<?xml version="1.0" encoding="UTF-8"?>
<!DOCTYPE article PUBLIC "-//NLM//DTD JATS (Z39.96) Journal Publishing DTD v1.3 20210610//EN" "JATS-journalpublishing1-3.dtd">
<article article-type="research-article" dtd-version="1.3" xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" xml:lang="ru"><front><journal-meta><journal-id journal-id-type="publisher-id">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/VJ16.193</article-id><article-id custom-type="elpub" pub-id-type="custom">vavilov-849</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>Анализ регуляторных последовательностей</subject></subj-group><subj-group subj-group-type="section-heading" xml:lang="en"><subject>Analysis of regulatory sequences</subject></subj-group></article-categories><title-group><article-title>Анализ геномного распределения сайтов связывания транскрипционных факторов GAGA и CNC в развитии Drosophila melanogaster</article-title><trans-title-group xml:lang="en"><trans-title>Analysis of whole-genome binding patterns of GAGA and CNC transcription factors during Drosophila melanogaster development</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>Брусенцов</surname><given-names>И. И.</given-names></name><name name-style="western" xml:lang="en"><surname>Brusentsov</surname><given-names>I. I.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Новосибирск, Россия</p></bio><bio xml:lang="en"><p>Novosibirsk, Russia</p></bio><email xlink:type="simple">brusentsovi@gmail.com</email><xref ref-type="aff" rid="aff-1"/></contrib><contrib contrib-type="author" corresp="yes"><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Карагодин</surname><given-names>Д. А.</given-names></name><name name-style="western" xml:lang="en"><surname>Karagodin</surname><given-names>D. A.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Новосибирск, Россия</p></bio><bio xml:lang="en"><p>Novosibirsk, Russia</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>Баричева</surname><given-names>Э. М.</given-names></name><name name-style="western" xml:lang="en"><surname>Baricheva</surname><given-names>E. M.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Новосибирск, Россия</p></bio><bio xml:lang="en"><p>Novosibirsk, Russia</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>Меркулова</surname><given-names>Т. И.</given-names></name><name name-style="western" xml:lang="en"><surname>Merkulova</surname><given-names>T. I.</given-names></name></name-alternatives><bio xml:lang="ru"><p>Новосибирск, Россия</p></bio><bio xml:lang="en"><p>Novosibirsk, Russia</p></bio><xref ref-type="aff" rid="aff-1"/></contrib></contrib-group><aff-alternatives id="aff-1"><aff xml:lang="ru">Федеральное государственное бюджетное научное учреждение «Федеральный исследовательский центр Институт цитологии и генетики Сибирского отделения Российской академии наук»<country>Россия</country></aff><aff xml:lang="en">Institute of Cytology and Genetics SB RAS<country>Russian Federation</country></aff></aff-alternatives><pub-date pub-type="collection"><year>2016</year></pub-date><pub-date pub-type="epub"><day>01</day><month>02</month><year>2017</year></pub-date><volume>20</volume><issue>6</issue><fpage>762</fpage><lpage>769</lpage><permissions><copyright-statement>Copyright &amp;#x00A9; Брусенцов И.И., Карагодин Д.А., Баричева Э.М., Меркулова Т.И., 2017</copyright-statement><copyright-year>2017</copyright-year><copyright-holder xml:lang="ru">Брусенцов И.И., Карагодин Д.А., Баричева Э.М., Меркулова Т.И.</copyright-holder><copyright-holder xml:lang="en">Brusentsov I.I., Karagodin D.A., Baricheva E.M., Merkulova T.I.</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/849">https://vavilov.elpub.ru/jour/article/view/849</self-uri><abstract><p>На основании доступных данных ChIP-seq и ChIP-chip экспериментов, выполненных с использованием антител к транскрипционным факторам GAGA и CNC, проведен анализ распределения по геному участков связывания этих факторов в эмбриогенезе (возраст эмбрионов 0–12 и 16–24 ч), а также на стадии белой предкуколки дрозофилы. Показано, что основная часть мест связывания GAGA и CNC попадает в промоторные районы и интроны генов, при этом максимальная плотность пиков связывания обоих факторов приходится на область старта транскрипции. Показано также, что у эмбрионов в возрасте как 0–12, так и 16–24 ч развития наблюдается неслучайная колокализация GAGA и CNC, в то время как на стадии белой предкуколки колокализации этих транскрипционных факторов в масштабе генома выявить не удается. Для того чтобы очертить круг генов, для которых возможна совместная регуляция GAGA и CNC, было осуществлено исследование их совместного распределения в аннотированных регуляторных районах (промоторная область и участки, соответствующие 5’-UTR и 3’-UTR мРНК). Оказалось, что совокупности генов, в регуляторных районах которых обнаружено связывание обоих факторов, сильно различаются на разных стадиях. Если у эмбрионов в возрасте 0–12 ч 353 гена характеризуются пересечением пиков ChIP-seq GAGA и CNC, а в возрасте 16–24 ч число таких генов составляет 611, то всего лишь 61 ген является «общим» для обеих стадий. Предполагается, что различные подгруппы генов-мишеней этих факторов регулируются разными сочетаниями изоформ GAGA и CNC, паттерны экспрессии которых изменяются в ходе эмбриогенеза дрозофилы. Функциональный анализ генов, в регуляторных районах которых найдена колокализация GAGA и CNC на всех исследованных стадиях развития, показывает обогащение генами, контролирующими эмбриогенез, развитие нервной системы и крыла дрозофилы.</p></abstract><trans-abstract xml:lang="en"><p>On the basis of available data of ChIP-seq and ChIPchip experiments performed using antibodies against GAGA and CNC transcription factors, genome-wide binding mapping of these factors at hours 0–12 and 16–24 of Drosophila embryogenesis, as well as on white pre-pupae stage, was conducted. It was shown that the bulk of GAGA and CNC binding falls into promoter regions and introns, with the maximal density of peaks in the vicinity of the transcription start site. Moreover, in both 0–12 and 16–24 hour old embryos GAGA and CNC are frequently co-localized, while on white pre-pupae stage there is no co-localization of these factors on a genome–wide scale. In order to select a set of genes potentially co-regulated by GAGA and CNC, the study of their co-binding in annotated regulatory regions (promoter areas and segments corresponding to the 5’-UTR and 3’-UTR of mRNA) was performed. The results obtained clearly demonstrated that the sets of genes characterized by co-binding of both factors vary greatly at different stages. Thus from 353 genes with overlapped GAGA and CNC binding loci on the 0–12 hour old embryos and 611 genes on the 0–12 hour old embryos only 61 genes “belong” to both stages. For an explanation it is proposed that different sets of target genes are regulated by combinations of various GAGA and CNC isoforms, which are characterized by distinct expression patterns during drosophila embryogenesis. Functional annotation analysis of genes, in whose regulatory regions both GAGA and CNC were found at all investigated stages, demonstrates enrichment by genes controlling embryogenesis, neurogenesis and wing development. The data obtained suggest the interaction of GAGA and CNC during D. melanogaster embryogenesis.</p></trans-abstract><kwd-group xml:lang="ru"><kwd>транскрипционные факторы</kwd><kwd>GAGA</kwd><kwd>CNC</kwd><kwd>развитие</kwd><kwd>D. melanogaster</kwd></kwd-group><kwd-group xml:lang="en"><kwd>transcription factors</kwd><kwd>GAGA</kwd><kwd>CNC</kwd><kwd>development</kwd><kwd>D. melanogaster</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">Adryan B., Teichmann S.A. The developmental expression dynamics of Drosophila melanogaster transcription factors. Genome Biol. 2010; 11(4):R40.</mixed-citation><mixed-citation xml:lang="en">Adryan B., Teichmann S.A. The developmental expression dynamics of Drosophila melanogaster transcription factors. Genome Biol. 2010; 11(4):R40.</mixed-citation></citation-alternatives></ref><ref id="cit2"><label>2</label><citation-alternatives><mixed-citation xml:lang="ru">Arnold C.D., Gerlach D., Stelzer C., Boryn L.M., Rath M., Stark A. Genome-wide quantitative enhancer activity maps identified by STARR-seq. Science. 2013;339(6123):1074-1077.</mixed-citation><mixed-citation xml:lang="en">Arnold C.D., Gerlach D., Stelzer C., Boryn L.M., Rath M., Stark A. Genome-wide quantitative enhancer activity maps identified by STARR-seq. Science. 2013;339(6123):1074-1077.</mixed-citation></citation-alternatives></ref><ref id="cit3"><label>3</label><citation-alternatives><mixed-citation xml:lang="ru">Benyajati C., Mueller L., Xu N., Pappano M., Gao J., Mosammaparast M., Conklin D., Granok H., Craig, C., Elgin S. Multiple isoforms of GAGA factor, a critical component of chromatin structure. Nucl. Acids Res. 1997;25(16):3345-3353.</mixed-citation><mixed-citation xml:lang="en">Benyajati C., Mueller L., Xu N., Pappano M., Gao J., Mosammaparast M., Conklin D., Granok H., Craig, C., Elgin S. Multiple isoforms of GAGA factor, a critical component of chromatin structure. Nucl. Acids Res. 1997;25(16):3345-3353.</mixed-citation></citation-alternatives></ref><ref id="cit4"><label>4</label><citation-alternatives><mixed-citation xml:lang="ru">Bhat K.M., Farkas G., Karch F., Gyurkovics H., Gausz J., Schedl P. The GAGA factor is required in the early Drosophila embryo not only for transcriptional regulation but also for nuclear division. Development (Cambridge, England). 1996;122(4):1113-1124.</mixed-citation><mixed-citation xml:lang="en">Bhat K.M., Farkas G., Karch F., Gyurkovics H., Gausz J., Schedl P. The GAGA factor is required in the early Drosophila embryo not only for transcriptional regulation but also for nuclear division. Development (Cambridge, England). 1996;122(4):1113-1124.</mixed-citation></citation-alternatives></ref><ref id="cit5"><label>5</label><citation-alternatives><mixed-citation xml:lang="ru">Bolger A.M., Lohse M., Usadel B. Trimmomatic: a flexible trimmer for Illumina sequence data. Bioinformatics (Oxford, England). 2014; 30(15):2114-2120.</mixed-citation><mixed-citation xml:lang="en">Bolger A.M., Lohse M., Usadel B. Trimmomatic: a flexible trimmer for Illumina sequence data. Bioinformatics (Oxford, England). 2014; 30(15):2114-2120.</mixed-citation></citation-alternatives></ref><ref id="cit6"><label>6</label><citation-alternatives><mixed-citation xml:lang="ru">Bulger M., Groudine M. Functional and mechanistic diversity of distal transcription enhancers. Cell. 2011;144(3):327-339.</mixed-citation><mixed-citation xml:lang="en">Bulger M., Groudine M. Functional and mechanistic diversity of distal transcription enhancers. Cell. 2011;144(3):327-339.</mixed-citation></citation-alternatives></ref><ref id="cit7"><label>7</label><citation-alternatives><mixed-citation xml:lang="ru">Campos-Ortega J.A., Hartenstein V. The Embryonic Development of Drosophila Melanogaster. Berlin; Heidelberg: Springer, 1997.</mixed-citation><mixed-citation xml:lang="en">Campos-Ortega J.A., Hartenstein V. The Embryonic Development of Drosophila Melanogaster. Berlin; Heidelberg: Springer, 1997.</mixed-citation></citation-alternatives></ref><ref id="cit8"><label>8</label><citation-alternatives><mixed-citation xml:lang="ru">Charoensawan V., Wilson D., Teichmann S.A. Genomic repertoires of DNA-binding transcription factors across the tree of life. Nucl. Acids Res. 2010;38(21):7364-7377.</mixed-citation><mixed-citation xml:lang="en">Charoensawan V., Wilson D., Teichmann S.A. Genomic repertoires of DNA-binding transcription factors across the tree of life. Nucl. Acids Res. 2010;38(21):7364-7377.</mixed-citation></citation-alternatives></ref><ref id="cit9"><label>9</label><citation-alternatives><mixed-citation xml:lang="ru">Diehl A.G., Boyle A.P. Deciphering ENCODE. Trends Genet. 2016; 32(4):238-249.</mixed-citation><mixed-citation xml:lang="en">Diehl A.G., Boyle A.P. Deciphering ENCODE. Trends Genet. 2016; 32(4):238-249.</mixed-citation></citation-alternatives></ref><ref id="cit10"><label>10</label><citation-alternatives><mixed-citation xml:lang="ru">Espinas M.L., Jimenez-Garcia E., Vaquero A., Canudas S., Bernues J., Azorin F. The N-terminal POZ domain of GAGA mediates the formation of oligomers that bind DNA with high affinity and specificity. J. Biol. Chemistry. 1999;274:16461-16469.</mixed-citation><mixed-citation xml:lang="en">Espinas M.L., Jimenez-Garcia E., Vaquero A., Canudas S., Bernues J., Azorin F. The N-terminal POZ domain of GAGA mediates the formation of oligomers that bind DNA with high affinity and specificity. J. Biol. Chemistry. 1999;274:16461-16469.</mixed-citation></citation-alternatives></ref><ref id="cit11"><label>11</label><citation-alternatives><mixed-citation xml:lang="ru">Farkas G., Gausz J., Galloni M., Reuter G., Gyurkovics H., Karch F. The Trithorax-like gene encodes the Drosophila GAGA factor. Nature. 1994;371(6500):806-808.</mixed-citation><mixed-citation xml:lang="en">Farkas G., Gausz J., Galloni M., Reuter G., Gyurkovics H., Karch F. The Trithorax-like gene encodes the Drosophila GAGA factor. Nature. 1994;371(6500):806-808.</mixed-citation></citation-alternatives></ref><ref id="cit12"><label>12</label><citation-alternatives><mixed-citation xml:lang="ru">Farnham P.J. Insights from genomic profiling of transcription factors. Nat. Rev. Genet. 2009;10(9):605-616.</mixed-citation><mixed-citation xml:lang="en">Farnham P.J. Insights from genomic profiling of transcription factors. Nat. Rev. Genet. 2009;10(9):605-616.</mixed-citation></citation-alternatives></ref><ref id="cit13"><label>13</label><citation-alternatives><mixed-citation xml:lang="ru">Filion G.J., van Bemmel J.G., Braunschweig U., Talhout W., Kind J., Ward L.D., Brugman W., de Castro I.J., Kerkhoven R.M., Bussemaker H.J., van Steensel B. Systematic protein location mapping reveals five principal chromatin types in Drosophila cells. Cell. 2010; 143(2):212- 224.</mixed-citation><mixed-citation xml:lang="en">Filion G.J., van Bemmel J.G., Braunschweig U., Talhout W., Kind J., Ward L.D., Brugman W., de Castro I.J., Kerkhoven R.M., Bussemaker H.J., van Steensel B. Systematic protein location mapping reveals five principal chromatin types in Drosophila cells. Cell. 2010; 143(2):212- 224.</mixed-citation></citation-alternatives></ref><ref id="cit14"><label>14</label><citation-alternatives><mixed-citation xml:lang="ru">Frith M.C., Valen E., Krogh A., Hayashizaki Y., Carninci P., Sandelin A. A code for transcription initiation in mammalian genomes. Genome Res. 2008;18(1):1-12.</mixed-citation><mixed-citation xml:lang="en">Frith M.C., Valen E., Krogh A., Hayashizaki Y., Carninci P., Sandelin A. A code for transcription initiation in mammalian genomes. Genome Res. 2008;18(1):1-12.</mixed-citation></citation-alternatives></ref><ref id="cit15"><label>15</label><citation-alternatives><mixed-citation xml:lang="ru">Granok H., Leibovitch B.A., Shaffer C.D., Elgin S.C. Chromatin. Gaga over GAGA factor. Curr. Biol. 1995;5(3):238-241.</mixed-citation><mixed-citation xml:lang="en">Granok H., Leibovitch B.A., Shaffer C.D., Elgin S.C. Chromatin. Gaga over GAGA factor. Curr. Biol. 1995;5(3):238-241.</mixed-citation></citation-alternatives></ref><ref id="cit16"><label>16</label><citation-alternatives><mixed-citation xml:lang="ru">Hammonds A.S., Bristow C.A., Fisher W.W., Weiszmann R., Wu S., Hartenstein V., Kellis M., Yu B., Frise E., Celniker S.E. Spatial expression of transcription factors in Drosophila embryonic organ development. Genome Biol. 2013;14(12):R140.</mixed-citation><mixed-citation xml:lang="en">Hammonds A.S., Bristow C.A., Fisher W.W., Weiszmann R., Wu S., Hartenstein V., Kellis M., Yu B., Frise E., Celniker S.E. Spatial expression of transcription factors in Drosophila embryonic organ development. Genome Biol. 2013;14(12):R140.</mixed-citation></citation-alternatives></ref><ref id="cit17"><label>17</label><citation-alternatives><mixed-citation xml:lang="ru">Harbison C.T., Gordon D.B., Lee T.I., Rinaldi N.J., Macisaac K.D., Danford T.W., Hannett N.M., Tagne J.-B., Reynolds D.B., Yoo J., Jennings E.G., Zeitlinger J., Pokholok D.K., Kellis M., Rolfe P.A., Takusagawa K.T., Lander E.S., Gifford D.K., Fraenkel E., Young R.A. Transcriptional regulatory code of a eukaryotic genome. Nature. 2004;431(7004):99-104.</mixed-citation><mixed-citation xml:lang="en">Harbison C.T., Gordon D.B., Lee T.I., Rinaldi N.J., Macisaac K.D., Danford T.W., Hannett N.M., Tagne J.-B., Reynolds D.B., Yoo J., Jennings E.G., Zeitlinger J., Pokholok D.K., Kellis M., Rolfe P.A., Takusagawa K.T., Lander E.S., Gifford D.K., Fraenkel E., Young R.A. Transcriptional regulatory code of a eukaryotic genome. Nature. 2004;431(7004):99-104.</mixed-citation></citation-alternatives></ref><ref id="cit18"><label>18</label><citation-alternatives><mixed-citation xml:lang="ru">Hoskins R.A., Carlson J.W., Wan K.H., Park S., Mendez I., Galle S.E., Booth B.W., Pfeiffer B.D., George R.A., Svirskas R., Krzywinski M., Schein J., Accardo M.C., Damia E., Messina G., Méndez-Lago M., de Pablos B., Demakova O.V., Andreyeva E.N., Boldyreva L.V., Marra M., Carvalho A.B., Dimitri P., Villasante A., Zhimulev I.F., Rubin G.M., Karpen G.H., Celniker S.E. The Release 6 reference sequence of the Drosophila melanogaster genome. Genome Res. 2015;25(3):445-458.</mixed-citation><mixed-citation xml:lang="en">Hoskins R.A., Carlson J.W., Wan K.H., Park S., Mendez I., Galle S.E., Booth B.W., Pfeiffer B.D., George R.A., Svirskas R., Krzywinski M., Schein J., Accardo M.C., Damia E., Messina G., Méndez-Lago M., de Pablos B., Demakova O.V., Andreyeva E.N., Boldyreva L.V., Marra M., Carvalho A.B., Dimitri P., Villasante A., Zhimulev I.F., Rubin G.M., Karpen G.H., Celniker S.E. The Release 6 reference sequence of the Drosophila melanogaster genome. Genome Res. 2015;25(3):445-458.</mixed-citation></citation-alternatives></ref><ref id="cit19"><label>19</label><citation-alternatives><mixed-citation xml:lang="ru">Ingham P.W. The molecular genetics of embryonic pattern formation in Drosophila. Nature. 1988;335(6185):25-34.</mixed-citation><mixed-citation xml:lang="en">Ingham P.W. The molecular genetics of embryonic pattern formation in Drosophila. Nature. 1988;335(6185):25-34.</mixed-citation></citation-alternatives></ref><ref id="cit20"><label>20</label><citation-alternatives><mixed-citation xml:lang="ru">Iyer E.P.R., Iyer S.C., Sullivan L., Wang D., Meduri R., Graybeal L.L., Cox D.N. Functional genomic analyses of two morphologically distinct classes of Drosophila sensory neurons: post-mitotic roles of transcription factors in dendritic patterning. PloS ONE. 2013;8(8):e72434.</mixed-citation><mixed-citation xml:lang="en">Iyer E.P.R., Iyer S.C., Sullivan L., Wang D., Meduri R., Graybeal L.L., Cox D.N. Functional genomic analyses of two morphologically distinct classes of Drosophila sensory neurons: post-mitotic roles of transcription factors in dendritic patterning. PloS ONE. 2013;8(8):e72434.</mixed-citation></citation-alternatives></ref><ref id="cit21"><label>21</label><citation-alternatives><mixed-citation xml:lang="ru">Karagodin D.A., Omelina E.S., Fedorova E.V., Baricheva E.M. Identification of functionally significant elements in the second intron of the Drosophila melanogaster Trithorax-like gene. Gene. 2013;520(2): 178-184.</mixed-citation><mixed-citation xml:lang="en">Karagodin D.A., Omelina E.S., Fedorova E.V., Baricheva E.M. Identification of functionally significant elements in the second intron of the Drosophila melanogaster Trithorax-like gene. Gene. 2013;520(2): 178-184.</mixed-citation></citation-alternatives></ref><ref id="cit22"><label>22</label><citation-alternatives><mixed-citation xml:lang="ru">Kvon E.Z., Stampfel G., Yanez-Cuna J.O., Dickson B.J., Stark A. HOT regions function as patterned developmental enhancers and have a distinct cis-regulatory signature. Genes Develop. 2012;26(9): 908-913.</mixed-citation><mixed-citation xml:lang="en">Kvon E.Z., Stampfel G., Yanez-Cuna J.O., Dickson B.J., Stark A. HOT regions function as patterned developmental enhancers and have a distinct cis-regulatory signature. Genes Develop. 2012;26(9): 908-913.</mixed-citation></citation-alternatives></ref><ref id="cit23"><label>23</label><citation-alternatives><mixed-citation xml:lang="ru">Langmead B., Salzberg S.L. Fast gapped-read alignment with Bowtie 2. Nature Methods. 2012;9(4):357-359.</mixed-citation><mixed-citation xml:lang="en">Langmead B., Salzberg S.L. Fast gapped-read alignment with Bowtie 2. Nature Methods. 2012;9(4):357-359.</mixed-citation></citation-alternatives></ref><ref id="cit24"><label>24</label><citation-alternatives><mixed-citation xml:lang="ru">Lelli K.M., Slattery M., Mann R.S. Disentangling the many layers of eukaryotic transcriptional regulation. Ann. Rev. Genet. 2012;46(1):43-68.</mixed-citation><mixed-citation xml:lang="en">Lelli K.M., Slattery M., Mann R.S. Disentangling the many layers of eukaryotic transcriptional regulation. Ann. Rev. Genet. 2012;46(1):43-68.</mixed-citation></citation-alternatives></ref><ref id="cit25"><label>25</label><citation-alternatives><mixed-citation xml:lang="ru">Li H., Handsaker B., Wysoker A., Fennell T., Ruan J., Homer N., Marth G., Abecasis G., Durbin R. &amp; 1000 Genome Project Data Processing Subgroup The Sequence Alignment/Map format and SAMtools. Bioinformatics (Oxford, England). 2009;25(16):2078-2079.</mixed-citation><mixed-citation xml:lang="en">Li H., Handsaker B., Wysoker A., Fennell T., Ruan J., Homer N., Marth G., Abecasis G., Durbin R. &amp; 1000 Genome Project Data Processing Subgroup The Sequence Alignment/Map format and SAMtools. Bioinformatics (Oxford, England). 2009;25(16):2078-2079.</mixed-citation></citation-alternatives></ref><ref id="cit26"><label>26</label><citation-alternatives><mixed-citation xml:lang="ru">McGinnis N., Ragnhildstveit E., Veraksa A., McGinnis W. A cap “n” collar protein isoform contains a selective Hox repressor function. Development (Cambridge, England). 1998;125(22):4553-4564.</mixed-citation><mixed-citation xml:lang="en">McGinnis N., Ragnhildstveit E., Veraksa A., McGinnis W. A cap “n” collar protein isoform contains a selective Hox repressor function. Development (Cambridge, England). 1998;125(22):4553-4564.</mixed-citation></citation-alternatives></ref><ref id="cit27"><label>27</label><citation-alternatives><mixed-citation xml:lang="ru">Merkulova T.I., Ananko E.A., Ignat’eva E.V., Kolchanov N.A. Regulatory transcription codes in eukaryotic genomes. Genetika = Genetics (Moscow). 2013;49(1):37-54. (in Russian)</mixed-citation><mixed-citation xml:lang="en">Merkulova T.I., Ananko E.A., Ignat’eva E.V., Kolchanov N.A. Regulatory transcription codes in eukaryotic genomes. Genetika = Genetics (Moscow). 2013;49(1):37-54. (in Russian)</mixed-citation></citation-alternatives></ref><ref id="cit28"><label>28</label><citation-alternatives><mixed-citation xml:lang="ru">Mohler J., Mahaffey J.W., Deutsch E., Vani K. Control of Drosophila head segment identity by the bZIP homeotic gene cnc. Development (Cambridge, England). 1995;121(1):237-247.</mixed-citation><mixed-citation xml:lang="en">Mohler J., Mahaffey J.W., Deutsch E., Vani K. Control of Drosophila head segment identity by the bZIP homeotic gene cnc. Development (Cambridge, England). 1995;121(1):237-247.</mixed-citation></citation-alternatives></ref><ref id="cit29"><label>29</label><citation-alternatives><mixed-citation xml:lang="ru">Moorman C., Sun L.V., Wang J., de Wit E., Talhout W., Ward L.D., Greil F., Lu X.-J., White K.P., Bussemaker H.J., van Steensel B. Hotspots of transcription factor colocalization in the genome of Drosophila melanogaster. Proc. Natl. Acad. Sci. 2006;103:12027-12032.</mixed-citation><mixed-citation xml:lang="en">Moorman C., Sun L.V., Wang J., de Wit E., Talhout W., Ward L.D., Greil F., Lu X.-J., White K.P., Bussemaker H.J., van Steensel B. Hotspots of transcription factor colocalization in the genome of Drosophila melanogaster. Proc. Natl. Acad. Sci. 2006;103:12027-12032.</mixed-citation></citation-alternatives></ref><ref id="cit30"><label>30</label><citation-alternatives><mixed-citation xml:lang="ru">Nègre N., Brown C.D., Ma L., Bristow C.A., Miller S.W., Wagner U., … White K.P. A cis- regulatory map of the Drosophila genome. Nature. 2011;471(7339):527-531. http://doi.org/10.1038/nature09990.</mixed-citation><mixed-citation xml:lang="en">Nègre N., Brown C.D., Ma L., Bristow C.A., Miller S.W., Wagner U., … White K.P. A cis- regulatory map of the Drosophila genome. Nature. 2011;471(7339):527-531. http://doi.org/10.1038/nature09990.</mixed-citation></citation-alternatives></ref><ref id="cit31"><label>31</label><citation-alternatives><mixed-citation xml:lang="ru">Nègre N., Brown C.D., Shah P.K., Kheradpour P., Morrison C.A., Henikoff J.G., … White K.P. A comprehensive map of insulator elements for the Drosophila genome. PLoS Genetics. 2010;6(1): e1000814. http://doi.org/10.1371/journal.pgen.1000814.</mixed-citation><mixed-citation xml:lang="en">Nègre N., Brown C.D., Shah P.K., Kheradpour P., Morrison C.A., Henikoff J.G., … White K.P. A comprehensive map of insulator elements for the Drosophila genome. PLoS Genetics. 2010;6(1): e1000814. http://doi.org/10.1371/journal.pgen.1000814.</mixed-citation></citation-alternatives></ref><ref id="cit32"><label>32</label><citation-alternatives><mixed-citation xml:lang="ru">O’Brien T., Wilkins R.C., Giardina C., Lis J.T. Distribution of GAGA protein on Drosophila genes in vivo. Genes Development. 1995;9(9): 1098-1110.</mixed-citation><mixed-citation xml:lang="en">O’Brien T., Wilkins R.C., Giardina C., Lis J.T. Distribution of GAGA protein on Drosophila genes in vivo. Genes Development. 1995;9(9): 1098-1110.</mixed-citation></citation-alternatives></ref><ref id="cit33"><label>33</label><citation-alternatives><mixed-citation xml:lang="ru">Ochoa-Espinosa A., Small S. Developmental mechanisms and cis-regulatory codes.Curr. Opin. Gen. Dev. 2006;16(2):165-170.</mixed-citation><mixed-citation xml:lang="en">Ochoa-Espinosa A., Small S. Developmental mechanisms and cis-regulatory codes.Curr. Opin. Gen. Dev. 2006;16(2):165-170.</mixed-citation></citation-alternatives></ref><ref id="cit34"><label>34</label><citation-alternatives><mixed-citation xml:lang="ru">Pagans S., Ortiz-Lombardia M., Espinas M.L., Bernues J., Azorin F. The Drosophila transcription factor tramtrack (TTK) interacts with Trithorax-like (GAGA) and represses GAGA-mediated activation. Nucl. Acids Res. 2002;30(20):4406-4413.</mixed-citation><mixed-citation xml:lang="en">Pagans S., Ortiz-Lombardia M., Espinas M.L., Bernues J., Azorin F. The Drosophila transcription factor tramtrack (TTK) interacts with Trithorax-like (GAGA) and represses GAGA-mediated activation. Nucl. Acids Res. 2002;30(20):4406-4413.</mixed-citation></citation-alternatives></ref><ref id="cit35"><label>35</label><citation-alternatives><mixed-citation xml:lang="ru">Perelygina L.M., Baricheva E.M., Sebeleva T.E., Kokoza V.A. The evolutionarily conserved gene Nc70F expressed in nerve tissue of Drosophila melanogaster encodes a protein homologous to the mouse delta transcription factor. Genetika = Genetics (Moscow). 1993;29(10):1597-1607.</mixed-citation><mixed-citation xml:lang="en">Perelygina L.M., Baricheva E.M., Sebeleva T.E., Kokoza V.A. The evolutionarily conserved gene Nc70F expressed in nerve tissue of Drosophila melanogaster encodes a protein homologous to the mouse delta transcription factor. Genetika = Genetics (Moscow). 1993;29(10):1597-1607.</mixed-citation></citation-alternatives></ref><ref id="cit36"><label>36</label><citation-alternatives><mixed-citation xml:lang="ru">Rivera-Pomar R., Jãckle H. From gradients to stripes in Drosophila embryogenesis: filling in the gaps. Trends Genet. 1996;12(11):478-483.</mixed-citation><mixed-citation xml:lang="en">Rivera-Pomar R., Jãckle H. From gradients to stripes in Drosophila embryogenesis: filling in the gaps. Trends Genet. 1996;12(11):478-483.</mixed-citation></citation-alternatives></ref><ref id="cit37"><label>37</label><citation-alternatives><mixed-citation xml:lang="ru">Roy S., Ernst J., Kharchenko P.V., Kheradpour P., Negre N., Eaton M.L., Landolin J.M., Bristow C.A., Ma L.J., Lin M.F., … Kellis M. Iden-tification of functional elements and regulatory circuits by Drosophila modENCODE. Science. 2010;330(6012):1787-1797.</mixed-citation><mixed-citation xml:lang="en">Roy S., Ernst J., Kharchenko P.V., Kheradpour P., Negre N., Eaton M.L., Landolin J.M., Bristow C.A., Ma L.J., Lin M.F., … Kellis M. Iden-tification of functional elements and regulatory circuits by Drosophila modENCODE. Science. 2010;330(6012):1787-1797.</mixed-citation></citation-alternatives></ref><ref id="cit38"><label>38</label><citation-alternatives><mixed-citation xml:lang="ru">Schertel C., Albarca M., Rockel-Bauer C., Kelley N.W., Bischof J., Hens K., van Nimwegen E., Basler K., Deplancke B. A large-scale, in vivo transcription factor screen defines bivalent chromatin as a key property of regulatory factors mediating Drosophila wing development. Genome Res. 2015;25(4):514-523.</mixed-citation><mixed-citation xml:lang="en">Schertel C., Albarca M., Rockel-Bauer C., Kelley N.W., Bischof J., Hens K., van Nimwegen E., Basler K., Deplancke B. A large-scale, in vivo transcription factor screen defines bivalent chromatin as a key property of regulatory factors mediating Drosophila wing development. Genome Res. 2015;25(4):514-523.</mixed-citation></citation-alternatives></ref><ref id="cit39"><label>39</label><citation-alternatives><mixed-citation xml:lang="ru">Segal E., Raveh-Sadka T., Schroeder M., Unnerstall U., Gaul U. Predicting expression patterns from regulatory sequence in Drosophila segmentation. Nature. 2008;451(7178):535-540.</mixed-citation><mixed-citation xml:lang="en">Segal E., Raveh-Sadka T., Schroeder M., Unnerstall U., Gaul U. Predicting expression patterns from regulatory sequence in Drosophila segmentation. Nature. 2008;451(7178):535-540.</mixed-citation></citation-alternatives></ref><ref id="cit40"><label>40</label><citation-alternatives><mixed-citation xml:lang="ru">Slattery M., Ma L., Spokony R.F., Arthur R.K., Kheradpour P., Kundaje A., Nègre N., Crofts A., Ptashkin R., Zieba J., Ostapenko A., Suchy S., Victorsen A., Jameel N., Grundstad A.J., Gao W., Moran J.R., Rehm E.J., Grossman R.L., Kellis M., White K.P. Diverse patterns of genomic targeting by transcriptional regulators in Drosophila melanogaster. Genome Res. 2014;24(7):1224-1235.</mixed-citation><mixed-citation xml:lang="en">Slattery M., Ma L., Spokony R.F., Arthur R.K., Kheradpour P., Kundaje A., Nègre N., Crofts A., Ptashkin R., Zieba J., Ostapenko A., Suchy S., Victorsen A., Jameel N., Grundstad A.J., Gao W., Moran J.R., Rehm E.J., Grossman R.L., Kellis M., White K.P. Diverse patterns of genomic targeting by transcriptional regulators in Drosophila melanogaster. Genome Res. 2014;24(7):1224-1235.</mixed-citation></citation-alternatives></ref><ref id="cit41"><label>41</label><citation-alternatives><mixed-citation xml:lang="ru">Soeller W.C., Oh C.E., Kornberg T.B. Isolation of cDNAs encoding the Drosophila GAGA transcription factor. Mol. Cell. Biol. 1993; 13(12):7961-7970.</mixed-citation><mixed-citation xml:lang="en">Soeller W.C., Oh C.E., Kornberg T.B. Isolation of cDNAs encoding the Drosophila GAGA transcription factor. Mol. Cell. Biol. 1993; 13(12):7961-7970.</mixed-citation></citation-alternatives></ref><ref id="cit42"><label>42</label><citation-alternatives><mixed-citation xml:lang="ru">Tomancak P., Ohler U. Mapping the complexity of transcription control in higher eukaryotes. Genome Biol. 2010;11(4):115.</mixed-citation><mixed-citation xml:lang="en">Tomancak P., Ohler U. Mapping the complexity of transcription control in higher eukaryotes. Genome Biol. 2010;11(4):115.</mixed-citation></citation-alternatives></ref><ref id="cit43"><label>43</label><citation-alternatives><mixed-citation xml:lang="ru">Veraksa A., McGinnis N., Li X., Mohler J., McGinnis W. Cap “n” collar B cooperates with a small Maf subunit to specify pharyngeal development and suppress deformed homeotic function in the Drosophila head. Development (Cambridge, England). 2000;127(18): 4023- 4037.</mixed-citation><mixed-citation xml:lang="en">Veraksa A., McGinnis N., Li X., Mohler J., McGinnis W. Cap “n” collar B cooperates with a small Maf subunit to specify pharyngeal development and suppress deformed homeotic function in the Drosophila head. Development (Cambridge, England). 2000;127(18): 4023- 4037.</mixed-citation></citation-alternatives></ref><ref id="cit44"><label>44</label><citation-alternatives><mixed-citation xml:lang="ru">Zhang Y., Liu T., Meyer C.A., Eeckhoute J., Johnson D.S., Bernstein B.E., Nusbaum C., Myers R.M., Brown M., Li W., Liu X.S. Modelbased analysis of ChIP-Seq (MACS). Genome Biol. 2008;9(9):R137.</mixed-citation><mixed-citation xml:lang="en">Zhang Y., Liu T., Meyer C.A., Eeckhoute J., Johnson D.S., Bernstein B.E., Nusbaum C., Myers R.M., Brown M., Li W., Liu X.S. Modelbased analysis of ChIP-Seq (MACS). Genome Biol. 2008;9(9):R137.</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>
