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Using this approach, we interrogated the exoproteome of the anaerobic cellulose-degrading bacterium\n                  <jats:italic>Clostridium thermocellum<\/jats:italic>\n                  as it carried out biomass degradation. Mass spectrometry indicated a broad exoproteome composition, including cellulose degrading machinery CelA and CipA. A focus on large exoproteome assemblies revealed abundant protein filaments and pleomorphic vesicular structures. Analysis of the most abundant protein filaments yielded an \u223c4 \u25a1 resolution native structure that, aided by mass spectrometry,\n                  <jats:italic>de novo<\/jats:italic>\n                  modeling, and structural searching, was found to be the aldehyde-alcohol dehydrogenase (AdhE) spirosome. AdhE contained both NAD\n                  <jats:sup>+<\/jats:sup>\n                  and Fe in their expected binding sites and biochemical and structural analyses of enriched spirosome preparations indicated they were functional. Altered NADH solution concentrations triggered conformational changes in the exoproteomic spirosomes, and the constituent AdhE remained capable of ethanol production. Although the basis for functional extracellular spirosome accumulation in live anaerobic\n                  <jats:italic>C. thermocellum<\/jats:italic>\n                  cultures remains unclear, their abundance in crude exoproteomes suggests their presence could influence biomass fueled\n                  <jats:italic>C. thermocellum<\/jats:italic>\n                  growth.\n                <\/jats:p>","DOI":"10.64898\/2026.05.21.726627","type":"posted-content","created":{"date-parts":[[2026,5,23]],"date-time":"2026-05-23T05:00:13Z","timestamp":1779512413000},"source":"Crossref","is-referenced-by-count":0,"title":["Visual exoproteomics of\n                  <i>Clostridium thermocellum<\/i>\n                  during anaerobic biomass-degradation identifies functional spirosomes"],"prefix":"10.64898","author":[{"given":"Matthew P.","family":"Agdanowski","sequence":"first","affiliation":[{"name":"Department of Chemistry and Biochemistry; UCLA-DOE Institute for Genomics and Proteomics; University of California, Los Angeles (UCLA), Los Angeles, CA 90095, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Matthew J.","family":"Kensil","sequence":"additional","affiliation":[{"name":"Department of Chemistry and Biochemistry; UCLA-DOE Institute for Genomics and Proteomics; University of California, Los Angeles (UCLA), Los Angeles, CA 90095, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Trevor H.","family":"Moser","sequence":"additional","affiliation":[{"name":"Environmental Molecular Sciences Laboratory (EMSL); Pacific Northwest National Laboratory, Richland, WA 99354, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ethan","family":"Humm","sequence":"additional","affiliation":[{"name":"Department of Microbiology Immunology and Molecular Genetics; University of California, Los Angeles (UCLA), Los Angeles, CA 90095, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yaneli I.","family":"Guandique","sequence":"additional","affiliation":[{"name":"Department of Chemistry and Biochemistry; UCLA-DOE Institute for Genomics and Proteomics; University of California, Los Angeles (UCLA), Los Angeles, CA 90095, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kayleigh","family":"Mason-Chalmers","sequence":"additional","affiliation":[{"name":"Department of Chemistry and Biochemistry; UCLA-DOE Institute for Genomics and Proteomics; University of California, Los Angeles (UCLA), Los Angeles, CA 90095, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tracy","family":"al-Set","sequence":"additional","affiliation":[{"name":"Department of Chemistry and Biochemistry; UCLA-DOE Institute for Genomics and Proteomics; University of California, Los Angeles (UCLA), Los Angeles, CA 90095, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rachel R.","family":"Ogorzalek Loo","sequence":"additional","affiliation":[{"name":"Department of Chemistry and Biochemistry; UCLA-DOE Institute for Genomics and Proteomics; University of California, Los Angeles (UCLA), Los Angeles, CA 90095, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"James E.","family":"Evans","sequence":"additional","affiliation":[{"name":"Environmental Molecular Sciences Laboratory (EMSL); Pacific Northwest National Laboratory, Richland, WA 99354, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Robert P.","family":"Gunsalus","sequence":"additional","affiliation":[{"name":"Department of Microbiology Immunology and Molecular Genetics; University of California, Los Angeles (UCLA), Los Angeles, CA 90095, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9989-1437","authenticated-orcid":false,"given":"Joseph A.","family":"Loo","sequence":"additional","affiliation":[{"name":"Department of Chemistry and Biochemistry; UCLA-DOE Institute for Genomics and Proteomics; University of California, Los Angeles (UCLA), Los Angeles, CA 90095, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0248-4964","authenticated-orcid":false,"given":"Jose A.","family":"Rodriguez","sequence":"additional","affiliation":[{"name":"Department of Chemistry and Biochemistry; UCLA-DOE Institute for Genomics and Proteomics; University of California, Los Angeles (UCLA), Los Angeles, CA 90095, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"54368","reference":[{"key":"2026052609200928000_2026.05.21.726627v1.1","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1016\/bs.aambs.2020.07.004","article-title":"Clostridium thermocellum: A microbial platform for high-value chemical production from lignocellulose","volume":"113","year":"2020","journal-title":"Adv. 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