{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,30]],"date-time":"2026-09-30T19:06:42Z","timestamp":1790795202198,"version":"4.1.0"},"reference-count":67,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2017,3,16]],"date-time":"2017-03-16T00:00:00Z","timestamp":1489622400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2017,3,16]],"date-time":"2017-03-16T00:00:00Z","timestamp":1489622400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"abstract":"<jats:title>Abstract<\/jats:title><jats:p>Gliadins, specified by six compound chromosomal loci (<jats:italic>Gli-A1\/B1\/D1<\/jats:italic> and <jats:italic>Gli-A2\/B2\/D2<\/jats:italic>) in hexaploid bread wheat, are the dominant carriers of celiac disease (CD) epitopes. Because of their complexity, genome-wide characterization of gliadins is a strong challenge. Here, we approached this challenge by combining transcriptomic, proteomic and bioinformatic investigations. Through third-generation RNA sequencing, full-length transcripts were identified for 52 gliadin genes in the bread wheat cultivar Xiaoyan 81. Of them, 42 were active and predicted to encode 25 \u03b1-, 11 \u03b3-, one \u03b4- and five \u03c9-gliadins. Comparative proteomic analysis between Xiaoyan 81 and six newly-developed mutants each lacking one <jats:italic>Gli<\/jats:italic> locus indicated the accumulation of 38 gliadins in the mature grains. A novel group of \u03b1-gliadins (the CSTT group) was recognized to contain very few or no CD epitopes. The \u03b4-gliadins identified here or previously did not carry CD epitopes. Finally, the mutant lacking <jats:italic>Gli-D2<\/jats:italic> showed significant reductions in the most celiac-toxic \u03b1-gliadins and derivative CD epitopes. The insights and resources generated here should aid further studies on gliadin functions in CD and the breeding of healthier wheat.<\/jats:p>","DOI":"10.1038\/srep44609","type":"journal-article","created":{"date-parts":[[2017,3,16]],"date-time":"2017-03-16T10:18:42Z","timestamp":1489659522000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":58,"title":["Genome-wide analysis of complex wheat gliadins, the dominant carriers of celiac disease epitopes"],"prefix":"10.1038","volume":"7","author":[{"given":"Da-Wei","family":"Wang","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Da","family":"Li","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Junjun","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yue","family":"Zhao","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhaojun","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Guidong","family":"Yue","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xin","family":"Liu","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Huanju","family":"Qin","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kunpu","family":"Zhang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lingli","family":"Dong","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Daowen","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2017,3,16]]},"reference":[{"key":"BFsrep44609_CR1","unstructured":"Food and Agriculture Organization of the United Nations Statistics Division. FAOSTAT, http:\/\/faostat3.fao.org\/download\/Q\/QC\/E (2016)."},{"key":"BFsrep44609_CR2","unstructured":"International Maize and Wheat Improvement Center (CIMMYT). Wheat Research, http:\/\/www.cimmyt.org\/global-wheat-research\/ (2016)."},{"key":"BFsrep44609_CR3","doi-asserted-by":"crossref","first-page":"238","DOI":"10.1097\/ACI.0b013e32832aa5bc","volume":"9","author":"N Inomata","year":"2009","unstructured":"Inomata, N. Wheat allergy. Curr. Opin. Allergy Clin. Immunol. 9, 238\u2013243 (2009).","journal-title":"Curr. Opin. Allergy Clin. Immunol."},{"key":"BFsrep44609_CR4","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1186\/1741-7015-10-13","volume":"10","author":"A Sapone","year":"2012","unstructured":"Sapone, A. et al. Spectrum of gluten-related disorders: consensus on new nomenclature and classification. BMC Med. 10, 13 (2012).","journal-title":"BMC Med."},{"key":"BFsrep44609_CR5","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1016\/j.jcs.2015.06.005","volume":"67","author":"PR Shewry","year":"2016","unstructured":"Shewry, P. R. & Tatham, A. S. Improving wheat to remove celiac epitopes but retain functionality. J. Cereal Sci. 67, 12\u201321 (2016).","journal-title":"J. Cereal Sci."},{"key":"BFsrep44609_CR6","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1016\/S0065-2660(03)01003-4","volume":"49","author":"PR Shewry","year":"2003","unstructured":"Shewry, P. R., Halford, N. G. & Lafiandra, D. Genetics of wheat gluten proteins. Adv. Genet. 49, 111\u2013184 (2003).","journal-title":"Adv. Genet"},{"key":"BFsrep44609_CR7","first-page":"796","volume":"40","author":"PF Qi","year":"2006","unstructured":"Qi, P. F., Wei, Y. M., Yue, Y. W., Yan, Z. H. & Zheng, Y. L. Biochemical and molecular characterization of gliadins. Mol Biol (Mosk). 40, 796\u2013807 (2006).","journal-title":"Mol Biol (Mosk)"},{"key":"BFsrep44609_CR8","doi-asserted-by":"crossref","unstructured":"Juhasz, A., Bekes, F. & Wrigley, C. W. Wheat proteins. In Applied Food Protein Chemistry (ed Ustunol, Z. ) 219\u2013303 (John Wiley & Sons, Ltd., 2015).","DOI":"10.1002\/9781118860588.ch11"},{"key":"BFsrep44609_CR9","doi-asserted-by":"crossref","first-page":"1537","DOI":"10.1093\/jxb\/erp058","volume":"60","author":"PR Shewry","year":"2009","unstructured":"Shewry, P. R. Wheat. J. Exp. Bot. 60, 1537\u20131553 (2009).","journal-title":"J. Exp. Bot."},{"key":"BFsrep44609_CR10","doi-asserted-by":"crossref","first-page":"2","DOI":"10.1016\/j.jcs.2015.07.008","volume":"67","author":"KA Scherf","year":"2016","unstructured":"Scherf, K. A., Koehler, P. & Wieser, H. Gluten and wheat sensitivities - An overview. J. Cereal Sci. 67, 2\u201311 (2016).","journal-title":"J. Cereal Sci."},{"key":"BFsrep44609_CR11","doi-asserted-by":"crossref","first-page":"365","DOI":"10.1016\/j.bpg.2015.05.004","volume":"29","author":"E Lionetti","year":"2015","unstructured":"Lionetti, E., Gatti, S., Pulvirenti, A. & Catassi, C. Celiac disease from a global perspective. Best Pract. Res. Clin. Gastroenterol. 29, 365\u2013379 (2015).","journal-title":"Best Pract. Res. Clin. Gastroenterol"},{"key":"BFsrep44609_CR12","doi-asserted-by":"crossref","first-page":"1770","DOI":"10.1007\/s00125-014-3265-1","volume":"57","author":"JC Antvorskov","year":"2014","unstructured":"Antvorskov, J. C., Josefsen, K., Engkilde, K., Funda, D. P. & Buschard, K. Dietary gluten and the development of type 1 diabetes. Diabetologia 57, 1770\u20131780 (2014).","journal-title":"Diabetologia"},{"key":"BFsrep44609_CR13","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1186\/s40348-015-0029-z","volume":"3","author":"MV Barone","year":"2016","unstructured":"Barone, M. V. & Zimmer, K. P. Endocytosis and transcytosis of gliadin peptides. Mol. Cell Pediatr. 3, 8 (2016).","journal-title":"Mol. Cell Pediatr"},{"key":"BFsrep44609_CR14","doi-asserted-by":"crossref","first-page":"1250092","DOI":"10.1126\/science.1250092","volume":"345","author":"T Marcussen","year":"2014","unstructured":"Marcussen, T. et al. Ancient hybridizations among the ancestral genomes of bread wheat. Science 345, 1250092 (2014).","journal-title":"Science"},{"key":"BFsrep44609_CR15","doi-asserted-by":"crossref","first-page":"341","DOI":"10.1007\/s10142-012-0269-0","volume":"12","author":"K Kawaura","year":"2012","unstructured":"Kawaura, K. et al. Genome change in wheat observed through the structure and expression of \u03b1\/\u03b2-gliadin genes. Funct. Integr. Genomics 12, 341\u2013355 (2012).","journal-title":"Funct. Integr. Genomics"},{"key":"BFsrep44609_CR16","doi-asserted-by":"crossref","first-page":"495","DOI":"10.1111\/tpj.13214","volume":"87","author":"L Dong","year":"2016","unstructured":"Dong, L. et al. Rapid evolutionary dynamics in a 2.8-Mb chromosomal region containing multiple prolamin and resistance gene families in Aegilops tauschii . Plant J. 87, 495\u2013506 (2016).","journal-title":"Plant J."},{"key":"BFsrep44609_CR17","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1186\/1471-2229-10-7","volume":"10","author":"SB Altenbach","year":"2010","unstructured":"Altenbach, S. B., Vensel, W. H. & Dupont, F. M. Analysis of expressed sequence tags from a single wheat cultivar facilitates interpretation of tandem mass spectrometry data and discrimination of gamma gliadin proteins that may play different functional roles in flour. BMC Plant Biol. 10, 7 (2010).","journal-title":"BMC Plant Biol"},{"key":"BFsrep44609_CR18","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1186\/1477-5956-9-10","volume":"9","author":"FM Dupont","year":"2011","unstructured":"Dupont, F. M., Vensel, W. H., Tanaka, C. K., Hurkman, W. J. & Altenbach, S. B. Deciphering the complexities of the wheat flour proteome using quantitative two-dimensional electrophoresis, three proteases and tandem mass spectrometry. Proteome Sci. 9, 10 (2011).","journal-title":"Proteome Sci."},{"key":"BFsrep44609_CR19","doi-asserted-by":"crossref","first-page":"1250091","DOI":"10.1126\/science.1250091","volume":"345","author":"M Pfeifer","year":"2014","unstructured":"Pfeifer, M. et al. Genome interplay in the grain transcriptome of hexaploid bread wheat. Science 345, 1250091 (2014).","journal-title":"Science"},{"key":"BFsrep44609_CR20","unstructured":"International Wheat Genome Sequencing Consortium. A chromosome-based draft sequence of the hexaploid bread wheat (Triticum aestivum) genome. Science 345, 1251788 (2014)."},{"key":"BFsrep44609_CR21","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1146\/annurev.immunol.18.1.53","volume":"18","author":"LM Sollid","year":"2000","unstructured":"Sollid, L. M. Molecular basis of celiac disease. Annu. Rev. Immunol. 18, 53\u201381 (2000).","journal-title":"Annu. Rev. Immunol."},{"key":"BFsrep44609_CR22","doi-asserted-by":"crossref","first-page":"343","DOI":"10.1016\/j.smim.2015.11.001","volume":"27","author":"J Stamnaes","year":"2015","unstructured":"Stamnaes, J. & Sollid, L. M. Celiac disease: Autoimmunity in response to food antigen. Semin. Immunol. 27, 343\u2013352 (2015).","journal-title":"Semin. Immunol."},{"key":"BFsrep44609_CR23","doi-asserted-by":"crossref","first-page":"413","DOI":"10.1016\/j.bpg.2015.04.001","volume":"29","author":"MF du Pr\u00e9","year":"2015","unstructured":"du Pr\u00e9, M. F. & Sollid, L. M. T-cell and B-cell immunity in celiac disease. Best Pract. Res. Clin. Gastroenterol. 29, 413\u2013423 (2015).","journal-title":"Best Pract. Res. Clin. Gastroenterol"},{"key":"BFsrep44609_CR24","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1186\/1471-2164-7-1","volume":"7","author":"T Van Herpen","year":"2006","unstructured":"Van Herpen, T. et al. Alpha-gliadin genes from the A, B, and D genomes of wheat contain different sets of celiac disease epitopes. BMC Genomics 7, 1\u201313 (2006).","journal-title":"BMC Genomics"},{"key":"BFsrep44609_CR25","doi-asserted-by":"crossref","first-page":"1239","DOI":"10.1007\/s00122-010-1384-8","volume":"121","author":"Z Xie","year":"2010","unstructured":"Xie, Z. et al. Molecular characterization of the celiac disease epitope domains in \u03b1-gliadin genes in Aegilops tauschii and hexaploid wheats (Triticum aestivum L.). Theor. Appl. Genet. 121, 1239\u20131251 (2010).","journal-title":"Theor. Appl. Genet."},{"key":"BFsrep44609_CR26","doi-asserted-by":"crossref","first-page":"1371","DOI":"10.1007\/s00122-012-1917-4","volume":"125","author":"S Wang","year":"2012","unstructured":"Wang, S. et al. Molecular characterization and dynamic expression patterns of two types of \u03b3-gliadin genes from Aegilops and Triticum species. Theor. Appl. Genet. 125, 1371\u20131384 (2012).","journal-title":"Theor. Appl. Genet."},{"key":"BFsrep44609_CR27","doi-asserted-by":"crossref","first-page":"277","DOI":"10.1186\/1471-2164-13-277","volume":"13","author":"EM Salentijn","year":"2012","unstructured":"Salentijn, E. M. et al. Celiac disease T-cell epitopes from gamma-gliadins: immunoreactivity depends on the genome of origin, transcript frequency, and flanking protein variation. BMC Genomics 13, 277 (2012).","journal-title":"BMC Genomics"},{"key":"BFsrep44609_CR28","doi-asserted-by":"crossref","first-page":"161","DOI":"10.1093\/jxb\/ers318","volume":"64","author":"Y Wan","year":"2013","unstructured":"Wan, Y., Shewry, P. R. & Hawkesford, M. J. A novel family of \u03b3-gliadin genes are highly regulated by nitrogen supply in developing wheat grain. J. Exp. Bot. 64, 161\u2013168 (2013).","journal-title":"J. Exp. Bot."},{"key":"BFsrep44609_CR29","doi-asserted-by":"crossref","first-page":"e52139","DOI":"10.1371\/journal.pone.0052139","volume":"7","author":"OD Anderson","year":"2012","unstructured":"Anderson, O. D., Dong, L., Huo, N. & Gu, Y. Q. A new class of wheat gliadin genes and proteins. PLoS One 7, e52139 (2012).","journal-title":"PLoS One"},{"key":"BFsrep44609_CR30","doi-asserted-by":"crossref","first-page":"359","DOI":"10.1007\/s00122-013-2224-4","volume":"127","author":"Y Yang","year":"2014","unstructured":"Yang, Y. et al. Efficient isolation of ion beam-induced mutants for homoeologous loci in common wheat and comparison of the contributions of Glu-1 loci to gluten functionality. Theor. Appl. Genet. 127, 359\u2013372 (2014).","journal-title":"Theor. Appl. Genet."},{"key":"BFsrep44609_CR31","doi-asserted-by":"crossref","first-page":"405","DOI":"10.1186\/gb-2013-14-6-405","volume":"14","author":"RJ Roberts","year":"2013","unstructured":"Roberts, R. J., Carneiro, M. O. & Schatz, M. C. The advantages of SMRT sequencing. Genome Biol. 14, 405 (2013).","journal-title":"Genome Biol."},{"key":"BFsrep44609_CR32","doi-asserted-by":"crossref","first-page":"1009","DOI":"10.1038\/nbt.2705","volume":"31","author":"D Sharon","year":"2013","unstructured":"Sharon, D., Tilgner, H., Grubert, F. & Snyder, M. A single-molecule long-read survey of the human transcriptome. Nat. Biotechnol. 31, 1009\u20131014 (2013).","journal-title":"Nat. Biotechnol."},{"key":"BFsrep44609_CR33","doi-asserted-by":"crossref","first-page":"541","DOI":"10.1016\/j.gene.2013.10.009","volume":"533","author":"W Zhang","year":"2014","unstructured":"Zhang, W., Ciclitira, P. & Messing, J. PacBio sequencing of gene families \u2013 A case study with wheat gluten genes. Gene 533, 541\u2013546 (2014).","journal-title":"Gene"},{"key":"BFsrep44609_CR34","doi-asserted-by":"crossref","first-page":"951","DOI":"10.1111\/tpj.12865","volume":"82","author":"Z Xu","year":"2015","unstructured":"Xu, Z. et al. Full-length transcriptome sequences and splice variants obtained by a combination of sequencing platforms applied to different root tissues of Salvia miltiorrhiza and tanshinone biosynthesis. Plant J. 82, 951\u2013961 (2015).","journal-title":"Plant J."},{"key":"BFsrep44609_CR35","doi-asserted-by":"crossref","first-page":"184","DOI":"10.1186\/s13059-015-0729-7","volume":"16","author":"AE Minoche","year":"2015","unstructured":"Minoche, A. E. et al. Exploiting single-molecule transcript sequencing for eukaryotic gene prediction. Genome Biol. 16, 184 (2015).","journal-title":"Genome Biol."},{"key":"BFsrep44609_CR36","doi-asserted-by":"crossref","first-page":"693","DOI":"10.1038\/nbt.2280","volume":"30","author":"S Koren","year":"2012","unstructured":"Koren, S. et al. Hybrid error correction and de novo assembly of single-molecule sequencing reads. Nat. Biotechnol. 30, 693\u2013700 (2012).","journal-title":"Nat. Biotechnol."},{"key":"BFsrep44609_CR37","doi-asserted-by":"crossref","first-page":"3506","DOI":"10.1093\/bioinformatics\/btu538","volume":"30","author":"L Salmela","year":"2014","unstructured":"Salmela, L. & Rivals, E. LoRDEC: accurate and efficient long read error correction. Bioinformatics 30, 3506\u20133514 (2014).","journal-title":"Bioinformatics"},{"key":"BFsrep44609_CR38","doi-asserted-by":"crossref","first-page":"1039","DOI":"10.1186\/s12864-015-2257-y","volume":"16","author":"L Dong","year":"2015","unstructured":"Dong, L. et al. Single-molecule real-time transcript sequencing facilitates common wheat genome annotation and grain transcriptome research. BMC Genomics 16, 1039 (2015).","journal-title":"BMC Genomics"},{"key":"BFsrep44609_CR39","first-page":"59","volume":"95","author":"OD Anderson","year":"1997","unstructured":"Anderson, O. D. & Greene, F. C. The \u03b1-gliadin gene family. II. DNA and protein sequence variation, subfamily structure, and origins of pseudogenes. Theor. Appl. Genet. 95, 59\u201365 (1997).","journal-title":"DNA and protein sequence variation, subfamily structure, and origins of pseudogenes. Theor. Appl. Genet"},{"key":"BFsrep44609_CR40","doi-asserted-by":"crossref","first-page":"323","DOI":"10.1007\/s00122-001-0551-3","volume":"103","author":"OD Anderson","year":"2001","unstructured":"Anderson, O. D., Hsia, C. C. & Torres, V. The wheat \u03b3-gliadin genes: characterization of ten new sequences and further understanding of \u03b3-gliadin gene family structure. Theor. Appl. Genet. 103, 323\u2013330 (2001).","journal-title":"Theor. Appl. Genet."},{"key":"BFsrep44609_CR41","doi-asserted-by":"crossref","first-page":"261","DOI":"10.1007\/s10142-013-0321-8","volume":"13","author":"OD Anderson","year":"2013","unstructured":"Anderson, O. D., Huo, N. & Gu, Y. Q. The gene space in wheat: the complete \u03b3-gliadin gene family from the wheat cultivar Chinese Spring. Funct. Integr. Genomics 13, 261\u2013273 (2013).","journal-title":"Funct. Integr. Genomics"},{"key":"BFsrep44609_CR42","doi-asserted-by":"crossref","first-page":"397","DOI":"10.1007\/s10142-009-0122-2","volume":"9","author":"OD Anderson","year":"2009","unstructured":"Anderson, O. D., Gu, Y. Q., Kong, X., Lazo, G. R. & Wu, J. The wheat \u03c9-gliadin genes: structure and EST analysis. Funct. Integr. Genomics 9, 397\u2013410 (2009).","journal-title":"Funct. Integr. Genomics"},{"key":"BFsrep44609_CR43","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1016\/j.jcs.2011.10.013","volume":"55","author":"AS Tatham","year":"2012","unstructured":"Tatham, A. S. & Shewry, P. R. The S-poor prolamins of wheat, barley and rye: revisited. J. Cereal Sci. 55, 79\u201399 (2012).","journal-title":"J. Cereal Sci."},{"key":"BFsrep44609_CR44","doi-asserted-by":"crossref","first-page":"2275","DOI":"10.1126\/science.1074129","volume":"297","author":"L Shan","year":"2002","unstructured":"Shan, L. et al. Structural basis for gluten intolerance in celiac spruce. Science 297, 2275\u20132279 (2002).","journal-title":"Science"},{"key":"BFsrep44609_CR45","doi-asserted-by":"crossref","unstructured":"Tye-Din, J. A. et al. Comprehensive, quantitative mapping of T cell epitopes in gluten in celiac disease. Sci. Transl. Med. 2, 41ra51 (2010).","DOI":"10.1126\/scitranslmed.3001012"},{"key":"BFsrep44609_CR46","doi-asserted-by":"crossref","first-page":"e2294","DOI":"10.1371\/journal.pone.0002294","volume":"3","author":"B Mor\u00f3n","year":"2008","unstructured":"Mor\u00f3n, B. et al. Toward the assessment of food toxicity for celiac patients: characterization of monoclonal antibodies to a main immunogenic gluten peptide. PLoS One 3, e2294 (2008).","journal-title":"PLoS One"},{"key":"BFsrep44609_CR47","doi-asserted-by":"crossref","first-page":"372","DOI":"10.5740\/jaoacint.SGE_Halbmayr-Jech","volume":"95","author":"E Halbmayr-Jech","year":"2012","unstructured":"Halbmayr-Jech, E. et al. Characterization of G12 sandwich ELISA, a next-generation immunoassay for gluten toxicity. J AOAC Int. 95, 372\u2013376 (2012).","journal-title":"J AOAC Int"},{"key":"BFsrep44609_CR48","doi-asserted-by":"crossref","first-page":"1033","DOI":"10.1038\/nature09144","volume":"465","author":"L Poliseno","year":"2010","unstructured":"Poliseno, L. et al. A coding-independent function of gene and pseudogene mRNAs regulates tumor biology. Nature 465, 1033\u20131038 (2010).","journal-title":"Nature"},{"key":"BFsrep44609_CR49","doi-asserted-by":"crossref","first-page":"350","DOI":"10.1016\/j.biocel.2014.05.008","volume":"54","author":"JN Groen","year":"2014","unstructured":"Groen, J. N., Capraro, D. & Morris, K. V. The emerging role of pseudogene expressed non-coding RNAs in cellular functions. Int. J. Biochem. Cell Biol. 54, 350\u2013355 (2014).","journal-title":"Int. J. Biochem. Cell Biol."},{"key":"BFsrep44609_CR50","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1016\/j.jcs.2010.04.006","volume":"52","author":"SB Altenbach","year":"2010","unstructured":"Altenbach, S. B., Vensel, W. H. & Dupont, F. M. Integration of proteomic and transcriptomic data from a single wheat cultivar provides new tools for understanding the roles of individual alpha gliadin proteins in flour quality and celiac disease. J. Cereal Sci. 52, 143\u2013151 (2010).","journal-title":"J. Cereal Sci."},{"key":"BFsrep44609_CR51","doi-asserted-by":"crossref","first-page":"725","DOI":"10.1007\/s12041-014-0441-5","volume":"93","author":"GR Li","year":"2014","unstructured":"Li, G. R., Lang, T., Yang, E. N., Liu, C. & Yang, Z. J. Characterization and phylogenetic analysis of \u03b1-gliadin gene sequences reveals significant genomic divergence in Triticeae species. J. Genet. 93, 725\u2013731 (2014).","journal-title":"J. Genet."},{"key":"BFsrep44609_CR52","doi-asserted-by":"crossref","first-page":"1908","DOI":"10.1002\/pmic.200500236","volume":"6","author":"P Ferrante","year":"2006","unstructured":"Ferrante, P. et al. proteomic approach to verify in vivo expression of a novel \u03b3-gliadin containing an extra cysteine residue. Proteomics 6, 1908\u20131914 (2006).","journal-title":"Proteomics"},{"key":"BFsrep44609_CR53","doi-asserted-by":"crossref","first-page":"169","DOI":"10.1016\/j.jcs.2007.02.001","volume":"46","author":"SB Altenbach","year":"2007","unstructured":"Altenbach, S. B. & Kothari, K. M. Omega gliadin genes expressed in Triticum aestivum cv. Butte 86: Effects of post-anthesis fertilizer on transcript accumulation during grain development. J. Cereal Sci. 46, 169\u2013177 (2007).","journal-title":"J. Cereal Sci."},{"key":"BFsrep44609_CR54","doi-asserted-by":"crossref","first-page":"11708","DOI":"10.1038\/ncomms11708","volume":"7","author":"B Wang","year":"2016","unstructured":"Wang, B. et al. Unveiling the complexity of the maize transcriptome by single-molecule long-read sequencing. Nat. Commun. 7, 11708 (2016).","journal-title":"Nat. Commun."},{"key":"BFsrep44609_CR55","doi-asserted-by":"crossref","first-page":"11706","DOI":"10.1038\/ncomms11706","volume":"7","author":"SE Abdel-Ghany","year":"2016","unstructured":"Abdel-Ghany, S. E. et al. A survey of the sorghum transcriptome using single-molecule long reads. Nat. Commun. 7, 11706 (2016).","journal-title":"Nat. Commun."},{"key":"BFsrep44609_CR56","doi-asserted-by":"crossref","first-page":"17023","DOI":"10.1073\/pnas.1007773107","volume":"107","author":"J Gil-Humanes","year":"2010","unstructured":"Gil-Humanes, J., Pist\u00f3n, F., Tollefsen, S., Sollid, L. M. & Barro, F. Effective shutdown in the expression of celiac disease-related wheat gliadin T-cell epitopes by RNA interference. Proc. Natl. Acad. Sci. USA 107, 17023\u201317028 (2010).","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"BFsrep44609_CR57","doi-asserted-by":"crossref","first-page":"136","DOI":"10.1186\/1471-2229-13-136","volume":"13","author":"F Pist\u00f3n","year":"2013","unstructured":"Pist\u00f3n, F., Gil-Humanes, J. & Barro, F. Integration of promoters, inverted repeat sequences and proteomic data into a model for high silencing efficiency of celiac disease related gliadins in bread wheat. BMC Plant Biol. 13, 136 (2013).","journal-title":"BMC Plant Biol"},{"key":"BFsrep44609_CR58","doi-asserted-by":"crossref","first-page":"393","DOI":"10.1186\/s12870-014-0393-1","volume":"14","author":"SB Altenbach","year":"2014","unstructured":"Altenbach, S. B., Tanaka, C. K. & Seabourn, B. W. Silencing of omega-5 gliadins in transgenic wheat eliminates a major source of environmental variability and improves dough mixing properties of flour. BMC Plant Biol. 14, 393 (2014).","journal-title":"BMC Plant Biol"},{"key":"BFsrep44609_CR59","doi-asserted-by":"crossref","unstructured":"Hilscher, J., B\u00fcrstmayr, H. & Stoger, E. Targeted modification of plant genomes for precision crop breeding. Biotechnol. J. doi: 10.1002\/biot.201600173 (2016).","DOI":"10.1002\/biot.201600173"},{"key":"BFsrep44609_CR60","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1016\/S0923-1811(03)00156-7","volume":"33","author":"E Morita","year":"2003","unstructured":"Morita, E. et al. Fast \u03c9-gliadin is a major allergen in wheat-dependent excise-induced anaphylaxis. J. Derm. Sci. 33, 99\u2013104 (2003).","journal-title":"J. Derm. Sci"},{"key":"BFsrep44609_CR61","doi-asserted-by":"crossref","first-page":"912","DOI":"10.1016\/S0091-6749(99)70438-0","volume":"103","author":"K Palosuo","year":"1999","unstructured":"Palosuo, K. et al. A novel wheat gliadin as a cause of excise-induced anaphylaxis. J. Allergy Clin. Immunol. 103, 912\u2013917 (1999).","journal-title":"J. Allergy Clin. Immunol."},{"key":"BFsrep44609_CR62","doi-asserted-by":"crossref","first-page":"1195","DOI":"10.1053\/j.gastro.2014.12.049","volume":"148","author":"A Fasano","year":"2015","unstructured":"Fasano, A., Sapone, A., Zevallos, V. & Schuppan, D. Nonceliac gluten sensitivity. Gastroenterology 148, 1195\u2013204 (2015).","journal-title":"Gastroenterology"},{"key":"BFsrep44609_CR63","doi-asserted-by":"crossref","first-page":"214","DOI":"10.1186\/1471-2229-10-214","volume":"10","author":"C Yu","year":"2010","unstructured":"Yu, C. et al. Molecular analysis of phosphomannomutase (PMM) genes reveals a unique PMM duplication event in diverse Triticeae species and the main PMM isozymes in bread wheat tissues. BMC Plant Biol. 10, 214 (2010).","journal-title":"BMC Plant Biol"},{"key":"BFsrep44609_CR64","doi-asserted-by":"crossref","first-page":"418","DOI":"10.1111\/j.1469-8137.2011.03715.x","volume":"191","author":"GF Wang","year":"2011","unstructured":"Wang, G. F. et al. Molecular analysis of common wheat genes encoding three types of cytosolic heat shock protein 90 (Hsp90): functional involvement of cytosolic Hsp90s in the control of wheat seedling growth and disease resistance. New Phytol. 191, 418\u2013431 (2011).","journal-title":"New Phytol"},{"key":"BFsrep44609_CR65","doi-asserted-by":"crossref","first-page":"166","DOI":"10.1093\/bioinformatics\/btu638","volume":"31","author":"S Anders","year":"2015","unstructured":"Anders, S., Pyl, P. T. & Huber, W. HTSeq - A Python framework to work with high-throughput sequencing data. Bioinformatics 31, 166\u2013169 (2015).","journal-title":"Bioinformatics"},{"key":"BFsrep44609_CR66","doi-asserted-by":"crossref","first-page":"1154","DOI":"10.1016\/j.phytochem.2011.01.002","volume":"72","author":"WH Vensel","year":"2011","unstructured":"Vensel, W. H., Dupont, F. M., Sloane, S. & Altenbach, S. B. Effect of cleavage enzyme, search algorithm and decoy database on mass spectrometric identification of wheat gluten proteins. Phytochemistry 72, 1154\u20131161 (2011).","journal-title":"Phytochemistry"},{"key":"BFsrep44609_CR67","doi-asserted-by":"crossref","first-page":"1658","DOI":"10.1093\/bioinformatics\/btl158","volume":"22","author":"WZ Li","year":"2006","unstructured":"Li, W. Z. & Godzik, A. Cd-hit: a fast program for clustering and comparing large sets of protein or nucleotide sequences. Bioinformatics 22, 1658\u20131659 (2006).","journal-title":"Bioinformatics"}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/srep44609.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/srep44609","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/srep44609.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,12,24]],"date-time":"2022-12-24T00:14:31Z","timestamp":1671840871000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/srep44609"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2017,3,16]]},"references-count":67,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2017,4,20]]}},"alternative-id":["BFsrep44609"],"URL":"https:\/\/doi.org\/10.1038\/srep44609","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2017,3,16]]},"assertion":[{"value":"21 October 2016","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"9 February 2017","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"16 March 2017","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing financial interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"44609"}}