{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,16]],"date-time":"2026-10-16T12:59:24Z","timestamp":1792155564641,"version":"4.4.0"},"reference-count":49,"publisher":"Wiley","issue":"6","license":[{"start":{"date-parts":[[2026,9,22]],"date-time":"2026-09-22T00:00:00Z","timestamp":1790035200000},"content-version":"vor","delay-in-days":21,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T00:00:00Z","timestamp":1788220800000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/100010661","name":"Horizon 2020 Framework Programme","doi-asserted-by":"publisher","award":["825730"],"award-info":[{"award-number":["825730"]}],"id":[{"id":"10.13039\/100010661","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004543","name":"China Scholarship Council","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100004543","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["The Plant Journal"],"published-print":{"date-parts":[[2026,9]]},"abstract":"<jats:title>SUMMARY<\/jats:title>\n                  <jats:p>\n                    Soapwort (\n                    <jats:italic>Saponaria officinalis<\/jats:italic>\n                    ) synthesizes a vast repertoire of specialized metabolites with bioactive properties. Among those, a specific type of highly glycosylated oleanane\u2010derived triterpenoid saponins has potential applications in the pharmaceutical industry due to their anticancer activity and enhancement of endosomal escape for targeted therapeutics. Saponarioside A and saponarioside B are representative of this group and the major bioactive saponins in soapwort. Recently, the enzymes involved in the biosynthetic pathway of saponarioside B have been characterized. SO1861, another soapwort saponin known for its endosomal escape properties, shares common structural features with these saponins, suggesting saponariosides may represent SO1861 precursors. However, the late\u2010pathway biosynthetic steps converting saponariosides into SO1861 remain unknown. Here, we report the identification and function of the uridine diphosphate (UDP)\u2010glycosyltransferase (UGT), UGT73EU1, and uncover an additional function of the previously characterized UGT73M2, both involved in the late biosynthetic pathway leading to SO1861 and its structural derivatives. We demonstrate their activity and utility through\n                    <jats:italic>in vitro<\/jats:italic>\n                    assays with recombinant UGT proteins produced in a cell\u2010free tobacco system. Furthermore, overexpression of\n                    <jats:italic>UGT73EU1<\/jats:italic>\n                    in hairy root cultures of\n                    <jats:italic>Gypsophila elegans<\/jats:italic>\n                    , a close relative of\n                    <jats:italic>Saponaria<\/jats:italic>\n                    , led to the accumulation of triterpenoid saponins typically not produced by this species. Our work showcases the potential of producing high\u2010value saponins in plants through genetic engineering approaches.\n                  <\/jats:p>","DOI":"10.1111\/tpj.71109","type":"journal-article","created":{"date-parts":[[2026,9,22]],"date-time":"2026-09-22T12:02:08Z","timestamp":1790078528000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Harnessing two\n                    <scp>UDP<\/scp>\n                    \u2010glycosyltransferases from\n                    <i>Saponaria officinalis<\/i>\n                    toward heterologous production of endosomal escape enhancer saponins in\n                    <i>Gypsophila elegans<\/i>\n                    hairy roots"],"prefix":"10.1111","volume":"127","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-0050-3620","authenticated-orcid":false,"given":"Tongtong","family":"Qu","sequence":"first","affiliation":[{"name":"Department of Plant Biotechnology and Bioinformatics Ghent University  9052 Ghent Belgium"},{"name":"VIB Center for Plant Systems Biology  9052 Ghent Belgium"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5739-618X","authenticated-orcid":false,"given":"Helga","family":"Schinkel","sequence":"additional","affiliation":[{"name":"Fraunhofer Institute for Molecular Biology and Applied Ecology IME  Forckenbeckstrasse 6 52074 Aachen Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9366-4727","authenticated-orcid":false,"given":"Tessa","family":"Moses","sequence":"additional","affiliation":[{"name":"EdinOmics, RRID:SCR_021838 University of Edinburgh  Max Born Crescent Edinburgh UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4866-2359","authenticated-orcid":false,"given":"Kaylan","family":"Reddy","sequence":"additional","affiliation":[{"name":"Department of Plant Biotechnology and Bioinformatics Ghent University  9052 Ghent Belgium"},{"name":"VIB Center for Plant Systems Biology  9052 Ghent Belgium"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0003-9871-3913","authenticated-orcid":false,"given":"Ren\u00e9","family":"de Vaumas","sequence":"additional","affiliation":[{"name":"Extrasynthese  Impasse Jacquard 69730 Genay France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1896-4575","authenticated-orcid":false,"given":"Stefan","family":"Schillberg","sequence":"additional","affiliation":[{"name":"Fraunhofer Institute for Molecular Biology and Applied Ecology IME  Forckenbeckstrasse 6 52074 Aachen Germany"},{"name":"Institute for Molecular Biotechnology RWTH Aachen University  Worringerweg 1 52074 Aachen Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1599-551X","authenticated-orcid":false,"given":"Alain","family":"Goossens","sequence":"additional","affiliation":[{"name":"Department of Plant Biotechnology and Bioinformatics Ghent University  9052 Ghent Belgium"},{"name":"VIB Center for Plant Systems Biology  9052 Ghent Belgium"},{"name":"Department of Botany and Zoology Stellenbosch University  Stellenbosch South Africa"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1598-8950","authenticated-orcid":false,"given":"Elia","family":"Lacchini","sequence":"additional","affiliation":[{"name":"Department of Plant Biotechnology and Bioinformatics Ghent University  9052 Ghent Belgium"},{"name":"VIB Center for Plant Systems Biology  9052 Ghent Belgium"},{"name":"Department of Plant Molecular Biology University of Lausanne  1015 Lausanne Switzerland"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,9,22]]},"reference":[{"key":"e_1_2_10_2_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.plaphy.2023.107847"},{"key":"e_1_2_10_3_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-1-0716-2241-4_8"},{"key":"e_1_2_10_4_1","doi-asserted-by":"publisher","DOI":"10.1002\/bit.25502"},{"key":"e_1_2_10_5_1","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2019.01523"},{"key":"e_1_2_10_6_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-020-19399-0"},{"key":"e_1_2_10_7_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.biotechadv.2023.108214"},{"key":"e_1_2_10_8_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.pbi.2025.102701"},{"key":"e_1_2_10_9_1","doi-asserted-by":"publisher","DOI":"10.1105\/tpc.114.133827"},{"key":"e_1_2_10_10_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-023-38039-x"},{"key":"e_1_2_10_11_1","doi-asserted-by":"publisher","DOI":"10.1016\/0031-9422(91)85281-4"},{"key":"e_1_2_10_12_1","article-title":"Glycosylated 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