{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,17]],"date-time":"2026-10-17T00:10:38Z","timestamp":1792195838368,"version":"4.4.0"},"reference-count":56,"publisher":"MDPI AG","issue":"17","license":[{"start":{"date-parts":[[2026,8,31]],"date-time":"2026-08-31T00:00:00Z","timestamp":1788134400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"China Agriculture Research System of MOF and MARA","award":["CARS-25"],"award-info":[{"award-number":["CARS-25"]}]},{"award":["CARS-25"],"award-info":[{"award-number":["CARS-25"]}],"id":[{"id":"https:\/\/ror.org\/03p5eky76","id-type":"ROR","asserted-by":"publisher"}]},{"name":"The Natural Science Foundation of Liaoning Province of China","award":["2025-MS-327"],"award-info":[{"award-number":["2025-MS-327"]}]},{"DOI":"10.13039\/501100012421","name":"The Agricultural Science and Technology Innovation Program","doi-asserted-by":"crossref","award":["ASTIP"],"award-info":[{"award-number":["ASTIP"]}],"id":[{"id":"10.13039\/501100012421","id-type":"DOI","asserted-by":"crossref"}]},{"name":"Doctoral Startup Project of the Natural Science Foundation of Liaoning Province","award":["2025-BS-0924"],"award-info":[{"award-number":["2025-BS-0924"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Plants"],"abstract":"<jats:p>The homeodomain\u2013leucine zipper IV (HD-Zip IV) transcription factor subfamily plays essential roles in epidermal development, cuticle formation, lipid metabolism, and environmental adaptation in plants. Despite its biological importance, the HD-Zip IV family has not been systematically characterized in apple (Malus domestica). Here, we identified 17 apple HD-Zip IV genes and named them MdHDZIV1\u2013MdHDZIV17 based on their locations on the chromosomes. The 17 genes showed a nonuniform distribution on eight chromosomes, while the occurrence of both tandem and segmental duplications indicated that family expansion involved more than one duplication mechanism. All MdHDZIV proteins contained the conserved HD, LZ, START, and SAD domains but lacked the MEKHLA domain, consistent with typical HD-Zip IV structural features. Phylogenetic analysis classified MdHDZIV proteins into five groups together with HD-Zip IV members from Arabidopsis thaliana and rice, indicating evolutionary conservation of this subfamily. Collinearity and Ka\/Ks analyses revealed that duplicated MdHDZIV gene pairs were mainly subjected to purifying selection. Promoter scanning revealed diverse cis-regulatory motifs associated with hormonal signaling, environmental stress, light response, and epidermal regulation, including ABRE, ARE, W-box, MYC, G-box, and L1-box motifs. Integration of transcriptomic profiling with qRT-PCR validation revealed pronounced tissue-dependent differences in the expression of MdHDZIV genes in leaf, fruit skin, and branch bark. Under PEG6000-induced osmotic stress and NaCl-induced salt stress, 10 candidate MdHDZIV genes displayed gene-specific and stress type-specific expression patterns, with MdHDZIV3 showing strong induction under PEG6000 treatment. Functional validation in apple calli showed that MdHDZIV3 overexpression enhanced PEG tolerance, increased fresh weight, elevated SOD and POD activities, and reduced MDA accumulation under osmotic stress. These findings provide a genome-wide framework for understanding the apple HD-Zip IV gene family.<\/jats:p>","DOI":"10.3390\/plants15172670","type":"journal-article","created":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T01:05:26Z","timestamp":1788224726000},"page":"2670","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Genome-Wide Characterization of the Apple HD-Zip IV Gene Family and Functional Validation of MdHDZIV3 Under PEG-Induced Osmotic Stress"],"prefix":"10.3390","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0009-0001-6909-1081","authenticated-orcid":false,"given":"An","family":"Yang","sequence":"first","affiliation":[{"name":"Key Laboratory of Horticulture Crops Germplasm Resources Utilization, Research Institute of Pomology, Chinese Academy of Agricultural Sciences (CAAS), Xingcheng 125100, China"},{"name":"Key Laboratory of Horticultural Crops Germplasm Resources Utilization, Ministry of Agriculture and Rural Affairs of the People\u2019s Republic of China, Xingcheng 125100, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xuqing","family":"Tong","sequence":"additional","affiliation":[{"name":"Key Laboratory of Horticulture Crops Germplasm Resources Utilization, Research Institute of Pomology, Chinese Academy of Agricultural Sciences (CAAS), Xingcheng 125100, China"},{"name":"Key Laboratory of Horticultural Crops Germplasm Resources Utilization, Ministry of Agriculture and Rural Affairs of the People\u2019s Republic of China, Xingcheng 125100, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Songya","family":"Ma","sequence":"additional","affiliation":[{"name":"Key Laboratory of Horticulture Crops Germplasm Resources Utilization, Research Institute of Pomology, Chinese Academy of Agricultural Sciences (CAAS), Xingcheng 125100, China"},{"name":"Key Laboratory of Horticultural Crops Germplasm Resources Utilization, Ministry of Agriculture and Rural Affairs of the People\u2019s Republic of China, Xingcheng 125100, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chenglong","family":"Liu","sequence":"additional","affiliation":[{"name":"Key Laboratory of Horticulture Crops Germplasm Resources Utilization, Research Institute of Pomology, Chinese Academy of Agricultural Sciences (CAAS), Xingcheng 125100, China"},{"name":"Key Laboratory of Horticultural Crops Germplasm Resources Utilization, Ministry of Agriculture and Rural Affairs of the People\u2019s Republic of China, Xingcheng 125100, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Bichen","family":"Wang","sequence":"additional","affiliation":[{"name":"Key Laboratory of Horticulture Crops Germplasm Resources Utilization, Research Institute of Pomology, Chinese Academy of Agricultural Sciences (CAAS), Xingcheng 125100, China"},{"name":"Key Laboratory of Horticultural Crops Germplasm Resources Utilization, Ministry of Agriculture and Rural Affairs of the People\u2019s Republic of China, Xingcheng 125100, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lin","family":"Guo","sequence":"additional","affiliation":[{"name":"Key Laboratory of Horticulture Crops Germplasm Resources Utilization, Research Institute of Pomology, Chinese Academy of Agricultural Sciences (CAAS), Xingcheng 125100, China"},{"name":"Key Laboratory of Horticultural Crops Germplasm Resources Utilization, Ministry of Agriculture and Rural Affairs of the People\u2019s Republic of China, Xingcheng 125100, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hui","family":"Li","sequence":"additional","affiliation":[{"name":"Key Laboratory of Horticulture Crops Germplasm Resources Utilization, Research Institute of Pomology, Chinese Academy of Agricultural Sciences (CAAS), Xingcheng 125100, China"},{"name":"Key Laboratory of Horticultural Crops Germplasm Resources Utilization, Ministry of Agriculture and Rural Affairs of the People\u2019s Republic of China, Xingcheng 125100, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zitong","family":"He","sequence":"additional","affiliation":[{"name":"Key Laboratory of Horticulture Crops Germplasm Resources Utilization, Research Institute of Pomology, Chinese Academy of Agricultural Sciences (CAAS), Xingcheng 125100, China"},{"name":"Key Laboratory of Horticultural Crops Germplasm Resources Utilization, Ministry of Agriculture and Rural Affairs of the People\u2019s Republic of China, Xingcheng 125100, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Liqun","family":"Kang","sequence":"additional","affiliation":[{"name":"Key Laboratory of Horticulture Crops Germplasm Resources Utilization, Research Institute of Pomology, Chinese Academy of Agricultural Sciences (CAAS), Xingcheng 125100, China"},{"name":"Key Laboratory of Horticultural Crops Germplasm Resources Utilization, Ministry of Agriculture and Rural Affairs of the People\u2019s Republic of China, Xingcheng 125100, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaolei","family":"Han","sequence":"additional","affiliation":[{"name":"Key Laboratory of Horticulture Crops Germplasm Resources Utilization, Research Institute of Pomology, Chinese Academy of Agricultural Sciences (CAAS), Xingcheng 125100, China"},{"name":"Key Laboratory of Horticultural Crops Germplasm Resources Utilization, Ministry of Agriculture and Rural Affairs of the People\u2019s Republic of China, Xingcheng 125100, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Caixia","family":"Zhang","sequence":"additional","affiliation":[{"name":"Key Laboratory of Horticulture Crops Germplasm Resources Utilization, Research Institute of Pomology, Chinese Academy of Agricultural Sciences (CAAS), Xingcheng 125100, China"},{"name":"Key Laboratory of Horticultural Crops Germplasm Resources Utilization, Ministry of Agriculture and Rural Affairs of the People\u2019s Republic of China, Xingcheng 125100, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,8,31]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1104\/pp.113.222737","article-title":"The formation and function of plant cuticles","volume":"163","author":"Yeats","year":"2013","journal-title":"Plant Physiol."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Xue, D., Zhang, X., Lu, X., Chen, G., and Chen, Z.H. 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