{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,25]],"date-time":"2026-08-25T09:19:04Z","timestamp":1787649544483,"version":"build-2736575974"},"reference-count":75,"publisher":"MDPI AG","issue":"17","license":[{"start":{"date-parts":[[2026,8,25]],"date-time":"2026-08-25T00:00:00Z","timestamp":1787616000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100012226","name":"Fundamental Research Funds for the Central Universities","doi-asserted-by":"publisher","award":["SWU120072"],"award-info":[{"award-number":["SWU120072"]}],"id":[{"id":"10.13039\/501100012226","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Plants"],"abstract":"<jats:p>Root hairs are tubular protrusions of root epidermal cells that expand the root surface area to facilitate water and nutrient uptake. The target of rapamycin (TOR) kinase has been identified as a positive regulator of root hair elongation, and the RHD6-RSL4 bHLH transcriptional cascade is well established as a core module that governs root hair morphogenesis. However, whether TOR signaling acts upstream of the RHD6-RSL4 pathway and how glucose signals are integrated into this transcriptional regulatory network during root hair development remain incompletely understood. In this study, transcriptome profiling combined with pharmacological and genetic functional assays was performed to elucidate the TOR-mediated transcriptional regulatory pathway of root hair elongation in Arabidopsis. Chemical inhibition of TOR triggered genome-wide transcriptional reprogramming in seedling roots, including disruption of auxin and ethylene signal transduction and pronounced downregulation of hundreds of genes related to root hair development. Glucose-activated TOR signaling modulates the expression of root hair-specific (RHS) genes mainly through the core RHD6-RSL4 transcriptional cascade. The transcription of RSL1\u2013RSL5 was strongly dependent on functional TOR activity, whereas RHD6 transcript abundance was specifically induced by glucose\u2013TOR signaling under carbon-starvation recovery conditions. Genetic overexpression of either RHD6 or RSL4 partially rescued root hair elongation defects caused by TOR suppression, confirming that the RHD6-RSL4 cascade functions as a critical downstream transcriptional module of glucose\u2013TOR signaling. Collectively, this work establishes a transcriptional framework in which glucose\u2013TOR signals modulate root hair elongation via transcriptional activation of the master bHLH regulators RHD6 and RSL4.<\/jats:p>","DOI":"10.3390\/plants15172586","type":"journal-article","created":{"date-parts":[[2026,8,25]],"date-time":"2026-08-25T08:00:48Z","timestamp":1787644848000},"page":"2586","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Glucose\u2013TOR Signaling Regulates Root Hair Elongation in Arabidopsis via the RHD6-RSL4 Transcriptional Cascade"],"prefix":"10.3390","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0009-0001-6675-4783","authenticated-orcid":false,"given":"Bingru","family":"Wang","sequence":"first","affiliation":[{"name":"College of Agronomy and Biotechnology, Southwest University, Chongqing 400715, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jueru","family":"Zhang","sequence":"additional","affiliation":[{"name":"College of Agronomy and Biotechnology, Southwest University, Chongqing 400715, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wei","family":"Yan","sequence":"additional","affiliation":[{"name":"College of Agronomy and Biotechnology, Southwest University, Chongqing 400715, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiumei","family":"Dai","sequence":"additional","affiliation":[{"name":"College of Agronomy and Biotechnology, Southwest University, Chongqing 400715, China"},{"name":"Engineering Research Center of South Upland Agriculture, Ministry of Education, Chongqing 400715, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6309-488X","authenticated-orcid":false,"given":"Jiankui","family":"Zhang","sequence":"additional","affiliation":[{"name":"College of Agronomy and Biotechnology, Southwest University, Chongqing 400715, China"},{"name":"Engineering Research Center of South Upland Agriculture, Ministry of Education, Chongqing 400715, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tian","family":"Zhang","sequence":"additional","affiliation":[{"name":"Chongqing Medical and Pharmaceutical College, Chongqing 401331, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6847-7487","authenticated-orcid":false,"given":"Kexuan","family":"Deng","sequence":"additional","affiliation":[{"name":"College of Agronomy and Biotechnology, Southwest University, Chongqing 400715, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,8,25]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"e0060","DOI":"10.1199\/tab.0060","article-title":"Root hairs","volume":"1","author":"Grierson","year":"2002","journal-title":"Arab. Book"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"365","DOI":"10.1146\/annurev.arplant.59.032607.092949","article-title":"A genetic regulatory network in the development of trichomes and root hairs","volume":"59","author":"Ishida","year":"2008","journal-title":"Annu. Rev. Plant Biol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"1848","DOI":"10.1242\/dev.132845","article-title":"The regulation and plasticity of root hair patterning and morphogenesis","volume":"143","author":"Schmidt","year":"2016","journal-title":"Development"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"115","DOI":"10.1016\/j.semcdb.2017.10.003","article-title":"Regulation and functional diversification of root hairs","volume":"83","author":"Cui","year":"2018","journal-title":"Semin. Cell Dev. Biol."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"1253","DOI":"10.1242\/dev.122.4.1253","article-title":"The homeobox gene GLABRA2 is required for position-dependent cell differentiation in the root epidermis of Arabidopsis thaliana","volume":"122","author":"Masucci","year":"1996","journal-title":"Development"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"1515","DOI":"10.1093\/jxb\/ern339","article-title":"The gene regulatory network for root epidermal cell-type pattern formation in Arabidopsis","volume":"60","author":"Schiefelbein","year":"2009","journal-title":"J. Exp. Bot."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"473","DOI":"10.1016\/S0092-8674(00)81536-6","article-title":"WEREWOLF, a MYB-related protein in Arabidopsis, is a position-dependent regulator of epidermal cell patterning","volume":"99","author":"Lee","year":"1999","journal-title":"Cell"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"740","DOI":"10.1006\/dbio.1994.1352","article-title":"The TTG gene is required to specify epidermal cell fate and cell patterning in the Arabidopsis root","volume":"166","author":"Galway","year":"1994","journal-title":"Dev. Biol."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"6431","DOI":"10.1242\/dev.00880","article-title":"The bHLH genes GLABRA3 (GL3) and ENHANCER OF GLABRA3 (EGL3) specify epidermal cell fate in the Arabidopsis root","volume":"130","author":"Bernhardt","year":"2003","journal-title":"Development"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"4859","DOI":"10.1242\/dev.00681","article-title":"A network of redundant bHLH proteins functions in all TTG1-dependent pathways of Arabidopsis","volume":"130","author":"Zhang","year":"2003","journal-title":"Development"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"5387","DOI":"10.1242\/dev.02139","article-title":"Cell-to-cell movement of the CAPRICE protein in Arabidopsis root epidermal cell differentiation","volume":"132","author":"Kurata","year":"2005","journal-title":"Development"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"1196","DOI":"10.1104\/pp.111.185785","article-title":"Cell fate in the Arabidopsis root epidermis is determined by competition between WEREWOLF and CAPRICE","volume":"157","author":"Song","year":"2011","journal-title":"Plant Physiol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"291","DOI":"10.1242\/dev.01565","article-title":"The bHLH genes GL3 and EGL3 participate in an intercellular regulatory circuit that controls cell patterning in the Arabidopsis root epidermis","volume":"132","author":"Bernhardt","year":"2005","journal-title":"Development"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"e02525","DOI":"10.7554\/eLife.02525","article-title":"Brassinosteroids control root epidermal cell fate via direct regulation of a MYB-bHLH-WD40 complex by GSK3-like kinases","volume":"3","author":"Cheng","year":"2014","journal-title":"eLife"},{"key":"ref_15","first-page":"1505","article-title":"Hormones act downstream of TTG and GL2 to promote root hair outgrowth during epidermis development in the Arabidopsis root","volume":"8","author":"Masucci","year":"1996","journal-title":"Plant Cell"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1335","DOI":"10.1104\/pp.106.4.1335","article-title":"The rhd6 Mutation of Arabidopsis thaliana Alters Root-Hair Initiation through an Auxin- and Ethylene-Associated Process","volume":"106","author":"Masucci","year":"1994","journal-title":"Plant Physiol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"1477","DOI":"10.1126\/science.1142618","article-title":"An ancient mechanism controls the development of cells with a rooting function in land plants","volume":"316","author":"Menand","year":"2007","journal-title":"Science"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"9571","DOI":"10.1073\/pnas.1305457110","article-title":"Recruitment and remodeling of an ancient gene regulatory network during land plant evolution","volume":"110","author":"Pires","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"264","DOI":"10.1038\/ng.529","article-title":"A basic helix-loop-helix transcription factor controls cell growth and size in root hairs","volume":"42","author":"Yi","year":"2010","journal-title":"Nat. Genet."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"1049","DOI":"10.1105\/tpc.19.00617","article-title":"Arabidopsis JAZ Proteins Interact with and Suppress RHD6 Transcription Factor to Regulate Jasmonate-Stimulated Root Hair Development","volume":"32","author":"Han","year":"2020","journal-title":"Plant Cell"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1105\/tpc.16.00732","article-title":"Tracheophytes Contain Conserved Orthologs of a Basic Helix-Loop-Helix Transcription Factor That Modulate ROOT HAIR SPECIFIC Genes","volume":"29","author":"Hwang","year":"2017","journal-title":"Plant Cell"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"13834","DOI":"10.1073\/pnas.1711723115","article-title":"Ethylene promotes root hair growth through coordinated EIN3\/EIL1 and RHD6\/RSL1 activity in Arabidopsis","volume":"114","author":"Feng","year":"2017","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"944","DOI":"10.1111\/nph.14095","article-title":"ROOT HAIR DEFECTIVE SIX-LIKE4 (RSL4) promotes root hair elongation by transcriptionally regulating the expression of genes required for cell growth","volume":"212","author":"Vijayakumar","year":"2016","journal-title":"New Phytol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"5289","DOI":"10.1073\/pnas.1701536114","article-title":"Molecular link between auxin and ROS-mediated polar growth","volume":"114","author":"Mangano","year":"2017","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"dev160887","DOI":"10.1242\/dev.160887","article-title":"TOR signaling in plants: Conservation and innovation","volume":"145","author":"Shi","year":"2018","journal-title":"Development"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"727","DOI":"10.7164\/antibiotics.28.727","article-title":"Rapamycin (AY-22,989), a new antifungal antibiotic. II. Fermentation, isolation and characterization","volume":"28","author":"Sehgal","year":"1975","journal-title":"J. Antibiot."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"585","DOI":"10.1016\/0092-8674(93)90144-F","article-title":"Target of rapamycin in yeast, TOR2, is an essential phosphatidylinositol kinase homolog required for G1 progression","volume":"73","author":"Kunz","year":"1993","journal-title":"Cell"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1016\/0092-8674(94)90570-3","article-title":"RAFT1: A mammalian protein that binds to FKBP12 in a rapamycin-dependent fashion and is homologous to yeast TORs","volume":"78","author":"Sabatini","year":"1994","journal-title":"Cell"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"274","DOI":"10.1016\/j.cell.2012.03.017","article-title":"mTOR signaling in growth control and disease","volume":"149","author":"Laplante","year":"2012","journal-title":"Cell"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"6422","DOI":"10.1073\/pnas.092141899","article-title":"Expression and disruption of the Arabidopsis TOR (target of rapamycin) gene","volume":"99","author":"Menand","year":"2002","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"499","DOI":"10.1104\/pp.113.229948","article-title":"The Role of Target of Rapamycin Signaling Networks in Plant Growth and Metabolism","volume":"164","author":"Xiong","year":"2014","journal-title":"Plant Physiol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"511","DOI":"10.1046\/j.1365-313X.1998.00232.x","article-title":"Molecular characterization of a plant FKBP12 that does not mediate action of FK506 and rapamycin","volume":"15","author":"Xu","year":"1998","journal-title":"Plant J."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"864","DOI":"10.1038\/sj.embor.7401043","article-title":"The Arabidopsis TOR kinase links plant growth, yield, stress resistance and mRNA translation","volume":"8","author":"Deprost","year":"2007","journal-title":"EMBO Rep."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"181","DOI":"10.1038\/nature12030","article-title":"Glucose-TOR signalling reprograms the transcriptome and activates meristems","volume":"496","author":"Xiong","year":"2013","journal-title":"Nature"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"4850","DOI":"10.1105\/tpc.112.107144","article-title":"Target of rapamycin signaling regulates metabolism, growth, and life span in Arabidopsis","volume":"24","author":"Ren","year":"2012","journal-title":"Plant Cell"},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Sormani, R., Yao, L., Menand, B., Ennar, N., Lecampion, C., Meyer, C., and Robaglia, C. (2007). Saccharomyces cerevisiae FKBP12 binds Arabidopsis thaliana TOR and its expression in plants leads to rapamycin susceptibility. BMC Plant Biol., 7.","DOI":"10.1186\/1471-2229-7-26"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"897","DOI":"10.1111\/tpj.12080","article-title":"Systemic analysis of inducible target of rapamycin mutants reveal a general metabolic switch controlling growth in Arabidopsis thaliana","volume":"73","author":"Caldana","year":"2013","journal-title":"Plant J."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"4361","DOI":"10.1093\/jxb\/ert242","article-title":"ATP-competitive mTOR kinase inhibitors delay plant growth by triggering early differentiation of meristematic cells but no developmental patterning change","volume":"64","author":"Menand","year":"2013","journal-title":"J. Exp. Bot."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"233","DOI":"10.1111\/nph.14118","article-title":"Brassinosteriod Insensitive 2 (BIN2) acts as a downstream effector of the Target of Rapamycin (TOR) signaling pathway to regulate photoautotrophic growth in Arabidopsis","volume":"213","author":"Xiong","year":"2017","journal-title":"New Phytol."},{"key":"ref_40","doi-asserted-by":"crossref","unstructured":"Deng, K., Dong, P., Wang, W., Feng, L., Xiong, F., Wang, K., Zhang, S., Feng, S., Wang, B., and Zhang, J. (2017). The TOR Pathway Is Involved in Adventitious Root Formation in Arabidopsis and Potato. Front. Plant Sci., 8.","DOI":"10.3389\/fpls.2017.00784"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"316","DOI":"10.1038\/s41477-019-0378-z","article-title":"Capturing the phosphorylation and protein interaction landscape of the plant TOR kinase","volume":"5","author":"Han","year":"2019","journal-title":"Nat. Plants"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"261","DOI":"10.1146\/annurev-arplant-043014-114648","article-title":"TOR Signaling and Nutrient Sensing","volume":"67","author":"Dobrenel","year":"2016","journal-title":"Annu. Rev. Plant Biol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"100","DOI":"10.1016\/j.molcel.2017.12.002","article-title":"Reciprocal Regulation of the TOR Kinase and ABA Receptor Balances Plant Growth and Stress Response","volume":"69","author":"Wang","year":"2018","journal-title":"Mol. Cell"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"2765","DOI":"10.1073\/pnas.1618782114","article-title":"Differential TOR activation and cell proliferation in Arabidopsis root and shoot apexes","volume":"114","author":"Li","year":"2017","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"886","DOI":"10.15252\/embj.201694816","article-title":"GTPase ROP2 binds and promotes activation of target of rapamycin, TOR, in response to auxin","volume":"36","author":"Schepetilnikov","year":"2017","journal-title":"EMBO J."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"185","DOI":"10.1105\/tpc.110.074005","article-title":"The PP2A regulatory subunit Tap46, a component of the TOR signaling pathway, modulates growth and metabolism in plants","volume":"23","author":"Ahn","year":"2011","journal-title":"Plant Cell"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"477","DOI":"10.1105\/tpc.105.035931","article-title":"Arabidopsis TARGET OF RAPAMYCIN interacts with RAPTOR, which regulates the activity of S6 kinase in response to osmotic stress signals","volume":"18","author":"Mahfouz","year":"2006","journal-title":"Plant Cell"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"3696","DOI":"10.1016\/j.celrep.2019.05.074","article-title":"Mutations of the AtYAK1 Kinase Suppress TOR Deficiency in Arabidopsis","volume":"27","author":"Forzani","year":"2019","journal-title":"Cell Rep."},{"key":"ref_49","doi-asserted-by":"crossref","unstructured":"Dobrenel, T., Mancera-Martinez, E., Forzani, C., Azzopardi, M., Davanture, M., Moreau, M., Schepetilnikov, M., Chicher, J., Langella, O., and Zivy, M. (2016). The Arabidopsis TOR Kinase Specifically Regulates the Expression of Nuclear Genes Coding for Plastidic Ribosomal Proteins and the Phosphorylation of the Cytosolic Ribosomal Protein S6. Front. Plant Sci., 7.","DOI":"10.3389\/fpls.2016.01611"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"827","DOI":"10.1093\/jxb\/eru438","article-title":"Overexpression of the PP2A regulatory subunit Tap46 leads to enhanced plant growth through stimulation of the TOR signalling pathway","volume":"66","author":"Ahn","year":"2015","journal-title":"J. Exp. Bot."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"1854","DOI":"10.1016\/j.cub.2016.05.005","article-title":"TOR Signaling Promotes Accumulation of BZR1 to Balance Growth with Carbon Availability in Arabidopsis","volume":"26","author":"Zhang","year":"2016","journal-title":"Curr. Biol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"2836","DOI":"10.1074\/jbc.M111.300749","article-title":"Rapamycin and glucose-target of rapamycin (TOR) protein signaling in plants","volume":"287","author":"Xiong","year":"2012","journal-title":"J. Biol. Chem."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"1898","DOI":"10.1105\/tpc.109.073007","article-title":"The TOR pathway modulates the structure of cell walls in Arabidopsis","volume":"22","author":"Leiber","year":"2010","journal-title":"Plant Cell"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"2507736","DOI":"10.1080\/15592324.2025.2507736","article-title":"Target of Rapamycin is involved in root hair development in Phaseolus vulgaris","volume":"20","author":"Arthikala","year":"2025","journal-title":"Plant Signal. Behav."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"553","DOI":"10.1046\/j.1365-313x.1998.00321.x","article-title":"Auxin and ethylene promote root hair elongation in Arabidopsis","volume":"16","author":"Pitts","year":"1998","journal-title":"Plant J."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"28","DOI":"10.1104\/pp.112.206441","article-title":"iRootHair: A comprehensive root hair genomics database","volume":"161","author":"Kwasniewski","year":"2013","journal-title":"Plant Physiol."},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"Bruex, A., Kainkaryam, R.M., Wieckowski, Y., Kang, Y.H., Bernhardt, C., Xia, Y., Zheng, X., Wang, J.Y., Lee, M.M., and Benfey, P. (2012). A gene regulatory network for root epidermis cell differentiation in Arabidopsis. PLoS Genet., 8.","DOI":"10.1371\/journal.pgen.1002446"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"1128","DOI":"10.1101\/gad.200201","article-title":"The chimeric leucine-rich repeat\/extensin cell wall protein LRX1 is required for root hair morphogenesis in Arabidopsis thaliana","volume":"15","author":"Baumberger","year":"2001","journal-title":"Genes Dev."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"442","DOI":"10.1038\/nature01485","article-title":"Reactive oxygen species produced by NADPH oxidase regulate plant cell growth","volume":"422","author":"Foreman","year":"2003","journal-title":"Nature"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"648","DOI":"10.1016\/j.molp.2019.04.004","article-title":"A Single-Cell RNA Sequencing Profiles the Developmental Landscape of Arabidopsis Root","volume":"12","author":"Zhang","year":"2019","journal-title":"Mol. Plant"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"2958","DOI":"10.1105\/tpc.106.045229","article-title":"Functional conservation of a root hair cell-specific cis-element in angiosperms with different root hair distribution patterns","volume":"18","author":"Kim","year":"2006","journal-title":"Plant Cell"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"3237","DOI":"10.1105\/tpc.006437","article-title":"Regulation of root hair initiation and expansin gene expression in Arabidopsis","volume":"14","author":"Cho","year":"2002","journal-title":"Plant Cell"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"1087","DOI":"10.1038\/emboj.2013.61","article-title":"TOR and S6K1 promote translation reinitiation of uORF-containing mRNAs via phosphorylation of eIF3h","volume":"32","author":"Schepetilnikov","year":"2013","journal-title":"EMBO J."},{"key":"ref_64","doi-asserted-by":"crossref","unstructured":"Deng, K., Yu, L., Zheng, X., Zhang, K., Wang, W., Dong, P., Zhang, J., and Ren, M. (2016). Target of Rapamycin Is a Key Player for Auxin Signaling Transduction in Arabidopsis. Front. Plant Sci., 7.","DOI":"10.3389\/fpls.2016.00291"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"32223","DOI":"10.1073\/pnas.2015400117","article-title":"Glucose-TOR signaling regulates PIN2 stability to orchestrate auxin gradient and cell expansion in Arabidopsis root","volume":"117","author":"Yuan","year":"2020","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_66","doi-asserted-by":"crossref","unstructured":"Retzer, K., and Weckwerth, W. (2021). The TOR-Auxin Connection Upstream of Root Hair Growth. Plants, 10.","DOI":"10.3390\/plants10010150"},{"key":"ref_67","doi-asserted-by":"crossref","unstructured":"Zhuo, F., Xiong, F., Deng, K., Li, Z., and Ren, M. (2020). Target of Rapamycin (TOR) Negatively Regulates Ethylene Signals in Arabidopsis. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21082680"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"288","DOI":"10.1038\/s41586-021-03310-y","article-title":"The TOR\u2013EIN2 axis mediates nuclear signalling to modulate plant growth","volume":"591","author":"Fu","year":"2021","journal-title":"Nature"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"6081","DOI":"10.1111\/pce.70601","article-title":"Auxin-Dependent Activation of RHD6-RSL4 Cascade Promotes Root Hair Growth Under Boron Deficiency in Arabidopsis Primary Root Apices","volume":"49","author":"Bahamonde","year":"2026","journal-title":"Plant Cell Environ."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"228","DOI":"10.1016\/j.cub.2021.10.044","article-title":"KAI2 promotes Arabidopsis root hair elongation at low external phosphate by controlling local accumulation of AUX1 and PIN2","volume":"32","author":"Maisch","year":"2022","journal-title":"Curr. Biol."},{"key":"ref_71","doi-asserted-by":"crossref","unstructured":"Song, L., Yu, H., Dong, J., Che, X., Jiao, Y., and Liu, D. (2016). The Molecular Mechanism of Ethylene-Mediated Root Hair Development Induced by Phosphate Starvation. PLoS Genet., 12.","DOI":"10.1371\/journal.pgen.1006194"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"389","DOI":"10.1046\/j.1365-313x.1999.00607.x","article-title":"The RCN1-encoded A subunit of protein phosphatase 2A increases phosphatase activity in vivo","volume":"20","author":"Deruere","year":"1999","journal-title":"Plant J."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"641","DOI":"10.1038\/nprot.2006.97","article-title":"Agrobacterium-mediated transformation of Arabidopsis thaliana using the floral dip method","volume":"1","author":"Zhang","year":"2006","journal-title":"Nat. Protoc."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"550","DOI":"10.1186\/s13059-014-0550-8","article-title":"Moderated estimation of fold change and dispersion for RNA-seq data with DESeq2","volume":"15","author":"Love","year":"2014","journal-title":"Genome Biol."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"139","DOI":"10.1093\/bioinformatics\/btp616","article-title":"edgeR: A Bioconductor package for differential expression analysis of digital gene expression data","volume":"26","author":"Robinson","year":"2010","journal-title":"Bioinformatics"}],"container-title":["Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/17\/2586\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,25]],"date-time":"2026-08-25T08:23:43Z","timestamp":1787646223000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/17\/2586"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,25]]},"references-count":75,"journal-issue":{"issue":"17","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["plants15172586"],"URL":"https:\/\/doi.org\/10.3390\/plants15172586","relation":{},"ISSN":["2223-7747"],"issn-type":[{"value":"2223-7747","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,25]]}}}