{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T14:01:09Z","timestamp":1790863269542,"version":"4.1.0"},"reference-count":39,"publisher":"MDPI AG","issue":"18","license":[{"start":{"date-parts":[[2026,9,17]],"date-time":"2026-09-17T00:00:00Z","timestamp":1789603200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100004281","name":"NCN OPUS project","doi-asserted-by":"publisher","award":["2011\/01\/B\/NZ1\/00217"],"award-info":[{"award-number":["2011\/01\/B\/NZ1\/00217"]}],"id":[{"id":"10.13039\/501100004281","id-type":"DOI","asserted-by":"publisher"}]},{"name":"University of Wroclaw and Wroclaw Medical University institutional fund","id":[{"id":"https:\/\/ror.org\/01qpw1b93","id-type":"ROR","asserted-by":"crossref"}]},{"id":[{"id":"https:\/\/ror.org\/01qpw1b93","id-type":"ROR","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Int. J. Mol. Sci.","IJMS"],"abstract":"<jats:p>Tocopherols are essential lipid-soluble antioxidants that play important roles in plant protection against oxidative stress and are closely connected with isoprenoid metabolism. To the best of our knowledge, this is the first report of stable constitutive expression of Arabidopsis thaliana VTE2 (AtVTE2) in flax (Linum usitatissimum L.) and its effects on plant metabolism and development. As a dual-purpose oil and fibre crop, flax combines PUFA-rich seeds with industrially important fibre-rich stems, providing a useful system for assessing the broader consequences of antioxidant-pathway engineering. Three independent transgenic lines were characterized by RT-qPCR analysis of genes associated with tocopherol, MEP, MVA, and carotenoid biosynthesis, targeted metabolite profiling, antioxidant-capacity assays, field phenotyping, and biochemical and spectroscopic analyses of mature straw. Three independent transgenic lines were characterized using gene-expression analyses, metabolic profiling, antioxidant assays, phenotypic evaluation, and biochemical analyses of mature stems. Constitutive AtVTE2 expression was associated with \u03b1-tocopherol accumulation and was accompanied by coordinated transcriptional changes in genes associated with the plastidial methylerythritol phosphate and cytosolic mevalonate pathways. These changes were associated with modifications in isoprenoid and phenylpropanoid metabolism, enhanced antioxidant capacity in selected transgenic lines, and altered plant architecture under field conditions. In mature straw, transgenic plants exhibited changes in phytosterol composition, lignin accumulation, and pectin distribution, whereas cellulose content remained unchanged. FT-IR spectra retained the same principal bands but showed differences in the relative intensities of selected cell-wall-associated bands. Collectively, these findings demonstrate that constitutive AtVTE2 expression is associated with tissue-dependent metabolic responses extending beyond vitamin E accumulation and affecting interconnected pathways involved in secondary metabolism, antioxidant capacity, and cell-wall organization.<\/jats:p>","DOI":"10.3390\/ijms27188295","type":"journal-article","created":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T13:18:26Z","timestamp":1790860706000},"page":"8295","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Constitutive Expression of AtVTE2 in Flax Is Associated with Changes in Isoprenoid Metabolism and Stem Cell-Wall Composition"],"prefix":"10.3390","volume":"27","author":[{"given":"Karolina","family":"Hasiewicz-Derkacz","sequence":"first","affiliation":[{"name":"Department of Genetic Biochemistry, Faculty of Biotechnology, University of Wroclaw, Przybyszewskiego 63, 51-148 Wroc\u0142aw, Poland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7338-8654","authenticated-orcid":false,"given":"Iwan","family":"Zalewski","sequence":"additional","affiliation":[{"name":"Department of Dietetics and Bromatology, Wroclaw Medical University, Borowska 211, 50-556 Wroc\u0142aw, Poland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1298-8100","authenticated-orcid":false,"given":"Aleksandra","family":"Boba","sequence":"additional","affiliation":[{"name":"Department of Genetic Biochemistry, Faculty of Biotechnology, University of Wroclaw, Przybyszewskiego 63, 51-148 Wroc\u0142aw, Poland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8044-6835","authenticated-orcid":false,"given":"Lucyna","family":"Dymi\u0144ska","sequence":"additional","affiliation":[{"name":"Department of Bioorganic Chemistry, Faculty of Production Engineering, Wroclaw University of Economics and Business, Komandorska 118\/120, 53-345 Wroc\u0142aw, Poland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dzmitry","family":"Lysh","sequence":"additional","affiliation":[{"name":"Department of Genetic Biochemistry, Faculty of Biotechnology, University of Wroclaw, Przybyszewskiego 63, 51-148 Wroc\u0142aw, Poland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tadeusz","family":"Czuj","sequence":"additional","affiliation":[{"name":"Department of Genetic Biochemistry, Faculty of Biotechnology, University of Wroclaw, Przybyszewskiego 63, 51-148 Wroc\u0142aw, Poland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4620-8953","authenticated-orcid":false,"given":"Anna","family":"Prescha","sequence":"additional","affiliation":[{"name":"Department of Dietetics and Bromatology, Wroclaw Medical University, Borowska 211, 50-556 Wroc\u0142aw, Poland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Katarzyna","family":"Sk\u00f3rkowska-Telichowska","sequence":"additional","affiliation":[{"name":"Medical Faculty, Wroc\u0142aw University of Science and Technology, Grunwaldzki 11, 51-377 Wroc\u0142aw, Poland"},{"name":"University College of Professional Education in Wroclaw, Plac Powsta\u0144c\u00f3w \u015al\u0105skich 1, 53-329 Wroc\u0142aw, Poland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jan","family":"Szopa","sequence":"additional","affiliation":[{"name":"Department of Genetic Biochemistry, Faculty of Biotechnology, University of Wroclaw, Przybyszewskiego 63, 51-148 Wroc\u0142aw, Poland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5073-901X","authenticated-orcid":false,"given":"Anna","family":"Kulma","sequence":"additional","affiliation":[{"name":"Department of Genetic Biochemistry, Faculty of Biotechnology, University of Wroclaw, Przybyszewskiego 63, 51-148 Wroc\u0142aw, Poland"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,9,17]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"200","DOI":"10.1016\/j.plaphy.2018.03.033","article-title":"An altered tocopherol composition in chloroplasts reduces plant resistance to Botrytis cinerea","volume":"127","author":"Cela","year":"2018","journal-title":"Plant Physiol. Biochem."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"743","DOI":"10.1016\/j.jplph.2005.04.022","article-title":"The role of alpha-tocopherol in plant stress tolerance","volume":"162","year":"2005","journal-title":"J. Plant Physiol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"632","DOI":"10.1104\/pp.015222","article-title":"Homogentisate phytyltransferase activity is limiting for tocopherol biosynthesis in Arabidopsis","volume":"131","author":"Collakova","year":"2003","journal-title":"Plant Physiol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"1419","DOI":"10.1105\/tpc.021360","article-title":"Vitamin E is essential for seed longevity and for preventing lipid peroxidation during germination","volume":"16","author":"Sattler","year":"2004","journal-title":"Plant Cell"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"321","DOI":"10.1016\/j.phytochem.2010.12.013","article-title":"Ectopic expression of apple fruit homogentisate phytyltransferase gene (MdHPT1) increases tocopherol in transgenic tomato (Solanum lycopersicum cv. Micro-Tom) leaves and fruits","volume":"72","author":"Seo","year":"2011","journal-title":"Phytochemistry"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"6866","DOI":"10.1073\/pnas.1031755100","article-title":"Crosstalk between cytosolic and plastidial pathways of isoprenoid biosynthesis in Arabidopsis thaliana","volume":"100","author":"Laule","year":"2003","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_7","first-page":"2846","article-title":"Remobilization of Phytol from Chlorophyll Degradation Is Essential for Tocopherol Synthesis and Growth of Arabidopsis","volume":"27","author":"Plohmann","year":"2015","journal-title":"Plant Cell"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"713","DOI":"10.1104\/pp.104.054908","article-title":"Alterations in Tocopherol Cyclase Activity in Transgenic and Mutant Plants of Arabidopsis Affect Tocopherol Content, Tocopherol Composition, and Oxidative Stress","volume":"137","author":"Kanwischer","year":"2005","journal-title":"Plant Physiol."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"369","DOI":"10.1016\/j.plaphy.2020.09.008","article-title":"Tocopherols mutual balance is a key player for maintaining Arabidopsis thaliana growth under salt stress","volume":"156","author":"Potocka","year":"2020","journal-title":"Plant Physiol. Biochem."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"4108","DOI":"10.1111\/ijfs.15039","article-title":"Biosynthesis and profiles of fatty acids, vitamin E and carotenoids during flax (Linum usitatissimum L.) seed capsule maturation","volume":"56","author":"Wang","year":"2021","journal-title":"Int. J. Food Sci. Technol."},{"key":"ref_11","first-page":"353","article-title":"Flax terpenoid pathway as a source of health promoting compounds","volume":"13","author":"Styrczewska","year":"2013","journal-title":"Mini-Rev. Med. Chem."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"100573","DOI":"10.1016\/j.focha.2023.100573","article-title":"Flaxseed (Linum usitatissimum); phytochemistry, pharmacological characteristics and functional food applications","volume":"4","author":"Kauser","year":"2024","journal-title":"Food Chem. Adv."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"862391","DOI":"10.1155\/2015\/862391","article-title":"Flax Fiber Hydrophobic Extract Inhibits Human Skin Cells Inflammation and Causes Remodeling of Extracellular Matrix and Wound Closure Activation","volume":"2015","author":"Styrczewska","year":"2015","journal-title":"Biomed. Res. Int."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1016\/j.indcrop.2015.05.005","article-title":"Linseed, the multipurpose plant","volume":"75","author":"Zuk","year":"2015","journal-title":"Ind. Crops Prod."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"3685","DOI":"10.1021\/jf047987z","article-title":"Pleiotropic effect of phenolic compounds content increases in transgenic flax plant","volume":"53","author":"Amarowicz","year":"2005","journal-title":"J. Agric. Food Chem."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"123","DOI":"10.17221\/104\/2015-CJGPB","article-title":"Transgenic flax\/linseed (Linum usitatissimum L.)\u2014Expectations and reality","volume":"51","author":"Griga","year":"2015","journal-title":"Czech J. Genet. Plant Breed."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"301","DOI":"10.1016\/S1016-8478(23)07343-0","article-title":"Overexpression of Arabidopsis homogentisate phytyltransferase or tocopherol cyclase elevates vitamin E content by increasing gamma-tocopherol level in lettuce (Lactuca sativa L.)","volume":"24","author":"Lee","year":"2007","journal-title":"Mol. Cells"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1016\/j.plaphy.2007.10.009","article-title":"Tocopherol content and isomers\u2019 composition in selected plant species","volume":"46","author":"Kruk","year":"2008","journal-title":"Plant Physiol. Biochem."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"784","DOI":"10.1111\/pbi.12828","article-title":"Improved fruit \u03b1-tocopherol, carotenoid, squalene and phytosterol contents through manipulation of Brassica juncea 3-HYDROXY-3-METHYLGLUTARYL-COA SYNTHASE1 in transgenic tomato","volume":"16","author":"Liao","year":"2018","journal-title":"Plant Biotechnol. J."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1016\/j.pbi.2015.04.001","article-title":"Breaking new ground in the regulation of the early steps of plant isoprenoid biosynthesis","volume":"25","author":"Boronat","year":"2015","journal-title":"Curr. Opin. Plant Biol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"3194","DOI":"10.1105\/tpc.106.049817","article-title":"Manipulation of phytoene levels in tomato fruit: Effects on isoprenoids, plastids, and intermediary metabolism","volume":"19","author":"Fraser","year":"2007","journal-title":"Plant Cell"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"3451","DOI":"10.1105\/tpc.105.037036","article-title":"Vitamin E protects against photoinhibition and photooxidative stress in Arabidopsis thaliana","volume":"17","author":"Havaux","year":"2005","journal-title":"Plant Cell"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"309","DOI":"10.1007\/s00425-007-0618-8","article-title":"Overexpression of CYP710A1 and CYP710A4 in transgenic Arabidopsis plants increases the level of stigmasterol at the expense of sitosterol","volume":"227","author":"Arnqvist","year":"2008","journal-title":"Planta"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"219","DOI":"10.1111\/j.1467-7652.2005.00174.x","article-title":"Enhanced seed phytosterol accumulation through expression of a modified HMG-CoA reductase","volume":"4","author":"Hey","year":"2006","journal-title":"Plant Biotechnol. J."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"707","DOI":"10.1007\/s00425-002-0913-3","article-title":"Co-ordinate regulation of sterol biosynthesis enzyme activity during accumulation of sterols in developing rape and tobacco seed","volume":"216","author":"Harker","year":"2003","journal-title":"Planta"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"92","DOI":"10.1104\/pp.103.032441","article-title":"Engineering plant shikimate pathway for production of tocotrienol and improving herbicide resistance","volume":"134","author":"Rippert","year":"2004","journal-title":"Plant Physiol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"1655","DOI":"10.1021\/jf803537k","article-title":"Antioxidant Assays for Plant and Food Components","volume":"57","author":"Moon","year":"2009","journal-title":"J. Agric. Food Chem."},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Maeda, H., Song, W., Sage, T., and Dellapenna, D. (2014). Role of callose synthases in transfer cell wall development in tocopherol deficient Arabidopsis mutants. Front. Plant Sci., 5.","DOI":"10.3389\/fpls.2014.00046"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"148","DOI":"10.1105\/tpc.106.044495","article-title":"Flavonoid accumulation in Arabidopsis repressed in lignin synthesis affects auxin transport and plant growth","volume":"19","author":"Besseau","year":"2007","journal-title":"Plant Cell"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1007\/s00425-020-03339-9","article-title":"Fusarium oxysporum infection activates the plastidial branch of the terpenoid biosynthesis pathway in flax, leading to increased ABA synthesis","volume":"251","author":"Boba","year":"2020","journal-title":"Planta"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"965","DOI":"10.1007\/s11746-002-0588-1","article-title":"Camelina oil and its unusual cholesterol content","volume":"79","author":"Shukla","year":"2002","journal-title":"J. Am. Oil Chem. Soc."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"1633","DOI":"10.1016\/j.foodchem.2006.04.033","article-title":"Phytosterol content of sea buckthorn (Hippophae rhamnoides L.) seed oil: Extraction and identification","volume":"101","author":"Li","year":"2007","journal-title":"Food Chem."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"1204","DOI":"10.1002\/ejlt.201200009","article-title":"Phytosterols in pumpkin seed oil extracted by organic solvents and supercritical CO2","volume":"114","author":"Hrabovski","year":"2012","journal-title":"Eur. J. Lipid Sci. Technol."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"1917","DOI":"10.1007\/s11032-014-0149-5","article-title":"Crossbreeding of transgenic flax plants overproducing flavonoids and glucosyltransferase results in progeny with improved antifungal and antioxidative properties","volume":"34","author":"Mierziak","year":"2014","journal-title":"Mol. Breed."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"3526392","DOI":"10.1155\/2017\/3526392","article-title":"Optimization of Phenolic Compounds Extraction from Flax Shives and Their Effect on Human Fibroblasts","volume":"2017","author":"Czemplik","year":"2017","journal-title":"Evid. Based Complement. Altern. Med."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"77","DOI":"10.1016\/j.postharvbio.2007.09.017","article-title":"Cell wall modifications in chilling-injured plum fruit (Prunus salicina)","volume":"48","author":"Manganaris","year":"2008","journal-title":"Postharvest Biol. Technol."},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Wojtasik, W., Kulma, A., Dymi\u0144ska, L., Hanuza, J., \u017bebrowski, J., and Szopa, J. (2013). Fibres from flax overproducing \u03b2-1,3-glucanase show increased accumulation of pectin and phenolics and thus higher antioxidant capacity. BMC Biotechnol., 13.","DOI":"10.1186\/1472-6750-13-10"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1002\/jsfa.2740510202","article-title":"Determination of lignin in herbaceous plants by an improved acetyl bromide procedure","volume":"51","author":"Iiyama","year":"1990","journal-title":"J. Sci. Food Agric."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"70","DOI":"10.1006\/abio.1996.0292","article-title":"The ferric reducing ability of plasma (FRAP) as a measure of \u201cantioxidant power\u201d: The FRAP assay","volume":"239","author":"Benzie","year":"1996","journal-title":"Anal. Biochem."}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/18\/8295\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T13:21:08Z","timestamp":1790860868000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/18\/8295"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,17]]},"references-count":39,"journal-issue":{"issue":"18","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["ijms27188295"],"URL":"https:\/\/doi.org\/10.3390\/ijms27188295","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,17]]}}}