{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T10:07:26Z","timestamp":1790158046774,"version":"4.1.0"},"reference-count":40,"publisher":"MDPI AG","issue":"19","license":[{"start":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T00:00:00Z","timestamp":1790121600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Scientific Research Operating Expenses Project for Provincial-Level Research Institutes of Heilongjiang Province","award":["CZKYF2026-1-B013"],"award-info":[{"award-number":["CZKYF2026-1-B013"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Int. J. Mol. Sci.","IJMS"],"abstract":"<jats:p>As a primary apple rootstock extensively cultivated across Northeast China, Malus baccata (L.) Borkh. constantly suffers from low temperature, drought and salt damage throughout its growth cycle. NAC transcription factors take critical parts in coordinating plant adaptation to various adverse environments, yet relevant functional research on this woody rootstock species remains limited. In this work, a NAC family gene designated MbNAC40 was cloned from M. baccata, and its biological function was explored via heterologous transformation in Arabidopsis thaliana. Sequence analysis revealed that the open reading frame of MbNAC40 spans 1467 bp and encodes a 488-amino-acid protein targeted to the cell nucleus. After 24 h cold acclimation at 4 \u00b0C, plants were subjected to \u22124 \u00b0C freezing stress for 12 h, then recovered at 25 \u00b0C for 3 d. Transgenic Arabidopsis lines overexpressing MbNAC40 exhibited much stronger cold tolerance relative to wild-type plants. Transgenic materials retained more chlorophyll and synthesized more osmoprotective proline, accompanied by enhanced activity of three antioxidant enzymes (SOD, POD and CAT). Meanwhile, multiple stress-marker genes (AtCAT1, AtCSD1, AtAPX6, and AtP5CS1) associated with ROS and osmotic adjustment were significantly activated in overexpression lines. Taken together, these results confirm that MbNAC40 acts as a positive modulator of cold resistance in Arabidopsis, which provides a valuable gene resource for molecular breeding of cold-resistant M. baccata rootstock varieties.<\/jats:p>","DOI":"10.3390\/ijms27198480","type":"journal-article","created":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T09:20:46Z","timestamp":1790155246000},"page":"8480","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Overexpression of a Malus baccata NAC Transcription Factor Gene MbNAC40 Increases Cold Tolerance in Arabidopsis"],"prefix":"10.3390","volume":"27","author":[{"given":"Kuibao","family":"Jiao","sequence":"first","affiliation":[{"name":"Heilongjiang Modern Agriculture Research Institute (Institute of Rural Revitalization Technology, Heilongjiang Academy of Agricultural Sciences), Harbin 150028, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"conceptualization","vocabulary":"credit"},{"role":"methodology","vocabulary":"credit"},{"role":"formal-analysis","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"}]},{"given":"Yuze","family":"Li","sequence":"additional","affiliation":[{"name":"Heilongjiang Modern Agriculture Research Institute (Institute of Rural Revitalization Technology, Heilongjiang Academy of Agricultural Sciences), Harbin 150028, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"software","vocabulary":"credit"},{"role":"validation","vocabulary":"credit"}]},{"given":"Penghui","family":"Song","sequence":"additional","affiliation":[{"name":"Heilongjiang Modern Agriculture Research Institute (Institute of Rural Revitalization Technology, Heilongjiang Academy of Agricultural Sciences), Harbin 150028, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"software","vocabulary":"credit"},{"role":"validation","vocabulary":"credit"}]},{"given":"Wenhui","family":"Li","sequence":"additional","affiliation":[{"name":"Key Laboratory of Biology and Genetic Improvement of Horticultural Crops (Northeast Region), Ministry of Agriculture and Rural Affairs, National\u2013Local Joint Engineering Research Center for Development and Utilization of Small Fruits in Cold Regions, College of Horticulture & Landscape Architecture, Northeast Agricultural University, Harbin 150030, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"writing-review-editing","vocabulary":"credit"}]},{"given":"Zhe","family":"Zhang","sequence":"additional","affiliation":[{"name":"Heilongjiang Modern Agriculture Research Institute (Institute of Rural Revitalization Technology, Heilongjiang Academy of Agricultural Sciences), Harbin 150028, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"writing-original-draft","vocabulary":"credit"}]},{"given":"Teng","family":"Wang","sequence":"additional","affiliation":[{"name":"Heilongjiang Modern Agriculture Research Institute (Institute of Rural Revitalization Technology, Heilongjiang Academy of Agricultural Sciences), Harbin 150028, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"visualization","vocabulary":"credit"}]},{"given":"Xingguo","family":"Li","sequence":"additional","affiliation":[{"name":"Key Laboratory of Biology and Genetic Improvement of Horticultural Crops (Northeast Region), Ministry of Agriculture and Rural Affairs, National\u2013Local Joint Engineering Research Center for Development and Utilization of Small Fruits in Cold Regions, College of Horticulture & Landscape Architecture, Northeast Agricultural University, Harbin 150030, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"investigation","vocabulary":"credit"},{"role":"resources","vocabulary":"credit"},{"role":"supervision","vocabulary":"credit"},{"role":"project-administration","vocabulary":"credit"},{"role":"funding-acquisition","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0132-3018","authenticated-orcid":false,"given":"Deguo","family":"Han","sequence":"additional","affiliation":[{"name":"Key Laboratory of Biology and Genetic Improvement of Horticultural Crops (Northeast Region), Ministry of Agriculture and Rural Affairs, National\u2013Local Joint Engineering Research Center for Development and Utilization of Small Fruits in Cold Regions, College of Horticulture & Landscape Architecture, Northeast Agricultural University, Harbin 150030, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"data-curation","vocabulary":"credit"},{"role":"funding-acquisition","vocabulary":"credit"}]}],"member":"1968","published-online":{"date-parts":[[2026,9,23]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"571","DOI":"10.1146\/annurev.arplant.50.1.571","article-title":"PLANT COLD ACCLIMATION: Freezing Tolerance Genes and Regulatory Mechanisms","volume":"50","author":"Thomashow","year":"1999","journal-title":"Annu. 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