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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/VJ19.481</article-id><article-id custom-type="elpub" pub-id-type="custom">vavilov-1935</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>РЕГУЛЯЦИЯ ГЕНОВ И ГЕНОМОВ</subject></subj-group><subj-group subj-group-type="section-heading" xml:lang="en"><subject>GENOME AND GENE REGULATION</subject></subj-group></article-categories><title-group><article-title>Ген Non3 необходим для формирования  ядрышка у Drosophila</article-title><trans-title-group xml:lang="en"><trans-title>Non3 is an essential Drosophila gene  required for proper nucleolus assembly</trans-title></trans-title-group></title-group><contrib-group><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0001-6997-0492</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Андреева</surname><given-names>Е. Н.</given-names></name><name name-style="western" xml:lang="en"><surname>Andreyeva</surname><given-names>E. N.</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">andreeva@mcb.nsc.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/0000-0002-0896-1899</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Огиенко</surname><given-names>А. А.</given-names></name><name name-style="western" xml:lang="en"><surname>Ogienko</surname><given-names>A. 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"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-9037-6866</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Юшкова</surname><given-names>А. А.</given-names></name><name name-style="western" xml:lang="en"><surname>Yushkova</surname><given-names>A. 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"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0001-8800-8645</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Попова</surname><given-names>Ю. В.</given-names></name><name name-style="western" xml:lang="en"><surname>Popova</surname><given-names>J. V.</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"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0001-7361-5039</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Павлова</surname><given-names>Г. А.</given-names></name><name name-style="western" xml:lang="en"><surname>Pavlova</surname><given-names>G. 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-1"/></contrib><contrib contrib-type="author" corresp="yes"><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Кожевникова</surname><given-names>Е. Н.</given-names></name><name name-style="western" xml:lang="en"><surname>Kozhevnikova</surname><given-names>E. N.</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"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0003-3309-6363</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Иванкин</surname><given-names>А. В.</given-names></name><name name-style="western" xml:lang="en"><surname>Ivankin</surname><given-names>A. V.</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 contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0003-3777-300X</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname> Гатти</surname><given-names>М.</given-names></name><name name-style="western" xml:lang="en"><surname>Gatti</surname><given-names>M.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Новосибирск; Рим.</p></bio><bio xml:lang="en"><p>Novosibirsk; Rome.</p></bio><xref ref-type="aff" rid="aff-4"/></contrib><contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid">https://orcid.org/0000-0001-6959-0641</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Пиндюрин</surname><given-names>А. В.</given-names></name><name name-style="western" xml:lang="en"><surname>Pindyurin</surname><given-names>A. 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">a.pindyurin@mcb.nsc.ru</email><xref ref-type="aff" rid="aff-5"/></contrib></contrib-group><aff-alternatives id="aff-1"><aff xml:lang="ru">Институт молекулярной и клеточной биологии Сибирского отделения Российской академии наук.<country>Россия</country></aff><aff xml:lang="en">Institute of Molecular and Cellular Biology, SB RAS.<country>Russian Federation</country></aff></aff-alternatives><aff-alternatives id="aff-2"><aff xml:lang="ru">Институт молекулярной и клеточной биологии Сибирского отделения Российской академии наук; Новосибирский национальный исследовательский государственный университет.<country>Россия</country></aff><aff xml:lang="en">Institute of Molecular and Cellular Biology, SB RAS;  Novosibirsk State University.<country>Russian Federation</country></aff></aff-alternatives><aff-alternatives id="aff-3"><aff xml:lang="ru">Институт молекулярной и клеточной биологии Сибирского отделения Российской академии наук; Федеральный исследовательский центр Институт цитологии и генетики Сибирского отделения Российской академии наук.<country>Россия</country></aff><aff xml:lang="en">Institute of Molecular and Cellular Biology, SB RAS;  Institute of Cytology and Genetics, SB RAS.<country>Russian Federation</country></aff></aff-alternatives><aff-alternatives id="aff-4"><aff xml:lang="ru">Институт молекулярной и клеточной биологии Сибирского отделения Российской академии наук; Институт молекулярной биологии и патологии Национального исследовательского совета и Отделение биологии и биотехнологии,  Римский университет Ла Сапиенса.<country>Россия</country></aff><aff xml:lang="en">Institute of Molecular and Cellular Biology, SB RAS; IBPM CNR and Department of Biology and Biotechnology, Sapienza University of Rome.<country>Russian Federation</country></aff></aff-alternatives><aff-alternatives id="aff-5"><aff xml:lang="ru">Институт молекулярной и клеточной биологии Сибирского отделения Российской академии наук; Новосибирский национальный исследовательский государственный университет;  Федеральный исследовательский центр Институт цитологии и генетики Сибирского отделения Российской академии наук.<country>Россия</country></aff><aff xml:lang="en">Institute of Molecular and Cellular Biology, SB RAS;  Novosibirsk State University; Institute of Cytology and Genetics, SB RAS.<country>Russian Federation</country></aff></aff-alternatives><pub-date pub-type="collection"><year>2019</year></pub-date><pub-date pub-type="epub"><day>30</day><month>03</month><year>2019</year></pub-date><volume>23</volume><issue>2</issue><fpage>190</fpage><lpage>198</lpage><permissions><copyright-statement>Copyright &amp;#x00A9; Андреева Е.Н., Огиенко А.А., Юшкова А.А., Попова Ю.В., Павлова Г.А., Кожевникова Е.Н., Иванкин А.В.,  Гатти М., Пиндюрин А.В., 2019</copyright-statement><copyright-year>2019</copyright-year><copyright-holder xml:lang="ru">Андреева Е.Н., Огиенко А.А., Юшкова А.А., Попова Ю.В., Павлова Г.А., Кожевникова Е.Н., Иванкин А.В.,  Гатти М., Пиндюрин А.В.</copyright-holder><copyright-holder xml:lang="en">Andreyeva E.N., Ogienko A.A., Yushkova A.A., Popova J.V., Pavlova G.A., Kozhevnikova E.N., Ivankin A.V., Gatti M., Pindyurin A.V.</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/1935">https://vavilov.elpub.ru/jour/article/view/1935</self-uri><abstract><p>Ядрышко представляет собой динамичную немембранную внутриядерную органеллу, которая играет ключевую роль как в биогенезе рибосом, так и в других клеточных процессах. Нарушение функции ядрышка ассоциировано со многими заболеваниями человека, в том числе с дегенеративными патологиями нервной и сердечно-сосудистой систем, а также с образованием злокачественных опухолей. В данной работе нами впервые охарактеризована функция гена Non3 (Novel nucleolar protein 3) у плодовой мушки Drosophila melanogaster, который кодирует гомолог Brix домен-содержащего белка человека Rpf2, участвующего в процессе созревания рибосомной РНК (рРНК). С помощью метода неточной эксцизии Р-элемента мы получили набор из четырех мутаций по гену Non3. Эти мутации формируют аллельный ряд, который включает в себя как жизнеспособные, так и летальные аллели. Non3∆600 – это нуль-аллель, который несет делецию большей части гена и является ранней рецессивной леталью. Мухи генотипа Non3∆600/+ демонстрируют очень слабое уменьшение длины и толщины торакальных щетинок, но при этом развиваются нормально и фертильны. Гетероаллельные комбинации жизнеспособных мутаций гена Non3 (Non3197, Non3310 и Non3259 ) имеют укороченные и тонкие торакальные щетинки, также у них наблюдается некоторое замедление онтогенеза. Такой же паттерн нарушений был ранее описан в литературе как Minute-like фенотип, который характерен при дефектах биогенеза рибосом. Кроме того, мы обнаружили, что белок Non3 является компонентом ядрышка в клетках вентрально-мозгового ганглия личинок третьего возраста и необходим для локализации белка Fibrillarin, важного для пост-трансляционной модификации и процессинга рРНК, в этом немембранном внутриядерном субкомпартменте. Полученный нами набор генетических и биохимических инструментов, использованных в ходе данной работы для первичной характеристики гена Non3, будет полезен также для исследований функции этого гена и его белкового продукта в будущем.</p></abstract><trans-abstract xml:lang="en"><p>The nucleolus is a dynamic non-membrane-bound nuclear organelle, which plays key roles not only in ribosome biogenesis but also in many other cellular processes. Consistent with its multiple functions, the nucleolus has been implicated in many human diseases, including cancer and degenerative pathologies of the nervous system and heart. Here, we report the characterization of the Drosophila Non3 (Novel nucleolar protein 3) gene, which encodes a protein homologous to the human Brix domain-containing Rpf2 that has been shown to control ribosomal RNA (rRNA) processing. We used imprecise P-element excision to generate four new mutant alleles in the Non3 gene. Complementation and phenotypic analyses showed that these Non3 mutations can be arranged in an allelic series that includes both viable and lethal alleles. The strongest lethal allele (Non3∆600) is a genetically null allele that carries a large deletion of the gene and exhibits early lethality when homozygous. Flies heterozygous for Non3∆600 occasionally exhibit a mild reduction in the bristle size, but develop normally and are fertile. However, heteroallelic combinations of viable Non3 mutations (Non3197, Non3310 and Non3259) display a Minute-like phenotype, consisting in delayed development and short and thin bristles, suggesting that they are defective in ribosome biogenesis. We also demonstrate that the Non3 protein localizes to the nucleolus of larval brain cells and it is required for proper nucleolar localization of Fibrillarin, a protein important for post-translational modification and processing of rRNAs. In summary, we generated a number of genetic and biochemical tools that were exploited for an initial characterization of Non3, and will be instrumental for future functional studies on this gene and its protein product.</p></trans-abstract><kwd-group xml:lang="ru"><kwd>Drosophila melanogaster</kwd><kwd>ядрышко</kwd><kwd>фенотип Minute-like</kwd><kwd>Non3</kwd><kwd>Fibrillarin</kwd><kwd>Rpf2</kwd><kwd>Brix домен</kwd><kwd>биогенез рибосом</kwd></kwd-group><kwd-group xml:lang="en"><kwd>Drosophila melanogaster</kwd><kwd>nucleolus</kwd><kwd>Minute-like phenotype</kwd><kwd>Non3</kwd><kwd>Fibrillarin</kwd><kwd>Rpf2</kwd><kwd>Brix domain</kwd><kwd>ribosome biogenesis</kwd></kwd-group></article-meta></front><back><ref-list><title>References</title><ref id="cit1"><label>1</label><citation-alternatives><mixed-citation xml:lang="ru">Biedka S., Micic J., Wilson D., Brown H., Diorio-Toth L., Wool- ford J.L., Jr. Hierarchical recruitment of ribosomal proteins and as-sembly factors remodels nucleolar pre-60S ribosomes. 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