Photosynthesis Research

, Volume 126, Issue 2–3, pp 449–463 | Cite as

Repetitive light pulse-induced photoinhibition of photosystem I severely affects CO2 assimilation and photoprotection in wheat leaves

  • Marek Zivcak
  • Marian BresticEmail author
  • Kristyna Kunderlikova
  • Oksana Sytar
  • Suleyman I. AllakhverdievEmail author
Regular Paper


It was previously found that photosystem I (PSI) photoinhibition represents mostly irreversible damage with a slow recovery; however, its physiological significance has not been sufficiently characterized. The aim of the study was to assess the effect of PSI photoinhibition on photosynthesis in vivo. The inactivation of PSI was done by a series of short light saturation pulses applied by fluorimeter in darkness (every 10 s for 15 min), which led to decrease of both PSI (~60 %) and photosystem II (PSII) (~15 %) photochemical activity. No PSI recovery was observed within 2 days, whereas the PSII was fully recovered. Strongly limited PSI electron transport led to an imbalance between PSII and PSI photochemistry, with a high excitation pressure on PSII acceptor side and low oxidation of the PSI donor side. Low and delayed light-induced NPQ and P700+ rise in inactivated samples indicated a decrease in formation of transthylakoid proton gradient (ΔpH), which was confirmed also by analysis of electrochromic bandshift (ECSt) records. In parallel with photochemical parameters, the CO2 assimilation was also strongly inhibited, more in low light (~70 %) than in high light (~45 %); the decrease was not caused by stomatal closure. PSI electron transport limited the CO2 assimilation at low to moderate light intensities, but it seems not to be directly responsible for a low CO2 assimilation at high light. In this regard, the possible effects of PSI photoinhibition on the redox signaling in chloroplast and its role in downregulation of Calvin cycle activity are discussed.


PSI photoinactivation Transthylakoid proton gradient Non-photochemical quenching Electrochromic bandshift P700 


\(A_{{{\text{CO}}_{2} }}\)

CO2 assimilation rate


Cyclic electron transport

cyt b6/f

Cytochrome b 6/f


Electrochromic shift


Apparent electron transport rate


Minimum fluorescence from dark-adapted leaf (PSII centers open)


Minimum fluorescence from light-adapted leaf

Fm, \(F_{\rm m}^{\prime}\)

Maximum fluorescence from dark- or light-adapted leaf respectively (PS II centers closed)


Ferredoxin NADP+ oxidoreductase


Maximum quantum yield of PSII photochemistry


Transthylakoid proton conductivity


Light emitting diode


Light harvesting complex


Non-photochemical quenching


P700 absorbance at given light intensity


Primary electron donor of PSI (reduced form)


Primary electron donor of PSI (oxidized form)


Pulse-amplitude modulated


Photosynthetic active radiation

Pm, \(P_{\rm m}^{\prime}\)

Maximum P700 signal in dark- or light-adapted state


Proton motive force


Photosystem I


Photosystem II


Primary PSII acceptor


Total Redox poise of the primary electron acceptor of PSII (1 − qP)


PH dependent energy dissipation


‘Lake’ model photochemical quenching coefficient


‘Puddle’ model photochemical quenching coefficient


Saturation light pulse


Transthylakoid pH gradient


Osmotic component of proton motive force


Quantum yield of non-photochemical energy dissipation in PSI due to acceptor side limitation


Quantum yield of non-photochemical energy dissipation in PSI due to donor side limitation


Quantum efficiency of non-regulated energy dissipation in PSII


Quantum yield of pH-dependent energy dissipation in PSII


Effective quantum yield (efficiency) of PSI photochemistry at given actinic light intensity


Actual quantum yield (efficiency) of PSII photochemistry


Transmembrane electric potential


Electric component of proton motive force



This work was supported by the European Community under the project no. 26220220180: “Construction of the “AgroBioTech” Research Centre and  project “Center of Excellence for Agrobiodiversity Conservation and Use, ECOVA”.” SIA was supported by Grants from the Russian Foundation for Basic Research (Nos. 14-04-01549, 14-04-92690), and by Molecular and Cell Biology Programs of the Russian Academy of Sciences.

Supplementary material

11120_2015_121_MOESM1_ESM.pdf (392 kb)
Supplementary material 1 (PDF 392 kb)


  1. Alboresi A, Ballottari M, Hienerwadel R, Giacometti GM, Morosinotto T (2009) Antenna complexes protect photosystem I from photoinhibition. BMC Plant Biol 9:71PubMedCentralCrossRefPubMedGoogle Scholar
  2. Allakhverdiev SI, Murata N (2004) Environmental stress inhibits the synthesis de novo of proteins involved in the photodamage-repair cycle of Photosystem II in Synechocystis sp. PCC 6803. Biochim Biophys Acta 1657:23–32CrossRefPubMedGoogle Scholar
  3. Allakhverdiev SI, Murata N (2008) Salt stress inhibits photosystems II and I in cyanobacteria. Photosynth Res 98:529–539CrossRefPubMedGoogle Scholar
  4. Apostol S, Briantais JM, Moise N, Cerovic Z, Moya I (2001) Photoinactivation of the photosynthetic electron transport chain by accumulation of over-saturating light pulses given to dark adapted pea leaves. Photosynth Res 67:215–227CrossRefPubMedGoogle Scholar
  5. Baker NR (2008) Chlorophyll fluorescence: a probe of photosynthesis in vivo. Annu Rev Plant Biol 59:89–113CrossRefPubMedGoogle Scholar
  6. Barber J, Andersson B (1994) Revealing the blueprint of photosynthesis. Nature 370:31–34CrossRefGoogle Scholar
  7. Brestic M, Zivcak M, Olsovska K, Shao HB, Kalaji HM, Allakhverdiev SI (2014) Reduced glutamine synthetase activity plays a role in control of photosynthetic responses to high light in barley leaves. Plant Physiol Biochem 81:74–83CrossRefPubMedGoogle Scholar
  8. Briantais JM, Vernotte C, Picaud M, Krause GH (1979) A quantitive study of the slow decline of chlorophyll a fluorescence in isolated chloroplasts. Biochim Biophys Acta 548:128–138CrossRefPubMedGoogle Scholar
  9. Buchanan BB, Schürmann P, Wolosiuk RA, Jacquot JP (2002) The ferredoxin/thioredoxin system: from discovery to molecular structures and beyond. Photosynth Res 73:215–222CrossRefPubMedGoogle Scholar
  10. Bukhov NG, Carpentier R (2003) Measurement of photochemical quenching of absorbed quanta in photosystem I of intact leaves using simultaneous measurements of absorbance changes at 830 nm and thermal dissipation. Planta 216:630–638PubMedGoogle Scholar
  11. Bukhov N, Carpentier R (2004) Alternative photosystem I-driven electron transport routes: mechanisms and functions. Photosynth Res 82:17–33CrossRefPubMedGoogle Scholar
  12. Cardol P, Forti G, Finazzi G (2011) Regulation of electron transport in microalgae. Biochim Biophys Acta 1807:912–918CrossRefPubMedGoogle Scholar
  13. Chazdon R (1988) Sunflecks and their importance to forest understory plants. Adv Ecol Res 18:1–63CrossRefGoogle Scholar
  14. Chibani K, Couturier J, Selles B, Jacquot JP, Rouhier N (2010) The chloroplastic thiol reducing systems: dual functions in the regulation of carbohydrate metabolism and regeneration of antioxidant enzymes, emphasis on the poplar redoxin equipment. Photosyn Res 104:75–99CrossRefPubMedGoogle Scholar
  15. Demmig-Adams B (1990) Carotenoids and photoprotection in plants: a role for the xanthophyll zeaxanthin. Biochim Biophys Acta 1020:1–24CrossRefGoogle Scholar
  16. Dietz KJ, Pfannschmidt T (2011) Novel regulators in photosynthetic redox control of plant metabolism and gene expression. Plant Physiol 155:1477–1485PubMedCentralCrossRefPubMedGoogle Scholar
  17. Fischer S, Gräber P (1999) Comparison of ΔpH- and Δφ-driven ATP synthesis catalyzed by H + -ATPases from Escherichia coli or chloroplasts reconstituted into liposomes. FEBS Lett 457:327–332CrossRefPubMedGoogle Scholar
  18. Goltsev V, Zaharieva I, Chernev P, Kouzmanova M, Kalaji MH, Yordanov I, Krasteva V, Alexandrov V, Stefanov D, Allakhverdiev SI, Strasser RJ (2012) Drought-induced modifications of photosynthetic electron transport in intact leaves: analysis and use of neural networks as a tool for a rapid non-invasive estimation. Biochim Biophys Acta 1817:1490–1498CrossRefPubMedGoogle Scholar
  19. Grieco M, Tikkanen M, Paakkarinen V, Kangasjärvi S, Aro EM (2012) Steady-state phosphorylation of light-harvesting complex II proteins preserves Photosystem I under fluctuating white light. Plant Physiol 160:1896–1910PubMedCentralCrossRefPubMedGoogle Scholar
  20. Heber U, Neimanis S, Dietz KJ (1988) Fractional control of photosynthesis by the QB-protein, the cytochrome-f cytochrome-B6 complex and other components of the photosynthetic apparatus. Planta 173:267–274CrossRefPubMedGoogle Scholar
  21. Ivanov AG, Morgan R, Gray G, Velitchkova M, Huner NP (1998) Temperature/light dependent development of selective resistance to photoinhibition of photosystem I. FEBS Lett 430:288–292CrossRefPubMedGoogle Scholar
  22. Ivanov AG, Krol M, Zeinalov Y, Huner NPA, Sane PV (2008) The lack of LHCII proteins modulates excitation energy partitioning and PSII charge recombination in Chlorina F2 mutant of barley. Physiol Mol Biol Plants 14:205–215PubMedCentralCrossRefPubMedGoogle Scholar
  23. Joliot P, Johnson GN (2011) Regulation of cyclic and linear electron flow in higher plants. Proc Natl Acad Sci USA 108:13317–13322PubMedCentralCrossRefPubMedGoogle Scholar
  24. Joliot P, Joliot A (2002) Cyclic electron transfer in plant leaf. Proc Natl Acad Sci USA 99:10209–10214PubMedCentralCrossRefPubMedGoogle Scholar
  25. Jones HG (1985) Partitioning stomatal and non-stomatal limitations to photosynthesis. Plant Cell Environ 8:95–104CrossRefGoogle Scholar
  26. Kalaji HM, Schansker G, Ladle RJ, Goltsev V, Bosa K, Allakhverdiev SI et al (2014) Frequently asked questions about in vivo chlorophyll fluorescence: practical issues. Photosynth Res 122:121–158PubMedCentralCrossRefPubMedGoogle Scholar
  27. Klughammer C, Schreiber U (1994) Saturation pulse method for assessment of energy conversion in PS I. Planta 192:261–268CrossRefGoogle Scholar
  28. Klughammer C, Siebke K, Schreiber U (2013) Continuous ECS-indicated recording of the proton-motive charge flux in leaves. Photosynth Res 117:471–487PubMedCentralCrossRefPubMedGoogle Scholar
  29. Kono M, Terashima I (2014) Long-term and short-term responses of the photosynthetic electron transport to fluctuating light. J Photochem Photobiol B 137:89–99CrossRefPubMedGoogle Scholar
  30. Kono M, Noguchi K, Terashima I (2014) Roles of the cyclic electron flow around PSI (CEF-PSI) and O2-dependent alternative pathways in regulation of the photosynthetic electron flow in short-term fluctuating light in Arabidopsis thaliana. Plant Cell Physiol 55:990–1004CrossRefPubMedGoogle Scholar
  31. Kramer DM, Evans JR (2011) The importance of energy balance in improving photosynthetic productivity. Plant Physiol 155:70–78PubMedCentralCrossRefPubMedGoogle Scholar
  32. Kramer DM, Cruz JA, Kanazawa A (2003) Balancing the central roles of the thylakoid proton gradient. Trends Plant Sci 8:27–32CrossRefPubMedGoogle Scholar
  33. Kramer DM, Johnson G, Kiirats O, Edwards GE (2004) New fluorescence parameters for the determination of QA redox state and excitation energy fluxes. Photosynth Res 79:209–218CrossRefPubMedGoogle Scholar
  34. Kudoh H, Sonoike K (2002) Irreversible damage to photosystem I by chilling in the light: cause of the degradation of chlorophyll after returning to normal growth temperature. Planta 215:541–548CrossRefPubMedGoogle Scholar
  35. Lemaire SD, Michelet L, Zaffagnini M, Massot V, Issakidis-Bourguet E (2007) Thioredoxins in chloroplasts. Curr Genet 51:343–365CrossRefPubMedGoogle Scholar
  36. Li XP, Björkman O, Shih C, Grossman AR, Rosenquist M, Jansson S, Niyogi KK (2000) A pigment-binding protein essential for regulation of photosynthetic light harvesting. Nature 403:391–395CrossRefPubMedGoogle Scholar
  37. Long SP, Drake BG (1991) Effect of the long-term elevation of CO2 concentration in the field on the quantum yield of photosynthesis of the C3 sedge, Scirpus-olneyi. Plant Physiol 96:221–226PubMedCentralCrossRefPubMedGoogle Scholar
  38. Meyer Y, Belin C, Delorme-Hinoux V, Reichheld JP, Riondet C (2012) Thioredoxin and glutaredoxin systems in plants: molecular mechanisms, crosstalks, and functional significance. Antioxid Redox Signal 17:1124–1160CrossRefPubMedGoogle Scholar
  39. Mitchell P (1966) Chemiosmotic coupling in oxidative and photosynthetic phosphorylation. Biol Rev 41:445–502CrossRefPubMedGoogle Scholar
  40. Miyake C (2010) Alternative electron flows (water–water cycle and cyclic electron flow around PSI) in photosynthesis: molecular mechanisms and physiological functions. Plant Cell Physiol 51:1951–1963CrossRefPubMedGoogle Scholar
  41. Miyake C, Miyata M, Shinzaki Y, Tomizawa KI (2005) CO2 response of cyclic electron flow around PSI (CEF-PSI) in tobacco leaves—relative electron fluxes through PSI and PSII determine the magnitude of non-photochemical quenching (NPQ) of Chl fluorescence. Plant Cell Physiol 46:629–637CrossRefPubMedGoogle Scholar
  42. Munekage Y, Hashimoto M, Miyake C, Tomizawa KI, Endo T, Tasaka M, Shikanai T (2004) Cyclic electron flow around photosystem I is essential for photosynthesis. Nature 429:579–582CrossRefPubMedGoogle Scholar
  43. Murata N, Takahashi S, Nishiyama Y, Allakhverdiev SI (2007) Photoinhibition of photosystem II under environmental stress. Biochim Biophys Acta 1767:414–421CrossRefPubMedGoogle Scholar
  44. Ögren E, Evans JR (1993) Photosynthetic light-response curves. Planta 189:182–190CrossRefGoogle Scholar
  45. Okegawa Y, Kagawa Y, Kobayashi Y, Shikanai T (2008) Characterization of factors affecting the activity of photosystem I cyclic electron transport in chloroplasts. Plant Cell Physiol 49:825–834CrossRefPubMedGoogle Scholar
  46. Pearcy RW, Chazdon RL, Gross LJ, Mott KA (1994) Photosynthetic utilization of sunflecks: a temporally patchy resource on a time scale of seconds to minutes. In: Caldwell MM, Pearcy RW (eds) Exploitation of environmental heterogeneity by plants. Academic Press, San Diego, pp 175–208CrossRefGoogle Scholar
  47. Powles SB (1984) Photoinhibition of Photosynthesis Induced by Visible Light. Annu Rev Plant Physiol 35:15–44CrossRefGoogle Scholar
  48. Purcell M, Carpentier R (1994) Homogeneous photobleaching of chlorophyll holochromes in a photosystem I reaction center complex. Photochem Photobiol 59:215–218CrossRefGoogle Scholar
  49. Rajagopal S, Bukhov NG, Carpentier R (2002) Changes in the structure of chlorophyll-protein complexes and excitation energy transfer during photoinhibitory treatment of isolated photosystem I submembrane particles. J Photochem Photobiol B 62:194–200CrossRefGoogle Scholar
  50. Rajagopal S, Bukhov NG, Carpentier R (2003) Photoinhibitory light-induced changes in the composition of chlorophyll-protein complexes and photochemical activity in photosystem-I submembrane fractions. Photochem Photobiol 77:284–291CrossRefPubMedGoogle Scholar
  51. Ruban AV, Walters RG, Horton P (1992) The molecular mechanism of the control of excitation energy dissipation in chloroplast membranes inhibition of pH-dependent quenching of chlorophyll fluorescence by dicyclohexylcarbodiimide. FEBS Lett 309:175–179CrossRefPubMedGoogle Scholar
  52. Ruelland E, Miginiac-Maslow M (1999) Regulation of chloroplast enzyme activities by thioredoxins: activation or relief from inhibition? Trends Plant Sci 4:136–141CrossRefPubMedGoogle Scholar
  53. Sacksteder CA, Kramer DM (2000) Dark-interval relaxation kinetics (DIRK) of absorbance changes as a quantitative probe of steady-state electron transfer. Photosynth Res 66:145–158CrossRefPubMedGoogle Scholar
  54. Sacksteder CA, Kanazawa A, Jacoby ME, Kramer DM (2000) The proton to electron stoichiometry of steady-state photosynthesis in living plants: a proton-pumping Q-cycle is continuously engaged. Proc Natl Acad Sci USA 97:14283–14288PubMedCentralCrossRefPubMedGoogle Scholar
  55. Scheller HV, Haldrup A (2005) Photoinhibition of photosystem I. Planta 221:5–8CrossRefPubMedGoogle Scholar
  56. Schürmann P, Buchanan BB (2008) The ferredoxin/thioredoxin system of oxygenic photosynthesis. Antioxid Redox Signal 10:1235–1274CrossRefPubMedGoogle Scholar
  57. Sejima T, Takagi D, Fukayama H, Makino A, Miyake C (2014) Repetitive short-pulse light mainly inactivates photosystem I in sunflower leaves. Plant Cell Physiol. doi: 10.1093/pcp/pcu061 PubMedGoogle Scholar
  58. Shen Y-K, Chow WS, Park Y-I, Anderson JM (1996) Photoinactivation of Photosystem II by cumulative exposure to short light pulses during the induction period of photosynthesis. Photosynth Res 47:51–59CrossRefPubMedGoogle Scholar
  59. Shikanai T (2012) Cyclic electron transport around photosystem I; genetic approaches. Annu Rev Plant Biol 58:199–217CrossRefGoogle Scholar
  60. Shikanai T (2014) Central role of cyclic electron transport around photosystem I in the regulation of photosynthesis. Curr Opin Biotechnol 26:25–30CrossRefPubMedGoogle Scholar
  61. Sonoike K (1996) Photoinhibition of photosystem I: its physiological significance in the chilling sensitivity of plants. Plant Cell Physiol 37:239–247CrossRefGoogle Scholar
  62. Sonoike K (2011) Photoinhibition of photosystem I. Physiol Plant 142:56–64CrossRefPubMedGoogle Scholar
  63. Sonoike K, Terashima I (1994) Mechanism of photosystem-I photoinhibition in leaves of Cucumis sativus L. Planta 194:287–293CrossRefGoogle Scholar
  64. Sonoike K, Kamo M, Hihara Y, Hiyama T, Enami I (1997) The mechanism of the degradation of PsaB gene product, one of the photosynthetic reaction centre subunits of photosystem I, upon photoinhibition. Photosynth Res 53:55–63CrossRefGoogle Scholar
  65. Suorsa M, Jarvi S, Grieco M, Nurmi M, Pietrzykowska M, Rantala M, Kangasjarvi S, Paakkarinen V, Tikkanen M, Jansson S, Aro EM (2012) Proton gradient regulation5 is essential for proper acclimation of Arabidopsis photosystem I to naturally and artificially fluctuating light conditions. Plant Cell 24:2934–2948PubMedCentralCrossRefPubMedGoogle Scholar
  66. Szyszka B, Ivanov AG, Huner NP (2007) Psychrophily is associated with differential energy partitioning, photosystem stoichiometry and polypeptide phosphorylation in Chlamydomonas raudensis. Biochim Biophys Acta 1767:789–800CrossRefPubMedGoogle Scholar
  67. Teicher HB, Møller BL, Scheller HV (2000) Photoinhibition of photosystem I in field-grown barley (Hordeum vulgare L.): induction, recovery and acclimation. Photosynth Res 64:53–61CrossRefGoogle Scholar
  68. Terashima I, Funayama S, Sonoike K (1994) The site of photoinhibition in leaves of Cucumis sativus L. at low temperatures is photosystem I, not photosystem II. Planta 193:300–306CrossRefGoogle Scholar
  69. Thiele A, Winter K, Krause GH (1997) Low inactivation of D1 protein of photosystem II in young canopy leaves of Anacardium excelsum under high-light stress. J Plant Physiol 151:286–292CrossRefGoogle Scholar
  70. Tikkanen M, Grieco M, Nurmi M, Rantala M, Suorsa M, Aro EM (2012) Regulation of the photosynthetic apparatus under fluctuating growth light. Philos Trans Roy Soc B 367:3486–3493CrossRefGoogle Scholar
  71. Tikkanen M, Mekala NR, Aro EM (2014) Photosystem II photoinhibition-repair cycle protects Photosystem I from irreversible damage. Biochim Biophys Acta 1837:210–215CrossRefPubMedGoogle Scholar
  72. Tjus SE, Møller BL (1998) Scheller HV (1998) Photosystem I is an early target of photoinhibition in barley illuminated at chilling temperatures. Plant Physiol 116:755–764CrossRefPubMedGoogle Scholar
  73. Tjus SE, Møller BL, Scheller HV (1999) Photoinhibition of photosystem I damages both reaction centre proteins PSI-A and PSI-B and acceptor-side located small photosystem I polypeptides. Photosynth Res 60:75–86CrossRefGoogle Scholar
  74. Wang LF, Chen YY (2013) Characterization of a wide leaf mutant of rice (Oryza sativa L.) with high yield potential in field. Pak J Bot 45:921–926Google Scholar
  75. Wang C, Yamamoto H, Shikanai T (2015) Role of cyclic electron transport around photosystem I in regulating proton motive force. Biochim Biophys Acta. doi: 10.1016/j.bbabio.2014.11.013 Google Scholar
  76. Weis E, Ball JR, Berry J (1987) Photosynthetic control of electron transport in leaves of Phaseolus vulgaris. Evidence for regulation of PSII by the proton gradient. In: Biggins J (ed) Progress in photosynthesis research. Kluwer, Dordrecht, pp 553–556CrossRefGoogle Scholar
  77. Witt HT (1979) Energy conversion in the functional membrane of photosynthesis. Analysis by light pulse and electric pulse methods the central role of the electric field. Biochim Biophys Acta 505:355–427CrossRefPubMedGoogle Scholar
  78. Wolosiuk RA, Buchanan BB (1977) Thioredoxin and glutathione regulate photosynthesis in chloroplasts. Nature 266:565–567CrossRefGoogle Scholar
  79. Zhang S, Scheller HV (2004) Photoinhibition of photosystem I at chilling temperature and subsequent recovery in Arabidopsis thaliana. Plant Cell Physiol 45:1595–1602CrossRefPubMedGoogle Scholar
  80. Zhang ZS, Jia YJ, Gao HY, Zhang LT, Li HD, Meng QW (2011) Characterization of PSI recovery after chilling-induced photoinhibition in cucumber (Cucumis sativus L.) leaves. Planta 234:883–889CrossRefPubMedGoogle Scholar
  81. Zivcak M, Brestic M, Balatová Z, Drevenaková P, Olsovska K, Kalaji HM, Allakhverdiev SI (2013) Photosynthetic electron transport and specific photoprotective responses in wheat leaves under drought stress. Photosynth Res 117:529–546CrossRefPubMedGoogle Scholar
  82. Zivcak M, Brestic M, Kalaji HM, Govindjee (2014a) Photosynthetic responses of sun-and shade-grown barley leaves to high light: is the lower PSII connectivity in shade leaves associated with protection against excess of light? Photosynth Res 119:339–354PubMedCentralCrossRefPubMedGoogle Scholar
  83. Zivcak M, Kalaji HM, Shao HB, Olšovská K, Brestič M (2014b) Photosynthetic proton and electron transport in wheat leaves under prolonged moderate drought stress. J Photochem Photobiol B 137:107–115CrossRefPubMedGoogle Scholar

Copyright information

© Springer Science+Business Media Dordrecht 2015

Authors and Affiliations

  • Marek Zivcak
    • 1
  • Marian Brestic
    • 1
    Email author
  • Kristyna Kunderlikova
    • 1
  • Oksana Sytar
    • 1
    • 2
  • Suleyman I. Allakhverdiev
    • 3
    • 4
    • 5
    Email author
  1. 1.Department of Plant PhysiologySlovak Agricultural UniversityNitraSlovak Republic
  2. 2.Department of Plant Physiology and EcologyTaras Shevchenko National University of KyivKyivUkraine
  3. 3.Institute of Plant PhysiologyRussian Academy of SciencesMoscowRussia
  4. 4.Institute of Basic Biological ProblemsRussian Academy of SciencesPushchinoRussia
  5. 5.Department of Plant Physiology, Faculty of BiologyM.V. Lomonosov Moscow State UniversityMoscowRussia

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