{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,25]],"date-time":"2026-08-25T07:19:03Z","timestamp":1787642343054,"version":"build-2736575974"},"reference-count":89,"publisher":"Oxford University Press (OUP)","issue":"9","license":[{"start":{"date-parts":[[2026,4,16]],"date-time":"2026-04-16T00:00:00Z","timestamp":1776297600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,9,1]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Strawberry aroma is a key component of fruit quality, influencing consumer preferences and playing important ecological roles, including plant defense. However, the genetic basis of volatile organic compound (VOC) biosynthesis remains only partially understood, particularly in the wild woodland strawberry Fragaria vesca, which has enormous potential to uncover genetic diversity within the genus for improving the commercial strawberries. Here, we performed Genome-Wide Association Studies (GWAS) across a diverse European collection of F. vesca accessions. We identified multiple novel candidate genes involved in the biosynthesis of diverse volatile esters, lactones, terpenoids, and methyl ketones. Among them, we characterized FvJMT2, a SABATH family methyltransferase, which we found associated with natural variation in benzenoid ester content. Transient expression in Nicotiana benthamiana and strawberry fruit confirmed its role in methyl benzoate biosynthesis, while enzymatic assays demonstrated that FvJMT2 encodes a promiscuous enzyme capable of methylating not only benzoic acid, but also cinnamic, salicylic, and jasmonic acids. Behavioral assays revealed that methyl benzoate at physiologically relevant concentrations significantly reduced the attraction of Drosophila suzukii flies, supporting a dual role of this VOC in both flavor and pest deterrence. Finally, natural variation analyses in wild Fragaria species and F. \u00d7 ananassa cultivars showed that benzenoid esters have been largely lost in modern cultivars but retained in ancient and wild accessions. Altogether, this study provides novel insights into the genetics and ecological relevance of strawberry volatiles and identifies candidate loci and alleles for future studies.<\/jats:p>","DOI":"10.1093\/hr\/uhag148","type":"journal-article","created":{"date-parts":[[2026,4,16]],"date-time":"2026-04-16T15:58:18Z","timestamp":1776355098000},"source":"Crossref","is-referenced-by-count":0,"title":["Uncovering the genetic basis of fruit volatiles in\n                    <i>Fragaria vesca<\/i>\n                    through GWAS reveals FvJMT2 as a methyl benzoate biosynthesis gene with bioactivity against\n                    <i>Drosophila suzukii<\/i>"],"prefix":"10.1093","volume":"13","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-3165-3189","authenticated-orcid":false,"given":"Raquel","family":"Jim\u00e9nez-Mu\u00f1oz","sequence":"first","affiliation":[{"name":"Grupo de Biotecnolog\u00eda en el Desarrollo del Fruto y su Calidad, Instituto de Hortofruticultura Subtropical y Mediterr\u00e1nea La Mayora, Universidad de M\u00e1laga-Consejo Superior de Investigaciones Cient\u00edficas , M\u00e1laga ,","place":["Spain"]},{"name":"Departamento de Fisiolog\u00eda Vegetal, Facultad de Ciencias, Universidad de Granada , Granada,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1418-7964","authenticated-orcid":false,"given":"Mar\u00eda","family":"Urrutia","sequence":"additional","affiliation":[{"name":"Grupo de Biotecnolog\u00eda en el Desarrollo del Fruto y su Calidad, Instituto de Hortofruticultura Subtropical y Mediterr\u00e1nea La Mayora, Universidad de M\u00e1laga-Consejo Superior de Investigaciones Cient\u00edficas , M\u00e1laga ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2244-4967","authenticated-orcid":false,"given":"Victoriano","family":"Meco","sequence":"additional","affiliation":[{"name":"Grupo de Biotecnolog\u00eda en el Desarrollo del Fruto y su Calidad, Instituto de Hortofruticultura Subtropical y Mediterr\u00e1nea La Mayora, Universidad de M\u00e1laga-Consejo Superior de Investigaciones Cient\u00edficas , M\u00e1laga ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4704-2534","authenticated-orcid":false,"given":"Jos\u00e9 L","family":"Rambla","sequence":"additional","affiliation":[{"name":"Instituto de Biolog\u00eda Molecular y Celular de Plantas (IBMCP), Consejo Superior de Investigaciones Cient\u00edficas, Universitat Polit\u00e8cnica de Val\u00e9ncia (CSIC-UPV) , Valencia ,","place":["Spain"]},{"name":"Departamento de Biolog\u00eda, Bioqu\u00edmica y Ciencias Naturales, Universitat Jaume I , Castell\u00f3n ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2817-8121","authenticated-orcid":false,"given":"Tuomas","family":"Toivainen","sequence":"additional","affiliation":[{"name":"Department of Agricultural Sciences, Viikki Plant Science Centre, University of Helsinki , Helsinki ,","place":["Finland"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3411-0761","authenticated-orcid":false,"given":"Meritxell","family":"P\u00e9rez-Hedo","sequence":"additional","affiliation":[{"name":"Instituto Valenciano de Investigaciones Agrarias (IVIA), Centro de Protecci\u00f3n Vegetal y Biotecnolog\u00eda , Moncada, Valencia ,","place":["Spain"]},{"name":"Instituto de Biolog\u00eda Molecular y Celular de Plantas (IBMCP), Consejo Superior de Investigaciones Cient\u00edficas, Universitat Polit\u00e8cnica de Val\u00e9ncia (CSIC-UPV) , Valencia ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6690-7196","authenticated-orcid":false,"given":"Jos\u00e9 F","family":"S\u00e1nchez-Sevilla","sequence":"additional","affiliation":[{"name":"Junta de Andaluc\u00eda, Instituto Andaluz de Investigaci\u00f3n y Formaci\u00f3n Agraria y Pesquera (IFAPA) , M\u00e1laga ,","place":["Spain"]},{"name":"Unidad Asociada de I+D+i IFAPA-CSIC Biotecnolog\u00eda y Mejora en Fresa , M\u00e1laga ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5986-3685","authenticated-orcid":false,"given":"Alberto","family":"Urbaneja","sequence":"additional","affiliation":[{"name":"Instituto Valenciano de Investigaciones Agrarias (IVIA), Centro de Protecci\u00f3n Vegetal y Biotecnolog\u00eda , Moncada, Valencia ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3259-2587","authenticated-orcid":false,"given":"Joaqu\u00edn J","family":"Salas","sequence":"additional","affiliation":[{"name":"Departamento de Bioqu\u00edmica y Biolog\u00eda Molecular de Productos Vegetales, Instituto de la Grasa (CSIC) , Sevilla ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5231-4031","authenticated-orcid":false,"given":"Timo","family":"Hyt\u00f6nen","sequence":"additional","affiliation":[{"name":"Department of Agricultural Sciences, Viikki Plant Science Centre, University of Helsinki , Helsinki ,","place":["Finland"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4266-9581","authenticated-orcid":false,"given":"Antonio","family":"Granell","sequence":"additional","affiliation":[{"name":"Instituto de Biolog\u00eda Molecular y Celular de Plantas (IBMCP), Consejo Superior de Investigaciones Cient\u00edficas, Universitat Polit\u00e8cnica de Val\u00e9ncia (CSIC-UPV) , Valencia ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9305-3096","authenticated-orcid":false,"given":"Carmen","family":"Mart\u00edn-Pizarro","sequence":"additional","affiliation":[{"name":"Grupo de Biotecnolog\u00eda en el Desarrollo del Fruto y su Calidad, Instituto de Hortofruticultura Subtropical y Mediterr\u00e1nea La Mayora, Universidad de M\u00e1laga-Consejo Superior de Investigaciones Cient\u00edficas , M\u00e1laga ,","place":["Spain"]},{"name":"Unidad Asociada de I+D+i IFAPA-CSIC Biotecnolog\u00eda y Mejora en Fresa , M\u00e1laga ,","place":["Spain"]},{"name":"Departamento de Biolog\u00eda Molecular y Bioqu\u00edmica, Facultad de Ciencias, Universidad de M\u00e1laga , M\u00e1laga,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3332-4661","authenticated-orcid":false,"given":"David","family":"Pos\u00e9","sequence":"additional","affiliation":[{"name":"Grupo de Biotecnolog\u00eda en el Desarrollo del Fruto y su Calidad, Instituto de Hortofruticultura Subtropical y Mediterr\u00e1nea La Mayora, Universidad de M\u00e1laga-Consejo Superior de Investigaciones Cient\u00edficas , M\u00e1laga ,","place":["Spain"]},{"name":"Unidad Asociada de I+D+i IFAPA-CSIC Biotecnolog\u00eda y Mejora en Fresa , M\u00e1laga ,","place":["Spain"]},{"name":"Departamento de Biolog\u00eda Molecular y Bioqu\u00edmica, Facultad de Ciencias, Universidad de M\u00e1laga , M\u00e1laga,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,4,16]]},"reference":[{"key":"2026082502262735000_ref57","doi-asserted-by":"crossref","first-page":"1704","DOI":"10.1105\/tpc.014548","article-title":"Structural basis for substrate recognition in the salicylic acid carboxyl methyltransferase family","volume":"15","author":"Zubieta","year":"2003","journal-title":"Plant Cell"},{"key":"2026082502262735000_ref33","doi-asserted-by":"crossref","first-page":"1315","DOI":"10.1016\/j.jplph.2014.06.004","article-title":"Expression of a functional jasmonic acid carboxyl methyltransferase is negatively correlated with strawberry fruit development","volume":"171","author":"Preu\u00df","year":"2014","journal-title":"J Plant Physiol"},{"key":"2026082502262735000_ref32","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1006\/abbi.1999.1255","article-title":"S-Adenosyl-l-methionine:salicylic acid carboxyl methyltransferase, an enzyme involved in floral scent production and plant defense, represents a new class of plant methyltransferases","volume":"367","author":"Ross","year":"1999","journal-title":"Arch Biochem Biophys"},{"key":"2026082502262735000_ref89","doi-asserted-by":"crossref","first-page":"61","DOI":"10.1038\/s41438-019-0142-6","article-title":"Updated annotation of the wild strawberry Fragaria vesca V4 genome","volume":"6","author":"Li","year":"2019","journal-title":"Hortic Res"},{"key":"2026082502262735000_ref64","doi-asserted-by":"crossref","first-page":"530","DOI":"10.1104\/pp.104.041566","article-title":"The role of the jasmonate response in plant susceptibility to diverse pathogens with a range of lifestyles","volume":"135","author":"Thaler","year":"2004","journal-title":"Plant Physiol"},{"key":"2026082502262735000_ref65","first-page":"1","article-title":"Strawberry sweetness and consumer preference are enhanced by specific volatile compounds","volume":"8","author":"Fan","year":"2021","journal-title":"Hortic Res"},{"key":"2026082502262735000_ref1","doi-asserted-by":"crossref","first-page":"1238","DOI":"10.1111\/tpj.15876","article-title":"Towards smart and sustainable development of modern berry cultivars in Europe","volume":"111","author":"Senger","year":"2022","journal-title":"Plant J"},{"key":"2026082502262735000_ref2","doi-asserted-by":"crossref","first-page":"1195","DOI":"10.1016\/j.foodres.2012.11.004","article-title":"Functional properties and quality characteristics of bioactive compounds in berries: biochemistry, biotechnology, and genomics","volume":"54","author":"Jimenez-Garcia","year":"2013","journal-title":"Food Res Int"},{"key":"2026082502262735000_ref3","doi-asserted-by":"crossref","first-page":"11","DOI":"10.3233\/JBR-140067","article-title":"Diversity of metabolite patterns and sensory characters in wild and cultivated strawberries 1","volume":"4","author":"Ulrich","year":"2014","journal-title":"J Berry Res"},{"key":"2026082502262735000_ref4","article-title":"Identification of volatile compounds correlated with consumer acceptability of strawberry preserves: untargeted GC\u2013MS analysis","volume":"378","author":"Dubrow","year":"2022","journal-title":"Food Chem"},{"key":"2026082502262735000_ref5","doi-asserted-by":"crossref","first-page":"1185","DOI":"10.1007\/s10722-006-9009-4","article-title":"Diversity of aroma patterns in wild and cultivated Fragaria accessions","volume":"54","author":"Ulrich","year":"2007","journal-title":"Genet Resour Crop Evol"},{"key":"2026082502262735000_ref6","doi-asserted-by":"crossref","first-page":"615749","DOI":"10.3389\/fpls.2021.615749","article-title":"Genetic analysis of methyl anthranilate, mesifurane, linalool, and other flavor compounds in cultivated strawberry (Fragaria \u00d7 ananassa)","volume":"12","author":"Barbey","year":"2021","journal-title":"Front Plant Sci"},{"key":"2026082502262735000_ref7","doi-asserted-by":"crossref","first-page":"851","DOI":"10.1104\/pp.111.188318","article-title":"Genetic analysis of strawberry fruit aroma and identification of O-methyltransferase FaOMT as the locus controlling natural variation in mesifurane content","volume":"159","author":"Zorrilla-Fontanesi","year":"2012","journal-title":"Plant Physiol"},{"key":"2026082502262735000_ref8","doi-asserted-by":"crossref","first-page":"959155","DOI":"10.3389\/fpls.2022.959155","article-title":"Inheritance of esters and other volatile compounds responsible for the fruity aroma in strawberry","volume":"13","author":"Rey-Serra","year":"2022","journal-title":"Front Plant Sci"},{"key":"2026082502262735000_ref9","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1016\/j.plaphy.2017.10.015","article-title":"Genetic analysis of the wild strawberry (Fragaria vesca) volatile composition","volume":"121","author":"Urrutia","year":"2017","journal-title":"Plant Physiol Biochem"},{"key":"2026082502262735000_ref10","doi-asserted-by":"crossref","DOI":"10.1093\/hr\/uhaf163","article-title":"The role of Fragaria vesca homolog of a (Z)-3:(E)-2-hexenal isomerase in the development of green-leafy fruit aroma","volume":"12","author":"Zhang","year":"2025","journal-title":"Hortic Res"},{"key":"2026082502262735000_ref11","doi-asserted-by":"crossref","first-page":"7382","DOI":"10.3390\/ijms21197382","article-title":"Terpenes and terpenoids in plants: interactions with environment and insects","volume":"21","author":"Boncan","year":"2020","journal-title":"Int J Mol Sci"},{"key":"2026082502262735000_ref12","doi-asserted-by":"crossref","first-page":"5467","DOI":"10.1073\/pnas.0610266104","article-title":"Within-plant signaling by volatiles leads to induction and priming of an indirect plant defense in nature","volume":"104","author":"Heil","year":"2007","journal-title":"Proc Natl Acad 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Agric. 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one of the key aroma compounds in strawberry fruits","volume":"31","author":"Wein","year":"2002","journal-title":"Plant J"},{"key":"2026082502262735000_ref35","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0183384","article-title":"Population genetic analysis of a global collection of Fragaria vesca using microsatellite markers","volume":"12","author":"Hilmarsson","year":"2017","journal-title":"PLoS One"},{"key":"2026082502262735000_ref36","doi-asserted-by":"crossref","first-page":"1201","DOI":"10.1111\/tpj.16404","article-title":"Diversity of the volatilome and the fruit size and shape in European woodland strawberry (Fragaria vesca)","volume":"116","author":"Urrutia","year":"2023","journal-title":"Plant J"},{"key":"2026082502262735000_ref37","doi-asserted-by":"crossref","first-page":"263","DOI":"10.1038\/s42003-026-09539-5","article-title":"The Late Quaternary climate impact on the genome of the woodland strawberry (Fragaria vesca), a perennial 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analysis","volume":"11","author":"Dujak","year":"2024","journal-title":"Hortic Res"},{"key":"2026082502262735000_ref41","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1007\/s11295-018-1238-0","article-title":"Genome-wide association mapping of fruit-quality traits using genotyping-by-sequencing approach in citrus landraces, modern cultivars, and breeding lines in Japan","volume":"14","author":"Imai","year":"2018","journal-title":"Tree Genet Genomes"},{"key":"2026082502262735000_ref42","doi-asserted-by":"crossref","first-page":"1089","DOI":"10.1111\/nph.18416","article-title":"A multi-omics framework reveals strawberry flavor genes and their regulatory elements","volume":"236","author":"Fan","year":"2022","journal-title":"New Phytol"},{"key":"2026082502262735000_ref43","doi-asserted-by":"crossref","DOI":"10.1002\/tpg2.20509","article-title":"Genome-wide association studies in a diverse strawberry collection unveil loci controlling agronomic and fruit quality 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