References

vavilov

Вавиловский журнал генетики и селекции

Vavilov Journal of Genetics and Breeding

2500-3259

Institute of Cytology and Genetics of Siberian Branch of the RAS

10.18699/VJGB-22-64

vavilov-3473

Research Article

СЕЛЕКЦИЯ РАСТЕНИЙ НА ИММУНИТЕТ И ПРОДУКТИВНОСТЬ

PLANT BREEDING FOR IMMUNITY AND PERFORMANCE

Голозерный ячмень: систематика, селекция и перспективы использования

Naked barley: taxonomy, breeding, and prospects of utilization

https://orcid.org/0000-0001-5477-8684

Лукина

К. А.

Lukina

K. A.

Санкт-Петербург

St. Petersburg

k.lukina@vir.nw.ru

https://orcid.org/0000-0002-3990-6526

Ковалева

О. Н.

Kovaleva

O. N.

Санкт-Петербург

St. Petersburg

https://orcid.org/0000-0002-9250-7225

Лоскутов

И. Г.

Loskutov

I. G.

Санкт-Петербург

St. Petersburg

Федеральный исследовательский центр Всероссийский институт генетических ресурсов растений им. Н.И. Вавилова (ВИР)РоссияFederal Research Center the N.I. Vavilov All-Russian Institute of Plant Genetic Resources (VIR)Russian Federation

2022

09102022

266524536

2022

Лукина К.А., Ковалева О.Н., Лоскутов И.Г.

Lukina K.A., Kovaleva O.N., Loskutov I.G.

This work is licensed under a Creative Commons Attribution 4.0 License.

https://vavilov.elpub.ru/jour/article/view/3473

В обзоре освещены вопросы современного состояния систематики, происхождения и перспектив использования голозерного ячменя. Представлены положения систематики культурного ячменя Hordeum vulgare L., в которой выделяются группы многорядного (сonvar. coeleste (L.) A. Trof.) и двурядного (сonvar. nudum (L.) A. Trof.) голозерного ячменя. Группы включают разновидности согласно особенностям строения колоса, остей, цветковых и колосковых чешуй и цвету зерновок. Отражены вопросы очагов формообразования голозерного ячменя, по данным археологии, палеоэтноботаники, разнообразию форм и сортов, а также рассмотрены вопросы происхождения голозерного ячменя по результатам ДНК-маркирования. Приведены основные зоны возделывания и возможные причины такого преимущественного или исключительного распространения голозерных ячменей в районах высокогорья. Обсуждается вопрос наследования признака голозерности ячменя и механизмов его проявления в соответствии с новыми данными в области генетики. Представлены характеристики биохимического состава зерна по белку, некоторым заменимым и незаменимым аминокислотам, β-глюканам, витаминам, антиоксидантам. Показано, что голозерный ячмень – ценный источник уникальных комбинаций растворимых и нерастворимых пищевых волокон и полисахаридов. Подчеркнуто значение параметров голозерного ячменя, которые ограничивают широкое распространение этой культуры в мире, и выделены направления селекционной работы для устранения этих недостатков. Приведены примеры образцов голозерного ячменя, устойчивых к вредным организмам, которые являются перспективными источниками для повышения урожайности и качества зерна, а также сохранения экологической чистоты и безопасности продукции. Отражены этапы и направления селекционных работ с голозерным ячменем и показано значение мировой коллекции ВИР как основного источника генетического материала для развития селекции.

This review surveys the current state of taxonomy, origin, and utilization prospects for naked barley. The cultivated barley Hordeum vulgare L. incorporates the covered and naked barley groups. Naked barleys are divided into six-row naked barley (convar. сoeleste (L.) A. Trof.) and two-row naked barley (convar. nudum (L.) A. Trof.). The groups include botanical varieties differing in the structural features of spikes, awns, floret and spikelet glumes, and the color of kernels. The centers of morphogenesis for naked barley are scrutinized employing archeological and paleoethnobotanical data, and the diversity of its forms. Hypotheses on the centers of its origin are discussed using DNA marker data. The main areas of its cultivation are shown, along with possible reasons for such a predominating or exclusive distribution of naked barley in highland areas. Inheritance of nakedness and mechanisms of its manifestation are considered in the context of new data in genetics. The biochemical composition of barley grain in protein, some essential and nonessential amino acids, β-glucans, vitamins, and antioxidants is described. Naked barley is shown to be a valuable source of unique combinations of soluble and insoluble dietary fibers and polysaccharides. The parameters limiting wider distribution of naked barley over the world are emphasized, and breeding efforts that could mitigate them are proposed. Pathogen-resistant naked barley accessions are identified to serve as promising sources for increasing grain yield and quality. Main stages and trends of naked barley breeding are considered and the importance of the VIR global germplasm collection as the richest repository of genetic material for the development of breeding is shown.

голозерный ячменьсистематикапроисхождениегенетикакачество зернаустойчивость к болезнямурожайностьселекция

naked barleytaxonomyorigingeneticsgrain qualitydisease resistanceyieldbreeding

The work was financially supported by the project of the Ministry of Science and Higher Education of the Russian Federation National Network Collection of Plant Genetic Resources for Effective Scientific and Technological Development of the Russian Federation in the Field of Genetic Technologies under Agreement No. 075-15-2021-1050 dated September 28, 2021.

References1

Agarwal M., Shrivastava N., Padh H. Advances in molecular marker techniques and their applications in plant sciences. Plant Cell Rep. 2008;27(4):617-631. DOI 10.1007/s00299-008-0507-z.

Alpateva N.V., Abdullaev R.A., Anisimova I.N., Gubareva N.K., Kovaleva O.N., Radchenko E.E. Local barley accessions from Ethiopia resistant to powdery mildew. Trudy po Prikladnoy Botanike, Genetike i Selektsii = Proceedings on Applied Botany, Genetics, and Breeding. 2016;177(4):70-78. DOI 10.30901/2227-8834-2016-4-70-78. (in Russian)

Aniskov N.I., Popolzukhin P.V., Nikolayev P.N., Safonova I.V. Agrobiological value of Omskiy Golozerny 1 and Omskiy Golozerny 2 cultivars of spring barley. Sibirskiy Vestnik Selskokhozyaystvennoy Nauki = Siberian Herald of Agricultural Sciences. 2015;6:24-29. (in Russian)

Atanasova-Penichon V., Barreau C., Richard-Forget F. Antioxidant secondary metabolites in cereals: potential involvement in resistance to Fusarium and mycotoxin accumulation. Front. Microbiol. 2016;7:566. DOI 10.3389/fmicb.2016.00566.

Avdeichik O., Guretskaya B., Semenova T., Shishlov M., Shishlova A. Naked barley: creation, prospects and use. Nauka i Innovatsii = Science and Innovation. 2009;3:15-19. (in Russian)

Badr A., Muller K., Schafer-Pregl R., El Rabey H., Effgen S., Ibrahim H.H., Pozzi C., Rohde W., Salamini F. On the origin and domestication history of barley (Hordeum vulgare L.). Mol. Biol. Evol. 2000;17(4):499-510. DOI 10.1093/oxfordjournals.molbev. a026330.

Bakhteev F.Kh. Barley. Moscow; Leningrad: Sel’khozgiz Publ., 1955. (in Russian)

Bekhtol’d N.P., Orlova E.A. Injuriousness of the barley smut agent under conditions of the forest-steppe of the Ob region. Dostizheniya Nauki i Tekhniki APK = Achievements of Science and Technology of AIC. 2018;32(7):36-39. DOI 10.24411/0235-2451-2018-10708. (in Russian)

Bhatty R.S. Production of food malt from hull-less barley. Cereal Chem. 1996;73(1):75-80.

Blake T., Blake V., Bowman J., Abdel-Haleem H. Barley feed uses and quality improvement. In: Ullrich S.E. (Ed.). Barley: Production, Improvement and Uses. Oxford: Wiley-Blackwell, 2011; 522-531.

Bogdanova T.M., Terent’eva I.A., Khokhlova A.P., Ivanova N.N. Breeding value of naked barley. In: Abstracts from the international scientific and practical conference “Genetic resources of cultivated plants”, St.-Petersburg: VIR Publ., 2001;215-217. (in Russian)

Borisonik Z.B. Features of the technology of barley cultivation. In: Breeding of Barley and Oats. Moscow: Kolos Publ., 1971;218-230. (in Russian)

Bozbulut R., Sanlier N. Promising effects of β-glucans on glyceamic control in diabetes. Trends Food Sci. Technol. 2019;83(1): 159-166. DOI 10.1016/j.tifs.2018.11.018.

Buerstmayr H., Legzdina L., Steiner B., Lemmens M. Variation for resistance to Fusarium head blight in spring barley. Euphytica. 2004;137:279-290.

Characteristics of Plant Varieties Included for the First Time in 2020 in the State Register of Selection Achievements Authorized for Use for Production Purposes (official publication). Moscow: Rosinformagrotekh Publ., 2020. (in Russian)

Chen W., Turkington T.K., Levesque C.A., Bamforth J.M., Patrick S.K., Lewis C.T., Chapados J.T., Gaba D., Tittlemier S., Macleod A., Grafenhan T. Geography and agronomical practices drive diversification of the epiphytic mycoflora associated with barley and its malt end product in western Canada. Agric. Ecosyst. Environ. 2016;226:43-55. DOI 10.1016/j.agee.2016.03.030.

Chen X., Long H., Gao P., Deng G., Pan Z., Liang J., Tang Y., Tashi N., Yu M. Transcriptome assembly and analysis of Tibetan hulless barley (Hordeum vulgare L. var. nudum) developing grains, with emphasis on quality properties. PLoS One. 2014; 9(5):e98144. DOI 10.1371/journal.pone.0098144.

Dadashko V.V., Romashko A.K., Zinkevich L.V. Productive qualities of hens with the use of hulless barley variety in mixed fodders. Izvestiya Natsionalnoj Akademii Nauk Belarusi. Seriya Agrarnykh Nauk = Proceedings of the National Academy of Sciences of Belarus. Agrarian Series. 2010;2:94-97. (in Russian)

Doroshenko E.S., Doroshenko Ed.S. Immunological assessment of the hulled barley collection according to its resistance to leaf diseases in the southern part of the Rostov region. Zernovoe Khozyajstvo Rossii = Grain Economy of Russia. 2018;4(58):66-69. DOI 10.31367/2079-8725-2018-58-4-66-69. (in Russian)

Doroshenko Ed.S., Filippov E.G., Doncova A.A., Sidorenko V.S. Study of naked spring barley varieties under the conditions of the North Caucasus. Zernobobovye i Krupyanye Kul’tury = Legumes and Groat Crops. 2019;2(30):131-139. DOI 10.24411/2309-348X-2019-11103. (in Russian)

Fernandes G.C., Sonawane K.S., Arya S.S. Cereal based functional beverages: a review. J. Microbiol. Biotechnol. Food Sci. 2018/19; 8(3):914-919. DOI 10.15414/jmbfs.2018-19.8.3.914-919.

Gagkaeva T.Yu., Gavrilova O.P. Fusarial head blight of barley ear and grain. Trudy po Prikladnoy Botanike, Genetike i Selektsii = Proceedings on Applied Botany, Genetics, and Breeding. 2009; 165:39-44. (in Russian)

Gasparis S., Kała M., Przyborowski M., Łyżnik L.A., Orczyk W., Nadolska-Orczyk A. A simple and efficient CRISPR/Cas9 platform for induction of single and multiple, heritable mutations in barley (Hordeum vulgare L.). Plant Methods. 2018;14:111. DOI 10.1186/s13007-018-0382-8.

Gauthier L., Atanasova-Penichon V., Chéreau S., Richard-Forget F. Metabolomics to decipher the chemical defense of cereals against Fusarium graminearum and deoxynivalenol accumulation. Int. J. Mol. Sci. 2015;16(10):24839-24872. DOI 10.3390/ijms161024839.

Ge X., Jing L., Zhao K., Su C., Zhang B., Zhang Q., Han L., Yu X., Li W. The phenolic compounds profile, quantitative analysis and antioxidant activity of four naked barley grains with different color. Food Chem. 2021;335:127655. DOI 10.1016/j.foodchem. 2020.127655.

Gerasimova S.V., Hertig C., Korotkova A.M., Kolosovskaya E.V., Otto I., Hiekel S., Kochetov A.V., Khlestkina E.K., Kumlehn J. Conversion of hulled into naked barley by Cas endonucleasemediated knockout of the NUD gene. BMC Plant Biol. 2020; 20(Suppl. 1):255. DOI 10.1186/s12870-020-02454-9.

Gladkikh A.V., Rendov N.A., Nekrasova E.V., Mozyleva S.I. Influence of elements of cultivation technology on the yielding capacity of hulless barley grain. Vestnik Omskogo Gosudarstvennogo Agrarnogo Universiteta = Bulletin of the Omsk State Agrarian University. 2019;2(34):19-23. (in Russian)

Gryaznov A.A. Naked Barley under Conditions of Unstable Moisture. Kurtamysh: Kurtamyshskaya Tipografiya Publ., 2014. (in Russian)

Gryaznov A.A. Nudum 95: a high-protein variety of naked barley. APK Rossii = The Russian Agroindustrial Complex. 2016;75(1): 175-180. (in Russian)

Gryaznov A.A., Letyago J.A., Belkina R.I., Ponomareva E.I. Production of bread with the use of mixtures of wheat flour of the highest grade and wholemeal from pigmented barley variety Granal 32. Vestnik Voronezhskogo Gosudarstvennogo Universiteta Inzhenernykh Tekhnologij = Proceedings of the Voronezh State University of Engineering Technologies. 2019;81(1): 196-200. DOI 10.20914/2310-1202-2019-1-196-200. (in Russian)

Harlan J.R. The origin of barley. In: Barley: Botany, Culture, Winter Hardiness, Genetics, Utilization, Pests. Agricultural Handbook No. 338. Washington: Agricultural Research Service, U.S. Dept. of Agriculture, 1979;10-36.

Helback H. Domestication of food plants in the old world. Science. 1959;153:365-372.

Helbaek H. Commentary on the phylogenesis of Triticum and Hordeum. Econ. Bot. 1966;20:350-360.

Hernandez J., Meints B., Hayes P. Introgression breeding in barley: Perspectives and case studies. Front. Plant Sci. 2020;11:761. DOI 10.3389/fpls.2020.00761.

Hoad S.P., Brennan M., Wilson G.W., Cochrane P.M. Hull to caryopsis adhesion and grain skinning in malting barley: identification of key growth stages in the adhesion process. J. Cereal Sci. 2016;68:8-15. DOI 10.1016/j.jcs.2015.10.007.

Huth M., Dongowski G., Gebhart E., Flamme W. Functional properties of dietary fibre enriched exudates from barley. J. Cereal Sci. 2002;32:115-117.

Ingledew W.M., Jones A.M., Bhatty R.S., Rossnagel B.G. Fuel alcohol production from hull-less barley. Cereal Chem. 1995;72(2): 147-150.

Jaganathan D., Bohra A., Thudi M., Varshney R.K. Fine mapping and gene cloning in the post-NGS era: advances and prospects. Theor. Appl. Genet. 2020;133(5):1791-1810. DOI 10.1007/s00122-020-03560-w.

Khlestkina E.K. Molecular markers in genetic studies and breeding. Vavilovskii Zhurnal Genetiki i Selektsii = Vavilov Journal of Genetics and Breeding. 2013;17(4/2):1044-1054. (in Russian)

Khod’kov L.E. Naked and Awnless Barley. Leningrad, 1985. (in Russian)

Kikuchi S., Taketa S., Ichii M., Kawasaki S. Efficient fine mapping of naked caryopsis gene (nud ) by HEGS (High Efficiency Genome Scanning)/AFLP in barley. Theor. Appl. Genet. 2003; 108(1):73-78. DOI 10.1007/s00122-003-1413-y.

Kusch S., Panstruga R. mlo-based resistance: An apparently universal “weapon” to defeat powdery mildew disease. Mol. Plant Microbe Interact. 2017;30(3):179-189. DOI 10.1094/MPMI-12-16-0255-CR.

Kyrdoglo E.K., Polyshchuk S.S., Chervonis M.V. Methodology and results of breeding barley for food end-use. Trudy po Prikladnoy Botanike, Genetike i Selektsii = Proceedings on Applied Botany, Genetics, and Breeding. 2013;171:240-253. (in Russian)

Letyago Yu.A., Belkina R.I. The development of bread recipes with the addition of the flour from barley and triticale grain. Vestnik KrasGAU = Bulletin of the Krasnoyarsk State Agrarian University. 2019;12:176-182. DOI 10.36718/1819-4036-2019-12-176-182. (in Russian)

Loskutov I.G., Khlestkina E.K. Wheat, barley, and oat breeding for health benefit components in grain. Plants. 2021;10(1):86. DOI 10.3390/plants10010086.

Luk’yanova M.V., Trofimovskaya A.Ya., Gudkova G.N., Terent’eva I.A., Yarosh N.P. Flora of Cultivated Plants of the USSR. Vol. 2. Part 2. Barley. Leningrad: Agropromizdat Publ., 1990. (in Russian)

Madakemohekar A.H., Talekar N.S., Kamboj A.D., Thakur G. Scope of hulless barley (Hordeum vulgare L.) as a nutritious and medicinal food: a review. Acta Sci. Agric. 2018;2(12):11-13.

Malashkina M.S. Morphological parameters, biochemical and technological properties of naked barley for breeding in the Kemerovo region: Cand. Sci. (Agric.) Dissertation. St. Petersburg, 2008. (in Russian)

Martin C., Schöneberg T., Vogelgsang S., Morisoli R., Bertossa M., Mauch-Mani B., Mascher F. Resistance against Fusarium graminearum and the relationship to β-glucan content in barley grains. Eur. J. Plant Pathol. 2018;152:621-634. DOI 10.1007/ s10658-018-1506-8.

Martin C., Schöneberg T., Vogelgsang S., Vincenti J., Bertossa M., Mauch-Mani B., Mascher F. Factors of wheat grain resistance to fusarium head blight. Phytopathol. Mediterr. 2017;56:154-166. DOI 10.14601/Phytopathol.

Martínez-Subirà M., Romero M.P., Puig E., Macià A., Romagosa I., Moralejo M . Purple, high β-glucan, hulless barley as valuable ingredient for functional food. LWT Food Sci. Technol. 2020; 131(9):109582. DOI 10.1016/j.lwt.2020.109582.

Meints B., Vallejos C., Hayes P. Multi-use naked barley: A new frontier. J. Cereal Sci. 2021;102:103370. DOI 10.1016/j.jcs.2021.103370.

Meshkova L.V., Sabaeva O.B. Resistance to fungous diseases in barley in the Irtysh river area of the Omsk region. Trudy po Prikladnoy Botanike, Genetike i Selektsii = Proceedings on Applied Botany, Genetics, and Breeding. 2009;165:159-163. (in Russian)

Moreau R.A., Flores R.A., Hicks K.B. Composition of functional lipids in hulled and hulless barley in fractions obtained by scarification and in barley oil. Cereal Chem. 2007;84(1):1-5.

Moza J., Gujral H.S. Starch digestibility and bioactivity of high altitude hulless barley. Food Chem. 2016;194:561-568. DOI 10.1016/ j.foodchem.2015.07.149.

Nevo E. Origin, evolution, population genetics and resources for breeding of wild barley, Hordeum spontaneum, in the Fertile Crescent. In: Barley: Biochemistry, Molecular Biology and Biotechnology. Wallingford: C.A.B. International, 1992; 19-43.

Omarov D.S. Genetic resources of barley in Dagestan. In: Botanical and Genetic Resources of Dagestan Flora. Makhachkala: Dag. FAN SSSR Publ., 1981;104-115. (in Russian)

Orlov A.A. Barley. In: Flora of Cultivated Plants of the USSR. Vol. 2. Moscow; Leningrad: Gossel’khozizdat Publ., 1936;97-332. (in Russian)

Polisenska I., Jirsa O., Vaculova K., Pospichalova M., Wawroszova S., Frydrych J. Fusarium mycotoxins in two hulless oat and barley cultivars used for food purposes. Foods. 2020;9(8):1037. DOI 10.3390/foods9081037.

Polonskiy V.I., Surin N.A., Gerasimov S.A., Lipshin A.G., Sumina A.V., Zute S.A. Evaluation of barley genotypes for the content of β-glucans in grain and other valuable features in Eastern Siberia. Trudy po Prikladnoy Botanike, Genetike i Selektsii = Proceedings on Applied Botany, Genetics, and Breeding. 2021; 182(1):48-58. DOI 10.30901/2227-8834-2021-1-48-58. (in Russian)

Pomortsev A.A., Kalabushkin B.A., Blank M.L., Bakhronov A. Investigation of natural selection in artificial hybrid populations of spring barley. Russ. J. Genet. 1996;32(11):1333-1341.

Radchеnko E.E., Abdullaev R.A., Anisimova I.N. Genetic diversity of cereal crops for powdery mildew resistance. Ekologicheskaya Genetika = Ecological Genetics. 2020;18(1):59-78. DOI 10.17816/ecogen14530. (in Russian)

Rozanova I.V., Khlestkina E.K. NGS sequencing in barley breeding and genetic studies. Vavilovskii Zhurnal Genetiki i Selektsii = Vavilov Journal of Genetics and Breeding. 2020;24(4):348-355. DOI 10.18699/VJ20.627. (in Russian)

Shakib M.-C.R., Gabrial S.G.N. Post-prandial responses to different bread products based on wheat, barley and fenugreek or ginger or both in healthy volunteers and their effect on the glycemic index of such products. J. Am. Sci. 2010;6(10):89-96.

Shen Y., Zhang H., Cheng L., Wang L., Qian H., Qi X. In vitro and in vivo antioxidant activity of polyphenols extracted from black highland barley. Food Chem. 2016;194(1):1003-1012. DOI 10.1016/j.foodchem.2015.08.083.

Siranidou E., Kang Z., Buchenauer H. Studies on symptom development, phenolic compounds and morphological defence responses in wheat cultivars differing in resistance to fusarium head blight. J. Phytopathol. 2002;150:200-208.

State Register of Selection Achievements Authorized for Use for Production Purposes. Vol. 1. Plant Varieties (official publication). Moscow: Rosinformagrotekh Publ., 2021. (in Russian)

Surin N.A. Adaptive Potential of Varieties of Grain Crops Bred in Siberia and Ways of Improving it (Wheat, Barley, Oats). Novosibirsk, 2011. (in Russian)

Taketa S., Amano S., Tsujino Y., Sato T., Saisho D., Kakeda K., Nomura M., Suzuki T., Matsumoto T., Sato K., Kanamori H., Kawasaki S., Takeda K. Barley grain with adhering hulls is controlled by an ERF family transcription factor gene regulating a lipid biosynthesis pathway. Proc. Natl. Acad. Sci. 2008;105:4062-4067. DOI 10.1073/pnas.0711034105.

Taketa S., Kikuchi S., Awayama T., Yamamoto S., Ichii M., Kawasaki S. Monophyletic origin of naked barley inferred from molecular analyses of a marker closely linked to the naked caryopsis gene (nud ). Theor. Appl. Genet. 2004;108:1236-1242. DOI 10.1007/s00122-003-1560-1.

Tatarkina N.I. Use of nutritional substances of rations by young repair large white breed of pigs. Vestnik Kurganskoj GSKHA = Bulletin of the Kurgan State Agricultural Academy. 2019;2: 55-57. (in Russian)

Teimouri H., Zarghi H., Golian A. Evaluation of Hull-Less Barley with or without enzyme cocktail in the finisher diets of broiler chickens. J. Agric. Sci. Technol. 2018;20:469-483.

Tetyannikov N.V., Bome N.A. Sources of characters useful for breeding in hulless barley. Trudy po Prikladnoy Botanike, Genetike i Selektsii = Proceedings on Applied Botany, Genetics, and Breeding. 2020;181(3):49-55. DOI 10.30901/2227-8834-2020-3-49-55. (in Russian)

Toropova N.A., Sukhanova S.F. The use of hull-less barley for feeding geese broilers. Vestnik Kurganskoy GSKHA = Bulletin of the Kurgan State Agricultural Academy. 2013;1:36-39. (in Russian)

Trofimovskaya A.Ya. Barley: Evolution, Classification, and Breeding. Leningrad: Kolos Publ., 1972. (in Russian)

Tyaglyi S.V. Resistant to lodging in naked barley. In: Modern Approaches and Methods in Barley Breeding. Krasnodar: Prosveshchenie-Yug Publ., 2007;162-164. (in Russian)

Vavilov N.I. World Resources of Cereals and Flax. Moscow; Leningrad: AN SSSR Publ., 1957. (in Russian)

Vavilov N.I. Centers of Origin of Cultivated Plants. Selected Works. Moscow; Leningrad: Nauka Publ., 1965. (in Russian)

Warzecha T., Adamski T., Kaczmarek Z., Surma M., Goliński P., Perkowski J., Chełkowski J., Wiśniewska H., Krystkowiak K., Kuczyńska A. Susceptibility of hulled and hulless barley doubled haploids to Fusarium culmorum head blight. Cereal Res. Commun. 2010;38(2):220-232. DOI 10.1556/CRC.38.2010.2.8.

Wirkijowska A., Rzedzicki Z., Kasprzak M., Błaszczak W. Distribution of (1-3)(1-4)-β-d-glucans in kernels of selected cultivars of naked and hulled barley. J. Cereal Sci. 2012;56(2):496-503. DOI 10.1016/j.jcs.2012.05.002.

Xifeng R., Eviatar N., Dongfa S., Genlou S. Tibet as a potential domestication center of cultivated barley of China. PLoS One. 2013;8(5):e62700. DOI 10.1371/journal.pone.0062700.

Yu S., Long H., Deng G., Pan Z., Liang J., Zeng X., Tang Y., Tashi Y., Yu M. A single nucleotide polymorphism of nud converts the caryopsis type of barley (Hordeum vulgare L.). Plant Mol. Biol. Rep. 2016;34:242-248. DOI 10.1007/s11105-015-0911-9.

Zaushintsena A.V., Chernova E.V., Malashkina M.S. Comparative characteristics of two-row and six-row varieties of naked barley. Vestnik KrasGAU = Bulletin of the Krasnoyarsk State Agrarian University. 2007;6:72-75. (in Russian)

Zeng X., Guo Y., Xu Q., Mascher M., Guo G., Li S., Mao L., Liu Q., Xia Z., Zhou J., Yuan H., Tai S., Wang Y., Wei Z., Song L., Zha S., Li S., Tang Y., Bai L., Zhuang Z., He W., Zhao S., Fang X., Gao Q., Yin Y., Wang J., Yang H., Zhang J., Henry R.J., Stein N., Tashi N. Origin and evolution of qingke barley in Tibet. Nat. Commun. 2018; 9:5433. DOI 10.1038/s41467-018-07920-5.

Zheleznov A.V., Kukoeva T.V., Zheleznova N.B. Naked barley: origin, distribution and prospects of utilisation. Vavilovskii Zhurnal Genetiki i Selektsii = Vavilov Journal of Genetics and Breeding. 2013;17(2):286-297. (in Russian)

Zhichkina L.N., Stolpivskaya E.V. Resistance of spring barley varieties to loose smut. Izvestiya Samarskoj Gosudarstvennoj Sel’skokhozyaistvennoj Akademii = Bulletin of the Samara State Agricultural Academy. 2015;4:49-52. (in Russian)

Zveinek I.A., Kovaleva O.N. Genetic control of ultra-earliness in Chinese barley landraces. Trudy po Prikladnoy Botanike, Genetike i Selektsii = Proceedings on Applied Botany, Genetics, and Breeding. 2017;178(3):91-96. DOI 10.30901/2227-8834-2017-3-91-96. (in Russian)

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