{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,24]],"date-time":"2026-09-24T06:04:09Z","timestamp":1790229849153,"version":"4.1.0"},"reference-count":27,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,9,24]],"date-time":"2026-09-24T00:00:00Z","timestamp":1790208000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Plant Sci."],"abstract":"<jats:sec>\n                    <jats:title>Introduction<\/jats:title>\n                    <jats:p>The stay-green trait is crucial for enhancing maize yield and stress tolerance, yet its genetic basis in new germplasm remains underexplored. This study aimed to dissect the molecular mechanisms underlying stay-green in the inbred line T01 using an integrated approach.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods<\/jats:title>\n                    <jats:p>We constructed an F\u2082:\u2083 population from T01 and the non-stay-green line Mo17, and combined BSA-seq to map candidate loci, RNA-seq at three developmental stages to profile transcriptomic dynamics, and WGCNA to identify co-expression networks.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>\n                      BSA-seq mapped two candidate QTL regions on chromosomes 5 and 9, with the former representing a potentially novel locus. Transcriptomic analysis revealed stage-specific regulation, with enrichment in reactive oxygen species metabolism, photosynthesis, and chloroplast maintenance at 10, 20, and 30 days after pollination (DAP), respectively. WGCNA identified six hub genes functionally involved in transcriptional regulation, ROS scavenging, and photosynthesis. Notably, three genes (\n                      <jats:italic>ZmFSD3<\/jats:italic>\n                      ,\n                      <jats:italic>ZmWRKY62<\/jats:italic>\n                      , and\n                      <jats:italic>ZmNAC29<\/jats:italic>\n                      ) were located within BSA-defined intervals and supported by WGCNA analysis, with their expression patterns validated by RT-qPCR.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Discussion<\/jats:title>\n                    <jats:p>Our findings reveal a complex, multi-pathway regulatory network in which WRKY and NAC transcription factors are highly connected candidate nodes. This provides new genetic resources and a theoretical basis for molecular breeding of stay-green maize.<\/jats:p>\n                  <\/jats:sec>","DOI":"10.3389\/fpls.2026.1936883","type":"journal-article","created":{"date-parts":[[2026,9,24]],"date-time":"2026-09-24T05:37:41Z","timestamp":1790228261000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Integrated transcriptomic and BSA-seq analyses identify novel QTL and candidate genes for stay-green in maize"],"prefix":"10.3389","volume":"17","author":[{"given":"Ran","family":"Zheng","sequence":"first","affiliation":[{"name":"Hunan Medical College","place":["Huaihua, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dan","family":"Lv","sequence":"additional","affiliation":[{"name":"Department of Biology and Food Engineering, Lyuliang University","place":["Lvliang, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kunyi","family":"Shen","sequence":"additional","affiliation":[{"name":"Hunan Medical College","place":["Huaihua, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qinggui","family":"Wu","sequence":"additional","affiliation":[{"name":"Hunan Agricultural University","place":["Changsha, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hongbing","family":"Luo","sequence":"additional","affiliation":[{"name":"Hunan Agricultural University","place":["Changsha, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tao","family":"Chang","sequence":"additional","affiliation":[{"name":"Hunan Medical College","place":["Huaihua, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,9,24]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"701","DOI":"10.1007\/s11032-014-0068-5","article-title":"Molecular mapping across three populations reveals a QTL hotspot region on chromosome 3 for secondary traits associated with drought tolerance in tropical maize","volume":"34","author":"Almeida","year":"2014","journal-title":"Mol. Breed."},{"key":"B2","doi-asserted-by":"publisher","first-page":"1934","DOI":"10.1105\/tpc.19.00465","article-title":"Senescence: The genetics behind stay-green corn","volume":"31","author":"Celine","year":"2019","journal-title":"Plant Cell"},{"key":"B3","doi-asserted-by":"publisher","DOI":"10.3390\/genes10110944","article-title":"Dissecting the regulatory network of leaf premature senescence in maize (Zea mays L.) using transcriptome analysis of ZmELS5 mutant","volume":"10","author":"Chai","year":"2019","journal-title":"Genes"},{"key":"B4","doi-asserted-by":"publisher","first-page":"1174","DOI":"10.1016\/j.molp.2017.07.008","article-title":"Arabidopsis WRKY45 interacts with the DELLA protein RGL1 to positively regulate age-triggered leaf senescence","volume":"10","author":"Chen","year":"2017","journal-title":"Mol. Plant"},{"key":"B5","first-page":"260","article-title":"Genetic analysis and assessment of stay-green traits in hybrids of temperate and tropical maize germplasm","volume":"58","author":"Chen","year":"2013","journal-title":"Maydica"},{"key":"B6","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2024.1399072","article-title":"Impact of irrigation, nitrogen fertilization, and plant density on stay-green and its effects on agronomic traits in maize","volume":"15","author":"Chibane","year":"2024","journal-title":"Front. Plant Sci."},{"key":"B7","doi-asserted-by":"publisher","first-page":"478","DOI":"10.1186\/s12870-024-05199-x","article-title":"Transcriptome-based identification and validation of reference genes for corn growth stages, different tissues, and drought stress in taro (Colocasia esculenta)","volume":"24","author":"Dong","year":"2024","journal-title":"BMC Plant Biol."},{"key":"B8","doi-asserted-by":"publisher","first-page":"182","DOI":"10.6048\/j.issn.1001-4330.2025.01.021","article-title":"QTL mapping and genetic analysis of soluble solids content in the center flesh of muskmelon","volume":"62","author":"Fan","year":"2025","journal-title":"Xinjiang. Agric. Sci."},{"key":"B9","doi-asserted-by":"publisher","DOI":"10.3389\/fphgy.2024.1331421","article-title":"Influence of high temperature and drought stress at jointing stage on crop physiological responses and growth in summer maize plants (Zea mays L.)","volume":"2","author":"Fan","year":"2024","journal-title":"Frontier. Plant Biochem."},{"key":"B10","doi-asserted-by":"publisher","first-page":"1558","DOI":"10.1016\/j.molp.2022.08.009","article-title":"Natural variations of ZmSRO1d modulate the trade-off between drought resistance and yield by affecting ZmRBOHC-mediated stomatal ROS production in maize","volume":"15","author":"Gao","year":"2022","journal-title":"Mol. Plant"},{"key":"B11","doi-asserted-by":"publisher","first-page":"830","DOI":"10.1016\/j.plaphy.2023.02.034","article-title":"Within-leaf chloroplasts and nitrogen allocation to thylakoids in relation to photosynthesis during grain filling in maize","volume":"196","author":"Gao","year":"2023","journal-title":"Plant Physiol. Biochem."},{"key":"B12","doi-asserted-by":"publisher","DOI":"10.1038\/srep09311","article-title":"Effects of nitrogen stress on the photosynthetic CO2 assimilation, chlorophyll fluorescence, and sugar-nitrogen ratio in corn","volume":"5","author":"Jin","year":"2015","journal-title":"Sci. Rep."},{"key":"B13","doi-asserted-by":"publisher","DOI":"10.3390\/ijms20235837","article-title":"Stay-green trait: A prospective approach for yield potential, and drought and heat stress adaptation in globally important cereals","volume":"20","author":"Kamal","year":"2019","journal-title":"Int. J. Mol. Sci."},{"key":"B14","doi-asserted-by":"publisher","first-page":"463","DOI":"10.1007\/s10681-015-1575-0","article-title":"Stay-green QTLs in temperate elite maize","volume":"207","author":"Kante","year":"2016","journal-title":"Euphytica"},{"key":"B15","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1007\/s10725-021-00758-w","article-title":"Stay-green trait serves as yield stability attribute under combined heat and drought stress in wheat (Triticum aestivum L.)","volume":"96","author":"Kumar","year":"2022","journal-title":"Plant Growth Regul."},{"key":"B16","doi-asserted-by":"publisher","DOI":"10.1016\/j.cpb.2024.100428","article-title":"Transcription factors-regulated leaf senescence in major crops: Insights, applications, and challenges","volume":"40","author":"Li","year":"2024","journal-title":"Curr. Plant Biol."},{"key":"B17","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2026.1810580","article-title":"Genetics of polymorphism in nitrogen-induced-susceptibility of rice to Magnaporthe oryzae","volume":"17","author":"Liu","year":"2026","journal-title":"Front. Plant Sci."},{"key":"B18","doi-asserted-by":"publisher","first-page":"1747","DOI":"10.1104\/pp.17.00542","article-title":"A rice NAC transcription factor promotes leaf senescence via ABA biosynthesis","volume":"174","author":"Mao","year":"2017","journal-title":"Plant Physiol."},{"key":"B19","doi-asserted-by":"publisher","first-page":"619","DOI":"10.1007\/s00344-018-9877-9","article-title":"Action of N-succinyl and N,O-dicarboxymethyl chitosan derivatives on chlorophyll photosynthesis and fluorescence in drought-sensitive maize","volume":"38","author":"Reis","year":"2019","journal-title":"J. Plant Growth Regul."},{"key":"B20","doi-asserted-by":"publisher","first-page":"208","DOI":"10.1186\/s12915-022-01398-w","article-title":"Improved biomarker discovery through a plot twist in transcriptomic data analysis","volume":"20","author":"S\u00e1nchez-Baiz\u00e1n","year":"2022","journal-title":"BMC Biol."},{"key":"B21","doi-asserted-by":"publisher","first-page":"1968","DOI":"10.1105\/tpc.18.00930","article-title":"Integrated genome-scale analysis identifies novel genes and networks underlying senescence in maize","volume":"31","author":"Sekhon","year":"2019","journal-title":"Plant Cell"},{"key":"B22","doi-asserted-by":"publisher","DOI":"10.3390\/plants14010012","article-title":"Identification of ZmSNAC06, a maize NAC family transcription factor with multiple transcripts conferring drought tolerance in Arabidopsis","volume":"14","author":"Wang","year":"2024","journal-title":"Plants"},{"key":"B23","doi-asserted-by":"publisher","first-page":"111774","DOI":"10.2139\/ssrn.4374093","article-title":"The NAC transcription factor ZmNAC132 regulates leaf senescence and male fertility in maize","volume":"334","author":"Yuan","year":"2023","journal-title":"Plant Sci."},{"key":"B24","doi-asserted-by":"publisher","first-page":"2272","DOI":"10.1111\/pbi.13139","article-title":"Identification and characterization of a novel stay-green QTL that increases yield in maize","volume":"17","author":"Zhang","year":"2019","journal-title":"Plant Biotechnol. J."},{"key":"B25","doi-asserted-by":"publisher","DOI":"10.3390\/ijms242417617","article-title":"Combined BSA-Seq and RNA-Seq reveal genes associated with the visual stay-green of maize (Zea mays L.)","volume":"24","author":"Zheng","year":"2023","journal-title":"Int. J. Mol. Sci."},{"key":"B26","doi-asserted-by":"publisher","first-page":"e0303602","DOI":"10.1371\/journal.pone.0303602","article-title":"Genetic analysis of stay green related traits in maize with major gene plus polygenes mixed model","volume":"19","author":"Zheng","year":"2024","journal-title":"PloS One"},{"key":"B27","doi-asserted-by":"publisher","first-page":"244","DOI":"10.1159\/000495205","article-title":"Weighted gene correlation network analysis (WGCNA) detected loss of MAGI2 promotes chronic kidney disease (CKD) by podocyte damage","volume":"51","author":"Zuo","year":"2018","journal-title":"Cell. Physiol. Biochem."}],"container-title":["Frontiers in Plant Science"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fpls.2026.1936883\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,24]],"date-time":"2026-09-24T05:37:43Z","timestamp":1790228263000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fpls.2026.1936883\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,24]]},"references-count":27,"alternative-id":["10.3389\/fpls.2026.1936883"],"URL":"https:\/\/doi.org\/10.3389\/fpls.2026.1936883","relation":{},"ISSN":["1664-462X"],"issn-type":[{"value":"1664-462X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,24]]},"article-number":"1936883"}}