{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,14]],"date-time":"2026-10-14T19:21:54Z","timestamp":1792005714379,"version":"4.4.0"},"reference-count":41,"publisher":"Wiley","issue":"10","license":[{"start":{"date-parts":[[2024,6,17]],"date-time":"2024-06-17T00:00:00Z","timestamp":1718582400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/onlinelibrary.wiley.com\/termsAndConditions#vor"}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Plant Cell &amp; Environment"],"published-print":{"date-parts":[[2024,10]]},"abstract":"<jats:title>Abstract<\/jats:title><jats:p>Proanthocyanidins (PAs) are important\u00a0metabolites that enhance freezing tolerance of plants. <jats:italic>Actinidia arguta<\/jats:italic>, especially freezing\u2010tolerant germplasms, accumulate abundant PAs in dormant shoots and thereby enhance freezing tolerance, but the underlying mechanism is unknown. In this study, we used two <jats:italic>A. arguta<\/jats:italic> with contrasting cold\u2010resistant phenotypes,\u00a0KL and RB, to explore the mechanisms in response to cold tolerance. We determined that a leucoanthocyanidin reductase gene (<jats:italic>AaLAR1<\/jats:italic>) was more highly expressed in freezing\u2010tolerant KL than in freezing\u2010sensitive RB. Moreover, overexpressing <jats:italic>AaLAR1<\/jats:italic> in kiwifruit promoted PAs biosynthesis and enhanced cold tolerance. The <jats:italic>AaLAR1<\/jats:italic> promoters of various <jats:italic>A. arguta<\/jats:italic> germplasms differ due to the presence of a 60\u2010bp deletion in cold\u2010tolerant genotypes that forms a functional binding site for MYC\u2010type transcription factor. Yeast one\u2010hybrid and two\u2010hybrid, dual\u2010luciferase reporter, bimolecular fluorescence complementation\u00a0and coimmunoprecipitation assays indicated that the AaMYC2a binds to the MYC\u2010core <jats:italic>cis<\/jats:italic>\u2010element in the <jats:italic>AaLAR1<\/jats:italic> promoter with the assistance of AaMYB5a, thereby promoting PAs accumulation in the shoots of cold\u2010tolerant kiwifruit. We conclude that the variation in the <jats:italic>AaLAR1<\/jats:italic> promoter and the AaMYC2a\u2013AaMYB5a\u2013<jats:italic>AaLAR1<\/jats:italic> module shape freezing tolerance in <jats:italic>A. arguta<\/jats:italic>. The identification of a key structural variation in the <jats:italic>AaLAR1<\/jats:italic> promoter offers a new target for resistance breeding of kiwifruit.<\/jats:p>","DOI":"10.1111\/pce.15003","type":"journal-article","created":{"date-parts":[[2024,6,17]],"date-time":"2024-06-17T11:36:33Z","timestamp":1718624193000},"page":"4048-4066","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":21,"title":["A structural variation in the promoter of the leucoanthocyanidin reductase gene <i>AaLAR1<\/i> enhances freezing tolerance by modulating proanthocyanidin accumulation in kiwifruit (<i>Actinidia arguta<\/i>)"],"prefix":"10.1111","volume":"47","author":[{"given":"Shihang","family":"Sun","sequence":"first","affiliation":[{"name":"National Key Laboratory for Germplasm Innovation and Utilization of Horticultural Crops Zhengzhou Fruit Research Institute, Chinese Academy of Agricultural Sciences Zhengzhou China"},{"name":"Zhongyuan Research Center Chinese Academy of Agricultural Sciences Xinxiang China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiujuan","family":"Qi","sequence":"additional","affiliation":[{"name":"National Key Laboratory for Germplasm Innovation and Utilization of Horticultural Crops Zhengzhou Fruit Research Institute, Chinese Academy of Agricultural Sciences Zhengzhou China"},{"name":"Zhongyuan Research Center Chinese Academy of Agricultural Sciences Xinxiang China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhenzhen","family":"Zhang","sequence":"additional","affiliation":[{"name":"National Key Laboratory for Germplasm Innovation and Utilization of Horticultural Crops Zhengzhou Fruit Research Institute, Chinese Academy of Agricultural Sciences Zhengzhou China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Leiming","family":"Sun","sequence":"additional","affiliation":[{"name":"National Key Laboratory for Germplasm Innovation and Utilization of Horticultural Crops Zhengzhou Fruit Research Institute, Chinese Academy of Agricultural Sciences Zhengzhou China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ran","family":"Wang","sequence":"additional","affiliation":[{"name":"National Key Laboratory for Germplasm Innovation and Utilization of Horticultural Crops Zhengzhou Fruit Research Institute, Chinese Academy of Agricultural Sciences Zhengzhou China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yukuo","family":"Li","sequence":"additional","affiliation":[{"name":"National Key Laboratory for Germplasm Innovation and Utilization of Horticultural Crops Zhengzhou Fruit Research Institute, Chinese Academy of Agricultural Sciences Zhengzhou China"},{"name":"Zhongyuan Research Center Chinese Academy of Agricultural Sciences Xinxiang China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jinyong","family":"Chen","sequence":"additional","affiliation":[{"name":"National Key Laboratory for Germplasm Innovation and Utilization of Horticultural Crops Zhengzhou Fruit Research Institute, Chinese Academy of Agricultural Sciences Zhengzhou China"},{"name":"Zhongyuan Research Center Chinese Academy of Agricultural Sciences Xinxiang China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hong","family":"Gu","sequence":"additional","affiliation":[{"name":"National Key Laboratory for Germplasm Innovation and Utilization of Horticultural Crops Zhengzhou Fruit Research Institute, Chinese Academy of Agricultural Sciences Zhengzhou China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jinbao","family":"Fang","sequence":"additional","affiliation":[{"name":"National Key Laboratory for Germplasm Innovation and Utilization of Horticultural Crops Zhengzhou Fruit Research Institute, Chinese Academy of Agricultural Sciences Zhengzhou China"},{"name":"Zhongyuan Research Center Chinese Academy of Agricultural Sciences Xinxiang China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2023-7657","authenticated-orcid":false,"given":"Miaomiao","family":"Lin","sequence":"additional","affiliation":[{"name":"National Key Laboratory for Germplasm Innovation and Utilization of Horticultural Crops Zhengzhou Fruit Research Institute, Chinese Academy of Agricultural Sciences Zhengzhou China"},{"name":"Zhongyuan Research Center Chinese Academy of Agricultural Sciences Xinxiang China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2024,6,17]]},"reference":[{"key":"e_1_2_9_2_1","doi-asserted-by":"publisher","DOI":"10.1104\/pp.109.146985"},{"key":"e_1_2_9_3_1","doi-asserted-by":"publisher","DOI":"10.1111\/tpj.14050"},{"key":"e_1_2_9_4_1","doi-asserted-by":"publisher","DOI":"10.1007\/s11103-011-9827-4"},{"key":"e_1_2_9_5_1","doi-asserted-by":"publisher","DOI":"10.1021\/acs.jafc.6b05609"},{"key":"e_1_2_9_6_1","doi-asserted-by":"publisher","DOI":"10.15252\/embj.201899819"},{"key":"e_1_2_9_7_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469-8137.2004.01217.x"},{"key":"e_1_2_9_8_1","doi-asserted-by":"publisher","DOI":"10.1186\/1471-2229-12-22"},{"key":"e_1_2_9_9_1","doi-asserted-by":"publisher","DOI":"10.1093\/mp\/ssp118"},{"key":"e_1_2_9_10_1","doi-asserted-by":"publisher","DOI":"10.1111\/pce.13609"},{"key":"e_1_2_9_11_1","first-page":"1","article-title":"Peach MYB7 activates transcription of the proanthocyanidin pathway gene encoding leucoanthocyanidin reductase, but not anthocyanidin reductase","volume":"6","author":"Hui Z.","year":"2015","journal-title":"Frontiers in Plant Science"},{"key":"e_1_2_9_12_1","doi-asserted-by":"publisher","DOI":"10.1104\/pp.16.01962"},{"key":"e_1_2_9_13_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.envexpbot.2018.07.027"},{"key":"e_1_2_9_14_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.plantsci.2018.02.009"},{"key":"e_1_2_9_15_1","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/eru213"},{"key":"e_1_2_9_16_1","doi-asserted-by":"publisher","DOI":"10.1139\/cjfr-2017-0377"},{"key":"e_1_2_9_17_1","doi-asserted-by":"publisher","DOI":"10.3390\/ijms22063026"},{"key":"e_1_2_9_18_1","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/erab401"},{"key":"e_1_2_9_19_1","doi-asserted-by":"publisher","DOI":"10.1186\/1471-2229-13-202"},{"key":"e_1_2_9_20_1","doi-asserted-by":"publisher","DOI":"10.1006\/meth.2001.1262"},{"key":"e_1_2_9_21_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.plantsci.2016.01.007"},{"key":"e_1_2_9_22_1","doi-asserted-by":"publisher","DOI":"10.1104\/pp.109.139071"},{"key":"e_1_2_9_23_1","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2021.677611"},{"key":"e_1_2_9_24_1","doi-asserted-by":"publisher","DOI":"10.1080\/10715762.2020.1843651"},{"key":"e_1_2_9_25_1","first-page":"682","article-title":"Molecular cloning and functional identification of proanthocyanidin synthesis related genes VcLAR and VcANR of blueberry","volume":"35","author":"Song Y.","year":"2019","journal-title":"Jiangsu Journal of Agricultural Sciences"},{"key":"e_1_2_9_26_1","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2021.628969"},{"key":"e_1_2_9_27_1","doi-asserted-by":"publisher","DOI":"10.3390\/plants9040515"},{"key":"e_1_2_9_28_1","doi-asserted-by":"publisher","DOI":"10.1186\/s12870-021-03152-w"},{"key":"e_1_2_9_29_1","doi-asserted-by":"publisher","DOI":"10.1104\/pp.108.131862"},{"key":"e_1_2_9_30_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.envexpbot.2022.104978"},{"key":"e_1_2_9_31_1","doi-asserted-by":"publisher","DOI":"10.1111\/tpj.13487"},{"key":"e_1_2_9_32_1","doi-asserted-by":"crossref","unstructured":"Wang \u00a0T. \u00a0Atkinson \u00a0R.\u00a0&\u00a0Janssen \u00a0B.\u00a0(2007)\u00a0The choice of\u00a0Agrobacterium\u00a0strain for transformation of kiwifruit. In:\u00a0Ferguson \u00a0A.R. \u00a0Hewett \u00a0E.W. \u00a0Gunson \u00a0F.A.\u00a0&\u00a0Hale \u00a0C.N. (Eds.)\u00a0Proceedings of the 6th International Symposium on Kiwifruit Vols. 1 and 2.\u00a0Belgium:International Society for Horticultural Science. pp.\u00a0227\u2013232.","DOI":"10.17660\/ActaHortic.2007.753.26"},{"key":"e_1_2_9_33_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.hpj.2020.08.006"},{"key":"e_1_2_9_34_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-3040.2012.02523.x"},{"key":"e_1_2_9_35_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.plantsci.2020.110473"},{"key":"e_1_2_9_36_1","doi-asserted-by":"publisher","DOI":"10.1093\/pcp\/pcn009"},{"key":"e_1_2_9_37_1","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/err425"},{"key":"e_1_2_9_38_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.gene.2019.02.037"},{"key":"e_1_2_9_39_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.devcel.2017.09.024"},{"key":"e_1_2_9_40_1","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/ery335"},{"key":"e_1_2_9_41_1","doi-asserted-by":"publisher","DOI":"10.1007\/s10535-016-0663-x"},{"key":"e_1_2_9_42_1","doi-asserted-by":"publisher","DOI":"10.1111\/pbi.14016"}],"container-title":["Plant, Cell &amp; Environment"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1111\/pce.15003","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,11,19]],"date-time":"2024-11-19T11:57:56Z","timestamp":1732017476000},"score":1,"resource":{"primary":{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/10.1111\/pce.15003"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2024,6,17]]},"references-count":41,"journal-issue":{"issue":"10","published-print":{"date-parts":[[2024,10]]}},"alternative-id":["10.1111\/pce.15003"],"URL":"https:\/\/doi.org\/10.1111\/pce.15003","archive":["Portico"],"relation":{},"ISSN":["0140-7791","1365-3040"],"issn-type":[{"value":"0140-7791","type":"print"},{"value":"1365-3040","type":"electronic"}],"subject":[],"published":{"date-parts":[[2024,6,17]]},"assertion":[{"value":"2023-12-03","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2024-05-27","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2024-06-17","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}]}}