{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,13]],"date-time":"2026-10-13T05:45:18Z","timestamp":1791870318949,"version":"4.3.4"},"reference-count":31,"publisher":"Wiley","license":[{"start":{"date-parts":[[2026,10,12]],"date-time":"2026-10-12T00:00:00Z","timestamp":1791763200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2026,10,12]],"date-time":"2026-10-12T00:00:00Z","timestamp":1791763200000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/501100003725","name":"National Research Foundation of Korea","doi-asserted-by":"publisher","award":["2020R1A2C108800"],"award-info":[{"award-number":["2020R1A2C108800"]}],"id":[{"id":"10.13039\/501100003725","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100003725","name":"National Research Foundation of Korea","doi-asserted-by":"publisher","award":["RS\u20102023\u201000208020"],"award-info":[{"award-number":["RS\u20102023\u201000208020"]}],"id":[{"id":"10.13039\/501100003725","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100003725","name":"National Research Foundation of Korea","doi-asserted-by":"publisher","award":["NRF\u20102022K2A9A2A06037703"],"award-info":[{"award-number":["NRF\u20102022K2A9A2A06037703"]}],"id":[{"id":"10.13039\/501100003725","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["scijournals.onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["J Sci Food Agric"],"abstract":"Abstract<\/jats:title>\n \n BACKGROUND<\/jats:title>\n \n Downy mildew, caused by\n Pseudoperonospora cubensis<\/jats:italic>\n , is one of the most destructive diseases affecting cucumber and remains difficult to manage sustainably through fungicide\u2010dependent control strategies. Although several resistance loci have been reported, many previously mapped regions remain too broad for efficient marker\u2010assisted selection or reliable candidate\u2010gene prioritisation. This study aimed to refine three TH118FLM\u2010derived downy mildew resistance loci and to identify biologically plausible candidate genes associated with resistance by integrating advanced recombinant populations, QTL\u2010seq, parent\u2013bulk concordance analysis, and candidate\u2010gene evaluation.\n <\/jats:p>\n <\/jats:sec>\n \n RESULTS<\/jats:title>\n \n Two recombinant inbred line populations, dm2.1\u2010F\n 7<\/jats:sub>\n and dm2.2\u2010F\n 7<\/jats:sub>\n , and one near\u2010isogenic\/backcross\u2010derived population, dm5.1\u2010BC\n 4<\/jats:sub>\n F\n 2<\/jats:sub>\n , were developed from a cross between TH118FLM and WMEJ. Phenotypic distributions in all three populations showed a higher frequency of susceptible disease\u2010index classes, supporting the involvement of multiple resistance\u2010associated quantitative loci rather than a single dominant resistance factor. QTL\u2010seq reduced the\n dm2.1<\/jats:italic>\n ,\n dm2.2<\/jats:italic>\n , and\n dm5.1<\/jats:italic>\n regions to 333, 296, and 694\u2009kb, respectively. Subsequent parent\u2013bulk concordance analysis of SNPs and INDELs further refined these loci to 61.7, 84.7, and 80.0\u2009kb, containing seven, ten, and five annotated genes, respectively. By integrating positional, sequence, transcriptional, and predicted structural evidence,\n CsACO<\/jats:italic>\n ,\n CsCA<\/jats:italic>\n , and\n CsSTPK<\/jats:italic>\n were prioritised as the candidate genes for\n dm2.1<\/jats:italic>\n ,\n dm2.2<\/jats:italic>\n , and\n dm5.1<\/jats:italic>\n , respectively. Expression and physiological analyses further showed that these candidates responded to pathogen infection and ethephon treatment, while the resistant background exhibited reduced stomatal aperture and altered stomatal conductance following ethephon application, supporting a testable ethylene\u2010associated stomatal\u2010defence hypothesis.\n <\/jats:p>\n <\/jats:sec>\n \n CONCLUSION<\/jats:title>\n \n This study refined three cucumber downy mildew resistance loci into compact, marker\u2010ready intervals and identified\n CsACO<\/jats:italic>\n ,\n CsCA<\/jats:italic>\n , and\n CsSTPK<\/jats:italic>\n as biologically credible candidate genes. The findings provide practical genomic resources for marker\u2010assisted cucumber resistance breeding and a testable biological framework, suggesting that TH118FLM\u2010derived resistance may involve ethylene\u2010associated stomatal defence. \u00a9 2026 The Author(s).\n Journal of the Science of Food and Agriculture<\/jats:italic>\n published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.\n <\/jats:p>\n <\/jats:sec>","DOI":"10.1002\/jsfa.71159","type":"journal-article","created":{"date-parts":[[2026,10,13]],"date-time":"2026-10-13T05:00:30Z","timestamp":1791867630000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Fine mapping of cucumber downy mildew resistance loci identifies candidate genes linked to ethylene\u2010associated stomatal defence"],"prefix":"10.1002","author":[{"given":"Mahdi Badri","family":"Anarjan","sequence":"first","affiliation":[{"name":"Plant Genomics Laboratory, Department of Bio\u2010resource Engineering, College of Life Sciences Sejong University Seoul Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Saeed","family":"Omrani","sequence":"additional","affiliation":[{"name":"Plant Genomics Laboratory, Department of Bio\u2010resource Engineering, College of Life Sciences Sejong University Seoul Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chunying","family":"Zhang","sequence":"additional","affiliation":[{"name":"Plant Genomics Laboratory, Department of Bio\u2010resource Engineering, College of Life Sciences Sejong University Seoul Republic of Korea"},{"name":"College of Agronomy Jilin Agricultural University Changchun China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shahida","family":"Begum","sequence":"additional","affiliation":[{"name":"Plant Genomics Laboratory, Department of Bio\u2010resource Engineering, College of Life Sciences Sejong University Seoul Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Khin Thanda","family":"Win","sequence":"additional","affiliation":[{"name":"Plant Genomics Laboratory, Department of Bio\u2010resource Engineering, College of Life Sciences Sejong University Seoul Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Haisu","family":"Li","sequence":"additional","affiliation":[{"name":"Plant Genomics Laboratory, Department of Bio\u2010resource Engineering, College of Life Sciences Sejong University Seoul Republic of Korea"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chunhua","family":"Wang","sequence":"additional","affiliation":[{"name":"College of Horticulture Hunan Agricultural University Changsha China"},{"name":"Key Laboratory for Evaluation and Utilization of Gene Resources of Horticultural Crops (Vegetables, Tea, etc.) Ministry of Agriculture and Rural Affairs of China Changsha China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tao","family":"Wu","sequence":"additional","affiliation":[{"name":"College of Horticulture Hunan Agricultural University Changsha China"},{"name":"Key Laboratory for Evaluation and Utilization of Gene Resources of Horticultural Crops (Vegetables, Tea, etc.) Ministry of Agriculture and Rural Affairs of China Changsha China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7684-2325","authenticated-orcid":false,"given":"Sanghyeob","family":"Lee","sequence":"additional","affiliation":[{"name":"Plant Genomics Laboratory, Department of Bio\u2010resource Engineering, College of Life Sciences Sejong University Seoul Republic of Korea"},{"name":"Plant Engineering Research Institute Sejong University Seoul Republic of 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