal-title":"Appl Biochem Biotechnol"},{"key":"2708_CR34","doi-asserted-by":"publisher","first-page":"152","DOI":"10.1016\/S0076-6879(99)99017-1","volume":"299","author":"VL Singleton","year":"1999","unstructured":"Singleton VL, Orthofer R, Lamuela-Ravent\u00f3s RM. Analysis of total phenols and other oxidation substrates and antioxidants by means of Folin-Ciocalteu reagent. Methods Enzymol. 1999;299:152\u201378.","journal-title":"Methods Enzymol"}],"container-title":["Biotechnology for Biofuels and Bioproducts"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s13068-025-02708-4.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s13068-025-02708-4\/fulltext.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s13068-025-02708-4.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,8]],"date-time":"2025-10-08T07:34:50Z","timestamp":1759908890000},"score":1,"resource":{"primary":{"URL":"https:\/\/biotechnologyforbiofuels.biomedcentral.com\/articles\/10.1186\/s13068-025-02708-4"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,10,8]]},"references-count":34,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2025,12]]}},"alternative-id":["2708"],"URL":"https:\/\/doi.org\/10.1186\/s13068-025-02708-4","relation":{},"ISSN":["2731-3654"],"issn-type":[{"value":"2731-3654","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,10,8]]},"assertion":[{"value":"17 July 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"19 September 2025","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"8 October 2025","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"Not applicable.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethics approval and consent to participate"}},{"value":"Not applicable.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent for publication"}},{"value":"LJ is co-inventor of patents in the area biomass conversion. The other authors declare that they have no competing interests.","order":4,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"103"}}