{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,22]],"date-time":"2026-10-22T09:56:52Z","timestamp":1792663012394,"version":"4.5.0"},"reference-count":73,"publisher":"Oxford University Press (OUP)","issue":"14","license":[{"start":{"date-parts":[[2026,3,10]],"date-time":"2026-03-10T00:00:00Z","timestamp":1773100800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,7,29]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>The growing prevalence of gluten-related disorders in humans has driven the development of wheat varieties with reduced immunogenic gluten. This study aimed to integrate RNA interference (RNAi) and CRISPR genome editing within a doubled haploid (DH) platform to overcome challenges of gene redundancy and polyploidy in wheat gliadins. We generated DH lines from crosses between RNAi and CRISPR lines and elite wheat cultivars, enabling stable fixation of multiple genetic modifications in a single generation. Deep sequencing analysis of \u03b1-gliadin amplicons was conducted using a custom bioinformatics pipeline optimized for complex, repetitive gene families. Gluten protein profiles were evaluated using RP-HPLC and R5 monoclonal antibody. Several DH lines presented &amp;gt;70% reduction in immunogenic epitopes in \u03b1-gliadins, with lines outperforming both parents. Editing frequency was influenced by sgRNA efficiency and parental background. Silencing and editing combined led to nearly depleted gliadins in some lines, often with compensatory increases in other storage proteins linked to bread-making quality, such as high-molecular-weight glutenin subunits. Kernel and specific weight traits were largely maintained. This work demonstrates that combining RNAi and CRISPR in a DH platform enables efficient, heritable reduction of immunogenic gluten, providing a viable strategy for breeding wheat lines safer for individuals with gluten-related disorders.<\/jats:p>","DOI":"10.1093\/jxb\/erag131","type":"journal-article","created":{"date-parts":[[2026,3,9]],"date-time":"2026-03-09T12:21:59Z","timestamp":1773058919000},"page":"4417-4434","source":"Crossref","is-referenced-by-count":6,"title":["Engineering ultra-low-gliadin wheat for celiac disease using an integrated RNAi, CRISPR, and doubled haploid strategy"],"prefix":"10.1093","volume":"77","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-0040-1053","authenticated-orcid":false,"given":"Miriam","family":"Mar\u00edn-Sanz","sequence":"first","affiliation":[{"name":"Functional Genomics Laboratory, Plant Breeding Department, Institute for Sustainable Agriculture, Spanish National Research Council (IAS-CSIC) , Av. Men\u00e9ndez Pidal s\/n, C\u00f3rdoba 14004 ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jos\u00e9 A","family":"Berlanga-Torres","sequence":"additional","affiliation":[{"name":"Functional Genomics Laboratory, Plant Breeding Department, Institute for Sustainable Agriculture, Spanish National Research Council (IAS-CSIC) , Av. Men\u00e9ndez Pidal s\/n, C\u00f3rdoba 14004 ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0032-0058","authenticated-orcid":false,"given":"Mar\u00eda H","family":"Guzm\u00e1n-L\u00f3pez","sequence":"additional","affiliation":[{"name":"Functional Genomics Laboratory, Plant Breeding Department, Institute for Sustainable Agriculture, Spanish National Research Council (IAS-CSIC) , Av. Men\u00e9ndez Pidal s\/n, C\u00f3rdoba 14004 ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Susana","family":"S\u00e1nchez-Le\u00f3n","sequence":"additional","affiliation":[{"name":"Functional Genomics Laboratory, Plant Breeding Department, Institute for Sustainable Agriculture, Spanish National Research Council (IAS-CSIC) , Av. Men\u00e9ndez Pidal s\/n, C\u00f3rdoba 14004 ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mar\u00eda P","family":"Vall\u00e9s","sequence":"additional","affiliation":[{"name":"Department of Genetics and Plant Breeding, Aula Dei Experimental Station, Spanish National Research Council (EEAD-CSIC) , Avda. Monta\u00f1ana 1005, Zaragoza 50059 ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ana M","family":"Castillo","sequence":"additional","affiliation":[{"name":"Department of Genetics and Plant Breeding, Aula Dei Experimental Station, Spanish National Research Council (EEAD-CSIC) , Avda. Monta\u00f1ana 1005, Zaragoza 50059 ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7652-229X","authenticated-orcid":false,"given":"Francisco","family":"Barro","sequence":"additional","affiliation":[{"name":"Functional Genomics Laboratory, Plant Breeding Department, Institute for Sustainable Agriculture, Spanish National Research Council (IAS-CSIC) , Av. Men\u00e9ndez Pidal s\/n, C\u00f3rdoba 14004 ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,3,10]]},"reference":[{"key":"2026072903365731000_erag131-B1","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1007\/s001220050531","article-title":"The \u03b1-gliadin gene family. I. Characterization of ten new wheat \u03b1-gliadin genomic clones, evidence for limited sequence conservation of flanking DNA, and Southern analysis of the gene family","volume":"95","author":"Anderson","year":"1997","journal-title":"Theoretical and Applied Genetics"},{"key":"2026072903365731000_erag131-B2","doi-asserted-by":"crossref","first-page":"603","DOI":"10.1084\/jem.191.4.603","article-title":"The intestinal T cell response to \u03b1-gliadin in adult celiac disease is focused on a single deamidated glutamine targeted by tissue transglutaminase","volume":"191","author":"Arentz-Hansen","year":"2000","journal-title":"Journal of Experimental Medicine"},{"key":"2026072903365731000_erag131-B3","doi-asserted-by":"crossref","first-page":"986","DOI":"10.1111\/pbi.12455","article-title":"Targeting of prolamins by RNAi in bread wheat: effectiveness of seven silencing-fragment combinations for obtaining lines devoid of coeliac disease epitopes from highly immunogenic gliadins","volume":"14","author":"Barro","year":"2016","journal-title":"Plant Biotechnology Journal"},{"key":"2026072903365731000_erag131-B4","first-page":"23","article-title":"Protein composition and techno-functional properties of transgenic wheat with reduced \u03b1-gliadin content obtained by RNA interference","volume":"85","author":"Becker","year":"2012","journal-title":"Journal of Applied Botany and Food Quality"},{"key":"2026072903365731000_erag131-B5","doi-asserted-by":"crossref","first-page":"6502","DOI":"10.1038\/s41598-018-24690-8","article-title":"Targeted mutagenesis in wheat microspores using CRISPR\/Cas9","volume":"8","author":"Bhowmik","year":"2018","journal-title":"Scientific Reports"},{"key":"2026072903365731000_erag131-B6","doi-asserted-by":"crossref","first-page":"2376","DOI":"10.1111\/pbi.13415","article-title":"Site-directed mutagenesis in bread and durum wheat via pollination by cas9\/guide RNA-transgenic maize used as haploidy inducer","volume":"18","author":"Budhagatapalli","year":"2020","journal-title":"Plant Biotechnology Journal"},{"key":"2026072903365731000_erag131-B7","volume-title":"BBMap: a fast, accurate, splice-aware aligner","author":"Bushnell","year":"2014"},{"key":"2026072903365731000_erag131-B8","doi-asserted-by":"crossref","first-page":"142","DOI":"10.1186\/s12916-019-1380-z","article-title":"Celiac disease: a comprehensive current review","volume":"17","author":"Caio","year":"2019","journal-title":"BMC Medicine"},{"key":"2026072903365731000_erag131-B9","doi-asserted-by":"crossref","first-page":"402","DOI":"10.3389\/fpls.2015.00402","article-title":"Effect of ovary induction on bread wheat anther culture: ovary genotype and developmental stage, and candidate gene association","volume":"6","author":"Castillo","year":"2015","journal-title":"Frontiers in Plant Science"},{"key":"2026072903365731000_erag131-B10","doi-asserted-by":"crossref","first-page":"227","DOI":"10.1007\/978-1-0716-1315-3_11","volume-title":"Doubled haploid technology. Volume 1: general topics, Alliaceae, cereals","author":"Castillo","year":"2021"},{"key":"2026072903365731000_erag131-B11","doi-asserted-by":"crossref","first-page":"S7","DOI":"10.1097\/01.mpg.0000450393.23156.59","article-title":"The new epidemiology of celiac disease","volume":"59","author":"Catassi","year":"2014","journal-title":"Journal of Pediatric Gastroenterology and Nutrition"},{"key":"2026072903365731000_erag131-B12","doi-asserted-by":"crossref","first-page":"224","DOI":"10.1038\/s41587-019-0032-3","article-title":"CRISPResso2 provides accurate and rapid genome editing sequence analysis","volume":"37","author":"Clement","year":"2019","journal-title":"Nature Biotechnology"},{"key":"2026072903365731000_erag131-B13","doi-asserted-by":"crossref","first-page":"1258096","DOI":"10.1126\/science.1258096","article-title":"The new frontier of genome engineering with CRISPR-Cas9","volume":"346","author":"Doudna","year":"2014","journal-title":"Science"},{"key":"2026072903365731000_erag131-B14","volume-title":"Usearch","author":"Edgar","year":"2010"},{"key":"2026072903365731000_erag131-B15","doi-asserted-by":"crossref","first-page":"108007","DOI":"10.1016\/j.biotechadv.2022.108007","article-title":"Developments and prospects for doubled haploid wheat","volume":"60","author":"Eliby","year":"2022","journal-title":"Biotechnology Advances"},{"key":"2026072903365731000_erag131-B16","doi-asserted-by":"crossref","first-page":"806","DOI":"10.1038\/35888","article-title":"Potent and specific genetic interference by double-stranded RNA in Caenorhabditis elegans","volume":"391","author":"Fire","year":"1998","journal-title":"Nature"},{"key":"2026072903365731000_erag131-B17","doi-asserted-by":"crossref","first-page":"102","DOI":"10.1016\/j.jprot.2017.06.010","article-title":"Comparative proteomic analysis of two transgenic low-gliadin wheat lines and non-transgenic wheat control","volume":"165","author":"Garc\u00eda-Molina","year":"2017","journal-title":"Journal of Proteomics"},{"key":"2026072903365731000_erag131-B18","doi-asserted-by":"crossref","first-page":"161","DOI":"10.1016\/j.jcs.2012.02.009","article-title":"Significant down-regulation of \u03b3-gliadins has minor effect on gluten and starch properties of bread wheat","volume":"56","author":"Gil-Humanes","year":"2012","journal-title":"Journal of Cereal Science"},{"key":"2026072903365731000_erag131-B19","doi-asserted-by":"crossref","first-page":"17023","DOI":"10.1073\/pnas.1007773107","article-title":"Effective shutdown in the expression of celiac disease-related wheat gliadin T-cell epitopes by RNA interference","volume":"107","author":"Gil-Humanes","year":"2010","journal-title":"Proceedings of the National Academy of Sciences, USA"},{"key":"2026072903365731000_erag131-B20","doi-asserted-by":"crossref","first-page":"2968","DOI":"10.1093\/bioinformatics\/btu427","article-title":"Genome editing assessment using CRISPR Genome Analyzer (CRISPR-GA)","volume":"30","author":"G\u00fcell","year":"2014","journal-title":"Bioinformatics"},{"key":"2026072903365731000_erag131-B21","doi-asserted-by":"crossref","first-page":"13076","DOI":"10.3390\/ijms222313076","article-title":"A bioinformatic workflow for InDel analysis in the wheat multi-copy \u03b1-gliadin gene family engineered with CRISPR\/Cas9","volume":"22","author":"Guzm\u00e1n-L\u00f3pez","year":"2021","journal-title":"International Journal of Molecular Sciences"},{"key":"2026072903365731000_erag131-B22","doi-asserted-by":"crossref","first-page":"4548","DOI":"10.3390\/nu13124548","article-title":"Oral consumption of bread from an RNAi wheat line with strongly silenced gliadins elicits no immunogenic response in a pilot study with celiac disease patients","volume":"13","author":"Guzm\u00e1n-L\u00f3pez","year":"2021","journal-title":"Nutrients"},{"key":"2026072903365731000_erag131-B23","doi-asserted-by":"crossref","first-page":"520","DOI":"10.1007\/BF01142316","article-title":"Improvement of isolated microspore culture of wheat (Triticum aestivum L.) through ovary co-culture","volume":"16","author":"Hul","year":"1997","journal-title":"Plant Cell Reports"},{"key":"2026072903365731000_erag131-B24","author":"Hunter","year":"1988"},{"key":"2026072903365731000_erag131-B25","doi-asserted-by":"crossref","first-page":"673","DOI":"10.3389\/fpls.2018.00673","article-title":"Gene duplication and evolution dynamics in the homeologous regions harboring multiple prolamin and resistance gene families in hexaploid wheat","volume":"9","author":"Huo","year":"2018","journal-title":"Frontiers in Plant Science"},{"key":"2026072903365731000_erag131-B26","doi-asserted-by":"crossref","first-page":"5181","DOI":"10.1038\/s41598-018-23570-5","article-title":"Dynamic evolution of \u03b1-gliadin prolamin gene family in homeologous genomes of hexaploid wheat","volume":"8","author":"Huo","year":"2018","journal-title":"Scientific Reports"},{"key":"2026072903365731000_erag131-B27","doi-asserted-by":"crossref","first-page":"1521","DOI":"10.1111\/nph.19021","article-title":"Combining multiplex gene editing and doubled haploid technology in maize","volume":"239","author":"Impens","year":"2023","journal-title":"New Phytologist"},{"key":"2026072903365731000_erag131-B28","volume-title":"Applied and fundamental aspects of plant cell, tissue, and organ culture","author":"Jensen","year":"1977"},{"key":"2026072903365731000_erag131-B29","doi-asserted-by":"crossref","first-page":"102903","DOI":"10.1016\/j.jcs.2019.102903","article-title":"Optimisation of droplet digital PCR for determining copy number variation of \u03b1-gliadin genes in mutant and gene-edited polyploid bread wheat","volume":"92","author":"Jouanin","year":"2020","journal-title":"Journal of Cereal Science"},{"key":"2026072903365731000_erag131-B30","doi-asserted-by":"crossref","first-page":"554","DOI":"10.1094\/MPMI-01-23-0004-SC","article-title":"Targeting disease susceptibility genes in wheat through wide hybridization with maize expressing Cas9 and guide RNA","volume":"36","author":"Karmacharya","year":"2023","journal-title":"Molecular Plant-Microbe Interactions"},{"key":"2026072903365731000_erag131-B31","doi-asserted-by":"crossref","first-page":"287","DOI":"10.1038\/s41587-019-0038-x","article-title":"One-step genome editing of elite crop germplasm during haploid induction","volume":"37","author":"Kelliher","year":"2019","journal-title":"Nature Biotechnology"},{"key":"2026072903365731000_erag131-B32","doi-asserted-by":"crossref","first-page":"507","DOI":"10.14309\/ajg.0000000000000523","article-title":"Incidence of celiac disease is increasing over time: a systematic review and meta-analysis","volume":"115","author":"King","year":"2020","journal-title":"American Journal of Gastroenterology"},{"key":"2026072903365731000_erag131-B33","doi-asserted-by":"crossref","first-page":"lqaa012","DOI":"10.1093\/nargab\/lqaa012","article-title":"CRISPRpic: fast and precise analysis for CRISPR-induced mutations via prefixed index counting","volume":"2","author":"Lee","year":"2020","journal-title":"NAR Genomics and Bioinformatics"},{"key":"2026072903365731000_erag131-B34","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1016\/j.jcs.2009.09.001","article-title":"Stacking HMW-GS transgenes in bread wheat: combining subunit 1Dy10 gives improved mixing properties and dough functionality","volume":"51","author":"Le\u00f3n","year":"2010","journal-title":"Journal of Cereal Science"},{"key":"2026072903365731000_erag131-B35","doi-asserted-by":"crossref","first-page":"148","DOI":"10.1016\/j.jcs.2008.08.002","article-title":"Mixing properties and dough functionality of transgenic lines of a commercial wheat cultivar expressing the 1Ax1, 1Dx5 and 1Dy10 HMW glutenin subunit genes","volume":"49","author":"Le\u00f3n","year":"2009","journal-title":"Journal of Cereal Science"},{"key":"2026072903365731000_erag131-B36","doi-asserted-by":"crossref","first-page":"344","DOI":"10.1016\/j.jcs.2010.02.002","article-title":"Pasting properties of transgenic lines of a commercial bread wheat expressing combinations of HMW glutenin subunit genes","volume":"51","author":"Le\u00f3n","year":"2010","journal-title":"Journal of Cereal Science"},{"key":"2026072903365731000_erag131-B37","doi-asserted-by":"publisher","author":"Li","year":"2013","DOI":"10.48550\/arXiv.1303.3997"},{"key":"2026072903365731000_erag131-B38","doi-asserted-by":"crossref","first-page":"21451","DOI":"10.1038\/srep21451","article-title":"Selection of highly efficient sgRNAs for CRISPR\/Cas9-based plant genome editing","volume":"6","author":"Liang","year":"2016","journal-title":"Scientific Reports"},{"key":"2026072903365731000_erag131-B39","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1186\/s13059-019-1762-8","article-title":"Modulating chromatin accessibility by transactivation and targeting proximal dsgRNAs enhances Cas9 editing efficiency in vivo","volume":"20","author":"Liu","year":"2019","journal-title":"Genome Biology"},{"key":"2026072903365731000_erag131-B40","doi-asserted-by":"crossref","first-page":"1210","DOI":"10.1136\/gutjnl-2013-306578","article-title":"Diagnosis and management of adult coeliac disease: guidelines from the British Society of Gastroenterology","volume":"63","author":"Ludvigsson","year":"2014","journal-title":"Gut"},{"key":"2026072903365731000_erag131-B41","doi-asserted-by":"crossref","first-page":"1171882","DOI":"10.3389\/fpls.2023.1171882","article-title":"Unraveling the celiac disease-related immunogenic complexes in a set of wheat and tritordeum genotypes: implications for low-gluten precision breeding in cereal crops","volume":"14","author":"Mar\u00edn-Sanz","year":"2023","journal-title":"Frontiers in Plant Science"},{"key":"2026072903365731000_erag131-B42","doi-asserted-by":"crossref","first-page":"314","DOI":"10.3389\/fpls.2020.00314","article-title":"Prolamin content and grain weight in RNAi silenced wheat lines under different conditions of temperature and nitrogen availability","volume":"11","author":"Mar\u00edn-Sanz","year":"2020","journal-title":"Frontiers in Plant Science"},{"key":"2026072903365731000_erag131-B43","doi-asserted-by":"crossref","first-page":"194","DOI":"10.1016\/j.cj.2021.04.009","article-title":"New transcriptomic insights in two RNAi wheat lines with the gliadins strongly down-regulated by two endosperm specific promoters","volume":"10","author":"Mar\u00edn-Sanz","year":"2022","journal-title":"The Crop Journal"},{"key":"2026072903365731000_erag131-B44","doi-asserted-by":"crossref","first-page":"3798","DOI":"10.1111\/pbi.70200","article-title":"Cas9-and Cas12a-mediated excision and replacement of the celiac disease-related \u03b1-gliadin immunogenic complex in hexaploid wheat","volume":"23","author":"Mar\u00edn-Sanz","year":"2025","journal-title":"Plant Biotechnology Journal"},{"key":"2026072903365731000_erag131-B45","doi-asserted-by":"crossref","first-page":"33","DOI":"10.1016\/j.talanta.2011.12.073","article-title":"Comprehensive analysis of gluten in processed foods using a new extraction method and a competitive ELISA based on the R5 antibody","volume":"91","author":"Mena","year":"2012","journal-title":"Talanta"},{"key":"2026072903365731000_erag131-B46","doi-asserted-by":"crossref","first-page":"4321","DOI":"10.1093\/nar\/8.19.4321","article-title":"Rapid isolation of high molecular weight plant DNA","volume":"8","author":"Murray","year":"1980","journal-title":"Nucleic Acids Research"},{"key":"2026072903365731000_erag131-B47","doi-asserted-by":"crossref","first-page":"587","DOI":"10.3109\/07853890.2010.505931","article-title":"The prevalence of celiac disease in Europe: results of a centralized, international mass screening project","volume":"42","author":"Mustalahti","year":"2010","journal-title":"Annals of Medicine"},{"key":"2026072903365731000_erag131-B48","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1016\/j.jcs.2018.12.017","article-title":"Contribution of \u03b1-gliadin alleles to the extensibility of flour dough in Japanese wheat cultivars","volume":"86","author":"Noma","year":"2019","journal-title":"Journal of Cereal Science"},{"key":"2026072903365731000_erag131-B49","doi-asserted-by":"crossref","first-page":"794","DOI":"10.1111\/tpj.12851","article-title":"Diversification of the celiac disease \u03b1-gliadin complex in wheat: a 33-mer peptide with six overlapping epitopes, evolved following polyploidization","volume":"82","author":"Ozuna C","year":"2015","journal-title":"Plant Journal"},{"key":"2026072903365731000_erag131-B50","doi-asserted-by":"crossref","first-page":"141","DOI":"10.1146\/annurev.pp.38.060187.001041","article-title":"Genetics of wheat storage proteins and the effect of allelic variation on bread-making quality","volume":"38","author":"Payne","year":"1987","journal-title":"Annual Review of Plant Physiology"},{"key":"2026072903365731000_erag131-B51","doi-asserted-by":"crossref","first-page":"136","DOI":"10.1186\/1471-2229-13-136","article-title":"Integration of promoters, inverted repeat sequences and proteomic data into a model for high silencing efficiency of coeliac disease related gliadins in bread wheat","volume":"13","author":"Pist\u00f3n","year":"2013","journal-title":"BMC Plant Biology"},{"key":"2026072903365731000_erag131-B52","doi-asserted-by":"crossref","first-page":"655","DOI":"10.1007\/s11032-009-9263-1","article-title":"Analysis of the activity of a \u03b3-gliadin promoter in transgenic wheat and characterization of gliadin synthesis in wheat by MALDI-TOF during grain development","volume":"23","author":"Pist\u00f3n","year":"2009","journal-title":"Molecular Breeding"},{"key":"2026072903365731000_erag131-B53","doi-asserted-by":"crossref","first-page":"77","DOI":"10.1007\/s00122-007-0542-0","article-title":"D hordeins of Hordeum chilense: a novel source of variation for improvement of wheat","volume":"115","author":"Pist\u00f3n","year":"2007","journal-title":"Theoretical and Applied Genetics"},{"key":"2026072903365731000_erag131-B54","volume-title":"R: A Language and Environment for Statistical Computing","author":"R Core Development Team","year":"2020"},{"key":"2026072903365731000_erag131-B55","doi-asserted-by":"crossref","first-page":"94","DOI":"10.1007\/s00122-025-04882-3","article-title":"Deletion of wheat alpha-gliadins from chromosome 6D improves gluten strength and reduces immunodominant celiac disease epitopes","volume":"138","author":"Rottersman","year":"2025","journal-title":"Theoretical and Applied Genetics"},{"key":"2026072903365731000_erag131-B56","doi-asserted-by":"crossref","first-page":"14","DOI":"10.1104\/pp.108.129437","article-title":"The development of endosperm in grasses","volume":"149","author":"Sabelli","year":"2009","journal-title":"Plant Physiology"},{"key":"2026072903365731000_erag131-B57","doi-asserted-by":"crossref","first-page":"209","DOI":"10.1007\/BF00226253","article-title":"Characterization and organization of gene families at the Gli-1 loci of bread and durum wheats by restriction fragment analysis","volume":"83","author":"Sabelli","year":"1991","journal-title":"Theoretical and Applied Genetics"},{"key":"2026072903365731000_erag131-B58","doi-asserted-by":"crossref","first-page":"48","DOI":"10.1186\/1471-2164-10-48","article-title":"Tetraploid and hexaploid wheat varieties reveal large differences in expression of alpha-gliadins from homoeologous Gli-2 loci","volume":"10","author":"Salentijn","year":"2009","journal-title":"BMC Genomics"},{"key":"2026072903365731000_erag131-B59","doi-asserted-by":"crossref","first-page":"902","DOI":"10.1111\/pbi.12837","article-title":"Low-gluten, nontransgenic wheat engineered with CRISPR\/Cas9","volume":"16","author":"S\u00e1nchez-Le\u00f3n","year":"2018","journal-title":"Plant Biotechnology Journal"},{"key":"2026072903365731000_erag131-B60","doi-asserted-by":"crossref","first-page":"742","DOI":"10.3389\/fpls.2021.663653","article-title":"The \u03b1-gliadins in bread wheat: effect of nitrogen treatment on the expression of the major celiac disease immunogenic complex in two RNAi low-gliadin lines","volume":"12","author":"S\u00e1nchez-Le\u00f3n","year":"2021","journal-title":"Frontiers in Plant Science"},{"key":"2026072903365731000_erag131-B61","doi-asserted-by":"crossref","first-page":"2933","DOI":"10.3390\/nu11122933","article-title":"Stimulatory response of celiac disease peripheral blood mononuclear cells induced by RNAi wheat lines differing in grain protein composition","volume":"11","author":"S\u00e1nchez-Le\u00f3n","year":"2019","journal-title":"Nutrients"},{"key":"2026072903365731000_erag131-B62","doi-asserted-by":"crossref","first-page":"7079","DOI":"10.1093\/jxb\/erae376","article-title":"CRISPR\/Cas9-mediated multiplex gene editing of gamma and omega gliadins, paving the way for gliadin-free wheat","volume":"75","author":"S\u00e1nchez-Le\u00f3n","year":"2024","journal-title":"Journal of Experimental Botany"},{"key":"2026072903365731000_erag131-B63","doi-asserted-by":"crossref","first-page":"101","DOI":"10.3389\/fnut.2019.00101","article-title":"What is gluten\u2014why is it special?","volume":"6","author":"Shewry","year":"2019","journal-title":"Frontiers in Nutrition"},{"key":"2026072903365731000_erag131-B64","doi-asserted-by":"crossref","first-page":"1537","DOI":"10.1093\/jxb\/erp058","article-title":"Wheat","volume":"60","author":"Shewry","year":"2009","journal-title":"Journal of Experimental Botany"},{"key":"2026072903365731000_erag131-B65","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1016\/S0065-2660(03)01003-4","article-title":"Genetics of wheat gluten proteins","volume":"49","author":"Shewry","year":"2003","journal-title":"Advances in Genetics"},{"key":"2026072903365731000_erag131-B66","doi-asserted-by":"crossref","first-page":"85","DOI":"10.1007\/s00251-019-01141-w","article-title":"Update 2020: nomenclature and listing of celiac disease\u2013relevant gluten epitopes recognized by CD4+ T cells","volume":"72","author":"Sollid","year":"2020","journal-title":"Immunogenetics"},{"key":"2026072903365731000_erag131-B67","doi-asserted-by":"crossref","first-page":"805","DOI":"10.1007\/s00299-007-0500-y","article-title":"Enhanced induction of microspore embryogenesis after n-butanol treatment in wheat (Triticum aestivum L.) anther culture","volume":"27","author":"Soriano","year":"2008","journal-title":"Plant Cell Reports"},{"key":"2026072903365731000_erag131-B68","doi-asserted-by":"crossref","first-page":"1026","DOI":"10.1038\/nbt.3988","article-title":"MMseqs2 enables sensitive protein sequence searching for the analysis of massive data sets","volume":"35","author":"Steinegger","year":"2017","journal-title":"Nature Biotechnology"},{"key":"2026072903365731000_erag131-B69","doi-asserted-by":"crossref","first-page":"258","DOI":"10.1186\/s12876-020-01390-0","article-title":"An updated overview of spectrum of gluten-related disorders: clinical and diagnostic aspects","volume":"20","author":"Taraghikhah","year":"2020","journal-title":"BMC Gastroenterology"},{"key":"2026072903365731000_erag131-B70","doi-asserted-by":"crossref","first-page":"41ra51","DOI":"10.1126\/scitranslmed.3001012","article-title":"Comprehensive, quantitative mapping of T cell epitopes in gluten in celiac disease","volume":"2","author":"Tye-Din","year":"2010","journal-title":"Science Translational Medicine"},{"key":"2026072903365731000_erag131-B71","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1186\/1471-2164-7-1","article-title":"Alpha-gliadin genes from the A, B, and D genomes of wheat contain different sets of celiac disease epitopes","volume":"7","author":"van Herpen","year":"2006","journal-title":"BMC Genomics"},{"key":"2026072903365731000_erag131-B72","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1089\/crispr.2017.0010","article-title":"Transgenerational CRISPR-cas9 activity facilitates multiplex gene editing in allopolyploid wheat","volume":"1","author":"Wang","year":"2018","journal-title":"The CRISPR Journal"},{"key":"2026072903365731000_erag131-B73","doi-asserted-by":"crossref","first-page":"892","DOI":"10.1111\/pbi.14231","article-title":"CRISPR-based editing of the \u03c9-and \u03b3-gliadin gene clusters reduces wheat immunoreactivity without affecting grain protein quality","volume":"22","author":"Yu","year":"2024","journal-title":"Plant Biotechnology Journal"}],"container-title":["Journal of Experimental Botany"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/jxb\/advance-article-pdf\/doi\/10.1093\/jxb\/erag131\/67286823\/erag131.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/jxb\/article-pdf\/77\/14\/4417\/67286823\/erag131.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/jxb\/article-pdf\/77\/14\/4417\/67286823\/erag131.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,29]],"date-time":"2026-07-29T07:37:33Z","timestamp":1785310653000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/jxb\/article\/77\/14\/4417\/8512773"}},"subtitle":[],"editor":[{"given":"Diane","family":"Beckles","sequence":"additional","affiliation":[{"name":"University of California, Davis, USA"}],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2026,3,10]]},"references-count":73,"journal-issue":{"issue":"14","published-online":{"date-parts":[[2026,3,10]]},"published-print":{"date-parts":[[2026,7,29]]}},"URL":"https:\/\/doi.org\/10.1093\/jxb\/erag131","relation":{},"ISSN":["0022-0957","1460-2431"],"issn-type":[{"value":"0022-0957","type":"print"},{"value":"1460-2431","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2026,7,29]]},"published":{"date-parts":[[2026,3,10]]}}}