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Disruptions in vascular bundle formation can therefore lead to severe physiological and developmental defects. In maize, ethyl methanesulfonate (EMS)\u2010induced dominant nonallelic\n                    <jats:italic>Wilty<\/jats:italic>\n                    mutants exhibit a pronounced wilting phenotype even under well\u2010watered conditions, indicating underlying defects in vascular function. In this study, we characterized the\n                    <jats:italic>Wi3<\/jats:italic>\n                    mutant, identified as\n                    <jats:italic>ZmDrought\u2010Overly\u2010Sensitive1\/DOS1<\/jats:italic>\n                    , and compared it with the previously described\n                    <jats:italic>Wi2<\/jats:italic>\n                    mutant to uncover shared mechanisms underlying their phenotypes. We provide evidence, by bulk segregant resequencing linkage disequilibrium of SNPs adjacent to the causal\n                    <jats:italic>Wilty<\/jats:italic>\n                    SNPs in respective \u00df\u2010 and \u03b1\u2010tubulin genes, for the personal communication from Gerry Neuffer that\n                    <jats:italic>Wi2\/\u00df\u2010tub6<\/jats:italic>\n                    provenance is from ACR\u2010related stock, whereas\n                    <jats:italic>Wi3\/\u03b1\u2010tub4<\/jats:italic>\n                    allele is from Mo17, not B73 as claimed by the authors who cloned\n                    <jats:italic>Dos1<\/jats:italic>\n                    . Histochemical staining and Fourier\u2010transform infrared (FTIR) spectroscopy of vascular bundles in\n                    <jats:italic>Wi3<\/jats:italic>\n                    indicated apparent alterations in cellulose and lignin content consistent with those observed in\n                    <jats:italic>Wi2<\/jats:italic>\n                    . Transcriptome analysis of shoot meristems further indicated that similar sets of genes and pathways are differentially expressed in both mutants, suggesting convergence on common biological pathways. Using bulk\u2010segregant whole\u2010genome resequencing, we identified\n                    <jats:italic>alpha\u2010tubulin4<\/jats:italic>\n                    (\n                    <jats:italic>TUA4<\/jats:italic>\n                    ) as the causal gene in\n                    <jats:italic>Wi3<\/jats:italic>\n                    (\n                    <jats:italic>ZmDOS1<\/jats:italic>\n                    ), harboring a C\u2010to\u2010T substitution within the N\u2010terminal GTPase\u2010binding domain. This mutation results in a glutamic acid196\u2010to\u2010lysine substitution. Given that \u03b1\u2010 and \u03b2\u2010tubulin subunits heterodimerize, and in many plants and animal mutant alleles are dominant\u2010negative gains\u2010of\u2010function, we infer\n                    <jats:italic>Wi2<\/jats:italic>\n                    ,\n                    <jats:italic>Wi3<\/jats:italic>\n                    , and likely\n                    <jats:italic>Wi4<\/jats:italic>\n                    , based on very similar FTIR biophysical difference spectra, may act as effectors of vascular bundle cell wall deposition, potentially involving vesicle trafficking as recently shown for asymmetric cell divisions in maize stomatal development. Together, these findings highlight the functional interdependence of tubulin subunits and provide a plausible mechanistic framework for the striking biophysical, transcriptomic, and phenotypic similarities observed between\n                    <jats:italic>Wi2<\/jats:italic>\n                    ,\n                    <jats:italic>Wi3<\/jats:italic>\n                    \/\n                    <jats:italic>ZmDOS1<\/jats:italic>\n                    , and\n                    <jats:italic>Wi4<\/jats:italic>\n                    mutants.\n                  <\/jats:p>","DOI":"10.1002\/pld3.70189","type":"journal-article","created":{"date-parts":[[2026,9,2]],"date-time":"2026-09-02T14:39:25Z","timestamp":1788359965000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["<i>\n                      <scp>Zea mays<\/scp>\n                      Drought\u2010Overly Sensitive1\/TUBA4\n                    <\/i>\n                    Is\n                    <i>Wilty3<\/i>\n                    , and Transcriptome Co\u2010Expression Analysis of Shoot Meristem Mutant Tissues Reveals\n                    <i>Wilty2\/TUB6:Wi3<\/i>\n                    Interactions Associated With Stem Vascular Bundle Development"],"prefix":"10.1002","volume":"10","author":[{"given":"Anuradha","family":"Dhingra","sequence":"first","affiliation":[{"name":"Department of Biological Sciences Texas Tech University  Lubbock Texas USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Indra","family":"Adhikari","sequence":"additional","affiliation":[{"name":"Department of Biological Sciences Texas Tech University  Lubbock Texas USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2090-0650","authenticated-orcid":false,"given":"Christopher D.","family":"Rock","sequence":"additional","affiliation":[{"name":"Department of Biological Sciences Texas Tech University  Lubbock Texas USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,9,2]]},"reference":[{"key":"e_1_2_10_2_1","doi-asserted-by":"publisher","DOI":"10.1534\/g3.116.029660"},{"key":"e_1_2_10_3_1","doi-asserted-by":"publisher","DOI":"10.1007\/BF00365615"},{"key":"e_1_2_10_4_1","doi-asserted-by":"publisher","DOI":"10.1105\/tpc.110.074781"},{"key":"e_1_2_10_5_1","doi-asserted-by":"publisher","DOI":"10.4161\/psb.6.8.15793"},{"key":"e_1_2_10_6_1","doi-asserted-by":"publisher","DOI":"10.1128\/MCB.00473\u201013"},{"key":"e_1_2_10_7_1","doi-asserted-by":"publisher","DOI":"10.1093\/genetics\/23.3.307"},{"key":"e_1_2_10_8_1","doi-asserted-by":"publisher","DOI":"10.1093\/g3journal\/jkae281"},{"key":"e_1_2_10_9_1","doi-asserted-by":"publisher","DOI":"10.1111\/tpj.14415"},{"key":"e_1_2_10_10_1","doi-asserted-by":"publisher","DOI":"10.1093\/aob\/mcu040"},{"key":"e_1_2_10_11_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.2517\u20106161.1995.tb02031.x"},{"key":"e_1_2_10_12_1","doi-asserted-by":"publisher","DOI":"10.1111\/nan.12688"},{"key":"e_1_2_10_13_1","doi-asserted-by":"publisher","DOI":"10.1093\/mp\/ssp009"},{"key":"e_1_2_10_14_1","doi-asserted-by":"publisher","DOI":"10.1002\/tpg2.20114"},{"key":"e_1_2_10_15_1","doi-asserted-by":"publisher","DOI":"10.1038\/nbt.3519"},{"key":"e_1_2_10_16_1","doi-asserted-by":"publisher","DOI":"10.1111\/tpj.12243"},{"key":"e_1_2_10_17_1","doi-asserted-by":"publisher","DOI":"10.1093\/gigascience\/gix134"},{"key":"e_1_2_10_18_1","doi-asserted-by":"publisher","DOI":"10.1046\/j.1365-313X.1999.00386.x"},{"key":"e_1_2_10_19_1","doi-asserted-by":"publisher","DOI":"10.1523\/ENEURO.0045\u201020.2020"},{"key":"e_1_2_10_20_1","doi-asserted-by":"publisher","DOI":"10.1093\/mp\/ssn060"},{"key":"e_1_2_10_21_1","doi-asserted-by":"publisher","DOI":"10.1046\/j.1365\u2010313x.1998.00301.x"},{"key":"e_1_2_10_22_1","doi-asserted-by":"publisher","DOI":"10.1038\/ng.2313"},{"key":"e_1_2_10_23_1","doi-asserted-by":"publisher","DOI":"10.4161\/fly.19695"},{"key":"e_1_2_10_24_1","doi-asserted-by":"publisher","DOI":"10.1104\/pp.104.044602"},{"key":"e_1_2_10_25_1","doi-asserted-by":"publisher","DOI":"10.1105\/tpc.108.065334"},{"key":"e_1_2_10_26_1","doi-asserted-by":"publisher","DOI":"10.3791\/62031"},{"key":"e_1_2_10_27_1","doi-asserted-by":"publisher","DOI":"10.1186\/s12864\u2010018\u20104490\u20107"},{"key":"e_1_2_10_28_1","unstructured":"Dhingra A.2021. \u201cNext Generation Sequencing\u2010Based Cloning of Zea mays Mutants and MIRNA\u2010Guided Regulation of Heat Stress Response in Peanut.\u201d Phd. 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