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«Антипролиферативное действие карнозина и его производных на опухолевые клетки нейрального происхождения ...»

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ADDIN EN.CITE.DATA [19, 217]. Гистидин и имидазол также демонстрировали защитное действие, в то время как -аланин и ГАМК не оказывали какого либо эффекта PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5Lb2hlbjwvQXV0aG9yPjxZZWFyPjE5ODg8L1llYXI+PFJl

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ADDIN EN.CITE.DATA [15, 19].

Одним из проявлений антиоксидантных свойств карнозина является его радиомодифицирующий эффект. Повреждающее действие ионизирующего излучения связано, главным образом, с развитием мощнейшего окислительного стресса в результате радиолиза воды. Пероральное введение мышам карнозина в течение 20 суток перед облучением способствовало повышению выживаемости животных ADDIN EN.CITE <EndNote><Cite><Author>Северин С. Е.</Author><Year>1990</Year><RecNum>167</RecNum><DisplayText>[218]</DisplayText><record><rec-number>167</rec-number><foreign-keys><key app="EN" db-id="w2wfxafe6va9z5eev2l55aa8sxsa599eev2s">167</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author><style face="normal" font="default" charset="204" size="100%">Северин С. Е., Болдырев А. А., Стволинский С. Л., Бордюков, М.М., Гончаренко, Е.Н., Деев, Л.И., Малинина, И.Е., Кудряшов, Ю.Е. </style></author></authors></contributors><titles><title><style face="normal" font="default" charset="204" size="100%">О радиомодифицирующих свойствах карнозина</style></title><secondary-title><style face="normal" font="default" charset="204" size="100%">Радиобиология</style></secondary-title></titles><periodical><full-title>Радиобиология</full-title></periodical><pages><style face="normal" font="default" charset="204" size="100%">765-768</style></pages><volume><style face="normal" font="default" charset="204" size="100%">30</style></volume><number><style face="normal" font="default" charset="204" size="100%">6</style></number><dates><year><style face="normal" font="default" charset="204" size="100%">1990</style></year></dates><urls></urls></record></Cite></EndNote>[218]. Радиомодифицирующее действие карнозина было подтверждено с помощью метода эндогенных селезеночных колоний. Было обнаружено увеличение эффективности образования колоний гематопоэтическими стволовыми клетками из костного мозга мышей, которым перед облучением вводили карнозин, по сравнению с контрольными животными PEVuZE5vdGU+PENpdGU+PEF1dGhvcj7QmtGD0YDQtdC70LvQsDwvQXV0aG9yPjxZZWFyPjE5OTE8

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ADDIN EN.CITE.DATA [219, 220]. 1.3.3. Эффект карнозина и его производных на пролиферацию нормальных и опухолевых клетокПродолжительное культивирование нормальных человеческих фибробластов с карнозином (10 - 50 мМ) приводило к удлинению продолжительности жизни клеток, что выражалось в увеличении количества популяционных удвоений, т.н. предела Хейфлика ADDIN EN.CITE <EndNote><Cite><Author>Hayflick</Author><Year>1965</Year><RecNum>226</RecNum><DisplayText>[221]</DisplayText><record><rec-number>226</rec-number><foreign-keys><key app="EN" db-id="w2wfxafe6va9z5eev2l55aa8sxsa599eev2s">226</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Hayflick, L.</author></authors></contributors><titles><title>THE LIMITED IN VITRO LIFETIME OF HUMAN DIPLOID CELL STRAINS</title><secondary-title>Exp Cell Res</secondary-title><alt-title>Experimental cell research</alt-title></titles><periodical><full-title>Exp Cell Res</full-title><abbr-1>Experimental cell research</abbr-1></periodical><alt-periodical><full-title>Exp Cell Res</full-title><abbr-1>Experimental cell research</abbr-1></alt-periodical><pages>614-36</pages><volume>37</volume><edition>1965/03/01</edition><keywords><keyword>Aging</keyword><keyword>Cell Division</keyword><keyword>Chromosome Aberrations</keyword><keyword>Fetus</keyword><keyword>Lung</keyword><keyword>Tissue Culture Techniques</keyword></keywords><dates><year>1965</year><pub-dates><date>Mar</date></pub-dates></dates><isbn>0014-4827 (Print)&#xD;0014-4827 (Linking)</isbn><accession-num>14315085</accession-num><urls></urls><remote-database-provider>NLM</remote-database-provider><language>eng</language></record></Cite></EndNote>[221]. Кроме того, отмечали задержку развития старческого фенотипа в виде усиления зернистости и потери вытянутой фибробластоподобной формы ADDIN EN.CITE <EndNote><Cite><Author>McFarland</Author><Year>1994</Year><RecNum>162</RecNum><DisplayText>[22]</DisplayText><record><rec-number>162</rec-number><foreign-keys><key app="EN" db-id="w2wfxafe6va9z5eev2l55aa8sxsa599eev2s">162</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>McFarland, G. A.</author><author>Holliday, R.</author></authors></contributors><auth-address>CSIRO Division of Biomolecular Engineering, Sydney Laboratory, NSW, Australia.</auth-address><titles><title>Retardation of the senescence of cultured human diploid fibroblasts by carnosine</title><secondary-title>Exp Cell Res</secondary-title><alt-title>Experimental cell research</alt-title></titles><periodical><full-title>Exp Cell Res</full-title><abbr-1>Experimental cell research</abbr-1></periodical><alt-periodical><full-title>Exp Cell Res</full-title><abbr-1>Experimental cell research</abbr-1></alt-periodical><pages>167-75</pages><volume>212</volume><number>2</number><edition>1994/06/01</edition><keywords><keyword>Carnosine/ pharmacology</keyword><keyword>Cell Aging/ drug effects</keyword><keyword>Fibroblasts</keyword><keyword>Humans</keyword></keywords><dates><year>1994</year><pub-dates><date>Jun</date></pub-dates></dates><isbn>0014-4827 (Print)&#xD;0014-4827 (Linking)</isbn><accession-num>8187813</accession-num><urls></urls><electronic-resource-num>10.1006/excr.1994.1132</electronic-resource-num><remote-database-provider>NLM</remote-database-provider><language>eng</language></record></Cite></EndNote>[22]. Добавление карнозина к клеткам долгое время культивируемым на среде без карнозина приводило к обращению старческого фенотипа и способствовало увеличению продолжительности их жизни в сравнении с контролем. Удаление карнозина из среды нивелировало описанный эффект ADDIN EN.CITE <EndNote><Cite><Author>McFarland</Author><Year>1999</Year><RecNum>161</RecNum><DisplayText>[222]</DisplayText><record><rec-number>161</rec-number><foreign-keys><key app="EN" db-id="w2wfxafe6va9z5eev2l55aa8sxsa599eev2s">161</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>McFarland, G. A.</author><author>Holliday, R.</author></authors></contributors><auth-address>CSIRO Division of Molecular Science, Sydney Laboratory, North Ryde, Australia.</auth-address><titles><title>Further evidence for the rejuvenating effects of the dipeptide L-carnosine on cultured human diploid fibroblasts</title><secondary-title>Exp Gerontol</secondary-title><alt-title>Experimental gerontology</alt-title></titles><periodical><full-title>Exp Gerontol</full-title><abbr-1>Experimental gerontology</abbr-1></periodical><alt-periodical><full-title>Exp Gerontol</full-title><abbr-1>Experimental gerontology</abbr-1></alt-periodical><pages>35-45</pages><volume>34</volume><number>1</number><edition>1999/04/10</edition><keywords><keyword>Carnosine/ pharmacology</keyword><keyword>Cell Adhesion/drug effects</keyword><keyword>Cell Aging/ drug effects</keyword><keyword>Cell Division/drug effects</keyword><keyword>Cells, Cultured</keyword><keyword>Fibroblasts/drug effects/physiology</keyword><keyword>Humans</keyword></keywords><dates><year>1999</year><pub-dates><date>Jan</date></pub-dates></dates><isbn>0531-5565 (Print)&#xD;0531-5565 (Linking)</isbn><accession-num>10197726</accession-num><urls></urls><remote-database-provider>NLM</remote-database-provider><language>eng</language></record></Cite></EndNote>[222]. Карнозин также способствовал увеличению эффективности посева фибробластов. При посеве клеток в низких разведениях, в чашках инкубируемых с карнозином образовывалось больше колоний, чем в контрольных, что указывает на активацию пролиферативных процессов фибробластов под действием карнозина ADDIN EN.CITE <EndNote><Cite><Author>McFarland</Author><Year>1999</Year><RecNum>161</RecNum><DisplayText>[222]</DisplayText><record><rec-number>161</rec-number><foreign-keys><key app="EN" db-id="w2wfxafe6va9z5eev2l55aa8sxsa599eev2s">161</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>McFarland, G. A.</author><author>Holliday, R.</author></authors></contributors><auth-address>CSIRO Division of Molecular Science, Sydney Laboratory, North Ryde, Australia.</auth-address><titles><title>Further evidence for the rejuvenating effects of the dipeptide L-carnosine on cultured human diploid fibroblasts</title><secondary-title>Exp Gerontol</secondary-title><alt-title>Experimental gerontology</alt-title></titles><periodical><full-title>Exp Gerontol</full-title><abbr-1>Experimental gerontology</abbr-1></periodical><alt-periodical><full-title>Exp Gerontol</full-title><abbr-1>Experimental gerontology</abbr-1></alt-periodical><pages>35-45</pages><volume>34</volume><number>1</number><edition>1999/04/10</edition><keywords><keyword>Carnosine/ pharmacology</keyword><keyword>Cell Adhesion/drug effects</keyword><keyword>Cell Aging/ drug effects</keyword><keyword>Cell Division/drug effects</keyword><keyword>Cells, Cultured</keyword><keyword>Fibroblasts/drug effects/physiology</keyword><keyword>Humans</keyword></keywords><dates><year>1999</year><pub-dates><date>Jan</date></pub-dates></dates><isbn>0531-5565 (Print)&#xD;0531-5565 (Linking)</isbn><accession-num>10197726</accession-num><urls></urls><remote-database-provider>NLM</remote-database-provider><language>eng</language></record></Cite></EndNote>[222]. Позже Shao et al. продемонстрировали замедление скорости укорачивания теломер и снижение количества дефектов в ДНК теломер под действием карнозина PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5TaGFvPC9BdXRob3I+PFllYXI+MjAwNDwvWWVhcj48UmVj

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ADDIN EN.CITE.DATA [223]. Укорачивание теломер представляется одним из основных факторов ограничивающий количество клеточных делений ADDIN EN.CITE <EndNote><Cite><Author>Levy</Author><Year>1992</Year><RecNum>227</RecNum><DisplayText>[224]</DisplayText><record><rec-number>227</rec-number><foreign-keys><key app="EN" db-id="w2wfxafe6va9z5eev2l55aa8sxsa599eev2s">227</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Levy, M. Z.</author><author>Allsopp, R. C.</author><author>Futcher, A. B.</author><author>Greider, C. W.</author><author>Harley, C. B.</author></authors></contributors><auth-address>Department of Biochemistry, McMaster University Hamilton, Ontario, Canada.</auth-address><titles><title>Telomere end-replication problem and cell aging</title><secondary-title>J Mol Biol</secondary-title><alt-title>Journal of molecular biology</alt-title></titles><periodical><full-title>J Mol Biol</full-title><abbr-1>Journal of molecular biology</abbr-1></periodical><alt-periodical><full-title>J Mol Biol</full-title><abbr-1>Journal of molecular biology</abbr-1></alt-periodical><pages>951-60</pages><volume>225</volume><number>4</number><edition>1992/06/20</edition><keywords><keyword>Adult</keyword><keyword>Base Sequence</keyword><keyword>Cell Division</keyword><keyword>Cells, Cultured</keyword><keyword>Chromosome Deletion</keyword><keyword>Chromosomes, Human/ physiology</keyword><keyword>DNA/genetics/ metabolism</keyword><keyword>DNA Replication</keyword><keyword>Fibroblasts/cytology/physiology</keyword><keyword>Humans</keyword><keyword>Kinetics</keyword><keyword>Models, Genetic</keyword><keyword>Oligonucleotide Probes</keyword><keyword>Repetitive Sequences, Nucleic Acid</keyword><keyword>Skin Physiological Phenomena</keyword><keyword>Telomere/ physiology</keyword></keywords><dates><year>1992</year><pub-dates><date>Jun 20</date></pub-dates></dates><isbn>0022-2836 (Print)&#xD;0022-2836 (Linking)</isbn><accession-num>1613801</accession-num><urls></urls><remote-database-provider>NLM</remote-database-provider><language>eng</language></record></Cite></EndNote>[224], замедление этого процесса может быть одним из механизмов удлинения предела Хейфлика и задержки формирования старческого фенотипа с помощью карнозина.

В 1986 году, японские ученые Nagai и Suda впервые описали эффект карнозина на опухолевый клетки, который в корне отличался от действия карнозина на нормальные клетки. Подкожные инъекции карнозина мышам линии ddY, которым предварительно вживляли клетки саркомы, способствовали снижению интенсивности пролиферации опухолевых клеток и увеличивали выживаемость животных ADDIN EN.CITE <EndNote><Cite><Author>Nagai</Author><Year>1986</Year><RecNum>88</RecNum><DisplayText>[20]</DisplayText><record><rec-number>88</rec-number><foreign-keys><key app="EN" db-id="w2wfxafe6va9z5eev2l55aa8sxsa599eev2s">88</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Nagai, K.</author><author>Suda, T.</author></authors></contributors><titles><title>[Antineoplastic effects of carnosine and beta-alanine--physiological considerations of its antineoplastic effects]</title><secondary-title>Nihon Seirigaku Zasshi</secondary-title><alt-title>Nihon seirigaku zasshi. Journal of the Physiological Society of Japan</alt-title></titles><periodical><full-title>Nihon Seirigaku Zasshi</full-title><abbr-1>Nihon seirigaku zasshi. Journal of the Physiological Society of Japan</abbr-1></periodical><alt-periodical><full-title>Nihon Seirigaku Zasshi</full-title><abbr-1>Nihon seirigaku zasshi. Journal of the Physiological Society of Japan</abbr-1></alt-periodical><pages>741-7</pages><volume>48</volume><number>11</number><edition>1986/01/01</edition><keywords><keyword>Alanine/ pharmacology</keyword><keyword>Animals</keyword><keyword>Antineoplastic Agents</keyword><keyword>Carnosine/administration &amp; dosage/ pharmacology/therapeutic use</keyword><keyword>Dipeptides/ pharmacology</keyword><keyword>Male</keyword><keyword>Mice</keyword><keyword>Mitomycin</keyword><keyword>Mitomycins/administration &amp; dosage</keyword><keyword>Picibanil/administration &amp; dosage</keyword><keyword>Sarcoma 180/drug therapy</keyword><keyword>beta-Alanine/administration &amp; dosage/ pharmacology/therapeutic use</keyword></keywords><dates><year>1986</year></dates><isbn>0031-9341 (Print)&#xD;0031-9341 (Linking)</isbn><accession-num>3102721</accession-num><urls></urls><remote-database-provider>NLM</remote-database-provider><language>jpn</language></record></Cite></EndNote>[20]. В независимом исследовании Renner et al. на мышах с ксенографами HER2/neu NIH3T3 фибробластов показали, что ежедневные внутрибрюшинные инъекции карнозина существенно подавляют пролиферацию злокачественных клеток и опухолевый рост. Снижение скорости пролиферации опухолевых клеток под действием карнозина сопровождалось снижением синтеза АФК и активности лактатдегидрогеназы (ЛДГ). Кроме того, в опухолях мышей, которым делали инъекции карнозина карнозина было обнаружено меньшее количество митозов, чем у контрольных животных PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5SZW5uZXI8L0F1dGhvcj48WWVhcj4yMDEwPC9ZZWFyPjxS

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ADDIN EN.CITE.DATA [21]. Сходные изменения наблюдали в первичной культуре клеток мультиформной глиобластомы (Glioblastoma multiforme). Обработка клеток карнозином (20-50 мМ) в течение 4 дней приводила к уменьшению внутриклеточного уровня АТФ и ЛДГ, а также к снижению синтеза ДНК ADDIN EN.CITE <EndNote><Cite><Author>Renner</Author><Year>2008</Year><RecNum>91</RecNum><DisplayText>[208]</DisplayText><record><rec-number>91</rec-number><foreign-keys><key app="EN" db-id="w2wfxafe6va9z5eev2l55aa8sxsa599eev2s">91</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Renner, C.

, Seyffarth, A., Garcia de Arriba, S., Meixensberger, J., Gebhardt, R., Gaunitz, F.</author></authors></contributors><titles><title>Carnosine Inhibits Growth of Cells Isolated from Human Glioblastoma Multiforme</title><secondary-title>International Journal of Peptide Research and Therapeutics </secondary-title></titles><periodical><full-title>International Journal of Peptide Research and Therapeutics</full-title></periodical><pages>127-135</pages><volume>14</volume><section>127</section><dates><year>2008</year></dates><urls></urls></record></Cite></EndNote>[208]. Ингибирование митохондриального дыхания с помощью KCN в клетках обработанных карнозином не влияло на выработку АТФ или выживаемость клеток. Обработка клеток глиобластомы оксаматом – ингибитором ЛДГ – приводила к резкому снижению концентрации АТФ, при этом карнозин приводил к дальнейшему снижению концентрации АТФ. Исходя из полученных данных авторами был сделан вывод о том, что карнозин индуцирует изменения на уровне гликолиза, которые приводят к ингибированию клеточной пролиферации ADDIN EN.CITE <EndNote><Cite><Author>Renner</Author><Year>2010</Year><RecNum>71</RecNum><DisplayText>[225]</DisplayText><record><rec-number>71</rec-number><foreign-keys><key app="EN" db-id="w2wfxafe6va9z5eev2l55aa8sxsa599eev2s">71</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Renner, C.</author><author>Asperger, A.</author><author>Seyffarth, A.</author><author>Meixensberger, J.</author><author>Gebhardt, R.</author><author>Gaunitz, F.</author></authors></contributors><auth-address>Klinik und Poliklinik fur Neurochirurgie, Medizinische Fakultat der Universitat Leipzig, Leipzig, Germany.</auth-address><titles><title>Carnosine inhibits ATP production in cells from malignant glioma</title><secondary-title>Neurol Res</secondary-title><alt-title>Neurological research</alt-title></titles><periodical><full-title>Neurol Res</full-title><abbr-1>Neurological research</abbr-1></periodical><alt-periodical><full-title>Neurol Res</full-title><abbr-1>Neurological research</abbr-1></alt-periodical><pages>101-5</pages><volume>32</volume><number>1</number><edition>2009/11/17</edition><keywords><keyword>Adenosine Triphosphate/ metabolism</keyword><keyword>Antineoplastic Agents/ pharmacology</keyword><keyword>Carnosine/ pharmacology</keyword><keyword>Cell Line, Tumor</keyword><keyword>Cells, Cultured</keyword><keyword>Glioma/ drug therapy/ metabolism</keyword><keyword>Glucose/metabolism</keyword><keyword>Glycolysis/drug effects</keyword><keyword>Humans</keyword><keyword>Mitochondria/drug effects/metabolism</keyword><keyword>Organic Chemicals/pharmacology</keyword><keyword>Potassium Cyanide/pharmacology</keyword></keywords><dates><year>2010</year><pub-dates><date>Feb</date></pub-dates></dates><isbn>1743-1328 (Electronic)&#xD;0161-6412 (Linking)</isbn><accession-num>19909581</accession-num><urls></urls><electronic-resource-num>10.1179/016164109x12518779082237</electronic-resource-num><remote-database-provider>NLM</remote-database-provider><language>eng</language></record></Cite></EndNote>[225]. Holliday и McFarland наглядно продемонстрировали избирательность ингибирующего эффекта карнозина на рост опухолевых клеток PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5Ib2xsaWRheTwvQXV0aG9yPjxZZWFyPjE5OTY8L1llYXI+

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ADDIN EN.CITE.DATA [23, 226]. Добавление карнозина к смеси опухолевых клеток (HeLa) и нормальных человеческих фибробластов (MRC-5) приводило к выборочному уничтожению опухолевых HeLa клеток и способствовало удлинению продолжительности жизни фибробластов ADDIN EN.CITE <EndNote><Cite><Author>Holliday</Author><Year>1996</Year><RecNum>73</RecNum><DisplayText>[23]</DisplayText><record><rec-number>73</rec-number><foreign-keys><key app="EN" db-id="w2wfxafe6va9z5eev2l55aa8sxsa599eev2s">73</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Holliday, R.</author><author>McFarland, G. A.</author></authors></contributors><auth-address>CSIRO Division of Biomolecular Engineering, Sydney Laboratory, NSW, Australia.</auth-address><titles><title>Inhibition of the growth of transformed and neoplastic cells by the dipeptide carnosine</title><secondary-title>Br J Cancer</secondary-title><alt-title>British journal of cancer</alt-title></titles><periodical><full-title>Br J Cancer</full-title><abbr-1>British journal of cancer</abbr-1></periodical><alt-periodical><full-title>Br J Cancer</full-title><abbr-1>British journal of cancer</abbr-1></alt-periodical><pages>966-71</pages><volume>73</volume><number>8</number><edition>1996/04/01</edition><keywords><keyword>Animals</keyword><keyword>Antineoplastic Agents/ pharmacology</keyword><keyword>Carnosine/ pharmacology</keyword><keyword>Cell Line, Transformed</keyword><keyword>Culture Media</keyword><keyword>Glycolysis/drug effects</keyword><keyword>HeLa Cells</keyword><keyword>Humans</keyword><keyword>Pyruvates/metabolism</keyword><keyword>Pyruvic Acid</keyword><keyword>Simian virus 40</keyword><keyword>Tumor Cells, Cultured</keyword></keywords><dates><year>1996</year><pub-dates><date>Apr</date></pub-dates></dates><isbn>0007-0920 (Print)&#xD;0007-0920 (Linking)</isbn><accession-num>8611433</accession-num><urls></urls><custom2>PMC2075811</custom2><remote-database-provider>NLM</remote-database-provider><language>eng</language></record></Cite></EndNote>[23]. Интересно отметить, что действие карнозина на две активно пролиферирующие культуры – эмбриональные стволовые (ЭС) клетки мыши и иммортальные клетки эмбриональной терато-карциномы (ЭК) мыши также было различно. Карнозин препятствовал росту злокачественных ЭК-клеток, не затрагивая роста ЭС-клеток ADDIN EN.CITE <EndNote><Cite><Author>McFarland</Author><Year>1999</Year><RecNum>160</RecNum><DisplayText>[226]</DisplayText><record><rec-number>160</rec-number><foreign-keys><key app="EN" db-id="w2wfxafe6va9z5eev2l55aa8sxsa599eev2s">160</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>McFarland, G.

</author><author>Holliday, R.</author></authors></contributors><titles><title>Differential response of embryonic stem cells and teratocarcinoma cells to carnosine</title><secondary-title>In Vitro Cell Dev Biol Anim</secondary-title><alt-title>In vitro cellular &amp; developmental biology. Animal</alt-title></titles><periodical><full-title>In Vitro Cell Dev Biol Anim</full-title><abbr-1>In vitro cellular &amp; developmental biology. Animal</abbr-1></periodical><alt-periodical><full-title>In Vitro Cell Dev Biol Anim</full-title><abbr-1>In vitro cellular &amp; developmental biology. Animal</abbr-1></alt-periodical><pages>15-6</pages><volume>35</volume><number>1</number><edition>1999/09/04</edition><keywords><keyword>Animals</keyword><keyword>Carnosine/ pharmacology</keyword><keyword>Cell Division/drug effects</keyword><keyword>Embryo, Mammalian</keyword><keyword>Mice</keyword><keyword>Stem Cells/ drug effects/metabolism</keyword><keyword>Teratocarcinoma/ metabolism</keyword></keywords><dates><year>1999</year><pub-dates><date>Jan</date></pub-dates></dates><isbn>1071-2690 (Print)&#xD;1071-2690 (Linking)</isbn><accession-num>10475249</accession-num><urls></urls><electronic-resource-num>10.1007/s11626-999-0037-1</electronic-resource-num><remote-database-provider>NLM</remote-database-provider><language>eng</language></record></Cite></EndNote>[226]. Полученные данные указывали на то, что ни степень дифференцировки, ни скорость пролиферации клеток не являются определяющими факторами избирательного действия карнозина на опухолевые клетки.

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ADDIN EN.CITE.DATA [227]. Интересно отметить, что VBP1, BAG2 и GrpEL участвуют в регуляции активности Hypoxia-inducible factor 1 (HIF-1) PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5Uc3VjaGl5YTwvQXV0aG9yPjxZZWFyPjE5OTY8L1llYXI+

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ADDIN EN.CITE.DATA [232, 233]. Приведенные данные указывают на возможное участие регуляторных каскадов HIF-1 и Hsp70 в антипролиферативном эффекте карнозина. В нашей лаборатории было показано, что карнозин замедляет рост уровня фосфо-ERK1/2 и JNK в гранулярных клетках мозжечка в условиях окислительного стресса, индуцированного активацией глутаматных рецепторов ADDIN EN.CITE <EndNote><Cite><Author>Rybakova</Author><Year>2012</Year><RecNum>345</RecNum><DisplayText>[234]</DisplayText><record><rec-number>345</rec-number><foreign-keys><key app="EN" db-id="w2wfxafe6va9z5eev2l55aa8sxsa599eev2s">345</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Rybakova, Y.</author><author>Akkuratov, E.</author><author>Kulebyakin, K.</author><author>Brodskaya, O.</author><author>Dizhevskaya, A.</author><author>Boldyrev, A.</author></authors></contributors><auth-address>International Biotechnological Center of MV Lomonosov Moscow State University, Moscow, Russia. julie_ribakova@yahoo.com</auth-address><titles><title>Receptor-mediated oxidative stress in murine cerebellar neurons is accompanied by phosphorylation of MAP (ERK 1/2) kinase</title><secondary-title>Curr Aging Sci</secondary-title><alt-title>Current aging science</alt-title></titles><periodical><full-title>Curr Aging Sci</full-title><abbr-1>Current aging science</abbr-1></periodical><alt-periodical><full-title>Curr Aging Sci</full-title><abbr-1>Current aging science</abbr-1></alt-periodical><pages>225-30</pages><volume>5</volume><number>3</number><edition>2013/02/08</edition><dates><year>2012</year><pub-dates><date>Dec</date></pub-dates></dates><isbn>1874-6128 (Electronic)</isbn><accession-num>23387889</accession-num><urls></urls><remote-database-provider>NLM</remote-database-provider><language>eng</language></record></Cite></EndNote>[234]. Как было отмечено выше ERK1/2 представляет собой редокс-чувствительную МАР-киназу, регулирующую множество внутриклеточных функций, в том числе и пролиферативные процессы. Было показано, что ингибирование образования Н2О2 препятствует активации ERK1/2 и развитию пролиферативного ответа PEVuZE5vdGU+PENpdGU+PEF1dGhvcj5Eb25nPC9BdXRob3I+PFllYXI+MjAwNDwvWWVhcj48UmVj

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