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However, lignin recalcitrance limits the enzymatic conversion of wood into fermentable sugars, posing a major challenge for biomass deconstruction. To address this problem, we aimed at incorporating the coumarin scopoletin into the lignin polymer of poplar (\n                    <jats:italic>\n                      <jats:styled-content style=\"fixed-case\">Populus tremula<\/jats:styled-content>\n                      \u00d7\n                      <jats:styled-content style=\"fixed-case\">P<\/jats:styled-content>\n                    <\/jats:italic>\n                    <jats:styled-content style=\"fixed-case\">\n                      <jats:italic>. alba<\/jats:italic>\n                    <\/jats:styled-content>\n                    ) by expressing\n                    <jats:italic>FERULOYL\u2010CoA 6\u2032\u2010HYDROXYLASE 1<\/jats:italic>\n                    (\n                    <jats:italic>F6\u2032H1<\/jats:italic>\n                    ) and\n                    <jats:italic>COUMARIN SYNTHASE<\/jats:italic>\n                    (\n                    <jats:italic>COSY<\/jats:italic>\n                    ) in lignifying cells. Three constructs were evaluated: two bicistronic constructs, SCOP1 (\n                    <jats:italic>COSY<\/jats:italic>\n                    followed by\n                    <jats:italic>F6\u2032H1<\/jats:italic>\n                    ) and SCOP2 (\n                    <jats:italic>F6\u2032H1<\/jats:italic>\n                    followed by\n                    <jats:italic>COSY<\/jats:italic>\n                    ), and one monocistronic, SCOP3 (only\n                    <jats:italic>F6\u2032H1<\/jats:italic>\n                    ). SCOP1 poplars produced most free scopoletin without altering overall lignin, cellulose or hemicellulose content. SCOP2 poplars were overall less efficient in scopoletin production and most of these lines showed a severe biomass yield penalty, whereas SCOP3 caused plant lethality. NMR and metabolic analyses confirmed that scopoletin cross\u2010coupled with G and S monomers during lignification in SCOP1 lines. In addition to scopoletin, the detection of benzodioxane structures revealed the incorporation of dihydroxycoumarins. Overall coumarin incorporation in lignin amounted up to 2.3%. After alkaline pretreatment, wood from greenhouse\u2010grown SCOP1 poplars released up to 29% more glucose compared to the wild type upon limited saccharification. Field\u2010testing of three SCOP1 lines showed a 6 to 11% increase in saccharification efficiency, with the line containing the lowest scopoletin levels maintaining normal growth. These results demonstrate that engineering lignin composition in poplar can improve saccharification, and emphasize the importance of construct design, translational research and field validation.\n                  <\/jats:p>","DOI":"10.1111\/pbi.70739","type":"journal-article","created":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T06:51:53Z","timestamp":1787035913000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Simultaneous Overexpression of\n                    <i>FERULOYL\u2010CoA 6\u2032\u2010HYDROXYLASE 1<\/i>\n                    and\n                    <i>COUMARIN SYNTHASE<\/i>\n                    Leads to Coumarin\u2010Enriched Lignin and Improved Saccharification in Greenhouse\u2010 and Field\u2010Grown Poplar"],"prefix":"10.1111","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-2636-0261","authenticated-orcid":false,"given":"Nette","family":"De Ridder","sequence":"first","affiliation":[{"name":"Department of Plant Biotechnology and Bioinformatics Ghent University  Ghent Belgium"},{"name":"Center for Plant Systems Biology VIB  Ghent Belgium"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7101-734X","authenticated-orcid":false,"given":"Lennart","family":"Hoengenaert","sequence":"additional","affiliation":[{"name":"Department of Plant Biotechnology and Bioinformatics Ghent University  Ghent Belgium"},{"name":"Center for Plant Systems Biology VIB  Ghent Belgium"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jan","family":"Van Doorsselaere","sequence":"additional","affiliation":[{"name":"VIVES  Roeselare Belgium"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wim","family":"De Clercq","sequence":"additional","affiliation":[{"name":"Research Institute for Nature and Forest (INBO)  Geraardsbergen Belgium"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Geert","family":"Goeminne","sequence":"additional","affiliation":[{"name":"Department of Plant Biotechnology and Bioinformatics Ghent University  Ghent Belgium"},{"name":"Center for Plant Systems Biology VIB  Ghent Belgium"},{"name":"VIB Metabolomics Core  Ghent Belgium"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sarah","family":"Liu","sequence":"additional","affiliation":[{"name":"Great Lakes Bioenergy Research Center and the Wisconsin Energy Institute University of Wisconsin\u2010Madison  Madison Wisconsin USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8230-9129","authenticated-orcid":false,"given":"Fachuang","family":"Lu","sequence":"additional","affiliation":[{"name":"Great Lakes Bioenergy Research Center and the Wisconsin Energy Institute University of Wisconsin\u2010Madison  Madison Wisconsin USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6093-4521","authenticated-orcid":false,"given":"John","family":"Ralph","sequence":"additional","affiliation":[{"name":"Great Lakes Bioenergy Research Center and the Wisconsin Energy Institute University of Wisconsin\u2010Madison  Madison Wisconsin USA"},{"name":"Department of Biochemistry University of Wisconsin  Madison Wisconsin USA"},{"name":"New Zealand Institute for Bioeconomy Science (BSI), BSI Scion  Rotorua New Zealand"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5848-3138","authenticated-orcid":false,"given":"Ruben","family":"Vanholme","sequence":"additional","affiliation":[{"name":"Department of Plant Biotechnology and Bioinformatics Ghent University  Ghent Belgium"},{"name":"Center for Plant Systems Biology VIB  Ghent Belgium"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1495-510X","authenticated-orcid":false,"given":"Wout","family":"Boerjan","sequence":"additional","affiliation":[{"name":"Department of Plant Biotechnology and Bioinformatics Ghent University  Ghent Belgium"},{"name":"Center for Plant Systems Biology VIB  Ghent Belgium"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,8,17]]},"reference":[{"key":"e_1_2_11_2_1","doi-asserted-by":"publisher","DOI":"10.3389\/fenrg.2018.00141"},{"key":"e_1_2_11_3_1","doi-asserted-by":"publisher","DOI":"10.1111\/pbi.14144"},{"key":"e_1_2_11_4_1","doi-asserted-by":"publisher","DOI":"10.1007\/s41061-018-0212-5"},{"key":"e_1_2_11_5_1","first-page":"1","article-title":"Non\u2010Conventional Lignans: Coumarinolignans, Flavonolignans, and Stilbenolignans","volume":"93","author":"Begum S. 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