vavilovВавиловский журнал генетики и селекцииVavilov Journal of Genetics and Breeding2500-3259Institute of Cytology and Genetics of Siberian Branch of the RAS10.18699/VJ16.082vavilov-694Research ArticleЭволюция и видообразование. ОБЗОРEvolution and speciation. REVIEWМежвидовая несовместимость при отдаленной гибридизации растений и возможности ее преодоленияInterspecific incompatibility in wide hybridization of plants and ways to overcomeПершинаЛ. А.PershinaL. A.

Новосибирск

Novosibirsk

pershina@bionet.nsc.ruТрубачееваН. В.TrubacheevaN. V.

Новосибирск

Novosibirsk

Федеральное государственное бюджетное научное учреждение «Федеральный исследовательский центр Институт цитологии и генетики Сибирского отделения Российской академии наук»РоссияInstitute of Cytology and Genetics SB RASRussian FederationФедеральное государственное автономное образовательное учреждение высшего образования «Новосибирский национальный исследовательский государственный университет»РоссияNovosibirsk State UniversityRussian Federation201625092016204416425Copyright © Першина Л.А., Трубачеева Н.В., 20162016Першина Л.А., Трубачеева Н.В.Pershina L.A., Trubacheeva N.V.This work is licensed under a Creative Commons Attribution 4.0 License.https://vavilov.elpub.ru/jour/article/view/694

Отдаленная гибридизация вносит существенный вклад в видообразование покрытосеменных растений, обеспечивая перенос генов между видами и расширяя их потенциал к адаптации. В экспериментальных условиях отдаленная гибридизация используется для увеличения генетического разнообразия культурных растений. В процессе видообразования определяющее значение имеет возникновение репродуктивной изоляции, а выявляемая у растений способность к отдаленным скрещиваниям определяется возможностью ее преодоления. В обзоре рассматриваются основные типы несовместимости при отдаленных скрещиваниях растений, а также факторы и методические подходы, способствующие преодолению репродуктивных барьеров. Пример генетического контроля презиготических барьеров – роль генов Kr, определяющих нескрещиваемость между пшеницей и рожью. Постзиготическая несовместимость может быть связана с сильно выраженной генетической и эпигенетической изменчивостью, индуцируемой в результате отдаленных скрещиваний. Ранние этапы постзиготического периода являются критическими для развивающихся гибридных семян из-за гибели зародышей, в том числе связанной с однородительской элиминацией хромосом из гибридных клеток и аномальным развитием эндосперма. Рассматриваются возможности использования методов культивирования in vitro для преодоления презиготической и эмбриональной несовместимости у гибридов. Подчеркивается, что депрессия и летальность гибридов F1 обусловлены результатом взаимодействия комплементарных генов, вызывающих гибридные некроз, хлороз и карликовость. Обращено внимание на причины стерильности гибридов F1. В качестве одного из механизмов несовместимости при отдаленных скрещиваниях рассматриваются ядерно-цитоплазматические взаимодействия. Подчеркиваются проблемы в работе с реципрокными гибридами и аллоплазматическими линиями – основными моделями, используемыми при изучении эффектов цитоплазмы и ядерно-цитоплазматических взаимодействий. Приведены примеры, указывающие на то, что аллоплазматические линии не являются результатом простого сочетания ядерного генома одного вида, а цитоплазмы – другого. Процесс формирования аллоплазматических линий связан со структурно-функциональной изменчивостью ядерных и органельных геномов.

Wide hybridization is an important factor of angiosperm speciation and provides an introgression of genes between species. In experimental conditions wide hybridization is used to increase the genetic diversity of cultivated plants. Since the emergence of reproductive barriers is of great importance for speciation, plant capacity for wide crosses is determined by a possibility of overcoming these barriers. The review discusses the main types of incompatibility in wide crossing plants, as well as factors and methodological approaches that contribute to overcoming them. The role of Kr genes, which determine incompatibility between wheat and rye, is one of the examples of prezygotic isolation mechanism. Postzygotic incompatibility may be associated with a highly pronounced genetic and epigenetic variability induced by wide crossings. Early stages of the postzygotic period are crucial for developing hybrid seeds due to embryo’s death, including those associated with uniparental chromosome elimination in hybrid cells and abnormal development of the endosperm. A depression and a lethality of F1 hybrids may be the result of interaction between complementary genes, which cause hybrid necrosis, hybrid chlorosis, and hybrid dwarfness. The causes of hybrid sterility are discussed. Nuclearcytoplasmic interactions are regarded as one of the mechanisms of incompatibility in wide crosses. Reciprocal hybrids and alloplasmic lines are the main models for studying cytoplasm effects and nuclearcytoplasmic interactions. Problems concerning work with their models are emphasized. There are some examples underlining the fact that alloplasmic lines are not a plain combination of nuclear genome and cytoplasm of different species. Development of alloplasmatic lines is connected with structural and functional variability of nuclear and organelle genomes.

отдаленная гибридизацияпокрытосеменные растениямежвидовая несовместимостьпреодоление несовместимостиwide hybridizationangiosperm plantsinterspecific incompatibilityovercoming incompatibilityРФФИReferencesБадаева Е.Д., Першина Л.А., Бильданова Л.Л. Цитогенетическое исследование нестабильных по проявлению фертильности и жизнеспособности аллоплазматических рекомбинантных линий (Hordeum vulgare)-Triticum aestivum. Генетика. 2006;42(2): 198-209.Abbott R.J., Ireland H.I., Joseph L., Davies M.S., Rogers H.J. Recent plant speciation in Britain and Ireland: origins, establishment and evolution of four new hybrid species. Proc. Roy. Irish Acad. 2005;105:173-183.Банникова В.П. Межвидовая несовместимость у растений. Киев: Наук. думка, 1986.Ahmed T.A., Tsujimoto H., Sasakuma T. QTL analysis of fertility-restoration against cytoplasmic male sterility in wheat. Genes Genet. Syst. 2001;76:33-38.Будашкина Е.Б. Цитогенетика межвидовых гибридов пшеницы. Цитогенетика пшеницы и ее гибридов. Новосибирск: Наука, 1971:196-221.Ainouche M.L., Baumel A., Salmon A. Spartina anglica C.E. Hubbard: a natural model system for analyzing early evolutionary changes that affect allopolyploid genomes. Biol. J. Linn. Soc. 2004;82:475-484.Вавилов Н.И. Научные основы селекции пшеницы. Л.: Сельхозгиз, 1935.Aksyonova E., Sinyavskaya M., Danilenko N., Pershina L., Nakamura C., Davydenko O. Heteroplasmy and paternally oriented shift of the organellar DNA composition in barley-wheat hybrids during backcrosses with wheat parents. Genome. 2005;48:761-769.Даниленко Н.Г., Давыденко О.Г. Миры геномов органелл. Минск: Тэхналогiя, 2003.Aleza P., Juárez J., Cuenca J., Ollitrault P., Navarro L. Extensive citrus triploid hybrid production by 2x × 4x sexual hybridizations and parent-effect on the length of the juvenile phase. Plant Cell Rep. 2012;31:1723-1735.Карпеченко Г.Д. Теория отдаленной гибридизации. Теоретические основы селекции растений. Т. 1. М.; Л.: Сельхозгиз, 1935.Alvarez J.B., Ballesteros J., Sillero J.A., Martin L.M. Tritordeum: a new crop of potential importance in the food industry. Hereditas. 1992;116:193-197.Карпеченко Г.Д. Тетраплоидные ячмени, полученные действием высокой температуры. Биол. журнал. 1938;7(2).Ananiev E.V., Riera-Lizarazu O., Rines H.W., Phillips R.L. Oat-maize chromosome addition lines: A new system for mapping the maize genome. Proc. Natl Acad. Sci. USA. 1997;94:3524-3529.Коропачинский И.Ю., Милютин Л.И. Естественная гибридизация древесных растений. Новосибирск: Акад. изд-во «Гео», 2006.Anderson E. Introgressive hybridization. Biol. Rev. 1953;28:280-307.Лайкова Л.И., Белан И.А., Бадаева Е.Д., Россеева Л.П., Шепелев С.С., Шумный В.К., Першина Л.А. Создание и изучение сорта яровой мягкой пшеницы «Памяти Майстренко» с интрогрессией генетического материала от синтетического гексаплоида Triticum timopheevii Zhuk. × Aegilops tauschii Coss. Генетика. 2013;49(1):103-112.Asakura N., Nakamura C., Ohtsuka I. A nuclear compatibility gene, Ncc-tmp, of Triticum timopheevi for the cytoplasm of Aegilops squarrosa. Genes Genet. Syst. 1997;72:71-78.Нумерова О.М., Першина Л.А., Шумный В.К. Особенности мейоза ячменно-пшеничных гибридов H. vulgare × T. timopheevii Zhuk. и их потомков от чужеродных скрещиваний. Цитология и генетика. 1990;24(3):22-26.Badaeva E.D., Pershina L.A., Bil’danova L.L. Cytogenetic analysis of alloplasmic recombinant lines (H. vulgare)-T. aestivum with unstable fertility and viability. Russ. J. Genet. 2006;42:140-149.Першина Л.А. Хромосомная инженерия – направление биотехнологии. Вавиловский журнал генетики и селекции. 2014;18(1): 138-146.Banga S.S., Deol J.S., Banga S.K. Alloplasmic male-sterile Brassica juncea with Enarthrocarpus lyratus cytoplasm and the introgression of gene(s) for fertility restoration from cytoplasm donor species. Theor. Appl. Genet. 2003;106:1390-1395.Першина Л.А., Белова Л.И., Нумерова О.М., Шумный В.К. Особенности влияния генотипического разнообразия Hordeum vulgare L. и Secale cereale L. на скрещиваемость, развитие и жизнеспособность гибридных зародышей и растений. Генетика. 2000;36(4):520-526.Bannikova V.P. Mezhvidovaya nesovmestimost u rasteniy [Interspecies incompatibility in plants]. Kiev, Naukova Dumka Publ., 1986.Першина Л.А., Девяткина Э.П., Трубачеева Н.В., Кравцова Л.А., Добровольская О.Б. Особенности восстановления фертильности аллоплазматических линий, полученных на основе гибридизации самоопыленного потомка ячменно-пшеничного амфиплоида (Hordeum vulgare L. × Triticum aestivum L.) с сортами мягкой пшеницы Саратовская 29 и Пиротрикс 28. Генетика. 2012;48(12):1372-1379.Bentley K.E., Mande J.R., McCauley D.E. Paternal leakage and heteroplasmy of mitochondrial genomes in Silene vulgaris: evidence from experimental crosses. Genetics. 2010;185:961-968.Першина Л.А., Трубачеева Н.В., Синявская М.Г., Девяткина Э.П., Кравцова Л.А. Ядерно-цитоплазматическая совместимость и состояние районов митохондриальной и хлоропластной ДНК у аллоплазматических рекомбинантных и интрогрессивных линий (H. vulgare) – T. aestivum. Генетика. 2014;50(10):1154-1162.Bertin I., Fish L., Foote T., Knight E., Snape J., Moore G. Development of consistently crossable wheat genotypes for alien wheat gene transfer through fine- mapping of the Kr1 locus. Theor. Appl. Genet. 2009;119:1371-1381.Пухальский В.А., Мартынов С.П., Добротворская Т.В. Гены гибридного некроза пшениц (теория вопроса и каталог носителей летальных генов). М.: Изд-во МСХА, 2002.Blakeslee A.F., Avery A.C. Methods of inducing doubling of chromosomes in plants. J. Hered. 1937;28(12):393-411.Пухальский В.А., Билинская Е.А., Мартынов С.П., Добротворская Т.В., Оболенкова Г.А. Новые данные по распространению генов гибридного некроза в сортах озимой мягкой пшеницы (Triticum aestivum L.). Генетика. 2008;44(2):209-218.Bogdanova V.S. Inheritance of organelle DNA markers in a pea cross associated with nuclear-cytoplasmic incompatibility. Theor. Appl. Genet. 2007;114:333-339.Трубачеева Н.В., Ефремова Т.Т., Бадаева Е.Д., Кравцова Л.А., Белова Л.И., Девяткина Э.П., Першина Л.А. Получение аллоплазматических и эуплазматических пшенично-ячменных дителосомных замещенных линий 7H1Lmar(7D) и изучение 18S/5S митохондриального повтора у этих линий. Генетика. 2009; 45(12):1627-1633.Bomblies K., Lempe J., Epple P., Warthmann N., Lanz C., Dangl J.L., Weigel D. Autoimmune response as a mechanism for a Dobzhansky- Muller-type incompatibility syndrome in plants. PLoS Biol. 2007; 5(9):e236.Трубачеева Н.В., Кравцова Л.А., Девяткина Э.П., Ефремова Т.Т., Синявская М.Г., Шумный В.К., Першина Л.А. Гетероплазматическое и гомоплазматическое состояние митохондриальной и хлоропластной ДНК у потомков отдаленных гибридов мягкой пшеницы разного происхождения. Вавиловский журнал генетики и селекции. 2012;16(1):160-169.Bomblies K., Weigel D. Hybrid necrosis: autoimmunity as a potential gene-flow barrier in plant species. Nat. Rev. Genet. 2007;8:382-393.Шулындин А.Ф. Генетические закономерности расщепления отдаленных гибридов растений. Проблемы отдаленной гибридизации растений. София: Болгарская Академия наук. 1978: 33-44.Brautigam K., Dietzel I., Pfannschmidt T. Plastid-nucleus communication: anterograde and retrograde signaling in the development and function of plastids. Cell Mol. Biol. Plastids. 2007;19:409-455.Abbott R.J., Ireland H.I., Joseph L., Davies M.S., Rogers H.J. Recent plant speciation in Britain and Ireland: origins, establishment and evolution of four new hybrid species. Proc. Roy. Irish Acad. 2005;105:173-183.Budashkina E.B. Tsitogenetika mezhvidovykh gibridov pshenitsy [Cytogenetics interspecific hybrids of wheat]. Tsitogenetika pshenitsy i ee gibridov [Cytogenetics of wheat and its hybrids]. Novosibirsk, Nauka Publ., 1971:196-221.Ahmed T.A., Tsujimoto H., Sasakuma T. QTL analysis of fertility-restoration against cytoplasmic male sterility in wheat. Genes Genet. Syst. 2001;76:33-38.Buggs R.J.A., Zhang L., Miles N., Tate J.A., Gao L., Wei W., Schnable P.S., Barbazuk W.B., Soltis P.S., Soltis D.E. Transcriptomic shock generates evolutionary novelty in a newly formed, natural allopolyploid plant. Curr. Biol. 2011;21:551-556.Ainouche M.L., Baumel A., Salmon A. Spartina anglica C.E. Hubbard: a natural model system for analyzing early evolutionary changes that affect allopolyploid genomes. Biol. J. Linn. Soc. 2004;82:475-484.Bushell C., Spielman M., Scott R.J. The basis of natural and artificial postzygotic hybridization barriers in Arabidopsis species. Plant Cell. 2003;15:1430-1442.Aksyonova E., Sinyavskaya M., Danilenko N., Pershina L., Nakamura C., Davydenko O. Heteroplasmy and paternally oriented shift of the organellar DNA composition in barley-wheat hybrids during backcrosses with wheat parents. Genome. 2005;48:761-769.Comai L. Genetic and epigenetic interactions in allopolyploids plants. Plant Mol. Biol. 2000;43:387-399.Aleza P., Juárez J., Cuenca J., Ollitrault P., Navarro L. Extensive citrus triploid hybrid production by 2x × 4x sexual hybridizations and parent-effect on the length of the juvenile phase. Plant Cell Rep. 2012;31:1723-1735.Crismani W., Girard C., Mercier R. Tinkering withmeiosis. J. Exp. Bot. 2013;64:55-65.Alvarez J.B., Ballesteros J., Sillero J.A., Martin L.M. Tritordeum: a new crop of potential importance in the food industry. Hereditas. 1992;116:193-197.Crosatti C., Quansah L., Maré C., Giusti L., Roncaglia E., Atienza S.G., Cattivelli L., Fait A. Cytoplasmic genome substitution in wheat affects the nuclear-cytoplasmic cross-talk leading to transcript and metabolite alterations. BMC Genom. 2013;14:868- 889.Ananiev E.V., Riera-Lizarazu O., Rines H.W., Phillips R.L. Oat-maize chromosome addition lines: A new system for mapping the maize genome. Proc. Natl Acad. Sci. USA. 1997;94:3524-3529.Danilenko N.G., Davidenko O.G. Miry genomov organell [World of organelle genomes]. Minsk, Tekhnologia Publ., 2003.Anderson E. Introgressive hybridization. Biol. Rev. 1953;28:280-307.Davies D.R. Chromosome elimination in inter-specific hybrids. Heredity. 1974;32:267- 270.Asakura N., Nakamura C., Ohtsuka I. A nuclear compatibility gene, Ncc-tmp, of Triticum timopheevi for the cytoplasm of Aegilops squarrosa. Genes Genet. Syst. 1997;72:71-78.Devaux P. The Hordeum bulbosum (L.) method. Doubled Haploid Production in Crop Plants: A Manual. Eds M. Maluszynski, K.J. Kasha, B.P. Forster, I. Szarejko. Dordrecht: Kluwer Acad. Publ. 2003; 15-19.Banga S.S., Deol J.S., Banga S.K. Alloplasmic male-sterile Brassica juncea with Enarthrocarpus lyratus cytoplasm and the introgression of gene(s) for fertility restoration from cytoplasm donor species. Theor. Appl. Genet. 2003;106:1390-1395.Dilkes B.P., Comai L. A differential dosage hypothesis for parental effects in seed development. Plant Cell. 2004;16:3174-3180.Bentley K.E., Mande J.R., McCauley D.E. Paternal leakage and heteroplasmy of mitochondrial genomes in Silene vulgaris: evidence from experimental crosses. Genetics. 2010;185:961-968.Eilam T., Anikster Y., Millet E., Manisterski J., Feldman M. Nuclear DNA amount and genome downsizing in natural and synthetic allopolyploids of the genera Aegilops and Triticum. Genome. 2008;51: 616-627.Bertin I., Fish L., Foote T., Knight E., Snape J., Moore G. Development of consistently crossable wheat genotypes for alien wheat gene transfer through fine-mapping of the Kr1 locus. Theor. Appl. Genet. 2009;119:1371-1381.Ellis J.R., Bentley K.E., McCauley D.E. Detection of rare paternal chloroplast inheritance in controlled crosses of the endangered sunflower Helianthus verticillatus. Heredity. 2008;100:574-580.Blakeslee A.F., Avery A.C. Methods of inducing doubling of chromosomes in plants. J. Hered. 1937;28(12):393-411.Ellstrand N.C., Schierenbeck K.A. Hybridization as stimulus for the evolution of invasivеness in plants? Proc. Natl Acad. Sci. USA. 2000;97:7043-7050.Bogdanova V.S. Inheritance of organelle DNA markers in a pea cross associated with nuclear-cytoplasmic incompatibility. Theor. Appl. Genet. 2007;114:333-339.Farbos I., Mouras A., Bereterbide A., Glimelius K. Defective cell proliferation in the floral meristem of alloplasmic plants of Nicotiana tabacum leads to abnormal floral organ development and male sterility. Plant J. 2001;26:131-142.Bomblies K., Lempe J., Epple P., Warthmann N., Lanz C., Dangl J.L., Weigel D. Autoimmune response as a mechanism for a DobzhanskyMuller-type incompatibility syndrome in plants. PLoS Biol. 2007; 5(9):e236.Feldman M., Levy A.A. Allopolyploidy – a shaping force in the evolution of wheat genomes. Cytogenet. Genome Res. 2005;109:250-258.Bomblies K., Weigel D. Hybrid necrosis: autoimmunity as a potential gene-flow barrier in plant species. Nat. Rev. Genet. 2007;8:382-393.Fujii S., Toriyama K. Genome barriers between nuclei and mitochondria exemplified by cytoplasmic male sterility. Plant Cell Physiol. 2008;49:1484-1494.Brautigam K., Dietzel I., Pfannschmidt T. Plastid-nucleus communication: anterograde and retrograde signaling in the development and function of plastids. Cell Mol. Biol. Plastids. 2007;19:409-455.Gaeta R.T., Yoo S.Y., Pires J.C., Doerge R.W., Chen Z.J., Osborn T.C. Analysis of gene expression in resynthesized Brassica napus allopolyploids using Arabidopsis 70 mer oligo microarrays. PLoS ONE. 2009;4:e4760.Buggs R.J.A., Zhang L., Miles N., Tate J.A., Gao L., Wei W., Schnable P.S., Barbazuk W.B., Soltis P.S., Soltis D.E. Transcriptomic shock generates evolutionary novelty in a newly formed, natural allopolyploid plant. Curr. Biol. 2011;21:551-556.Geerts P., Toussaint A., Mergeai G., Baudoin J.P. Phaseolus immature embryo rescue technology. Method. Mol. Biol. 2011;710:117-129.Bushell C., Spielman M., Scott R.J. The basis of natural and artificial postzygotic hybridization barriers in Arabidopsis species. Plant Cell. 2003;15:1430-1442.Gernand D., Houben A., Pickering R., Rutten T. Elimination of chromosomes in Hordeum vulgare × H. bulbosum crosses at mitosis and interphase involves micronucleus formation and progressive heterochromatinization. Cytogenet. Genome Res. 2006;114:169-174.Comai L. Genetic and epigenetic interactions in allopolyploids plants. Plant Mol. Biol. 2000;43:387-399.Gernand D., Rutten T., Varshney A., Rubtsova M., Prodanovic S., Brüß C., Kumlehn J., Matzk F., Houben A. Uniparental chromosome elimination at mitosis and interphase in wheat and pearl millet crosses involves micronucleus formation, progressive heterochromatinization, and DNA fragmentation. Plant Cell. 2005;17:2431-2438.Crismani W., Girard C., Mercier R. Tinkering withmeiosis. J. Exp. Bot. 2013;64:55-65.Gerstel D.U., Burns J.A., Burk L.G. Cytoplasmic male sterility in Nicotiana, restoration of fertility, and the nucleolus. Genetics. 1978; 89:157-169.Crosatti C., Quansah L., Maré C., Giusti L., Roncaglia E., Atienza S.G., Cattivelli L., Fait A. Cytoplasmic genome substitution in wheat affects the nuclear-cytoplasmic cross-talk leading to transcript and metabolite alterations. BMC Genom. 2013;14:868-889.Hattori N., Kitagawa K., Takumi S., Nakamura C. Mitochondrial DNA heteroplasmy in wheat, Aegilops and their nucleus-cytoplasm hybrids. Genetics. 2002;160(4):1619- 1630.Davies D.R. Chromosome elimination in inter-specific hybrids. Heredity. 1974;32:267-270.He G., Zhu X., Elling A.A., Chen L., Wang X., Guo L., Liang M., He H., Zhang H., Chen F., Qi Y., Chen R., Denga X-Q. Global epigenetic and transcriptional trends among two rice subspecies and their reciprocal hybrids. Plant Cell. 2010;22:17-33.Devaux P. The Hordeum bulbosum (L.) method. Doubled Haploid Production in Crop Plants: A Manual. Eds M. Maluszynski, K.J. Kasha, B.P. Forster, I. Szarejko. Dordrecht: Kluwer Acad. Publ. 2003; 15-19.Ho K.M., Kasha K.J. Genetic control of chromosome elimination during haploid formation in barley. Genome. 1975;81:263-273.Dilkes B.P., Comai L. A differential dosage hypothesis for parental effects in seed development. Plant Cell. 2004;16:3174-3180.Hossain K.G., Riera-Lizarazu O., Kalavacharia V., Vales M.I., Rust J.L., Maan S.S., Kianian S.F. Molecular cytogenetic characterization of an alloplasmic durum wheat line with a portion of chromosome 1D of Triticum aestivum carrying the scs ae gene. Genome. 2004;47: 206-214.Eilam T., Anikster Y., Millet E., Manisterski J., Feldman M. Nuclear DNA amount and genome downsizing in natural and synthetic allopolyploids of the genera Aegilops and Triticum. Genome. 2008;51: 616-627.Houben A., Sanei M., Pickering R. Barley doubled-haploid production by uniparental chromosome elimination. Plant Cell Tiss. Org. 2011;104:321-327.Ellis J.R., Bentley K.E., McCauley D.E. Detection of rare paternal chloroplast inheritance in controlled crosses of the endangered sunflower Helianthus verticillatus. Heredity. 2008;100:574-580.Inagaki M.N., Mujeeb-Kazi A. Production of polyhaploids of hexaploid wheat using stored pearl millet pollen. Euphytica. 1998;100(1-3): 253-259.Ellstrand N.C., Schierenbeck K.A. Hybridization as stimulus for the evolution of invasivеness in plants? Proc. Natl Acad. Sci. USA. 2000;97:7043-7050.Ishii T., Ueda T., Tanaka H., Tsujimoto H. Chromosome elimination by wide hybridization between Triticeae or oat plant and pearl millet: pearl chromosome dynamics in hybrid embryo cells. Chromosome Res. 2010;18:821-883.Farbos I., Mouras A., Bereterbide A., Glimelius K. Defective cell proliferation in the floral meristem of alloplasmic plants of Nicotiana tabacum leads to abnormal floral organ development and male sterility. Plant J. 2001;26:131-142.Ito T., Konno I., Kubota S., Ochiai T., Sonoda T., Hayashi Y., Fukuda T., Yokoyama J., Nakayama H., Kameya T., Kanno A. Production and characterization of interspecific hybrids between Asparagus kiusianus Makino and A. officinalis L. Euphytica. 2011;182:285-294.Feldman M., Levy A.A. Allopolyploidy – a shaping force in the evolution of wheat genomes. Cytogenet. Genome Res. 2005;109:250-258.Janeja H.S., Banga S.K., Bhaskar P.B., Banga S.S. Alloplasmic male sterile Brassica napus with Enarthrocarpus lyratus cytoplasm: introgression and molecular mapping of an E. lyratus chromosome segment carrying a fertility restoring gene. Genome. 2003;46:792-797.Fujii S., Toriyama K. Genome barriers between nuclei and mitochondria exemplified by cytoplasmic male sterility. Plant Cell Physiol. 2008;49:1484-1494.Jansky S. Overcoming hybridization barriers in potato. Plant Breeding. 2006;125:1-12.Gaeta R.T., Yoo S.Y., Pires J.C., Doerge R.W., Chen Z.J., Osborn T.C. Analysis of gene expression in resynthesized Brassica napus allopolyploids using Arabidopsis 70 mer oligo microarrays. PLoS ONE. 2009;4:e4760.Jesse D., Woodson J.D., Chory J. Coordination of gene expression between organellar and nuclear genomes. Nat. Rev. Genet. 2008;9: 383-395.Geerts P., Toussaint A., Mergeai G., Baudoin J.P. Phaseolus immature embryo rescue technology. Method. Mol. Biol. 2011;710:117-129.Jiang J., Chen P., Friebe B., Raupp W.J., Gill B.S. Alloplasmic wheat – Elymus ciliaris chromosome addition lines. Genome. 1993;36: 327-333.Gernand D., Houben A., Pickering R., Rutten T. Elimination of chromosomes in Hordeum vulgare × H. bulbosum crosses at mitosis and interphase involves micronucleus formation and progressive heterochromatinization. Cytogenet. Genome Res. 2006;114:169-174.Karpechenko G.D. Teoriya otdalennoy gibridizatsii [Theory of remote hybridization]. Teoreticheskie osnovy selektsii rasteniy [Theoretical Foundations of Plant Breeding]. Moscow; Leningrad, Selhozgiz,1935;1.Gernand D., Rutten T., Varshney A., Rubtsova M., Prodanovic S., Brüß C., Kumlehn J., Matzk F., Houben A. Uniparental chromosome elimination at mitosis and interphase in wheat and pearl millet crosses involves micronucleus formation, progressive heterochromatinization, and DNA fragmentation. Plant Cell. 2005;17:2431-2438.Karpechenko G.D. Tetraploid barleys obtained using high temperatures. Biologicheskiy zhurnal = Biological Journal. 1938;7(2).Gerstel D.U., Burns J.A., Burk L.G. Cytoplasmic male sterility in Nicotiana, restoration of fertility, and the nucleolus. Genetics. 1978; 89:157-169.Kaul M.L.H. Male Sterility in Higher Plants. Berlin; Heidelberg; N.Y.; London; Paris; Toronto: Springer-Verlag, 1988.Hattori N., Kitagawa K., Takumi S., Nakamura C. Mitochondrial DNA heteroplasmy in wheat, Aegilops and their nucleus-cytoplasm hybrids. Genetics. 2002;160(4):1619-1630.Kiang A.S., Connolly V., McConnell D.J., Kavanagh T.A. Paternal inheritance of mitochondria and chloroplasts in Festuca pratensis- Lolium perenne intergeneric hybrids. Theor. Appl. Genet. 1994;87: 681-688.He G., Zhu X., Elling A.A., Chen L., Wang X., Guo L., Liang M., He H., Zhang H., Chen F., Qi Y., Chen R., Denga X-Q. Global epigenetic and transcriptional trends among two rice subspecies and their reciprocal hybrids. Plant Cell. 2010;22:17-33.Kitagawa K., Takumi S., Nakamura C. Evidence of paternal transmission of mitochondrial DNA in a nucleus-cytoplasm hybrid of timopheevi wheat. Genes. Genet. Syst. 2002;77:243-250.Ho K.M., Kasha K.J. Genetic control of chromosome elimination during haploid formation in barley. Genome. 1975;81:263-273.Koba T., Shimada T. Variations in the crossability of common wheat cultivars with cultivated barley. Hereditas. 1992;116:187-192.Hossain K.G., Riera-Lizarazu O., Kalavacharia V., Vales M.I., Rust J.L., Maan S.S., Kianian S.F. Molecular cytogenetic characterization of an alloplasmic durum wheat line with a portion of chromosome 1D of Triticum aestivum carrying the scs ae gene. Genome. 2004;47: 206-214.Koropachinsky I.Y., Milutin L.I. Estestvennaya gibridizatsiya drevesnykh rasteniy [Natural hybridization of woody plants]. Novosibirsk, Geo, 2006.Houben A., Sanei M., Pickering R. Barley doubled-haploid production by uniparental chromosome elimination. Plant Cell Tiss. Org. 2011;104:321-327.Kruse A. An in vivo/in vitro embryo culture technique. Hereditas. 1974; 77:219-224.Inagaki M.N., Mujeeb-Kazi A. Production of polyhaploids of hexaploid wheat using stored pearl millet pollen. Euphytica. 1998;100(1-3): 253-259.Lai Z., Nakazato T., Salmaso M., Burke J.M., Tang S., Knapp S.J., Rieseberg L.H. Extensive chromosomal repatterning and evolution of sterility barriers in hybrid sunflower species. Genetics. 2005;171: 291-303.Ishii T., Ueda T., Tanaka H., Tsujimoto H. Chromosome elimination by wide hybridization between Triticeae or oat plant and pearl millet: pearl chromosome dynamics in hybrid embryo cells. Chromosome Res. 2010;18:821-883.Laibach F. Das taubwerden der bastardsamen und die künstliche anfzucht früh absterbender bastardembryonen. Ztschr. Bot. 1925;17: 417-485.Ito T., Konno I., Kubota S., Ochiai T., Sonoda T., Hayashi Y., Fukuda T., Yokoyama J., Nakayama H., Kameya T., Kanno A. Production and characterization of interspecific hybrids between Asparagus kiusianus Makino and A. officinalis L. Euphytica. 2011;182:285-294.Laikova L.I., Belan I.A., Badaeva E.D., Rosseeva L.P., Shepelev S.S., Shumny V.K., Pershina L.A. Development and study of spring bread wheat variety “Pamyati Maystrenko” with introgression of genetic material from synthetic hexaploid Triticum timopheevii Zhuk. × Aegilops tauschii Coss. Russ. J. Genet. 2013;49:89-97.Janeja H.S., Banga S.K., Bhaskar P.B., Banga S.S. Alloplasmic male sterile Brassica napus with Enarthrocarpus lyratus cytoplasm: introgression and molecular mapping of an E. lyratus chromosome segment carrying a fertility restoring gene. Genome. 2003;46: 792-797.Landgren M., Zetterstrand M., Sundberg E., Glimelius K. Alloplasmic male-sterile Brassica lines containing B. tournefortii mitochondria express an ORF 3 of the atp6 gene and a 32 kDa protein. Plant Mol. Biol. 1996;32:879-890.Jansky S. Overcoming hybridization barriers in potato. Plant Breeding. 2006;125:1-12.Lange L., Wojciechowska B. The crossing of common wheat (T. aestivum L.) with cultivated rye (S. cereale L.). I. Crossability, pollen grain germination and pollen tube growth. Euphytica. 1976;25:609-620.Jesse D., Woodson J.D., Chory J. Coordination of gene expression between organellar and nuclear genomes. Nat. Rev. Genet. 2008;9: 383-395.Laurie D.A., Bennett M.D. The effect of the crossability loci Kr1 and Kr2 on fertilization frequency in hexapoid wheat × maize crosses. Theor. Appl. Genet. 1987;73:403-409.Jiang J., Chen P., Friebe B., Raupp W.J., Gill B.S. Alloplasmic wheat – Elymus ciliaris chromosome addition lines. Genome. 1993;36: 327-333.Lee J.A. Genetics of D3 complementary lethality in Gossypium hirsutum and G. barbadense. J. Hered. 1981;72:299-300.Kaul M.L.H. Male Sterility in Higher Plants. Berlin; Heidelberg; N.Y.; London; Paris; Toronto: Springer-Verlag, 1988.Levin D.A. The cytoplamic factor in plant speciation. Syst. Bot. 2003; 28(1):5-11.Kiang A.S., Connolly V., McConnell D.J., Kavanagh T.A. Paternal inheritance of mitochondria and chloroplasts in Festuca pratensisLolium perenne intergeneric hybrids. Theor. Appl. Genet. 1994;87: 681-688.Li G.R., Ji W., Wang G., Zhang J.X., Wang Y.J. An improved embryorescue protocol for hybrid progeny from seedless Vitis vinifera grapes × wild Chinese Vitis species. In Vitro Cell. Dev. Biol. Plant. 2014a;50:110-120.Kitagawa K., Takumi S., Nakamura C. Evidence of paternal transmission of mitochondrial DNA in a nucleus-cytoplasm hybrid of timopheevi wheat. Genes. Genet. Syst. 2002;77:243-250.Li J., Sun Q., Yu N., Zhu J., Zou X., Qi Z., Ghani M.A., Chen L. The role of small RNAs on phenotypes in reciprocal hybrids between Solanum lycopersicum and S. pimpinellifolium. BMC Plant Biol. 2014b;14:296-306.Koba T., Shimada T. Variations in the crossability of common wheat cultivars with cultivated barley. Hereditas. 1992;116:187-192.Liu H.Y., Xu C.G., Zhang Q. Male and female gamete abortions, and reduced affinity between the uniting gametes as the causes for sterility in an indica/japonica hybrid in rice. Sex. Plant Reprod. 2004; 17:55-62.Kruse A. An in vivo/in vitro embryo culture technique. Hereditas. 1974; 77:219-224.Maan S.S. Transfer of the species specific cytoplasm (scs) from Triticum timopheevii to Triticum turgidum. Genome. 1992;35:238-243.Lai Z., Nakazato T., Salmaso M., Burke J.M., Tang S., Knapp S.J., Rieseberg L.H. Extensive chromosomal repatterning and evolution of sterility barriers in hybrid sunflower species. Genetics. 2005;171: 291-303.Mahé L., Le Pierrès D., Combes M.-C., Lashermes P. Introgressive hybridization between the allotetraploid Coffea arabica and one of its diploid ancestors, Coffea canephora, in an exceptional sympatric zone in New Caledonia. Genome. 2007;50:316- 324.Laibach F. Das taubwerden der bastardsamen und die künstliche anfzucht früh absterbender bastardembryonen. Ztschr. Bot. 1925;17: 417-485.Michalak de Jimenez M.K., Bassi F.M., Ghavami F., Simons K., Dizon R., Seetan R.I., Alnemer L.M., Denton A.M., Doğramaci M., Šimková H., Doležel J., Seth K., Luo M.-C., Dvorak J., Gu Y.Q., Kianian S.F. A radiation hybrid map of chromosome 1D reveals synteny conservation at a wheat speciation locus. Funct. Integr. Genomics. 2013;13:19-32.Landgren M., Zetterstrand M., Sundberg E., Glimelius K. Alloplasmic male-sterile Brassica lines containing B. tournefortii mitochondria express an ORF 3 of the atp6 gene and a 32 kDa protein. Plant Mol. Biol. 1996;32:879-890.Mizuno N., Hosog N., Park P., Takum S. Hypersensitive response-like reaction is associated with hybrid necrosis in interspecific crosses between tetraploid wheat and Aegilops tauschii Coss. PLoS ONE. 2010;5:1-16.Lange L., Wojciechowska B. The crossing of common wheat (T. aestivum L.) with cultivated rye (S. cereale L.). I. Crossability, pollen grain germination and pollen tube growth. Euphytica. 1976;25: 609-620.Molnár-Láng M., Linc G., Logojan A., Sutka J. Production and meiotic pairing behaviour of new hybrids of winter wheat (Triticum aestivum) × winter barley (Hordeum vulgare). Genome. 2000;43: 1045-1054.Laurie D.A., Bennett M.D. The effect of the crossability loci Kr1 and Kr2 on fertilization frequency in hexapoid wheat × maize crosses. Theor. Appl. Genet. 1987;73:403-409.Molnár-Láng M., Linc G., Sutka J. Transfer of the recessive crossability allele kr1 from Chinese Spring into the winter wheat variety Martonvásári 9. Euphytica. 1996;90:301-305.Lee J.A. Genetics of D3 complementary lethality in Gossypium hirsutum and G. barbadense. J. Hered. 1981;72:299-300.Moreno P.E., Caetano C.M., Olaya C.A., Agrono T.C., Torres E.A. Chromosome elimination in intergeneric hybrid of Oryza sativa × Luziola peruviana. Agr. Sci. 2014;5:1344-1350.Levin D.A. The cytoplamic factor in plant speciation. Syst. Bot. 2003; 28(1):5-11.Moreno E.M.S., Speranza P.R., Laque J.M.R., Neffa V.G.S. Natural hybridization among subspecies of Turnera sidoides L. (Passifloraceae) revealed by morphological and genetic evidence. Plant Syst. Evol. 2015;301:883-892.Li G.R., Ji W., Wang G., Zhang J.X., Wang Y.J. An improved embryorescue protocol for hybrid progeny from seedless Vitis vinifera grapes × wild Chinese Vitis species. In Vitro Cell. Dev. Biol. Plant. 2014a;50:110-120.Nakano N., Mizuno N., Tosa Y., Yoshida K., Park P., Takumi S. Accelerated senescence and enhanced disease resistance in hybrid chlorosis lines derived from interspecific crosses between tetraploid wheat and Aegilops tauschii. PLoS ONE. 2015;10. DOI 10.1371/journal.pone.0121583.Li J., Sun Q., Yu N., Zhu J., Zou X., Qi Z., Ghani M.A., Chen L. The role of small RNAs on phenotypes in reciprocal hybrids between Solanum lycopersicum and S. pimpinellifolium. BMC Plant Biol. 2014b;14:296-306.Ng D.W.-K., Lu J., Chen Z.J. Big roles for small RNAs in polyploidy, hybrid vigor, and hybrid incompatibility. Curr. Opin. Plant Biol. 2012;15:154-161.Liu H.Y., Xu C.G., Zhang Q. Male and female gamete abortions, and reduced affinity between the uniting gametes as the causes for sterility in an indica/japonica hybrid in rice. Sex. Plant Reprod. 2004; 17:55-62.Niroula R.K., Bimb H.P. Overview of wheat × maize system of crosses for dihaploid induction in wheat. World Appl. Sci. J. 2009;7:1037-1045.Maan S.S. Transfer of the species specific cytoplasm (scs) from Triticum timopheevii to Triticum turgidum. Genome. 1992;35:238-243.Niu Z., Jiang A., Hammad W.A., Oladzadabbasabadi A., Xu S.S., Mergoum M., Elias E.M. Review of doubled haploid production in durum and common wheat through wheat × maize hybridization. Plant Breeding. 2014;133:313-320.Mahé L., Le Pierrès D., Combes M.-C., Lashermes P. Introgressive hybridization between the allotetraploid Coffea arabica and one of its diploid ancestors, Coffea canephora, in an exceptional sympatric zone in New Caledonia. Genome. 2007;50:316-324.Numerova O.M., Pershina L.A., Shumny V.K. Meiosis in barley-wheat hybrids H. vulgare × T. timopheevii Zhuk and their descendants from alien crossings. Tsitologiya i genetika = Cytology and Genetics. 1990;24(3):22-26.Michalak de Jimenez M.K., Bassi F.M., Ghavami F., Simons K., Dizon R., Seetan R.I., Alnemer L.M., Denton A.M., Doğramaci M., Šimková H., Doležel J., Seth K., Luo M.-C., Dvorak J., Gu Y.Q., Kianian S.F. A radiation hybrid map of chromosome 1D reveals synteny conservation at a wheat speciation locus. Funct. Integr. Genomics. 2013;13:19-32.Pathania A., Bhat S.R., Dinesh Kumar V., Ashutosh K.P.B., Prakash S., Chopra V.L. Cytoplasmic male sterility in alloplasmic Brassica juncea carrying Diplotaxis catholica cytoplasm: molecular characterization and genetics of fertility restoration. Theor. Appl. Genet. 2003;107:455-461.Mizuno N., Hosog N., Park P., Takum S. Hypersensitive response-like reaction is associated with hybrid necrosis in interspecific crosses between tetraploid wheat and Aegilops tauschii Coss. PLoS ONE. 2010;5:1-16.Pelletier G., Budar F. The molecular biology of cytoplasmically inherited male sterility and prospects for its engineering. Curr. Opin. Biotechnol. 2007;18:121-125.Molnár-Láng M., Linc G., Logojan A., Sutka J. Production and meiotic pairing behaviour of new hybrids of winter wheat (Triticum aestivum) × winter barley (Hordeum vulgare). Genome. 2000;43: 1045-1054.Pershina L.A. Plant chromosome engineering is an area of biotechnology. Russ. J. Genet.: Appl. Res. 2014;4(4):311-317.Molnár-Láng M., Linc G., Sutka J. Transfer of the recessive crossability allele kr1 from Chinese Spring into the winter wheat variety Martonvásári 9. Euphytica. 1996;90:301-305.Pershina L.A., Belova L.I., Numerova O.M., Shumny V.K. Effect of the genotypic diversity of Hordeum vulgare L. and Secale cereale L. on their crossability and the development and viability of hybrid embryos and plants. Genetika = Genetics (Moscow). 2000;36(4): 520-526.Moreno P.E., Caetano C.M., Olaya C.A., Agrono T.C., Torres E.A. Chromosome elimination in intergeneric hybrid of Oryza sativa × Luziola peruviana. Agr. Sci. 2014;5:1344-1350.Pershina L.A., Devyatkina E.P., Trubacheeva N.V., Kravtsova L.A., Dobrovol’skaya O.B. Characterization of fertility restoration in alloplasmic lines derived from hybridization of self-fertilized offspring of barley–wheat (Hordeum vulgare L. × Triticum aestivum L.) amphiploid with common wheat varieties Saratovskaya 29 and Pyrotrix 28. Russ. J. Genet. 2012;48(12):1184-1190.Moreno E.M.S., Speranza P.R., Laque J.M.R., Neffa V.G.S. Natural hybridization among subspecies of Turnera sidoides L. (Passifloraceae) revealed by morphological and genetic evidence. Plant Syst. Evol. 2015;301:883-892.Pershina L.A., Numerova O.M., Belova L.I., Devyatkina E.P. Biotechnological and cytogenetic aspects of producing new wheat genotypes using hybrids. Euphytica. 1998;1006(1-3):239-244.Nakano N., Mizuno N., Tosa Y., Yoshida K., Park P., Takumi S. Accelerated senescence and enhanced disease resistance in hybrid chlorosis lines derived from interspecific crosses between tetraploid wheat and Aegilops tauschii. PLoS ONE. 2015;10. DOI 10.1371/journal.pone.0121583.Pershina L.A., Shumny V.K. Simulating processes of species formation and rise of biodiversity on the base of wide hybridization of cereals. Biodiversity and dynamics of ecosystems in North Eurasia. V. 1. Basic problems of species and ecosystems evolution. Part 2. Biodiversity and dynamics of ecosystems in North Eurasia: Informational Technologies and Modelling. Novosibirsk: ICG. 2000;88-91.Ng D.W.-K., Lu J., Chen Z.J. Big roles for small RNAs in polyploidy, hybrid vigor, and hybrid incompatibility. Curr. Opin. Plant Biol. 2012;15:154-161.Pershina L.A., Trubacheeva N.V., Sinyavskaya M.G., Devyatkina E.P., Kravtsova L.A. Nuclear-cytoplasmic compatibility and the state of mitochondrial and chloroplast DNA regions in alloplasmic recombinant and introgressive lines (H. vulgare)–T. aestivum. Russ. J. Genet. 2014;50(10):1017-1024.Niroula R.K., Bimb H.P. Overview of wheat × maize system of crosses for dihaploid induction in wheat. World Appl. Sci. J. 2009;7: 1037-1045.Pires J.C., Lim K.Y., Kovarík A., Matyásek R., Boyd A., Leitch A.R., Leitch I.J., Bennett M.D., Soltis P.S., Soltis D.E. Molecular cytogenetic analysis of recently evolved Tragopogon (Asteraceae) allopolyploids reveal a karyotype that is additive of the diploid progenitors. Am. J. Bot. 2004;91:1022-1035.Niu Z., Jiang A., Hammad W.A., Oladzadabbasabadi A., Xu S.S., Mergoum M., Elias E.M. Review of doubled haploid production in durum and common wheat through wheat × maize hybridization. Plant Breeding. 2014;133:313-320.Prakash S., Ahuja I., Upreti C., Kumar V.D., Bhat S.R., Kirti P.B., Chopra V.L. Expression of male sterility in alloplasmic Brassica juncea with Erucastrum canariense cytoplasm and the development of a fertility restoration system. Plant Breeding. 2001;120:479-482.Pathania A., Bhat S.R., Dinesh Kumar V., Ashutosh K.P.B., Prakash S., Chopra V.L. Cytoplasmic male sterility in alloplasmic Brassica juncea carrying Diplotaxis catholica cytoplasm: molecular characterization and genetics of fertility restoration. Theor. Appl. Genet. 2003;107:455-461.Pukhalskiy V.A., Bilinskaya E.A., Martynov S.P., Dobrotvorskaya T.V., Obolenkova G.A. New data on the distribution of hybrid necrosis genes in winter bread wheat (Triticum aestivum L.) cultivars. Russ. J. Genet. 2008;44(2):171-179.Pelletier G., Budar F. The molecular biology of cytoplasmically inherited male sterility and prospects for its engineering. Curr. Opin. Biotechnol. 2007;18:121-125.Pukhalskiy V.A., Martynov S.P., Dobrotvorskaya T.V. Geny gibridnogo nekroza pshenits (teoriya voprosa i katalog nositeley letalnykh genov) [Hybrid Necrosis Genes in Wheats (Theory of Matter and Catalog of Lethal Genes Carriers)]. Moscow, MSHA Publ., 2002.Pershina L.A., Numerova O.M., Belova L.I., Devyatkina E.P. Biotechnological and cytogenetic aspects of producing new wheat genotypes using hybrids. Euphytica. 1998;1006(1-3):239-244.Riera-Lizarazu O., Rines H.W., Phillips R.L. Cytological and molecular characterization of out × maize partial hybrids. Theor. Appl. Genet. 1996;93:123-135.Pershina L.A., Shumny V.K. Simulating processes of species formation and rise of biodiversity on the base of wide hybridization of cereals. Biodiversity and dynamics of ecosystems in North Eurasia. V. 1. Basic problems of species and ecosystems evolution. Part 2. Biodiversity and dynamics of ecosystems in North Eurasia: Informational Technologies and Modelling. Novosibirsk: ICG. 2000;88-91.Rieseberg L.H., Carney S.E. Plant hybridization. New Phytol. 1998; 140:599-624.Pires J.C., Lim K.Y., Kovarík A., Matyásek R., Boyd A., Leitch A.R., Leitch I.J., Bennett M.D., Soltis P.S., Soltis D.E. Molecular cytogenetic analysis of recently evolved Tragopogon (Asteraceae) allopolyploids reveal a karyotype that is additive of the diploid progenitors. Am. J. Bot. 2004;91:1022-1035.Rieseberg L.H., Wendel J.F. Introgression and its consequences in plants. Ed. R.G. Harrison. Hybrid Zones and the Evolutionary Process. Oxford Univ. Press, 1993;70-109.Prakash S., Ahuja I., Upreti C., Kumar V.D., Bhat S.R., Kirti P.B., Chopra V.L. Expression of male sterility in alloplasmic Brassica juncea with Erucastrum canariense cytoplasm and the development of a fertility restoration system. Plant Breeding. 2001;120:479-482.Rieseberg L.H., Wills J.H. Plant speciation. Science. 2007;317:910-913.Riera-Lizarazu O., Rines H.W., Phillips R.L. Cytological and molecular characterization of out × maize partial hybrids. Theor. Appl. Genet. 1996;93:123-135.Sanei M., Pickering R., Kumke K., Nasuda S., Houben A. Loss of centromeric histone H3 (CENH3) from centromeres precedes uniparental chromosome elimination in interspecific barley hybrids. Proc. Natl Acad. Sci. USA. 2011;108:498-505.Rieseberg L.H., Carney S.E. Plant hybridization. New Phytol. 1998; 140:599-624.Seehausen O. Hybridization and adaptive radiation. Trends Ecol. Evol. 2004;19:198- 207.Rieseberg L.H., Wendel J.F. Introgression and its consequences in plants. Ed. R.G. Harrison. Hybrid Zones and the Evolutionary Process. Oxford Univ. Press, 1993;70-109.Sharma G., Srivalli B., Khanna-Chopra R. Hybrid necrosis in wheat – a genetic system showing reduced capacity to detoxify reactive oxygen species leading to programmed cell death. Indian J. Biotechnol. 2003;2(1):17-25.Rieseberg L.H., Wills J.H. Plant speciation. Science. 2007;317:910-913.Shinada T., Kikuchi Y., Fujimoto R., Kishitani S. An alloplasmic malesterile line of Brassica oleracea harboring the mitochondria from Diplotaxis muralis expresses a novel chimeric open reading frame, orf72. Plant Cell Physiol. 2006;47:549-553.Sanei M., Pickering R., Kumke K., Nasuda S., Houben A. Loss of centromeric histone H3 (CENH3) from centromeres precedes uniparental chromosome elimination in interspecific barley hybrids. Proc. Natl Acad. Sci. USA. 2011;108:498-505.Shulyndin A.F. Geneticheskie zakonomernosti rasshchepleniya otdalennykh gibridov rasteniy [Genetic regularity of remote hybrid segregation]. Problemy otdalennoy gibridizatsii rasteniy [Problems of Remote Plant Hybridization]. Sofia, Bolgarskaya Akademiya nauk, 1978:33-44.Seehausen O. Hybridization and adaptive radiation. Trends Ecol. Evol. 2004;19:198-207.Silkova O.G., Shchapova A.I., Shumny V.K. Patterns of meiosis in ABDR amphihaploids depend on the specific type of univalent chromosome division. Euphytica; 2011;178:415-426.Sharma G., Srivalli B., Khanna-Chopra R. Hybrid necrosis in wheat – a genetic system showing reduced capacity to detoxify reactive oxygen species leading to programmed cell death. Indian J. Biotechnol. 2003;2(1):17-25.Sinha P., Tomar S.M.S., Vinod Singh V.K., Balyan H.S. Genetic analysis and molecular mapping of new fertility restorer gene Rf8 for Triticum timopheevii cytoplasm in wheat (Triticum aestivum L.) using SSR markers. Genetics. 2013;141:431-441.Shinada T., Kikuchi Y., Fujimoto R., Kishitani S. An alloplasmic malesterile line of Brassica oleracea harboring the mitochondria from Diplotaxis muralis expresses a novel chimeric open reading frame, orf72. Plant Cell Physiol. 2006;47:549-553.Sobel J.M., Chen G.F., Watt L.R., Schemske D.W. The biology of speciation. Evolution. 2010;64:295-315.Silkova O.G., Shchapova A.I., Shumny V.K. Patterns of meiosis in ABDR amphihaploids depend on the specific type of univalent chromosome division. Euphytica; 2011;178:415-426.Soliman K., Fedak G., Allard R.W. Inheritance of organelle DNA in barley and Hordeum × Secale intergeneric hybrids. Genome. 1987; 29:867-872.Sinha P., Tomar S.M.S., Vinod Singh V.K., Balyan H.S. Genetic analysis and molecular mapping of new fertility restorer gene Rf8 for Triticum timopheevii cytoplasm in wheat (Triticum aestivum L.) using SSR markers. Genetics. 2013;141:431-441.Subrahmanyam N.C., Bothmer R. von. Interspecific hybridization with Hordeum bulbosum and development of hybrids and haploids. Hereditas. 1987;106:119-127.Sobel J.M., Chen G.F., Watt L.R., Schemske D.W. The biology of speciation. Evolution. 2010;64:295-315.Suzuki T., Nakamura C., Mori N., Kaneda C. Overexpression of mitochondrial genes in alloplasmic common wheat with a cytoplasm of wheatgrass (Agropyron trichophorum) showing depressed vigor and male sterility. Plant Mol. Biol. 1995;27:553-565.Soliman K., Fedak G., Allard R.W. Inheritance of organelle DNA in barley and Hordeum × Secale intergeneric hybrids. Genome. 1987; 29:867-872.Tate J.A., Ni Z., Scheen A.-C., Koh J., Gilbert C.A., Lefkowitz D., Chen Z.J., Soltis P.S., Soltis D.E. Evolution and expression of homeologous loci in Tragopogon miscellus (Asteraceae), a recent and reciprocally formed allopolyploid. Genetics. 2006;173:1599-1611.Subrahmanyam N.C., Bothmer R. von. Interspecific hybridization with Hordeum bulbosum and development of hybrids and haploids. Hereditas. 1987;106:119-127.Tikhenko N., Rutten T., Tsvetkova N., Dobrovolskaya O., Zaynali Nezhad K., Roder M.S., Börner A. Hybrid dwarfness in crosses between wheat (Triticum aestivum L.) and rye (Secale cereale L.): a new look at an old phenomenon. Plant Biol. (Stuttg). 2015;17: 320-326.Suzuki T., Nakamura C., Mori N., Kaneda C. Overexpression of mitochondrial genes in alloplasmic common wheat with a cytoplasm of wheatgrass (Agropyron trichophorum) showing depressed vigor and male sterility. Plant Mol. Biol. 1995;27:553-565.Trubacheeva N.V., Efremova T.T., Badaeva E.D., Kravtsova L.A., Belova L.I., Devyatkina E.P., Pershina L.A. Production of alloplasmic and euplasmic wheat-barley ditelosomic substitution lines 7H1Lmar (7D) and analysis of the 18S/5S mitochondrial repeat in these lines. Russ. J. Genet. 2009;45(12):1438-1443.Tate J.A., Ni Z., Scheen A.-C., Koh J., Gilbert C.A., Lefkowitz D., Chen Z.J., Soltis P.S., Soltis D.E. Evolution and expression of homeologous loci in Tragopogon miscellus (Asteraceae), a recent and reciprocally formed allopolyploid. Genetics. 2006;173:1599-1611.Trubacheeva N.V., Kravtsova L.A., Devyatkina E.P., Efremova T.T., Sinyavskaya M.G., Shumny V.K., Pershina L.A. Heteroplasmic and homoplasmic states of mitochondrial and chloroplast DNA regions in progenies of distant common wheat hybrids of different origins. Russ. J. Genet.: Appl. Res. 2012;2(6):494-500.Tikhenko N., Rutten T., Tsvetkova N., Dobrovolskaya O., Zaynali Nezhad K., Roder M.S., Börner A. Hybrid dwarfness in crosses between wheat (Triticum aestivum L.) and rye (Secale cereale L.): a new look at an old phenomenon. Plant Biol. (Stuttg). 2015;17: 320-326.Tsukamoto N., Asakura N., Hattori N., Takumi S., Mori N., Nakamura C. Identification of paternal mitochondrial DNA sequences in the nucleus-cytoplasm hybrid of tetraploid and hexaploid wheat with D and D2 plasmon from Aegilops species. Curr. Genet. 2000;38: 208-221.Tsukamoto N., Asakura N., Hattori N., Takumi S., Mori N., Nakamura C. Identification of paternal mitochondrial DNA sequences in the nucleus-cytoplasm hybrid of tetraploid and hexaploid wheat with D and D2 plasmon from Aegilops species. Curr. Genet. 2000;38: 208-221.Tsunewaki K. Aneuploid analyses of hybrid necrosis and hybrid chlorosis in tetraploid wheats using the D genome chromosome substitution lines of durum wheat. Genome. 1992;35:594-601.Tsunewaki K. Aneuploid analyses of hybrid necrosis and hybrid chlorosis in tetraploid wheats using the D genome chromosome substitution lines of durum wheat. Genome. 1992;35:594- 601.Tsunewaki K. Plasmon analysis as the counterpart of genome analysis. Methods of Genome Analysis in Plant. Ed. P.P. Jauhar. Boca Raton; New York; London; Tokyo: CRC Press, 1996;271-299.Tsunewaki K. Plasmon analysis as the counterpart of genome analysis. Methods of Genome Analysis in Plant. Ed. P.P. Jauhar. Boca Raton; New York; London; Tokyo: CRC Press, 1996;271-299.Tu Y., Sun J., Ge X., Li Z. Chromosome elimination, addition and introgression in intertribal partial hybrids between Brassica rapa and Isatis indigotica. Ann. Bot. 2009;103:1039-1048.Tu Y., Sun J., Ge X., Li Z. Chromosome elimination, addition and introgression in intertribal partial hybrids between Brassica rapa and Isatis indigotica. Ann. Bot. 2009;103:1039-1048.Van Tuyl J.M., Van Diën M.P., Van Creij M.G.M., Van Kleinwee T.C.M., Franken J., Bino R.J. Application of in vitro pollination, ovary culture, ovule culture and embryo rescue for overcoming incongruity barriers in interspecific Lilium crosses. Plant Sci. 1991;74: 115-126.Van Tuyl J.M., Van Diën M.P., Van Creij M.G.M., Van Kleinwee T.C.M., Franken J., Bino R.J. Application of in vitro pollination, ovary culture, ovule culture and embryo rescue for overcoming incongruity barriers in interspecific Lilium crosses. Plant Sci. 1991;74:115-126.Vavilov N.I. Nauchnye osnovy selektsii pshenitsy [Scientific foundations of wheat breeding]. Leningrad, Selhozgiz Publ., 1935.Vinkenoog V., Bushell C., Spielman M., Adams S., Dickinson H.G., Rod J., Scott R.J. Genomic imprinting and endosperm development in flowering plants. Mol. Biotechnol. 2003;25:149-184.Vinkenoog V., Bushell C., Spielman M., Adams S., Dickinson H.G., Rod J., Scott R.J. Genomic imprinting and endosperm development in flowering plants. Mol. Biotechnol. 2003;25:149-184.Vu H.Q., Iwata M., Yamauchi N., Shigyo M. Production of novel alloplasmic male sterile lines in Allium cepa harbouring the cytoplasm from Allium roylei. Plant Breeding. 2011;130:469-475.Vu H.Q., Iwata M., Yamauchi N., Shigyo M. Production of novel alloplasmic male sterile lines in Allium cepa harbouring the cytoplasm from Allium roylei. Plant Breeding. 2011;130:469-475.Wang H., Jiang J., Chen S., Qi X., Fang W., Guan Z., Teng N., Liao Y., Chen F. Rapid genetic and epigenetic alterations under intergeneric genomic shock in newly synthesized Chrysanthemum morifolium × Leucanthemum paludosum hybrids (Asteraceae). Genome Biol. Evol. 2014;6:247-259.Wang H., Jiang J., Chen S., Qi X., Fang W., Guan Z., Teng N., Liao Y., Chen F. Rapid genetic and epigenetic alterations under intergeneric genomic shock in newly synthesized Chrysanthemum morifolium × Leucanthemum paludosum hybrids (Asteraceae). Genome Biol. Evol. 2014;6:247-259.Widmer A., Lexer C., Cozzolino S. Evolution of reproductive isolation in plants. Heredity. 2009;102:31-38.Widmer A., Lexer C., Cozzolino S. Evolution of reproductive isolation in plants. Heredity. 2009;102:31-38.Yang D., Li W., Li S., Yang X., Wu J., Cao Z. In vitro embryo rescue culture of F1 progenies from crosses between diploid and tetraploid grape varieties. Plant Growth Regul. 2007;51:63-71.Yang D., Li W., Li S., Yang X., Wu J., Cao Z. In vitro embryo rescue culture of F1 progenies from crosses between diploid and tetraploid grape varieties. Plant Growth Regul. 2007;51:63-71.Yang J., Zhang M., Yu J. Mitochondrial retrograde regulation tuning fork in nuclear genes expressions of higher plants. J. Genet. Genomics. 2008;35:65-71.Yang J., Zhang M., Yu J. Mitochondrial retrograde regulation tuning fork in nuclear genes expressions of higher plants. J. Genet. Genomics. 2008;35:65-71.Zhou W.M., Yoshida K., Shintaku Y., Takeda G. The use of IAA to overcome interspecific hybrid inviability in reciprocal crosses between Nicotiana tabacum L. and N. repanda Willd. Theor. Appl. Genet. 1991;82:657-661.Zhou W.M., Yoshida K., Shintaku Y., Takeda G. The use of IAA to overcome interspecific hybrid inviability in reciprocal crosses between Nicotiana tabacum L. and N. repanda Willd. Theor. Appl. Genet. 1991;82:657-661.Zhu Y., Saraike T., Yamamoto Y., Hagita H., Takumi S., Murai K. Orf-260cra, a novel mitochondrial gene, is associated with the homeotic transformation of stamens into pistil-like structures (pistillody) in alloplasmic wheat. Plant Cell Physiol. 2008;49:1723-1733.Zhu Y., Saraike T., Yamamoto Y., Hagita H., Takumi S., Murai K. Orf-260cra, a novel mitochondrial gene, is associated with the homeotic transformation of stamens into pistil-like structures (pistillody) in alloplasmic wheat. Plant Cell Physiol. 2008;49:1723-1733.

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