<?xml version="1.0" encoding="UTF-8"?>
<!DOCTYPE article PUBLIC "-//NLM//DTD JATS (Z39.96) Journal Publishing DTD v1.3 20210610//EN" "JATS-journalpublishing1-3.dtd">
<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">vir-nw</journal-id><journal-title-group><journal-title xml:lang="ru">Труды по прикладной ботанике, генетике и селекции</journal-title><trans-title-group xml:lang="en"><trans-title>Proceedings on applied botany, genetics and breeding</trans-title></trans-title-group></journal-title-group><issn pub-type="ppub">2227-8834</issn><issn pub-type="epub">2619-0982</issn><publisher><publisher-name>N.I. Vavilov All-Russian Institute of Plant Genetic Resources</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="doi">10.30901/2227-8834-2022-4-172-180</article-id><article-id custom-type="elpub" pub-id-type="custom">vir-nw-1398</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>ИДЕНТИФИКАЦИЯ ГЕНЕТИЧЕСКОГО РАЗНООБРАЗИЯ КУЛЬТУРНЫХ РАСТЕНИЙ И ИХ ДИКИХ РОДИЧЕЙ ДЛЯ РЕШЕНИЯ ФУНДАМЕНТАЛЬНЫХ И ПРИКЛАДНЫХ ПРОБЛЕМ</subject></subj-group><subj-group subj-group-type="section-heading" xml:lang="en"><subject>IDENTIFICATION OF THE DIVERSITY OF CULTIVATED PLANTS AND THEIR WILD RELATIVES FOR SOLVING FUNDAMENTAL AND APPLIED PROBLEMS</subject></subj-group></article-categories><title-group><article-title>Изменчивость морфологических признаков и содержания ядерной ДНК гаплоидов и удвоенных гаплоидов в андрогенных каллусных линиях риса (Oryza sativa L.)</article-title><trans-title-group xml:lang="en"><trans-title>Variability of morphological features and nuclear DNA content in haploids and doubled haploids of androgenic callus lines of rice (Oryza sativa L.)</trans-title></trans-title-group></title-group><contrib-group><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0001-7042-8641</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Илюшко</surname><given-names>М. В.</given-names></name><name name-style="western" xml:lang="en"><surname>Ilyushko</surname><given-names>M. V.</given-names></name></name-alternatives><bio xml:lang="ru"><p>кандидат биологических наук, ведущий научный сотрудник, </p><p>692539 Уссурийск, пос. Тимирязевский, ул. Воложенина, 30</p></bio><bio xml:lang="en"><p>Cand. Sci. (Biology), Leading Researcher,</p><p>30 Volozhenina St., Timiryazevsky Settlem., Ussuriysk 692539</p></bio><email xlink:type="simple">ilyushkoiris@mail.ru</email><xref ref-type="aff" rid="aff-1"/></contrib><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-4884-0768</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Скапцов</surname><given-names>М. В.</given-names></name><name name-style="western" xml:lang="en"><surname>Skaptsov</surname><given-names>M. V.</given-names></name></name-alternatives><bio xml:lang="ru"><p>кандидат биологических наук, старший научный сотрудник, </p><p>656049 Барнаул, пр. Ленина, 61</p></bio><bio xml:lang="en"><p>Cand. Sci. (Biology), Senior Researcher,</p><p>61 Lenina Ave., Barnaul 656049</p></bio><email xlink:type="simple">mr.skaprsov@mail.ru</email><xref ref-type="aff" rid="aff-2"/></contrib><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-7426-8523</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Ромашова</surname><given-names>М. В.</given-names></name><name name-style="western" xml:lang="en"><surname>Romashova</surname><given-names>M. V.</given-names></name></name-alternatives><bio xml:lang="ru"><p>кандидат сельскохозяйственных наук, старший научный сотрудник, </p><p>692539 Уссурийск, пос. Тимирязевский, ул. Воложенина, 30</p></bio><bio xml:lang="en"><p>Cand. Sci. (Agriculture), Senior Researcher,</p><p>30 Volozhenina St., Timiryazevsky Settlem., Ussuriysk 692539</p></bio><email xlink:type="simple">romashova_1969@mail.ru</email><xref ref-type="aff" rid="aff-1"/></contrib></contrib-group><aff-alternatives id="aff-1"><aff xml:lang="ru"><institution>Федеральный научный центр агробиотехнологий Дальнего Востока им. А.К. Чайки</institution><country>Россия</country></aff><aff xml:lang="en"><institution>Federal Scientific Center of Agricultural Biotechnology of the Far East named after A.K. Chaika</institution><country>Russian Federation</country></aff></aff-alternatives><aff-alternatives id="aff-2"><aff xml:lang="ru"><institution>Алтайский государственный университет, Южно-Сибирский ботанический сад</institution><country>Россия</country></aff><aff xml:lang="en"><institution>Altai State University, South-Siberian Botanical Garden</institution><country>Russian Federation</country></aff></aff-alternatives><pub-date pub-type="collection"><year>2022</year></pub-date><pub-date pub-type="epub"><day>20</day><month>12</month><year>2022</year></pub-date><volume>183</volume><issue>4</issue><fpage>172</fpage><lpage>180</lpage><permissions><copyright-statement>Copyright &amp;#x00A9; Илюшко М.В., Скапцов М.В., Ромашова М.В., 2022</copyright-statement><copyright-year>2022</copyright-year><copyright-holder xml:lang="ru">Илюшко М.В., Скапцов М.В., Ромашова М.В.</copyright-holder><copyright-holder xml:lang="en">Ilyushko M.V., Skaptsov M.V., Romashova M.V.</copyright-holder><license xml:lang="ru" license-type="creative-commons-attribution" xlink:href="https://creativecommons.org/licenses/by/4.0/" xlink:type="simple"><license-p>Данная работа распространяется под лицензией Creative Commons Attribution 4.0.</license-p></license><license xml:lang="en" 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://elpub.vir.nw.ru/jour/article/view/1398">https://elpub.vir.nw.ru/jour/article/view/1398</self-uri><abstract><p>Работа актуальна для понимания эволюционных процессов у видов растений. Исследование проведено на гибридах F1 риса (Oryza sativa L.). Получено 12 каллусных линий с множественной регенерацией (более 60 растений на линии) гаплоидов и удвоенных гаплоидов. Регенерантные растения одной каллусной линии, полученной из одного пыльника риса, разделяли на две–четыре группы в зависимости от объема выборки в порядке их дифференциации на каллусе и высадки на среду укоренения. В первую группу входили растения с порядковым номером 1–30, во вторую – 31–60, в третью – 61–90, в четвертую – 91–120. Измеряли биометрические показатели и содержание ядерной ДНК. Между группами гаплоидов и удвоенных гаплоидов внутри каллусной линии выявлены различия по одному, двум или трем морфологическим признакам (p &lt; 0,05). Внутрикаллусная изменчивость признаков гаплоидов чаще сопровождалась уменьшением значений морфологических признаков с увеличением порядкового номера. На пяти каллусных линиях не обнаружено изменчивости, то есть такое явление не является правилом. Среднее содержание ядерной ДНК удвоенных гаплоидов в четырех группах одной каллусной линии составило 1,03–1,09 пг, у гаплоидов – 0,53–0,58 пг. Внутрикаллусная изменчивость содержания ядерной ДНК выявлена между группами у гаплоидов на одной линии и среди удвоенных гаплоидов одной линии. Обнаружены достоверные отличия гаплоидов одной каллусной линии от трех других каллусных линий гибрида Садко × Кубояр по содержанию ядерной ДНК в сторону увеличения (p &lt; 0,0015). Рассматривается теоретическая возможность появления внутривидовой изменчивости среди растений с небольшим числом хромосом. Предлагается схема геномных преобразований у таких видов: исходное растение (2n) → анеугаплоидные растения (n + 1) → мегаспорогенез и микроспорогенез по типу 0-n, формирование фертильной пыльцы (n + 1) → диплоидное растение (2n + 2). Анеугаплоидная эволюция объясняет внутривидовую изменчивость чисел хромосом среди видов растений с низкой плоидностью. </p></abstract><trans-abstract xml:lang="en"><p>The work is relevant for understanding evolutionary processes in plant species. Twelve callus lines with multiple regeneration of haploids and doubled haploids were obtained in F1 hybrids of Oryza sativa L. through in vitro androgenesis. Intracallus variability of the morphological features of haploids was often accompanied by a decrease in the values of morphological features with an increase in the serial number (p &lt; 0.05). The number of panicles on a plant and the number of flowers on a panicle on two callus lines in the second or third group were increased. No variability was detected in five callus lines, i.e., such a phenomenon was not a rule. The nuclear DNA content of doubled haploids in four groups of the same callus line was 1.03– 1.09 pg, and for haploids it was 0.53–0.58 pg. Intracallus variability of nuclear DNA content was detected between groups of haploids of the same line and among doubled haploids of the same line. Significant differences were found between the haploids of one callus line and the three other callus lines of the Sadko × Kuboyar hybrid towards an increase of nuclear DNA content (p &lt; 0.0015). The theoretical possibility of the appearance of intraspecific variability among plants with a small number of chromosomes is considered. A scheme of genomic reorganization is proposed for such species: initial plant (2n) → aneuhaploid plants (n + 1) → megasporogenesis and microsporogenesis of the 0-n type, formation of fertile pollen (n + 1) → diploid plant (2n + 2). Aneuhaploid evolution explains the intraspecific variability of chromosome numbers among plant species with low ploidy. Aneuploid technologies can help in the artificial formation of new polyploid crops, and rice is given a primary role. </p></trans-abstract><kwd-group xml:lang="ru"><kwd>андрогенез  in vitro</kwd><kwd>проточная цитометрия</kwd><kwd>внутрикаллусная изменчивость гаплоидов и удвоенных гаплоидов</kwd><kwd>анеугаплоидная эволюция растений</kwd></kwd-group><kwd-group xml:lang="en"><kwd>in vitro androgenesis</kwd><kwd>flow cytometry</kwd><kwd>intracallus variability</kwd><kwd>aneuhaploid plant evolution</kwd></kwd-group><funding-group><funding-statement xml:lang="en">The work was carried out within the theme “To develop new genotypes of agricultural crops with high productivity, resistant to abiotic and biotic environmental factors” (FNGW-2022-0008), registration number NIOKTR 122022600066-1. The authors express their gratitude to P. I. Kostylev (Donskoy Agricultural Research Center, Russia) for the provided seeds of the Sadko × Kuboyar hybrid and S. S. Guchenko (Federal Scientific Center of Agricultural Biotechnology of the Far East named after A. K. Chaika, Russia) for theDubrava × Atlant hybrid, S. O. Baturin (Institute of Cytology and Genetics, Russia) for the consultation on terminology in the plant reproduction field.</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">Агапова Н.Д., Архарова К.Б., Вахтина Л.И., Земскова Е.А., Тарвис Л.В. Числа хромосом цветковых растений флоры СССР: Семейства Aceraceae–Menyanthacea / под ред. А.Л. Тахтаджяна. Ленинград: Наука; 1990.</mixed-citation><mixed-citation xml:lang="en">Agapova N.D., Arkharova K.B., Vakhtina L.I., Zemskova E.A., Tarvis L.V. Numeri chromosomatum Magnoliophytorum florae URSS: Aceraceae–Menyanthacea. A.L. Takhtajan (ed.). Leningrad: Nauka; 1990. [in Russian].</mixed-citation></citation-alternatives></ref><ref id="cit2"><label>2</label><citation-alternatives><mixed-citation xml:lang="ru">Агапова Н.Д., Архарова К.Б., Вахтина Л.И., Земскова Е.А., Тарвис Л.В., Сафонова И.Н. Числа хромосом цветковых растений флоры СССР: Семейства Moraceae–Zygophyllaceae / под ред. А.Л. Тахтаджяна. Санкт-Петербург: Наука; 1993.</mixed-citation><mixed-citation xml:lang="en">Agapova N.D., Arkharova K.B., Vakhtina L.I., Zemskova E.A., Tarvis L.V., Saphonova I.N. Numeri chromosomatum Magnoliophytorum florae URSS: Moraceae–Zygophyllacea. A.L. Takhtajan (ed.). St. Petersburg: Nauka; 1993. [in Russian].</mixed-citation></citation-alternatives></ref><ref id="cit3"><label>3</label><citation-alternatives><mixed-citation xml:lang="ru">Bai C., Alverson W.S., Follansbee A., Waller D.M. New reports of nuclear DNA content for 407 vascular plant taxa from the United States. Annals of Botany. 2012;110(8):1623-1629. DOI: 10.1093/aob/mcs222</mixed-citation><mixed-citation xml:lang="en">Bai C., Alverson W.S., Follansbee A., Waller D.M. New reports of nuclear DNA content for 407 vascular plant taxa from the United States. Annals of Botany. 2012;110(8):1623-1629. DOI: 10.1093/aob/mcs222</mixed-citation></citation-alternatives></ref><ref id="cit4"><label>4</label><citation-alternatives><mixed-citation xml:lang="ru">Barow M., Jovtchev G. Endopolyploidy in plants and its analysis by flow cytometry. In: J. Doležel, J. Greilhuber, J. Suda (eds). Flow Cytometry with Plant Cells: Analysis of Genes, Chromosomes and Genomes. Weinhaim: Wiley-VCH; 2007. p.349-372. DOI: 10.1002/9783527610921.ch15</mixed-citation><mixed-citation xml:lang="en">Barow M., Jovtchev G. Endopolyploidy in plants and its analysis by flow cytometry. In: J. Doležel, J. Greilhuber, J. Suda (eds). Flow Cytometry with Plant Cells: Analysis of Genes, Chromosomes and Genomes. Weinhaim: Wiley-VCH; 2007. p.349-372. DOI: 10.1002/9783527610921.ch15</mixed-citation></citation-alternatives></ref><ref id="cit5"><label>5</label><citation-alternatives><mixed-citation xml:lang="ru">Bennett M.D., Smith J.B. Nuclear DNA amounts in angiosperms. Philosophical Transactions of the Royal Society B: Biological Sciences. 1991;334(1271):309-345. DOI: 10.1098/rstb.1991.0120</mixed-citation><mixed-citation xml:lang="en">Bennett M.D., Smith J.B. Nuclear DNA amounts in angiosperms. Philosophical Transactions of the Royal Society B: Biological Sciences. 1991;334(1271):309-345. DOI: 10.1098/rstb.1991.0120</mixed-citation></citation-alternatives></ref><ref id="cit6"><label>6</label><citation-alternatives><mixed-citation xml:lang="ru">Cistué L., Soriano M., Castillo A.M., Vallés M.P., Sanz J.M., Echávarri B. Production of doubled haploids in durum wheat (Triticum turgidum L.) through isolated microspore culture. Plant Cell Reports. 2006;25(4):257-264. DOI: 10.1007/s00299-005-0047-8</mixed-citation><mixed-citation xml:lang="en">Cistué L., Soriano M., Castillo A.M., Vallés M.P., Sanz J.M., Echávarri B. Production of doubled haploids in durum wheat (Triticum turgidum L.) through isolated microspore culture. Plant Cell Reports. 2006;25(4):257-264. DOI: 10.1007/s00299-005-0047-8</mixed-citation></citation-alternatives></ref><ref id="cit7"><label>7</label><citation-alternatives><mixed-citation xml:lang="ru">De Wet J.M.J. Reversible tetraploidy as an evolutionary mechanism. Evolution. 1971;25:545-548. DOI: 10.1111/j.1558-5646.1971.tb01914.x</mixed-citation><mixed-citation xml:lang="en">De Wet J.M.J. Reversible tetraploidy as an evolutionary mechanism. Evolution. 1971;25:545-548. DOI: 10.1111/j.1558- 5646.1971.tb01914.x</mixed-citation></citation-alternatives></ref><ref id="cit8"><label>8</label><citation-alternatives><mixed-citation xml:lang="ru">Grant V. Plant speciation. New York, NY: Columbia University Press; 1981. DOI: 10.7312/gran92318</mixed-citation><mixed-citation xml:lang="en">Grant V. Plant speciation. New York, NY: Columbia University Press; 1981. DOI: 10.7312/gran92318</mixed-citation></citation-alternatives></ref><ref id="cit9"><label>9</label><citation-alternatives><mixed-citation xml:lang="ru">He Y., Wei Q., Ge J., Jiang A., Gan L., Song Z. et al. Genome duplication effects on pollen development and the interrelated physiological substances in tetraploid rice with polyploidy meiosis stability. Planta. 2010;232(5):1219-1228. DOI: 10.1007/s00425-010-1249-z</mixed-citation><mixed-citation xml:lang="en">He Y., Wei Q., Ge J., Jiang A., Gan L., Song Z. et al. Genome duplication effects on pollen development and the interrelated physiological substances in tetraploid rice with polyploidy meiosis stability. Planta. 2010;232(5):1219-1228. DOI: 10.1007/s00425-010-1249-z</mixed-citation></citation-alternatives></ref><ref id="cit10"><label>10</label><citation-alternatives><mixed-citation xml:lang="ru">Ilyushko M.V., Romashova M.V. Formation of rice tetraploids in in vitro androgenesis. Russian Agricultural Sciences. 2020;46(4):332-336. DOI: 10.3103/S1068367420040084</mixed-citation><mixed-citation xml:lang="en">Ilyushko M.V., Romashova M.V. Formation of rice tetraploids in in vitro androgenesis. Russian Agricultural Sciences. 2020;46(4):332-336. DOI: 10.3103/S1068367420040084</mixed-citation></citation-alternatives></ref><ref id="cit11"><label>11</label><citation-alternatives><mixed-citation xml:lang="ru">Ilyushko M.V., Romashova M.V. Variability of rice haploids obtained from in vitro anther culture. Russian Agricultural Sciences. 2019;45(3):243-246. DOI: 10.3103/S1068367419030108</mixed-citation><mixed-citation xml:lang="en">Ilyushko M.V., Romashova M.V. Variability of rice haploids obtained from in vitro anther culture. Russian Agricultural Sciences. 2019;45(3):243-246. DOI: 10.3103/S1068367419030108</mixed-citation></citation-alternatives></ref><ref id="cit12"><label>12</label><citation-alternatives><mixed-citation xml:lang="ru">Ilyushko M.V., Romashova M.V., Zhang J.M., Deng L.W., Liu D.J., Zhang R., Guchenko S.S. Intra-callus variability of rice doubled haploids generated through in vitro androgenesis. Agricultural Biology. 2020;55(3):533-543. DOI: 10.15389/agrobiology.2020.3.533eng</mixed-citation><mixed-citation xml:lang="en">Ilyushko M.V., Romashova M.V., Zhang J.M., Deng L.W., Liu D.J., Zhang R., Guchenko S.S. Intra-callus variability of rice doubled haploids generated through in vitro androgenesis. Agricultural Biology. 2020;55(3):533-543. DOI: 10.15389/agrobiology.2020.3.533eng</mixed-citation></citation-alternatives></ref><ref id="cit13"><label>13</label><citation-alternatives><mixed-citation xml:lang="ru">Ilyushko M.V., Skaptsov M.V., Romashova M.V. Nuclear DNA content in rice (Oryza sativa L.) regenerants derived from anther culture in vitro. Agricultural Biology. 2018;53(3):531-538. DOI: 10.15389/agrobiology.2018.3.531eng</mixed-citation><mixed-citation xml:lang="en">Ilyushko M.V., Skaptsov M.V., Romashova M.V. Nuclear DNA content in rice (Oryza sativa L.) regenerants derived from anther culture in vitro. Agricultural Biology. 2018;53(3):531-538. DOI: 10.15389/agrobiology.2018.3.531eng</mixed-citation></citation-alternatives></ref><ref id="cit14"><label>14</label><citation-alternatives><mixed-citation xml:lang="ru">Jauhar P.P., Xu S.S., Baenziger P.S. Haploidy in cultivated wheats: Induction and utility in basic and applied research. Crop Science. 2009;49(3):737-755. DOI: 10.2135/cropsci2008.08.0462</mixed-citation><mixed-citation xml:lang="en">Jauhar P.P., Xu S.S., Baenziger P.S. Haploidy in cultivated wheats: Induction and utility in basic and applied research. Crop Science. 2009;49(3):737-755. DOI: 10.2135/cropsci2008.08.0462</mixed-citation></citation-alternatives></ref><ref id="cit15"><label>15</label><citation-alternatives><mixed-citation xml:lang="ru">Kasha K.J., Hu T.C., Oro R., Simion E., Shim Y.S. Nuclear fusion leads to chromosome doubling during mannitol pretreatment of barley (Hordeum vulgare L.) microspores. Journal of Experimental Botany. 2001;52(359):1227-1238. DOI: 10.1093/jxb/52.359.1227</mixed-citation><mixed-citation xml:lang="en">Kasha K.J., Hu T.C., Oro R., Simion E., Shim Y.S. Nuclear fusion leads to chromosome doubling during mannitol pretreatment of barley (Hordeum vulgare L.) microspores. Journal of Experimental Botany. 2001;52(359):1227-1238. DOI: 10.1093/jxb/52.359.1227</mixed-citation></citation-alternatives></ref><ref id="cit16"><label>16</label><citation-alternatives><mixed-citation xml:lang="ru">Хохлов С.С., Тырнов В.С., Гришина Е.В., Давоян Н.И., Зайцева М.И., Звержанская Л.С., Селиванов А.С., Суханов В,М., Шишкинская H.A., Гусева А.И. Гаплоидия и селекция. Москва: Наука; 1976.</mixed-citation><mixed-citation xml:lang="en">Khokhlov S.S., Tyrvov V.S., Grishina E.V., Davoyan N.I., Zaitseva M.I., Zverzhanskaya L.S., Selivanov A.S., Sukhanov V.M., Shishkinskaya N.A., Guseva A.I. Haploidy and breeding (Haploidiya i selektsiya). Мoscow: Nauka; 1976. [in Russian].</mixed-citation></citation-alternatives></ref><ref id="cit17"><label>17</label><citation-alternatives><mixed-citation xml:lang="ru">Kikuchi S., Takaiwa F., Oono K. Variable copy number DNA sequences in rice. Molecular and General Genetics. 1987;210(3):373-380. DOI: 10.1007/BF00327185</mixed-citation><mixed-citation xml:lang="en">Kikuchi S., Takaiwa F., Oono K. Variable copy number DNA sequences in rice. Molecular and General Genetics. 1987;210(3):373-380. DOI: 10.1007/BF00327185</mixed-citation></citation-alternatives></ref><ref id="cit18"><label>18</label><citation-alternatives><mixed-citation xml:lang="ru">Кириллова Г.А. Явление гаплоидии у покрытосеменных растений. Генетика. 1966;2:137-147.</mixed-citation><mixed-citation xml:lang="en">Kirillova G.A. Phenomenon of haploidy in angiospermous plants (Yavleniye gaploidii u pokrytosemennykh rasteniy). Genetika = Genetics. 1966;2:137-147. [in Russian].</mixed-citation></citation-alternatives></ref><ref id="cit19"><label>19</label><citation-alternatives><mixed-citation xml:lang="ru">Кунах В.А. Геномная изменчивость соматических клеток растений. 2. Изменчивость в природе. Биополимеры и клетка. 1995;11(6):5-40.</mixed-citation><mixed-citation xml:lang="en">Kunakh V.A. Genome variability in plant somatic cells. 2. Natural variability (Genomnaya izmenchivost somaticheskikh kletok rasteniy. 2. Izmenchivost v prirode). Biopolymers and Cell. 1995;11(6):5-40. [in Russian].</mixed-citation></citation-alternatives></ref><ref id="cit20"><label>20</label><citation-alternatives><mixed-citation xml:lang="ru">Кунах В.А. Геномная изменчивость соматических клеток растений. 4. Изменчивость в процессе дедифференцировки и каллусообразования in vitro. Биополимеры и клетка. 1998;14(4):298-319. DOI: 10.7124/bc.0004DB</mixed-citation><mixed-citation xml:lang="en">Kunakh V.A. Genome variability in plant somatic cells. 4. Variability in the process of dedifferentiation and callus formation in vitro (Genomnaya izmenchivost somaticheskikh kletok rasteniy. 4. Izmenchivost v protsesse dedifferentsirovki i kallusoobrazovaniya in vitro). Biopolymers and Cell. 1998;14(4):298-319. [in Russian]. DOI: 10.7124/bc.0004DB</mixed-citation></citation-alternatives></ref><ref id="cit21"><label>21</label><citation-alternatives><mixed-citation xml:lang="ru">Kuznetsova O.I., Ash O.A., Gostimsky S.A. The effect of duration of callus culture on the accumulation of genetic alteration in pea Pisum sativum L. Russian Journal of Genetics. 2006;42(5):555-562. DOI: 10.1134/s1022795406050139</mixed-citation><mixed-citation xml:lang="en">Kuznetsova O.I., Ash O.A., Gostimsky S.A. The effect of duration of callus culture on the accumulation of genetic alteration in pea Pisum sativum L. Russian Journal of Genetics. 2006;42(5):555-562. DOI: 10.1134/s1022795406050139</mixed-citation></citation-alternatives></ref><ref id="cit22"><label>22</label><citation-alternatives><mixed-citation xml:lang="ru">Levin D.A. Why polyploidy exceptionalism is not accompanied by reduced extinction rates. Plant Systematics and Evolution. 2019;305(1):1-11. DOI: 10.1007/s00606-018-1552-x</mixed-citation><mixed-citation xml:lang="en">Levin D.A. Why polyploidy exceptionalism is not accompanied by reduced extinction rates. Plant Systematics and Evolution. 2019;305(1):1-11. DOI: 10.1007/s00606-018-1552-x</mixed-citation></citation-alternatives></ref><ref id="cit23"><label>23</label><citation-alternatives><mixed-citation xml:lang="ru">Lewis H. Speciation in flowering plants. Science. 1966;152(3719):167-172. DOI: 1026/science.152.3719.167</mixed-citation><mixed-citation xml:lang="en">Lewis H. Speciation in flowering plants. Science. 1966;152(3719):167-172. DOI: 1026/science.152.3719.167</mixed-citation></citation-alternatives></ref><ref id="cit24"><label>24</label><citation-alternatives><mixed-citation xml:lang="ru">Ochatt S.J. Flow cytometry in plant breeding. Cytometry. 2008;73(7):581-598. DOI: 10.1002/cyto.a.20562</mixed-citation><mixed-citation xml:lang="en">Ochatt S.J. Flow cytometry in plant breeding. Cytometry. 2008;73(7):581-598. DOI: 10.1002/cyto.a.20562</mixed-citation></citation-alternatives></ref><ref id="cit25"><label>25</label><citation-alternatives><mixed-citation xml:lang="ru">Raven P.H., Thompson H.J. Haploidy and angiosperm evolution. The American Naturalist. 1964;98(901):251-252. DOI: 10.1086/282324</mixed-citation><mixed-citation xml:lang="en">Raven P.H., Thompson H.J. Haploidy and angiosperm evolution. The American Naturalist. 1964;98(901):251-252. DOI: 10.1086/282324</mixed-citation></citation-alternatives></ref><ref id="cit26"><label>26</label><citation-alternatives><mixed-citation xml:lang="ru">Scowcroff W.R. Somaclonal variation: The myth of clonal uniformity. In: B. Hohn, E.S. Dennis (eds). Genetic Flux in Planta. Wien: Springer; 1985. p.217-245. DOI: 10.1007/978-3-7091-8765-4</mixed-citation><mixed-citation xml:lang="en">Scowcroff W.R. Somaclonal variation: The myth of clonal uniformity. In: B. Hohn, E.S. Dennis (eds). Genetic Flux in Planta. Wien: Springer; 1985. p.217-245. DOI: 10.1007/978-3-7091-8765-4</mixed-citation></citation-alternatives></ref><ref id="cit27"><label>27</label><citation-alternatives><mixed-citation xml:lang="ru">Скапцов М.В., Смирнов С.В., Куцев М.Г., Шмаков А.И. Проблемы стандартизации в проточной цитометрии растений. Turczaninowia. 2016;19(3):120-122. DOI: 10.14258/turczaninowia.19.3.9</mixed-citation><mixed-citation xml:lang="en">Skaptsov M.V., Smirmov S.V., Kutsev M.G., Shmakov A.I. Problems of a standardization in plant flow cytometry. Turczaninowia. 2016;19(3):120-122. [in Russian]. DOI: 10.14258/turczaninowia.19.3.9</mixed-citation></citation-alternatives></ref><ref id="cit28"><label>28</label><citation-alternatives><mixed-citation xml:lang="ru">Stebbins G.L. A brief summary of my ideas on evolution. American Journal of Botany. 1999; 86(8):1207-1208. DOI: 10.2307/2656985</mixed-citation><mixed-citation xml:lang="en">Stebbins G.L. A brief summary of my ideas on evolution. American Journal of Botany. 1999; 86(8):1207-1208. DOI: 10.2307/2656985</mixed-citation></citation-alternatives></ref><ref id="cit29"><label>29</label><citation-alternatives><mixed-citation xml:lang="ru">Stebbins G.L. Chromosomal variation and evolution. Science. 1966; 152(3728):1463-1469. DOI: 10.1126/science.152.3728.1463</mixed-citation><mixed-citation xml:lang="en">Stebbins G.L. Chromosomal variation and evolution. Science. 1966; 152(3728):1463-1469. DOI: 10.1126/science.152.3728.1463</mixed-citation></citation-alternatives></ref><ref id="cit30"><label>30</label><citation-alternatives><mixed-citation xml:lang="ru">Stebbins G.L. Polyploidy, hybridization, and the invasion of new habitats. Annals of the Missouri Botanical Garden. 1985;72(4):824-832. DOI: 10.2307/2399224</mixed-citation><mixed-citation xml:lang="en">Stebbins G.L. Polyploidy, hybridization, and the invasion of new habitats. Annals of the Missouri Botanical Garden. 1985;72(4):824-832. DOI: 10.2307/2399224</mixed-citation></citation-alternatives></ref><ref id="cit31"><label>31</label><citation-alternatives><mixed-citation xml:lang="ru">Takhtajan A. Flowering plants. Dordrecht: Springer; 2009. DOI: 10.1007/978-1-4020-9609-9</mixed-citation><mixed-citation xml:lang="en">Takhtajan A. Flowering plants. Dordrecht: Springer; 2009. DOI: 10.1007/978-1-4020-9609-9</mixed-citation></citation-alternatives></ref><ref id="cit32"><label>32</label><citation-alternatives><mixed-citation xml:lang="ru">Тахтаджян А.Л. Макроэволюционные процессы в истории растительного мира. Ботанический журнал. 1983;68(12):1593- 1603.</mixed-citation><mixed-citation xml:lang="en">Takhtajan A.L. Macroevolutionary processes in the history of the plant world (Makroevolyutsionnye protsessy v istorii rastitelnogo mira). Botanicheskii zhurnal = Botanical Journal. 1983;68(12):1593-1603. [in Russian].</mixed-citation></citation-alternatives></ref><ref id="cit33"><label>33</label><citation-alternatives><mixed-citation xml:lang="ru">Цаценко Л.В., Мосунов С.А. Гаметы с соматическим числом хромосом: механизмы их формирования и роль в эволюции автополиплоидных растений (обзор иностранной литературы). Сельскохозяйственная биология. 2008;43(1):16-25.</mixed-citation><mixed-citation xml:lang="en">Tsatsenko L.V., Mosunov S.A. Gametes with somatic number of chromosomes: the mechanisms of their formation and their role in evolution of autopolyploid plants (review). Agricultural Biology. 2008;43(1):16-25. [in Russian].</mixed-citation></citation-alternatives></ref><ref id="cit34"><label>34</label><citation-alternatives><mixed-citation xml:lang="ru">Тырнов В.С. Гаплоидия у растений. Саратов: Саратовский университет; 2005.</mixed-citation><mixed-citation xml:lang="en">Tyrnov V.S. Haploidy in plants (Haploidiya u rasteniy). Saratov: Saratov University; 2005. [in Russian].</mixed-citation></citation-alternatives></ref><ref id="cit35"><label>35</label><citation-alternatives><mixed-citation xml:lang="ru">Wu Y., Sun Y., Sun S., Li G., Wang J., Wang B. et al. Aneuploidization under segmental allotetraploidy in rice and its phenotypic manifestation. Theoretical and Applied Genetics. 2018;131(6):1273-1285. DOI: 10.1007/s00122-018-3077-7</mixed-citation><mixed-citation xml:lang="en">Wu Y., Sun Y., Sun S., Li G., Wang J., Wang B. et al. Aneuploidization under segmental allotetraploidy in rice and its phenotypic manifestation. Theoretical and Applied Genetics. 2018;131(6):1273-1285. DOI: 10.1007/s00122-018-3077-7</mixed-citation></citation-alternatives></ref><ref id="cit36"><label>36</label><citation-alternatives><mixed-citation xml:lang="ru">Zagorska N.A., Shtereva L.A., Kruleva M.M., Sotirova V.G., Baralieva D.L., Dimotrov B.D. Induced androgenesis in tomato (Lycopersicon esculentum Mill.). III. Characterization of the regenerants. Plant Cell Reports. 2004;22(7):449- 456. DOI: 10.1007/s00299-003-0720-8</mixed-citation><mixed-citation xml:lang="en">Zagorska N.A., Shtereva L.A., Kruleva M.M., Sotirova V.G., Baralieva D.L., Dimotrov B.D. Induced androgenesis in tomato (Lycopersicon esculentum Mill.). III. Characterization of the regenerants. Plant Cell Reports. 2004;22(7):449- 456. DOI: 10.1007/s00299-003-0720-8</mixed-citation></citation-alternatives></ref><ref id="cit37"><label>37</label><citation-alternatives><mixed-citation xml:lang="ru">Zeyl C., Vanderford T., Carter M. An evolutionary advantage of haploidy in large yeast populations. Science. 2003;299(5606):555-558. DOI: 10.1126/science.1078417</mixed-citation><mixed-citation xml:lang="en">Zeyl C., Vanderford T., Carter M. An evolutionary advantage of haploidy in large yeast populations. Science. 2003;299(5606):555-558. DOI: 10.1126/science.1078417</mixed-citation></citation-alternatives></ref><ref id="cit38"><label>38</label><citation-alternatives><mixed-citation xml:lang="ru">Zheng K.L., Castiglione S., Biasini M.G., Biroli A., Morandi C., Sala F. Nuclear DNA amplification in cultured cells of Oryza sativa L. Theoretical and Applied Genetics. 1987;74(1):65-70. DOI: 10.1007/BF00290085</mixed-citation><mixed-citation xml:lang="en">Zheng K.L., Castiglione S., Biasini M.G., Biroli A., Morandi C., Sala F. Nuclear DNA amplification in cultured cells of Oryza sativa L. Theoretical and Applied Genetics. 1987;74(1):65-70. DOI: 10.1007/BF00290085</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>
