{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T16:31:20Z","timestamp":1791304280185,"version":"4.3.1"},"reference-count":72,"publisher":"Oxford University Press (OUP)","issue":"10","license":[{"start":{"date-parts":[[2026,4,29]],"date-time":"2026-04-29T00:00:00Z","timestamp":1777420800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32572351"],"award-info":[{"award-number":["32572351"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100012166","name":"National Key Research and Development Program of China","doi-asserted-by":"publisher","award":["2022YFD1200104"],"award-info":[{"award-number":["2022YFD1200104"]}],"id":[{"id":"10.13039\/501100012166","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100018522","name":"Jiangsu Collaborative Innovation Center for Modern Crop Production","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100018522","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,5,21]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>High-quality seeds necessitate both appropriate dormancy to prevent preharvest sprouting and robust storability to preserve germination vigor, which serve as the fundamental guarantee for the successful production and safe storage of crops. As complex and quantitative traits, the improvement of seed quality has been impeded by limited understanding of their regulatory genes. Here, in rice, we identify SEED-AGING GENE 9 (SAG9), encoding a nuclear protein, as a negative regulator underlying the major seed-storability quantitative trait locus qSS-9, which also exerts a moderate effect on dormancy. Indeed, SAG9 physically interacts with selected bZIP transcription factors expressed in seeds, inhibiting their transcriptional activation of Late Embryogenesis Abundant genes\u2014revealing a shared genetic pathway coordinating dormancy and storability. This has driven the differential local adaptation of SAG9 distinct haplotypes, thereby promoting balancing selection during rice domestication. Notably, the elite SAG9 haplotype Hap\u2212 has been largely lost during domestication, which is one of the causes for the reduction in seed dormancy and storability of modern varieties. Furthermore, strong-dormancy alleles (SD1, SDR3.1, Sdr4, SD6, and Rc) enhance storability and show additive effects with SAG9 Hap\u2212. Diverse pyramiding combinations of these genes across regions reflect synergistic adaptation to environmental or breeding demands. Collectively, our study elucidates the molecular and domestication basis of seed quality traits, providing targets for precision breeding in modern rice varieties.<\/jats:p>","DOI":"10.1093\/nsr\/nwag248","type":"journal-article","created":{"date-parts":[[2026,4,29]],"date-time":"2026-04-29T22:27:56Z","timestamp":1777501676000},"source":"Crossref","is-referenced-by-count":2,"title":["Superior alleles of\n                    <i>SEED-AGING GENE 9<\/i>\n                    enhance seed storability and dormancy in rice"],"prefix":"10.1093","volume":"13","author":[{"ORCID":"https:\/\/orcid.org\/0009-0001-4753-4988","authenticated-orcid":false,"given":"Fulin","family":"Zhang","sequence":"first","affiliation":[{"name":"State Key Laboratory for Crop Genetics and Germplasm Enhancement, Jiangsu Nanjing National Field Scientific Observation and Research Station for Rice Germplasm, Nanjing Agricultural University , Nanjing 210095 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wenchao","family":"Chi","sequence":"additional","affiliation":[{"name":"State Key Laboratory for Crop Genetics and Germplasm Enhancement, Jiangsu Nanjing National Field Scientific Observation and Research Station for Rice Germplasm, Nanjing Agricultural University , Nanjing 210095 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0009-1443-2318","authenticated-orcid":false,"given":"Jiawei","family":"Song","sequence":"additional","affiliation":[{"name":"State Key Laboratory for Crop Genetics and Germplasm Enhancement, Jiangsu Nanjing National Field Scientific Observation and Research Station for Rice Germplasm, Nanjing Agricultural University , Nanjing 210095 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0006-2442-9727","authenticated-orcid":false,"given":"Xingjie","family":"Zhu","sequence":"additional","affiliation":[{"name":"State Key Laboratory for Crop Genetics and Germplasm Enhancement, Jiangsu Nanjing National Field Scientific Observation and Research Station for Rice Germplasm, Nanjing Agricultural University , Nanjing 210095 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0001-0547-6396","authenticated-orcid":false,"given":"Ping","family":"Wang","sequence":"additional","affiliation":[{"name":"State Key Laboratory for Crop Genetics and Germplasm Enhancement, Jiangsu Nanjing National Field Scientific Observation and Research Station for Rice Germplasm, Nanjing Agricultural University , Nanjing 210095 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0005-6669-8565","authenticated-orcid":false,"given":"Yingying","family":"Chen","sequence":"additional","affiliation":[{"name":"State Key Laboratory for Crop Genetics and Germplasm Enhancement, Jiangsu Nanjing National Field Scientific Observation and Research Station for Rice Germplasm, Nanjing Agricultural University , Nanjing 210095 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0002-9070-5102","authenticated-orcid":false,"given":"Tengfei","family":"Ma","sequence":"additional","affiliation":[{"name":"State Key Laboratory for Crop Genetics and Germplasm Enhancement, Jiangsu Nanjing National Field Scientific Observation and Research Station for Rice Germplasm, Nanjing Agricultural University , Nanjing 210095 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0001-4377-3260","authenticated-orcid":false,"given":"Ziyan","family":"Zhu","sequence":"additional","affiliation":[{"name":"State Key Laboratory for Crop Genetics and Germplasm Enhancement, Jiangsu Nanjing National Field Scientific Observation and Research Station for Rice Germplasm, Nanjing Agricultural University , Nanjing 210095 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0003-0137-9403","authenticated-orcid":false,"given":"Qixian","family":"Hao","sequence":"additional","affiliation":[{"name":"State Key Laboratory for Crop Genetics and Germplasm Enhancement, Jiangsu Nanjing National Field Scientific Observation and Research Station for Rice Germplasm, Nanjing Agricultural University , Nanjing 210095 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qingkai","family":"Wang","sequence":"additional","affiliation":[{"name":"State Key Laboratory for Crop Genetics and Germplasm Enhancement, Jiangsu Nanjing National Field Scientific Observation and Research Station for Rice Germplasm, Nanjing Agricultural University , Nanjing 210095 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ye","family":"Chen","sequence":"additional","affiliation":[{"name":"State Key Laboratory for Crop Genetics and Germplasm Enhancement, Jiangsu Nanjing National Field Scientific Observation and Research Station for Rice Germplasm, Nanjing Agricultural University , Nanjing 210095 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kunhao","family":"Li","sequence":"additional","affiliation":[{"name":"State Key Laboratory for Crop Genetics and Germplasm Enhancement, Jiangsu Nanjing National Field Scientific Observation and Research Station for Rice Germplasm, Nanjing Agricultural University , Nanjing 210095 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Boliang","family":"Lu","sequence":"additional","affiliation":[{"name":"State Key Laboratory for Crop Genetics and Germplasm Enhancement, Jiangsu Nanjing National Field Scientific Observation and Research Station for Rice Germplasm, Nanjing Agricultural University , Nanjing 210095 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yiwan","family":"Wu","sequence":"additional","affiliation":[{"name":"State Key Laboratory for Crop Genetics and Germplasm Enhancement, Jiangsu Nanjing National Field Scientific Observation and Research Station for Rice Germplasm, Nanjing Agricultural University , Nanjing 210095 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0007-9361-6400","authenticated-orcid":false,"given":"Changling","family":"Mou","sequence":"additional","affiliation":[{"name":"State Key Laboratory for Crop Genetics and Germplasm Enhancement, Jiangsu Nanjing National Field Scientific Observation and Research Station for Rice Germplasm, Nanjing Agricultural University , Nanjing 210095 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yunshuai","family":"Huang","sequence":"additional","affiliation":[{"name":"State Key Laboratory for Crop Genetics and Germplasm Enhancement, Jiangsu Nanjing National Field Scientific Observation and Research Station for Rice Germplasm, Nanjing Agricultural University , Nanjing 210095 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hongming","family":"Wu","sequence":"additional","affiliation":[{"name":"State Key Laboratory for Crop Genetics and Germplasm Enhancement, Jiangsu Nanjing National Field Scientific Observation and Research Station for Rice Germplasm, Nanjing Agricultural University , Nanjing 210095 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0000-9199-1740","authenticated-orcid":false,"given":"Rong","family":"Miao","sequence":"additional","affiliation":[{"name":"State Key Laboratory for Crop Genetics and Germplasm Enhancement, Jiangsu Nanjing National Field Scientific Observation and Research Station for Rice Germplasm, Nanjing Agricultural University , Nanjing 210095 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qiuyun","family":"Lin","sequence":"additional","affiliation":[{"name":"State Key Laboratory for Crop Genetics and Germplasm Enhancement, Jiangsu Nanjing National Field Scientific Observation and Research Station for Rice Germplasm, Nanjing Agricultural University , Nanjing 210095 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Penghui","family":"Cao","sequence":"additional","affiliation":[{"name":"State Key Laboratory for Crop Genetics and Germplasm Enhancement, Jiangsu Nanjing National Field Scientific Observation and Research Station for Rice Germplasm, Nanjing Agricultural University , Nanjing 210095 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3268-6097","authenticated-orcid":false,"given":"Shijia","family":"Liu","sequence":"additional","affiliation":[{"name":"State Key Laboratory for Crop Genetics and Germplasm Enhancement, Jiangsu Nanjing National Field Scientific Observation and Research Station for Rice Germplasm, Nanjing Agricultural University , Nanjing 210095 ,","place":["China"]},{"name":"Zhongshan Biological Breeding Laboratory , Nanjing 210095 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1464-5881","authenticated-orcid":false,"given":"Ling","family":"Jiang","sequence":"additional","affiliation":[{"name":"State Key Laboratory for Crop Genetics and Germplasm Enhancement, Jiangsu Nanjing National Field Scientific Observation and Research Station for Rice Germplasm, Nanjing Agricultural University , Nanjing 210095 ,","place":["China"]},{"name":"Zhongshan Biological Breeding Laboratory , Nanjing 210095 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jianmin","family":"Wan","sequence":"additional","affiliation":[{"name":"State Key Laboratory for Crop Genetics and Germplasm Enhancement, Jiangsu Nanjing National Field Scientific Observation and Research Station for Rice Germplasm, Nanjing Agricultural University , Nanjing 210095 ,","place":["China"]},{"name":"Institute of Crop Science, Chinese Academy of Agricultural Sciences National Key Facility for Crop Gene Resources and Genetic Improvement, , Beijing 100081 ,","place":["China"]},{"name":"Zhongshan Biological Breeding Laboratory , Nanjing 210095 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,4,29]]},"reference":[{"key":"2026060918345108900_bib1","doi-asserted-by":"publisher","first-page":"507","DOI":"10.1146\/annurev-arplant-042811-105550","article-title":"Seed germination and vigor","volume":"63","author":"Rajjou","year":"2012","journal-title":"Annu Rev Plant Biol"},{"key":"2026060918345108900_bib2","doi-asserted-by":"publisher","first-page":"1440","DOI":"10.1111\/j.1365-3040.2012.02501.x","article-title":"Temporal profiling of the heat-stable proteome during late maturation of Medicago truncatula seeds identifies a restricted subset of late embryogenesis abundant proteins associated with longevity","volume":"35","author":"Chatelain","year":"2012","journal-title":"Plant Cell Environ"},{"key":"2026060918345108900_bib3","doi-asserted-by":"publisher","first-page":"960","DOI":"10.1016\/j.molp.2025.05.010","article-title":"Seed dormancy and germination in rice: molecular regulatory mechanisms and breeding","volume":"18","author":"Xu","year":"2025","journal-title":"Mol Plant"},{"key":"2026060918345108900_bib4","doi-asserted-by":"publisher","first-page":"3857","DOI":"10.1093\/plcell\/koae211","article-title":"The phosphatidylethanolamine-binding proteins OsMFT1 and OsMFT2 regulate seed dormancy in rice","volume":"36","author":"Shen","year":"2024","journal-title":"Plant Cell"},{"key":"2026060918345108900_bib5","doi-asserted-by":"publisher","first-page":"e2026355119","DOI":"10.1073\/pnas.2026355119","article-title":"A multiomic study uncovers a bZIP23-PER1A-mediated detoxification pathway to enhance seed vigor in rice","volume":"119","author":"Wang","year":"2022","journal-title":"Proc Natl Acad Sci USA"},{"key":"2026060918345108900_bib6","doi-asserted-by":"publisher","first-page":"660","DOI":"10.1093\/pcp\/pcv186","article-title":"Staying alive: molecular aspects of seed longevity","volume":"57","author":"Sano","year":"2016","journal-title":"Plant Cell Physiol"},{"key":"2026060918345108900_bib7","doi-asserted-by":"publisher","first-page":"20972","DOI":"10.1021\/acs.jafc.5c06276","article-title":"Improvement of rice seed storability by regulating lipid metabolism using a CRISPR\/Cas9 system","volume":"73","author":"Zhou","year":"2025","journal-title":"J Agric Food Chem"},{"key":"2026060918345108900_bib8","doi-asserted-by":"publisher","first-page":"112215","DOI":"10.1016\/j.plantsci.2024.112215","article-title":"Rice seed storability: from molecular mechanisms to agricultural practices","volume":"348","author":"Yan","year":"2024","journal-title":"Plant Sci"},{"key":"2026060918345108900_bib9","doi-asserted-by":"publisher","first-page":"1357","DOI":"10.3390\/ijms22031357","article-title":"Regulation of seed dormancy and germination mechanisms in a changing environment","volume":"22","author":"Klupczynska","year":"2021","journal-title":"Int J Mol Sci"},{"key":"2026060918345108900_bib10","doi-asserted-by":"publisher","first-page":"450","DOI":"10.1038\/s41437-022-00497-2","article-title":"Seed germination and vigor: ensuring crop sustainability in a changing climate","volume":"128","author":"Reed","year":"2022","journal-title":"Heredity"},{"key":"2026060918345108900_bib11","doi-asserted-by":"publisher","first-page":"437","DOI":"10.1007\/s10342-023-01633-1","article-title":"Global warming could shorten the seed lifespan of pioneer tree species and thus natural regeneration window of damaged areas","volume":"143","author":"Tiebel","year":"2024","journal-title":"Eur J Forest Res"},{"key":"2026060918345108900_bib12","doi-asserted-by":"publisher","first-page":"177","DOI":"10.1042\/BCJ20220246","article-title":"Impact of climate perturbations on seeds and seed quality for global agriculture","volume":"480","author":"Bailly","year":"2023","journal-title":"Biochem J"},{"key":"2026060918345108900_bib13","doi-asserted-by":"publisher","first-page":"2735","DOI":"10.1105\/tpc.16.00470","article-title":"ABI5 is a regulator of seed maturation and longevity in legumes","volume":"28","author":"Zinsmeister","year":"2016","journal-title":"Plant Cell"},{"key":"2026060918345108900_bib14","doi-asserted-by":"publisher","first-page":"2472","DOI":"10.1111\/pbi.14362","article-title":"A genome-wide association study uncovers a ZmRap2.7-ZCN9\/ZCN10 module to regulate ABA signalling and seed vigour in maize","volume":"22","author":"Guo","year":"2024","journal-title":"Plant Biotechnol J"},{"key":"2026060918345108900_bib15","doi-asserted-by":"publisher","first-page":"1134","DOI":"10.1038\/s41467-024-45402-z","article-title":"A mediator of OsbZIP46 deactivation and degradation negatively regulates seed dormancy in rice","volume":"15","author":"Guo","year":"2024","journal-title":"Nat Commun"},{"key":"2026060918345108900_bib16","doi-asserted-by":"publisher","first-page":"532","DOI":"10.1111\/tpj.14748","article-title":"OsMFT2 is involved in the regulation of ABA signaling-mediated seed germination through interacting with OsbZIP23\/66\/72 in rice","volume":"103","author":"Song","year":"2020","journal-title":"Plant J"},{"key":"2026060918345108900_bib17","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1017\/S0960258511000079","article-title":"The reduction of seed-specific dehydrins reduces seed longevity in Arabidopsis thaliana","volume":"21","author":"Hundertmark","year":"2011","journal-title":"Seed Sci Res"},{"key":"2026060918345108900_bib18","doi-asserted-by":"publisher","first-page":"1185","DOI":"10.1104\/pp.102.4.1185","article-title":"Acquisition of desiccation tolerance and longevity in seeds of Arabidopsis thaliana (a comparative study using abscisic acid-insensitive abi3 mutants)","volume":"102","author":"Ooms","year":"1993","journal-title":"Plant Physiol"},{"key":"2026060918345108900_bib19","doi-asserted-by":"publisher","first-page":"448","DOI":"10.1111\/j.0031-9317.2004.00339.x","article-title":"Genetic differences in seed longevity of various Arabidopsis mutants","volume":"121","author":"Clerkx","year":"2004","journal-title":"Physiol Plant"},{"key":"2026060918345108900_bib20","doi-asserted-by":"publisher","first-page":"451","DOI":"10.1111\/tpj.13118","article-title":"The Arabidopsis DELAY OF GERMINATION 1 gene affects ABSCISIC ACID INSENSITIVE 5 (ABI5) expression and genetically interacts with ABI3 during Arabidopsis seed development","volume":"85","author":"Dekkers","year":"2016","journal-title":"Plant J"},{"key":"2026060918345108900_bib21","doi-asserted-by":"publisher","first-page":"17042","DOI":"10.1073\/pnas.0607877103","article-title":"Cloning of DOG1, a quantitative trait locus controlling seed dormancy in Arabidopsis","volume":"103","author":"Bentsink","year":"2006","journal-title":"Proc Natl Acad Sci USA"},{"key":"2026060918345108900_bib22","doi-asserted-by":"publisher","first-page":"373","DOI":"10.1146\/annurev.arplant.55.031903.141701","article-title":"Reactive oxygen species: metabolism, oxidative stress, and signal transduction","volume":"55","author":"Apel","year":"2004","journal-title":"Annu Rev Plant Biol"},{"key":"2026060918345108900_bib23","doi-asserted-by":"publisher","first-page":"kiae642","DOI":"10.1093\/plphys\/kiae642","article-title":"The BRAHMA-associated SWI\/SNF chromatin remodeling complex controls Arabidopsis seed quality and physiology","volume":"197","author":"Wrona","year":"2024","journal-title":"Plant Physiol"},{"key":"2026060918345108900_bib24","doi-asserted-by":"publisher","first-page":"669","DOI":"10.1093\/aob\/mcv108","article-title":"Glutathione redox state, tocochromanols, fatty acids, antioxidant enzymes and protein carbonylation in sunflower seed embryos associated with after-ripening and ageing","volume":"116","author":"Morscher","year":"2015","journal-title":"Ann Bot"},{"key":"2026060918345108900_bib25","doi-asserted-by":"publisher","first-page":"6047","DOI":"10.1093\/jxb\/erae163","article-title":"Plant quiescence strategy and seed dormancy under hypoxia","volume":"75","author":"Pucciariello","year":"2024","journal-title":"J Exp Bot"},{"key":"2026060918345108900_bib26","doi-asserted-by":"publisher","first-page":"e2402407121","DOI":"10.1073\/pnas.2402407121","article-title":"GPCR-MAPK signaling pathways underpin fitness trade-offs in whitefly","volume":"121","author":"Fu","year":"2024","journal-title":"Proc Natl Acad Sci USA"},{"key":"2026060918345108900_bib27","doi-asserted-by":"publisher","first-page":"728","DOI":"10.1126\/science.aar3780","article-title":"Wicked evolution: can we address the sociobiological dilemma of pesticide resistance?","volume":"360","author":"Gould","year":"2018","journal-title":"Science"},{"key":"2026060918345108900_bib28","first-page":"305","article-title":"Sickle cell disease","volume":"377","author":"Piel","year":"2017","journal-title":"N Engl J Med"},{"key":"2026060918345108900_bib29","doi-asserted-by":"publisher","first-page":"104","DOI":"10.1038\/ncomms1104","article-title":"Global distribution of the sickle cell gene and geographical confirmation of the malaria hypothesis","volume":"1","author":"Piel","year":"2010","journal-title":"Nat Commun"},{"key":"2026060918345108900_bib30","doi-asserted-by":"publisher","first-page":"7051","DOI":"10.1073\/pnas.1505665112","article-title":"Malaria continues to select for sickle cell trait in Central Africa","volume":"112","author":"Elguero","year":"2015","journal-title":"Proc Natl Acad Sci USA"},{"key":"2026060918345108900_bib31","doi-asserted-by":"publisher","first-page":"74","DOI":"10.1038\/nature01588","article-title":"Fitness costs of R-gene-mediated resistance in Arabidopsis thaliana","volume":"423","author":"Tian","year":"2003","journal-title":"Nature"},{"key":"2026060918345108900_bib32","doi-asserted-by":"publisher","first-page":"1069","DOI":"10.1016\/j.tplants.2017.09.010","article-title":"Balancing immunity and yield in crop plants","volume":"22","author":"Ning","year":"2017","journal-title":"Trends Plant Sci"},{"key":"2026060918345108900_bib33","doi-asserted-by":"publisher","first-page":"1246","DOI":"10.1111\/jipb.13266","article-title":"Sdr4 dominates pre-harvest sprouting and facilitates adaptation to local climatic condition in Asian cultivated rice","volume":"64","author":"Zhao","year":"2022","journal-title":"J Integr Plant Biol"},{"key":"2026060918345108900_bib34","doi-asserted-by":"publisher","first-page":"1972","DOI":"10.1038\/s41588-022-01240-7","article-title":"Antagonistic control of seed dormancy in rice by two bHLH transcription factors","volume":"54","author":"Xu","year":"2022","journal-title":"Nat Genet"},{"key":"2026060918345108900_bib35","doi-asserted-by":"publisher","first-page":"293","DOI":"10.1111\/pbr.12264","article-title":"Fine mapping of qSS-9, a major and stable quantitative trait locus, for seed storability in rice (Oryza sativa L.)","volume":"134","author":"Lin","year":"2015","journal-title":"Plant Breed"},{"key":"2026060918345108900_bib36","doi-asserted-by":"publisher","first-page":"769","DOI":"10.1007\/s00122-015-2471-7","article-title":"Fine mapping of a major quantitative trait locus, qLG-9, that controls seed longevity in rice (Oryza sativa L.)","volume-title":"Theor Appl Genet","author":"Sasaki","year":"2015"},{"key":"2026060918345108900_bib37","doi-asserted-by":"publisher","first-page":"333","DOI":"10.1104\/pp.107.112821","article-title":"Genomic survey and gene expression analysis of the basic leucine zipper transcription factor family in rice","volume":"146","author":"Nijhawan","year":"2008","journal-title":"Plant Physiol"},{"key":"2026060918345108900_bib38","doi-asserted-by":"publisher","first-page":"kiae545","DOI":"10.1093\/plphys\/kiae545","article-title":"OsPUB75-OsHDA716 mediates deactivation and degradation of OsbZIP46 to negatively regulate drought tolerance in rice","volume":"197","author":"Sun","year":"2024","journal-title":"Plant Physiol"},{"key":"2026060918345108900_bib39","doi-asserted-by":"publisher","first-page":"1297","DOI":"10.1016\/j.molp.2021.05.001","article-title":"Nuclear translocation of OsMFT1 that is impeded by OsFTIP1 promotes drought tolerance in rice","volume":"14","author":"Chen","year":"2021","journal-title":"Mol Plant"},{"key":"2026060918345108900_bib40","doi-asserted-by":"publisher","first-page":"1390","DOI":"10.1016\/j.molp.2025.07.011","article-title":"A novel OsCRK14-OsRLCK57-MAPK signaling module activates OsbZIP66 to confer drought resistance in rice","volume":"18","author":"Ye","year":"2025","journal-title":"Mol Plant"},{"key":"2026060918345108900_bib41","doi-asserted-by":"publisher","first-page":"605","DOI":"10.1007\/s00425-008-0857-3","article-title":"Identification of OsbZIP72 as a positive regulator of ABA response and drought tolerance in rice","volume":"229","author":"Lu","year":"2009","journal-title":"Planta"},{"key":"2026060918345108900_bib42","doi-asserted-by":"publisher","first-page":"723","DOI":"10.1111\/jipb.12850","article-title":"Synergistic regulation of drought-responsive genes by transcription factor OsbZIP23 and histone modification in rice","volume":"62","author":"Zong","year":"2020","journal-title":"J Integr Plant Biol"},{"key":"2026060918345108900_bib43","doi-asserted-by":"publisher","first-page":"1031","DOI":"10.1111\/tpj.13739","article-title":"Rice SUMO protease overly tolerant to salt 1 targets the transcription factor, OsbZIP23 to promote drought tolerance in rice","volume":"92","author":"Srivastava","year":"2017","journal-title":"Plant J"},{"key":"2026060918345108900_bib44","doi-asserted-by":"publisher","first-page":"201","DOI":"10.1016\/j.devcel.2016.06.011","article-title":"OsREM4.1 interacts with OsSERK1 to coordinate the interlinking between abscisic acid and brassinosteroid signaling in rice","volume":"38","author":"Gui","year":"2016","journal-title":"Dev Cell"},{"key":"2026060918345108900_bib45","doi-asserted-by":"publisher","first-page":"2161","DOI":"10.1105\/tpc.16.00171","article-title":"MODD mediates deactivation and degradation of OsbZIP46 to negatively regulate aba signaling and drought resistance in rice","volume":"28","author":"Tang","year":"2016","journal-title":"Plant Cell"},{"key":"2026060918345108900_bib46","doi-asserted-by":"publisher","first-page":"1397","DOI":"10.1104\/pp.111.173427","article-title":"Rice ABI5-Like1 regulates abscisic acid and auxin responses by affecting the expression of ABRE-containing genes","volume":"156","author":"Yang","year":"2011","journal-title":"Plant Physiol"},{"key":"2026060918345108900_bib47","doi-asserted-by":"publisher","first-page":"1512","DOI":"10.1016\/j.jplph.2010.05.008","article-title":"The ABRE-binding bZIP transcription factor OsABF2 is a positive regulator of abiotic stress and ABA signaling in rice","volume":"167","author":"Hossain","year":"2010","journal-title":"J Plant Physiol"},{"key":"2026060918345108900_bib48","doi-asserted-by":"publisher","first-page":"1336","DOI":"10.1111\/nph.16774","article-title":"Abscisic acid promotes jasmonic acid biosynthesis via a \u2018SAPK10-bZIP72-AOC\u2019 pathway to synergistically inhibit seed germination in rice (Oryza sativa)","volume":"228","author":"Wang","year":"2020","journal-title":"New Phytol"},{"key":"2026060918345108900_bib49","doi-asserted-by":"publisher","first-page":"939","DOI":"10.1111\/j.1365-313X.2005.02583.x","article-title":"Abscisic acid-activated SNRK2 protein kinases function in the gene-regulation pathway of ABA signal transduction by phosphorylating ABA response element-binding factors","volume":"44","author":"Kobayashi","year":"2005","journal-title":"Plant J"},{"key":"2026060918345108900_bib50","doi-asserted-by":"publisher","first-page":"15348","DOI":"10.1073\/pnas.96.26.15348","article-title":"A bZIP factor, TRAB1, interacts with VP1 and mediates abscisic acid-induced transcription","volume":"96","author":"Hobo","year":"1999","journal-title":"Proc Natl Acad Sci USA"},{"key":"2026060918345108900_bib51","doi-asserted-by":"crossref","first-page":"257","DOI":"10.1016\/j.rsci.2021.04.005","article-title":"OsbZIP72 is involved in transcriptional gene-regulation pathway of abscisic acid signal transduction by activating rice high-affinity potassium transporter OsHKT1;1","volume":"28","author":"Wang","year":"2021","journal-title":"Rice Sci"},{"key":"2026060918345108900_bib52","doi-asserted-by":"publisher","first-page":"112","DOI":"10.1016\/j.plaphy.2021.11.005","article-title":"bZIP72 promotes submerged rice seed germination and coleoptile elongation by activating ADH1","volume":"169","author":"Wang","year":"2021","journal-title":"Plant Physiol Biochem"},{"key":"2026060918345108900_bib53","doi-asserted-by":"publisher","first-page":"koae304","DOI":"10.1093\/plcell\/koae304","article-title":"Maternally expressed FERTILIZATION- INDEPENDENT ENDOSPERM1 regulates seed dormancy and aleurone development in rice","volume":"37","author":"Cheng","year":"2024","journal-title":"Plant Cell"},{"key":"2026060918345108900_bib54","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1038\/s41586-018-0063-9","article-title":"Genomic variation in 3,010 diverse accessions of Asian cultivated rice","volume":"557","author":"Wang","year":"2018","journal-title":"Nature"},{"key":"2026060918345108900_bib55","doi-asserted-by":"crossref","unstructured":"Li JY, Wang J, Zeigler RS. The 3,000 rice genomes project: new opportunities and challenges for future rice research. Gigascience \u00a02014; 3: 8.10.1186\/2047-217X-3-8","DOI":"10.1186\/2047-217X-3-8"},{"key":"2026060918345108900_bib56","doi-asserted-by":"publisher","first-page":"D1018","DOI":"10.1093\/nar\/gku894","article-title":"RiceVarMap: a comprehensive database of rice genomic variations","volume":"43","author":"Zhao","year":"2015","journal-title":"Nucleic Acids Res"},{"key":"2026060918345108900_bib57","first-page":"812","article-title":"Genome-wide association study reveals the QTLs for seed storability in world rice core collections","volume":"10","author":"Wu","year":"2021","journal-title":"Plants (Basel)"},{"key":"2026060918345108900_bib58","doi-asserted-by":"publisher","first-page":"3553","DOI":"10.1007\/s00122-021-03911-1","article-title":"Integrating GWAS and transcriptomics to identify genes involved in seed dormancy in rice","volume":"134","author":"Shi","year":"2021","journal-title":"Theor Appl Genet"},{"key":"2026060918345108900_bib59","doi-asserted-by":"publisher","first-page":"5792","DOI":"10.1073\/pnas.0911965107","article-title":"Molecular cloning of Sdr4, a regulator involved in seed dormancy and domestication of rice","volume":"107","author":"Sugimoto","year":"2010","journal-title":"Proc Natl Acad Sci USA"},{"key":"2026060918345108900_bib60","doi-asserted-by":"publisher","first-page":"1515","DOI":"10.1534\/genetics.111.131169","article-title":"Association between seed dormancy and pericarp color is controlled by a pleiotropic gene that regulates abscisic acid and flavonoid synthesis in weedy red rice","volume":"189","author":"Gu","year":"2011","journal-title":"Genetics"},{"key":"2026060918345108900_bib61","doi-asserted-by":"publisher","first-page":"469","DOI":"10.1093\/plphys\/kiab059","article-title":"OsGRETCHENHAGEN3-2 modulates rice seed storability via accumulation of abscisic acid and protective substances","volume":"186","author":"Yuan","year":"2021","journal-title":"Plant Physiol"},{"key":"2026060918345108900_bib62","doi-asserted-by":"publisher","first-page":"497","DOI":"10.1038\/nature11532","article-title":"A map of rice genome variation reveals the origin of cultivated rice","volume":"490","author":"Huang","year":"2012","journal-title":"Nature"},{"key":"2026060918345108900_bib63","doi-asserted-by":"publisher","first-page":"1938","DOI":"10.1104\/pp.108.128199","article-title":"Characterization of OsbZIP23 as a key player of the basic leucine zipper transcription factor family for conferring abscisic acid sensitivity and salinity and drought tolerance in rice","volume":"148","author":"Xiang","year":"2008","journal-title":"Plant Physiol"},{"key":"2026060918345108900_bib64","doi-asserted-by":"publisher","first-page":"3633","DOI":"10.1073\/pnas.1220589110","article-title":"Evidence for participation of the methionine sulfoxide reductase repair system in plant seed longevity","volume":"110","author":"Chatelain","year":"2013","journal-title":"Proc Natl Acad Sci USA"},{"key":"2026060918345108900_bib65","doi-asserted-by":"publisher","first-page":"5664","DOI":"10.1038\/s41467-022-33320-x","article-title":"Regain flood adaptation in rice through a 14-3-3 protein OsGF14h","volume":"13","author":"Sun","year":"2022","journal-title":"Nat Commun"},{"key":"2026060918345108900_bib66","doi-asserted-by":"publisher","first-page":"5665","DOI":"10.1038\/s41467-022-33318-5","article-title":"Genome-wide association study identifies a gene responsible for temperature-dependent rice germination","volume":"13","author":"Yoshida","year":"2022","journal-title":"Nat Commun"},{"key":"2026060918345108900_bib67","doi-asserted-by":"publisher","first-page":"415","DOI":"10.1046\/j.1365-313x.2000.00797.x","article-title":"Quantitative nature of the prolamin-box, ACGT and AACA motifs in a rice glutelin gene promoter: minimal cis-element requirements for endosperm-specific gene expression","volume":"23","author":"Wu","year":"2000","journal-title":"Plant J"},{"key":"2026060918345108900_bib68","doi-asserted-by":"publisher","first-page":"4070","DOI":"10.3390\/plants12234070","article-title":"OsLEA1b modulates starch biosynthesis at high temperatures in rice","volume":"12","author":"Li","year":"2023","journal-title":"Plants"},{"key":"2026060918345108900_bib69","doi-asserted-by":"publisher","first-page":"3765","DOI":"10.1093\/jxb\/erz168","article-title":"NF-YC12 is a key multi-functional regulator of accumulation of seed storage substances in rice","volume":"70","author":"Xiong","year":"2019","journal-title":"J Exp Bot"},{"key":"2026060918345108900_bib70","doi-asserted-by":"crossref","first-page":"962","DOI":"10.3390\/life15060962","article-title":"OsMYBR1, a 1R-MYB family transcription factor regulates starch biosynthesis in rice endosperm","volume":"15","author":"Huang","year":"2025","journal-title":"Life"},{"key":"2026060918345108900_bib71","doi-asserted-by":"publisher","first-page":"e2321612121","DOI":"10.1073\/pnas.2321612121","article-title":"Autophagy maintains endosperm quality during seed storage to preserve germination ability in Arabidopsis","volume":"121","author":"Shinozaki","year":"2024","journal-title":"Proc Natl Acad Sci USA"},{"key":"2026060918345108900_bib72","doi-asserted-by":"publisher","first-page":"406","DOI":"10.1038\/nature12878","article-title":"D14-SCFD3-dependent degradation of D53 regulates strigolactone signalling","volume":"504","author":"Zhou","year":"2013","journal-title":"Nature"}],"container-title":["National Science Review"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/nsr\/advance-article-pdf\/doi\/10.1093\/nsr\/nwag248\/68189466\/nwag248.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/nsr\/article-pdf\/13\/10\/nwag248\/68189466\/nwag248.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/nsr\/article-pdf\/13\/10\/nwag248\/68189466\/nwag248.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,6,9]],"date-time":"2026-06-09T22:34:58Z","timestamp":1781044498000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/nsr\/article\/doi\/10.1093\/nsr\/nwag248\/8665133"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,4,29]]},"references-count":72,"journal-issue":{"issue":"10","published-print":{"date-parts":[[2026,5,21]]}},"URL":"https:\/\/doi.org\/10.1093\/nsr\/nwag248","relation":{},"ISSN":["2095-5138","2053-714X"],"issn-type":[{"value":"2095-5138","type":"print"},{"value":"2053-714X","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2026,5]]},"published":{"date-parts":[[2026,4,29]]},"article-number":"nwag248"}}