{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,16]],"date-time":"2026-10-16T10:50:58Z","timestamp":1792147858527,"version":"4.4.0"},"reference-count":44,"publisher":"Wiley","issue":"6","license":[{"start":{"date-parts":[[2026,9,30]],"date-time":"2026-09-30T00:00:00Z","timestamp":1790726400000},"content-version":"vor","delay-in-days":29,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T00:00:00Z","timestamp":1788220800000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32572844"],"award-info":[{"award-number":["32572844"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["The Plant Journal"],"published-print":{"date-parts":[[2026,9]]},"abstract":"<jats:title>SUMMARY<\/jats:title>\n                  <jats:p>\n                    Previously, the crucial role of calcium\u2010dependent protein kinase 3 (CPK3) in Cu\n                    <jats:sup>2+<\/jats:sup>\n                    \u2010triggered immunity (CuTI) has been established in Arabidopsis. CPK3 promotes the phosphorylation of basic helix\u2013loop\u2013helix 107 (bHLH107), thereby activating the expression of\n                    <jats:italic>1\u2010aminocyclopropanecarboxylic acid synthesis 8<\/jats:italic>\n                    (\n                    <jats:italic>ACS8<\/jats:italic>\n                    ) and downstream defense responses. However, the precise mechanisms underlying the fine\u2010tuning of CPK3\u2010mediated phosphorylation of bHLH107 remain unclear. To elucidate the mechanism underlying the inactivation of the CPK3\u2010bHLH107 module, we identified a 14\u20103\u20103 protein, GRF1 (GF14\u03c7), among candidate bHLH107 interactors. Notably, the interaction between GRF1 and bHLH107 was weaker following Cu\n                    <jats:sup>2+<\/jats:sup>\n                    treatment than under control conditions. In\n                    <jats:italic>grf1<\/jats:italic>\n                    mutant plants, the transcript level of\n                    <jats:italic>ACS8<\/jats:italic>\n                    was slightly increased, concomitant with a modest increase in resistance to\n                    <jats:italic>Pst<\/jats:italic>\n                    DC3000. GRF1 interacts with both CPK3 and bHLH107 to maintain an \u2018OFF\u2019 state. Cu\n                    <jats:sup>2+<\/jats:sup>\n                    mediates the phosphorylation of CPK3, attenuating its interaction with GRF1 and promoting its dissociation from GRF1. Subsequently, CPK3 phosphorylates bHLH107 and GRF1, thereby switching the immune response to the \u2018ON\u2019 state. Furthermore, wild\u2010type GRF1, the quadruple phospho\u2010defective mutant GRF1\n                    <jats:sup>S72\/88\/125\/156A<\/jats:sup>\n                    and the quadruple phosphomimetic mutant GRF1\u00a0\n                    <jats:sup>S72\/88\/125\/156D<\/jats:sup>\n                    directly interact with unphosphorylated bHLH107, but not with the double phosphomimetic mutant protein bHLH107\n                    <jats:sup>S62\/72D<\/jats:sup>\n                    . We also revealed that GRF1 attenuates CPK3 kinase activity. Collectively, our findings indicated that CuTI is precisely orchestrated through GRF1\u2010mediated modulation of the CPK3\u2010bHLH107 module in Arabidopsis.\n                  <\/jats:p>","DOI":"10.1111\/tpj.71150","type":"journal-article","created":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T04:47:42Z","timestamp":1790830062000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["<scp>GRF1<\/scp>\n                    fine\u2010tunes the copper\u2010responsive transcription module of\n                    <scp>CPK3<\/scp>\n                    \u2010\n                    <scp>bHLH107<\/scp>\n                    by manipulating interaction and phosphorylation"],"prefix":"10.1111","volume":"127","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-2389-1710","authenticated-orcid":false,"given":"Yue","family":"Yu","sequence":"first","affiliation":[{"name":"State Key Laboratory of Hybrid Rice, Hubei Provincial Research Center for Basic Biological Sciences, College of Life Sciences Wuhan University  Wuhan 430072 China"},{"name":"Hubei Hongshan Laboratory  Wuhan 430070 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0007-6332-2380","authenticated-orcid":false,"given":"Haoran","family":"Xia","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Hybrid Rice, Hubei Provincial Research Center for Basic Biological Sciences, College of Life Sciences Wuhan University  Wuhan 430072 China"},{"name":"Hubei Hongshan Laboratory  Wuhan 430070 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ran","family":"Dong","sequence":"additional","affiliation":[{"name":"Shandong Institute of Pomology  Tai'an 271000 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0537-930X","authenticated-orcid":false,"given":"Haifeng","family":"Liu","sequence":"additional","affiliation":[{"name":"College of Agronomy Shandong Agricultural University  Taian 271018 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4311-8001","authenticated-orcid":false,"given":"Xiangsong","family":"Chen","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Hybrid Rice, Hubei Provincial Research Center for Basic Biological Sciences, College of Life Sciences Wuhan University  Wuhan 430072 China"},{"name":"Hubei Hongshan Laboratory  Wuhan 430070 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8320-7872","authenticated-orcid":false,"given":"Zhaohui","family":"Chu","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Hybrid Rice, Hubei 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China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,9,30]]},"reference":[{"key":"e_1_2_9_2_1","doi-asserted-by":"publisher","DOI":"10.4161\/psb.5.10.12957"},{"key":"e_1_2_9_3_1","doi-asserted-by":"publisher","DOI":"10.1104\/pp.110.153593"},{"key":"e_1_2_9_4_1","doi-asserted-by":"publisher","DOI":"10.1007\/s11033-022-07330-3"},{"key":"e_1_2_9_5_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.rsci.2025.02.001"},{"key":"e_1_2_9_6_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00709-012-0437-z"},{"key":"e_1_2_9_7_1","doi-asserted-by":"publisher","DOI":"10.1093\/plcell\/koad088"},{"key":"e_1_2_9_8_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.molp.2023.04.002"},{"key":"e_1_2_9_9_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.devcel.2007.06.009"},{"key":"e_1_2_9_10_1","doi-asserted-by":"publisher","DOI":"10.1111\/tpj.12677"},{"key":"e_1_2_9_11_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-022-28395-5"},{"key":"e_1_2_9_12_1","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.31636"},{"key":"e_1_2_9_13_1","doi-asserted-by":"publisher","DOI":"10.1105\/tpc.010188"},{"key":"e_1_2_9_14_1","doi-asserted-by":"publisher","DOI":"10.1038\/cdd.2012.114"},{"key":"e_1_2_9_15_1","doi-asserted-by":"publisher","DOI":"10.1021\/bi1003487"},{"key":"e_1_2_9_16_1","doi-asserted-by":"publisher","DOI":"10.1007\/s13593-018-0503-9"},{"key":"e_1_2_9_17_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.devcel.2024.12.036"},{"key":"e_1_2_9_18_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-313X.2012.04986.x"},{"key":"e_1_2_9_19_1","doi-asserted-by":"publisher","DOI":"10.1111\/ppl.70196"},{"key":"e_1_2_9_20_1","doi-asserted-by":"publisher","DOI":"10.1111\/mpp.12919"},{"key":"e_1_2_9_21_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.molp.2015.07.008"},{"key":"e_1_2_9_22_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-020-20007-4"},{"key":"e_1_2_9_23_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-313X.2010.04257.x"},{"key":"e_1_2_9_24_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.devcel.2025.06.020"},{"key":"e_1_2_9_25_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.tplants.2016.11.007"},{"key":"e_1_2_9_26_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00122-023-04455-2"},{"key":"e_1_2_9_27_1","doi-asserted-by":"publisher","DOI":"10.1111\/nph.17079"},{"key":"e_1_2_9_28_1","first-page":"S1","article-title":"The 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