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VOGiS"],"abstract":"<jats:p>\n                    To assess the possibility of integrating extracellular double-stranded DNA fragments into the recipient genome of hematopoietic stem cells, a complex substrate was constructed consisting of the entire M13F-\n                    <jats:italic>AluI<\/jats:italic>\n                    -M13R fragment and its two restrictive derivatives, appearing after hydrolysis with restriction endonucleases EcoRI and HindIII: M13F\n                    <jats:italic>AluI<\/jats:italic>\n                    -EcoRI and M13R-\n                    <jats:italic>AluI<\/jats:italic>\n                    -HindIII. The substrate contained a pBlueScript+ plasmid polylinker sequence, absent in the human genome, which framed the human\n                    <jats:italic>AluI<\/jats:italic>\n                    fragment cloned at the EcoRV site. Human bone marrow cells were treated with the DNA of the constructed complex substrate; taking into account the repair time of pangenomic single-strand breaks, preparations of metaphase plates were obtained. FISH revealed specific fluorescent signals. Simultaneously, DNA isolated from colonies obtained from bone marrow cells treated with a complex substrate was sequenced. Two rounds of sequencing were carried out: whole-genome and selective after targeted hybridization on metal beads. The results obtained indicate that homologous exchange between extrachromosomal and chromosomal DNA is possible. Integration into the genome via the single-strand annealing mechanism, involving microhomologies, is also possible. Intermediates were discovered that suggest the existence of an unusual integration into the genome at the nick of one end of the fragment and the other end of the fragment hanging freely into the interchromosomal space. A direct assessment of the possibility of integrating TAMRA-labeled fragments of fragmented human DNA and\n                    <jats:italic>E. coli<\/jats:italic>\n                    DNA into the genome of recipient cells was carried out using a human bone marrow cell model. The results obtained indicate that specific signals of homologous DNA are distributed throughout the chromosome body (human bone marrow cell model). Signals from nonhomologous\n                    <jats:italic>E. coli<\/jats:italic>\n                    DNA are predominantly concentrated in the centromeric regions of chromosomes. The ratio of the number of obtained reads with integration elements and FISH signals suggested the existence of a strong interaction between extracellular fragments and chromosomal DNA. Experiments have been conducted showing that linear plasmid DNA, after internalization into hematopoietic stem cells, forms a monomer ring. Internalized into the intracellular space, extracellular plasmid DNA is isolated together with chromosomal DNA after stringent purification and fractionation procedures. This fact suggests the existence of a strong ring associate of plasmid DNA and chromosome DNA formed without the participation of a protein framework in the form of a looped chromosomal strand.\n                  <\/jats:p>","DOI":"10.18699\/vjgb-26-18","type":"journal-article","created":{"date-parts":[[2026,4,8]],"date-time":"2026-04-08T17:20:01Z","timestamp":1775668801000},"page":"163-180","source":"Crossref","is-referenced-by-count":1,"title":["Concept of natural genome reconstruction. Part 4. Integration of extracellular double-stranded DNA fragments into the genome of hematopoietic stem cells and the formation of extrachromosomal intermediates"],"prefix":"10.18699","volume":"30","author":[{"given":"S. G.","family":"Oshikhmina","sequence":"first","affiliation":[{"name":"Institute of Cytology and Genetics of the Siberian Branch of the Russian Academy of Sciences;\r\nNovosibirsk State University"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"V. S.","family":"Ruzanova","sequence":"additional","affiliation":[{"name":"Institute of Cytology and Genetics of the Siberian Branch of the Russian Academy of Sciences"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"G. S.","family":"Ritter","sequence":"additional","affiliation":[{"name":"Institute of Cytology and Genetics of the Siberian Branch of the Russian Academy of Sciences"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"E. V.","family":"Dolgova","sequence":"additional","affiliation":[{"name":"Institute of Cytology and Genetics of the Siberian Branch of the Russian Academy of Sciences"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"S. S.","family":"Kirikovich","sequence":"additional","affiliation":[{"name":"Institute of Cytology and Genetics of the Siberian Branch of the Russian Academy of Sciences"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"E. V.","family":"Levites","sequence":"additional","affiliation":[{"name":"Institute of Cytology and Genetics of the Siberian Branch of the Russian Academy of Sciences"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Y. R.","family":"Efremov","sequence":"additional","affiliation":[{"name":"Institute of Cytology and Genetics of the Siberian Branch of the Russian Academy of Sciences"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"T. V.","family":"Karamysheva","sequence":"additional","affiliation":[{"name":"Institute of Cytology and Genetics of the Siberian Branch of the Russian Academy of Sciences"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"A. S.","family":"Molodtseva","sequence":"additional","affiliation":[{"name":"Institute of Molecular and Cellular Biology of the Siberian Branch of the Russian Academy of Sciences"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Y. V.","family":"Raitsina","sequence":"additional","affiliation":[{"name":"Institute of Cytology and Genetics of the Siberian Branch of the Russian Academy of Sciences;\r\nNovosibirsk State University"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"O. S.","family":"Taranov","sequence":"additional","affiliation":[{"name":"State Scientific Center of Virology and Biotechnology \u201cVector\u201d of Rospotrebnadzor"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"S. V.","family":"Sidorov","sequence":"additional","affiliation":[{"name":"Novosibirsk State University;\r\nCity Clinical Hospital No. 1"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"S. D.","family":"Nikonov","sequence":"additional","affiliation":[{"name":"Novosibirsk Tuberculosis Research Institute"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"O. Y.","family":"Leplina","sequence":"additional","affiliation":[{"name":"Research Institute of Fundamental and Clinical Immunology"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"A. A.","family":"Ostanin","sequence":"additional","affiliation":[{"name":"Research Institute of Fundamental and Clinical Immunology"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"E. R.","family":"Chernykh","sequence":"additional","affiliation":[{"name":"Research Institute of Fundamental and Clinical Immunology"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"N. 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Mol Cell. 2001;8(6):1163-1174. doi 10.1016\/S1097-2765(01)00419-1","DOI":"10.1016\/S1097-2765(01)00419-1"},{"key":"ref2","doi-asserted-by":"crossref","unstructured":"Dolgova E.V., Nikolin V.P., Popova N.A., Proskurina A.S., Orishenko K.E., Alyamkina E.A., Efremov Y.R., \u2026 Taranov O.S., Rogachev V.A., Sidorov S.V., Bogachev S.S., Shurdov M.A. Internalization of exogenous DNA into internal compartments of murine bone marrow cells. Russ J Genet Appl Res. 2012;2(6):440-452. doi 10.1134\/S2079059712060056","DOI":"10.1134\/S2079059712060056"},{"key":"ref3","doi-asserted-by":"crossref","unstructured":"Dolgova E.V., Efremov Y.R., Orishchenko K.E., Andrushkevich O.M., Alyamkina E.A., Proskurina A.S., Bayborodin S.I., \u2026 Omigov V.V., Minkevich A.M., Rogachev V.A., Bogachev S.S., Shurdov M.A. Delivery and processing of exogenous double-stranded DNA in mouse CD34+ hematopoietic progenitor cells and their cell cycle changes upon combined treatment with cyclophosphamide and doublestranded DNA. Gene. 2013;528(2):74-83. doi 10.1016\/j.gene.2013.06.058","DOI":"10.1016\/j.gene.2013.06.058"},{"key":"ref4","doi-asserted-by":"crossref","unstructured":"Farzaneh F., Zalin R., Brill D., Shall S. DNA strand breaks and ADP-ribosyl transferase activation during cell differentiation. Nature. 1982; 300(5890):362-366. doi 10.1038\/300362A0","DOI":"10.1038\/300362a0"},{"key":"ref5","unstructured":"Glover D.M. (Ed.) DNA Cloning. A Practical Approach. IRL Press, 1988"},{"key":"ref6","doi-asserted-by":"crossref","unstructured":"Hagan C.R., Sheffield R.F., Rudin C.M. Human Alu element retrotransposition induced by genotoxic stress. Nat Genet. 2003;35(3): 219-220. doi 10.1038\/ng1259","DOI":"10.1038\/ng1259"},{"key":"ref7","doi-asserted-by":"crossref","unstructured":"Hastings P.J., McGill C., Shafer B., Strathern J.N. Ends-in vs. endsout recombination in yeast. Genetics. 1993;135(4):973-980. doi 10.1093\/genetics\/135.4.973","DOI":"10.1093\/genetics\/135.4.973"},{"key":"ref8","doi-asserted-by":"crossref","unstructured":"Hastings P.J., Ira G., Lupski J.R. A microhomology-mediated breakinduced replication model for the origin of human copy number variation. PLoS Genet. 2009;5(1):e1000327. doi 10.1371\/journal.pgen.1000327","DOI":"10.1371\/journal.pgen.1000327"},{"key":"ref9","doi-asserted-by":"crossref","unstructured":"Holmgren L., Szeles A., Rajnav\u00f6lgyi E., Folkman J., Klein G., Ernberg I., Falk K.I. Horizontal transfer of DNA by the uptake of apoptotic bodies. Blood. 1999;93(11):3956-3963. doi 10.1182\/blood.V93.11.3956","DOI":"10.1182\/blood.V93.11.3956"},{"key":"ref10","doi-asserted-by":"crossref","unstructured":"Jacobson G.K., Pinon R., Esposito R.E., Esposito M.S. Single-strand scissions of chromosomal DNA during commitment to recombination at meiosis. Proc Natl Acad Sci USA. 1975;72(5):1887-1891. doi 10.1073\/pnas.72.5.1887","DOI":"10.1073\/pnas.72.5.1887"},{"key":"ref11","doi-asserted-by":"crossref","unstructured":"Johnstone A.P., Williams G.T. Role of DNA breaks and ADP-ribosyl transferase activity in eukaryotic differentiation demonstrated in human lymphocytes. Nature. 1982;300(5890):368-370. doi 10.1038\/300368A0","DOI":"10.1038\/300368a0"},{"key":"ref12","doi-asserted-by":"crossref","unstructured":"Kucherlapati R.S., Eves E.M., Song K.Y., Morse B.S., Smithies O. Homologous recombination between plasmids in mammalian cells can be enhanced by treatment of input DNA. Proc Natl Acad Sci USA. 1984;81(10):3153-3157. doi 10.1073\/pnas.81.10.3153","DOI":"10.1073\/pnas.81.10.3153"},{"key":"ref13","doi-asserted-by":"crossref","unstructured":"Kumagai A., Dunphy W.G. Claspin, a novel protein required for the activation of Chk1 during a DNA replication checkpoint response in Xenopus egg extracts. Mol Cell. 2000;6(4):839-849. doi 10.1016\/S1097-2765(05)00092-4","DOI":"10.1016\/S1097-2765(05)00092-4"},{"key":"ref14","doi-asserted-by":"crossref","unstructured":"Leung W., Malkova A., Haber J.E. Gene targeting by linear duplex DNA frequently occurs by assimilation of a single strand that is subject to preferential mismatch correction. Proc Natl Acad Sci USA. 1997;94(13):6851-6856. doi 10.1073\/pnas.94.13.6851","DOI":"10.1073\/pnas.94.13.6851"},{"key":"ref15","doi-asserted-by":"crossref","unstructured":"Li J., Read L.R., Baker M.D. The mechanism of mammalian gene replacement is consistent with the formation of long regions of heteroduplex DNA associated with two crossing-over events. Mol Cell Biol. 2001;21(2):501-510. doi 10.1128\/mcb.21.2.501-510.2001","DOI":"10.1128\/MCB.21.2.501-510.2001"},{"key":"ref16","unstructured":"Likhacheva A.S., Rogachev V.A., Nikolin V.P., Popova N.A., Shilov A.G., SebelevaT.E., Strunkin D.N., Chernykh E.R., Gel\u2019fgat E.L., Bogachev S.S., Shurdov M.A. Involvement of exogenous DNA in the molecular processes in somatic cell. Informatsionnyy Vestnik VOGiS = Herald Vavilov Soc Genet Breed Sci. 2008;12(3):426-447 (in Russian)"},{"key":"ref17","doi-asserted-by":"crossref","unstructured":"Lin J., Krishnaraj R., Kemp R.G. Exogenous ATP enhances calcium influx in intact thymocytes. J Immunol. 1985;135(5):3403-3410","DOI":"10.4049\/jimmunol.135.5.3403"},{"key":"ref18","doi-asserted-by":"crossref","unstructured":"Lin Y., Waldman A.S. Capture of DNA sequences at double-strand breaks in mammalian chromosomes. Genetics. 2001;158(4):1665-1674. doi 10.1093\/genetics\/158.4.1665","DOI":"10.1093\/genetics\/158.4.1665"},{"key":"ref19","doi-asserted-by":"crossref","unstructured":"MacDougall C.A., Byun T.S., Van C., Yee M.C., Cimprich K.A. The structural determinants of checkpoint activation. Genes Dev. 2007; 21(8):898-903. doi 10.1101\/gad.1522607","DOI":"10.1101\/gad.1522607"},{"key":"ref20","doi-asserted-by":"crossref","unstructured":"Maizels N., Davis L. Initiation of homologous recombination at DNA nicks. Nucleic Acids Res. 2018;46(14):6962-6973. doi 10.1093\/nar\/gky588","DOI":"10.1093\/nar\/gky588"},{"key":"ref21","unstructured":"Maniatis T., Fritch E., Sambrook D. Methods of Genetic Engineering. Molecular Cloning. Moscow: Mir Publ., 1984 (in Russian)"},{"key":"ref22","doi-asserted-by":"crossref","unstructured":"Maricic T., Whitten M., P\u00e4\u00e4bo S. Multiplexed DNA sequence capture of mitochondrial genomes using PCR products. PLoS One. 2010;5(11):e14004. doi 10.1371\/journal.pone.0014004","DOI":"10.1371\/journal.pone.0014004"},{"key":"ref23","doi-asserted-by":"crossref","unstructured":"McVey M., Lee S.E. MMEJ repair of double-strand breaks (director\u2019s cut): deleted sequences and alternative endings. Trends Genet. 2008; 24(11):529-538. doi 10.1016\/j.tig.2008.08.007","DOI":"10.1016\/j.tig.2008.08.007"},{"key":"ref24","doi-asserted-by":"crossref","unstructured":"M\u00f8ller H.D., Mohiyuddin M., Prada-Luengo I., Sailani M.R., Halling J.F., Plomgaard P., Maretty L., Hansen A.J., Snyder M.P., Pilegaard H., Lam H.Y.K., Regenberg B. Circular DNA elements of chromosomal origin are common in healthy human somatic tissue. Nat Commun. 2018;9(1):1069. doi 10.1038\/S41467-018-03369-8","DOI":"10.1038\/s41467-018-03369-8"},{"key":"ref25","doi-asserted-by":"crossref","unstructured":"Murnane J.P., Yezzi M.J., Young B.R. Recombination events during integration of transfected DNA into normal human cells. Nucleic Acids Res. 1990;18(9):2733-2738. doi 10.1093\/nar\/18.9.2733","DOI":"10.1093\/nar\/18.9.2733"},{"key":"ref26","doi-asserted-by":"crossref","unstructured":"Pecorino L.T., Verhaak R.G.W., Henssen A., Mischel P.S. Extrachromosomal DNA (ecDNA): an origin of tumor heterogeneity, genomic remodeling, and drug resistance. Biochem Soc Trans. 2022;50(6): 1911-1920. doi 10.1042\/BST20221045","DOI":"10.1042\/BST20221045"},{"key":"ref27","doi-asserted-by":"crossref","unstructured":"Perucho M., Hanahan D., Wigler M. Genetic and physical linkage of exogenous sequences in transformed cells. Cell. 1980;22(1):309-317. doi 10.1016\/0092-8674(80)90178-6","DOI":"10.1016\/0092-8674(80)90178-6"},{"key":"ref28","doi-asserted-by":"crossref","unstructured":"Perucho M., Wigler M. Linkage and expression of foreign DNA in cultured animal cells. Cold Spring Harb Symp Quant Biol. 1981; 45(Pt.2):829-838. doi 10.1101\/sqb.1981.045.01.101","DOI":"10.1101\/SQB.1981.045.01.101"},{"key":"ref29","doi-asserted-by":"crossref","unstructured":"Petrova D.D., Dolgova E.V., Proskurina A.S., Ritter G.S., Ruzanova V.S., Efremov Y.R., Potter E.A., Kirikovich S.S., Levites E.V., Taranov O.S., Ostanin A.A., Chernykh E.R., Kolchanov N.A., Bogachev S.S. The new general biological property of stem-like tumor cells (Part II: surface molecules, which belongs to distinctive groups with particular functions, form a unique pattern characteristic of a certain type of tumor stem-like cells). Int J Mol Sci. 2022; 23(24):15800. doi 10.3390\/ijms232415800","DOI":"10.3390\/ijms232415800"},{"key":"ref30","doi-asserted-by":"crossref","unstructured":"Pierandrei S., Luchetti A., Sanchez M., Novelli G., Sangiuolo F., Lucarelli M. The gene targeting approach of small fragment homologous replacement (SFHR) alters the expression patterns of DNA repair and cell cycle control genes. Mol Ther Nucleic Acids. 2016; 5(4):e304. doi 10.1038\/mtna.2016.2","DOI":"10.1038\/mtna.2016.2"},{"key":"ref31","doi-asserted-by":"crossref","unstructured":"Potter E.A., Dolgova E.V., Proskurina A.S., Efremov Y.R., Minkevich A.M., Rozanov A.S., Peltek S.E., \u2026 Baiborodin S.I., Ostanin A.A., Chernykh E.R., Kolchanov N.A., Bogachev S.S. Gene expression profiling of tumor-initiating stem cells from mouse Krebs-2 carcinoma using a novel marker of poorly differentiated cells. Oncotarget. 2017;8(6):9425-9441. doi 10.18632\/oncotarget.14116","DOI":"10.18632\/oncotarget.14116"},{"key":"ref32","doi-asserted-by":"crossref","unstructured":"Potter E.A., Dolgova E.V., Proskurina A.S., Ruzanova V.S., Efremov Y.R., Kirikovich S.S., Oshikhmina S.G., \u2026 Grivtsova L.U., Kolchanov N.A., Ostanin A.A., Chernykh E.R., Bogachev S.S. Stimulation of mouse hematopoietic stem cells by angiogenin and DNA preparations. Braz J Med Biol Res. 2024;57:e13072. doi 10.1590\/1414-431X2024e13072","DOI":"10.1590\/1414-431x2024e13072"},{"key":"ref33","doi-asserted-by":"crossref","unstructured":"Ritter G.S., Dolgova E.V., Petrova D.D., Efremov Y.R., Proskurina A.S., Potter E.A., Ruzanova V.S., Kirikovich S.S., Levites E.V., Taranov O.S., Ostanin A.A., Chernykh E.R., Kolchanov N.A., Bogachev S.S. The new general biological property of stem-like tumor cells. Part I. Peculiarities of the process of the double-stranded DNA fragments internalization into stem-like tumor cells. Front Genet. 2022;13:954395. doi 10.3389\/fgene.2022.954395","DOI":"10.3389\/fgene.2022.954395"},{"key":"ref34","doi-asserted-by":"crossref","unstructured":"Rogachev V.A., Likhacheva A., Vratskikh O., Mechetina L.V., Sebeleva T.E., Bogachev S.S., Yakubov L.A., Shurdov M.A. Qualitative and quantitative characteristics of the extracellular DNA delivered to the nucleus of a living cell. Cancer Cell Int. 2006;6:23. doi 10.1186\/1475-2867-6-23","DOI":"10.1186\/1475-2867-6-23"},{"key":"ref35","doi-asserted-by":"crossref","unstructured":"Ruzanova V., Proskurina A., Efremov Y., Kirikovich S., Ritter G., Levites E., Dolgova E., Potter E., Babaeva O., Sidorov S., Taranov O., Ostanin A., Chernykh E., Bogachev S. Chronometric administration of cyclophosphamide and a double-stranded DNA-mix at interstrand crosslinks repair timing, called \u201cKaranahan\u201d therapy, is highly efficient in a weakly immunogenic Lewis carcinoma model. Pathol Oncol Res. 2022;28:1610180. doi 10.3389\/pore.2022.1610180","DOI":"10.3389\/pore.2022.1610180"},{"key":"ref36","doi-asserted-by":"crossref","unstructured":"Ruzanova V.S., Oshikhmina S.G., Proskurina A.S., Ritter G.S., Kirikovich S.S., Levites E.V., Efremov Y.R., Karamysheva T.V., Meschaninova M.I., Mamaev A.L., Taranov O.S., Bogachev A.S., Sidorov S.V., Nikonov S.D., Leplina O.Y., Ostanin A.A., Chernykh E.R., Kolchanov N.A., Dolgova E.V., Bogachev S.S. A concept of natural genome reconstruction. Part 2. Effect of extracellular double-stranded DNA fragments on hematopoietic stem cells. Vavilovskii Zhurnal Genetiki i Selektsii = Vavilov J Genet Breed. 2024;28(8):993-1007. doi 10.18699\/vjgb-24-106","DOI":"10.18699\/vjgb-24-106"},{"key":"ref37","doi-asserted-by":"crossref","unstructured":"Smith A.J.H., Berg P. Homologous recombination between defective neo genes in mouse 3T6 cells. Cold Spring Harb Symp Quant Biol. 1984;49:171-181. doi 10.1101\/sqb.1984.049.01.020","DOI":"10.1101\/SQB.1984.049.01.020"},{"key":"ref38","doi-asserted-by":"crossref","unstructured":"Smithies O., Gregg R.G., Boggs S.S., Koralewski M.A., Kucherlapati R.S. Insertion of DNA sequences into the human chromosomal beta-globin locus by homologous recombination. Nature. 1985; 317(6034):230-234. doi 10.1038\/317230A0","DOI":"10.1038\/317230a0"},{"key":"ref39","doi-asserted-by":"crossref","unstructured":"Spriggs A.I., Boddington M.M., Clarke C.M. Chromosomes of human cancer cells. Br Med J. 1962;2(5317):1431-1435. doi 10.1136\/bmj.2.5317.1431","DOI":"10.1136\/bmj.2.5317.1431"},{"key":"ref40","doi-asserted-by":"crossref","unstructured":"Thomas K.R., Folger K.R., Capecchi M.R. High frequency targeting of genes to specific sites in the mammalian genome. Cell. 1986;44(3): 419-428. doi 10.1016\/0092-8674(86)90463-0","DOI":"10.1016\/0092-8674(86)90463-0"},{"key":"ref41","doi-asserted-by":"crossref","unstructured":"Vriend L.E.M., Krawczyk P.M. Nick-initiated homologous recombination: protecting the genome, one strand at a time. DNA Repair. 2017; 50:1-13. doi 10.1016\/j.dnarep.2016.12.005","DOI":"10.1016\/j.dnarep.2016.12.005"},{"key":"ref42","doi-asserted-by":"crossref","unstructured":"Wu S., Turner K.M., Nguyen N., Raviram R., Erb M., Santini J., Luebeck J., \u2026 Furnari F.B., Chang H.Y., Ren B., Bafna V., Mischel P.S. Circular ecDNA promotes accessible chromatin and high oncogene expression. Nature. 2019;575(7784):699-703. doi 10.1038\/S41586-019-1763-5","DOI":"10.1038\/s41586-019-1763-5"},{"key":"ref43","doi-asserted-by":"crossref","unstructured":"Yakubov L.A., Deeva E.A., Zarytova V.F., Ivanova E.M., Ryte A.S., Yurchenko L.V., Vlassov V.V. Mechanism of oligonucleotide uptake by cells: involvement of specific receptors? Proc Natl Acad Sci USA. 1989;86(17):6454-6458. doi 10.1073\/pnas.86.17.6454","DOI":"10.1073\/pnas.86.17.6454"},{"key":"ref44","doi-asserted-by":"crossref","unstructured":"Yoo H.Y., Shevchenko A., Shevchenko A., Dunphy W.G. Mcm2 is a direct substrate of ATM and ATR during DNA damage and DNA replication checkpoint responses. J Biol Chem. 2004;279(51):53353-53364. doi 10.1074\/jbc.M408026200","DOI":"10.1074\/jbc.M408026200"},{"key":"ref45","doi-asserted-by":"crossref","unstructured":"Yoo H.Y., Jeong S.Y., Dunphy W.G. Site-specific phosphorylation of a checkpoint mediator protein controls its responses to different DNA structures. Genes Dev. 2006;20(7):772-783. doi 10.1101\/GAD.1398806","DOI":"10.1101\/gad.1398806"},{"key":"ref46","doi-asserted-by":"crossref","unstructured":"Zhu Y., Gujar A.D., Wong C.H., Tjong H., Ngan C.Y., Gong L., Chen Y.A., \u2026 Yi E., deCarvalho A.C., Ruan Y., Verhaak R.G.W., Wei C.L. Oncogenic extrachromosomal DNA functions as mobile enhancers to globally amplify chromosomal transcription. Cancer Cell. 2021;39(5):694-707.e7. doi 10.1016\/j.ccell.2021.03.006","DOI":"10.1016\/j.ccell.2021.03.006"},{"key":"ref47","doi-asserted-by":"crossref","unstructured":"Zou L. Single- and double-stranded DNA: building a trigger of ATRmediated DNA damage response. Genes Dev. 2007;21(8):879-885. doi 10.1101\/gad.1550307","DOI":"10.1101\/gad.1550307"}],"container-title":["Vavilov Journal of Genetics and Breeding"],"original-title":[],"link":[{"URL":"https:\/\/vavilov.elpub.ru\/jour\/article\/viewFile\/5029\/2068","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,4,8]],"date-time":"2026-04-08T17:20:16Z","timestamp":1775668816000},"score":1,"resource":{"primary":{"URL":"https:\/\/vavilov.elpub.ru\/jour\/article\/view\/5029"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,4,6]]},"references-count":47,"journal-issue":{"issue":"2","published-online":{"date-parts":[[2026,4,6]]}},"URL":"https:\/\/doi.org\/10.18699\/vjgb-26-18","relation":{},"ISSN":["2500-3259"],"issn-type":[{"value":"2500-3259","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,4,6]]}}}