{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T01:10:19Z","timestamp":1787015419313,"version":"build-2736575974"},"reference-count":100,"publisher":"Wiley","issue":"5","license":[{"start":{"date-parts":[[2025,8,31]],"date-time":"2025-08-31T00:00:00Z","timestamp":1756598400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["P21729"],"award-info":[{"award-number":["P21729"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["21H01202"],"award-info":[{"award-number":["21H01202"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["J Eukaryotic Microbiology"],"published-print":{"date-parts":[[2025,9]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>\n                    Benthic Foraminifera exhibit diverse adaptations to low oxygen (O\n                    <jats:sub>2<\/jats:sub>\n                    ) environments, including denitrification, a rare trait among eukaryotes. Denitrifying species store intracellular nitrate (NO\n                    <jats:sub>3<\/jats:sub>\n                    <jats:sup>\u2212<\/jats:sup>\n                    ), possibly within vacuoles, and contribute significantly to the global marine nitrogen (N) cycle. Additionally, widespread phosphate (PO\n                    <jats:sub>4<\/jats:sub>\n                    <jats:sup>3\u2212<\/jats:sup>\n                    ) accumulation suggests a role in supporting metabolism under O\n                    <jats:sub>2<\/jats:sub>\n                    \u2010depleted conditions. However, the organelles storing NO\n                    <jats:sub>3<\/jats:sub>\n                    <jats:sup>\u2212<\/jats:sup>\n                    and PO\n                    <jats:sub>4<\/jats:sub>\n                    <jats:sup>3\u2212<\/jats:sup>\n                    remain unknown, limiting the mechanistic understanding of these alternative metabolic pathways. To investigate the intracellular NO\n                    <jats:sub>3<\/jats:sub>\n                    <jats:sup>\u2212<\/jats:sup>\n                    and PO\n                    <jats:sub>4<\/jats:sub>\n                    <jats:sup>3\u2212<\/jats:sup>\n                    localization in the benthic foraminifera\n                    <jats:italic>Bolivina spissa<\/jats:italic>\n                    , experimental incubations under varying O\n                    <jats:sub>2<\/jats:sub>\n                    and NO\n                    <jats:sub>3<\/jats:sub>\n                    <jats:sup>\u2212<\/jats:sup>\n                    conditions followed by cryogenic fixation and scanning electron microscopy\u2010energy dispersive spectroscopy (SEM\u2010EDS) analyses were carried out. Most vacuoles were enriched in N relative to the surrounding cytoplasm, likely representing the intracellular NO\n                    <jats:sub>3<\/jats:sub>\n                    <jats:sup>\u2212<\/jats:sup>\n                    reservoir. The elemental mapping also confirmed phosphorus (P) enrichment in organelles resembling acidocalcisomes, likely as PO\n                    <jats:sub>4<\/jats:sub>\n                    <jats:sup>3\u2212<\/jats:sup>\n                    , which may serve as a readily available energy source used over NO\n                    <jats:sub>3<\/jats:sub>\n                    <jats:sup>\u2212<\/jats:sup>\n                    storage during the transition between aerobic and anaerobic respiration. Additionally, barium\u2010rich vacuoles of unknown function(s) display a unique spatial distribution. This study emphasizes the effectiveness of cryogenic techniques in elucidating metabolic processes in foraminifers and other large and\/or testate unicellular organisms, particularly for studying soluble compounds that have rarely been investigated.\n                  <\/jats:p>","DOI":"10.1111\/jeu.70044","type":"journal-article","created":{"date-parts":[[2025,9,1]],"date-time":"2025-09-01T03:55:39Z","timestamp":1756698939000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Different Elemental Compositions and Potential Functions of Vacuoles in\n                    <i>Bolivina spissa<\/i>\n                    (Foraminifera, Rhizaria) Based on Cryo\u2010SEM\u2010EDS Analyses"],"prefix":"10.1111","volume":"72","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-3618-836X","authenticated-orcid":false,"given":"Julien","family":"Richirt","sequence":"first","affiliation":[{"name":"SUGAR, X\u2010Star Japan Agency for Marine\u2010Earth Science and Technology (JAMSTEC)  Yokosuka Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5417-5955","authenticated-orcid":false,"given":"Satoshi","family":"Okada","sequence":"additional","affiliation":[{"name":"SUGAR, X\u2010Star Japan Agency for Marine\u2010Earth Science and Technology (JAMSTEC)  Yokosuka Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8941-6990","authenticated-orcid":false,"given":"Yoshiyuki","family":"Ishitani","sequence":"additional","affiliation":[{"name":"SUGAR, X\u2010Star Japan Agency for Marine\u2010Earth Science and Technology (JAMSTEC)  Yokosuka Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2782-8997","authenticated-orcid":false,"given":"Nicolaas","family":"Glock","sequence":"additional","affiliation":[{"name":"Institute for Geology University of Hamburg  Hamburg Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8461-5608","authenticated-orcid":false,"given":"Katsuyuki","family":"Uematsu","sequence":"additional","affiliation":[{"name":"Marine Works Japan Ltd.  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