vir-nwТруды по прикладной ботанике, генетике и селекцииProceedings on applied botany, genetics and breeding2227-88342619-0982N.I. Vavilov All-Russian Institute of Plant Genetic Resources10.30901/2227-8834-2021-1-168-185vir-nw-885Research ArticleОБЗОРЫSURVEYSСтепень влияния глютенинов на качество зерна как одного из сложных полигенных признаков рода Triticum (обзор)The effect of glutenins on grain quality as one of the complex polygenic traits in the genus Triticum (a review)https://orcid.org/0000-0001-8799-0681ВоротынцеваМ. В.VorotyntsevaM. V.

656910,  г. Барнаул, Научный городок, 35

35 Nauchny Gorodok, Barnaul 656910

masha.vorotintseva@yandex.ruФедеральный Алтайский научный центр агробиотехнологийРоссияFederal Altai Scientific Center of Agro-BioTechnologiesRussian Federation2021030420211821168185Copyright © Воротынцева М.В., 20212021Воротынцева М.В.Vorotyntseva M.V.Данная работа распространяется под лицензией Creative Commons Attribution 4.0.This work is licensed under a Creative Commons Attribution 4.0 License.https://elpub.vir.nw.ru/jour/article/view/885

Оценка селекционного материала на основе белковых маркеров дает возможность достаточно быстро и качественно проводить отбор и контролировать передачу желаемых признаков от родительских форм потомкам. Вместе с тем для селекции необходимо выявление новых и стабильных белковых маркеров для определения качества зерна пшеницы. Широко изученными белками пшеницы являются запасные белки глютенины. Полная характеристика отдельных фракций и компонентов белков отражена во многих научных исследованиях, поэтому изучение генетических закономерностей накопления белков в зерне различных сортов пшеницы и использование при идентификации генотипов в качестве маркеров компонентов высокомолекулярных (HMW-GS) и низкомолекулярных (LMW-GS) субъединиц глютенина является весьма актуальной проблемой. В данной статье проведен полный анализ научных статей о структуре и молекулярном строении глютенинов, а также сделан сравнительный анализ 22 статей о степени влияния данных белков на качественные показатели зерна: седиментационный объем (мл), содержание белка в зерне/муке (%; 14% m.b.; 12,5% m.b.), содержание белка в зерне, время смешивания (мин), устойчивость к замесу (мин; мм), объем буханки хлеба (см3; мл), cила теста (10-4 J), отношение упругости к растяжимости. Итогом статьи явилось выделение лучших по показателям качества аллелей глютенина, которыми стали Glu-A1а, Glu-B1(h, f, b), Glu-D1d, Glu-A3d, Glu-B3d.

Evaluation of plant breeding material, based on protein markers, gives an opportunity to perform rapid and reliable selection and control the transfer of desired traits from parents to their progeny. A search for new and stable protein markers is needed to identify genotypes with high grain quality. Such storage proteins in wheat as glutenins have been studied profoundly enough. Full characterization of individual protein fractions and components can be found in many scientific publications, while studying genetic patterns of protein accumulation in the grain of different wheat cultivars and using high-molecular-weight (HMW) and lowmolecular-weight (LMW) glutenin subunits (GS) for genotype identification remain high in the research agenda. This is a comprehensive review of scientific publications about the structure and molecular organization of glutenins and a comparative analysis of 22 research papers about the degree of their effect on grain quality indicators: SDS-sedimentation volume (ml), grain/ flour protein content (%; 14% m.b.; 12,5% m.b.), mixing time (min), mixing tolerance (min; mm), bread loaf volume (cm3; ml), dough strength  (10-4J), and P/L ratio. As a result of reviewing, the best alleles (subunits) of glutenin were identi fied, namely: Glu-A1а, Glu-B1(h, f, b), Glu-D1d, Glu-A3d, and Glu-B3d.

пшеницаглютенинбелковые маркерыкачественные показатели зернаwheatgluteninprotein markersgrain quality indicatorsРабота выполнена в рамках государственного задания Федерального Алтайского научного центра агробиотехнологий № 0534-2019-0013 «Использование молекулярно-генетических методов исследований в селекции растений».The research was implemented within the framework of the State Task for the Federal Altai Scientific Centre of Agro-BioTechnologies, No. 0534-2019-001 “The use of molecular genetic research methods in plant breeding”ReferencesAhn J.H., Kang C.S., Jeung J.U., Baik B.K., Peña R.J., Park C.S. Effect of allelic variations at the Glu-D1, Glu-A3, Glu-B3 and Pinb-D1 loci on flour characteristics and bread loaf volume. International Food Research Journal. 2014; 21(3):1141-1149.Ahn J.H., Kang C.S., Jeung J.U., Baik B.K., Peña R.J., Park C.S. Effect of allelic variations at the Glu-D1, Glu-A3, Glu-B3 and Pinb-D1 loci on flour characteristics and bread loaf volume. International Food Research Journal. 2014;21(3):1141-1149.An X., Li Q., Yan Y., Xiao Y., Xsam S.L.K., Zeller F.J. Genetic diversity of European spelt wheat (Triticum aestivum ssp. spelta L. em. Thell.) revealed by glutenin subunit variations at the Glu-1 and Gli-3 loci. Euphytica. 2005;146(3):193-201. DOI: 10.1007/s10681-005-9002-6An X., Li Q., Yan Y., Xiao Y., Xsam S.L.K., Zeller F.J. Genetic diversity of European spelt wheat (Triticum aestivum ssp. spelta L. em. Thell.) revealed by glutenin subunit variations at the Glu-1 and Gli-3 loci. Euphytica. 2005;146(3):193-201. DOI: 10.1007/s10681-005-9002-6An X., Zhang Q., Yan Y., Li Q., Zhang Y., Wang A. et al. Cloning and molecular characterization of three novel LMW-i glutenin subunit genes from cultivated einkorn (Triticum monococcum L.). Theoretical and Applied Genetics. 2006;113(3):383-395. DOI: 10.1007/s00122-006-0299-xAn X., Zhang Q., Yan Y., Li Q., Zhang Y., Wang A. et al. Cloning and molecular characterization of three novel LMW-i glutenin subunit genes from cultivated einkorn (Triticum monococcum L.). Theoretical and Applied Genetics. 2006;113(3):383-395. DOI: 10.1007/s00122-006-0299-xAndersen J.R., Lübberstedt T. Functional markers in plants. Trends in Plant Science. 2003;8(11):554-560. DOI: 10.1016/j. tplants.2003.09.010Andersen J.R., Lübberstedt T. Functional markers in plants. Trends in Plant Science. 2003;8(11):554-560. DOI: 10.1016/j. tplants.2003.09.010Anjum F.M., Khan M.R., Din A., Saeed M., Pasha I., Arshad M.U. Wheat gluten: high molecular weight glutenin subunits – structure, genetics, and relation to dough elasticity. Journal of Food Science. 2007;72(3):56-63. DOI: 10.1111/j.1750-3841.2007.00292.xAnjum F.M., Khan M.R., Din A., Saeed M., Pasha I., Arshad M.U. Wheat gluten: high molecular weight glutenin subunits – structure, genetics, and relation to dough elasticity. Journal of Food Science. 2007;72(3):56-63. DOI: 10.1111/j.1750-3841.2007.00292.xAxford D.W.E.; McDermott E.E.; Redman D.G. Small-scale tests of bread-making quality. Milling Feed and Fertilizer. 1978;161:18-20.Axford D.W.E.; McDermott E.E.; Redman D.G. Small-scale tests of bread-making quality. Milling Feed and Fertilizer. 1978;161:18-20.Barro F., Barceló P., Lazzeri P.A., Shewry P.R., Ballesteros J., Martin A. Functional properties of flours from field grown transgenic wheat lines expressing the HMW glutenin subunit 1Ax1 and 1Dx5 genes. Molecular Breeding. 2003;12(3):223-229. DOI: 10.1023/A:1026367214120Barro F., Barceló P., Lazzeri P.A., Shewry P.R., Ballesteros J., Martin A. Functional properties of flours from field grown transgenic wheat lines expressing the HMW glutenin subunit 1Ax1 and 1Dx5 genes. Molecular Breeding. 2003;12(3):223-229. DOI: 10.1023/A:1026367214120Branlard G., Dardevet R., Saccomano F., Lagoutte F., Gourdon J. Genetics diversity of wheat storage proteins and bread wheat quality. Euphytica. 2001;119(1-2):59-67. DOI: 10.1023/A:1017586220359Branlard G., Dardevet R., Saccomano F., Lagoutte F., Gourdon J. Genetics diversity of wheat storage proteins and bread wheat quality. Euphytica. 2001;119(1-2):59-67. DOI: 10.1023/A:1017586220359Brites C., Carrillo J.M. Influence of high molecular weight (HMW) and low molecular weight (LMW) glutenin subunits controlled by Glu-1 and Glu-3 loci on durum wheat quality. Cereal Chemistry. 2001;78(1):59-63. DOI: 10.1094/CCHEM.2001.78.1.59Brites C., Carrillo J.M. Influence of high molecular weight (HMW) and low molecular weight (LMW) glutenin subunits controlled by Glu-1 and Glu-3 loci on durum wheat quality. Cereal Chemistry. 2001;78(1):59-63. DOI: 10.1094/CCHEM.2001.78.1.59Caballero L., Martin L.M., Alvarez J.B. Relationships between the HMW- and LMW-glutenin subunits and SDS-sedimentation volume in Spanish hulled wheat lines. Czech Journal of Genetics and Plant Breeding. 2008a;44(3):114-117. DOI: 10.17221/8/2008-CJGPBCaballero L., Martin L.M., Alvarez J.B. Relationships between the HMW- and LMW-glutenin subunits and SDS-sedimentation volume in Spanish hulled wheat lines. Czech Journal of Genetics and Plant Breeding. 2008a;44(3):114-117. DOI: 10.17221/8/2008-CJGPBCaballero L., Martin L.M., Alvarez J.B. Variation of high molecular weight glutenin subunits in two neglected tetraploid wheat subspecies. Czech Journal of Genetics and Plant Breeding. 2008b;44(4):140-146. DOI: 10.17221/61/2008-CJGPBCaballero L., Martin L.M., Alvarez J.B. Variation of high molecular weight glutenin subunits in two neglected tetraploid wheat subspecies. Czech Journal of Genetics and Plant Breeding. 2008b;44(4):140-146. DOI: 10.17221/61/2008-CJGPBCarrillo J.M., Rousset M., Qualset C.O., Kasarda D.D. Use of recombinant inbred lines of wheat for study of association of high-molecular-weight glutenin subunit alleles to quantitative traits: 1. Grain yield and quality prediction tests. Theoretical and Applied Genetics. 1990;79(3):321-330. DOI: 10.1007/BF01186074Carrillo J.M., Rousset M., Qualset C.O., Kasarda D.D. Use of recombinant inbred lines of wheat for study of association of high-molecular-weight glutenin subunit alleles to quantitative traits: 1. Grain yield and quality prediction tests. Theoretical and Applied Genetics. 1990;79(3):321-330. DOI: 10.1007/BF01186074Chacón E.A., Vázquez F.J., Giraldo P., Carrillo J.M., Benavente E., Rodriguez-Quijano M. Allelic variation for prolamins in Spanish durum wheat landraces and its relationship with quality traits. Agronomy. 2020;10(1):136. DOI: 10.3390/agronomy10010136Chacón E.A., Vázquez F.J., Giraldo P., Carrillo J.M., Benavente E., Rodriguez-Quijano M. Allelic variation for prolamins in Spanish durum wheat landraces and its relationship with quality traits. Agronomy. 2020;10(1):136. DOI: 10.3390/agronomy10010136Cho S.W., Roy S.K., Chun J.B., Cho K., Park C.S. Overexpression of the Bx7 high molecular weight glutenin subunit on the Glu-B1 locus in a Korean wheat landrace. Plant Biotechnology Reports. 2017;11(2):97-105. DOI: 10.1007/s11816-017-0434-yCho S.W., Roy S.K., Chun J.B., Cho K., Park C.S. Overexpression of the Bx7 high molecular weight glutenin subunit on the Glu-B1 locus in a Korean wheat landrace. Plant Biotechnology Reports. 2017;11(2):97-105. DOI: 10.1007/s11816-017-0434-yЧуприна Ю.Ю. Характеристика коллекционных образцов пшеницы яровой в условиях восточной лесостепи Украины. В кн.: Молодые исследователи агропромышленного и лесного комплексов – регионам. Материалы V Международной молодежной научно-практической конференции, 23 апреля 2019 г. Вологда; 2019. С.161-166.Chuprina Yu.Yu. Characterization of spring wheat germplasm accessions in the environments of the eastern Ukainian forest-steppe (Kharakteristika kollektsionnykh obraztsov pshenitsy yarovoy v usloviyakh vostochnoy lesostepi Ukrainy). In: Young Researchers of Agro-Industrial and Forestry Sectors – for the Regions (Molodye issledovateli agropromyshlennogo i lesnogo kompleksov – regionam). Proceesings of the V International Youth-Oriented Scientific and Practical Conference, April 23, 2019. Vologda; 2019. p.161-166. [in Russian]Cloutier S., Rampitsch C., Penner G.A., Lukow O.M. Cloning and expression of a LMW-i glutenin gene. Journal of Cereal Science. 2001;33(2):143-154. DOI: 10.1006/jcrs.2000.0359Cloutier S., Rampitsch C., Penner G.A., Lukow O.M. Cloning and expression of a LMW-i glutenin gene. Journal of Cereal Science. 2001;33(2):143-154. DOI: 10.1006/jcrs.2000.0359D’Ovidio R., Marchitelli C., Cardelli L.E., Porceddu E., Hart G.E. Sequence similarity between allelic Glu-B3 genes related to quality properties of durum wheat. Theoretical and Applied Genetics. 1999;98(3-4);455-461. DOI: 10.1007/s001220051091D’Ovidio R., Marchitelli C., Cardelli L.E., Porceddu E., Hart G.E. Sequence similarity between allelic Glu-B3 genes related to quality properties of durum wheat. Theoretical and Applied Genetics. 1999;98(3-4);455-461. DOI: 10.1007/s001220051091Dick J.W., Quick J.S. A modified screening test for rapid estimation of gluten strength in early-generation durum wheat breeding lines. Cereal Chemistry. 1983;60(4):315-318.Dick J.W., Quick J.S. A modified screening test for rapid estimation of gluten strength in early-generation durum wheat breeding lines. Cereal Chemistry. 1983;60(4):315-318.Du Cros D.L. Glutenin proteins and gluten strength in durum wheat. Journal of Cereal Science. 1987;5(1):3-12. DOI: 10.1016/S0733-5210(87)80003-6Du Cros D.L. Glutenin proteins and gluten strength in durum wheat. Journal of Cereal Science. 1987;5(1):3-12. DOI: 10.1016/S0733-5210(87)80003-6Дуктова Н.А. Создание и оценка по хозяйственно-биологическим признакам новых образцов яровой твердой пшеницы (T. durum Desf.) различного генетического происхождения: дис. ... канд. с.-х. наук. Горки: Белорусская ГСХА; 2007.Duktova N.A. Development and evaluation for economically important traits of new spring durum wheat cultivars (T. durum Desf.) of various genetic origin (Sozdaniye i otsenka po khozyaystvenno-biologicheskim priznakam novykh obraztsov yarovoy tverdoy pshenitsy (T. durum Desf.) razlichnogo geneticheskogo proiskhozhdeniya) [dissertation]. Gorki: Belarussian State Agricultural Academy; 2007. [in Russian]Field J.M., Tatham A.S., Shewry P.R. The structure of a high-Mr subunit of durum-wheat (Triticum durum) gluten. Biochemical Journal. 1987;247(1):215-221. DOI: 10.1042/bj2470215Field J.M., Tatham A.S., Shewry P.R. The structure of a highMr subunit of durum-wheat (Triticum durum) gluten.Biochemical Journal. 1987;247(1):215-221. DOI: 10.1042/bj2470215Figueroa J.D.C., Maucher T., Reule W., Peña R.J. Influence of high molecular weight glutenins on viscoelastic properties of intact wheat kernel and relation to functional properties of wheat dough. Cereal Chemistry. 2009;86(2):139-144. DOI: 10.1094/CCHEM-86-2-0139Figueroa J.D.C., Maucher T., Reule W., Peña R.J. Influence of high molecular weight glutenins on viscoelastic properties of intact wheat kernel and relation to functional properties of wheat dough. Cereal Chemistry. 2009;86(2):139-144. DOI: 10.1094/CCHEM-86-2-0139Фомина Е.А., Кулинкович С.Н., Малышев С.В., Урбанович О.Ю. Идентификация полиморфизма HMW глютенинов у сортообразцов из коллекционных и селекционных питомников озимой пшеницы. Земледелие и селекция в Беларуси. 2014;50:326-335.Fomina E.A., Kulinkovich S.N., Malyshev S.V., Urbanovich O.Y. Identification of HMW glutenin polymorphism in variety samples from collection and breeding nurseries of winter wheat. Zemledeliye i selektsiya v Belarusi = Crop Husbandry and Plant Breeding in Belarus. 2014;50:326-335.[in Russian]Gianibelli M.C., Larroque O., MacRitchie F., Wrigley C.W. Biochemical, genetic, and molecular characterization of wheat endosperm proteins. Cereal Chemistry. 2001;78(6):635-646. DOI: 10.1094/CCHEM.2001.78.6.635Gianibelli M.C., Larroque O., MacRitchie F., Wrigley C.W. Biochemical, genetic, and molecular characterization of wheat endosperm proteins. Cereal Chemistry. 2001;78(6):635-646. DOI: 10.1094/CCHEM.2001.78.6.635Gilbert S.M., Wellner N., Belton P.S., Greenfield J.A., Siligardi G., Shewry P.R. et al. Expression and characterisation of a highly repetitive peptide derived from a wheat seed storage protein. Biochimica et Biophysica Acta (BBA) – Protein Structure and Molecular Enzymology. 2000;1479(1-2):135-146. DOI: 10.1016/S0167-4838(00)00059-5Gilbert S.M., Wellner N., Belton P.S., Greenfield J.A., Siligardi G., Shewry P.R. et al. Expression and characterisation of a highly repetitive peptide derived from a wheat seed storage protein. Biochimica et Biophysica Acta (BBA) – Protein Structure and Molecular Enzymology. 2000;1479(1-2):135-146. DOI: 10.1016/S0167-4838(00)00059-5Greene F.C., Anderson O.D., Yip R.E., Halford N.G., Romero J.M.M., Shewry P.R. Analysis of possible quality-related sequence variations in the 1D glutenin high molecular weight subunit genes of wheat. In: Proceedings of the Seventh International Wheat Genetics Symposium, held at Cambridge, UK, 13–19 July 1988. Cambridge; 1988. p.735-740.Greene F.C., Anderson O.D., Yip R.E., Halford N.G., Ro me ro J.M.M., Shewry P.R. Analysis of possible quality-related sequence variations in the 1D glutenin high molecular weight subunit genes of wheat. In: Proceedings of the Seventh International Wheat Genetics Symposium, held at Cambridge, UK, 13–19 July 1988. Cambridge; 1988. p.735-740.He Z.H., Lin Z.J., Wang L.J., Xiao Z.M., Wan F.S., Zhuang Q.S. Classification on Chinese wheat regions based on quality. Scientia Agricultura Sinica. 2002;35(4):359-364. [in Chinese]He Z.H., Lin Z.J., Wang L.J., Xiao Z.M., Wan F.S., Zhuang Q.S. Classification on Chinese wheat regions based on quality. Scientia Agricultura Sinica. 2002;35(4):359-364. [in Chinese]He Z.H., Liu L., Peña R.J. Allelic variation at the Glu-1 and Glu-3 loci, presence of 1BL/1RS translocation, and their effect on dough properties of Chinese bread wheats. In: D. Lafiandra, S. Masci, R. D’Ovidio (eds). The Gluten Proteins. The Proceedings of the 8th Gluten Workshop held on 8–10 September 2003 in Viterbo, Italy. Cambridge: The Royal Society of Chemistry; 2004. p.97-100.He Z.H., Liu L., Peña R.J. Allelic variation at the Glu-1 and Glu-3 loci, presence of 1BL/1RS translocation, and their effect on dough properties of Chinese bread wheats. In: D. Lafiandra, S. Masci, R. D’Ovidio (eds). The Gluten Proteins. The Proceedings of the 8th Gluten Workshop held on 8–10 September 2003 in Viterbo, Italy. Cambridge: The Royal Society of Chemistry; 2004. p.97-100.Horvat C.A., Jurković Z., Sudar R., Pavlinić D., Simić G. The relative amount of HMW glutenin subunits of OS wheat cultivars in relation to bread-making quality. Cereal Research Communications. 2002;30:415-422. DOI: 10.1007/BF03543438Horvat C.A., Jurković Z., Sudar R., Pavlinić D., Simić G. The relative amount of HMW glutenin subunits of OS wheat cultivars in relation to bread-making quality. Cereal Research Communications. 2002;30:415-422. DOI: 10.1007/BF03543438Huang Y.C., Lai H.M. Noodle quality affected by different cereal starches. Journal of Food Engineering. 2010;97(2):135-143. DOI: 10.1016/j.jfoodeng.2009.10.002Huang Y.C., Lai H.M. Noodle quality affected by different cereal starches. Journal of Food Engineering. 2010;97(2):135-143. DOI: 10.1016/j.jfoodeng.2009.10.002Johansson E., Svensson G. Contribution of the high molecular weight glutenin subunit 21* to breadmaking quality of Swedish wheats. Cereal Chemistry. 1995;72(3):287-290.Johansson E., Svensson G. Contribution of the high molecular weight glutenin subunit 21* to breadmaking quality of Swedish wheats. Cereal Chemistry. 1995;72(3):287-290.Kang C.S., Jeung J.U., Baik B.K., Park C.S. Effects of allelic variations in Wx-1, Glu-D1, Glu-B3, and Pinb-D1 loci on flour characteristics and white salted noodle-making quality of wheat flour. Cereal Chemistry. 2012;89(6):296-306. DOI: 10.1094/cchem-03-12-0034-rKang C.S., Jeung J.U., Baik B.K., Park C.S. Effects of allelic variations in Wx-1, Glu-D1, Glu-B3, and Pinb-D1 loci on flour characteristics and white salted noodle-making quality of wheat flour. Cereal Chemistry. 2012;89(6):296-306. DOI: 10.1094/cchem-03-12-0034-rKasarda D.D. Glutenin structure in relation to wheat quality. Wheat is Unique. 1989;277:302.Kasarda D.D. Glutenin structure in relation to wheat quality. Wheat is Unique. 1989;277:302.Kasarda D.D., Bernardin J.E., Nimmo C.C. Wheat proteins. Advances in Cereal Science and Technology. 1976;1:158-236.Kasarda D.D., Bernardin J.E., Nimmo C.C. Wheat proteins. Advances in Cereal Science and Technology. 1976;1:158-236.Kaya Y., Akcura M. Effects of genotype and environment on grain yield and quality traits in bread wheat (T. aestivum L.). Food Science and Technology. 2014;34(2):386-393. DOI: 10.1590/fst.2014.0041Kaya Y., Akcura M. Effects of genotype and environment on grain yield and quality traits in bread wheat (T. aestivum L.). Food Science and Technology. 2014;34(2):386-393. DOI: 10.1590/fst.2014.0041Kolpakova V.V., Molchanova E.N., Vasil’ev A.V., Chumikina L.V. Physicochemical properties of proteins from wheat grown under high contrast climatic conditions. Applied Biochemistry and Microbiology. 2007;43(3):347-355. DOI: 10.1134/S0003683807030180Kolpakova V.V., Molchanova E.N., Vasil’ev A.V., Chumikina L.V. Physicochemical properties of proteins from wheat grown under high contrast climatic conditions. Applied Biochemistry and Microbiology. 2007;43(3):347-355. DOI: 10.1134/S0003683807030180Кременевская Е.М., Гриб С.И., Буштевич В.Н. Cоздание яровых форм гексаплоидных тритикале с интрогрессией пшеничного локуса Glu-D1. Земледелие и селекция в Беларуси. 2013;49:299-311.Kremenevskaya E.M., Grib S.I., Bushtevich V.N. Development of spring forms of hexaploid triticale with introgression of Glu-D1 wheat locus. Zemledeliye i selektsiya v Belarusi = Crop Husbandry and Plant Breeding in Belarus. 2013;49:299-311. [in Russian]Krupnov V.A., Krupnova O.V. Genetic architecture of grain protein content in wheat. Russian Journal of Genetics. 2012;48(2):129-138. DOI: 10.1134/S1022795412010139Krupnov V.A., Krupnova O.V. Genetic architecture of grain protein content in wheat. Russian Journal of Genetics. 2012;48(2):129-138. DOI: 10.1134/S1022795412010139León E., Marín S., Giménez M.J., Piston F., Rodríguez-Quijano M., Shewry P.R. et al. Mixing properties and dough functionality of transgenic lines of a commercial wheat cultivar expressing the 1Aх1, 1D6х5 and 1Dy10 HMW glutenin subunit genes. Journal of Cereal Science. 2009;49(1):148-156. DOI: 10.1016/j.jcs.2008.08.002León E., Marín S., Giménez M.J., Piston F., Rodríguez-Quijano M., Shewry P.R. et al. Mixing properties and dough functionality of transgenic lines of a commercial wheat cultivar expressing the 1Aх1, 1D6х5 and 1Dy10 HMW glutenin subunit genes. Journal of Cereal Science. 2009;49(1):148-156. DOI: 10.1016/j.jcs.2008.08.002Lew E.J.L., Kuzmicky D.D., Kasarda D.D. Characterization of low-molecular-weight glutenin subunits by reversephase high-performance liquid chromatography, sodium dodecyl sulphate-polyacrylamide gel electrophoresis, and N-terminal amino acid sequencing. Cereal Chemistry. 1992;69(5):508-515.Lew E.J.L., Kuzmicky D.D., Kasarda D.D. Characterization of low-molecular-weight glutenin subunits by reversephase high-performance liquid chromatography, sodium dodecyl sulphate-polyacrylamide gel electrophoresis, and N-terminal amino acid sequencing. Cereal Chemistry. 1992;69(5):508-515.Li L., Jun Y., Yan Z., Zhonghu H., Peña-Bautista R.J., Li-Ping Z. Allelic variation at the Glu-1 and Glu-3 loci and presence of 1B/1R translocation, and their effects on processing quality in cultivars and advanced lines from autumnsown wheat regions in China. Scientia Agricultura Sinica. 2005;38(10):1944-1950. [in Chinese]Li L., Jun Y., Yan Z., Zhonghu H., Peña-Bautista R.J., Li-Ping Z. Allelic variation at the Glu-1 and Glu-3 loci and presence of 1B/1R translocation, and their effects on processing quality in cultivars and advanced lines from autumnsown wheat regions in China. Scientia Agricultura Sinica. 2005;38(10):1944-1950. [in Chinese]Li W., Zhang P., Fellers J.P., Friebe B., Gill B.S. Sequence composition, organization, and evolution of the core Triticeae genome. The Plant Journal. 2004;40(4):500-511. DOI: 10.1111/j.1365-313X.2004.02228.xLi W., Zhang P., Fellers J.P., Friebe B., Gill B.S. Sequence composition, organization, and evolution of the core Triticeae genome. The Plant Journal. 2004;40(4):500-511. DOI: 10.1111/j.1365-313X.2004.02228.xLi Y., Huang C., Sui X., Fan Q., Li G., Chu X. Genetic variation of wheat glutenin subunits between landraces and varieties and their contributions to wheat quality improvement in China. Euphytica. 2009;169(2):159-168. DOI: 10.1007/s10681-009-9905-8Li Y., Huang C., Sui X., Fan Q., Li G., Chu X. Genetic variation of wheat glutenin subunits between landraces and varieties and their contributions to wheat quality improvement in China. Euphytica. 2009;169(2):159-168. DOI: 10.1007/s10681-009-9905-8Li Y., Zhou R., Branlard G., Jia J. Development of introgression lines with 18 alleles of glutenin subunits and evaluation of the effects of various alleles on quality related traits in wheat (Triticum aestivum L.). Journal of Cereal Science. 2010; 51(1):127-133. DOI: 10.1016/j.jcs.2009.10.008Li Y., Zhou R., Branlard G., Jia J. Development of introgression lines with 18 alleles of glutenin subunits and evaluation of the effects of various alleles on quality related traits in wheat (Triticum aestivum L.). Journal of Cereal Science. 2010;51(1):127-133. DOI: 10.1016/j.jcs.2009.10.008Liu L., He Z., Yan J., Zhang Y., Xia X., Peña R.J. Allelic variation at the Glu-1 and Glu-3 loci, presence of the 1B.1R translocation, and their effects on mixographic properties in Chinese bread wheats. Euphytica. 2005;142(3):197-204. DOI: 10.1007/s10681-005-1682-4Liu L., He Z., Yan J., Zhang Y., Xia X., Peña R.J. Allelic variation at the Glu-1 and Glu-3 loci, presence of the 1B.1R translocation, and their effects on mixographic properties in Chinese bread wheats. Euphytica. 2005;142(3):197-204. DOI: 10.1007/s10681-005-1682-4Luo Z., Chen F., Feng D., Xia G. LMW-GS genes in Agropyron elongatum and their potential value in wheat breeding. Theoretical and Applied Genetics. 2005;111(2):272-280. DOI: 10.1007/s00122-005-2021-9Luo Z., Chen F., Feng D., Xia G. LMW-GS genes in Agropyron elongatum and their potential value in wheat breeding. Theoretical and Applied Genetics. 2005;111(2):272-280. DOI: 10.1007/s00122-005-2021-9Magallanes-López A.M., Ammar K., Morales-Dorantes A., González-Santoyo H., Crossa J., Guzmán C. Grain quality traits of commercial durum wheat varieties and their relationships with drought stress and glutenins composition. Journal of Cereal Science. 2017;75:1-9. DOI: 10.1016/j.jcs.2017.03.005Magallanes-López A.M., Ammar K., Morales-Dorantes A., González-Santoyo H., Crossa J., Guzmán C. Grain quality traits of commercial durum wheat varieties and their relationships with drought stress and glutenins composition. Journal of Cereal Science. 2017;75:1-9. DOI: 10.1016/j.jcs.2017.03.005Marcussen T., Sandve S.R., Heier L., Spannagl M., Pfeifer M.; The International Wheat Genome Sequencing Consortium et al. Ancient hybridizations among the ancestral genomes of bread wheat. Science. 2014;345(6194):1250092. DOI: 10.1126/science.1250092Marcussen T., Sandve S.R., Heier L., Spannagl M., Pfeifer M.; The International Wheat Genome Sequencing Consortium et al. Ancient hybridizations among the ancestral genomes of bread wheat. Science. 2014;345(6194):1250092. DOI: 10.1126/science.1250092Maruyama-Funatsuki W., Takata K., Nishio Z., Tabiki T., Yahata E., Kato A. et al. Identification of low-molecular weight glutenin subunits of wheat associated with bread-making quality. Plant Breeding. 2004;123(4):355-360. DOI: 10.1111/j.1439-0523.2004.00978.xMaruyama-Funatsuki W., Takata K., Nishio Z., Tabiki T., Yahata E., Kato A. et al. Identification of low-molecular weight glutenin subunits of wheat associated with bread-making quality. Plant Breeding. 2004;123(4):355-360. DOI: 10.1111/j.1439-0523.2004.00978.xMasci S., D’Ovidio R., Lafiandra D., Kasarda D.D. A 1B-coded low-molecular-weight glutenin subunit associated with quality in durum wheats shows strong similarity to a subunit present in some bread wheat cultivars. Theoretical and Applied Genetics. 2000;100(3):396-400. DOI: 10.1007/s001220050052Masci S., D’Ovidio R., Lafiandra D., Kasarda D.D. A 1B-coded low-molecular-weight glutenin subunit associated with quality in durum wheats shows strong similarity to a subunit present in some bread wheat cultivars. Theoretical and Applied Genetics. 2000;100(3):396-400. DOI: 10.1007/s001220050052Masci S., Rovelli L., Kasarda D.D., Vensel W.H., Lafiandra D. Characterisation and chromosomal localisation of C-type low-molecular-weight glutenin subunits in the bread wheat cultivar Chinese Spring. Theoretical and Applied Genetics. 2002;104(2-3):422-428. DOI: 10.1007/s001220100761Masci S., Rovelli L., Kasarda D.D., Vensel W.H., Lafiandra D. Characterisation and chromosomal localisation of C-type low-molecular-weight glutenin subunits in the bread wheat cultivar Chinese Spring. Theoretical and Applied Genetics. 2002;104(2-3):422-428. DOI: 10.1007/s001220100761Meppelink E.K. Einsatzmоglichkeit des Microbackversuches in der Weizenzüichtung. Getreide, Mehl und Brot. 1981;35:107-109. [in German]Meppelink E.K. Einsatzmоglichkeit des Microbackversuches in der Weizenzüichtung. Getreide, Mehl und Brot. 1981;35:107-109. [in German]Metakovsky E.V., Wrigley C.W., Bekes F., Gupta R.B. Gluten polypeptides as useful genetic markers of dough quality in Australian wheats. Australian Journal of Agricultural Research. 1990;41(2):289-306. DOI: 10.1071/AR9900289Metakovsky E.V., Wrigley C.W., Bekes F., Gupta R.B. Gluten polypeptides as useful genetic markers of dough quality in Australian wheats. Australian Journal of Agricultural Research. 1990;41(2):289-306. DOI: 10.1071/AR9900289Растительный белок / под ред. Т.П. Микулович. Москва: Агропромиздат; 1991.Mikulovich T.P. (ed.). Plant protein (Rastitelny belok). Moscow: Agropromizdat; 1991. [in Russian]Miles M.J., Carr H.J., McMaster T., l’Anson K.J., Belton P.S., Morris V.J. et al. Scanning tunneling microscopy of a wheat seed storage protein reveals details of an unusual supersecondary structure. Proceedings of the National Academy of Sciences of the United States of America. 1991;88(1):68-71. DOI: 10.1073/pnas.88.1.68Miles M.J., Carr H.J., McMaster T., l’Anson K.J., Belton P.S., Morris V.J. et al. Scanning tunneling microscopy of a wheat seed storage protein reveals details of an unusual supersecondary structure. Proceedings of the National Academy of Sciences of the United States of America. 1991;88(1):68-71. DOI: 10.1073/pnas.88.1.68Mohan D., Gupta R.K. Gluten characteristics imparting bread quality in wheats differing for high molecular weight glutenin subunits at Glu D1 locus. Physiology and Molecular Biology of Plants. 2015;21(3):447-451. DOI: 10.1007/s12298-015-0298-yMohan D., Gupta R.K. Gluten characteristics imparting bread quality in wheats differing for high molecular weight glutenin subunits at Glu D1 locus. Physiology and Molecular Biology of Plants. 2015;21(3):447-451. DOI: 10.1007/s12298-015-0298-yMoonen J.H.E., Scheepstra A., Graveland A.; Centraal Instituut voor Voedingsonderzoek TNO. The positive effects of the high molecular weight subunits 3+10 and 2* of glutenin on the bread-making quality of wheat cultivars. Euphytica. 1983;32(3):735-742. DOI: 10.1007/bf00042153Moonen J.H.E., Scheepstra A., Graveland A.; Centraal Instituut voor Voedingsonderzoek TNO. The positive effects of the high molecular weight subunits 3+10 and 2* of glutenin on the bread-making quality of wheat cultivars. Euphytica. 1983;32(3):735-742. DOI: 10.1007/bf00042153Moragues M., Zarco-Hernández J., Moralejo M.A., Royo С. Genetic diversity of glutenin protein subunits composition in durum wheat landraces [Triticum turgidum ssp. turgidum convar. durum (Desf.) MacKey] from the Mediterranean basin. Genetic Resources and Crop Evolution. 2006;53(5):993-1002. DOI: 10.1007/s10722-004-7367-3Moragues M., Zarco-Hernández J., Moralejo M.A., Royo С. Genetic diversity of glutenin protein subunits composition in durum wheat landraces [Triticum turgidum ssp. turgidum convar. durum (Desf.) MacKey] from the Mediterranean basin. Genetic Resources and Crop Evolution. 2006;53(5):993-1002. DOI: 10.1007/s10722-004-7367-3Nakamura H. Genetic diversity of high-molecular-weight glutenin subunit compositions in landraces of hexaploid wheat from Japan. Euphytica. 2000;120(2):227-234. DOI: 10.1023/A:1017514423061Nakamura H. Genetic diversity of high-molecular-weight glutenin subunit compositions in landraces of hexaploid wheat from Japan. Euphytica. 2000;120(2):227-234. DOI: 10.1023/A:1017514423061Нечаев А.П., Дубцова Г.Н., Колпакова В.В. Белки пшеницы. Технология получения и применения (состояние, проблемы, пути развития). Известия высших учебных заведений. Пищевая технология. 1995;1-2(224-225):28-30.Nechaev A.P., Dubtsova G.N., Kolpakova V.V. Wheat proteins. Production technology and application (state, problems, ways of development) (Belki pshenitsy. Tekhnologiya polucheniya i primeneniya [sostoyanie, problemy, puti razvitiya]). Food Technology. 1995;1-2(224-225):28-30. [in Russian]Орловская О.А., Вакула С.И., Хотылева Л.В., Кильчевский А.В. Качество зерна у линий мягкой пшеницы с генетическим материалом Triticum kiharae. Фактори експериментальної еволюції організмів. 2018;(23):108-113.Orlovskaya O.A., Vakula S.I., Khotyleva L.V., Kilchevsky A.V. Grain quality in soft wheat lines with genetic material of Triticum kiharae (Kachestvo zerna u liniy myagkoy pshenitsy s geneticheskim materialom Triticum kiharae). Faktory eksperimentalnoy evolyutsіi organіzmіv = Factors of Experimental Evolution of Organisms. 2018;(23):108-113. [in Russian]Park C.S., Kang C.S., Jeung J.U., Woo S.H. Influence of allelic variations in glutenin on the quality of pan bread and white salted noodles made from Korean wheat cultivars. Euphytica. 2011;180(2):235-250. DOI: 10.1007/s10681-011-0385-2Park C.S., Kang C.S., Jeung J.U., Woo S.H. Influence of allelic variations in glutenin on the quality of pan bread and white salted noodles made from Korean wheat cultivars. Euphytica. 2011;180(2):235-250. DOI: 10.1007/s10681-011-0385-2Payne P.I., Corfield K.G., Holt L.M., Blackman J.A. Correlations between the inheritance of certain high-molecular weight subunits of glutenin and bread-making quality in progenies of six crosses of bread wheat. Journal of the Science of Food and Agriculture. 1981;32(1):51-60. DOI: 10.1002/jsfa.2740320109Payne P.I., Corfield K.G., Holt L.M., Blackman J.A. Correlations between the inheritance of certain high-molecular weight subunits of glutenin and bread-making quality in progenies of six crosses of bread wheat. Journal of the Science of Food and Agriculture. 1981;32(1):51-60. DOI: 10.1002/jsfa.2740320109Payne P.I., Holt L.M., Jackson E.A., Law C.N. Wheat storage proteins: their genetics and their potential for manipulation by plant breeding. Philosophical Transactions of the Royal Society. Biological Sciences. 1984;304:359-371. DOI: 10.1098/rstb.1984.0031Payne P.I., Holt L.M., Jackson E.A., Law C.N. Wheat storage proteins: their genetics and their potential for manipulation by plant breeding. Philosophical Transactions of the Royal Society. Biological Sciences. 1984;304:359-371. DOI: 10.1098/rstb.1984.0031Payne P.I., Seekings J.A., Worland A.J., Jarvis M.G., Holt L.M. Allelic variation of glutenin subunits and gliadins and its effect on breadmaking quality in wheat: Analysis of F5 progeny from Chinese Spring × Chinese Spring (Hope 1A). Journal of Cereal Science. 1987;6(2):103-118. DOI: 10.1016/S0733-5210(87)80047-4Payne P.I., Seekings J.A., Worland A.J., Jarvis M.G., Holt L.M. Allelic variation of glutenin subunits and gliadins and its effect on breadmaking quality in wheat: Analysis of F5 progeny from Chinese Spring × Chinese Spring (Hope 1A). Journal of Cereal Science. 1987;6(2):103-118. DOI: 10.1016/S0733-5210(87)80047-4Peña R.J., Amaya A., Rajaram S., Mujeeb-Kazi A. Variation in quality characteristics associated with some spring 1B.1R translocation wheats. Journal of Cereal Science. 1990;12:105-112. DOI: 10.1016/S0733-5210(09)80092-1Peña R.J., Amaya A., Rajaram S., Mujeeb-Kazi A. Variation in quality characteristics associated with some spring 1B.1R translocation wheats. Journal of Cereal Science. 1990;12:105-112. DOI: 10.1016/S0733-5210(09)80092-1Подгорный С.В., Самофалов А.П., Скрипка О.В. Генетические источники высокого содержания и качества белка для селекции озимой мягкой пшеницы. Аграрный вестник Урала. 2 0 16; 6(148):51-55.Podgorny S.V., Samofalov A.P., Skripka O.V. Genetic sources of high content and quality of protein for soft winter wheat breeding. Agrarian Bulletin of the Urals. 2016;6(148):51-55. [in Russian]Попереля Ф.О. Три основні генетичні системи якості зерна озимої м’якої пшениці. В кн. Реалізація потенційних можливостей сортів та гібридів Селекційно-генетичного інституту в умовах України. Зб. наук. праць СГІ. Одеса; 1996. С. 117-132 [in Ukrainian]Poperelya F.O. Three main genetic systems for grain quality in winter bread wheat (Tri osnovnі genetichnі sistemi yakostі zerna ozimoy myakoy pshenitsі). In: Implementing the potential of cultivars and hybrids from the Breeding and Genetics Institute in the environments of Ukraine. Collection of scientific papers of the BGI (Realіzatsіya potentsіynikh mozhlivostey sortіv ta gіbridіv Selektsіynogenetichnogo іnstitutu v umovakh Ukraini. Zb. nauk. prats SGІ) . Odesa; 1996. p.117-132. [in Ukrainian]Popineau Y., Deshayes G., Lefebvre J., Fido R., Tatham A.S., Shewry P.R. Prolamin aggregation, gluten viscoelasticity, and mixing properties of transgenic wheat lines expressing 1Ax and 1Dx high molecular weight glutenin subunit transgenes. Journal of Agricultural and Food Chemistry. 2001;49(1):395-401. DOI: 10.1021/jf001015jPopineau Y., Deshayes G., Lefebvre J., Fido R., Tatham A.S., Shewry P.R. Prolamin aggregation, gluten viscoelasticity, and mixing properties of transgenic wheat lines expressing 1Ax and 1Dx high molecular weight glutenin subunit transgenes. Journal of Agricultural and Food Chemistry. 2001;49(1):395-401. DOI: 10.1021/jf001015jRabinovich S.V., Panchenko I.A., Parchomenko R.G., Bondarenko V.N. High-molecular weight glutenin subunit composition of spring bread wheats grown in the Ukraine and the Russian Federation between 1995-97 and its connection with pedigrees. Annual Wheat Newsletter. 1998;44:236-251.Rabinovich S.V., Panchenko I.A., Parchomenko R.G., Bondarenko V.N. High-molecular weight glutenin subunit composition of spring bread wheats grown in the Ukraine and the Russian Federation between 1995-97 and its connection with pedigrees. Annual Wheat Newsletter. 1998;44:236-251.Ram S. High molecular weight glutenin subunit composition of Indian wheats and their relationships with dough strength. Journal of Plant Biochemistry and Biotechnology. 2003;12(2):151-155. DOI: 10.1007/bf03263177Ram S. High molecular weight glutenin subunit composition of Indian wheats and their relationships with dough strength. Journal of Plant Biochemistry and Biotechnology. 2003;12(2):151-155. DOI: 10.1007/bf03263177Ribeirо M., Carvalho C., Carnide V., Guedes-Pinto H., Igrejas G. Towards allelic diversity in the storage proteins of old and currently growing tetraploid and hexaploid wheats in Portugal. Genetic Resources and Crop Evolution. 2011;58(7):1051-1073.Ribeirо M., Carvalho C., Carnide V., Guedes-Pinto H., Igrejas G. Towards allelic diversity in the storage proteins of old and currently growing tetraploid and hexaploid wheats in Portugal. Genetic Resources and Crop Evolution. 2011;58(7):1051-1073.Rosa Filho O. Effect of the six glutenin loci on selected bread quality traits in wheat [dissertation]. Corvallis: Oregon State University; 1997.Rosa Filho O. Effect of the six glutenin loci on selected bread quality traits in wheat [dissertation]. Corvallis: Oregon State University; 1997.Sabelli P.A., Shewry P.R. Characterization and organization of gene families at the Gli-1 loci of bread and durum wheats by restriction fragment analysis. Theoretical and Applied Genetics. 1991;83(2):209-216. DOI: 10.1007/BF00226253Sabelli P.A., Shewry P.R. Characterization and organization of gene families at the Gli-1 loci of bread and durum wheats by restriction fragment analysis. Theoretical and Applied Genetics. 1991;83(2):209-216. DOI: 10.1007/BF00226253Salmanowicz B.P., Langner M. Identification and characterization of high-molecular-weight glutenin genes in Polish triticale cultivars by PCR-based DNA markers. Journal of Applied Genetics. 2007;48(4):347-357. DOI: 10.1007/BF03195231Salmanowicz B.P., Langner M. Identification and characterization of high-molecular-weight glutenin genes in Polish triticale cultivars by PCR-based DNA markers. Journal of Applied Genetics. 2007;48(4):347-357. DOI: 10.1007/BF03195231Seilmeier W., Belitz H.D., Wieser H. Separation and quantitative determination of high molecular weight subunits of glutenin from different wheat varieties and genetic variants of the variety Sicco. Zeitschrift für Lebensmittel-Untersuchung und Forschung. 1991;192:124-129. DOI: 10.1007/BF01202625Seilmeier W., Belitz H.D., Wieser H. Separation and quantitative determination of high molecular weight subunits of glutenin from different wheat varieties and genetic variants of the variety Sicco. Zeitschrift für Lebensmittel-Untersuchung und Forschung. 1991;192:124-129. DOI: 10.1007/BF01202625Семенов О.Г., Дивашук М.Г., Шангешапувако Х.Г., Плющиков В.Г., Хупацария Т.И., Введенский В.В. и др. Специфика сочетаний качественных и количественных характеристик клейковины у генотипов аллоцитоплазматической яровой пшеницы с аллелем Wx-B1a. Вестник Российского университета дружбы народов. Серия: Агрономия и животноводство. 2018;13(1):14-25.Semenov O.G., Divashur M.G., Shangeshapvako H.G., Plushikov V.G., Hupacaria T.I., Vvedensky V.V. et al. Specificity of combinations of qualitative and quantitative characteristics of glucovine in genotypes of allocytoplasmatic spruce wheat with allel of wild type Wx-B1a. RUDN Journal of Agronomy and Animal Industries. 2018;13(1):14-25. [in Russian]Sharma S., Ram S., Gupta R. Relationship of high and low molecular weight glutenins with chemical and rheological properties of wheat flour. Journal of Wheat Research. 2012;4(1):74-78.Sharma S., Ram S., Gupta R. Relationship of high and low molecular weight glutenins with chemical and rheological properties of wheat flour. Journal of Wheat Research. 2012;4(1):74-78.Shewry P.R., Halford N.G., Tatham A.S. High molecular weight subunits of wheat glutenin. Journal of cereal science. 1992;15(2):105-120. DOI: 10.1016/S0733-5210(09)80062-3Shewry P.R., Halford N.G., Tatham A.S. High molecular weight subunits of wheat glutenin. Journal of cereal science. 1992;15(2):105-120. DOI: 10.1016/S0733-5210(09)80062-3Шмалько Н.А. Сравнительный анализ белково-протеиназного комплекса хлебопекарной пшеничной и амарантовой муки. Техника и технология пищевых производств. 2011;2(21):84-88.Shmalko N.A. Comparative analysis of protein-proteinase complex of breadwheat and amaranth flours. Food Processing: Techniques and Technology. 2011;2(21):84-88. [in Russian]Сичкарь С.Н., Великожон Л.Г., Дубровная О.В. Молекулярно-генетический анализ и хлебопекарные качества гибридов Triticum spelta L. × Triticum aestivum L. Молекулярная и прикладная генетика. 2017;22:96-103.Sichkar S.N., Velykozhon L.G., Dubrovna O.V. Molecular genetic analysis and baking properties of hybrids Triticum spelta L. × Triticum aestivum L. Molekulyarnaya i prikladnaya genetika = Molecular and Applied Genetics. 2017;22:96-103. [in Russian]Singh N.K., Shepherd K.W. Linkage mapping of genes controlling endosperm storage proteins in wheat. Theoretical and Applied Genetics. 1988;75(4):628-641. DOI: 10.1007/BF00289132Singh N.K., Shepherd K.W. Linkage mapping of genes controlling endosperm storage proteins in wheat. Theoretical and Applied Genetics. 1988;75(4):628-641. DOI: 10.1007/BF00289132Smak C. New approach to determine the brownness of bread crust. Correlation between crust colour and protein content. Cereal Chemistry. 1972;49:554-560.Smak C. New approach to determine the brownness of bread crust. Correlation between crust colour and protein content. Cereal Chemistry. 1972;49:554-560.Tanksley S.D. Mapping polygenes. Annual review of genetics. 1993;27(1):205-233. DOI: 10.1146/annurev.ge.27.120193.001225Tanksley S.D. Mapping polygenes. Annual review of genetics. 1993;27(1):205-233. DOI: 10.1146/annurev.ge.27.120193.001225Вафин Р.Р., Абдулина И.Р., Ржанова И.В., Гараева А.Л., Тюлькин С.В., Асхадуллин Д.Ф. и др. Молекулярные подходы к идентификации генотипов Triticum aestivum L. по аллельным вариантам Waxy-генов и HMW субъединиц глютенинов. Современные проблемы науки и образования. 2015;(2).556-556).Vafin R.R., Abdulina I.R., Rzhanova I.V., Garaeva A.L., Tyulkin S.V., Askhadullin D.F. et al. Molecular approaches for identification of Triticum aestivum L. genotypes by allelic variants of Waxy-genes and HMW glutenin subunits. Modern Problems of Science and Education. 2015;(2):556-556. [in Russian]Vancini C., Torres G.A.M., de Miranda M.Z., Consoli L., Bonow S., Grando M.F. Impact of high-molecular-weight glutenin alleles on wheat technological quality. Pesquisa Agropecuária Brasileira. 2019;54:e00639. DOI: 10.1590/S1678-3921.pab2019.v54.00639Vancini C., Torres G.A.M., de Miranda M.Z., Consoli L., Bonow S., Grando M.F. Impact of high-molecular-weight glutenin alleles on wheat technological quality. Pesquisa Agropecuária Brasileira. 2019;54:e00639. DOI: 10.1590/S1678-3921.pab2019.v54.00639Vázquez J.F., Ruiz M., Nieto-Taladriz M.T., Albuquerque M.M. Effects on gluten strength of low Mr glutenin subunits coded by alleles at Glu-A3 and Glu-B3 loci in durum wheat. Journal of Cereal Science.1996;24(2):125-130. DOI: 10.1006/jcrs.1996.0045Vázquez J.F., Ruiz M., Nieto-Taladriz M.T., Albuquerque M.M. Effects on gluten strength of low Mr glutenin subunits coded by alleles at Glu-A3 and Glu-B3 loci in durum wheat. Journal of Cereal Science.1996;24(2):125-130. DOI: 10.1006/jcrs.1996.0045Вьюшков А.А., Мальчиков П.Н., Сюков В.В., Шевченко С.Н. Селекционно-генетическое улучшение яровой пшеницы. Самара: Самарский научный центр РАН; 2012.Vyushkov A.A., Malchikov P.N., Syukov V.V., Shevchenko S.N. Breeding and genetic improvement of spring wheat (Selektsionno-geneticheskoye uluchsheniye yarovoy pshenitsy). Samara: Samara Scientific Center of the RAS; 2012 [in Russian]Wang H.Y., Wei Y.M., Yan Z.H., Zheng Y.L. Genetic variation of gliadin and HMW glutenins subunits in durum wheat. Journal of Agricultural Biotechnology. 2006;14:721- 727.Wang H.Y., Wei Y.M., Yan Z.H., Zheng Y.L. Genetic variation of gliadin and HMW glutenins subunits in durum wheat. Journal of Agricultural Biotechnology. 2006;14:721-727.Пшеница, пшеница твердая, пшеница мягкая. Информационно-аналитический портал для крестьянских фермерских хозяйств). 2012. URL: http://fermer.zol.ru/a/15665/?module=a&param1=15665 [дата обращения: 15.04.2015].Wheat, durum wheat, bread wheat (Pshenitsa, pshenitsa tverdaya, pshenitsa myagkaya). Analytical information web portal for homestead farming enterprises (Informatsionno-analiticheskiy portal dlya krestyanskikh fermerskikh hozyajytv). 2012. [in Russian]Zhang W., Gianibelli M.C., Ma W., Rampling L., Gale K.R. Identi fication of SNPs and development of allele-specific PCR markers for gamma-gliadin alleles in Triticum aestivum. Theoretical and Applied Genetics. 2003;107(1):130-138. DOI: 10.1007/s00122-003-1223-2Zhang W., Gianibelli M.C., Ma W., Rampling L., Gale K.R. Identi fication of SNPs and development of allele-speci fic PCR markers for gamma-gliadin alleles in Triticum aestivum. Theoretical and Applied Genetics. 2003;107(1):130-138. DOI: 10.1007/s00122-003-1223-2Zhang X., Zhang B.Q., Wu H.Y., Lu C.B., Lü G.F., Liu D.T. et al. Effect of high-molecular-weight glutenin subunit dele tion on soft wheat quality properties and sugar-snap cookie quality estimated through near-isogenic lines. Journal of Integrative Agriculture. 2018;17(5):1066-1073. DOI: 10.1016/S2095-3119(17)61729-5Zhang X., Zhang B.Q., Wu H.Y., Lu C.B., Lü G.F., Liu D.T. et al. Effect of high-molecular-weight glutenin subunit dele tion on soft wheat quality properties and sugar-snap cookie quality estimated through near-isogenic lines. Journal of Integrative Agriculture. 2018;17(5):1066-1073. DOI: 10.1016/S2095-3119(17)61729-5Zhao H., Wang R., Guo A., Hu S., Sun G. Development of primers specific for LMW-GS genes located on chromosome 1D and molecular characterization of a gene from Glu-D3 complex locus in bread wheat. Hereditas. 2004;141(3):193-198. DOI: 10.1111/j.1601-5223.2004.01852.xZhao H., Wang R., Guo A., Hu S., Sun G. Development of primers specific for LMW-GS genes located on chromosome 1D and molecular characterization of a gene from Glu-D3 complex locus in bread wheat. Hereditas. 2004;141(3):193-198. DOI: 10.1111/j.1601-5223.2004.01852.xЗлацька А.В. Ідентифікація алеля Glu-B1al високомолекулярних глютенінів та його вплив на ознаки хлібопекар ської якості у пшениць, придатних до поширення в Україні. Физиология и биохимия культурных растений. 2010;42(4):315-321.Zlatska A.V. Identification of the Glu-B1al allele of highmolecular weight glutenins and it impact on characteristics related to bread-making quality in wheats permitted for realization in Ukraine. Fiziologiya i biokhimiya kulturnykh rasteniy = Crop Physiology and Biochemistry. 2010;42(4):315-321. [in Ukrainian]

The authors declare that there are no conflicts of interest present.