{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,28]],"date-time":"2026-08-28T19:18:13Z","timestamp":1787944693175,"version":"build-2784847793"},"reference-count":52,"publisher":"MDPI AG","issue":"17","license":[{"start":{"date-parts":[[2026,8,28]],"date-time":"2026-08-28T00:00:00Z","timestamp":1787875200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Department of Science and Technology of Jilin Province, China","award":["YDZJ202601ZYTS328"],"award-info":[{"award-number":["YDZJ202601ZYTS328"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Plants"],"abstract":"<jats:p>Soil salinization poses a major environmental threat to global agriculture, affecting approximately 20% of cultivated land and 50% of irrigated land worldwide. Developing salt\u2013alkali tolerant plant varieties represents a sustainable strategy for utilizing these marginal lands. Phosphatidylinositol-specific phospholipase C (PI-PLC) is a key enzyme in the phosphoinositide signaling system and has been implicated in plant stress responses; however, its function under salt\u2013alkali stress remains poorly understood. In this study, the function of AtPLC1 in salt\u2013alkali tolerance was investigated, and only the atplc1 mutant exhibited a pronounced stress-sensitive phenotype, with AtPLC1 being predominantly expressed in roots and leaves, with peak expression at 6 h of treatment. Compared with wild-type, atplc1 mutants displayed significantly reduced seedling survival, retarded root growth, decreased biomass, water content, chlorophyll, and soluble sugar contents, yet accumulated higher levels of Na+, malondialdehyde, H2O2, and superoxide anions under salt\u2013alkali stress. Notably, atplc1 mutants showed increased stomatal conductance and decreased leaf surface temperature, as detected by thermal imaging, indicating impaired water regulation. Collectively, our findings demonstrate that AtPLC1 positively regulates salt\u2013alkali tolerance and provides a candidate gene for molecular breeding of stress-resistant crops.<\/jats:p>","DOI":"10.3390\/plants15172633","type":"journal-article","created":{"date-parts":[[2026,8,28]],"date-time":"2026-08-28T18:33:52Z","timestamp":1787942032000},"page":"2633","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Loss of AtPLC1 Impairs Salt\u2013Alkali Tolerance via Disruption of Stomatal Regulation and Redox Homeostasis in Arabidopsis thaliana"],"prefix":"10.3390","volume":"15","author":[{"given":"Xiang","family":"Li","sequence":"first","affiliation":[{"name":"College of Life Science, Jilin Agricultural University, Changchun 130118, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"methodology","vocabulary":"credit"},{"role":"software","vocabulary":"credit"},{"role":"validation","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"}]},{"given":"Yu","family":"Wang","sequence":"additional","affiliation":[{"name":"College of Life Science, Jilin Agricultural University, Changchun 130118, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"software","vocabulary":"credit"},{"role":"validation","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"},{"role":"visualization","vocabulary":"credit"}]},{"given":"Linhan","family":"Si","sequence":"additional","affiliation":[{"name":"College of Life Science, Jilin Agricultural University, Changchun 130118, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"formal-analysis","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6871-1026","authenticated-orcid":false,"given":"Daqian","family":"Sun","sequence":"additional","affiliation":[{"name":"College of Life Science, Jilin Agricultural University, Changchun 130118, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"formal-analysis","vocabulary":"credit"}]},{"given":"Nan","family":"Wang","sequence":"additional","affiliation":[{"name":"College of Life Science, Jilin Agricultural University, Changchun 130118, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"formal-analysis","vocabulary":"credit"}]},{"given":"Weican","family":"Liu","sequence":"additional","affiliation":[{"name":"College of Life Science, Jilin Agricultural University, Changchun 130118, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"resources","vocabulary":"credit"}]},{"given":"Yuanyuan","family":"Dong","sequence":"additional","affiliation":[{"name":"College of Life Science, Jilin Agricultural University, Changchun 130118, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"data-curation","vocabulary":"credit"},{"role":"project-administration","vocabulary":"credit"}]},{"given":"Xiaowei","family":"Li","sequence":"additional","affiliation":[{"name":"College of Life Science, Jilin Agricultural University, Changchun 130118, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"methodology","vocabulary":"credit"},{"role":"software","vocabulary":"credit"},{"role":"data-curation","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5733-2696","authenticated-orcid":false,"given":"Fawei","family":"Wang","sequence":"additional","affiliation":[{"name":"College of Life Science, Jilin Agricultural University, Changchun 130118, China"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"conceptualization","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"},{"role":"funding-acquisition","vocabulary":"credit"},{"role":"supervision","vocabulary":"credit"}]}],"member":"1968","published-online":{"date-parts":[[2026,8,28]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"3267","DOI":"10.4236\/ajps.2015.619318","article-title":"Enhancing the Salt Tolerance Potential of Watermelon (Citrullus lanatus) by Exogenous Application of Salicylic Acid","volume":"6","author":"Ayyub","year":"2015","journal-title":"Am. J. Plant Sci."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Wang, J., Zhang, Y., Yan, X., and Guo, J. (2020). Physiological and transcriptomic analyses of yellow horn (Xanthoceras sorbifolia) provide important insights into salt and saline-alkali stress tolerance. PLoS ONE, 15.","DOI":"10.1371\/journal.pone.0244365"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"125","DOI":"10.1007\/978-3-032-00190-0_6","article-title":"Soil Salinity and Climate Change: Strategies for Mitigation of Salt Stress to Sustainable Agriculture","volume":"Volume 1","author":"Zahid","year":"2025","journal-title":"Climate Resilient and Sustainable Agriculture: Volume 1. Advances in Global Change Research"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"1332","DOI":"10.1104\/pp.18.00499","article-title":"VAMP711 Is Required for Abscisic Acid-Mediated Inhibition of Plasma Membrane H+-ATPase Activity","volume":"178","author":"Xue","year":"2018","journal-title":"Plant Physiol."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"100079","DOI":"10.1016\/j.ncrops.2025.100079","article-title":"Current perspectives on improving soybean performance on saline-alkaline lands","volume":"3","author":"He","year":"2025","journal-title":"New Crops"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"S260","DOI":"10.1194\/jlr.R800098-JLR200","article-title":"Plant phospholipid signaling: \u201cin a nutshell\u201d","volume":"50","author":"Munnik","year":"2009","journal-title":"J. Lipid Res."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1016\/j.plipres.2016.01.002","article-title":"Plant phospholipases D and C and their diverse functions in stress responses","volume":"62","author":"Hong","year":"2016","journal-title":"Prog. Lipid Res."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1192","DOI":"10.1111\/mpp.13343","article-title":"Phospholipase C: Diverse functions in plant biotic stress resistance and fungal pathogenicity","volume":"24","author":"Fang","year":"2023","journal-title":"Mol. Plant Pathol."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"1172","DOI":"10.1111\/nph.14426","article-title":"A phosphoinositide-specific phospholipase C pathway elicits stress-induced Ca(2+) signals and confers salt tolerance to rice","volume":"214","author":"Li","year":"2017","journal-title":"New Phytol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"799","DOI":"10.1007\/s10529-005-5802-y","article-title":"Characterization of a novel phosphoinositide-specific phospholipase C from Zea mays and its expression in Escherichia coli","volume":"11","author":"Zhai","year":"2005","journal-title":"Biotechnol. Lett."},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Wang, F., Deng, Y., Zhou, Y., Dong, J., Chen, H., Dong, Y., Wang, N., Li, X., and Li, H. (2015). Genome-Wide Analysis and Expression Profiling of the Phospholipase C Gene Family in Soybean (Glycine max). PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0138467"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"534","DOI":"10.1016\/j.jplph.2010.09.004","article-title":"Phosphatidic acid production in chitosan-elicited tomato cells, via both phospholipase D and phospholipase C\/diacylglycerol kinase, requires nitric oxide","volume":"168","author":"Raho","year":"2011","journal-title":"J. Plant Physiol."},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Wang, H., Yu, J., Zhang, X., Zeng, Q., Zeng, T., Gu, L., Zhu, B., Yu, F., and Du, X. (2024). Genome-Wide Identification and Analysis of Phospholipase C Gene Family Reveals Orthologs, Co-Expression Networks, and Expression Profiling Under Abiotic Stress in Sorghum bicolor. Plants, 13.","DOI":"10.3390\/plants13212976"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"3519","DOI":"10.1105\/tpc.106.047373","article-title":"Pollen tube tip growth depends on plasma membrane polarization mediated by tobacco PLC3 activity and endocytic membrane recycling","volume":"18","author":"Helling","year":"2006","journal-title":"Plant Cell"},{"key":"ref_15","doi-asserted-by":"crossref","unstructured":"Kong, J., Chen, R., Liu, R., Wang, W., Wang, S., Zhang, J., and Yang, N. (2023). PLC1 mediated Cycloastragenol-induced stomatal movement by regulating the production of NO in Arabidopsis thaliana. BMC Plant Biol., 23.","DOI":"10.1186\/s12870-023-04555-7"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"469","DOI":"10.1093\/pcp\/pcx194","article-title":"Arabidopsis phospholipase C3 is involved in lateral root initiation and ABA responses in seed germination and stomatal closure","volume":"59","author":"Zhang","year":"2018","journal-title":"Plant Cell Physiol."},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Kanehara, K., Yu, C.Y., Cho, Y., Cheong, W.F., Torta, F., Shui, G., Wenk, M.R., and Nakamura, Y. (2015). Arabidopsis AtPLC2 Is a Primary Phosphoinositide-Specific Phospholipase C in Phosphoinositide Metabolism and the Endoplasmic Reticulum Stress Response. PLoS Genet., 11.","DOI":"10.1371\/journal.pgen.1005511"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"kiae534","DOI":"10.1093\/plphys\/kiae534","article-title":"Plant PI-PLC signaling in stress and development","volume":"197","author":"Laxalt","year":"2025","journal-title":"Plant Physiol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"101158","DOI":"10.1016\/j.plipres.2022.101158","article-title":"The functions of phospholipases and their hydrolysis products in plant growth, development and stress responses","volume":"86","author":"Ali","year":"2022","journal-title":"Prog. Lipid Res."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"9752","DOI":"10.1038\/s41598-024-60530-8","article-title":"Genetic interaction between TTG2 and AtPLC1 reveals a role for phosphoinositide signaling in a co-regulated suite of Arabidopsis epidermal pathways","volume":"14","author":"Goldberg","year":"2024","journal-title":"Sci. Rep."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1143","DOI":"10.1105\/tpc.13.5.1143","article-title":"Arabidopsis PLC1 Is Required for Secondary Responses to Abscisic Acid Signals","volume":"13","author":"Sanchez","year":"2001","journal-title":"Plant Cell"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1007\/s00425-024-04554-4","article-title":"Development and drought escape response in Arabidopsis thaliana are regulated by AtPLC1 in response to abscisic acid","volume":"260","author":"Wang","year":"2024","journal-title":"Planta"},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Fan, D., Ruan, J., Xu, Q., Ying, J., Wang, Y., Tong, X., Li, Z., Cheng, Y., Xue, D., and Zhang, J. (2026). From Model Plants to Staple Crops: Molecular Mechanisms of Plant Saline-Alkali Tolerance. Plants, 15.","DOI":"10.3390\/plants15040532"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1007\/s11032-023-01383-3","article-title":"Twenty years of mining salt tolerance genes in soybean","volume":"43","author":"Leung","year":"2023","journal-title":"Mol. Breed."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"14500","DOI":"10.1039\/C8RA00721G","article-title":"Effects of arbuscular mycorrhizal fungi on Leymus chinensis seedlings under salt-alkali stress and nitrogen deposition conditions: From osmotic adjustment and ion balance","volume":"8","author":"Lin","year":"2018","journal-title":"RSC Adv."},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Wang, X., Shen, H., and Yang, L. (2024). The Response of Hormones, Reactive Oxygen Species and Nitric Oxide in the Polyethylene-Glycol-Promoted, Salt-Alkali-Stress-Induced Embryo Germination of Sorbus pohuashanensis. Int. J. Mol. Sci., 25.","DOI":"10.3390\/ijms25105128"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"3903","DOI":"10.1073\/pnas.92.9.3903","article-title":"A gene encoding a phosphatidylinositol-specific phospholipase C is induced by dehydration and salt stress in Arabidopsis thaliana","volume":"92","author":"Hirayama","year":"1995","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"643","DOI":"10.1111\/j.1469-8137.2004.01069.x","article-title":"Gene-specific expression and calcium activation of Arabidopsis thaliana phospholipase C isoforms","volume":"162","author":"Hunt","year":"2004","journal-title":"New Phytol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"885","DOI":"10.1093\/pcp\/pcad123","article-title":"Ectopic Expression of Distinct PLC Genes Identifies \u2018Compactness\u2019 as a Possible Architectural Shoot Strategy to Cope with Drought Stress","volume":"65","author":"Vaseva","year":"2024","journal-title":"Plant Cell Physiol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"455","DOI":"10.1111\/nph.16538","article-title":"E2 conjugases UBC1 and UBC2 regulate MYB42-mediated SOS pathway in response to salt stress in Arabidopsis","volume":"227","author":"Sun","year":"2000","journal-title":"New Phytol."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"uhaf337","DOI":"10.1093\/hr\/uhaf337","article-title":"Response of plant roots to mechanical environmental stimuli","volume":"13","author":"Baiyin","year":"2025","journal-title":"Hortic. Res."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"651","DOI":"10.1146\/annurev.arplant.59.032607.092911","article-title":"Mechanisms of salinity tolerance","volume":"59","author":"Munns","year":"2008","journal-title":"Annu. Rev. Plant Biol."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"163","DOI":"10.1007\/s11099-013-0021-6","article-title":"Photosynthesis under stressful environments: An overview","volume":"51","author":"Ashraf","year":"2013","journal-title":"Photosynthetica"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"25113","DOI":"10.1038\/s41598-024-75434-w","article-title":"Comparative study of the photosynthetic efficiency and leaf structure of four Cotoneaster species","volume":"14","author":"Borkowska","year":"2024","journal-title":"Sci. Rep."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"e18804","DOI":"10.7717\/peerj.18804","article-title":"A plant-derived biostimulant Aminolom Enzimatico application stimulates chlorophyll content, electrolyte leakage, stomata density and root yield of radishes under salinity stress","volume":"13","author":"Kaya","year":"2025","journal-title":"Peer J."},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Fang, S., Hou, X., and Liang, X. (2021). Response Mechanisms of Plants Under Saline-Alkali Stress. Front. Plant Sci., 12.","DOI":"10.3389\/fpls.2021.667458"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1111\/j.1365-3040.2010.02232.x","article-title":"Glycinebetaine protects plants against abiotic stress: Mechanisms and biotechnological applications","volume":"34","author":"Chen","year":"2011","journal-title":"Plant Cell Environ."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"1746","DOI":"10.4161\/psb.6.11.17801","article-title":"Glycinebetaine and abiotic stress tolerance in plants","volume":"6","author":"Giri","year":"2011","journal-title":"Plant Signal. Behav."},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Sallam, A., Hasseb, N.M., Karam, M.A., B\u00f6rner, A., Zheng, X., and Moursi, Y.S. (2026). Differential Regulation of Gene Expression, Ion Homeostasis, and Antioxidant Defense Confers Salinity Tolerance During Seed Germination in Wheat. Plants, 15.","DOI":"10.3390\/plants15020230"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"112213","DOI":"10.1016\/j.plantsci.2024.112213","article-title":"Mechanical wounding improves salt tolerance by maintaining root ion homeostasis in a desert shrub","volume":"348","author":"Liu","year":"2024","journal-title":"Plant Sci."},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Zhang, M., Qin, S., Yan, J., Li, L., Xu, M., Liu, Y., and Zhang, W. (2023). Genome-wide identification and analysis of TCP family genes in Medicago sativa reveal their critical roles in Na+\/K+ homeostasis. BMC Plant Biol., 23.","DOI":"10.1186\/s12870-023-04318-4"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"100024","DOI":"10.1016\/j.ncrops.2024.100024","article-title":"ZmHAK17 encodes a Na+-selective transporter that promotes maize seed germination under salt conditions","volume":"1","author":"Wang","year":"2024","journal-title":"New Crops"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"358","DOI":"10.1016\/j.plantsci.2016.08.009","article-title":"The coordinated regulation of Na+ and K+ in Hordeum brevisubulatum responding to time of salt stress","volume":"252","author":"Wang","year":"2016","journal-title":"Plant Sci."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"1167","DOI":"10.1007\/s13258-021-01123-x","article-title":"Phosphoinositide-specific phospholipase C gene involved in heat and drought tolerance in wheat (Triticum aestivum L.)","volume":"43","author":"Wang","year":"2021","journal-title":"Genes Genom."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"374","DOI":"10.1007\/s00344-025-11869-3","article-title":"PLC1 Regulates Stomatal Closure and Enhances Arabidopsis Tolerance to Water Deficit by Modulating the ABA\u2013H2O2\u2013NO Pathway","volume":"1","author":"Wang","year":"2026","journal-title":"J. Plant Growth Regul."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1046\/j.1365-313X.2003.01698.x","article-title":"Phospholipase C is required for the control of stomatal aperture by ABA","volume":"34","author":"Hunt","year":"2003","journal-title":"Plant J."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"199","DOI":"10.1093\/jxb\/erh027","article-title":"The effects of manipulating phospholipase C on guard cell ABA-signalling","volume":"55","author":"Mills","year":"2004","journal-title":"J. Exp. Bot."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"151","DOI":"10.3389\/fpls.2014.00151","article-title":"Tolerance to drought and salt stress in plants: Unraveling the signaling networks","volume":"5","author":"Golldack","year":"2014","journal-title":"Front. Plant Sci."},{"key":"ref_49","doi-asserted-by":"crossref","unstructured":"Wang, J., Jiang, X., Zhao, C., Fang, Z., and Jiao, P. (2020). Transcriptomic and metabolomic analysis reveals the role of CoA in the salt tolerance of Zygophyllum spp.. BMC Plant Biol., 20.","DOI":"10.1186\/s12870-019-2226-8"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1111\/nph.15899","article-title":"How do stomata respond to water status?","volume":"224","author":"Buckley","year":"2019","journal-title":"New Phytol."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"e473","DOI":"10.1002\/pld3.473","article-title":"COP1 mutation causes low leaf temperature under various abiotic stresses in Arabidopsis thaliana","volume":"6","author":"Kim","year":"2022","journal-title":"Plant Direct"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1007\/s44154-024-00199-1","article-title":"CsLAC4, regulated by CsmiR397a, confers drought tolerance to the tea plant by enhancing lignin biosynthesis","volume":"4","author":"Yang","year":"2024","journal-title":"Stress Biol."}],"container-title":["Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/17\/2633\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,28]],"date-time":"2026-08-28T18:51:14Z","timestamp":1787943074000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/17\/2633"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,28]]},"references-count":52,"journal-issue":{"issue":"17","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["plants15172633"],"URL":"https:\/\/doi.org\/10.3390\/plants15172633","relation":{},"ISSN":["2223-7747"],"issn-type":[{"value":"2223-7747","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,28]]}}}