{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T08:55:46Z","timestamp":1791276946480,"version":"4.1.0"},"reference-count":85,"publisher":"Oxford University Press (OUP)","issue":"9","license":[{"start":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T00:00:00Z","timestamp":1788220800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/pages\/standard-publication-reuse-rights"}],"funder":[{"name":"National Technology Innovation Center for Prataculture Special fund for innovation platform construction","award":["TICPH2025RC01"],"award-info":[{"award-number":["TICPH2025RC01"]}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32370881"],"award-info":[{"award-number":["32370881"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32500723"],"award-info":[{"award-number":["32500723"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32570963"],"award-info":[{"award-number":["32570963"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"name":"National Natural Science Foundation of Shandong Province","award":["ZR2024QC30"],"award-info":[{"award-number":["ZR2024QC30"]}]},{"name":"Postdoctoral Fellowship Program of CPSF","award":["GZC20240930"],"award-info":[{"award-number":["GZC20240930"]}]},{"name":"Taishan Scholars Program","award":["TSTP20250711"],"award-info":[{"award-number":["TSTP20250711"]}]},{"DOI":"10.13039\/501100000038","name":"Natural Sciences and Engineering Research Council of Canada","doi-asserted-by":"publisher","award":["RGPIN-582 2019-05923"],"award-info":[{"award-number":["RGPIN-582 2019-05923"]}],"id":[{"id":"10.13039\/501100000038","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Canada Research Chairs program to I.M.","id":[{"id":"https:\/\/ror.org\/0517h6h17","id-type":"ROR","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,9,1]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Seed physical dormancy is a crucial adaptive trait conferred by the water-impermeable seed coat, particularly its specialized palisade layer. Although the structural components of this barrier, including the cuticle and cell wall polysaccharides, are well characterized, the molecular mechanisms coordinating their formation and remodeling remain poorly understood. Here, using single-nucleus RNA sequencing of Medicago truncatula, we identify a regulatory module essential for palisade layer formation and the establishment of physical dormancy. We show that MtKNOX4 interacts with cytoplasmic MtSAW1\/2 proteins and facilitates their nuclear recruitment, thereby enabling the formation of functional nuclear MtKNOX4\u2013MtSAW1\/2 complexes. These complexes activate the expression of downstream target genes, including MtATT1 (encoding a fatty acid \u03c9-hydroxylase belonging to the CYP86A subfamily of cytochrome P450 enzymes) and MtPAE8 (encoding a pectin acetylesterase). Thus, the MtKNOX4\u2013MtSAW1\/2 module coordinates two parallel barrier-forming pathways involving lipid polyester remodeling and pectin deacetylation, thereby reinforcing seed coat impermeability. Loss-of-function mutations in these regulators disrupt palisade layer integrity and impair physical dormancy. Collectively, our results reveal a regulatory module in which MtKNOX4-mediated nuclear recruitment of MtSAW1\/2 links transcriptional regulation of lipid metabolism and cell wall remodeling to seed coat barrier formation, providing potential targets for tuning hard-seededness in legume crops.<\/jats:p>","DOI":"10.1093\/plcell\/koag255","type":"journal-article","created":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T08:13:16Z","timestamp":1788250396000},"source":"Crossref","is-referenced-by-count":0,"title":["A regulatory module integrates lipid metabolism and cell wall remodeling to establish the seed coat palisade barrier"],"prefix":"10.1093","volume":"38","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-6122-9488","authenticated-orcid":false,"given":"Zhichao","family":"Lu","sequence":"first","affiliation":[{"name":"The Key Laboratory of Plant Development and Environmental Adaptation Biology, Ministry of Education, Shandong Key Laboratory of Precision Molecular Crop Design and Breeding, Shandong Provincial Key Laboratory of Plant Stress Biology and Genetic Improvement, School of Life Sciences, Shandong University , Qingdao 266237 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2247-0217","authenticated-orcid":false,"given":"Lu","family":"Han","sequence":"additional","affiliation":[{"name":"The Key Laboratory of Plant Development and Environmental Adaptation Biology, Ministry of Education, Shandong Key Laboratory of Precision Molecular Crop Design and Breeding, Shandong Provincial Key Laboratory of Plant Stress Biology and Genetic Improvement, School of Life Sciences, Shandong University , Qingdao 266237 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhiqun","family":"Gu","sequence":"additional","affiliation":[{"name":"The Key Laboratory of Plant Development and Environmental Adaptation Biology, Ministry of Education, Shandong Key Laboratory of Precision Molecular Crop Design and Breeding, Shandong Provincial Key Laboratory of Plant Stress Biology and Genetic Improvement, School of Life Sciences, Shandong University , Qingdao 266237 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Annika","family":"Sonntag","sequence":"additional","affiliation":[{"name":"Department of Biology, School of Life Sciences and the Environment, Algoma University , Sault Ste Marie, ON P6A 2G4 ,","place":["Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3450-893X","authenticated-orcid":false,"given":"Isabel","family":"Molina","sequence":"additional","affiliation":[{"name":"Department of Biology, School of Life Sciences and the Environment, Algoma University , Sault Ste Marie, ON P6A 2G4 ,","place":["Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hongjiao","family":"Jiang","sequence":"additional","affiliation":[{"name":"The Key Laboratory of Plant Development and Environmental Adaptation Biology, Ministry of Education, Shandong Key Laboratory of Precision Molecular Crop Design and Breeding, Shandong Provincial Key Laboratory of Plant Stress Biology and Genetic Improvement, School of Life Sciences, Shandong University , Qingdao 266237 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Min","family":"Wang","sequence":"additional","affiliation":[{"name":"The Key Laboratory of Plant Development and Environmental Adaptation Biology, Ministry of Education, Shandong Key Laboratory of Precision Molecular Crop Design and Breeding, Shandong Provincial Key Laboratory of Plant Stress Biology and Genetic Improvement, School of Life Sciences, Shandong University , Qingdao 266237 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0004-0083-9540","authenticated-orcid":false,"given":"Xue","family":"Zhang","sequence":"additional","affiliation":[{"name":"The Key Laboratory of Plant Development and Environmental Adaptation Biology, Ministry of Education, Shandong Key Laboratory of Precision Molecular Crop Design and Breeding, Shandong Provincial Key Laboratory of Plant Stress Biology and Genetic Improvement, School of Life Sciences, Shandong University , Qingdao 266237 ,","place":["China"]},{"name":"Advanced Medical Research Institute, Cheeloo College of Medicine, Shandong University , Jinan 250012 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5935-2727","authenticated-orcid":false,"given":"Hongfeng","family":"Wang","sequence":"additional","affiliation":[{"name":"The Key Laboratory of Plant Development and Environmental Adaptation Biology, Ministry of Education, Shandong Key Laboratory of Precision Molecular Crop Design and Breeding, Shandong Provincial Key Laboratory of Plant Stress Biology and Genetic Improvement, School of Life Sciences, Shandong University , Qingdao 266237 ,","place":["China"]},{"name":"Shandong Peanut Research Institute , Qingdao 266199 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4323-4697","authenticated-orcid":false,"given":"Yiteng","family":"Xu","sequence":"additional","affiliation":[{"name":"The Key Laboratory of Plant Development and Environmental Adaptation Biology, Ministry of Education, Shandong Key Laboratory of Precision Molecular Crop Design and Breeding, Shandong Provincial Key Laboratory of Plant Stress Biology and Genetic Improvement, School of Life Sciences, Shandong University , Qingdao 266237 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University , Qingdao 266237 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9915-0321","authenticated-orcid":false,"given":"Maofeng","family":"Chai","sequence":"additional","affiliation":[{"name":"Grassland Agri-Husbandry Research Center, College of Grassland Science, Qingdao Agricultural University , Qingdao 266109 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6835-7917","authenticated-orcid":false,"given":"Zeng-Yu","family":"Wang","sequence":"additional","affiliation":[{"name":"Grassland Agri-Husbandry Research Center, College of Grassland Science, Qingdao Agricultural University , Qingdao 266109 ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9805-5741","authenticated-orcid":false,"given":"Kirankumar S","family":"Mysore","sequence":"additional","affiliation":[{"name":"Institute of 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