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Volumn 66, Issue 4, 2008, Pages 361-378

Generation of superoxide anion in chloroplasts of Arabidopsis thaliana during active photosynthesis: A focus on rapidly induced genes

Author keywords

Antioxidant response; Chloroplast; Hsp; Oxidative stress; WRKY

Indexed keywords

CHLOROPHYLL; ELECTROLYTE; ENZYME; PLANT DNA; SUPEROXIDE; VEGETABLE PROTEIN;

EID: 43049145797     PISSN: 01674412     EISSN: None     Source Type: Journal    
DOI: 10.1007/s11103-007-9274-4     Document Type: Article
Times cited : (189)

References (59)
  • 1
    • 3242715114 scopus 로고    scopus 로고
    • Reactive oxygen species: Metabolism, oxidative stress, and signal transduction
    • Apel K, Hirt H (2004) Reactive oxygen species: metabolism, oxidative stress, and signal transduction. Annu Rev Plant Biol 55:373-399
    • (2004) Annu Rev Plant Biol , vol.55 , pp. 373-399
    • Apel, K.1    Hirt, H.2
  • 2
    • 0037298310 scopus 로고    scopus 로고
    • Redox regulation of thylakoid protein phosphorylation
    • Aro EM, Ohad I (2003) Redox regulation of thylakoid protein phosphorylation. Antioxid Redox Signal 5:55-67
    • (2003) Antioxid Redox Signal , vol.5 , pp. 55-67
    • Aro, E.M.1    Ohad, I.2
  • 4
    • 0000682385 scopus 로고
    • Production and scavenging of active oxygens in photosynthesis
    • Elsevier/North Holland Biomedical Press Amsterdam
    • Asada K, Takahashi M (1987) Production and scavenging of active oxygens in photosynthesis. In: Kyle DJ, Osmond CB, Arntzen DJ (eds) Photoinhibition. Elsevier/North Holland Biomedical Press, Amsterdam, pp 227-280
    • (1987) Photoinhibition , pp. 227-280
    • Asada, K.1    Takahashi, M.2    Kyle, D.J.3    Osmond, C.B.4    Arntzen, D.J.5
  • 5
    • 0028685490 scopus 로고
    • Fitting a mixture model by expectation maximization to discover motifs in biopolymers
    • Bailey TL, Elkan C (1994) Fitting a mixture model by expectation maximization to discover motifs in biopolymers. Proc Int Conf Intell Syst Mol Biol 2:28-36
    • (1994) Proc Int Conf Intell Syst Mol Biol , vol.2 , pp. 28-36
    • Bailey, T.L.1    Elkan, C.2
  • 6
    • 4043181966 scopus 로고    scopus 로고
    • Applications of chlorophyll fluorescence can improve crop production strategies: An examination of future possibilities
    • Baker NR, Rosenqvist E (2004) Applications of chlorophyll fluorescence can improve crop production strategies: an examination of future possibilities. J Exp Bot 55:1607-1621
    • (2004) J Exp Bot , vol.55 , pp. 1607-1621
    • Baker, N.R.1    Rosenqvist, E.2
  • 7
    • 0015153416 scopus 로고
    • Superoxide dismutase: Improved assays and an assay applicable to acrylamide gels
    • Beauchamp C, Fridovich I (1971) Superoxide dismutase: improved assays and an assay applicable to acrylamide gels. Anal Biochem 44:276-287
    • (1971) Anal Biochem , vol.44 , pp. 276-287
    • Beauchamp, C.1    Fridovich, I.2
  • 8
    • 0030267627 scopus 로고    scopus 로고
    • Molecular chaperones and protein folding in plants
    • Boston RS, Viitanen PV, Vierling E (1996) Molecular chaperones and protein folding in plants. Plant Mol Biol 32:191-222
    • (1996) Plant Mol Biol , vol.32 , pp. 191-222
    • Boston, R.S.1    Viitanen, P.V.2    Vierling, E.3
  • 9
    • 0036598460 scopus 로고    scopus 로고
    • Transcriptional profiling reveals novel interactions between wounding, pathogen, abiotic stress, and hormonal responses in Arabidopsis
    • Cheong YH, Chang HS, Gupta R, Wang X, Zhu T, Luan S (2002) Transcriptional profiling reveals novel interactions between wounding, pathogen, abiotic stress, and hormonal responses in Arabidopsis. Plant Physiol 129:661-677
    • (2002) Plant Physiol , vol.129 , pp. 661-677
    • Cheong, Y.H.1    Chang, H.S.2    Gupta, R.3    Wang, X.4    Zhu, T.5    Luan, S.6
  • 10
    • 0032447801 scopus 로고    scopus 로고
    • Floral dip: A simplified method for Agrobacterium-mediated transformation of Arabidopsis thaliana
    • Clough SJ, Bent AF (1998) Floral dip: a simplified method for Agrobacterium-mediated transformation of Arabidopsis thaliana. Plant J 16:735-743
    • (1998) Plant J , vol.16 , pp. 735-743
    • Clough, S.J.1    Bent, A.F.2
  • 11
    • 33344455077 scopus 로고    scopus 로고
    • The zinc-finger protein Zat12 plays a central role in reactive oxygen and abiotic stress signaling in Arabidopsis
    • Davletova S, Schlauch K, Coutu J, Mittler R (2005) The zinc-finger protein Zat12 plays a central role in reactive oxygen and abiotic stress signaling in Arabidopsis. Plant Physiol 139:847-856
    • (2005) Plant Physiol , vol.139 , pp. 847-856
    • Davletova, S.1    Schlauch, K.2    Coutu, J.3    Mittler, R.4
  • 12
    • 0034062690 scopus 로고    scopus 로고
    • Hydrogen peroxide-induced gene expression in Arabidopsis thaliana
    • Desikan R, Neill SJ, Hancock JT (2000) Hydrogen peroxide-induced gene expression in Arabidopsis thaliana. Free Radic Biol Med 28:773-778
    • (2000) Free Radic Biol Med , vol.28 , pp. 773-778
    • Desikan, R.1    Neill, S.J.2    Hancock, J.T.3
  • 15
    • 0036859947 scopus 로고    scopus 로고
    • Three redundant brassinosteroid early response genes encode putative bHLH transcription factors required for normal growth
    • Friedrichsen DM, Nemhauser J, Muramitsu T, Maloof JN, Alonso J, Ecker JR, Furuya M, Chory J (2002) Three redundant brassinosteroid early response genes encode putative bHLH transcription factors required for normal growth. Genetics 162:1445-1456
    • (2002) Genetics , vol.162 , pp. 1445-1456
    • Friedrichsen, D.M.1    Nemhauser, J.2    Muramitsu, T.3    Maloof, J.N.4    Alonso, J.5    Ecker, J.R.6    Furuya, M.7    Chory, J.8
  • 17
    • 0041363685 scopus 로고    scopus 로고
    • Oxidative stress and acclimation mechanisms in plants
    • Somerville C, Meyerowitz E (eds) American Society of Plant Biologists, Rockville, MD, doi: 10.1199/tab.0009
    • Grene R (2002) Oxidative stress and acclimation mechanisms in plants. In: Somerville C, Meyerowitz E (eds) The Arabidopsis book. American Society of Plant Biologists, Rockville, MD, doi: 10.1199/tab.0009, www.aspb.org/ publications/arabidopsis/
    • (2002) The Arabidopsis Book
    • Grene, R.1
  • 18
    • 0027141553 scopus 로고
    • Overexpression of superoxide dismutase protects plants from oxidative stress (Induction of ascorbate peroxidase in superoxide dismutase-overexpressing plants)
    • Gupta AS, Webb RP, Holaday AS, Allen RD (1993) Overexpression of superoxide dismutase protects plants from oxidative stress (Induction of ascorbate peroxidase in superoxide dismutase-overexpressing plants). Plant Physiol 103:1067-1073
    • (1993) Plant Physiol , vol.103 , pp. 1067-1073
    • Gupta, A.S.1    Webb, R.P.2    Holaday, A.S.3    Allen, R.D.4
  • 20
    • 0032918558 scopus 로고    scopus 로고
    • Plant cis-acting regulatory DNA elements (PLACE) database: 1999
    • Higo K, Ugawa Y, Iwamoto M, Korenaga T (1999) Plant cis-acting regulatory DNA elements (PLACE) database: 1999. Nucleic Acids Res 27:297-300
    • (1999) Nucleic Acids Res , vol.27 , pp. 297-300
    • Higo, K.1    Ugawa, Y.2    Iwamoto, M.3    Korenaga, T.4
  • 21
    • 0029746193 scopus 로고    scopus 로고
    • Initiation of runaway cell death in an Arabidopsis mutant by extracellular superoxide
    • Jabs T, Dietrich RA, Dangl JL (1996) Initiation of runaway cell death in an Arabidopsis mutant by extracellular superoxide. Science 273:1853-1856
    • (1996) Science , vol.273 , pp. 1853-1856
    • Jabs, T.1    Dietrich, R.A.2    Dangl, J.L.3
  • 22
    • 0000987360 scopus 로고    scopus 로고
    • Changes in the isozyme composition of antioxidant enzymes in response to aminotriazole in leaves of Arabidopsis thaliana
    • Kang KS, Lim CJ, Han TJ, Kim JC, Jin CD (1999) Changes in the isozyme composition of antioxidant enzymes in response to aminotriazole in leaves of Arabidopsis thaliana. J Plant Biol 42:187-193
    • (1999) J Plant Biol , vol.42 , pp. 187-193
    • Kang, K.S.1    Lim, C.J.2    Han, T.J.3    Kim, J.C.4    Jin, C.D.5
  • 24
    • 0042591328 scopus 로고    scopus 로고
    • Reaction mechanism of plant 2-Cys peroxiredoxin. Role of the C terminus and the quaternary structure
    • Konig J, Lotte K, Plessow R, Brockhinke A, Baier M, Dietz KJ (2003) Reaction mechanism of plant 2-Cys peroxiredoxin. Role of the C terminus and the quaternary structure. J Biol Chem 278:24409-24420
    • (2003) J Biol Chem , vol.278 , pp. 24409-24420
    • Konig, J.1    Lotte, K.2    Plessow, R.3    Brockhinke, A.4    Baier, M.5    Dietz, K.J.6
  • 27
    • 77957068711 scopus 로고
    • Chlorophylls and carotenoids: Pigments of photosynthetic biomembranes
    • Packer L, Douce R (eds)
    • Lichtenthaler HK (1987) Chlorophylls and carotenoids: pigments of photosynthetic biomembranes. In: Packer L, Douce R (eds) Methods Enzymol 148:350-382
    • (1987) Methods Enzymol , vol.148 , pp. 350-382
    • Lichtenthaler, H.K.1
  • 28
    • 34548503443 scopus 로고    scopus 로고
    • Chloroplast-generated reactive oxygen species are involved in hypersensitive response-like cell death mediated by a mitogen-activated protein kinase cascade
    • Liu Y, Ren D, Pike S, Pallardy S, Gassmann W, Zhang S (2007) Chloroplast-generated reactive oxygen species are involved in hypersensitive response-like cell death mediated by a mitogen-activated protein kinase cascade. Plant J 51:941-954
    • (2007) Plant J , vol.51 , pp. 941-954
    • Liu, Y.1    Ren, D.2    Pike, S.3    Pallardy, S.4    Gassmann, W.5    Zhang, S.6
  • 29
    • 0027217043 scopus 로고
    • Direct detection of singlet oxygen from isolated Photosystem II reaction centres
    • Macpherson AN, Telfer A, Barber J, Truscott TG (1993) Direct detection of singlet oxygen from isolated Photosystem II reaction centres. Biochim Biophys Acta 1143:301-309
    • (1993) Biochim Biophys Acta , vol.1143 , pp. 301-309
    • MacPherson, A.N.1    Telfer, A.2    Barber, J.3    Truscott, T.G.4
  • 30
    • 0024545871 scopus 로고
    • Superoxide-dependent nitroblue tetrazolium reduction and expression of cytochrome b-245 components by human tonsillar B lymphocytes and B cell lines
    • Maly FE, Nakamura M, Gauchat JF, Urwyler A, Walker C, Dahinden CA, Cross AR, Jones OT, De Weck AL (1989) Superoxide-dependent nitroblue tetrazolium reduction and expression of cytochrome b-245 components by human tonsillar B lymphocytes and B cell lines. J Immunol 142:1260-1267
    • (1989) J Immunol , vol.142 , pp. 1260-1267
    • Maly, F.E.1    Nakamura, M.2    Gauchat, J.F.3    Urwyler, A.4    Walker, C.5    Dahinden, C.A.6    Cross, A.R.7    Jones, O.T.8    De Weck, A.L.9
  • 31
    • 0003092129 scopus 로고
    • Studies on reactions of illuminated chloroplasts. I. Mechanism of the reduction of oxygen and other Hill reagents
    • Mehler AH (1951) Studies on reactions of illuminated chloroplasts. I. Mechanism of the reduction of oxygen and other Hill reagents. Arch Biochem 33:65-77
    • (1951) Arch Biochem , vol.33 , pp. 65-77
    • Mehler, A.H.1
  • 32
    • 0036728244 scopus 로고    scopus 로고
    • Oxidative stress, antioxidants and stress tolerance
    • Mittler R (2002) Oxidative stress, antioxidants and stress tolerance. Trends Plant Sci 7:405-410
    • (2002) Trends Plant Sci , vol.7 , pp. 405-410
    • Mittler, R.1
  • 34
    • 33750551504 scopus 로고    scopus 로고
    • Arabidopsis heat shock transcription factor A2 as a key regulator in response to several types of environmental stress
    • Nishizawa A, Yabuta Y, Yoshida E, Maruta T, Yoshimura K, Shigeoka S (2006) Arabidopsis heat shock transcription factor A2 as a key regulator in response to several types of environmental stress. Plant J 48:535-547
    • (2006) Plant J , vol.48 , pp. 535-547
    • Nishizawa, A.1    Yabuta, Y.2    Yoshida, E.3    Maruta, T.4    Yoshimura, K.5    Shigeoka, S.6
  • 35
    • 0033506468 scopus 로고    scopus 로고
    • Photoprotection revisited: Genetic and molecular approaches
    • Niyogi KK (1999) Photoprotection revisited: genetic and molecular approaches. Annu Rev Plant Physiol Plant Mol Biol 50:333-359
    • (1999) Annu Rev Plant Physiol Plant Mol Biol , vol.50 , pp. 333-359
    • Niyogi, K.K.1
  • 36
    • 0034804489 scopus 로고    scopus 로고
    • Arabidopsis and the heat stress transcription factor world: How many heat stress transcription factors do we need?
    • Nover L, Bharti K, Doring P, Mishra SK, Ganguli A, Scharf KD (2001) Arabidopsis and the heat stress transcription factor world: how many heat stress transcription factors do we need? Cell Stress Chaperones 6:177-189
    • (2001) Cell Stress Chaperones , vol.6 , pp. 177-189
    • Nover, L.1    Bharti, K.2    Doring, P.3    Mishra, S.K.4    Ganguli, A.5    Scharf, K.D.6
  • 38
    • 0032753804 scopus 로고    scopus 로고
    • The role of photosynthetic electron transport in the oxidative degradation of chloroplastic glutamine synthetase
    • Palatnik JF, Carrillo N, Valle EM (1999) The role of photosynthetic electron transport in the oxidative degradation of chloroplastic glutamine synthetase. Plant Physiol 121:471-478
    • (1999) Plant Physiol , vol.121 , pp. 471-478
    • Palatnik, J.F.1    Carrillo, N.2    Valle, E.M.3
  • 40
    • 0345306696 scopus 로고    scopus 로고
    • Precursors with altered affinity for Hsp70 in their transit peptides are efficiently imported into chloroplasts
    • Rial DV, Ottado J, Ceccarelli EA (2003) Precursors with altered affinity for Hsp70 in their transit peptides are efficiently imported into chloroplasts. J Biol Chem 278:46473-46481
    • (2003) J Biol Chem , vol.278 , pp. 46473-46481
    • Rial, D.V.1    Ottado, J.2    Ceccarelli, E.A.3
  • 41
    • 33750942076 scopus 로고    scopus 로고
    • Chloroplast Hsp70s are not involved in the import of ferredoxin-NADP+ reductase precursor
    • Rial DV, Arakaki AK, Almara AM, Orellano EG, Ceccarelli EA (2006) Chloroplast Hsp70s are not involved in the import of ferredoxin-NADP+ reductase precursor. Physiol Plant 128:618-632
    • (2006) Physiol Plant , vol.128 , pp. 618-632
    • Rial, D.V.1    Arakaki, A.K.2    Almara, A.M.3    Orellano, E.G.4    Ceccarelli, E.A.5
  • 42
    • 1642564552 scopus 로고    scopus 로고
    • The zinc finger protein Zat12 is required for cytosolic ascorbate peroxidase 1 expression during oxidative stress in Arabidopsis
    • Rizhsky L, Davletova S, Liang H, Mittler R (2004a) The zinc finger protein Zat12 is required for cytosolic ascorbate peroxidase 1 expression during oxidative stress in Arabidopsis. J Biol Chem 279:11736-11743
    • (2004) J Biol Chem , vol.279 , pp. 11736-11743
    • Rizhsky, L.1    Davletova, S.2    Liang, H.3    Mittler, R.4
  • 43
    • 1942533606 scopus 로고    scopus 로고
    • When defense pathways collide. the response of Arabidopsis to a combination of drought and heat stress
    • Rizhsky L, Liang H, Shuman J, Shulaev V, Davletova S, Mittler R (2004b) When defense pathways collide. The response of Arabidopsis to a combination of drought and heat stress. Plant Physiol 134:1683-1696
    • (2004) Plant Physiol , vol.134 , pp. 1683-1696
    • Rizhsky, L.1    Liang, H.2    Shuman, J.3    Shulaev, V.4    Davletova, S.5    Mittler, R.6
  • 44
    • 0034755053 scopus 로고    scopus 로고
    • A new member of the Arabidopsis WRKY transcription factor family, AtWRKY6, is associated with both senescence- and defence-related processes
    • Robatzek S, Somssich IE (2001) A new member of the Arabidopsis WRKY transcription factor family, AtWRKY6, is associated with both senescence- and defence-related processes. Plant J 28:123-133
    • (2001) Plant J , vol.28 , pp. 123-133
    • Robatzek, S.1    Somssich, I.E.2
  • 45
    • 0036851187 scopus 로고    scopus 로고
    • Global changes in gene expression in response to high light in Arabidopsis
    • Rossel JB, Wilson IW, Pogson BJ (2002) Global changes in gene expression in response to high light in Arabidopsis. Plant Physiol 130:1109-1120
    • (2002) Plant Physiol , vol.130 , pp. 1109-1120
    • Rossel, J.B.1    Wilson, I.W.2    Pogson, B.J.3
  • 47
    • 33846608278 scopus 로고    scopus 로고
    • Plant nutritional status modulates glutamine synthetase levels in ripe tomatoes (Solanum lycopersicum cv. Micro-Tom)
    • Scarpeci TE, Marro ML, Bortolotti S, Boggio SB, Valle EM (2007) Plant nutritional status modulates glutamine synthetase levels in ripe tomatoes (Solanum lycopersicum cv. Micro-Tom). J Plant Physiol 164:137-145
    • (2007) J Plant Physiol , vol.164 , pp. 137-145
    • Scarpeci, T.E.1    Marro, M.L.2    Bortolotti, S.3    Boggio, S.B.4    Valle, E.M.5
  • 48
    • 33646346645 scopus 로고    scopus 로고
    • The heat stress transcription factor HsfA2 serves as a regulatory amplifier of a subset of genes in the heat stress response in Arabidopsis
    • Schramm F, Ganguli A, Kiehlmann E, Englich G, Walch D, von Koskull-Doring P (2006) The heat stress transcription factor HsfA2 serves as a regulatory amplifier of a subset of genes in the heat stress response in Arabidopsis. Plant Mol Biol 60:759-772
    • (2006) Plant Mol Biol , vol.60 , pp. 759-772
    • Schramm, F.1    Ganguli, A.2    Kiehlmann, E.3    Englich, G.4    Walch, D.5    Von Koskull-Doring, P.6
  • 49
    • 0037136230 scopus 로고    scopus 로고
    • Small heat shock proteins and stress tolerance in plants
    • Sun W, Van Montagu M, Verbruggen N (2002) Small heat shock proteins and stress tolerance in plants. Biochim Biophys Acta 1577:1-9
    • (2002) Biochim Biophys Acta , vol.1577 , pp. 1-9
    • Sun, W.1    Van Montagu, M.2    Verbruggen, N.3
  • 50
    • 0034978496 scopus 로고    scopus 로고
    • Comprehensive expression profile analysis of the Arabidopsis Hsp70 gene family
    • Sung DY, Vierling E, Guy CL (2001) Comprehensive expression profile analysis of the Arabidopsis Hsp70 gene family. Plant Physiol 126:789-800
    • (2001) Plant Physiol , vol.126 , pp. 789-800
    • Sung, D.Y.1    Vierling, E.2    Guy, C.L.3
  • 51
    • 12144291195 scopus 로고    scopus 로고
    • MAPMAN: A user-driven tool to display genomics data sets onto diagrams of metabolic pathways and other biological processes
    • Thimm O, Blasing O, Gibon Y, Nagel A, Meyer S, Kruger P, Selbig J, Muller LA, Rhee SY, Stitt M (2004) MAPMAN: a user-driven tool to display genomics data sets onto diagrams of metabolic pathways and other biological processes. Plant J 37:914-939
    • (2004) Plant J , vol.37 , pp. 914-939
    • Thimm, O.1    Blasing, O.2    Gibon, Y.3    Nagel, A.4    Meyer, S.5    Kruger, P.6    Selbig, J.7    Muller, L.A.8    Rhee, S.Y.9    Stitt, M.10
  • 52
    • 0030748530 scopus 로고    scopus 로고
    • 2 accumulation in papillae and hypersensitive response during the barley-powdery mildew interaction
    • 2 accumulation in papillae and hypersensitive response during the barley-powdery mildew interaction. Plant J 11:1187-1194
    • (1997) Plant J , vol.11 , pp. 1187-1194
    • Thordal-Christensen, H.1    Zhang, Z.2    Wei, Y.3    Collinge, D.B.4
  • 55
    • 33644653602 scopus 로고    scopus 로고
    • Genome-wide analysis of hydrogen peroxide-regulated gene expression in Arabidopsis reveals a high light-induced transcriptional cluster involved in anthocyanin biosynthesis
    • Vanderauwera S, Zimmermann P, Rombauts S, Vandenabeele S, Langebartels C, Gruissem W, Inze D, Van Breusegem F (2005) Genome-wide analysis of hydrogen peroxide-regulated gene expression in Arabidopsis reveals a high light-induced transcriptional cluster involved in anthocyanin biosynthesis. Plant Physiol 139:806-821
    • (2005) Plant Physiol , vol.139 , pp. 806-821
    • Vanderauwera, S.1    Zimmermann, P.2    Rombauts, S.3    Vandenabeele, S.4    Langebartels, C.5    Gruissem, W.6    Inze, D.7    Van Breusegem, F.8
  • 57
    • 33745466148 scopus 로고    scopus 로고
    • Physical and functional interactions between pathogen-induced Arabidopsis WRKY18, WRKY40, and WRKY60 transcription factors
    • Xu X, Chen C, Fan B, Chen Z (2006) Physical and functional interactions between pathogen-induced Arabidopsis WRKY18, WRKY40, and WRKY60 transcription factors. Plant Cell 18:1310-1326
    • (2006) Plant Cell , vol.18 , pp. 1310-1326
    • Xu, X.1    Chen, C.2    Fan, B.3    Chen, Z.4
  • 58
    • 15744382453 scopus 로고    scopus 로고
    • Identification of a novel class of target genes and a novel type of binding sequence of heat shock transcription factor in Saccharomyces cerevisiae
    • Yamamoto A, Mizukami Y, Sakurai H (2005) Identification of a novel class of target genes and a novel type of binding sequence of heat shock transcription factor in Saccharomyces cerevisiae. J Biol Chem 280:11911-11919
    • (2005) J Biol Chem , vol.280 , pp. 11911-11919
    • Yamamoto, A.1    Mizukami, Y.2    Sakurai, H.3
  • 59
    • 13444264729 scopus 로고    scopus 로고
    • GENEVESTIGATOR. Arabidopsis microarray database and analysis toolbox
    • Zimmermann P, Hirsch-Hoffmann M, Hennig L, Gruissem W (2004) GENEVESTIGATOR. Arabidopsis microarray database and analysis toolbox. Plant Physiol 136:2621-2632
    • (2004) Plant Physiol , vol.136 , pp. 2621-2632
    • Zimmermann, P.1    Hirsch-Hoffmann, M.2    Hennig, L.3    Gruissem, W.4


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