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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/VJ21.026</article-id><article-id custom-type="elpub" pub-id-type="custom">vavilov-2981</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>ГЕНЕТИКА МИКРООРГАНИЗМОВ</subject></subj-group><subj-group subj-group-type="section-heading" xml:lang="en"><subject>MICROBIAL GENETICS</subject></subj-group></article-categories><title-group><article-title>Галофильные бактерии соленых озер и солончаковых почв Прикаспийской низменности (Республика Дагестан) и их биотехнологический потенциал</article-title><trans-title-group xml:lang="en"><trans-title>Halophilic bacteria of salt lakes and saline soils of the Peri-Caspian lowland (Republic of Daghestan) and their biotechnological potential</trans-title></trans-title-group></title-group><contrib-group><contrib contrib-type="author" corresp="yes"><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Халилова</surname><given-names>Э. А.</given-names></name><name name-style="western" xml:lang="en"><surname>Khalilova</surname><given-names>E. A.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Махачкала, Республика Дагестан</p></bio><bio xml:lang="en"><p>Makhachkala, Daghestan</p></bio><email xlink:type="simple">eslanda61@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/0000-0002-7099-873X</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Котенко</surname><given-names>С. Ц.</given-names></name><name name-style="western" xml:lang="en"><surname>Kotenko</surname><given-names>S. T.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Махачкала, Республика Дагестан</p></bio><bio xml:lang="en"><p>Makhachkala, Daghestan</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/0000-0002-9897-1256</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Исламмагомедова</surname><given-names>Э. А.</given-names></name><name name-style="western" xml:lang="en"><surname>Islammagomedova</surname><given-names>E. A.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Махачкала, Республика Дагестан</p></bio><bio xml:lang="en"><p>Makhachkala, Daghestan</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/0000-0003-0888-9580</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Абакарова</surname><given-names>А. А.</given-names></name><name name-style="western" xml:lang="en"><surname>Abakarova</surname><given-names>A. A.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Махачкала, Республика Дагестан</p></bio><bio xml:lang="en"><p>Makhachkala, Daghestan</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/0000-0003-3092-7539</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Черных</surname><given-names>Н. А.</given-names></name><name name-style="western" xml:lang="en"><surname>Chernyh</surname><given-names>N. A.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Москва</p></bio><bio xml:lang="en"><p>Moscow</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/0000-0002-4784-4548</contrib-id><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Аливердиева</surname><given-names>Д. А.</given-names></name><name name-style="western" xml:lang="en"><surname>Aliverdiyeva</surname><given-names>D. A.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Махачкала, Республика Дагестан</p></bio><bio xml:lang="en"><p>Makhachkala, Daghestan</p></bio><xref ref-type="aff" rid="aff-1"/></contrib></contrib-group><aff-alternatives id="aff-1"><aff xml:lang="ru">Прикаспийский институт биологических ресурсов Дагестанского федерального исследовательского центра Российской академии наук<country>Россия</country></aff><aff xml:lang="en">Precaspian Institute of Biological Resources of the Daghestan Federal Research Center of the Russian Academy of Sciences<country>Russian Federation</country></aff></aff-alternatives><aff-alternatives id="aff-2"><aff xml:lang="ru">Институт микробиологии им. С.Н. Виноградского, Федеральный исследовательский центр «Фундаментальные основы биотехнологии» Российской академии наук<country>Россия</country></aff><aff xml:lang="en">Winogradsky Institute of Microbiology, Federal Research Center “Fundamentals of Biotechnology” of the Russian Academy of Sciences<country>Russian Federation</country></aff></aff-alternatives><pub-date pub-type="collection"><year>2021</year></pub-date><pub-date pub-type="epub"><day>29</day><month>04</month><year>2021</year></pub-date><volume>25</volume><issue>2</issue><fpage>224</fpage><lpage>233</lpage><permissions><copyright-statement>Copyright &amp;#x00A9; Халилова Э.А., Котенко С.Ц., Исламмагомедова Э.А., Абакарова А.А., Черных Н.А., Аливердиева Д.А., 2021</copyright-statement><copyright-year>2021</copyright-year><copyright-holder xml:lang="ru">Халилова Э.А., Котенко С.Ц., Исламмагомедова Э.А., Абакарова А.А., Черных Н.А., Аливердиева Д.А.</copyright-holder><copyright-holder xml:lang="en">Khalilova E.A., Kotenko S.T., Islammagomedova E.A., Abakarova A.A., Chernyh N.A., Aliverdiyeva D.A.</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/2981">https://vavilov.elpub.ru/jour/article/view/2981</self-uri><abstract><p>Приведены результаты изучения биоразнообразия и биотехнологического потенциала галофильных микроорганизмов из термального высокоминерализованного Берикейского озера, соленого Тарумовского озера и солончаковых почв Прикаспийской низменности (Республика Дагестан). С использованием микробиологических методов и метода анализа генов 16S рРНК идентифицированы денитрифицирующие галофильные бактерии родов Halomonas и Virgibacillus. Выявлен новый вид Halomonas sp. G2 (MW386470) с 95 % уровнем сходства нуклеотидных последовательностей генов 16S рРНК. Штамм G2 – экстремальный галофил, способный расти в диапазоне 5–25 % NaCl (оптимум 25 %) и образовывать каротиноидный пигмент. Мезофил (30–37 °С, оптимум 30 °С), нейтрофил (рН 6–8, оптимум 7.2–7.4). Хемолитотроф; редуцирует нитрат или нитрит в качестве доноров электронов; каталазо-, амилазо-, протеазо- и β-галактозидазоположительный; липазо-, оксидазо- и уреазоотрицательный. Не способен гидролизовать инозит, индол; продуцирует лизин, желатин, эктоин; в качестве источника углерода и энергии использует цитрат и малат натрия; не продуцирует орнитин, H2S и кислоту из d-маннозы, сахарозы, глицерина, целлобиозы, кроме лактозы и d-глюкозы. Восприимчив к триметоприму, ципрофлоксацину, офлоксацину, канамицину, ванкомицину, рифампицину, цефуроксиму, ампициллину, цефтазидиму, фосфомицину, кларитромицину, цефепиму, цефаклору. Содержание G+C в ДНК 67.3 %. Отличительной характеристикой изолята являлось продуцирование промышленно значимых гидролитических ферментов, таких как амилаза, протеаза, β-галактозидаза и оксиредуктазы – каталазы при концентрации NaCl в среде 25 %. Местообитание: солончаковые почвы на территории Терско-Кумской низменности (Республика Дагестан, Россия). Остальные галофильные изоляты H. ventosae G1 (MW386469), H. elongata G3 (MW386471), V. salinarius B2 (MW386472) и V. salinarius B3 (MW386473) имели высокую степень сходства (100 %) с типовыми штаммами H. elongata DSM 2581Т и V. salarius DSM 18441Т; содержание G+C в ДНК составляло 65.8, 66.5, 42.8 и 37.3 % соответственно. Штаммы имели высокий биотехнологический потенциал при концентрации NaCl в среде 5 и 25 %. Полученные данные расширили представление о разнообразии и экологическом значении денитрифицирующих бактерий в функционировании засушливых экосистем и выявлении штаммов, продуцирующих ферменты промышленного значения.</p></abstract><trans-abstract xml:lang="en"><p>The article presents the results of studying the biodiversity and biotechnological potential of halophilic microorganisms from the thermal highly mineralized Berikey Lake, the salty Lake Tarumovskoye and saline soils of the Peri-Caspian Lowland (Republic of Daghestan). Denitrifying halophilic bacteria of the genus Halomonas and Virgibacillus were identified using microbiological methods and 16S rRNA gene analysis. A new species Halomonas sp. G2 (MW386470) with a similarity of the nucleotide sequences of the 16S rRNA genes is 95 %. Strain G2 is an extreme halophile capable of growing in the range of 5–25 % NaCl (optimum 25 %) and forming a carotenoid pigment. Mesophil, 30–37 °С (optimum 30 °С); neutrophil, pH 6–8 (optimum 7.2–7.4). Strain G2 chemolithotroph; reduces nitrate or nitrite as electron donors; catalase-, amylase-, protease- and β-galactosidase-positive; lipase-, oxidase- and urease-negative. Not able to hydrolyze inositol, indole; produces lysine, gelatin, ectoine; uses citrate and sodium malate as a source of carbon and energy; does not produce ornitin, H2S or acid from d-mannose, sucrose, glycerol, cellobiose, except for lactose and d-glucose. Susceptible to trimethoprim, ciprofloxacin, ofloxacin, kanamycin, vancomycin, rifampicin, cefuroxime, ampicillin, ceftazidime, fosfomycin, clarithromycin, cefepime, cefaclor. The G+C content in DNA is 67.3 %. A distinctive characteristic of the isolate was the production of industrially significant hydrolytic enzymes such as amylase, protease, β-galactosidase, and oxidoreductase (catalase) at a NaCl concentration of 25 % in the medium. Habitat: saline soils on the territory of the Tersko-Kumskaya lowland (Republic of Daghestan, Russia). The rest of the halophilic isolates of H. ventosae G1 (MW386469), H. elongata G3 (MW386471), V. salinarius B2 (MW386472), and V. salinarius B3 (MW386473) had a high degree of similarity (100 %) with the type strains of H. elongata DSM 2581Т and V. salarius DSM 18441Т; the content of G+C in DNA was 65.8, 66.5, 42.8 and 37.3 %, respectively. The strains had a high biotechnological potential at NaCl concentrations of 5 and 25 % in the medium. The data obtained expanded the understanding of the diversity and ecological significance of denitrifying bacteria in the functioning of arid ecosystems and make it possible to identify strains producing enzymes of industrial importance.</p></trans-abstract><kwd-group xml:lang="ru"><kwd>бактерии</kwd><kwd>род Halomonas</kwd><kwd>род Virgibacillus</kwd><kwd>соленые озера</kwd><kwd>почвы</kwd><kwd>солончаки</kwd><kwd>биотехнологический потенциал</kwd></kwd-group><kwd-group xml:lang="en"><kwd>bacteria of the genus Halomonas and Virgibacillus</kwd><kwd>salt lakes</kwd><kwd>salt marshes soils</kwd><kwd>biotechnological potential</kwd></kwd-group><funding-group xml:lang="en"><funding-statement>The work of N.A. Chernykh carried out with the partial financial support of the Ministry of Science and Higher Education of the Russian Federation.</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">Banciu H.L., Enach M., Rodriguez R.M., Oren A., Ventosa A. Ecology and physiology of halophilic microorganisms – Thematic issue based on papers presented at Halophiles 2019 – 12th International Conference on Halophilic Microorganisms, Cluj-Napoca, Romania, 24–28 June, 2019. FEMS Microbiol. Lett. 2020;366(23):1-4. DOI 10.1093/femsle/fnz250.</mixed-citation><mixed-citation xml:lang="en">Banciu H.L., Enach M., Rodriguez R.M., Oren A., Ventosa A. Ecology and physiology of halophilic microorganisms – Thematic issue based on papers presented at Halophiles 2019 – 12th International Conference on Halophilic Microorganisms, Cluj-Napoca, Romania, 24–28 June, 2019. FEMS Microbiol. Lett. 2020;366(23):1-4. DOI 10.1093/femsle/fnz250.</mixed-citation></citation-alternatives></ref><ref id="cit2"><label>2</label><citation-alternatives><mixed-citation xml:lang="ru">Вaumann P., Baumann L. The marine gram-negative Eubacteria. In: Starr M.P., Stolp H., Trüper H.G., Balows A., Schlegel H.G. (Eds.). The Prokaryotes. A handbook on habitats, isolation nd identification of bacteria. Berlin: Springer-Verlag, 1986;2:1302-1331.</mixed-citation><mixed-citation xml:lang="en">Вaumann P., Baumann L. The marine gram-negative Eubacteria. In: Starr M.P., Stolp H., Trüper H.G., Balows A., Schlegel H.G. (Eds.). The Prokaryotes. A handbook on habitats, isolation nd identification of bacteria. Berlin: Springer-Verlag, 1986;2:1302-1331.</mixed-citation></citation-alternatives></ref><ref id="cit3"><label>3</label><citation-alternatives><mixed-citation xml:lang="ru">Begmatov Sh.A., Selitskaya O.V., Vasileva L.V., Berestovskaja Yu.Yu., Manucharova N.A., Drenova N.V. Morphophysiological features of some cultivable bacteria from saline soils of the Aral Sea region. Eur. J. Soil Sci. 2020;53(1):90-96. DOI 10.1134/S1064229320010044.</mixed-citation><mixed-citation xml:lang="en">Begmatov Sh.A., Selitskaya O.V., Vasileva L.V., Berestovskaja Yu.Yu., Manucharova N.A., Drenova N.V. Morphophysiological features of some cultivable bacteria from saline soils of the Aral Sea region. Eur. J. Soil Sci. 2020;53(1):90-96. DOI 10.1134/S1064229320010044.</mixed-citation></citation-alternatives></ref><ref id="cit4"><label>4</label><citation-alternatives><mixed-citation xml:lang="ru">Birnboim H.C., Doly J. A rapid alkaline extraction procedure for screening recombinant plasmid DNA. Nucleic Acids Res. 1979; 7(6):1513-1523.</mixed-citation><mixed-citation xml:lang="en">Birnboim H.C., Doly J. A rapid alkaline extraction procedure for screening recombinant plasmid DNA. Nucleic Acids Res. 1979; 7(6):1513-1523.</mixed-citation></citation-alternatives></ref><ref id="cit5"><label>5</label><citation-alternatives><mixed-citation xml:lang="ru">Bonch-Osmolovskaya E.A., Atomi H. Editorial overview: Extremophiles: from extreme environments to highly stable catalysts. Curr. Opin. Microbiol. 2015;25:88-96.</mixed-citation><mixed-citation xml:lang="en">Bonch-Osmolovskaya E.A., Atomi H. Editorial overview: Extremophiles: from extreme environments to highly stable catalysts. Curr. Opin. Microbiol. 2015;25:88-96.</mixed-citation></citation-alternatives></ref><ref id="cit6"><label>6</label><citation-alternatives><mixed-citation xml:lang="ru">Bulygina E.S., Kuznetsov B.B., Marusina A.I., Kravchenko I.K., Bykova S.A., Kolganova T.V., Galchenko V.F. Study of nucleotide sequences of nif H genes in methanotrophic bacteria. Mikrobiologiya = Microbiology. 2002;71(4):425-432.</mixed-citation><mixed-citation xml:lang="en">Bulygina E.S., Kuznetsov B.B., Marusina A.I., Kravchenko I.K., Bykova S.A., Kolganova T.V., Galchenko V.F. Study of nucleotide sequences of nif H genes in methanotrophic bacteria. Mikrobiologiya = Microbiology. 2002;71(4):425-432.</mixed-citation></citation-alternatives></ref><ref id="cit7"><label>7</label><citation-alternatives><mixed-citation xml:lang="ru">Camacho C., Coulouris G., Avagyan V., Ma N., Papadopoulos J., Bealer K., Madden T.L. BLAST+: architecture and applications. BMC Bioinform. 2009;10(421):1-9. DOI 10.1186/1471-2105-10-421.</mixed-citation><mixed-citation xml:lang="en">Camacho C., Coulouris G., Avagyan V., Ma N., Papadopoulos J., Bealer K., Madden T.L. BLAST+: architecture and applications. BMC Bioinform. 2009;10(421):1-9. DOI 10.1186/1471-2105-10-421.</mixed-citation></citation-alternatives></ref><ref id="cit8"><label>8</label><citation-alternatives><mixed-citation xml:lang="ru">Chernousova E.Yu., Akimov V.N., Gridneva E.V., Dubinina G.A., Grabovich M.Yu. Phylogenetic in situ/ex situ analysis of a sulfur mat microbial community from a thermal sulfide spring in the North Caucasus. Mikrobiologiya = Microbiology. 2008;77(2):219-223. DOI 10.1134/S002626170802015X.</mixed-citation><mixed-citation xml:lang="en">Chernousova E.Yu., Akimov V.N., Gridneva E.V., Dubinina G.A., Grabovich M.Yu. Phylogenetic in situ/ex situ analysis of a sulfur mat microbial community from a thermal sulfide spring in the North Caucasus. Mikrobiologiya = Microbiology. 2008;77(2):219-223. DOI 10.1134/S002626170802015X.</mixed-citation></citation-alternatives></ref><ref id="cit9"><label>9</label><citation-alternatives><mixed-citation xml:lang="ru">Cira-Chávez L.A., Guevara-Luna J., Soto-Padilla M.Y., RománPonce B., Vásquez-Murrieta M.S., Estrada-Alvarado M.I. Kinetics of halophilic enzymes. In: Rajendran L., Fernandez C. (Eds.). Kinetics of Enzymatic Synthesis. IntOpen, 2018;1-25. DOI 10.5772/intechopen.81100.</mixed-citation><mixed-citation xml:lang="en">Cira-Chávez L.A., Guevara-Luna J., Soto-Padilla M.Y., RománPonce B., Vásquez-Murrieta M.S., Estrada-Alvarado M.I. Kinetics of halophilic enzymes. In: Rajendran L., Fernandez C. (Eds.). Kinetics of Enzymatic Synthesis. IntOpen, 2018;1-25. DOI 10.5772/intechopen.81100.</mixed-citation></citation-alternatives></ref><ref id="cit10"><label>10</label><citation-alternatives><mixed-citation xml:lang="ru">Corral P., Amoozegar M.A., Ventosa A. Halophiles and their biomolecules: recent advances and future applications in biomedicine. Marine Drugs. 2020;18:2-33. DOI 10.3390/md18010033.</mixed-citation><mixed-citation xml:lang="en">Corral P., Amoozegar M.A., Ventosa A. Halophiles and their biomolecules: recent advances and future applications in biomedicine. Marine Drugs. 2020;18:2-33. DOI 10.3390/md18010033.</mixed-citation></citation-alternatives></ref><ref id="cit11"><label>11</label><citation-alternatives><mixed-citation xml:lang="ru">Delgado-García M., Valdivia-Urdiales B., Aguilar-Gonzalez C.N., Contreras-Esquivel J.C., Rodriguez-Herrera R. Halophilic hydrolases as a new tool for the biotechnological industries. J. Sci. Food Agric. 2012;92(13):2575-2580. DOI 10.1002/jsfa.5860.</mixed-citation><mixed-citation xml:lang="en">Delgado-García M., Valdivia-Urdiales B., Aguilar-Gonzalez C.N., Contreras-Esquivel J.C., Rodriguez-Herrera R. Halophilic hydrolases as a new tool for the biotechnological industries. J. Sci. Food Agric. 2012;92(13):2575-2580. DOI 10.1002/jsfa.5860.</mixed-citation></citation-alternatives></ref><ref id="cit12"><label>12</label><citation-alternatives><mixed-citation xml:lang="ru">De Lourdes Moreno M., Pérez D., García M.T., Mellado E. Halophilic bacteria as a source of novel hydrolytic enzymes. Life. 2013;3(1): 38-51. DOI 10.3390/life3010038.</mixed-citation><mixed-citation xml:lang="en">De Lourdes Moreno M., Pérez D., García M.T., Mellado E. Halophilic bacteria as a source of novel hydrolytic enzymes. Life. 2013;3(1): 38-51. DOI 10.3390/life3010038.</mixed-citation></citation-alternatives></ref><ref id="cit13"><label>13</label><citation-alternatives><mixed-citation xml:lang="ru">Di Donato P., Buono A., Poli A., Finore I., Abbamondi R.G., Nicolaus B., Lama L. Exploring marine environments for the identification of extremophiles and their enzymes for sustainable and green bioprocesses. Sustainability. 2019;11:149-169. DOI 10.3390/su11010149.</mixed-citation><mixed-citation xml:lang="en">Di Donato P., Buono A., Poli A., Finore I., Abbamondi R.G., Nicolaus B., Lama L. Exploring marine environments for the identification of extremophiles and their enzymes for sustainable and green bioprocesses. Sustainability. 2019;11:149-169. DOI 10.3390/su11010149.</mixed-citation></citation-alternatives></ref><ref id="cit14"><label>14</label><citation-alternatives><mixed-citation xml:lang="ru">Franzmann P.D., Wehmeyer U., Stackebrandt E. Halomonadaceae fam. nov., a new family of the class Proteobacteria to accommodate the genera Halomonas and Deleya. Syst. Appl. Microbiol. 1988;11: 16-19.</mixed-citation><mixed-citation xml:lang="en">Franzmann P.D., Wehmeyer U., Stackebrandt E. Halomonadaceae fam. nov., a new family of the class Proteobacteria to accommodate the genera Halomonas and Deleya. Syst. Appl. Microbiol. 1988;11: 16-19.</mixed-citation></citation-alternatives></ref><ref id="cit15"><label>15</label><citation-alternatives><mixed-citation xml:lang="ru">Ghosh S., Kumar S., Kumar Khare S. Microbial diversity of saline habitats: an overview of biotechnological applications. In: Giri B., Varma A. (Eds.). Microorganisms in Saline Environments: Strategies and Functions (Ser. Soil Biology. 56). Cham: Springer, 2019;65-92. DOI 10.1007/978-3-030-18975-4_4.</mixed-citation><mixed-citation xml:lang="en">Ghosh S., Kumar S., Kumar Khare S. Microbial diversity of saline habitats: an overview of biotechnological applications. In: Giri B., Varma A. (Eds.). Microorganisms in Saline Environments: Strategies and Functions (Ser. Soil Biology. 56). Cham: Springer, 2019;65-92. DOI 10.1007/978-3-030-18975-4_4.</mixed-citation></citation-alternatives></ref><ref id="cit16"><label>16</label><citation-alternatives><mixed-citation xml:lang="ru">Gordon R.E., Smith M.M. Rapidly growing, acid fast bacteria. I. Species descriptions of Mycobacterium phlei Lehmann and Neumann and Mycobacterium smegmatis (Trevisan) Lehmann and Neumann. J. Bacteriol. 1953;66(1):41-48.</mixed-citation><mixed-citation xml:lang="en">Gordon R.E., Smith M.M. Rapidly growing, acid fast bacteria. I. Species descriptions of Mycobacterium phlei Lehmann and Neumann and Mycobacterium smegmatis (Trevisan) Lehmann and Neumann. J. Bacteriol. 1953;66(1):41-48.</mixed-citation></citation-alternatives></ref><ref id="cit17"><label>17</label><citation-alternatives><mixed-citation xml:lang="ru">Gridneva E.V., Grabovich M.Yu., Dubinina G.A., Chernousova E.Yu., Akimov V.N. Ecophysiology of lithotrophic sulfur-oxidizing Sphaerotilus species from sulfide springs in the Northern Caucasus. Mikrobiologiya = Microbiology. 2009;78(1):76-83.</mixed-citation><mixed-citation xml:lang="en">Gridneva E.V., Grabovich M.Yu., Dubinina G.A., Chernousova E.Yu., Akimov V.N. Ecophysiology of lithotrophic sulfur-oxidizing Sphaerotilus species from sulfide springs in the Northern Caucasus. Mikrobiologiya = Microbiology. 2009;78(1):76-83.</mixed-citation></citation-alternatives></ref><ref id="cit18"><label>18</label><citation-alternatives><mixed-citation xml:lang="ru">Heyndrickx M., Lebbe L., Kersters K., De Vos P., Forsyth G., Logan N.A. Virgibacillus: a new genus to accommodate Bacillus pantothenticus (Proom and Knight 1950). Emended description of Virgibacillus pantothenticus. Int. J. Syst. Bacteriol. 1998;48(1):99-106. DOI 10.1099/00207713-48-1-99.</mixed-citation><mixed-citation xml:lang="en">Heyndrickx M., Lebbe L., Kersters K., De Vos P., Forsyth G., Logan N.A. Virgibacillus: a new genus to accommodate Bacillus pantothenticus (Proom and Knight 1950). Emended description of Virgibacillus pantothenticus. Int. J. Syst. Bacteriol. 1998;48(1):99-106. DOI 10.1099/00207713-48-1-99.</mixed-citation></citation-alternatives></ref><ref id="cit19"><label>19</label><citation-alternatives><mixed-citation xml:lang="ru">Heyrman J., Logan N.A., Busse H.-J., Balcaen A., Lebbe L., RodriguezDiaz M., Swings J., De Vos P. Virgibacillus carmonensis sp. nov., Virgibacillus necropolis sp. nov. and Virgibacillus picturae sp. nov., three novel species isolated from deteriorated mural paintings, transfer of the species of the genus Salibacillus to Virgibacillus, as Virgibacillus marismortui comb. nov. and Virgibacillus salexigens comb. nov., and emended description of the genus Virgibacillus. Int. J. Syst. Evol. Microbiol. 2003;53(2):501-511. DOI 10.1099/ijs.0.02371-0.</mixed-citation><mixed-citation xml:lang="en">Heyrman J., Logan N.A., Busse H.-J., Balcaen A., Lebbe L., RodriguezDiaz M., Swings J., De Vos P. Virgibacillus carmonensis sp. nov., Virgibacillus necropolis sp. nov. and Virgibacillus picturae sp. nov., three novel species isolated from deteriorated mural paintings, transfer of the species of the genus Salibacillus to Virgibacillus, as Virgibacillus marismortui comb. nov. and Virgibacillus salexigens comb. nov., and emended description of the genus Virgibacillus. Int. J. Syst. Evol. Microbiol. 2003;53(2):501-511. DOI 10.1099/ijs.0.02371-0.</mixed-citation></citation-alternatives></ref><ref id="cit20"><label>20</label><citation-alternatives><mixed-citation xml:lang="ru">Holt J.R., Corey D.P., Eatock R.A. Mechanoelectrical transduction and adaptation in hair cells of the mouse utricle, a low-frequency vestibular organ. J. Neurosci. 1997;17:8739-8748.</mixed-citation><mixed-citation xml:lang="en">Holt J.R., Corey D.P., Eatock R.A. Mechanoelectrical transduction and adaptation in hair cells of the mouse utricle, a low-frequency vestibular organ. J. Neurosci. 1997;17:8739-8748.</mixed-citation></citation-alternatives></ref><ref id="cit21"><label>21</label><citation-alternatives><mixed-citation xml:lang="ru">Hua N.P., Hamza-Chaffai A., Vreeland R.H., Isoda H., Naganuma T. Virgibacillus salarius sp. nov., a halophilic bacterium isolated from a Saharan salt lake. Int. J. Syst. Evol. Microbiol. 2008;58:2409-2414. DOI 10.1099/ijs.0.65693-0.</mixed-citation><mixed-citation xml:lang="en">Hua N.P., Hamza-Chaffai A., Vreeland R.H., Isoda H., Naganuma T. Virgibacillus salarius sp. nov., a halophilic bacterium isolated from a Saharan salt lake. Int. J. Syst. Evol. Microbiol. 2008;58:2409-2414. DOI 10.1099/ijs.0.65693-0.</mixed-citation></citation-alternatives></ref><ref id="cit22"><label>22</label><citation-alternatives><mixed-citation xml:lang="ru">Kaitouni L.B.D., Anissi J., Sendide K., Hassouni M.E. Diversity of hydrolase-producing halophilic bacteria and evaluation of their enzymatic activities in submerged cultures. Annals Microbiol. 2020; 70:33. DOI 10.1186/s13213-020-01570-z.</mixed-citation><mixed-citation xml:lang="en">Kaitouni L.B.D., Anissi J., Sendide K., Hassouni M.E. Diversity of hydrolase-producing halophilic bacteria and evaluation of their enzymatic activities in submerged cultures. Annals Microbiol. 2020; 70:33. DOI 10.1186/s13213-020-01570-z.</mixed-citation></citation-alternatives></ref><ref id="cit23"><label>23</label><citation-alternatives><mixed-citation xml:lang="ru">Khalilova E.A., Kotenko S.Ts., Islammagomedova E.A., Hasanov R.Z., Abakarova A.A., Aliverdiyeva D.A. Extremophilic microbial communities of saline soils and their diversity in the regions of the Caspian depression. Aridnye Ekosistemy = Arid Ecosystems. 2017;7(2): 116-120. DOI 10.1134/S2079096117020068.</mixed-citation><mixed-citation xml:lang="en">Khalilova E.A., Kotenko S.Ts., Islammagomedova E.A., Hasanov R.Z., Abakarova A.A., Aliverdiyeva D.A. Extremophilic microbial communities of saline soils and their diversity in the regions of the Caspian depression. Aridnye Ekosistemy = Arid Ecosystems. 2017;7(2): 116-120. DOI 10.1134/S2079096117020068.</mixed-citation></citation-alternatives></ref><ref id="cit24"><label>24</label><citation-alternatives><mixed-citation xml:lang="ru">Khalilova E.A., Kotenko S.Ts., Islammagomedova E.A., Hasanov R.Z., Abakarova A.A., Aliverdiyeva D.A. Halophilic microbial communities and their biodiversity in the arid regions of the Caspian Lowland. Aridnye Ekosistemy = Arid Ecosystems. 2020;10(1):79-85. DOI 10.1134/S2079096120010084.</mixed-citation><mixed-citation xml:lang="en">Khalilova E.A., Kotenko S.Ts., Islammagomedova E.A., Hasanov R.Z., Abakarova A.A., Aliverdiyeva D.A. Halophilic microbial communities and their biodiversity in the arid regions of the Caspian Lowland. Aridnye Ekosistemy = Arid Ecosystems. 2020;10(1):79-85. DOI 10.1134/S2079096120010084.</mixed-citation></citation-alternatives></ref><ref id="cit25"><label>25</label><citation-alternatives><mixed-citation xml:lang="ru">Khalilova E.A., Nuratinov R.A., Kotenko S.Ts., Islammagomedova E.A. Hydrocarbon-oxidizing microorganisms of hot springs and their significance in the assessment of the biodiversity of microbial communities. Aridnye Ekosistemy = Arid Ecosystems. 2014;4(1): 25-30. DOI 10.1134/S2079096114010028.</mixed-citation><mixed-citation xml:lang="en">Khalilova E.A., Nuratinov R.A., Kotenko S.Ts., Islammagomedova E.A. Hydrocarbon-oxidizing microorganisms of hot springs and their significance in the assessment of the biodiversity of microbial communities. Aridnye Ekosistemy = Arid Ecosystems. 2014;4(1): 25-30. DOI 10.1134/S2079096114010028.</mixed-citation></citation-alternatives></ref><ref id="cit26"><label>26</label><citation-alternatives><mixed-citation xml:lang="ru">Kindzierski V., Raschke S., Knabe N., Siedler F., Scheffer B., Grau K.P., Pfeiffer F., Oesterhelt D., Marin-Sanguino A., Kunte H.J. Osmoregulation in the halophilic bacterium Halomonas elongata: a case study for integrative systems biology. PLoS One. 2017;12(1):1-22. DOI 10.1371/journal.pone.0168818.</mixed-citation><mixed-citation xml:lang="en">Kindzierski V., Raschke S., Knabe N., Siedler F., Scheffer B., Grau K.P., Pfeiffer F., Oesterhelt D., Marin-Sanguino A., Kunte H.J. Osmoregulation in the halophilic bacterium Halomonas elongata: a case study for integrative systems biology. PLoS One. 2017;12(1):1-22. DOI 10.1371/journal.pone.0168818.</mixed-citation></citation-alternatives></ref><ref id="cit27"><label>27</label><citation-alternatives><mixed-citation xml:lang="ru">Kolganova T.V., Kuznetsov B.B., Turova T.P. Designing and testing of oligonucleotide primers for amplification and sequencing of 16S rRNA genes of archaea. Mikrobiologiya = Microbiology. 2002; 71(2):243-246.</mixed-citation><mixed-citation xml:lang="en">Kolganova T.V., Kuznetsov B.B., Turova T.P. Designing and testing of oligonucleotide primers for amplification and sequencing of 16S rRNA genes of archaea. Mikrobiologiya = Microbiology. 2002; 71(2):243-246.</mixed-citation></citation-alternatives></ref><ref id="cit28"><label>28</label><citation-alternatives><mixed-citation xml:lang="ru">Kushner D.J., Kamekura M. Physiology of halophilic eubacteria. In: Rodríguez-Varela F. (Ed.). Halophilic Bacteria. Boca Raton, FL, USA: CRC Press, 1988;1:109-138.</mixed-citation><mixed-citation xml:lang="en">Kushner D.J., Kamekura M. Physiology of halophilic eubacteria. In: Rodríguez-Varela F. (Ed.). Halophilic Bacteria. Boca Raton, FL, USA: CRC Press, 1988;1:109-138.</mixed-citation></citation-alternatives></ref><ref id="cit29"><label>29</label><citation-alternatives><mixed-citation xml:lang="ru">Kuznetsov A.E., Kalenov S.V. Russian Federation Patent RU 2323226. Method for preparing halophilic bacterium biomass. Bull. No. 12. 2008;6. (in Russian)</mixed-citation><mixed-citation xml:lang="en">Kuznetsov A.E., Kalenov S.V. Russian Federation Patent RU 2323226. Method for preparing halophilic bacterium biomass. Bull. No. 12. 2008;6. (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit30"><label>30</label><citation-alternatives><mixed-citation xml:lang="ru">Lalov V.V., Osokina N.V., Piorunskij D.A., Chizhikov M.A. Russian Federation Patent RU 2115722 C1. Method of halophilic microorganism culturing. 1998;5 (in Russian)</mixed-citation><mixed-citation xml:lang="en">Lalov V.V., Osokina N.V., Piorunskij D.A., Chizhikov M.A. Russian Federation Patent RU 2115722 C1. Method of halophilic microorganism culturing. 1998;5 (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit31"><label>31</label><citation-alternatives><mixed-citation xml:lang="ru">Lane D.J. 16S/23S rRNA sequencing. Nucleic acid techniques in bacterial systematics. In: Stackebrandt E., Goodfellow M. (Eds.). New York: John Wiley &amp; Sons, 1991;115-175.</mixed-citation><mixed-citation xml:lang="en">Lane D.J. 16S/23S rRNA sequencing. Nucleic acid techniques in bacterial systematics. In: Stackebrandt E., Goodfellow M. (Eds.). New York: John Wiley &amp; Sons, 1991;115-175.</mixed-citation></citation-alternatives></ref><ref id="cit32"><label>32</label><citation-alternatives><mixed-citation xml:lang="ru">Lee S.-Y., Kang C.-H., Oh T.K., Yoon J.H. Virgibacillus campisalis sp. nov., from a marine solar saltern. Int. J. Syst. Evol. Microbiol. 2012;62:347-351. DOI 10.1099/ijs.0.033084-0.</mixed-citation><mixed-citation xml:lang="en">Lee S.-Y., Kang C.-H., Oh T.K., Yoon J.H. Virgibacillus campisalis sp. nov., from a marine solar saltern. Int. J. Syst. Evol. Microbiol. 2012;62:347-351. DOI 10.1099/ijs.0.033084-0.</mixed-citation></citation-alternatives></ref><ref id="cit33"><label>33</label><citation-alternatives><mixed-citation xml:lang="ru">Liu C., Baffoe D.K., Zhang M. Halophile, an essential platform for bioproduction. J. Microbiol. Methods. 2019;166:105704. DOI 10.1016/j.mimet.2019.105704.</mixed-citation><mixed-citation xml:lang="en">Liu C., Baffoe D.K., Zhang M. Halophile, an essential platform for bioproduction. J. Microbiol. Methods. 2019;166:105704. DOI 10.1016/j.mimet.2019.105704.</mixed-citation></citation-alternatives></ref><ref id="cit34"><label>34</label><citation-alternatives><mixed-citation xml:lang="ru">Marmur J. A procedure for the isolation of deoxyribonucleic acid from microorganisms. J. Mol. Biol. 1961;3:208-218. DOI 10.1016/S0022-2836(61)80047-8.</mixed-citation><mixed-citation xml:lang="en">Marmur J. A procedure for the isolation of deoxyribonucleic acid from microorganisms. J. Mol. Biol. 1961;3:208-218. DOI 10.1016/S0022-2836(61)80047-8.</mixed-citation></citation-alternatives></ref><ref id="cit35"><label>35</label><citation-alternatives><mixed-citation xml:lang="ru">Namsaraev Z.B., Babasanova O.B., Dunaevsky Y.E., Akimov V.N., Barkhutova D.D., Gorlenko V.M., Namsaraev B.B. Anoxybacillus mongoliensis sp. nov., a novel thermophilic proteinase producing bacterium isolated from alkaline hot spring, Central Mongolia. Microbiology. 2010;79(4):491-499.</mixed-citation><mixed-citation xml:lang="en">Namsaraev Z.B., Babasanova O.B., Dunaevsky Y.E., Akimov V.N., Barkhutova D.D., Gorlenko V.M., Namsaraev B.B. Anoxybacillus mongoliensis sp. nov., a novel thermophilic proteinase producing bacterium isolated from alkaline hot spring, Central Mongolia. Microbiology. 2010;79(4):491-499.</mixed-citation></citation-alternatives></ref><ref id="cit36"><label>36</label><citation-alternatives><mixed-citation xml:lang="ru">Netrusov A.I., Egorova M.A., Zakharchuk L.M. Laboratory Manual on Microbiology for college students. Moscow: Akademiya Publ., 2005. (in Russian)</mixed-citation><mixed-citation xml:lang="en">Netrusov A.I., Egorova M.A., Zakharchuk L.M. Laboratory Manual on Microbiology for college students. Moscow: Akademiya Publ., 2005. (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit37"><label>37</label><citation-alternatives><mixed-citation xml:lang="ru">Owen R.J., Hill L.R., Lapage S.P. Determination of DNA base composition from melting profiles in dilute buffers. Biopolimers. 1969; 7:503-516.</mixed-citation><mixed-citation xml:lang="en">Owen R.J., Hill L.R., Lapage S.P. Determination of DNA base composition from melting profiles in dilute buffers. Biopolimers. 1969; 7:503-516.</mixed-citation></citation-alternatives></ref><ref id="cit38"><label>38</label><citation-alternatives><mixed-citation xml:lang="ru">Sanger F., Nicklen S., Coulson A.R. DNA sequencing with chainterminating inhibitors. Proс. Natl. Acad. Sci. USA. 1977;84:5463-5467.</mixed-citation><mixed-citation xml:lang="en">Sanger F., Nicklen S., Coulson A.R. DNA sequencing with chainterminating inhibitors. Proс. Natl. Acad. Sci. USA. 1977;84:5463-5467.</mixed-citation></citation-alternatives></ref><ref id="cit39"><label>39</label><citation-alternatives><mixed-citation xml:lang="ru">Schwibbert K., Marin-Sanguino A., Bagyan I., Heidrich G., Lentzen G., Seitz H., Rampp M., Schuster S.C., Klenk H.P., Pfeiffer F., Oesterhelt D., Kunte H.J. A blueprint of ectoine metabolism from the genome of the industrial producer Halomonas elongata DSM 2581T. Environ. Microbiol. 2011;13:1973-1994. DOI 10.1111/j.1462-2920.2010.02336.</mixed-citation><mixed-citation xml:lang="en">Schwibbert K., Marin-Sanguino A., Bagyan I., Heidrich G., Lentzen G., Seitz H., Rampp M., Schuster S.C., Klenk H.P., Pfeiffer F., Oesterhelt D., Kunte H.J. A blueprint of ectoine metabolism from the genome of the industrial producer Halomonas elongata DSM 2581T. Environ. Microbiol. 2011;13:1973-1994. DOI 10.1111/j.1462-2920.2010.02336.</mixed-citation></citation-alternatives></ref><ref id="cit40"><label>40</label><citation-alternatives><mixed-citation xml:lang="ru">Steel K.J. The oxidase reaction as a taxonomic tool. J. Gen. Microbiol. 1961;25:297-306.</mixed-citation><mixed-citation xml:lang="en">Steel K.J. The oxidase reaction as a taxonomic tool. J. Gen. Microbiol. 1961;25:297-306.</mixed-citation></citation-alternatives></ref><ref id="cit41"><label>41</label><citation-alternatives><mixed-citation xml:lang="ru">Tamura K., Nei M. Estimation of the number of nucleotide substitutions in the control region of mitochondrial DNA in humans and chimpanzees. Mol. Biol. Evol. 1993;10:512-526.</mixed-citation><mixed-citation xml:lang="en">Tamura K., Nei M. Estimation of the number of nucleotide substitutions in the control region of mitochondrial DNA in humans and chimpanzees. Mol. Biol. Evol. 1993;10:512-526.</mixed-citation></citation-alternatives></ref><ref id="cit42"><label>42</label><citation-alternatives><mixed-citation xml:lang="ru">Tamura K., Stecher G., Peterson D., Filipski A., Kumar S. MEGA6: Molecular Evolutionary Genetics Analysis version 6.0. Mol. Biol. Evol. 2013;30:2725-2729.</mixed-citation><mixed-citation xml:lang="en">Tamura K., Stecher G., Peterson D., Filipski A., Kumar S. MEGA6: Molecular Evolutionary Genetics Analysis version 6.0. Mol. Biol. Evol. 2013;30:2725-2729.</mixed-citation></citation-alternatives></ref><ref id="cit43"><label>43</label><citation-alternatives><mixed-citation xml:lang="ru">Thomas J.C., Khour R., Neeley C.K., Akroush A.M., Davies E.C. A fast CTAB method of human DNA isolation for polymerase chain reaction applications. Biochem. Educ. 1997;25(4):233-235. DOI 10.1016/S0307-4412(97)00122-2.</mixed-citation><mixed-citation xml:lang="en">Thomas J.C., Khour R., Neeley C.K., Akroush A.M., Davies E.C. A fast CTAB method of human DNA isolation for polymerase chain reaction applications. Biochem. Educ. 1997;25(4):233-235. DOI 10.1016/S0307-4412(97)00122-2.</mixed-citation></citation-alternatives></ref><ref id="cit44"><label>44</label><citation-alternatives><mixed-citation xml:lang="ru">Van de Peer Y., De Wachter R. TREECON for Windows: a software package for the construction and drawing of evolutionary trees for the Microsoft Windows environment. Comput. Appl. Biosci. 1994; 10:569-570. DOI 10.1093/bioinformatics/10.5.569.</mixed-citation><mixed-citation xml:lang="en">Van de Peer Y., De Wachter R. TREECON for Windows: a software package for the construction and drawing of evolutionary trees for the Microsoft Windows environment. Comput. Appl. Biosci. 1994; 10:569-570. DOI 10.1093/bioinformatics/10.5.569.</mixed-citation></citation-alternatives></ref><ref id="cit45"><label>45</label><citation-alternatives><mixed-citation xml:lang="ru">Varrella S., Tangherlini M., Corinaldesi C. Deep hypersaline anoxic basins as untapped reservoir of polyextremophilic prokaryotes of biotechnological interest. Mar. Drugs. 2020;18(2):91. DOI 10.3390/md18020091.</mixed-citation><mixed-citation xml:lang="en">Varrella S., Tangherlini M., Corinaldesi C. Deep hypersaline anoxic basins as untapped reservoir of polyextremophilic prokaryotes of biotechnological interest. Mar. Drugs. 2020;18(2):91. DOI 10.3390/md18020091.</mixed-citation></citation-alternatives></ref><ref id="cit46"><label>46</label><citation-alternatives><mixed-citation xml:lang="ru">Vreeland R.H., Litchfield C.D., Martin E.L., Elliot E. Halomonas elongata, a new genus and species of extremely salt-tolerant bacteria. Int. J. Syst. Bacteriol. 1980;3(2):485-495. DOI 10.1099/00207713-30-2-485.</mixed-citation><mixed-citation xml:lang="en">Vreeland R.H., Litchfield C.D., Martin E.L., Elliot E. Halomonas elongata, a new genus and species of extremely salt-tolerant bacteria. Int. J. Syst. Bacteriol. 1980;3(2):485-495. DOI 10.1099/00207713-30-2-485.</mixed-citation></citation-alternatives></ref><ref id="cit47"><label>47</label><citation-alternatives><mixed-citation xml:lang="ru">Wang T., Zhang L., Bo L., Zhu Y., Tang X., Liu W. Simultaneous heterotrophic nitrification and aerobic denitrification at high concentrations of NaCl by Halomonas nacteria. IOP Conf. Ser: Earth. Environ. Sci. 2019;237(5):052033. DOI 10.1088/1755-1315/237/5/052033.</mixed-citation><mixed-citation xml:lang="en">Wang T., Zhang L., Bo L., Zhu Y., Tang X., Liu W. Simultaneous heterotrophic nitrification and aerobic denitrification at high concentrations of NaCl by Halomonas nacteria. IOP Conf. Ser: Earth. Environ. Sci. 2019;237(5):052033. DOI 10.1088/1755-1315/237/5/052033.</mixed-citation></citation-alternatives></ref><ref id="cit48"><label>48</label><citation-alternatives><mixed-citation xml:lang="ru">Zavarzin G.A. Microbial diversity studies at the Winogradsky Institute of Microbiology. Mikrobiologiya = Microbiology. 2004;73(5):509-522. DOI 10.1023/B:MICI.0000044242.93603.00.</mixed-citation><mixed-citation xml:lang="en">Zavarzin G.A. Microbial diversity studies at the Winogradsky Institute of Microbiology. Mikrobiologiya = Microbiology. 2004;73(5):509-522. DOI 10.1023/B:MICI.0000044242.93603.00.</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>
