{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,30]],"date-time":"2026-07-30T16:14:19Z","timestamp":1785428059540,"version":"3.56.0"},"reference-count":42,"publisher":"MDPI AG","issue":"9","license":[{"start":{"date-parts":[[2026,4,26]],"date-time":"2026-04-26T00:00:00Z","timestamp":1777161600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001809","name":"Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32470299"],"award-info":[{"award-number":["32470299"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"award":["32470299"],"award-info":[{"award-number":["32470299"]}],"id":[{"id":"https:\/\/ror.org\/01h0zpd94","id-type":"ROR","asserted-by":"publisher"}]},{"name":"Jiangsu Natural Science Foundation for Distinguished Young Scholars","award":["BK2022084"],"award-info":[{"award-number":["BK2022084"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>Osmotic stress limits rice productivity, yet the crosstalk between hydrogen sulfide signaling and redox regulation remains incompletely understood. We previously showed that redox-dependent oligomerization of the basic (region) leucine zippers transcription factor bZIP68 at Cys245 confers osmotic tolerance. However, the role of an adjacent cysteine, Cys171, was undefined. Here, we demonstrate that osmotic stress induces persulfidation of bZIP68 specifically at Cys171. This modification facilitates Cys245-mediated oxidation-dependent oligomerization, thereby enhancing bZIP68 transcriptional activity toward COLD-REGULATED413-THYLAKOID MEMBRANE1 (COR413-TM1). Transgenic complementation and physiological assays confirmed that Cys171 persulfidation is essential for full bZIP68 function in osmotic adaptation. Transcriptomic analysis further revealed that Cys171 is required for bZIP68-driven transcriptional reprogramming under stress. Our findings establish a hierarchical redox cascade wherein persulfidation primes bZIP68 for oxidative activation, highlighting a regulatory crosstalk between distinct post-translational modifications. These mechanistic insights expand our understanding of H2S signaling and identify the bZIP68 cysteine network as a potential target for improving crop stress resilience.<\/jats:p>","DOI":"10.3390\/ijms27093841","type":"journal-article","created":{"date-parts":[[2026,4,28]],"date-time":"2026-04-28T11:33:37Z","timestamp":1777376017000},"page":"3841","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Hydrogen Sulfide Primes bZIP68 via Persulfidation to Enhance Redox-Dependent Transcription and Adaptation to Osmotic Stress in Rice"],"prefix":"10.3390","volume":"27","author":[{"given":"Xiaoyun","family":"Ma","sequence":"first","affiliation":[{"name":"Laboratory Center of Life Sciences, College of Life Sciences, Nanjing Agricultural University, Nanjing 210095, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fengchao","family":"Zhai","sequence":"additional","affiliation":[{"name":"Laboratory Center of Life Sciences, College of Life Sciences, Nanjing Agricultural University, Nanjing 210095, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lingxi","family":"Geng","sequence":"additional","affiliation":[{"name":"Laboratory Center of Life Sciences, College of Life Sciences, Nanjing Agricultural University, Nanjing 210095, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Guojing","family":"Chen","sequence":"additional","affiliation":[{"name":"Laboratory Center of Life Sciences, College of Life Sciences, Nanjing Agricultural University, Nanjing 210095, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wenge","family":"Li","sequence":"additional","affiliation":[{"name":"Laboratory Center of Life Sciences, College of Life Sciences, Nanjing Agricultural University, Nanjing 210095, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0090-6665","authenticated-orcid":false,"given":"Mohammad Saidur","family":"Rhaman","sequence":"additional","affiliation":[{"name":"Department of Seed Science and Technology, Bangladesh Agricultural University, Mymensingh 2202, Bangladesh"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jing","family":"Zhang","sequence":"additional","affiliation":[{"name":"National Key Laboratory for the Development and Utilization of Forest Food Resources, Co-Innovation Center for Sustainable Forestry in Southern China, State Key Laboratory of Tree Genetics and Breeding, Key Laboratory of State Forestry and Grassland Administration on Subtropical Forest Biodiversity Conservation, College of Life Sciences, Nanjing Forestry University, Nanjing 210037, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3503-1267","authenticated-orcid":false,"given":"Yanjie","family":"Xie","sequence":"additional","affiliation":[{"name":"Laboratory Center of Life Sciences, College of Life Sciences, Nanjing Agricultural University, Nanjing 210095, China"},{"name":"National Key Laboratory for the Development and Utilization of Forest Food Resources, Co-Innovation Center for Sustainable Forestry in Southern China, State Key Laboratory of Tree Genetics and Breeding, Key Laboratory of State Forestry and Grassland Administration on Subtropical Forest Biodiversity Conservation, College of Life Sciences, Nanjing Forestry University, Nanjing 210037, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Heng","family":"Zhou","sequence":"additional","affiliation":[{"name":"Laboratory Center of Life Sciences, College of Life Sciences, Nanjing Agricultural University, Nanjing 210095, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,4,26]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"104","DOI":"10.1038\/s41576-021-00413-0","article-title":"Abiotic stress responses in plants","volume":"23","author":"Zhang","year":"2022","journal-title":"Nat. Rev. Genet."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"663","DOI":"10.1038\/s41580-022-00499-2","article-title":"Reactive oxygen species signalling in plant stress responses","volume":"23","author":"Mittler","year":"2022","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"1063","DOI":"10.1038\/s41467-018-03463-x","article-title":"Hydrogen peroxide positively regulates brassinosteroid signaling through oxidation of the BRASSINAZOLERESISTANT1 transcription factor","volume":"9","author":"Tian","year":"2018","journal-title":"Nat. Commun."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"768","DOI":"10.1111\/tpj.14476","article-title":"Redox sensitive bZIP68 plays a role in balancing stress tolerance with growth in Arabidopsis","volume":"100","author":"Li","year":"2019","journal-title":"Plant J."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"112","DOI":"10.1093\/plcell\/koad250","article-title":"H2O2-dependent oxidation of the transcription factor GmNTL1 promotes salt tolerance in soybean","volume":"36","author":"Zhang","year":"2023","journal-title":"Plant Cell"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"367","DOI":"10.1093\/plcell\/koad276","article-title":"S-nitrosylation of the transcription factor MYB30 facilitates nitric oxide-promoted seed germination in Arabidopsis","volume":"36","author":"Zhao","year":"2024","journal-title":"Plant Cell"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"415","DOI":"10.1016\/j.tplants.2022.11.007","article-title":"Cysteine thiol-based post-translational modification: What do we know about transcription factors?","volume":"28","author":"Zhou","year":"2023","journal-title":"Trends Plant Sci."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"651","DOI":"10.1016\/j.molp.2021.11.006","article-title":"Rice GLUTATHIONE PEROXIDASE1-mediated oxidation of bZIP68 positively regulates ABA-independent osmotic stress signaling","volume":"15","author":"Zhou","year":"2022","journal-title":"Mol. Plant."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"977","DOI":"10.1111\/j.1469-8137.2010.03465.x","article-title":"Hydrogen sulphide, a novel gasotransmitter involved in guard cell signalling","volume":"188","author":"Lamattina","year":"2010","journal-title":"New Phytol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"983","DOI":"10.1016\/j.tplants.2019.08.003","article-title":"Hydrogen Sulfide: A New Warrior against Abiotic Stress","volume":"24","author":"Corpas","year":"2019","journal-title":"Trends Plant Sci."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"146","DOI":"10.1111\/jipb.13022","article-title":"Hydrogen sulfide, a signaling molecule in plant stress responses","volume":"63","author":"Zhang","year":"2021","journal-title":"J. Integr. Plant Biol."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"4251","DOI":"10.1093\/jxb\/erz225","article-title":"Signaling by hydrogen sulfide and cyanide through post-translational modification","volume":"70","author":"Gotor","year":"2019","journal-title":"J. Exp. Bot."},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Aroca, A., Carrillo, R., Montesinos-Pereira, D., Luque, C., Romero, L.C., and Gotor, C. (2025). Unraveling the Protective Role of Sulfide Signaling in Plants against Climate-Induced Abiotic Stress. J. Exp. Bot., eraf527.","DOI":"10.1093\/jxb\/eraf527"},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Aroca, A., Gotor, C., and Romero, L.C. (2018). Hydrogen Sulfide Signaling in Plants: Emerging Roles of Protein Persulfidation. Front. Plant Sci., 9.","DOI":"10.3389\/fpls.2018.01369"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"1253","DOI":"10.1021\/acs.chemrev.7b00205","article-title":"Chemical Biology of H2S Signaling through Persulfidation","volume":"118","author":"Filipovic","year":"2018","journal-title":"Chem. Rev."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1431","DOI":"10.1111\/nph.18838","article-title":"Persulfidation protects from oxidative stress under nonphotorespiratory conditions in Arabidopsis","volume":"238","author":"Vignane","year":"2023","journal-title":"New Phytol."},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Romero, L.C., Carrillo, R., Montesinos-Pereira, D., Luque, C., Aroca, A., and Gotor, C. (2025). Protein persulfidation: A ubiquitous modification regulating a broad spectrum of biological processes. J. Exp. Bot., eraf464.","DOI":"10.1093\/jxb\/eraf464"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"4915","DOI":"10.1093\/jxb\/erx294","article-title":"Persulfidation proteome reveals the regulation of protein function by hydrogen sulfide in diverse biological processes in Arabidopsis","volume":"68","author":"Aroca","year":"2017","journal-title":"J. Exp. Bot."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"3902","DOI":"10.1105\/tpc.20.00766","article-title":"Abscisic Acid-Triggered Persulfidation of the Cys Protease ATG4 Mediates Regulation of Autophagy by Sulfide","volume":"32","author":"Aroca","year":"2020","journal-title":"Plant Cell"},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Jurado-Flores, A., Romero, L.C., and Gotor, C. (2021). Label-Free Quantitative Proteomic Analysis of Nitrogen Starvation in Arabidopsis Root Reveals New Aspects of H2S Signaling by Protein Persulfidation. Antioxidants, 10.","DOI":"10.3390\/antiox10040508"},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Jurado-Flores, A., Gotor, C., and Romero, L.C. (2023). Proteome Dynamics of Persulfidation in Leaf Tissue under Light\/Dark Conditions and Carbon Deprivation. Antioxidants, 12.","DOI":"10.3390\/antiox12040789"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"4654","DOI":"10.1093\/jxb\/erad165","article-title":"Sulfide promotes tolerance to drought through protein persulfidation in Arabidopsis","volume":"74","author":"Aroca","year":"2023","journal-title":"J. Exp. Bot."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"5130","DOI":"10.1093\/jxb\/erae249","article-title":"Analysis of sulfide signaling in rice highlights specific drought responses","volume":"75","author":"Zhang","year":"2024","journal-title":"J. Exp. Bot."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"732","DOI":"10.1016\/j.molp.2020.01.004","article-title":"Hydrogen Sulfide Positively Regulates Abscisic Acid Signaling through Persulfidation of SnRK2.6 in Guard Cells","volume":"13","author":"Chen","year":"2020","journal-title":"Mol. Plant."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"1000","DOI":"10.1105\/tpc.19.00826","article-title":"Persulfidation-based Modification of Cysteine Desulfhydrase and the NADPH Oxidase RBOHD Controls Guard Cell Abscisic Acid Signaling","volume":"32","author":"Shen","year":"2020","journal-title":"Plant Cell"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"921","DOI":"10.1016\/j.molp.2021.03.007","article-title":"Hydrogen sulfide-linked persulfidation of ABI4 controls ABA responses through the transactivation of MAPKKK18 in Arabidopsis","volume":"14","author":"Zhou","year":"2021","journal-title":"Mol. Plant"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"213","DOI":"10.1016\/j.plaphy.2020.07.038","article-title":"Hydrogen sulfide promotes rice drought tolerance via reestablishing redox homeostasis and activation of ABA biosynthesis and signaling","volume":"155","author":"Zhou","year":"2020","journal-title":"Plant Physiol. Biochem."},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Liu, Q., Luo, L., Wang, X., Shen, Z., and Zheng, L. (2017). Comprehensive Analysis of Rice Laccase Gene (OsLAC) Family and Ectopic Expression of OsLAC10 Enhances Tolerance to Copper Stress in Arabidopsis. Int. J. Mol. Sci., 18.","DOI":"10.3390\/ijms18020209"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"126771","DOI":"10.1016\/j.envpol.2025.126771","article-title":"Dose-dependent effects of butachlor accumulation and degradation on amino acid metabolism and lignin biosynthesis in rice","volume":"382","author":"Wang","year":"2025","journal-title":"Environ. Pollut."},{"key":"ref_30","doi-asserted-by":"crossref","unstructured":"Wang, Y., Du, F., Wang, J., Li, Y., Zhang, Y., Zhao, X., Zheng, T., Li, Z., Xu, J., and Wang, W. (2021). Molecular Dissection of the Gene OsGA2ox8 Conferring Osmotic Stress Tolerance in Rice. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms22179107"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"1144","DOI":"10.1111\/pce.12223","article-title":"Overexpression of stress-inducible OsBURP16, the \u03b2 subunit of polygalacturonase 1, decreases pectin content and cell adhesion and increases abiotic stress sensitivity in rice","volume":"37","author":"Liu","year":"2014","journal-title":"Plant Cell Environ."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"860","DOI":"10.1111\/tpj.15987","article-title":"The Germin-like protein OsGER4 is involved in promoting crown root development under exogenous jasmonic acid treatment in rice","volume":"112","author":"To","year":"2022","journal-title":"Plant J."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"113464","DOI":"10.1016\/j.envpol.2019.113464","article-title":"Identification of epigenetic mechanisms in paddy crop associated with lowering environmentally related cadmium risks to food safety","volume":"256","author":"Feng","year":"2020","journal-title":"Environ. Pollut."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"394","DOI":"10.1111\/jipb.13622","article-title":"How plants sense and respond to osmotic stress","volume":"66","author":"Yu","year":"2024","journal-title":"J. Integr. Plant Biol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"100005","DOI":"10.1016\/j.ncrops.2023.11.003","article-title":"The central role of transcription factors in bridging biotic and abiotic stress responses for plants\u2019 resilience","volume":"1","author":"Liu","year":"2024","journal-title":"New Crops"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"e14366","DOI":"10.1111\/ppl.14366","article-title":"Transcription factors as molecular switches regulating plant responses to drought stress","volume":"176","author":"Wei","year":"2024","journal-title":"Physiol. Plant."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"2005","DOI":"10.1111\/jipb.13932","article-title":"Recent advances in improving yield and immunity through transcription factor engineering","volume":"67","author":"Iswanto","year":"2025","journal-title":"J. Integr. Plant Biol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"2862","DOI":"10.1093\/jxb\/eraa093","article-title":"Hydrogen sulfide: A multi-tasking signal molecule in the regulation of oxidative stress responses","volume":"71","author":"Chen","year":"2020","journal-title":"J. Exp. Bot."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"335","DOI":"10.1016\/j.freeradbiomed.2024.08.041","article-title":"Systematic analysis of the global characteristics and reciprocal effects of S-nitrosylation and S-persulfidation in the human proteome","volume":"224","author":"Li","year":"2024","journal-title":"Free Radic. Biol. Med."},{"key":"ref_40","doi-asserted-by":"crossref","unstructured":"Zhou, M., Zhang, J., Zhou, H., Zhao, D., Duan, T., Wang, S., Yuan, X., and Xie, Y. (2022). Hydrogen Sulfide-Linked Persulfidation Maintains Protein Stability of ABSCISIC ACID-INSENSITIVE 4 and Delays Seed Germination. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms23031389"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"2796","DOI":"10.1038\/nprot.2006.469","article-title":"A protocol for Agrobacterium-mediated transformation in rice","volume":"1","author":"Nishimura","year":"2006","journal-title":"Nat Protoc."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"100012","DOI":"10.1016\/j.ncrops.2024.100012","article-title":"ZmMYB56 regulates stomatal closure and drought tolerance in maize seedlings through the transcriptional regulation of ZmTOM7","volume":"1","author":"Li","year":"2024","journal-title":"New Crops"}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/9\/3841\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,4,28]],"date-time":"2026-04-28T11:41:13Z","timestamp":1777376473000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/9\/3841"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,4,26]]},"references-count":42,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2026,5]]}},"alternative-id":["ijms27093841"],"URL":"https:\/\/doi.org\/10.3390\/ijms27093841","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,4,26]]}}}