{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,22]],"date-time":"2026-08-22T04:26:55Z","timestamp":1787372815526,"version":"build-2736575974"},"reference-count":98,"publisher":"MDPI AG","issue":"16","license":[{"start":{"date-parts":[[2026,8,20]],"date-time":"2026-08-20T00:00:00Z","timestamp":1787184000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Biology"],"abstract":"<jats:p>Cuticular wax is a critical component of the plant cuticle, playing an indispensable role in plant growth, development, and defense against environmental stresses. ECERIFERUM3 (CER3), a very-long-chain fatty acyl-CoA reductase, serves as a pivotal hub channeling carbon resources into the alkane- and alcohol-forming pathways respectively. Here, we systematically characterized the protein structural features and evolutionary patterns of CER3. We also comprehensively reviewed the multiple regulatory mechanisms governing CER3, encompassing transcriptional, post-transcriptional, translational, and epigenetic modifications, with particular emphasis on the definitive finding that CER3 modulates cuticular wax biosynthetic flux by assembling distinct protein complexes. Finally, existing knowledge gaps and future research directions are also discussed.<\/jats:p>","DOI":"10.3390\/biology15161437","type":"journal-article","created":{"date-parts":[[2026,8,20]],"date-time":"2026-08-20T16:05:19Z","timestamp":1787241919000},"page":"1437","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Protein Structure, Evolution and Regulation of CER3, a Core Enzyme Partitioning Carbon into Two Specific Wax Biosynthetic Pathways"],"prefix":"10.3390","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0009-0007-8252-8376","authenticated-orcid":false,"given":"Qingqing","family":"Niu","sequence":"first","affiliation":[{"name":"State Key Laboratory of Biocatalysis and Enzyme Engineering, School of Life Sciences, Hubei University, Wuhan 430062, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Limei","family":"Liu","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Biocatalysis and Enzyme Engineering, School of Life Sciences, Hubei University, Wuhan 430062, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chang","family":"Liu","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Biocatalysis and Enzyme Engineering, School of Life Sciences, Hubei University, Wuhan 430062, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0449-2471","authenticated-orcid":false,"given":"Shiyou","family":"L\u00fc","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Biocatalysis and Enzyme Engineering, School of Life Sciences, Hubei University, Wuhan 430062, China"},{"name":"Hubei Hongshan Laboratory, Wuhan 430074, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0006-3903-8307","authenticated-orcid":false,"given":"Hui","family":"Zhang","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Biocatalysis and Enzyme Engineering, School of Life Sciences, Hubei University, Wuhan 430062, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,8,20]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1104\/pp.113.222737","article-title":"The Formation and Function of Plant Cuticles","volume":"163","author":"Yeats","year":"2013","journal-title":"Plant Physiol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"1120","DOI":"10.1104\/pp.112.198473","article-title":"Composition and Physiological Function of the Wax Layers Coating Arabidopsis Leaves: \u03b2-Amyrin Negatively Affects the Intracuticular Water Barrier","volume":"160","author":"Buschhaus","year":"2012","journal-title":"Plant Physiol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"75","DOI":"10.1071\/FP02159","article-title":"The maize epicuticular wax layer provides UV protection","volume":"30","author":"Long","year":"2003","journal-title":"Funct. Plant Biol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"1649","DOI":"10.1104\/pp.105.070805","article-title":"Cuticular Lipid Composition, Surface Structure, and Gene Expression in Arabidopsis Stem Epidermis","volume":"139","author":"Suh","year":"2005","journal-title":"Plant Physiol."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"3015","DOI":"10.1105\/tpc.106.042044","article-title":"Wax-deficient anther1 is involved in cuticle and wax production in rice anther walls and is required for pollen development","volume":"18","author":"Jung","year":"2006","journal-title":"Plant Cell"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"1880","DOI":"10.1111\/nph.16741","article-title":"GDSL lipase occluded stomatal pore 1 is required for wax biosynthesis and stomatal cuticular ledge formation","volume":"228","author":"Tang","year":"2020","journal-title":"New Phytol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1046\/j.1365-313X.2003.01854.x","article-title":"The YORE-YORE gene regulates multiple aspects of epidermal cell differentiation in","volume":"36","author":"Kurata","year":"2003","journal-title":"Plant J."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1170","DOI":"10.1105\/tpc.010926","article-title":"Cloning and characterization of the WAX2 gene of Arabidopsis involved in cuticle membrane and wax production","volume":"15","author":"Chen","year":"2003","journal-title":"Plant Cell"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"841","DOI":"10.1093\/jxb\/erq366","article-title":"Composition differences between epicuticular and intracuticular wax substructures: How do plants seal their epidermal surfaces?","volume":"62","author":"Buschhaus","year":"2011","journal-title":"J. Exp. Bot."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"246","DOI":"10.1093\/mp\/sss159","article-title":"Recent Advances in Cuticular Wax Biosynthesis and Its Regulation in Arabidopsis","volume":"6","author":"Lee","year":"2013","journal-title":"Mol. Plant"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"159","DOI":"10.1038\/s41477-024-01897-4","article-title":"How plants adapt surface lipids to environmental changes","volume":"11","author":"Dang","year":"2025","journal-title":"Nat. Plants"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"3632","DOI":"10.1111\/pce.15376","article-title":"In a Different Light: Irradiation-Induced Cuticular Wax Accumulation Fails to Reduce Cuticular Transpiration","volume":"48","author":"Herzig","year":"2025","journal-title":"Plant Cell Env."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"557","DOI":"10.1007\/s00299-015-1772-2","article-title":"Advances in the understanding of cuticular waxes in Arabidopsis thaliana and crop species","volume":"34","author":"Lee","year":"2015","journal-title":"Plant Cell Rep."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"eadz0135","DOI":"10.1126\/sciadv.adz0135","article-title":"Molecular basis of very-long-chain fatty acid elongation by the CER6-GL2 enzyme complex in plant wax biosynthesis","volume":"11","author":"Liu","year":"2025","journal-title":"Sci. Adv."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"1164","DOI":"10.1104\/pp.112.201640","article-title":"Arabidopsis ECERIFERUM2 Is a Component of the Fatty Acid Elongation Machinery Required for Fatty Acid Extension to Exceptional Lengths","volume":"160","author":"Haslam","year":"2012","journal-title":"Plant Physiol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"485","DOI":"10.1111\/j.1365-313X.2007.03252.x","article-title":"Characterization of Arabidopsis ABCG11\/WBC11, an ATP binding cassette (ABC) transporter that is required for cuticular lipid secretion","volume":"52","author":"Bird","year":"2007","journal-title":"Plant J."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"3126","DOI":"10.1093\/jxb\/eraa046","article-title":"Drought stress modulates cuticular wax composition of the grape berry","volume":"71","author":"Dimopoulos","year":"2020","journal-title":"J. Exp. Bot."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"4643","DOI":"10.1111\/pce.15465","article-title":"The Function of PpKCS6 in Regulating Cuticular Wax Synthesis and Drought Resistance of Kentucky Bluegrass","volume":"48","author":"Jiang","year":"2025","journal-title":"Plant Cell Env."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"702","DOI":"10.1126\/science.1102331","article-title":"Plant cuticular lipid export requires an ABC transporter","volume":"306","author":"Pighin","year":"2004","journal-title":"Science"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"3012","DOI":"10.1111\/pce.15320","article-title":"Sec24C Participates in Cuticular Wax Transport by Facilitating Plasma Membrane Localization of ABCG5","volume":"48","author":"Zhao","year":"2025","journal-title":"Plant Cell Env."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"8518","DOI":"10.1111\/pce.70144","article-title":"XsLTPG31 Confers Leaf Cuticular Wax Deposition and Drought Resistance in Yellowhorn","volume":"48","author":"Xu","year":"2025","journal-title":"Plant Cell Env."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"3106","DOI":"10.1105\/tpc.112.099796","article-title":"Reconstitution of Plant Alkane Biosynthesis in Yeast Demonstrates That Arabidopsis ECERIFERUM1 and ECERIFERUM3 Are Core Components of a Very-Long-Chain Alkane Synthesis Complex","volume":"24","author":"Bernard","year":"2012","journal-title":"Plant Cell"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"459","DOI":"10.1046\/j.1365-313X.1996.10030459.x","article-title":"The CER3 gene of Arabidopsis thaliana is expressed in leaves, stems, roots, flowers and apical meristems","volume":"10","author":"Hannoufa","year":"2002","journal-title":"Plant J."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"3538","DOI":"10.1016\/j.febslet.2007.06.065","article-title":"The CER3 wax biosynthetic gene from Arabidopsis thaliana is allelic to WAX2\/YRE\/FLP1","volume":"581","author":"Rowland","year":"2007","journal-title":"FEBS Lett."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1038\/s41477-024-01892-9","article-title":"Deciphering the core shunt mechanism in Arabidopsis cuticular wax biosynthesis and its role in plant environmental adaptation","volume":"11","author":"Li","year":"2025","journal-title":"Nat. Plants"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"2799","DOI":"10.1093\/jxb\/erab509","article-title":"Regulatory mechanisms underlying cuticular wax biosynthesis","volume":"73","author":"Lee","year":"2022","journal-title":"J. Exp. Bot."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"698","DOI":"10.1111\/nph.16571","article-title":"Wax biosynthesis in response to danger: Its regulation upon abiotic and biotic stress","volume":"227","author":"Lewandowska","year":"2020","journal-title":"New Phytol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"4","DOI":"10.1007\/s11295-023-01637-3","article-title":"Insights into the evolution of the eucalypt CER1 and CER3 genes involved in the synthesis of alkane waxes","volume":"20","author":"Vaillancourt","year":"2024","journal-title":"Tree Genet. Genomes"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"405","DOI":"10.1146\/annurev.arplant.47.1.405","article-title":"Biochemistry and molecular biology of wax production in plants","volume":"47","author":"PostBeittenmiller","year":"1996","journal-title":"Annu. Rev. Plant Physiol. Plant Mol. Biol."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"107101","DOI":"10.1016\/j.ympev.2021.107101","article-title":"Origin and diversification of ECERIFERUM1 (CER1) and ECERIFERUM3 (CER3) genes in land plants and phylogenetic evidence that the ancestral CER1\/3 gene resulted from the fusion of pre-existing domains","volume":"159","author":"Chaudhary","year":"2021","journal-title":"Mol. Phylogenet. Evol."},{"key":"ref_31","first-page":"2115","article-title":"Molecular characterization of the CER1 gene of arabidopsis involved in epicuticular wax biosynthesis and pollen fertility","volume":"7","author":"Aarts","year":"1995","journal-title":"Plant Cell"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"611","DOI":"10.1146\/annurev.arplant.49.1.611","article-title":"Desaturation and related modifications of fatty acids","volume":"49","author":"Shanklin","year":"1998","journal-title":"Annu. Rev. Plant Physiol. Plant Mol. Biol."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"391","DOI":"10.1046\/j.1365-313X.1996.09030391.x","article-title":"Isolation and characterization of an Arabidopsis thaliana cDNA encoding a Delta(7)-sterol-C-5-desaturase by functional complementation of a defective yeast mutant","volume":"9","author":"Gachotte","year":"1996","journal-title":"Plant J."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"731","DOI":"10.1007\/s00425-012-1791-y","article-title":"The CER22 gene required for the synthesis of cuticular wax alkanes in Arabidopsis thaliana is allelic to CER1","volume":"237","author":"Sakuradani","year":"2013","journal-title":"Planta"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"3895","DOI":"10.1007\/s00018-008-8588-y","article-title":"The SDR superfamily: Functional and structural diversity within a family of metabolic and regulatory enzymes","volume":"65","author":"Kavanagh","year":"2008","journal-title":"Cell. Mol. Life Sci."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"uhac004","DOI":"10.1093\/hr\/uhac004","article-title":"Tomato SlCER1\u20131 catalyzes the synthesis of wax alkanes, increasing drought tolerance and fruit storability","volume":"9","author":"Wu","year":"2022","journal-title":"Hortic. Res."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1007\/s00425-011-1481-1","article-title":"Wax crystal-sparse leaf2, a rice homologue of WAX2\/GL1, is involved in synthesis of leaf cuticular wax","volume":"235","author":"Mao","year":"2011","journal-title":"Planta"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.compbiolchem.2019.02.009","article-title":"Molecular evolution of the plant ECERIFERUM1 and ECERIFERUM3 genes involved in aliphatic hydrocarbon production","volume":"80","author":"Wang","year":"2019","journal-title":"Comput. Biol. Chem."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"1311","DOI":"10.1111\/nph.71257","article-title":"The algal homolog of the plant CER1 and CER3 proteins is a bifunctional hydrocarbon-forming enzyme","volume":"251","author":"Six","year":"2026","journal-title":"New Phytol."},{"key":"ref_40","doi-asserted-by":"crossref","unstructured":"Youssif, N.E.E., Yang, B., Huang, H., Amar, M.H., Ezzat, M., Belal, M., Zaghlool, S.A.M., Zhao, H., Fu, D., and L\u00fc, S. (2026). Comparative Genomic Analysis and Functional Identification of CER1 and CER3 Homologs in Rice Wax Synthesis. Biology, 15.","DOI":"10.3390\/biology15020166"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"1646","DOI":"10.1111\/nph.71033","article-title":"Transcription factor bZIP48-MYB96 module positively regulates cuticular wax synthesis by directly activating KCSs and CER3 genes in response to cold stress","volume":"250","author":"Liu","year":"2026","journal-title":"New Phytol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"1264","DOI":"10.1111\/pce.70300","article-title":"BOUNDARY OF ROP DOMAIN3 Modulates Salt Tolerance by Mediating Cuticle Wax Synthesis","volume":"49","author":"Miao","year":"2026","journal-title":"Plant Cell Env."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"2223","DOI":"10.1105\/tpc.19.00152","article-title":"The F-Box Protein SAGL1 and ECERIFERUM3 Regulate Cuticular Wax Biosynthesis in Response to Changes in Humidity in Arabidopsis","volume":"31","author":"Kim","year":"2019","journal-title":"Plant Cell"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"110","DOI":"10.1016\/j.plipres.2012.10.002","article-title":"Arabidopsis cuticular waxes: Advances in synthesis, export and regulation","volume":"52","author":"Bernard","year":"2013","journal-title":"Prog. Lipid Res."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"1138","DOI":"10.1105\/tpc.111.083485","article-title":"The MYB96 Transcription Factor Regulates Cuticular Wax Biosynthesis under Drought Conditions in Arabidopsis","volume":"23","author":"Seo","year":"2011","journal-title":"Plant Cell"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"48","DOI":"10.1093\/pcp\/pcu142","article-title":"Cuticular Wax Biosynthesis is Up-Regulated by the MYB94 Transcription Factor in Arabidopsis","volume":"56","author":"Lee","year":"2015","journal-title":"Plant Cell Physiol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"2300","DOI":"10.1093\/pcp\/pcw147","article-title":"MYB94 and MYB96 Additively Activate Cuticular Wax Biosynthesis in Arabidopsis","volume":"57","author":"Lee","year":"2016","journal-title":"Plant Cell Physiol."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"752","DOI":"10.1105\/tpc.107.054858","article-title":"A MYB Transcription Factor Regulates Very-Long-Chain Fatty Acid Biosynthesis for Activation of the Hypersensitive Cell Death Response in Arabidopsis","volume":"20","author":"Raffaele","year":"2008","journal-title":"Plant Cell"},{"key":"ref_49","doi-asserted-by":"crossref","unstructured":"Yu, J., Lei, B., Zhao, H., Wang, B., Kakar, K.U., Guo, Y., Zhang, X., Jia, M., Yang, H., and Zhao, D. (2022). Cloning, characterization and functional analysis of NtMYB306a gene reveals its role in wax alkane biosynthesis of tobacco trichomes and stress tolerance. Front. Plant Sci., 13.","DOI":"10.3389\/fpls.2022.1005811"},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Wang, L., Xue, W., Li, X., Li, J., Wu, J., Xie, L., Kawabata, S., Li, Y., and Zhang, Y. (2020). EgMIXTA1, a MYB-Type Transcription Factor, Promotes Cuticular Wax Formation in Eustoma grandiflorum Leaves. Front. Plant Sci., 11.","DOI":"10.3389\/fpls.2020.524947"},{"key":"ref_51","first-page":"2133","article-title":"PtoMYB142, a poplar R2R3-MYB transcription factor, contributes to drought tolerance by regulating wax biosynthesis","volume":"42","author":"Song","year":"2022","journal-title":"Tree Physiol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"28","DOI":"10.1007\/s00425-022-03945-9","article-title":"AtMYB31 is a wax regulator associated with reproductive development in Arabidopsis","volume":"256","author":"Shi","year":"2022","journal-title":"Planta"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"179","DOI":"10.1111\/nph.17965","article-title":"Two R2R3-MYB genes cooperatively control trichome development and cuticular wax biosynthesis in Prunus persica","volume":"234","author":"Yang","year":"2022","journal-title":"New Phytol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"953","DOI":"10.1093\/jxb\/erab420","article-title":"CsMYB96 confers resistance to water loss in citrus fruit by simultaneous regulation of water transport and wax biosynthesis","volume":"73","author":"Zhang","year":"2022","journal-title":"J. Exp. Bot."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"2321","DOI":"10.1111\/nph.71095","article-title":"A cascade of BoABF2-BoMYB96-BoMIEL1 regulates wax accumulation in kale","volume":"250","author":"Li","year":"2026","journal-title":"New Phytol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"777","DOI":"10.1016\/j.plaphy.2021.06.045","article-title":"Ectopic expression of CsMYB30 from Citrus sinensis enhances salt and drought tolerance by regulating wax synthesis in Arabidopsis thaliana","volume":"166","author":"Wen","year":"2021","journal-title":"Plant Physiol. Biochem."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"120","DOI":"10.1016\/j.bbagrm.2011.09.002","article-title":"The role of WRKY transcription factors in plant abiotic stresses","volume":"1819","author":"Chen","year":"2012","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"2155","DOI":"10.1093\/jxb\/ert073","article-title":"Expression of wild soybean WRKY20 in Arabidopsis enhances drought tolerance and regulates ABA signalling","volume":"64","author":"Luo","year":"2013","journal-title":"J. Exp. Bot."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"170270","DOI":"10.1098\/rsob.170270","article-title":"Surveillance-ready transcription: Nuclear RNA decay as a default fate","volume":"8","author":"Bresson","year":"2018","journal-title":"Open Biol."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"518","DOI":"10.1038\/s41576-018-0013-2","article-title":"Controlling nuclear RNA levels","volume":"19","author":"Schmid","year":"2018","journal-title":"Nat. Rev. Genet"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"e1529","DOI":"10.1002\/wrna.1529","article-title":"Writing a wrong: Coupled RNA polymerase II transcription and RNA quality control","volume":"10","author":"Peck","year":"2019","journal-title":"Wiley Interdiscip. Rev."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"3125","DOI":"10.1016\/j.bbamcr.2016.09.023","article-title":"Cytoplasmic RNA decay pathways\u2014Enzymes and mechanisms","volume":"1863","author":"Labno","year":"2016","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"102014","DOI":"10.1016\/j.pbi.2021.102014","article-title":"The initiation of RNA interference (RNAi) in plants","volume":"61","author":"Hung","year":"2021","journal-title":"Curr. Opin. Plant Biol."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"1939","DOI":"10.1093\/pcp\/pcz067","article-title":"Transcription Is Just the Beginning of Gene Expression Regulation: The Functional Significance of RNA-Binding Proteins to Post-transcriptional Processes in Plants","volume":"60","author":"Prall","year":"2019","journal-title":"Plant Cell Physiol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"1340","DOI":"10.1016\/j.cell.2007.10.056","article-title":"Genome-wide high-resolution mapping of exosome substrates reveals hidden features in the Arabidopsis transcriptome","volume":"131","author":"Chekanova","year":"2007","journal-title":"Cell"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"409","DOI":"10.1016\/j.molcel.2012.08.018","article-title":"Extensive degradation of RNA precursors by the exosome in wild-type cells","volume":"48","author":"Gudipati","year":"2012","journal-title":"Mol. Cell"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"422","DOI":"10.1016\/j.molcel.2012.08.013","article-title":"Transcriptome-wide analysis of exosome targets","volume":"48","author":"Schneider","year":"2012","journal-title":"Mol. Cell"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"904","DOI":"10.1105\/tpc.106.049304","article-title":"A Core Subunit of the RNA-Processing\/Degrading Exosome Specifically Influences Cuticular Wax Biosynthesis in Arabidopsis","volume":"19","author":"Hooker","year":"2007","journal-title":"Plant Cell"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"1211","DOI":"10.1104\/pp.19.01002","article-title":"CER16 Inhibits Post-Transcriptional Gene Silencing of CER3 to Regulate Alkane Biosynthesis","volume":"182","author":"Yang","year":"2020","journal-title":"Plant Physiol."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"1385","DOI":"10.1104\/pp.112.199646","article-title":"RDR1 and SGS3, Components of RNA-Mediated Gene Silencing, Are Required for the Regulation of Cuticular Wax Biosynthesis in Developing Inflorescence Stems of Arabidopsis","volume":"159","author":"Lam","year":"2012","journal-title":"Plant Physiol."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"323","DOI":"10.1104\/pp.114.252825","article-title":"The Exosome and Trans-Acting Small Interfering RNAs Regulate Cuticular Wax Biosynthesis during Arabidopsis Inflorescence Stem Development","volume":"167","author":"Lam","year":"2015","journal-title":"Plant Physiol."},{"key":"ref_72","first-page":"960","article-title":"SUPERKILLER complex components are required for the RNA exosome-mediated control of cuticular wax biosynthesis in Arabidopsis inflorescence stems","volume":"171","author":"Zhao","year":"2016","journal-title":"Plant Physiol."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"3871","DOI":"10.1038\/s41467-019-11807-4","article-title":"RST1 and RIPR connect the cytosolic RNA exosome to the Ski complex in Arabidopsis","volume":"10","author":"Lange","year":"2019","journal-title":"Nat. Commun."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"950","DOI":"10.1016\/j.tplants.2020.06.009","article-title":"Degrade or Silence?\u2013RNA Turnover Takes Control of Epicuticular Wax Synthesis","volume":"25","year":"2020","journal-title":"Trends Plant Sci."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"kiaf153","DOI":"10.1093\/plphys\/kiaf153","article-title":"The SAGA histone acetyltransferase complex functions in concert with RNA processing machinery to regulate wheat wax biosynthesis","volume":"198","author":"Wang","year":"2025","journal-title":"Plant Physiol."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"112180","DOI":"10.1016\/j.plantsci.2024.112180","article-title":"A review on ubiquitin ligases: Orchestrators of plant resilience in adversity","volume":"347","author":"Suranjika","year":"2024","journal-title":"Plant Sci."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"e70846","DOI":"10.1111\/tpj.70846","article-title":"Ubiquitin ligases in plant immunity: Structural mechanisms and signaling","volume":"126","author":"Hamada","year":"2026","journal-title":"Plant J."},{"key":"ref_78","doi-asserted-by":"crossref","unstructured":"Liu, S., Tong, M., Zhao, L., Li, X., and Kunst, L. (2021). The ARRE RING-Type E3 Ubiquitin Ligase Negatively Regulates Cuticular Wax Biosynthesis in Arabidopsis thaliana by Controlling ECERIFERUM1 and ECERIFERUM3 Protein Levels. Front. Plant Sci., 12.","DOI":"10.3389\/fpls.2021.752309"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"1249","DOI":"10.1093\/pcp\/pcx065","article-title":"The MIEL1 E3 Ubiquitin Ligase Negatively Regulates Cuticular Wax Biosynthesis in Arabidopsis Stems","volume":"58","author":"Gil","year":"2017","journal-title":"Plant Cell Physiol."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"930","DOI":"10.1104\/pp.112.198697","article-title":"Arabidopsis ECERIFERUM9 Involvement in Cuticle Formation and Maintenance of Plant Water Status","volume":"159","author":"Zhao","year":"2012","journal-title":"Plant Physiol."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"102815","DOI":"10.1016\/j.pbi.2025.102815","article-title":"Plant histone acetyltransferase complexes: Conserved and plant-specific characteristics","volume":"88","author":"Xu","year":"2025","journal-title":"Curr. Opin. Plant Biol."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"413","DOI":"10.1007\/s11033-024-09300-3","article-title":"Recent advancements in the role of histone acetylation dynamics to improve stress responses in plants","volume":"51","author":"Bajpai","year":"2024","journal-title":"Mol. Biol. Rep."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"2911","DOI":"10.1093\/jxb\/ery077","article-title":"GCN5 contributes to stem cuticular wax biosynthesis by histone acetylation of CER3 in Arabidopsis","volume":"69","author":"Wang","year":"2018","journal-title":"J. Exp. Bot."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"1250","DOI":"10.1104\/pp.20.00603","article-title":"Epigenetic Activation of Enoyl-CoA Reductase By An Acetyltransferase Complex Triggers Wheat Wax Biosynthesis","volume":"183","author":"Kong","year":"2020","journal-title":"Plant Physiol."},{"key":"ref_85","first-page":"1640","article-title":"Global regulation of plant immunity by histone lysine methyl transferases","volume":"28","author":"Lee","year":"2016","journal-title":"Plant Cell"},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"455","DOI":"10.1093\/pcp\/pct182","article-title":"Histone H2B Monoubiquitination is Involved in the Regulation of Cutin and Wax Composition in Arabidopsis thaliana","volume":"55","author":"Verdier","year":"2014","journal-title":"Plant Cell Physiol."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"415","DOI":"10.1104\/pp.18.01075","article-title":"Arabidopsis CER1-LIKE1 Functions in a Cuticular Very-Long-Chain Alkane-Forming Complex","volume":"179","author":"Pascal","year":"2019","journal-title":"Plant Physiol."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"635","DOI":"10.1104\/pp.122.3.635","article-title":"Purification of a Jojoba Embryo Fatty Acyl-Coenzyme A Reductase and Expression of Its cDNA in High Erucic Acid Rapeseed","volume":"122","author":"Metz","year":"2000","journal-title":"Plant Physiol."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"866","DOI":"10.1104\/pp.106.086785","article-title":"CER4 encodes an alcohol-forming fatty acyl-coenzyme A reductase involved in cuticular wax production in Arabidopsis","volume":"142","author":"Rowland","year":"2006","journal-title":"Plant Physiol."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1104\/pp.111.172320","article-title":"Overexpression of Arabidopsis ECERIFERUM1 Promotes Wax Very-Long-Chain Alkane Biosynthesis and Influences Plant Response to Biotic and Abiotic Stresses","volume":"156","author":"Bourdenx","year":"2011","journal-title":"Plant Physiol."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"1574","DOI":"10.1111\/nph.18615","article-title":"Tree tobacco (Nicotiana glauca) cuticular wax composition is essential for leaf retention during drought, facilitating a speedy recovery following rewatering","volume":"237","author":"Negin","year":"2023","journal-title":"New Phytol."},{"key":"ref_92","doi-asserted-by":"crossref","unstructured":"Jenks, M.A., Hasegawa, P.M., and Jain, S.M. (2007). Eco-Physiological and Molecular-Genetic Determinants of Plant Cuticle Function in Drought and Salt Stress Tolerance. Advances in Molecular Breeding Toward Drought and Salt Tolerant Crops, Springer.","DOI":"10.1007\/978-1-4020-5578-2"},{"key":"ref_93","doi-asserted-by":"crossref","unstructured":"Borrego, E.J., and Kolomiets, M.V. (2016). Synthesis and Functions of Jasmonates in Maize. Plants, 5.","DOI":"10.3390\/plants5040041"},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"1619","DOI":"10.1111\/tpj.16701","article-title":"The Arabidopsis cytochrome P450 enzyme CYP96A4 is involved in the wound-induced biosynthesis of cuticular wax and cutin monomers","volume":"118","author":"Huang","year":"2024","journal-title":"Plant J."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"108877","DOI":"10.1016\/j.biotechadv.2026.108877","article-title":"From molecules to field: Integrated insights into cuticle-mediated drought tolerance in plants","volume":"89","author":"Khan","year":"2026","journal-title":"Biotechnol. Adv."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"118","DOI":"10.1016\/j.eng.2022.12.006","article-title":"Structures and Functions of Cuticular Wax in Postharvest Fruit and Its Regulation: A Comprehensive Review with Future Perspectives","volume":"23","author":"Wu","year":"2023","journal-title":"Engineering"},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"5369","DOI":"10.1093\/jxb\/erx341","article-title":"Breeding for cuticle-associated traits in crop species: Traits, targets, and strategies","volume":"68","author":"Petit","year":"2017","journal-title":"J. Exp. Bot."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"893","DOI":"10.1071\/FP06139","article-title":"Evans Review No. 3: Structure-function relationships of the plant cuticle and cuticular waxes\u2014A smart material?","volume":"33","author":"Bargel","year":"2006","journal-title":"Funct. Plant Biol."}],"container-title":["Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2079-7737\/15\/16\/1437\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,22]],"date-time":"2026-08-22T04:14:56Z","timestamp":1787372096000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2079-7737\/15\/16\/1437"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,20]]},"references-count":98,"journal-issue":{"issue":"16","published-online":{"date-parts":[[2026,8]]}},"alternative-id":["biology15161437"],"URL":"https:\/\/doi.org\/10.3390\/biology15161437","relation":{},"ISSN":["2079-7737"],"issn-type":[{"value":"2079-7737","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,20]]}}}