{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,3,30]],"date-time":"2026-03-30T10:59:03Z","timestamp":1774868343661,"version":"3.50.1"},"reference-count":42,"publisher":"MDPI AG","issue":"7","license":[{"start":{"date-parts":[[2026,3,30]],"date-time":"2026-03-30T00:00:00Z","timestamp":1774828800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"The National Key Research and Development Program of China","award":["2022YFD1200704-3"],"award-info":[{"award-number":["2022YFD1200704-3"]}]},{"name":"Local Financial Funds of National Agricultural Science and Technology Center","award":["NASC2024KY09"],"award-info":[{"award-number":["NASC2024KY09"]}]},{"name":"Scientific Research Fund Project of Chengdu Agricultural University","award":["23BS04"],"award-info":[{"award-number":["23BS04"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Plants"],"abstract":"<jats:p>ZmSIG2A is a nuclear-encoded plastid sigma factor 2A in maize (Zea mays L.) that is essential for plastid gene transcription and chloroplast biogenesis. As a key regulator of chloroplast development and function, ZmSIG2A may also contribute to the coordination of plant growth and environmental adaptation; however, its roles in root development and stress responses remain largely unclear. We compared two ZmSIG2A mutants, eal1-1 (hypomorphic) and ems110 (nonsense). eal1-1 had increased root number and longer roots, while ems110 had normal root number but shorter roots and failed to mature. The zmsig2aVal480del transcript was upregulated in eal1-1, and the root-promoting effect of OsSIG2A in rice suggests a conserved role in monocot root growth. DAP-seq indicated that zmsig2aVal480del targets are involved in metabolism, transport, signaling, and antioxidants, with Chr4 peak clustering near multiple LRR-RLKs, suggesting a ZmSIG2A\u2013LRR-RLK module in root development and stress integration. Physiologically, eal1-1 showed increased antioxidant enzyme activities and reduced MDA, indicating enhanced ROS scavenging, while ems110 exhibited decreased enzyme activities and elevated MDA, indicating compromised ROS detoxification. Upstream, Y1H and dual-luciferase assays demonstrated that the Mediator subunit ZmMed31 positively regulates transcription from the ZmSIG2A promoter. Given Mediator\u2019s role in bridging transcription factors and the core transcriptional machinery, ZmMed31 likely links hormone-responsive transcription factors to the ZmSIG2A regulatory network. Collectively, we propose a stress-responsive ZmMed31\u2013ZmSIG2A\u2013LRR-RLK module that underpins maize root development and drought adaptation, offering mechanistic insight and potential targets for stress-resilient breeding.<\/jats:p>","DOI":"10.3390\/plants15071057","type":"journal-article","created":{"date-parts":[[2026,3,30]],"date-time":"2026-03-30T09:57:11Z","timestamp":1774864631000},"page":"1057","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["ZmMed31\u2013ZmSIG2A Coordinates ROS Homeostasis and LRR-RLK Signaling to Regulate Root Development"],"prefix":"10.3390","volume":"15","author":[{"given":"Dan","family":"Jiang","sequence":"first","affiliation":[{"name":"State Key Laboratory of Crop Gene Exploration and Utilization in Southwest China, Maize Research Institute, Sichuan Agricultural University, Chengdu 611130, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Shengwei","family":"Guo","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Crop Gene Exploration and Utilization in Southwest China, Maize Research Institute, Sichuan Agricultural University, Chengdu 611130, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Xin","family":"Yuan","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Crop Gene Exploration and Utilization in Southwest China, Maize Research Institute, Sichuan Agricultural University, Chengdu 611130, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Sheng","family":"Zhang","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Crop Gene Exploration and Utilization in Southwest China, Maize Research Institute, Sichuan Agricultural University, Chengdu 611130, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yuxin","family":"Zhang","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Crop Gene Exploration and Utilization in Southwest China, Maize Research Institute, Sichuan Agricultural University, Chengdu 611130, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yuqi","family":"Ning","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Crop Gene Exploration and Utilization in Southwest China, Maize Research Institute, Sichuan Agricultural University, Chengdu 611130, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Fujian","family":"Qu","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Crop Gene Exploration and Utilization in Southwest China, Maize Research Institute, Sichuan Agricultural University, Chengdu 611130, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Qunkai","family":"Niu","sequence":"additional","affiliation":[{"name":"Chengdu Agricultural College, Chengdu 611130, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Moju","family":"Cao","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Crop Gene Exploration and Utilization in Southwest China, Maize Research Institute, Sichuan Agricultural University, Chengdu 611130, China"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1968","published-online":{"date-parts":[[2026,3,30]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/S0074-7696(05)44001-2","article-title":"Plastid RNA Polymerases, Promoters, and Transcription Regulators in Higher Plants","volume":"244","author":"Shiina","year":"2005","journal-title":"Int. Rev. Cytol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"1382","DOI":"10.1093\/pcp\/pcp069","article-title":"Arabidopsis Mutants Carrying Chimeric Sigma Factor Genes Reveal Regulatory Determinants for Plastid Gene Expression","volume":"50","author":"Schweer","year":"2009","journal-title":"Plant Cell Physiol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"845","DOI":"10.1007\/s00299-007-0318-7","article-title":"Plant Sigma Factors and Their Role in Plastid Transcription","volume":"26","author":"Lysenko","year":"2007","journal-title":"Plant Cell Rep."},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"Shimmura, S., Nozoe, M., Kitora, S., Kin, S., Matsutani, S., Ishizaki, Y., Nakahira, Y., and Shiina, T. (2017). Comparative Analysis of Chloroplast psbD Promoters in Terrestrial Plants. Front. Plant Sci., 8.","DOI":"10.3389\/fpls.2017.01186"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"357","DOI":"10.1093\/pcp\/pch050","article-title":"The Multiple-Stress Responsive Plastid Sigma Factor, SIG5, Directs Activation of the psbD Blue Light-Responsive Promoter (BLRP) in Arabidopsis Thaliana","volume":"45","author":"Nagashima","year":"2004","journal-title":"Plant Cell Physiol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"940","DOI":"10.1016\/j.ejcb.2010.06.016","article-title":"Role and Regulation of Plastid Sigma Factors and Their Functional Interactors during Chloroplast Transcription\u2014Recent Lessons from Arabidopsis Thaliana","volume":"89","author":"Schweer","year":"2010","journal-title":"Eur. J. Cell Biol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"642","DOI":"10.1104\/pp.106.085878","article-title":"Dual Temporal Role of Plastid Sigma Factor 6 in Arabidopsis Development","volume":"142","author":"Loschelder","year":"2006","journal-title":"Plant Physiol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"2773","DOI":"10.1093\/jxb\/ery053","article-title":"Abiotic Stress-Induced Chloroplast Proteome Remodelling: A Mechanistic Overview","volume":"69","author":"Watson","year":"2018","journal-title":"J. Exp. Bot."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"770","DOI":"10.1016\/j.bbabio.2015.01.001","article-title":"Plastid Sigma Factors: Their Individual Functions and Regulation in Transcription","volume":"1847","author":"Chi","year":"2015","journal-title":"Biochim. Biophys. Acta BBA Bioenerg."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"1046","DOI":"10.1016\/j.tplants.2025.04.007","article-title":"Hub Metabolites at the Root\u2013Microbiome Interface: Unlocking Plant Drought Resilience","volume":"30","author":"Pantigoso","year":"2025","journal-title":"Trends Plant Sci."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"3267","DOI":"10.1093\/jxb\/ery082","article-title":"Genetic Diversity of Root System Architecture in Response to Drought Stress in Grain Legumes","volume":"69","author":"Ye","year":"2018","journal-title":"J. Exp. Bot."},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Kou, X., Han, W., and Kang, J. (2022). Responses of Root System Architecture to Water Stress at Multiple Levels: A Meta-Analysis of Trials under Controlled Conditions. Front. Plant Sci., 13.","DOI":"10.3389\/fpls.2022.1085409"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"e20395","DOI":"10.1002\/tpg2.20395","article-title":"Understanding Role of Roots in Plant Response to Drought: Way Forward to Climate-resilient Crops","volume":"17","author":"Kalra","year":"2024","journal-title":"Plant Genome"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"102789","DOI":"10.1016\/j.redox.2023.102789","article-title":"Novel Insight into Functions of Ascorbate Peroxidase in Higher Plants: More than a Simple Antioxidant Enzyme","volume":"64","author":"Li","year":"2023","journal-title":"Redox Biol."},{"key":"ref_15","doi-asserted-by":"crossref","unstructured":"Cannea, F.B., and Padiglia, A. (2025). Antioxidant Defense Systems in Plants: Mechanisms, Regulation, and Biotechnological Strategies for Enhanced Oxidative Stress Tolerance. Life, 15.","DOI":"10.3390\/life15081293"},{"key":"ref_16","doi-asserted-by":"crossref","unstructured":"Chen, F., Ha, X., Ma, T., and Ma, H. (2024). Comparative Analysis of the Physiological and Transcriptomic Profiles Reveals Alfalfa Drought Resistance Mechanisms. BMC Plant Biol., 24.","DOI":"10.1186\/s12870-024-05671-8"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"295","DOI":"10.1007\/s11104-022-05582-3","article-title":"Arbuscular Mycorrhizal Fungi and Antioxidant Enzymes in Ameliorating Drought Stress: A Meta-Analysis","volume":"480","author":"Chandrasekaran","year":"2022","journal-title":"Plant Soil."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"dev185819","DOI":"10.1242\/dev.185819","article-title":"Flavonols Regulate Root Hair Development by Modulating Accumulation of Reactive Oxygen Species in the Root Epidermis","volume":"147","author":"Gayomba","year":"2020","journal-title":"Development"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"940","DOI":"10.1016\/j.tplants.2021.03.005","article-title":"How Roots and Shoots Communicate through Stressful Times","volume":"26","author":"Li","year":"2021","journal-title":"Trends Plant Sci."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"640","DOI":"10.1016\/j.cub.2015.12.066","article-title":"Shoot-to-Root Mobile Transcription Factor HY5 Coordinates Plant Carbon and Nitrogen Acquisition","volume":"26","author":"Chen","year":"2016","journal-title":"Curr. Biol."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"2201","DOI":"10.1016\/j.cub.2023.04.061","article-title":"Photosynthetic Sucrose Drives the Lateral Root Clock in Arabidopsis Seedlings","volume":"33","author":"Kircher","year":"2023","journal-title":"Curr. Biol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"111570","DOI":"10.1016\/j.plantsci.2022.111570","article-title":"The Role of Shoot-Derived RNAs Transported to Plant Root in Response to Abiotic Stresses","volume":"328","author":"Liu","year":"2023","journal-title":"Plant Sci."},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Gao, Y., Qu, Q., Liu, N., Sun, M., Liu, X., Cao, Z., and Dong, J. (2025). Genome Identification of the LRR-RLK Gene Family in Maize (Zea Mays) and Expression Analysis in Response to Fusarium Verticillioides Infection. BMC Plant Biol., 25.","DOI":"10.1186\/s12870-025-06495-w"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"3047","DOI":"10.1007\/s00122-019-03405-1","article-title":"Fine Mapping and Candidate Gene Analysis of a QTL Associated with Leaf Rolling Index on Chromosome 4 of Maize (Zea mays L.)","volume":"132","author":"Gao","year":"2019","journal-title":"Theor. Appl. Genet."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"3321","DOI":"10.1007\/s00122-019-03427-9","article-title":"Characterization and Fine Mapping of Qkrnw4, a Major QTL Controlling Kernel Row Number in Maize","volume":"132","author":"Nie","year":"2019","journal-title":"Theor. Appl. Genet."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"5063","DOI":"10.1093\/jxb\/eru271","article-title":"The ZmCLA4 Gene in the qLA4-1 QTL Controls Leaf Angle in Maize (Zea mays L.)","volume":"65","author":"Zhang","year":"2014","journal-title":"J. Exp. Bot."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"1233","DOI":"10.1175\/JAMC-D-18-0174.1","article-title":"Global Patterns of Crop Production Losses Associated with Droughts from 1983 to 2009","volume":"58","author":"Kim","year":"2019","journal-title":"J. Appl. Meteorol. Climatol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"831","DOI":"10.1038\/s43017-023-00491-0","article-title":"Climate Change Impacts on Crop Yields","volume":"4","author":"Rezaei","year":"2023","journal-title":"Nat. Rev. Earth Environ."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"733","DOI":"10.1146\/annurev-arplant-042817-040218","article-title":"The Physiological Basis of Drought Tolerance in Crop Plants: A Scenario-Dependent Probabilistic Approach","volume":"69","author":"Tardieu","year":"2018","journal-title":"Annu. Rev. Plant Biol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"973","DOI":"10.1271\/bbb.68.973","article-title":"Differential Expression of Three Plastidial Sigma Factors, OsSIG1, OsSIG2A, and OsSIG2B, during Leaf Development in Rice","volume":"68","author":"Kasai","year":"2004","journal-title":"Biosci. Biotechnol. Biochem."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1007\/s00425-021-03730-0","article-title":"A Maize Leucine-Rich Repeat Receptor-like Protein Kinase Mediates Responses to Fungal Attack","volume":"254","author":"Block","year":"2021","journal-title":"Planta"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"388","DOI":"10.1016\/j.jgg.2024.10.016","article-title":"An LRR-RLK Protein Modulates Drought- and Salt-Stress Responses in Maize","volume":"52","author":"Yang","year":"2025","journal-title":"J. Genet. Genom."},{"key":"ref_33","doi-asserted-by":"crossref","unstructured":"Dufayard, J.-F. (2017). Corrigendum: New Insights on Leucine-Rich Repeats Receptor-Like Kinase Orthologous Relationships in Angiosperms. Front. Plant. Sci., 8.","DOI":"10.3389\/fpls.2017.00916"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"2898","DOI":"10.1105\/tpc.112.098277","article-title":"The Arabidopsis Mediator Subunit MED25 Differentially Regulates Jasmonate and Abscisic Acid Signaling through Interacting with the MYC2 and ABI5 Transcription Factors","volume":"24","author":"Chen","year":"2012","journal-title":"Plant Cell"},{"key":"ref_35","first-page":"E5624","article-title":"Mediator Subunit MED31 Is Required for Radial Patterning of Arabidopsis Roots","volume":"115","author":"Zhang","year":"2018","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"1330","DOI":"10.1016\/j.cj.2021.01.005","article-title":"A Valine Residue Deletion in ZmSig2A, a Sigma Factor, Accounts for a Revertible Leaf-Color Mutation in Maize","volume":"9","author":"Li","year":"2021","journal-title":"Crop J."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"611","DOI":"10.1373\/clinchem.2008.112797","article-title":"The MIQE Guidelines: Minimum Information for Publication of Quantitative Real-Time PCR Experiments","volume":"55","author":"Bustin","year":"2009","journal-title":"Clin. Chem."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"211","DOI":"10.1007\/s00253-006-0465-8","article-title":"Overview of Bacterial Expression Systems for Heterologous Protein Production: From Molecular and Biochemical Fundamentals to Commercial Systems","volume":"72","author":"Terpe","year":"2006","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"1659","DOI":"10.1038\/nprot.2017.055","article-title":"Mapping Genome-Wide Transcription-Factor Binding Sites Using DAP-Seq","volume":"12","author":"Bartlett","year":"2017","journal-title":"Nat. Protoc."},{"key":"ref_40","doi-asserted-by":"crossref","unstructured":"Cao, Y., Bi, M., Yang, P., Song, M., He, G., Wang, J., Yang, Y., Xu, L., and Ming, J. (2021). Construction of Yeast One-Hybrid Library and Screening of Transcription Factors Regulating LhMYBSPLATTER Expression in Asiatic Hybrid Lilies (Lilium Spp.). BMC Plant Biol., 21.","DOI":"10.1186\/s12870-021-03347-1"},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Franco-Zorrilla, J.M., and Mart\u00edn-Pizarro, C. (2026). Plant Transcription Factors: Methods and Protocols, Springer. Methods in Molecular Biology.","DOI":"10.1007\/978-1-0716-4972-5"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"5165","DOI":"10.1021\/acsnano.0c10351","article-title":"Facile Bioself-Assembled Crystals in Plants Promote Photosynthesis and Salt Stress Resistance","volume":"15","author":"Chi","year":"2021","journal-title":"ACS Nano"}],"container-title":["Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/7\/1057\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,3,30]],"date-time":"2026-03-30T10:04:49Z","timestamp":1774865089000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/7\/1057"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,3,30]]},"references-count":42,"journal-issue":{"issue":"7","published-online":{"date-parts":[[2026,4]]}},"alternative-id":["plants15071057"],"URL":"https:\/\/doi.org\/10.3390\/plants15071057","relation":{},"ISSN":["2223-7747"],"issn-type":[{"value":"2223-7747","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,3,30]]}}}