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 <front>
  <journal-meta>
   <journal-id journal-id-type="publisher-id">Russian Journal of Biological Physics and Chemisrty</journal-id>
   <journal-title-group>
    <journal-title xml:lang="en">Russian Journal of Biological Physics and Chemisrty</journal-title>
    <trans-title-group xml:lang="ru">
     <trans-title>АКТУАЛЬНЫЕ ВОПРОСЫ БИОЛОГИЧЕСКОЙ ФИЗИКИ И ХИМИИ</trans-title>
    </trans-title-group>
   </journal-title-group>
   <issn publication-format="print">2499-9962</issn>
  </journal-meta>
  <article-meta>
   <article-id pub-id-type="publisher-id">54002</article-id>
   <article-categories>
    <subj-group subj-group-type="toc-heading" xml:lang="ru">
     <subject>Общая биофизика</subject>
    </subj-group>
    <subj-group subj-group-type="toc-heading" xml:lang="en">
     <subject>General biophysics</subject>
    </subj-group>
    <subj-group>
     <subject>Общая биофизика</subject>
    </subj-group>
   </article-categories>
   <title-group>
    <article-title xml:lang="en">INHIBITION OF PHOTOSYSTEM II ELECTRON-TRANSPORT CHAIN BY AMMONIA AND “DECOUPLING EFFECT”</article-title>
    <trans-title-group xml:lang="ru">
     <trans-title>РАЗОБЩЕНИЕ ФУНКЦИИ ВЫДЕЛЕНИЯ КИСЛОРОДА И ЭЛЕКТРОННОГО ТРАНСПОРТА В ФОТОСИСТЕМЕ 2 АММОНИЕМ</trans-title>
    </trans-title-group>
   </title-group>
   <contrib-group content-type="authors">
    <contrib contrib-type="author">
     <name-alternatives>
      <name xml:lang="ru">
       <surname>Ловягина</surname>
       <given-names>Е Р</given-names>
      </name>
      <name xml:lang="en">
       <surname>Lovyagina</surname>
       <given-names>E R</given-names>
      </name>
     </name-alternatives>
     <email>Elena.Lovyagina@gmail.com</email>
     <xref ref-type="aff" rid="aff-1"/>
    </contrib>
    <contrib contrib-type="author">
     <name-alternatives>
      <name xml:lang="ru">
       <surname>Белевич</surname>
       <given-names>Н П</given-names>
      </name>
      <name xml:lang="en">
       <surname>Belevich</surname>
       <given-names>N P</given-names>
      </name>
     </name-alternatives>
     <xref ref-type="aff" rid="aff-2"/>
    </contrib>
    <contrib contrib-type="author">
     <name-alternatives>
      <name xml:lang="ru">
       <surname>Сёмин</surname>
       <given-names>Б К</given-names>
      </name>
      <name xml:lang="en">
       <surname>Semin</surname>
       <given-names>B K</given-names>
      </name>
     </name-alternatives>
     <xref ref-type="aff" rid="aff-3"/>
    </contrib>
   </contrib-group>
   <aff-alternatives id="aff-1">
    <aff>
     <institution xml:lang="ru">ФГБОУ ВО «Московский государственный университет им. М.В. Ломоносова»</institution>
     <country>ru</country>
    </aff>
    <aff>
     <institution xml:lang="en">Moscow State University</institution>
     <country>ru</country>
    </aff>
   </aff-alternatives>
   <aff-alternatives id="aff-2">
    <aff>
     <institution xml:lang="ru">ФГБОУ ВО «Московский государственный университет им. М.В. Ломоносова»</institution>
     <country>ru</country>
    </aff>
    <aff>
     <institution xml:lang="en">Moscow State University</institution>
     <country>ru</country>
    </aff>
   </aff-alternatives>
   <aff-alternatives id="aff-3">
    <aff>
     <institution xml:lang="ru">ФГБОУ ВО «Московский государственный университет им. М.В. Ломоносова»</institution>
     <country>ru</country>
    </aff>
    <aff>
     <institution xml:lang="en">Moscow State University</institution>
     <country>ru</country>
    </aff>
   </aff-alternatives>
   <pub-date publication-format="print" date-type="pub" iso-8601-date="2016-06-25T20:22:29+03:00">
    <day>25</day>
    <month>06</month>
    <year>2016</year>
   </pub-date>
   <pub-date publication-format="electronic" date-type="pub" iso-8601-date="2016-06-25T20:22:29+03:00">
    <day>25</day>
    <month>06</month>
    <year>2016</year>
   </pub-date>
   <volume>1</volume>
   <issue>1</issue>
   <fpage>95</fpage>
   <lpage>98</lpage>
   <history>
    <date date-type="received" iso-8601-date="2016-06-20T20:22:29+03:00">
     <day>20</day>
     <month>06</month>
     <year>2016</year>
    </date>
    <date date-type="accepted" iso-8601-date="2016-06-20T20:22:29+03:00">
     <day>20</day>
     <month>06</month>
     <year>2016</year>
    </date>
   </history>
   <self-uri xlink:href="https://rusjbpc.ru/en/nauka/article/54002/view">https://rusjbpc.ru/en/nauka/article/54002/view</self-uri>
   <abstract xml:lang="ru">
    <p>Каталитическим центром кислород-выделяющего комплекса (КВК) фотосистемы 2 (ФС2) является кластер Mn4CaO5, структура которого в настоящее время расшифрована. Экстракция катионов кальция из КВК приводит к разобщению функции выделения молекулярного кислорода и электронного транспорта через ФС2 [Semin et al. Photosynth. Res. 98 (2008) 235] и одновременному появлению уширенного ЭПР сигнала с g = 2 (расщепленный «S3» сигнал). Не исключено, что это взаимосвязанные эффекты. Однако ЭПР сигнал «S3» наблюдается не только при удалении Ca2+ из КВК, но и после обработки нативных мембранных препаратов ФС2 анионами фтора, ацетата и хлоридом аммония. Ранее мы установили, что анионы фтора, подобно экстракции Ca2+, разобщают электрон-транспортные процессы и выделение кислорода. В представленной работе изучено действие хлорида и нитрата аммония на функциональную активность мембранных препаратов ФС2 с нативным КВК. Обнаружено, что как хлорид, так и нитрат аммония ингибирует выделение кислорода более эффективно, чем восстановление искусственного акцептора электронов 2,6-дихлорфенолиндофенола на акцепторной стороне ФС2, то есть действие аммония приводит к разобщению этих процессов. Предполагается, что ингибирующий эффект аммония связан с инактивацией функции Са2+.</p>
   </abstract>
   <trans-abstract xml:lang="en">
    <p>Catalytic center of oxygen-evolving complex (OEC) of photosystem II (PSII) is Mn4CaO5 cluster structure of which is determined now. Ca2+ extraction from OEC is accompanied by decoupling of oxygen evolution and electron transport processes in PSII and by the appearance of a broad EPR signal (so-called “S3” or “split signal”) at low temperatures. These data imply that decoupling and EPR effects can be interdependent. It is interesting that EPR signal «S3» is observed also after treatment of native PSII membranes by fluoride anions, acetate and chloride of ammonia. Previously we found that fluoride anions treatment of PSII like Ca2+ extraction from the OEC provide the appearance of decoupling effect. In the presented work we investigated the effect of ammonia chloride and ammonia nitrate on the functional activity of native PSII membranes to study the effect of anions. We found that ammonia nitrate like ammonia chloride inhibit the oxygen evolution more effectively than the reduction of artificial electron acceptor 2,6- dichlorophenolindophenol that is the action of ammonia decouples these processes. However the inhibition doesn’t depend on the anions. We suggest that ammonia inhibition effect is determined by inactivation of Ca2+ function.</p>
   </trans-abstract>
   <kwd-group xml:lang="ru">
    <kwd>фотосистема 2</kwd>
    <kwd>кислород-выделяющий комплекс</kwd>
    <kwd>аммоний</kwd>
   </kwd-group>
   <kwd-group xml:lang="en">
    <kwd>photosystem II</kwd>
    <kwd>oxygen-evolving complex</kwd>
    <kwd>ammonia</kwd>
   </kwd-group>
  </article-meta>
 </front>
 <body>
  <p></p>
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 <back>
  <ref-list>
   <ref id="B1">
    <label>1.</label>
    <citation-alternatives>
     <mixed-citation xml:lang="ru">Umena Y., Kawakami K., Shen J.-R., Kamiya N. Crystal structure of oxygen evolving photosystem II at a resolution of 1.9 Å. Nature, 2011, vol. 473, pp. 55-60.</mixed-citation>
     <mixed-citation xml:lang="en">Umena Y., Kawakami K., Shen J.-R., Kamiya N. Crystal structure of oxygen evolving photosystem II at a resolution of 1.9 Å. Nature, 2011, vol. 473, pp. 55-60.</mixed-citation>
    </citation-alternatives>
   </ref>
   <ref id="B2">
    <label>2.</label>
    <citation-alternatives>
     <mixed-citation xml:lang="ru">Suga M., Akita F., Hirata K., Ueno G., Murakami H., Nakajima Y., Shimizu T., Yamashita K., Yamamoto M., Ago H., Shen J.-R. Native structure of photosystem II at 1.95 Å resolution viewed by femtosecond X-ray pulses. Nature, 2015, vol. 517, pp. 99-103.</mixed-citation>
     <mixed-citation xml:lang="en">Suga M., Akita F., Hirata K., Ueno G., Murakami H., Nakajima Y., Shimizu T., Yamashita K., Yamamoto M., Ago H., Shen J.-R. Native structure of photosystem II at 1.95 Å resolution viewed by femtosecond X-ray pulses. Nature, 2015, vol. 517, pp. 99-103.</mixed-citation>
    </citation-alternatives>
   </ref>
   <ref id="B3">
    <label>3.</label>
    <citation-alternatives>
     <mixed-citation xml:lang="ru">Cardona T., Murray J.W., Rutherford A.W. Origin and Evolution of Water Oxidation before the Last Common Ancestor of the Cyanobacteria. Mol. Biol. Evol,. 2015, vol. 32, pp. 1310-1328.</mixed-citation>
     <mixed-citation xml:lang="en">Cardona T., Murray J.W., Rutherford A.W. Origin and Evolution of Water Oxidation before the Last Common Ancestor of the Cyanobacteria. Mol. Biol. Evol,. 2015, vol. 32, pp. 1310-1328.</mixed-citation>
    </citation-alternatives>
   </ref>
   <ref id="B4">
    <label>4.</label>
    <citation-alternatives>
     <mixed-citation xml:lang="ru">Ghanotakis D.F., Babcock G.T., Yocum C.F. Calcium reconstitutes high rates of oxygen evolution in polypeptide depleted photosystem II preparations. FEBS Lett., 1984, vol. 167, pp. 127-130.</mixed-citation>
     <mixed-citation xml:lang="en">Ghanotakis D.F., Babcock G.T., Yocum C.F. Calcium reconstitutes high rates of oxygen evolution in polypeptide depleted photosystem II preparations. FEBS Lett., 1984, vol. 167, pp. 127-130.</mixed-citation>
    </citation-alternatives>
   </ref>
   <ref id="B5">
    <label>5.</label>
    <citation-alternatives>
     <mixed-citation xml:lang="ru">Semin B.K., Davletshina L.N., Ivanov I.I., Rubin A.B., Seibert M. Decoupling of the processes of molecular oxygen synthesis and electron transport in Ca2+-depleted PSII membranes. Photosynth. Res., 2008, vol. 98, pp. 235-249.</mixed-citation>
     <mixed-citation xml:lang="en">Semin B.K., Davletshina L.N., Ivanov I.I., Rubin A.B., Seibert M. Decoupling of the processes of molecular oxygen synthesis and electron transport in Ca2+-depleted PSII membranes. Photosynth. Res., 2008, vol. 98, pp. 235-249.</mixed-citation>
    </citation-alternatives>
   </ref>
   <ref id="B6">
    <label>6.</label>
    <citation-alternatives>
     <mixed-citation xml:lang="ru">Semin B.K., Davletshina L.N., Timofeev K.N., Ivanov I.I., Rubin A.B., Seibert M. Production of reactive oxygen species in decoupled, Ca2+-depleted PSII and their use in assigning a function to chloride on both sides of PSII. Photosynth. Res., 2013, vol. 117, pp. 385-399.</mixed-citation>
     <mixed-citation xml:lang="en">Semin B.K., Davletshina L.N., Timofeev K.N., Ivanov I.I., Rubin A.B., Seibert M. Production of reactive oxygen species in decoupled, Ca2+-depleted PSII and their use in assigning a function to chloride on both sides of PSII. Photosynth. Res., 2013, vol. 117, pp. 385-399.</mixed-citation>
    </citation-alternatives>
   </ref>
   <ref id="B7">
    <label>7.</label>
    <citation-alternatives>
     <mixed-citation xml:lang="ru">Ono T., Inoue Y. Abnormal redox reactions in photosynthetic O2-evolving centers in NaCl/EDTA-washed PSII. A dark-stable EPR multiline signal and an unknown positive charge accumulator. Biochim. Biophys. Acta, 1990, vol. 1020, pp. 269-277.</mixed-citation>
     <mixed-citation xml:lang="en">Ono T., Inoue Y. Abnormal redox reactions in photosynthetic O2-evolving centers in NaCl/EDTA-washed PSII. A dark-stable EPR multiline signal and an unknown positive charge accumulator. Biochim. Biophys. Acta, 1990, vol. 1020, pp. 269-277.</mixed-citation>
    </citation-alternatives>
   </ref>
   <ref id="B8">
    <label>8.</label>
    <citation-alternatives>
     <mixed-citation xml:lang="ru">Boussac A., Zimmermann J.-L., Rutherford A.W. EPR signals from modified charge accumulation states of the oxygen evolving enzyme in Ca2+-deficient photosystem II. Biochemistry, 1989, vol. 28, pp. 8984-8989.</mixed-citation>
     <mixed-citation xml:lang="en">Boussac A., Zimmermann J.-L., Rutherford A.W. EPR signals from modified charge accumulation states of the oxygen evolving enzyme in Ca2+-deficient photosystem II. Biochemistry, 1989, vol. 28, pp. 8984-8989.</mixed-citation>
    </citation-alternatives>
   </ref>
   <ref id="B9">
    <label>9.</label>
    <citation-alternatives>
     <mixed-citation xml:lang="ru">Sivaraja M., Tso J., Dismukes G.C. A calcium-specific site influence the structure and activity of the manganese cluster responsible for photosynthetic water oxidation. Biochemistry, 1989, vol. 28, pp. 9459-9464.</mixed-citation>
     <mixed-citation xml:lang="en">Sivaraja M., Tso J., Dismukes G.C. A calcium-specific site influence the structure and activity of the manganese cluster responsible for photosynthetic water oxidation. Biochemistry, 1989, vol. 28, pp. 9459-9464.</mixed-citation>
    </citation-alternatives>
   </ref>
   <ref id="B10">
    <label>10.</label>
    <citation-alternatives>
     <mixed-citation xml:lang="ru">Hallahan B.J., Nugent J.H.A., Warden J.T., Evans M.C.W. Investigation of the origin of the &quot;S3&quot; EPR signal from the oxygen-evolving complex of photosystem 2: the role of tyrosine. Z. Biochemistry, 1992, vol. 31, pp. 4562-4573.</mixed-citation>
     <mixed-citation xml:lang="en">Hallahan B.J., Nugent J.H.A., Warden J.T., Evans M.C.W. Investigation of the origin of the &quot;S3&quot; EPR signal from the oxygen-evolving complex of photosystem 2: the role of tyrosine. Z. Biochemistry, 1992, vol. 31, pp. 4562-4573.</mixed-citation>
    </citation-alternatives>
   </ref>
   <ref id="B11">
    <label>11.</label>
    <citation-alternatives>
     <mixed-citation xml:lang="ru">Baumgarten M., Philo J.S., Dismukes G.C. Mechanism of photoinhibition of photosynthetic water oxidation by chloride depletion and fluoride substitution: oxidation of a protein residue. Biochemistry, 1990, vol. 29, pp. 10814-10822.</mixed-citation>
     <mixed-citation xml:lang="en">Baumgarten M., Philo J.S., Dismukes G.C. Mechanism of photoinhibition of photosynthetic water oxidation by chloride depletion and fluoride substitution: oxidation of a protein residue. Biochemistry, 1990, vol. 29, pp. 10814-10822.</mixed-citation>
    </citation-alternatives>
   </ref>
   <ref id="B12">
    <label>12.</label>
    <citation-alternatives>
     <mixed-citation xml:lang="ru">MacLachlan D.J., Nugent J.H.A. Investigation of the S3 Electron Paramagnetic Resonance Signal from the Oxygen-Evolving Complex of Photosystem 2: Effect of Inhibition of Oxygen Evolution by Acetate. Biochemistry, 1993, vol. 32, pp. 9772-9780.</mixed-citation>
     <mixed-citation xml:lang="en">MacLachlan D.J., Nugent J.H.A. Investigation of the S3 Electron Paramagnetic Resonance Signal from the Oxygen-Evolving Complex of Photosystem 2: Effect of Inhibition of Oxygen Evolution by Acetate. Biochemistry, 1993, vol. 32, pp. 9772-9780.</mixed-citation>
    </citation-alternatives>
   </ref>
   <ref id="B13">
    <label>13.</label>
    <citation-alternatives>
     <mixed-citation xml:lang="ru">Szalai V.A., Kühne H., Lakshmi K.V., Brudvig G.W. Characterization of the Interaction between Manganese and Tyrosine Z in Acetate-Inhibited Photosystem II. Biochemistry, 1998, vol. 37, pp. 13594-13603.</mixed-citation>
     <mixed-citation xml:lang="en">Szalai V.A., Kühne H., Lakshmi K.V., Brudvig G.W. Characterization of the Interaction between Manganese and Tyrosine Z in Acetate-Inhibited Photosystem II. Biochemistry, 1998, vol. 37, pp. 13594-13603.</mixed-citation>
    </citation-alternatives>
   </ref>
   <ref id="B14">
    <label>14.</label>
    <citation-alternatives>
     <mixed-citation xml:lang="ru">Ghanotakis D.F., Babcock G.T. Hydroxylamine as an inhibitor between Z and P680 in photosystem II. FEBS Lett., 1983, vol. 153, pp. 231-234.</mixed-citation>
     <mixed-citation xml:lang="en">Ghanotakis D.F., Babcock G.T. Hydroxylamine as an inhibitor between Z and P680 in photosystem II. FEBS Lett., 1983, vol. 153, pp. 231-234.</mixed-citation>
    </citation-alternatives>
   </ref>
   <ref id="B15">
    <label>15.</label>
    <citation-alternatives>
     <mixed-citation xml:lang="ru">Dunahay T.G., Staechelin L.A., Seibert M., Ogilvie P.D., Berg S.P. Structural biochemical and biophysical characterization of four oxygen-evolving Photosystem 2 preparations from spinach. Biochim. Biophys. Acta, 1984, vol. 764, pp. 179-193.</mixed-citation>
     <mixed-citation xml:lang="en">Dunahay T.G., Staechelin L.A., Seibert M., Ogilvie P.D., Berg S.P. Structural biochemical and biophysical characterization of four oxygen-evolving Photosystem 2 preparations from spinach. Biochim. Biophys. Acta, 1984, vol. 764, pp. 179-193.</mixed-citation>
    </citation-alternatives>
   </ref>
   <ref id="B16">
    <label>16.</label>
    <citation-alternatives>
     <mixed-citation xml:lang="ru">Polander B.C., Barry B.A. A hydrogen-bonding network plays a catalytic role in photosynthetic oxygen evolution. Proc. Natl. Acad. Sci. U.S.A., 2012, vol. 109, pp. 6112-6117.</mixed-citation>
     <mixed-citation xml:lang="en">Polander B.C., Barry B.A. A hydrogen-bonding network plays a catalytic role in photosynthetic oxygen evolution. Proc. Natl. Acad. Sci. U.S.A., 2012, vol. 109, pp. 6112-6117.</mixed-citation>
    </citation-alternatives>
   </ref>
   <ref id="B17">
    <label>17.</label>
    <citation-alternatives>
     <mixed-citation xml:lang="ru">Polander B.C., Barry B.A. Detection of an intermediary, protonated water cluster in photosynthetic oxygen evolution. Proc. Natl. Acad. Sci. U.S.A., 2013, vol. 110, pp. 10634-10639.</mixed-citation>
     <mixed-citation xml:lang="en">Polander B.C., Barry B.A. Detection of an intermediary, protonated water cluster in photosynthetic oxygen evolution. Proc. Natl. Acad. Sci. U.S.A., 2013, vol. 110, pp. 10634-10639.</mixed-citation>
    </citation-alternatives>
   </ref>
   <ref id="B18">
    <label>18.</label>
    <citation-alternatives>
     <mixed-citation xml:lang="ru">Kuntzleman T., Haddy A. Fluoride inhibition of photosystem II and the effect of removal of the PsbQ subunit. Photosynth. Res., 2009, vol. 102, pp. 7-19.</mixed-citation>
     <mixed-citation xml:lang="en">Kuntzleman T., Haddy A. Fluoride inhibition of photosystem II and the effect of removal of the PsbQ subunit. Photosynth. Res., 2009, vol. 102, pp. 7-19.</mixed-citation>
    </citation-alternatives>
   </ref>
   <ref id="B19">
    <label>19.</label>
    <citation-alternatives>
     <mixed-citation xml:lang="ru">Sandusky P.O., Yocum C.F. The chloride requirement for photosynthetic oxygen evolution: factors affecting nucleophylic displacement of chloride from the oxygen-evolving complex. Biochim. Biophys. Acta, 1986, vol. 849, pp. 85-93.</mixed-citation>
     <mixed-citation xml:lang="en">Sandusky P.O., Yocum C.F. The chloride requirement for photosynthetic oxygen evolution: factors affecting nucleophylic displacement of chloride from the oxygen-evolving complex. Biochim. Biophys. Acta, 1986, vol. 849, pp. 85-93.</mixed-citation>
    </citation-alternatives>
   </ref>
   <ref id="B20">
    <label>20.</label>
    <citation-alternatives>
     <mixed-citation xml:lang="ru">Tsuno M., Suzuki H., Kondo T., Mino H., Noguchi T. Interaction and Inhibitory Effect of Ammonium Cation in the Oxygen Evolving Center of Photosytem II. Biochemistry, 2011, vol. 50, pp. 2506-2514.</mixed-citation>
     <mixed-citation xml:lang="en">Tsuno M., Suzuki H., Kondo T., Mino H., Noguchi T. Interaction and Inhibitory Effect of Ammonium Cation in the Oxygen Evolving Center of Photosytem II. Biochemistry, 2011, vol. 50, pp. 2506-2514.</mixed-citation>
    </citation-alternatives>
   </ref>
   <ref id="B21">
    <label>21.</label>
    <citation-alternatives>
     <mixed-citation xml:lang="ru">Sandusky P.O., Yocum C.F. The chloride requirement for photosynthetic oxygen evolution. Analysis of the effects of chloride and other anions on amine inhibition of the oxygen-evolving complex. Biochim. Biophys. Acta, 1984, vol. 766, pp. 603-611.</mixed-citation>
     <mixed-citation xml:lang="en">Sandusky P.O., Yocum C.F. The chloride requirement for photosynthetic oxygen evolution. Analysis of the effects of chloride and other anions on amine inhibition of the oxygen-evolving complex. Biochim. Biophys. Acta, 1984, vol. 766, pp. 603-611.</mixed-citation>
    </citation-alternatives>
   </ref>
  </ref-list>
 </back>
</article>
