{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T18:29:11Z","timestamp":1787077751527,"version":"build-2736575974"},"reference-count":54,"publisher":"National Academy of Sciences","issue":"34","license":[{"start":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T00:00:00Z","timestamp":1787011200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"content-domain":{"domain":["www.pnas.org"],"crossmark-restriction":true},"short-container-title":["Proc. Natl. Acad. Sci. U.S.A."],"published-print":{"date-parts":[[2026,8,25]]},"abstract":"<jats:p>\n                    Extensive selection for yield and disease resistance during tomato crop improvement has led to flavor loss in modern commercial tomatoes in comparison with heirloom varieties. To investigate the chemical and genetic architecture of tomato flavor through both domestication and improvement, we analyzed flavor-related chemicals across 558 globally collected accessions comprising wild relatives, semidomesticated and domesticated populations, including uncharacterized Latin American accessions. Key flavor volatiles exhibit major differences across accessions. A genome-wide association study was used to detect associations between genetic loci and flavor-related chemical contents, including sugars, acids, and volatiles. Multiple genetic loci linked to known genes encoding flavor metabolism enzymes, as well as many new loci, were identified. Among the newly identified loci, a gene encoding a previously uncharacterized lipase (\n                    <jats:italic toggle=\"yes\">Sl-LIP100<\/jats:italic>\n                    ) was experimentally proven to have an important role in synthesis of lipid-derived flavor volatiles. This enzyme is responsible for synthesis of several important five- and six-carbon flavor volatiles. In sum, this study provides chemical and genetic insights into the evolution of tomato flavor during domestication and subsequent improvement, identifying the complexity of genetic control of fruit flavor chemicals as well as a number of new alleles that can be used to improve the contents of flavor-linked chemicals.\n                  <\/jats:p>","DOI":"10.1073\/pnas.2610136123","type":"journal-article","created":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T17:59:27Z","timestamp":1787075967000},"update-policy":"https:\/\/doi.org\/10.1073\/pnas.cm10313","source":"Crossref","is-referenced-by-count":0,"title":["The genetic architecture of tomato flavor variation through crop domestication and improvement"],"prefix":"10.1073","volume":"123","author":[{"given":"Xiang","family":"Li","sequence":"first","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0327f3359","id-type":"ROR","asserted-by":"publisher"}],"name":"School of Horticulture, Anhui Agricultural University","place":["Hefei, People\u2019s Republic of China"]},{"id":[{"id":"https:\/\/ror.org\/0327f3359","id-type":"ROR","asserted-by":"publisher"}],"name":"State Key Laboratory for Tea Plant Germplasm Innovation and Resource Utilization, Anhui Agricultural University","place":["Hefei, People\u2019s Republic of China"]},{"id":[{"id":"https:\/\/ror.org\/02y3ad647","id-type":"ROR","asserted-by":"publisher"}],"name":"Horticultural Sciences, University of Florida","place":["Gainesville"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Denise M.","family":"Tieman","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0327f3359","id-type":"ROR","asserted-by":"publisher"}],"name":"School of Horticulture, Anhui Agricultural University","place":["Hefei, People\u2019s Republic of China"]},{"id":[{"id":"https:\/\/ror.org\/02y3ad647","id-type":"ROR","asserted-by":"publisher"}],"name":"Horticultural Sciences, University of Florida","place":["Gainesville"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yue","family":"Huang","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0327f3359","id-type":"ROR","asserted-by":"publisher"}],"name":"School of Horticulture, Anhui Agricultural University","place":["Hefei, People\u2019s Republic of China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Luis Felipe V.","family":"Ferr\u00e3o","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/02y3ad647","id-type":"ROR","asserted-by":"publisher"}],"name":"Horticultural Sciences, University of Florida","place":["Gainesville"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Manoj","family":"Sapkota","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/02k3smh20","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Horticulture, University of Kentucky","place":["Lexington"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhuoliang","family":"Lang","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0327f3359","id-type":"ROR","asserted-by":"publisher"}],"name":"School of Horticulture, Anhui Agricultural University","place":["Hefei, People\u2019s Republic of China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6186-7164","authenticated-orcid":false,"given":"Anastasiya","family":"Kuhalskaya","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/01fbde567","id-type":"ROR","asserted-by":"publisher"}],"name":"Max Planck Institute of Molecular Plant Physiology","place":["Potsdam-Golm, Germany"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ronglin","family":"Hao","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0327f3359","id-type":"ROR","asserted-by":"publisher"}],"name":"School of Horticulture, Anhui Agricultural University","place":["Hefei, People\u2019s Republic of China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Pu","family":"Liu","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0327f3359","id-type":"ROR","asserted-by":"publisher"}],"name":"School of Horticulture, Anhui Agricultural University","place":["Hefei, People\u2019s Republic of China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jing","family":"Chen","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0327f3359","id-type":"ROR","asserted-by":"publisher"}],"name":"State Key Laboratory for Tea Plant Germplasm Innovation and Resource Utilization, Anhui Agricultural University","place":["Hefei, People\u2019s Republic of China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2067-5235","authenticated-orcid":false,"given":"Saleh","family":"Alseekh","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/01fbde567","id-type":"ROR","asserted-by":"publisher"}],"name":"Max Planck Institute of Molecular Plant Physiology","place":["Potsdam-Golm, Germany"]},{"id":[{"id":"https:\/\/ror.org\/0020pnp42","id-type":"ROR","asserted-by":"publisher"}],"name":"Center of Plant Systems Biology and Biotechnology","place":["Plovdiv, Bulgaria"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"suffix":"Jr.","given":"Marcio F. R.","family":"Resende","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/02y3ad647","id-type":"ROR","asserted-by":"publisher"}],"name":"Horticultural Sciences, University of Florida","place":["Gainesville"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Esther","family":"van der Knaap","sequence":"additional","affiliation":[{"name":"Institute of Plant Breeding, Genetics & Genomics, University of Georgia","place":["Athens"]},{"name":"Department of Horticulture, University of Georgia","place":["Athens"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xueren","family":"Yin","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/0327f3359","id-type":"ROR","asserted-by":"publisher"}],"name":"School of Horticulture, Anhui Agricultural University","place":["Hefei, People\u2019s Republic of China"]},{"id":[{"id":"https:\/\/ror.org\/0327f3359","id-type":"ROR","asserted-by":"publisher"}],"name":"State Key Laboratory for Tea Plant Germplasm Innovation and Resource Utilization, Anhui Agricultural University","place":["Hefei, People\u2019s Republic of China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3823-4518","authenticated-orcid":false,"given":"Harry J.","family":"Klee","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/02y3ad647","id-type":"ROR","asserted-by":"publisher"}],"name":"Horticultural Sciences, University of Florida","place":["Gainesville"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"341","published-online":{"date-parts":[[2026,8,18]]},"reference":[{"key":"e_1_3_4_1_2","doi-asserted-by":"publisher","DOI":"10.1126\/science.aal1556"},{"key":"e_1_3_4_2_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41576-018-0002-5"},{"key":"e_1_3_4_3_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.0602469103"},{"key":"e_1_3_4_4_2","doi-asserted-by":"publisher","DOI":"10.1111\/tpj.16310"},{"key":"e_1_3_4_5_2","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-313X.2008.03446.x"},{"key":"e_1_3_4_6_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1613910113"},{"key":"e_1_3_4_7_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2115865119"},{"key":"e_1_3_4_8_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-024-08186-2"},{"key":"e_1_3_4_9_2","first-page":"416","article-title":"Quantifying the source-sink balance and carbohydrate content in three tomato cultivars","volume":"6","author":"Li T.","year":"2015","unstructured":"T. Li, E. Heuvelink, L. F. M. Marcelis, Quantifying the source-sink balance and carbohydrate content in three tomato cultivars. Front. Plant Sci. 6, 416 (2015).","journal-title":"Front. Plant Sci."},{"key":"e_1_3_4_10_2","unstructured":"L. Bartoshuk T. A. Colquhoun C. A. Sims \u201cSweetness enhancing volatile compositions and methods of increasing perceived sweetness of a comestible.\u201d US Patent 11744273 (2023)."},{"key":"e_1_3_4_11_2","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0088446"},{"key":"e_1_3_4_12_2","doi-asserted-by":"publisher","DOI":"10.1104\/pp.104.041608"},{"key":"e_1_3_4_13_2","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/ert382"},{"key":"e_1_3_4_14_2","doi-asserted-by":"publisher","DOI":"10.1146\/annurev-arplant-050718-100353"},{"key":"e_1_3_4_15_2","doi-asserted-by":"publisher","DOI":"10.1104\/pp.19.00059"},{"key":"e_1_3_4_16_2","doi-asserted-by":"publisher","DOI":"10.1002\/jsfa.4076"},{"key":"e_1_3_4_17_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2020.05.021"},{"key":"e_1_3_4_18_2","doi-asserted-by":"publisher","DOI":"10.1105\/tpc.113.114231"},{"key":"e_1_3_4_19_2","doi-asserted-by":"publisher","DOI":"10.1093\/plphys\/kiac509"},{"key":"e_1_3_4_20_2","doi-asserted-by":"publisher","DOI":"10.1038\/ng.3117"},{"key":"e_1_3_4_21_2","doi-asserted-by":"publisher","DOI":"10.1093\/molbev\/msz297"},{"key":"e_1_3_4_22_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41588-019-0410-2"},{"key":"e_1_3_4_23_2","doi-asserted-by":"publisher","DOI":"10.1038\/ng.3596"},{"key":"e_1_3_4_24_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41588-019-0522-8"},{"key":"e_1_3_4_25_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2206076119"},{"key":"e_1_3_4_26_2","doi-asserted-by":"publisher","DOI":"10.1186\/s13059-023-02945-6"},{"key":"e_1_3_4_27_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41477-024-01873-y"},{"key":"e_1_3_4_28_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41588-025-02135-z"},{"key":"e_1_3_4_29_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41588-025-02401-0"},{"key":"e_1_3_4_30_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-019-09462-w"},{"key":"e_1_3_4_31_2","doi-asserted-by":"publisher","DOI":"10.1093\/plphys\/kiab230"},{"key":"e_1_3_4_32_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41588-023-01340-y"},{"key":"e_1_3_4_33_2","doi-asserted-by":"publisher","DOI":"10.1111\/tpj.16268"},{"key":"e_1_3_4_34_2","doi-asserted-by":"publisher","DOI":"10.1093\/pnasnexus\/pgaf401"},{"key":"e_1_3_4_35_2","doi-asserted-by":"publisher","DOI":"10.1105\/tpc.17.00211"},{"key":"e_1_3_4_36_2","first-page":"20160569","article-title":"Genetics of complex traits: Prediction of phenotype, identification of causal polymorphisms and genetic architecture","volume":"283","author":"Goddard M. E.","year":"2016","unstructured":"M. E. Goddard, K. E. Kemper, I. M. MacLeod, A. J. Chamberlain, B. J. Hayes, Genetics of complex traits: Prediction of phenotype, identification of causal polymorphisms and genetic architecture. Proc. Biol. Sci. 283, 20160569 (2016).","journal-title":"Proc. Biol. Sci."},{"key":"e_1_3_4_37_2","doi-asserted-by":"publisher","DOI":"10.1007\/s13253-017-0277-6"},{"key":"e_1_3_4_38_2","doi-asserted-by":"publisher","DOI":"10.1093\/genetics\/iyae164"},{"key":"e_1_3_4_39_2","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pgen.1010751"},{"key":"e_1_3_4_40_2","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1216515109"},{"key":"e_1_3_4_41_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.molp.2014.11.007"},{"key":"e_1_3_4_42_2","doi-asserted-by":"publisher","DOI":"10.1111\/tpj.14795"},{"key":"e_1_3_4_43_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.hpj.2022.09.015"},{"key":"e_1_3_4_44_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.febslet.2004.05.064"},{"key":"e_1_3_4_45_2","doi-asserted-by":"publisher","DOI":"10.1038\/nature11119"},{"key":"e_1_3_4_46_2","doi-asserted-by":"publisher","DOI":"10.3390\/agronomy11020349"},{"key":"e_1_3_4_47_2","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-022-33648-4"},{"key":"e_1_3_4_48_2","doi-asserted-by":"publisher","DOI":"10.15252\/embj.2019103256"},{"key":"e_1_3_4_49_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.molp.2018.06.003"},{"key":"e_1_3_4_50_2","doi-asserted-by":"publisher","DOI":"10.1111\/tpj.14951"},{"key":"e_1_3_4_51_2","doi-asserted-by":"publisher","DOI":"10.1146\/annurev-arplant-050718-100202"},{"key":"e_1_3_4_52_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.plipres.2022.101163"},{"key":"e_1_3_4_53_2","unstructured":"D. Tieman The genome re-sequencing dataset for tomato flavor study Raw sequence reads. National Center for Biotechnology Information BioProject. https:\/\/www.ncbi.nlm.nih.gov\/bioproject\/?term=PRJNA353161. Deposited 11 November 2016."},{"key":"e_1_3_4_54_2","unstructured":"100 Tomato Genome Sequencing Consortium . Umbrella Study linking 5 studies Exploring genetic variation in the tomato (Solanum section Lycopersicon) clade by whole-genome sequencing. National Center for Biotechnology Information BioProject. https:\/\/www.ncbi.nlm.nih.gov\/bioproject\/237077. Deposited 31 January 2014."}],"container-title":["Proceedings of the National Academy of Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/pnas.org\/doi\/pdf\/10.1073\/pnas.2610136123","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/pnas.org\/doi\/pdf\/10.1073\/pnas.2610136123","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T17:59:50Z","timestamp":1787075990000},"score":1,"resource":{"primary":{"URL":"https:\/\/pnas.org\/doi\/10.1073\/pnas.2610136123"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,18]]},"references-count":54,"journal-issue":{"issue":"34","published-print":{"date-parts":[[2026,8,25]]}},"alternative-id":["10.1073\/pnas.2610136123"],"URL":"https:\/\/doi.org\/10.1073\/pnas.2610136123","relation":{},"ISSN":["0027-8424","1091-6490"],"issn-type":[{"value":"0027-8424","type":"print"},{"value":"1091-6490","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,18]]},"assertion":[{"value":"2026-03-22","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-07-17","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-08-18","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"e2610136123"}}