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<article article-type="research-article" dtd-version="1.3" xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" xml:lang="ru"><front><journal-meta><journal-id journal-id-type="publisher-id">vavilov</journal-id><journal-title-group><journal-title xml:lang="ru">Вавиловский журнал генетики и селекции</journal-title><trans-title-group xml:lang="en"><trans-title>Vavilov Journal of Genetics and Breeding</trans-title></trans-title-group></journal-title-group><issn pub-type="epub">2500-3259</issn><publisher><publisher-name>Institute of Cytology and Genetics of Siberian Branch of the RAS</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="doi">10.18699/vjgb-26-55</article-id><article-id custom-type="elpub" pub-id-type="custom">vavilov-5121</article-id><article-categories><subj-group subj-group-type="heading"><subject>Research Article</subject></subj-group><subj-group subj-group-type="section-heading" xml:lang="ru"><subject>БИОИНФОРМАТИКА И СИСТЕМНАЯ БИОЛОГИЯ</subject></subj-group><subj-group subj-group-type="section-heading" xml:lang="en"><subject>BIOINFORMATICS AND SYSTEMS BIOLOGY</subject></subj-group></article-categories><title-group><article-title>Генетический анализ архитектуры колоса пшеницы и его компьютерное фенотипирование у F2 гибридов тетраплоидных пшениц Triticum aethiopicum и T. carthlicum</article-title><trans-title-group xml:lang="en"><trans-title>Genetic analysis of wheat ear architecture in F2 hybrid of tetraploid wheats Triticum aethiopicum and T. carthlicum and its computer phenotyping</trans-title></trans-title-group></title-group><contrib-group><contrib contrib-type="author" corresp="yes"><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Кручинина</surname><given-names>Ю, В.</given-names></name><name name-style="western" xml:lang="en"><surname>Kruchinina</surname><given-names>Yu. V.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Новосибирск</p></bio><bio xml:lang="en"><p>Novosibirsk</p></bio><email xlink:type="simple">kruchinina2023@yandex.ru</email><xref ref-type="aff" rid="aff-1"/></contrib><contrib contrib-type="author" corresp="yes"><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Комышев</surname><given-names>Е. Г.</given-names></name><name name-style="western" xml:lang="en"><surname>Komyshev</surname><given-names>E. G.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Новосибирск</p></bio><bio xml:lang="en"><p>Novosibirsk</p></bio><xref ref-type="aff" rid="aff-2"/></contrib><contrib contrib-type="author" corresp="yes"><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Генаев</surname><given-names>М. А.</given-names></name><name name-style="western" xml:lang="en"><surname>Genaev</surname><given-names>M. A.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Новосибирск</p></bio><bio xml:lang="en"><p>Novosibirsk</p></bio><xref ref-type="aff" rid="aff-2"/></contrib><contrib contrib-type="author" corresp="yes"><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Коваль</surname><given-names>В. С.</given-names></name><name name-style="western" xml:lang="en"><surname>Koval</surname><given-names>V. S.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Новосибирск</p></bio><bio xml:lang="en"><p>Novosibirsk</p></bio><xref ref-type="aff" rid="aff-2"/></contrib><contrib contrib-type="author" corresp="yes"><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Афонников</surname><given-names>Д. А.</given-names></name><name name-style="western" xml:lang="en"><surname>Afonnikov</surname><given-names>D. A.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Новосибирск</p></bio><bio xml:lang="en"><p>Novosibirsk</p></bio><xref ref-type="aff" rid="aff-3"/></contrib><contrib contrib-type="author" corresp="yes"><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Гончаров</surname><given-names>Н. П.</given-names></name><name name-style="western" xml:lang="en"><surname>Goncharov</surname><given-names>N. P.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Новосибирск</p></bio><bio xml:lang="en"><p>Novosibirsk</p></bio><xref ref-type="aff" rid="aff-1"/></contrib></contrib-group><aff-alternatives id="aff-1"><aff xml:lang="ru">Федеральный исследовательский центр Институт цитологии и генетики Сибирского отделения Российской академии наук<country>Россия</country></aff><aff xml:lang="en">Institute of Cytology and Genetics of the Siberian Branch of the Russian Academy of Sciences<country>Russian Federation</country></aff></aff-alternatives><aff-alternatives id="aff-2"><aff xml:lang="ru">Федеральный исследовательский центр Институт цитологии и генетики Сибирского отделения Российской академии наук;&#13;
Курчатовский геномный центр ИЦиГ СО РАН<country>Россия</country></aff><aff xml:lang="en">Institute of Cytology and Genetics of the Siberian Branch of the Russian Academy of Sciences;&#13;
Kurchatov Genomic Center of ICG SB RAS<country>Russian Federation</country></aff></aff-alternatives><aff-alternatives id="aff-3"><aff xml:lang="ru">Федеральный исследовательский центр Институт цитологии и генетики Сибирского отделения Российской академии наук;&#13;
Курчатовский геномный центр ИЦиГ СО РАН;&#13;
Новосибирский национальный исследовательский государственный университет<country>Россия</country></aff><aff xml:lang="en">Institute of Cytology and Genetics of the Siberian Branch of the Russian Academy of Sciences;&#13;
Kurchatov Genomic Center of ICG SB RAS;&#13;
Novosibirsk State University<country>Russian Federation</country></aff></aff-alternatives><pub-date pub-type="collection"><year>2026</year></pub-date><pub-date pub-type="epub"><day>26</day><month>05</month><year>2026</year></pub-date><volume>30</volume><issue>3</issue><fpage>527</fpage><lpage>536</lpage><permissions><copyright-statement>Copyright &amp;#x00A9; Кручинина Ю.В., Комышев Е.Г., Генаев М.А., Коваль В.С., Афонников Д.А., Гончаров Н.П., 2026</copyright-statement><copyright-year>2026</copyright-year><copyright-holder xml:lang="ru">Кручинина Ю.В., Комышев Е.Г., Генаев М.А., Коваль В.С., Афонников Д.А., Гончаров Н.П.</copyright-holder><copyright-holder xml:lang="en">Kruchinina Y.V., Komyshev E.G., Genaev M.A., Koval V.S., Afonnikov D.A., Goncharov N.P.</copyright-holder><license license-type="creative-commons-attribution" xlink:href="https://creativecommons.org/licenses/by/4.0/" xlink:type="simple"><license-p>This work is licensed under a Creative Commons Attribution 4.0 License.</license-p></license></permissions><self-uri xlink:href="https://vavilov.elpub.ru/jour/article/view/5121">https://vavilov.elpub.ru/jour/article/view/5121</self-uri><abstract><p>Всестороннее описание фенотипов растений конкретных таксонов является важной задачей при описаниях родов, видов и построениях их естественных классификаций. Развитие современных технологий эффективного фенотипирования позволяет получать большое количество данных с количественным и/или качественным описанием различных признаков у растений, преимущественно на основе анализа их цифровых изображений. В исследовании проведено сравнение результатов оценки расщепления в F2 гибридов двух эндемичных тетраплоидных (2n = 4x = 28) видов пшениц – пшеницы эфиопской (Triticum aethiopicum Jakubz.) и пшеницы карталинской (T. carthlicum Nevski), полученных визуально и с помощью методов машинного обучения. В последнем случае для анализа расщеплений предложено использовать метод смеси гауссовых (нормальных) распределений в морфометрии растений для того, чтобы выделить группы, которые различаются по значениям признаков. Ввиду того что большинство таксономически значимых (видообразующих) признаков у пшениц контролируется олигогенно и имеет четкое фенотипическое проявление, гибридологический анализ был незаменимым и основным видом анализа для последующего детального фенотипирования колосьев видов пшениц с использованием методов машинного обучения. По ряду признаков оценки характера наследования таких признаков, полученные разными методами, совпадают. На основании результатов проведенного исследования мы можем утверждать, что признак «тетраостость» (наличие остей одновременно на цветковой и колосковой чешуях) видоспецифический (таксономически значим) для T. carthlicum и он может быть эффективно использован исследователями в таксономических целях как при проведении гибридологического анализа, так и в экспериментах с использованием машинного обучениям. Для T. aethiopicum таким видоспецифическим признаком является «характер (тип) остистости». Наша работа демонстрирует, что комбинация методов автоматического фенотипирования, модели смеси гауссовых распределений в принципе позволит проводить автоматический анализ выделения классов в F2 гибридов, различающихся по величинам фенотипических признаков. Это позволит, в свою очередь, выявлять наличие генов, ассоциированных с видоспецифическими признаками растений. Тем не менее требуется совершенствование применяемых алгоритмов искусственного интеллекта.</p></abstract><trans-abstract xml:lang="en"><p>A comprehensive description of plant phenotypes of certain taxa is an important task when describing genera and species, as well as when setting their natural taxonomies. The development of modern technologies of effective phenotyping makes it possible to obtain a large amount of data with a quantitative and/or qualitative description of various traits in plants, mainly based on the analysis of their digital images. The study compared the results of the F2 hybrids assessment – visually and using machine learning methods – of two endemic tetraploid (2n = 4x = 28) wheat species which are Ethiopian wheat (Triticum aethiopicum Jakubz.) and Kartalian or Dika wheat (T. carthlicum Nevski). In the latter case, it is proposed to use the method of a mixture of Gaussian (normal) distributions in plant morphometry in order to identify groups that differ in character values. Most taxonomically important (species-specific) traits are controlled oligogenically and have a clear phenotypic manifestation, so hybridological analysis was an indispensable and basic type of analysis for subsequent detailed phenotyping of wheat spikes using machine-learning methods. According to a number of criteria, the estimates of patterns of inheritance obtained by different methods coincide. Based on the conducted research, we can state that the trait “tetraaristatum” (the presence of awns on both flower and spike glumes) is species-specific (taxonomically important) for T. carthlicum and it can be effectively used for taxonomic purposes both in carrying out hybridological analysis and in experiments using machine learning. Such a species-specific character is the “character (type) of awnedness” for T. aethiopicum. Our study demonstrates that a combination of automatic phenotyping methods and a model of a mixture of Gaussian distributions can, in principle, lead to an automatic analysis of the allocation of classes in F2 hybrids. It allows, in turn, to detect the presence of genes associated with species-specific traits of wheat plants. Further, the improvement of the applied artificial intelligence (AI) algorithms is required.</p></trans-abstract><kwd-group xml:lang="ru"><kwd>тетраплоидные пшеницы</kwd><kwd>Triticum aethiopicum</kwd><kwd>T. carthlicum</kwd><kwd>генетический анализ</kwd><kwd>гибридизация</kwd><kwd>расщепление</kwd><kwd>цифровое фенотипирование</kwd><kwd>машинное обучение</kwd></kwd-group><kwd-group xml:lang="en"><kwd>tetraploid wheat</kwd><kwd>Triticum aethiopicum</kwd><kwd>T. carthlicum</kwd><kwd>genetic analysis</kwd><kwd>hybridization</kwd><kwd>cleavage</kwd><kwd>digital phenotyping</kwd><kwd>machine learning</kwd></kwd-group><funding-group xml:lang="en"><funding-statement>The preparation of data, development, and verification of the algorithm were supported by the Russian Science Foundation, project No. 23-14-00150</funding-statement></funding-group></article-meta></front><back><ref-list><title>References</title><ref id="cit1"><label>1</label><citation-alternatives><mixed-citation xml:lang="ru">Afonnikov D.A., Genaev M.A., Doroshkov A.V., Komyshev E.G., Pshenichnikova T.A. Methods of high-throughput plant phenotyping for large-scale breeding and genetic experiments. Russ J Genet. 2016;52(7):688-701. doi 10.1134/S1022795416070024</mixed-citation><mixed-citation xml:lang="en">Afonnikov D.A., Genaev M.A., Doroshkov A.V., Komyshev E.G., Pshenichnikova T.A. Methods of high-throughput plant phenotyping for large-scale breeding and genetic experiments. Russ J Genet. 2016;52(7):688-701. doi 10.1134/S1022795416070024</mixed-citation></citation-alternatives></ref><ref id="cit2"><label>2</label><citation-alternatives><mixed-citation xml:lang="ru">Artemenko N.V., Genaev M.A., Epifanov R.U., Komyshev E.G., Kruchinina Y.V., Koval V.S., Goncharov N.P., Afonnikov D.A. Imagebased classification of wheat spikes by glume pubescence using convolutional neural networks. Front Plant Sci. 2024;14:1336192. doi 10.3389/fpls.2023.1336192</mixed-citation><mixed-citation xml:lang="en">Artemenko N.V., Genaev M.A., Epifanov R.U., Komyshev E.G., Kruchinina Y.V., Koval V.S., Goncharov N.P., Afonnikov D.A. Imagebased classification of wheat spikes by glume pubescence using convolutional neural networks. Front Plant Sci. 2024;14:1336192. doi 10.3389/fpls.2023.1336192</mixed-citation></citation-alternatives></ref><ref id="cit3"><label>3</label><citation-alternatives><mixed-citation xml:lang="ru">Awada L., Phillips P.W.B., Bodan A.M. The evolution of plant phenomics: global insights, trends, and collaborations (2000-2021). Front Plant Sci. 2024;15:1410738. doi 10.3389/fpls.2024.1410738</mixed-citation><mixed-citation xml:lang="en">Awada L., Phillips P.W.B., Bodan A.M. The evolution of plant phenomics: global insights, trends, and collaborations (2000-2021). Front Plant Sci. 2024;15:1410738. doi 10.3389/fpls.2024.1410738</mixed-citation></citation-alternatives></ref><ref id="cit4"><label>4</label><citation-alternatives><mixed-citation xml:lang="ru">Bersimbaev R.I., Shulembaeva K.K. Cytogenetic studies of common wheat in Kazakhstan. Informatsionnyy Vestnik VOGiS. 2005; 9(3):317-323 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Bersimbaev R.I., Shulembaeva K.K. Cytogenetic studies of common wheat in Kazakhstan. Informatsionnyy Vestnik VOGiS. 2005; 9(3):317-323 (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit5"><label>5</label><citation-alternatives><mixed-citation xml:lang="ru">Bukasov S.M. Potato species system of section Tuberarium (Dum.) Buk. genus Solanim L. Proc Appl Bot Genet Breed. 1971;46(1):3-44 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Bukasov S.M. Potato species system of section Tuberarium (Dum.) Buk. genus Solanim L. Proc Appl Bot Genet Breed. 1971;46(1):3-44 (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit6"><label>6</label><citation-alternatives><mixed-citation xml:lang="ru">Chouhan S.S., Singh U.P., Sharma U., Jain S. Classification of different plant species using deep learning and machine learning algorithms. Wireless Pers Commun. 2024;136(4):2275-2298. doi 10.1007/s11277-024-11374-y</mixed-citation><mixed-citation xml:lang="en">Chouhan S.S., Singh U.P., Sharma U., Jain S. Classification of different plant species using deep learning and machine learning algorithms. Wireless Pers Commun. 2024;136(4):2275-2298. doi 10.1007/s11277-024-11374-y</mixed-citation></citation-alternatives></ref><ref id="cit7"><label>7</label><citation-alternatives><mixed-citation xml:lang="ru">Dobrovolskaya O.B., Dresvyannikova A.E., Badaeva E.D., Popova K.I., Travnickova M., Martinek P. The study of genetic factors that determine the awned glume trait in bread wheat. Vavilovskii Zhurnal Genetiki i Selektsii = Vavilov J Genet Breed. 2020;24(6):568-574. doi 10.18699/VJ20.650 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Dobrovolskaya O.B., Dresvyannikova A.E., Badaeva E.D., Popova K.I., Travnickova M., Martinek P. The study of genetic factors that determine the awned glume trait in bread wheat. Vavilovskii Zhurnal Genetiki i Selektsii = Vavilov J Genet Breed. 2020;24(6):568-574. doi 10.18699/VJ20.650 (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit8"><label>8</label><citation-alternatives><mixed-citation xml:lang="ru">Dolezel J., Greilhuber J., Suda J. Estimation of nuclear DNA content in plants using flow cytometry. Nat Protoc. 2007;2(9):2233-2244. doi 10.1038/nprot.2007.310</mixed-citation><mixed-citation xml:lang="en">Dolezel J., Greilhuber J., Suda J. Estimation of nuclear DNA content in plants using flow cytometry. Nat Protoc. 2007;2(9):2233-2244. doi 10.1038/nprot.2007.310</mixed-citation></citation-alternatives></ref><ref id="cit9"><label>9</label><citation-alternatives><mixed-citation xml:lang="ru">Dorofeev V.F. Cultural flora of the USSR. Vol. 1. Wheat. Leningrad, Kolos, 1979 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Dorofeev V.F. Cultural flora of the USSR. Vol. 1. Wheat. Leningrad, Kolos, 1979 (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit10"><label>10</label><citation-alternatives><mixed-citation xml:lang="ru">Eig A. Monographisch-kritische Ubersicht der Gattung Aegilops. Repertorium specierum novarum regni vegetabilis. Beihefte, Berlin, 1929</mixed-citation><mixed-citation xml:lang="en">Eig A. Monographisch-kritische Ubersicht der Gattung Aegilops. Repertorium specierum novarum regni vegetabilis. Beihefte, Berlin, 1929</mixed-citation></citation-alternatives></ref><ref id="cit11"><label>11</label><citation-alternatives><mixed-citation xml:lang="ru">Gandilian P.A. Spontaneous hybridization, mutations, and phylogeny issues in wheat. Genetika (Moscow). 1972;8(8):5-19 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Gandilian P.A. Spontaneous hybridization, mutations, and phylogeny issues in wheat. Genetika (Moscow). 1972;8(8):5-19 (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit12"><label>12</label><citation-alternatives><mixed-citation xml:lang="ru">Genaev M.A., Komyshev E.G., Fu Hao, Koval V.S., Goncharov N.P., Afonnikov D.A. SpikeDroidDB: an information system for annotation of morphometric characteristics of wheat spike. Vavilovskii Zhurnal Genetiki i Selektsii = Vavilov J Genet Breed. 2018; 22(1):132-140. doi 10.18699/VJ18.340 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Genaev M.A., Komyshev E.G., Fu Hao, Koval V.S., Goncharov N.P., Afonnikov D.A. SpikeDroidDB: an information system for annotation of morphometric characteristics of wheat spike. Vavilovskii Zhurnal Genetiki i Selektsii = Vavilov J Genet Breed. 2018; 22(1):132-140. doi 10.18699/VJ18.340 (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit13"><label>13</label><citation-alternatives><mixed-citation xml:lang="ru">Genaev M.A., Komyshev E.G., Kruchinina Y.V., Goncharov N.P.,Afonnikov D.A., Smirnov N.V. Morphometry of the wheat spike by analyzing 2D images. Agronomy. 2019;9(7):390. doi 10.3390/agronomy9070390</mixed-citation><mixed-citation xml:lang="en">Genaev M.A., Komyshev E.G., Kruchinina Y.V., Goncharov N.P.,Afonnikov D.A., Smirnov N.V. Morphometry of the wheat spike by analyzing 2D images. Agronomy. 2019;9(7):390. doi 10.3390/agronomy9070390</mixed-citation></citation-alternatives></ref><ref id="cit14"><label>14</label><citation-alternatives><mixed-citation xml:lang="ru">Golovnina K.A., Glushkov S.A., Blinov A.G., Mayorov V.I., Adkison L.R., Goncharov N.P. Molecular phylogeny of the genus Triticum L. Plant Syst Evol. 2007;264:195-216. doi 10.1007/s00606-006-0478-x</mixed-citation><mixed-citation xml:lang="en">Golovnina K.A., Glushkov S.A., Blinov A.G., Mayorov V.I., Adkison L.R., Goncharov N.P. Molecular phylogeny of the genus Triticum L. Plant Syst Evol. 2007;264:195-216. doi 10.1007/s00606-006-0478-x</mixed-citation></citation-alternatives></ref><ref id="cit15"><label>15</label><citation-alternatives><mixed-citation xml:lang="ru">Goncharov N.P., Mitina R.L., Anfilova N.A. Inheritance of awnlessness in tetraploid wheat species. Russ J Genet. 2003;39(4):463-466. doi 10.1023/A:1023326202320</mixed-citation><mixed-citation xml:lang="en">Goncharov N.P., Mitina R.L., Anfilova N.A. Inheritance of awnlessness in tetraploid wheat species. Russ J Genet. 2003;39(4):463-466. doi 10.1023/A:1023326202320</mixed-citation></citation-alternatives></ref><ref id="cit16"><label>16</label><citation-alternatives><mixed-citation xml:lang="ru">Goncharov N.P., Bannikova S.V., Kawahara T. Wheat artificial amphiploids involving Triticum timopheevii genome: their studies, preservation and reproduction. Genet Resour Crop Evol. 2007;54(7): 1507-1516. doi 10.1007/s10722-006-9141-1</mixed-citation><mixed-citation xml:lang="en">Goncharov N.P., Bannikova S.V., Kawahara T. Wheat artificial amphiploids involving Triticum timopheevii genome: their studies, preservation and reproduction. Genet Resour Crop Evol. 2007;54(7): 1507-1516. doi 10.1007/s10722-006-9141-1</mixed-citation></citation-alternatives></ref><ref id="cit17"><label>17</label><citation-alternatives><mixed-citation xml:lang="ru">Goncharov N.P. Manual book of common and durum wheat. Novosibirsk: RAS SB Publ. House, 2009 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Goncharov N.P. Manual book of common and durum wheat. Novosibirsk: RAS SB Publ. House, 2009 (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit18"><label>18</label><citation-alternatives><mixed-citation xml:lang="ru">Goncharov N.P. Genus Triticum L. taxonomy: The present and the future. Plant Syst Evol. 2011;295(1):1-11. doi 10.1007/s00606-011-0480-9</mixed-citation><mixed-citation xml:lang="en">Goncharov N.P. Genus Triticum L. taxonomy: The present and the future. Plant Syst Evol. 2011;295(1):1-11. doi 10.1007/s00606-011-0480-9</mixed-citation></citation-alternatives></ref><ref id="cit19"><label>19</label><citation-alternatives><mixed-citation xml:lang="ru">Goncharov N.P. Comparative genetics of wheat and its relatives. Novosibirsk: Geo Publ., 2012 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Goncharov N.P. Comparative genetics of wheat and its relatives. Novosibirsk: Geo Publ., 2012 (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit20"><label>20</label><citation-alternatives><mixed-citation xml:lang="ru">Goncharov N.P., Adonina I.G. A new hexaploid wheat species Triticum aminovii. Cereal Res Comm. 2025;53:2159-2165. doi 10.1007/s42976-025-00677-w</mixed-citation><mixed-citation xml:lang="en">Goncharov N.P., Adonina I.G. A new hexaploid wheat species Triticum aminovii. Cereal Res Comm. 2025;53:2159-2165. doi 10.1007/s42976-025-00677-w</mixed-citation></citation-alternatives></ref><ref id="cit21"><label>21</label><citation-alternatives><mixed-citation xml:lang="ru">Hammer K., Filatenko A.A., Pistrick K. Taxonomic remarks on Triticum L. and ×Triticosecale Wittm. Genet Resour Crop Evol. 2011; 58:3-10. doi 10.1007/s10722-010-9590-4</mixed-citation><mixed-citation xml:lang="en">Hammer K., Filatenko A.A., Pistrick K. Taxonomic remarks on Triticum L. and ×Triticosecale Wittm. Genet Resour Crop Evol. 2011; 58:3-10. doi 10.1007/s10722-010-9590-4</mixed-citation></citation-alternatives></ref><ref id="cit22"><label>22</label><citation-alternatives><mixed-citation xml:lang="ru">Hammer Ø., Harper D.A.T., Ryan P.D. PAST: Paleontological statistics software package for education and data analysis. Palaeontologia Electronica. 2001;4(1):1-9</mixed-citation><mixed-citation xml:lang="en">Hammer Ø., Harper D.A.T., Ryan P.D. PAST: Paleontological statistics software package for education and data analysis. Palaeontologia Electronica. 2001;4(1):1-9</mixed-citation></citation-alternatives></ref><ref id="cit23"><label>23</label><citation-alternatives><mixed-citation xml:lang="ru">Haque M.A., Takayama A., Watanabe N., Kuboyama T. Cytological and genetic mapping of the gene for four-awned phenotype in Triticum carthlicum Nevski. Genet Resour Crop Evol. 2011;58:1087-1093. doi 10.1007/s10722-010-9644-7</mixed-citation><mixed-citation xml:lang="en">Haque M.A., Takayama A., Watanabe N., Kuboyama T. Cytological and genetic mapping of the gene for four-awned phenotype in Triticum carthlicum Nevski. Genet Resour Crop Evol. 2011;58:1087-1093. doi 10.1007/s10722-010-9644-7</mixed-citation></citation-alternatives></ref><ref id="cit24"><label>24</label><citation-alternatives><mixed-citation xml:lang="ru">Hawkes J. A revision of the tuber-bearing Solanums. In: Record Scottish Plant Breeding Station. 1963;76-181</mixed-citation><mixed-citation xml:lang="en">Hawkes J. A revision of the tuber-bearing Solanums. In: Record Scottish Plant Breeding Station. 1963;76-181</mixed-citation></citation-alternatives></ref><ref id="cit25"><label>25</label><citation-alternatives><mixed-citation xml:lang="ru">Hodač L., Karbstein K., Tomasello S., Wäldchen J., Bradican J.P., Hörandl E. Geometric morphometric versus genomic patterns in a large polyploid plant species complex. Biology. 2023;12(3):418. doi 10.3390/biology12030418</mixed-citation><mixed-citation xml:lang="en">Hodač L., Karbstein K., Tomasello S., Wäldchen J., Bradican J.P., Hörandl E. Geometric morphometric versus genomic patterns in a large polyploid plant species complex. Biology. 2023;12(3):418. doi 10.3390/biology12030418</mixed-citation></citation-alternatives></ref><ref id="cit26"><label>26</label><citation-alternatives><mixed-citation xml:lang="ru">Kim B.H., Kwon H., Kim W. Deciphering individual triticale grain weight patterns: A gaussian mixture model approach. PloS One. 2024;19(11):e0313942. doi 10.1371/journal.pone.0313942</mixed-citation><mixed-citation xml:lang="en">Kim B.H., Kwon H., Kim W. Deciphering individual triticale grain weight patterns: A gaussian mixture model approach. PloS One. 2024;19(11):e0313942. doi 10.1371/journal.pone.0313942</mixed-citation></citation-alternatives></ref><ref id="cit27"><label>27</label><citation-alternatives><mixed-citation xml:lang="ru">Komyshev E.G., Genaev M.A., Kruchinina Yu.V., Koval V.S., Goncharov N.P., Afonnikov D.A. Evaluation of the spike diversity of seven hexaploid wheat species and an artificial amphidiploid using a quadrangle model obtained from 2D images. Plants. 2024;13(19): 2736. doi 10.3390/plants13192736</mixed-citation><mixed-citation xml:lang="en">Komyshev E.G., Genaev M.A., Kruchinina Yu.V., Koval V.S., Goncharov N.P., Afonnikov D.A. Evaluation of the spike diversity of seven hexaploid wheat species and an artificial amphidiploid using a quadrangle model obtained from 2D images. Plants. 2024;13(19): 2736. doi 10.3390/plants13192736</mixed-citation></citation-alternatives></ref><ref id="cit28"><label>28</label><citation-alternatives><mixed-citation xml:lang="ru">Kudryavtsev A.M., Popova T.A. Genetic linkage between gliadincoding genes and genes for color and pubescence of the ear in spring durum wheat (Triticum durum Desf.). Genetika (Moscow). 1994; 30(12):1587-1592 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Kudryavtsev A.M., Popova T.A. Genetic linkage between gliadincoding genes and genes for color and pubescence of the ear in spring durum wheat (Triticum durum Desf.). Genetika (Moscow). 1994; 30(12):1587-1592 (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit29"><label>29</label><citation-alternatives><mixed-citation xml:lang="ru">Lyapunova O.A. Intraspecific diversity of durum wheat (Triticum durum Desf.): a unified classification. Vavilovskii Zhurnal Genetiki i Selektsii = Vavilov J Genet Breed. 2021;25(3):260-268. doi 10.18699/VJ21.029</mixed-citation><mixed-citation xml:lang="en">Lyapunova O.A. Intraspecific diversity of durum wheat (Triticum durum Desf.): a unified classification. Vavilovskii Zhurnal Genetiki i Selektsii = Vavilov J Genet Breed. 2021;25(3):260-268. doi 10.18699/VJ21.029</mixed-citation></citation-alternatives></ref><ref id="cit30"><label>30</label><citation-alternatives><mixed-citation xml:lang="ru">Martinek P., Bednar J. Changes of spike morphology (multirow spike – MRS, long glumes – LG) in wheat (Triticum aestivum L.) and their importance for breeding. In: Proc. Intern. Conf. “Genetic collections, isogenic and alloplasmic lines”. Novosibirsk, 2001;192-194</mixed-citation><mixed-citation xml:lang="en">Martinek P., Bednar J. Changes of spike morphology (multirow spike – MRS, long glumes – LG) in wheat (Triticum aestivum L.) and their importance for breeding. In: Proc. Intern. Conf. “Genetic collections, isogenic and alloplasmic lines”. Novosibirsk, 2001;192-194</mixed-citation></citation-alternatives></ref><ref id="cit31"><label>31</label><citation-alternatives><mixed-citation xml:lang="ru">Merezhko A.F. The use of Mendelian principles in computer analysis of the inheritance of varying traits. In: Ecological genetics of cultivated plants. Krasnodar: All-Russian Research Institute of Rice Publ., 2005;107-117 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Merezhko A.F. The use of Mendelian principles in computer analysis of the inheritance of varying traits. In: Ecological genetics of cultivated plants. Krasnodar: All-Russian Research Institute of Rice Publ., 2005;107-117 (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit32"><label>32</label><citation-alternatives><mixed-citation xml:lang="ru">Migushova E.F., Zhukovsky P.M. Towards the knowledge of wheat T. ispahanicum Heslot. Proc Appl Bot Genet Breed. 1969;39(3): 71-90 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Migushova E.F., Zhukovsky P.M. Towards the knowledge of wheat T. ispahanicum Heslot. Proc Appl Bot Genet Breed. 1969;39(3): 71-90 (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit33"><label>33</label><citation-alternatives><mixed-citation xml:lang="ru">Mulugeta A.K., Sharma D.P., Mesfin A.H. Deep learning for medicinal plant species classification and recognition: a systematic review. Front Plant Sci. 2024;14:1286088. doi 10.3389/fpls.2023.1286088</mixed-citation><mixed-citation xml:lang="en">Mulugeta A.K., Sharma D.P., Mesfin A.H. Deep learning for medicinal plant species classification and recognition: a systematic review. Front Plant Sci. 2024;14:1286088. doi 10.3389/fpls.2023.1286088</mixed-citation></citation-alternatives></ref><ref id="cit34"><label>34</label><citation-alternatives><mixed-citation xml:lang="ru">Pronozin A.Yu., Paulish A.A., Zavarzin E.A., Prikhodko A.Yu., Prokhoshin N.M., Kruchinina Y.V., Goncharov N.P., Komyshev E.G., Genaev M.A. Automatic phenotyping of ear morphology of tetraand hexaploid wheat species by computer vision methods. Vavilovskii Zhurnal Genetiki i Selektsii = Vavilov J Genet Breed. 2021;25(1): 71-81. doi 10.18699/VJ21.009</mixed-citation><mixed-citation xml:lang="en">Pronozin A.Yu., Paulish A.A., Zavarzin E.A., Prikhodko A.Yu., Prokhoshin N.M., Kruchinina Y.V., Goncharov N.P., Komyshev E.G., Genaev M.A. Automatic phenotyping of ear morphology of tetraand hexaploid wheat species by computer vision methods. Vavilovskii Zhurnal Genetiki i Selektsii = Vavilov J Genet Breed. 2021;25(1): 71-81. doi 10.18699/VJ21.009</mixed-citation></citation-alternatives></ref><ref id="cit35"><label>35</label><citation-alternatives><mixed-citation xml:lang="ru">Ran Z., Xiao X., Zhou L., Yan C., Bai X., Ou J., Li Z. Phenotypic diversity analysis in the Sect. Tuberculate (Camellia L.) population, an endemic taxon in China. Plants. 2024;13(22):3210. doi 10.3390/plants13223210</mixed-citation><mixed-citation xml:lang="en">Ran Z., Xiao X., Zhou L., Yan C., Bai X., Ou J., Li Z. Phenotypic diversity analysis in the Sect. Tuberculate (Camellia L.) population, an endemic taxon in China. Plants. 2024;13(22):3210. doi 10.3390/plants13223210</mixed-citation></citation-alternatives></ref><ref id="cit36"><label>36</label><citation-alternatives><mixed-citation xml:lang="ru">Rechkin D.V. Analysis of quantitative traits segregation based on fundamental properties of the normal distribution. Pisma v Vavilovskii Zhurnal Genetiki i Selektsii = Lett Vavilov J Genet Breed. 2024; 10(4):199-203. doi 10.18699/letvjgb-2024-10-25 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Rechkin D.V. Analysis of quantitative traits segregation based on fundamental properties of the normal distribution. Pisma v Vavilovskii Zhurnal Genetiki i Selektsii = Lett Vavilov J Genet Breed. 2024; 10(4):199-203. doi 10.18699/letvjgb-2024-10-25 (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit37"><label>37</label><citation-alternatives><mixed-citation xml:lang="ru">Rozhkov R.V., Krivoruchenko R.V., Kovalenko I.V. Genetic control of the awned glume trait in wheat. Vestnik Khar’kovskogo Natsional’nogo Agrarnogo Universiteta = Herald of the Kharkiv National Agrarian University. 2014;2(32):70-76 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Rozhkov R.V., Krivoruchenko R.V., Kovalenko I.V. Genetic control of the awned glume trait in wheat. Vestnik Khar’kovskogo Natsional’nogo Agrarnogo Universiteta = Herald of the Kharkiv National Agrarian University. 2014;2(32):70-76 (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit38"><label>38</label><citation-alternatives><mixed-citation xml:lang="ru">Sobko T.A., Sozinov A.A. Genetic control of morphological features of the ear and the relationship of allelic variability of marker loci of chromosomes 1A and 1B of winter soft wheat. Tsitologiya i Genetika = Cytology and Genetics. 1993;27(5):15-22 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Sobko T.A., Sozinov A.A. Genetic control of morphological features of the ear and the relationship of allelic variability of marker loci of chromosomes 1A and 1B of winter soft wheat. Tsitologiya i Genetika = Cytology and Genetics. 1993;27(5):15-22 (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit39"><label>39</label><citation-alternatives><mixed-citation xml:lang="ru">Sourdille P., Cadalen T., Gay G., Bernard M. Molecular and physical mapping of genes affecting awning in wheat. Plant Breeding. 2002; 121(4):320-324. doi 10.1046/j.1439-0523.2002.728336.x</mixed-citation><mixed-citation xml:lang="en">Sourdille P., Cadalen T., Gay G., Bernard M. Molecular and physical mapping of genes affecting awning in wheat. Plant Breeding. 2002; 121(4):320-324. doi 10.1046/j.1439-0523.2002.728336.x</mixed-citation></citation-alternatives></ref><ref id="cit40"><label>40</label><citation-alternatives><mixed-citation xml:lang="ru">Tiburtini M., Scrucca L., Peruzzi L. Using Gaussian Mixture Models in plant morphometrics. Perspect Plant Ecol Evol Syst. 2025;69:125902. doi 10.1016/j.ppees.2025.125902</mixed-citation><mixed-citation xml:lang="en">Tiburtini M., Scrucca L., Peruzzi L. Using Gaussian Mixture Models in plant morphometrics. Perspect Plant Ecol Evol Syst. 2025;69:125902. doi 10.1016/j.ppees.2025.125902</mixed-citation></citation-alternatives></ref><ref id="cit41"><label>41</label><citation-alternatives><mixed-citation xml:lang="ru">Wang D., Yu K., Jin D., Sun L., Chu J., Wu W., Xin P., Gregová E., Li X., Sun J., Yang W., Zhan K., Zhang A., Liu D. Natural variations in the promoter of Awn Length Inhibitor 1 (ALI‐1) are associated with awn elongation and grain length in common wheat. Plant J. 2019;101(5):1075-1090. doi 10.1111/tpj.14575</mixed-citation><mixed-citation xml:lang="en">Wang D., Yu K., Jin D., Sun L., Chu J., Wu W., Xin P., Gregová E., Li X., Sun J., Yang W., Zhan K., Zhang A., Liu D. Natural variations in the promoter of Awn Length Inhibitor 1 (ALI‐1) are associated with awn elongation and grain length in common wheat. Plant J. 2019;101(5):1075-1090. doi 10.1111/tpj.14575</mixed-citation></citation-alternatives></ref><ref id="cit42"><label>42</label><citation-alternatives><mixed-citation xml:lang="ru">Wheat manual book / Eds V.F. Dorofeev, A.A. Filatenko, E.F. Migushova. Leningrad: VIR, 1980 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Wheat manual book / Eds V.F. Dorofeev, A.A. Filatenko, E.F. Migushova. Leningrad: VIR, 1980 (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit43"><label>43</label><citation-alternatives><mixed-citation xml:lang="ru">Zhukovsky P.M. A critical and systematic review of the species of the genus Aegilops. Proc Appl Bot Genet Breed. 1928;18(1):417-609 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Zhukovsky P.M. A critical and systematic review of the species of the genus Aegilops. Proc Appl Bot Genet Breed. 1928;18(1):417-609 (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit44"><label>44</label><citation-alternatives><mixed-citation xml:lang="ru">Zuev E.V., Brykova A.N., Kudryavtseva E.Yu. Results of analyzing the passport database ‘Spring bread wheat landraces in the VIR collection’. Proc Appl Bot Genet Breed. 2019;180(1):7-11. doi 10.30901/2227-8834-2019-1-7-11 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Zuev E.V., Brykova A.N., Kudryavtseva E.Yu. Results of analyzing the passport database ‘Spring bread wheat landraces in the VIR collection’. Proc Appl Bot Genet Breed. 2019;180(1):7-11. doi 10.30901/2227-8834-2019-1-7-11 (in Russian)</mixed-citation></citation-alternatives></ref></ref-list><fn-group><fn fn-type="conflict"><p>The authors declare that there are no conflicts of interest present.</p></fn></fn-group></back></article>
