Summary
Ascorbate, glutathione, and tocopherol are the three major low molecular weight antioxidants of plant cells. While tocopherol is hydrophobic and is found only in lipid membranes, ascorbate and glutathione are hydrophilic, accumulating to high concentrations in the chloroplast stroma and other compartments of the plant cell. Ascorbate and glutathione not only limit photo-oxidative damage but can also act independently as signal-transducing molecules regulating defense gene expression. Both metabolites transmit information concerning oxidative load and redoxbuffering capacity. Ascorbate modifies the expression of chloroplast genes. Net glutathione synthesis during stress restores the cellular redox state and allows orchestration of systemic acquired resistance. The degree of redox coupling between these antioxidants has profound implications for regulation, function, and signaling associated with the two major energy-generating systems, i.e. photosynthesis and respiration. Tocopherol fulfills an essential protective function, counter-acting the harmful effects of singlet oxygen production at photosystem II. Ascorbate reduces and thus regenerates oxidized tocopherol, but flux through this reaction is not sufficient to maintain the reduced tocopherol pool under high light stress. This may be because tocopherol regeneration draws on the ascorbate pool of the chloroplast lumen, which may be depleted under stress. Moreover, while glutathione always reduces oxidized ascorbate (dehydroascorbate), the degree of coupling between the ascorbate and glutathione redox couples is variable. The flexibility of coupling between these antioxidant pools is crucial to differential redox signaling, particularly by ascorbate and glutathione.
Access this chapter
Tax calculation will be finalised at checkout
Purchases are for personal use only
Preview
Unable to display preview. Download preview PDF.
References
Adir N, Zer H, Shochat S and Ohad I (2003) Photoinhibition—a historical perspective. Photosynth Res 76: 343–370
Agius F, Gonz’alez-Lamothe R, Caballero JL, Mu˜noz-Blanco J, Botella MA and Valpuesta V (2003) Engineering increased vitamin C levels in plants by overexpression of a D-galacturonic acid reductase. Nature Biotechnol 21: 177–181
Allen JF and Hall DO (1973) Superoxide reduction as a mechanism of ascorbate-stimulated oxygen uptake by isolated chloroplasts. Biochem Biophys Res Commun 52: 856–862
Alscher RG (1989) Biosynthesis and antioxidant function of glutathione in plants. Physiol Plant 77: 457–64
Alvarez ME, Penell RI, Meijer PJ, Ishikawa A, Dixon RA and Lamb C (1998) Reactive oxygen intermediates mediate a systemic signal network in the establishment of plant immunity. Cell 92: 773–784
Amako K, Chen G-X and Asada K (1994) Separate assays specific for ascorbate peroxidase and guaiacol peroxidase and for the chloroplastic and cytosolic isozymes of ascorbate peroxidase in plants. Plant Cell Physiol 35: 497–504
Anderson JW, Foyer CH and Walker DA (1983a) Lightdependent reduction of hydrogen peroxide by intact spinach chloroplasts. Biochim Biophys Acta 724: 69–74
Anderson JW, Foyer CH and Walker DA (1983b) Lightdependent reduction of dehydroascorbate and uptake of exogenous ascorbate by spinach chloroplasts. Planta 158: 442–450
Angerhofer A (1991) Chlorophyll triplets and radical pairs. In: Scheer H (ed), Chlorophylls, pp 945–992. CRC Press, Boca Raton
Aro E-M, Virgin I and Andersson B (1993) Photoinhibition of photosystem II: inactivation, protein damage and turnover. Biochem Biophys Acta 1143: 113–134
Arrigoni O and de Tullio MC (2000) The role of ascorbic acid in cell metabolism: between gene-directed functions and unpredictable chemical reactions. J Plant Physiol 157: 481–488
Asada K (1999) The water-water cycle in chloroplasts: Scavenging of active oxygens and dissipation of excess photons. Annu Rev Plant Physiol Plant Mol Biol 50: 601–639
Asard H, Kapila J, Verelst W and B’erczi A (2001) Higher-plant plasma membrane cytochrome b561: a protein in search of a function. Protoplasma 217: 77–93
Baier M and Dietz KJ (1997) The plant 2-cys peroxiredoxin BAS1 is a nuclear-encoded chloroplast protein: its expressional regulation, phylogenetic origin, and implications for its specific physiological function in plants. Plant J 12: 179–190
BaierMand Dietz K-J (1999a) Alkyl hydroperoxide reductases: the way out of the oxidative breakdown of lipids in chloroplasts. Trends Plant Sci 4: 166–168
Baier M and Dietz K-J (1999b) Protective function of chloroplast 2-cysteine peroxiredoxin in photosynthesis. Evidence from transgenic Arabidopsis. Plant Physiol 119: 1407–1414
Baier M, Noctor G, Foyer CH and Dietz KJ (2000) Antisense suppression of 2-cysteine peroxiredoxin in Arabidopsis specifically enhances the activities and expression of enzymes associated with ascorbate metabolism but not glutathione metabolism. Plant Physiol 124: 823–832
Ball L, Accotto G, Bechtold U, Creissen G, Funck D, Jimenez A, Kular B, Leyland N, Mejia-Carranza J, Reynolds H, Karpinski S and Mullineaux PM (2004) Evidence for a direct link between glutathione biosynthesis and stress defense gene expression in Arabidopsis. Plant Cell 16: 2448–2462
Bartling D, Radzio R, Steiner U and Weiler EW (1993) A glutathione-S-transferase with glutathione peroxidase activity from Arabidopsis thaliana. Molecular cloning and functional characterization. Eur J Biochem 216: 579–586
Bartoli CG, Pastori GM and Foyer CH (2000) Ascorbate biosynthesis in mitochondria is linked to the electron transport chain between complexes III and IV. Plant Physiol 123: 335–343
Bartoli CG, Guiamet JJ, Kiddle G, Pastori G, Di Cagno R, Theodoulou FL and Foyer CH (2004) The relationship between L-galactono-1, 4-lactone dehydrogenase (GalLDH) and ascorbate content in leaves under optimal and stress conditions. Plant Cell Environ, In press
Berczi A and Møller IM (1998) NADH-monodehydroascorbate oxidoreductase is one of the redox enzymes in spinach leaf plasma membranes. Plant Physiol 116: 1029–1036
Bergmüller E, Porfirova S and Dörmann P (2003) Characterization of an Arapidopsis mutant deficient in γ -tocopherol methyltransferase. Plant Mol Biol 52: 1181–1190
Beyer RE (1990) The participation of coenzyme Q in free radical production and antioxidation. Free Radic BiolMed8: 545–565
Bick JA, Setterdahl AT, Knaff DB, Chen Y, Pitcher LH, Zilinskas BA and Leustek T (2001) Regulation of the plant-type 5’-adenylylsulfate reductase by oxidative stress. Biochemistry 40: 9040–9048
Bishop NI and Wong J (1974) Photochemical characteristics of a vitamin E deficient mutant of Scenedesmus obliquus. Ber dtsch bot Ges 87: 359–371
Boveris A, Oshino N and Chance B (1972) The cellular production of hydrogen peroxide. Biochem J 128: 617–630
Bramley PM, Elmadf’a I, Kafatos A, Kelly FJ, Manios Y, Roxborough HE, Schuch W, Sheehy PJA and Wagner KH (2000) Vitamin E. J Sci Food Agric 80: 913–938
Booij-James I, Swegle W M, Edelman M and Mattoo A (2002) Phosphorylation of the D1 photosystem II reaction center protein is controlled by an endogenous circadian rhythm. Plant Physiol 130: 2069–2075
Bowsher CG, Hucklesby DP and Emes MJ (1989) Nitrite reduction and carbohydrate metabolism in plastids purified from roots of Pisum sativum L. Planta 177: 359–366
Breitenbach J, ZhuCand Sandmann G(2001) The bleaching herbicice norflurazon inhibits phytoene desaturase by competetion with the cofactors. J Agric Food Chem 49: 5270–5272
Buchanan BB (1991) Regulation of CO2 assimilation in oxygenic photosynthesis: the ferredoxin/thioredoxin system. Arch Biochem Biophys 228: 1–9
Bunkelmann JR and Trelease RN (1996) Ascorbate peroxidase. A prominent membrane protein in oilseed glyoxysomes. Plant Physiol 110: 589–598
BurkeyKO, EasonGand Fiscus EL (2003) Factors that affect leaf extracellular ascorbic acid content and redox status. Physiol Plant 117: 51–57
Chamnongpol S, Willekens H, Moeder W, Langebartels C, Sandermann H Jr, Van Montagu M, Inze D and Van Camp W (1998) Defense activation and enhanced pathogen tolerance induced by H2O2 in transgenic tobacco. Proc Natl Acad Sci USA 95: 5818–5823
Chen G and Asada K (1989) Ascorbate peroxidase in tea leaves: Occurrence of two isozymes and the differences in their enzymatic and molecular properties. Plant Cell Physiol 30: 987– 998
Chen Z and Gallie DR (2004). The ascorbic acid redox state controls guard cell signaling and stomatal movement. Plant Cell 16: 1143–1162
Chen Z, Young TE, Ling J, Chang SC and Gallie DR (2003) Increasing vitamin C content of plants through enhanced ascorbate recycling. Proc Natl Acad Sci USA 100: 3525–3530
Cogdell R and Frank HA (1996) Carotenoids in photosynthesis. Photochem Photobiol 63: 257–264
Collakova E and DellaPenna D (2001) Isolation and functional analysis of homogentisate phytyltransferase from Synechocystis sp. PCC 6803 and Arabidopsis. Plant Physiol 127: 1113– 1124
Collakova E and DellaPenna D (2003a) Homogentisate phytyltransferase activity is limiting for tocopherol biosynthesis in Arabidopsis. Plant Physiol 131: 632–642
Collakova E and DellaPenna D (2003b) The role of homogentisate phytyltransferase and other tocopherol pathway enzymes in the regulation of tocopherol synthesis during abiotic stress. Plant Physiol 131: 930–940
Collin V, Issakidis-Bourguet E, Marchand C, Hirasawa M, Lancelin JM, Knaff DB and Miginiac-Maslow M (2003) The Arabidopsis plastidial thioredoxins: new functions and new insights into specificity. J Biol Chem 278: 23747–23752
Collinson EJ, Wheeler GL, Garrido EO, Avery AM, Avery SV and Grant CM (2002) The yeast glutaredoxins are active as glutathione peroxidases. J Biol Chem 277: 16712–16717
Conklin PL and Barth C (2004) Ascorbic acid, a familiar small molecule intertwined in the response of plants to ozone, pathogens and the onset of senescence. Plant Cell Environ 27: 959–970
Conklin PL, Williams EH and Last RL (1996) Environmental stress sensitivity of an ascorbic acid-deficient Arabidopsis mutant. Proc Natl Acad Sci USA 93: 9970–9974
Dähnhardt D, Falk J, Appel J, van der Kooij TAW, Schulz- Friedrich R and Krupinska K (2002) The hydroxyphenylpyruvate dioxygenase from Synechocystis sp. PCC 6803 is not required for plastoquinone biosynthesis. FEBS Lett 523: 177– 181
Dat J, Vandenabeele S, Vranov’a E, Van Montagu M, Inz’e D and Van Breusegem F (2000) Dual action of AOS during plant stress responses. Cell Mol Life Sci 57: 779–795
Davey MW, Gilot C, Persiau G, Ostergaard J, Han Y, Bauw GC and Van Montagu MC (1999) Ascorbate biosynthesis in Arabidopsis cell suspension culture. Plant Physiol 121: 535–543
Delauney A, Pflieger D, Barrault MB, Vinh J and Toledano MB (2002) A thiol peroxidase is an H2O2 receptor and redoxtransducer in gene activation. Cell 111: 1–11
Demmig-Adams B and Adams WW III (1992) Photoprotection and other responses of plants to high light stress. Annu Rev Plant Physiol Plant Mol Biol 43: 599–626
Dietz KJ (2003) Plant peroxiredoxins. Annu Rev Plant Biol 54: 93–107
Dietz K-J, Stork T, Finkemeier I, Lamkemeyer P, Li W-X, El-Tayeb MA, Michel K-P, Pistorius E and Baier M (2005) The role of peroxiredoxins in oxygenic photosynthesis of cyanobacteria and higher plants: peroxide detoxification or redox sensing? In: Demmig-Adams B, Adams WW III and MattooAK(eds) Photoprotection, Photoinhibition, Gene Regulation, and Environment, pp 303–319. Springer, Dordrecht
Diner BA and Rappoport F (2002) Structure, dynamics, and energetics of the primary photochemistry of photosystem II of oxygenic photosynthesis. Annu Rev Plant Biol 53: 552– 580
Dixon DP, Davis BG and Edwards R (2002) Functional divergence in the glutathione transferase superfamily in plants. Identification of two classes with putative functions in redox homeostasis in Arabidopsis thaliana. J BiolChem277: 30859– 30869
Dron M, ClouseSD, Dixon RA, LawtonMAand Lamb CJ (1988) Glutathione and fungal elicitor regulation of a plant defense gene promoter in electroporated protoplasts. Proc Natl Acad Sci USA 85: 6738–6742
Du G, Mouithys-Mickalad A and Sluse FE (1998) Generation of superoxide anion by mitochondria and impairment of their functions during anoxia and reoxygenation in vitro. Free Radic Biol Med 25: 1066–1074
Duke SO, Lydon JM, Becerril TD, Sherman L and Matsumoto H (1991) Protoporphyrinogen oxidase inhibiting herbicides. Weed Sci 39: 465–473
Durrant JB, Giorgi LB, Barber J, Klug DR and Porter G (1990) Characterization of triplet states in isolated photosystem II reaction centers: oxygen quenching as a mechanism for photodamage. Biochim Biophys Acta 1017: 167–175
Dutilleul C, Driscoll S, Cornic G, De Paepe R, Foyer CH and Noctor G (2003a) Functional mitochondrial complex I is required by tobacco leaves for optimal photosynthetic performance in photorespiratory conditions and during transients. Plant Physiol 313: 264–275
Dutilleul C, Garmier M, Noctor G, Mathieu CD, Ch’etrit P, Foyer CH and De Paepe R (2003b) Leaf mitochondria modulate whole cell redox homeostasis, set antioxidant capacity and determine stress resistance through altered signaling and diurnal regulation. Plant Cell 15: 1212–1226
Elstner EF, Wagner GA and SchultzW(1998) Activated oxygen in green plants in relation to stress situation. Curr Top Plant Biochem Physiol 7: 159–187
Endo T and Asada K (2005) Photosystem I and photoprotection: cyclic electron flow and water-water cycle. In: Demmig-Adams B, Adams WW III and Mattoo AK (eds) Photoprotection, Photoinhibition, Gene Regulation, and Environment, pp 205–221. Springer, Dordrecht
Eshdat Y, Holland D, Faltin Z and Ben-Hayyim G (1997) Plant glutathione peroxidases. Physiol Plant 100: 234–240
Eskling M and Âkerlund HE (1998) Changes in the quantities of violaxanthin de-epoxidase, xanthophylls and ascorbate in spinach upon shift from low to high light. Photosynth Res 57: 41–50
Falk J, Krauβ N, Dähnhardt D and Krupinska K (2002) The senescence associated gene of barley encoding 4-hydroxyphenylpyruvate dioxygenase is expressed during oxidative stress. J Plant Physiol 159: 1245–1253
Fath A, Bethke PC and Jones RL (2001) Enzymes that scavenge reactive oxygen species are down-regulated prior to gibberellic acid-induced programmed cell death in barley aleurone. Plant Physiol 126: 156–166
Feierabend J and Dehne S (1996) Fate of the porphyrin cofactors during the light-dependent turnover of catalase and of the photosystem II reaction center protein D1 in mature rye leaves. Planta 198: 413–422
Ferreira KN, Iverson TM, Maghlaoui K, Barber J and Iwata S (2004) Architecture of the Photosynthetic Oxygen-Evolving Center Science 303: 1831–1838.
Foreman J, Demidchik V, Bothwell JHF, Mylona P, Miedema H, Torres MA, Linstead P, Costa S, Brownlee C, Jones JDG, Davies JM and Dolan L (2003) Reactive oxygen species produced by NADPH oxidase regulate plant cell growth. Nature 422: 442–446
Foyer CH and Halliwell B (1976) The presence of glutathione and glutathione reductase in chloroplasts: a proposed role in ascorbic acid metabolism. Planta 133: 21–25
Foyer CH and Harbinson J (1999) Relationships between antioxidant metabolism and carotenoids in the regulation of photosynthesis. In: Frank HA, Young AJ, Britton G and Cogdell RJ (eds) The Photochemistry of Carotenoids, pp 305–325. Kluwer Academic Publishers, Dordrecht
Foyer CH and Noctor G (2000) Oxygen processing in photosynthesis: regulation and signalling. New Phytol 146: 359–388
Foyer CH and Noctor G (2003) Redox sensing and signaling associated with reactive oxygen in chloroplasts, peroxisomes and mitochondria. Physiol Plant 119: 355–364
Foyer CH, Rowell J and Walker D (1983) Measurements of the ascorbate content of spinach leaf protoplasts and chloroplasts during illumination. Planta 157: 239–244
Foyer CH, Souriau N, Perret S, Lelandais M, Kunert KJ, Pruvost C and Jouanin L (1995) Overexpression of glutathione reductase but not glutathione synthetase leads to increases in antioxidant capacity and resistance to photoinhibition in poplar trees. Plant Physiol 109: 1047–1057
Foyer CH, GomezLDand van Heerden PDR. (2005) Glutathione. In: Smirnoff N (ed) Antioxidants and Reactive Species in Plants, pp 000–000. Blackwell Publishing, London
Fryer MJ (1992) The antioxidant effect of thylakoid vitamin E (α-tocopherol). Plant Cell Environ 15: 381–392
Fufezan C, Rutherford AW and Krieger-Liszkay A(2002) Singlet oxygen production in herbicide-treated photosystem II. FEBS Lett 532: 407–410
Genova ML, Ventura B, Giuliano G, Bovina C, Formiggini G, Castelli GP and Lenaz G (2001) The site of production of superoxide radical in mitochondrial Complex I is not a bound ubisemiquinone but presumably iron-sulfur cluster N2. FEBS Lett 505: 364–368
Gillham DJ and Dodge AD (1986) Hydrogen peroxide scavenging systems within pea chloroplasts. A quantitative study. Planta 167: 246–251
Gomez L, Vanacker H, Buchner P, NoctorGand Foyer CH (2004) Regulation of glutathione metabolism during the short-term chilling response of maize leaves. Plant Physiol 134: 1662– 1671
Graβes T, Grimm B, Koroleva O and Jahns P (2001) Loss of α-tocopherol in tobacco plants with decreased geranylgeranyl reductase activity does not modify photosynthesis in optimal growth conditions but increases sensitivity to high-light stress. Planta 213: 620–628
Green R and Fluhr R (1995) UV-B-induced PR-1 accumulation is mediated by active oxygen species. Plant Cell: 203–212
Gressel J (2002) Molecular Biology of Weed Control. Taylor & Francis, London
Groden D and Beck E (1979) H2O2 destruction by ascorbatedependent systems from chloroplasts. Biochim Biophys Acta 546: 426–435
Han D, Canali R, Rettori D and Cadenas E (2001) Production of superoxide into the intermembrane space and cytoplasm by heart mitochondria. Free Radic Biol Med 31: 45
Hancock JT, Desikan R and Neill SJ (2001) Does the redox status of cytochrome C act as a fail-safe mechanism in the regulation of programmed cell death? Free Rad Biol Medic 31: 697– 703
Havaux M, Lütz C and Grimm B (2003) Chloroplast membrane photostability in chlP transgenic tobacco plants deficient in tocopherols. Plant Physiol 132: 300–310
Heazlewood JL, Howell KA and Millar AH (2003) Mitochondrial complex I from Arabidopsis and rice: orthologs of mammalian and fungal components coupled with plant-specific subunits. Biochim Biophys Acta 1604: 159–169
Herbette S, Lenne C, Leblanc N, Julien JL, Drevet JR and Roeckel-Drevet P (2002) Two GPX-like proteins from Lycopersicon esculentum and Helianthus annuus are antioxidant enzymes with phospholipid hydroperoxide glutathione peroxidase and thioredoxin peroxidase activities. Eur J Biochem 269: 2414–2420
Hideg E, Spetea C and Vass I (1994) Singlet oxygen production in thylakoid membranes during photoinhibition as detected by ESR spectroscopy. Photosynth Res 39: 191–199
Hideg E, Kalai T, Hideg K and Vass I (1998) Photoinhibition of photosynthesis in vivo results in singlet oxygen production. Detection via nitroxide-induced fluorescence quenching in broad bean leaves. Biochemistry 37: 11405–11411
Hideg E, Barta C, Kalai T, Vass I, Hideg K and Asada K (2002) Detection of singlet oxygen and superoxide with fluorescent sensors in leaves under stress by photoinhibition or UV radiation. Plant Cell Physiol 43: 1154–1164
Hofius D and Sonnewald U (2003) Vitamin E biosynthesis: biochemistry meets cell biology. Trends Plant Sci 8: 6–8
Horemans N, Asard H, Van Gestelen P and Caubergs RJ (1998) Faciliated diffusion drives transport of oxidized molecules into purified plasma membrane vesicles of Phaseolus vulgaris. Physiol Plant 104: 783–789
HoremansN, Foyer CH and AsardH(2000) Transport and action of ascorbate at the plant plasma membrane. Trends Plant Sci 5: 263–267
Horling F, Lamkemeyer P, Konig J, Finkemeier I, Kandlbinder A, Baier M and Dietz KJ (2003) Divergent light-, ascorbate-, and oxidative stress-dependent regulation of expression of the peroxiredoxin gene family in Arabidopsis. Plant Physiol 131: 317–325
Hossain MA and Asada K (1984) Inactivation of ascorbate peroxidase in spinach chloroplasts on dark addition of hydrogen peroxide: its protection by ascorbate. Plant Cell Physiol 25: 1285–1295
Hou WC, Chen HJ and Lin YH (1999) Dioscorins, the major tuber storage proteins of yam (Dioscorea batatas Decne), with dehydroascorbate reductase and monodehydroascorbate reductase activities. Plant Sci 149: 151–156
Imai T, Karita S, Shiratori G, Hattori M, Nunome T, Oba K and Hirai M (1998) L-galactono-γ -lactone dehydrogenase from sweet potato: Purification and cDNA sequence analysis. Plant Cell Physiol 39: 1350–1358
Ito H, Iwabuchi M and Ogawa K (2003) The sugar-metabolic enzymes aldolase and triose-phosphate isomerase are targets of glutathionylation in Arabidopsis thaliana: Detection using biotinylated glutathione. Plant Cell Physiol 44: 655–660
Jiménez A, Hernández JA, del Río L and Sevilla F (1997) Evidence for the presence of the ascorbate-glutathione cycle in mitochondria and peroxisomes of pea leaves. Plant Physiol 114: 275–284
Joo BH, Bae YS and Lee JS (2001) Role of auxin-induced reactive oxygen species in root gravitropism. 126: 1055–1060
Jung BG, Lee KO, Lee SS, Chi YH, Jang HH, Kang SS, Lee K, Lim D, Yoon SC, Yun DJ, Inoue Y, Cho MJ and Lee SY (2002) A Chinese cabbage cDNA with high sequence identity to phospholipid hydroperoxide glutathione peroxidases encodes a novel isoform of thioredoxin-dependent peroxidases. J Biol Chem 277: 12572–12578
Kaiser S, DiMascio P, Murphy ME and Sies H (1990), Physical and chemical scavenging of singlet molecular oxygen by tocopherols. Arch Biochem Biophys 277: 101–108
Kamiya N and Shen J-R (2003) Crystal structure of oxygenevolving photosystem II from Thermosynechoccus vulcanus at 3.7 A resolution. Proc Natl Acad Sci USA 100: 98–103
Kanwischer M, Porfirova S, Bergmüller E andDörmann P (2005) Alterations in tocopherol cyclase activity in transgenic and mutant plants of Arapidosis affect tocopherol content, tocopherol composition, and oxidative stress. Plant Physiol 137: 713–723.
Karpinski S, Escobar C, Karprinska B, Creissen G and Mullineaux PM (1997) Photosynthetic electron transport regulates the expression of cytosolic ascorbate peroxidase genes in Arabidopsis during excess light stress. Plant Cell 9: 627–640
Karpinski S, Reynolds H, Karpinksa B, Wingsle G, Creissen G and Mullineaux PM (1999) Systemic signaling and acclimation in response to excess excitation energy in Arabidopsis. Science 284: 654–657
Kelly GJ and Latzko E (1979) Soluble ascorbate peroxidase. Naturwissenschaften 66: 377–382
Keren N, Gong. H and Ohad I (1995) Oscillations of reaction center II-D1 protein degradation in vivo induced by repetitive light flashes. J Biol Chem 270: 806–814
Keren N, Berg A, van Kann PJM, Levanon H and Ohad I (1997) Mechanism of photosystem II inactivation and D1 protein degradation at low light: the role of back electron flow. Proc Natl Acad Sci USA 94: 1579–1584
Kiddle G, Pastori GM, Bernard S, Pignocchi C, Antoniw J, Verrier PJ and Foyer CH (2003) Effects of ascorbate signaling on defense and photosynthesis genes. Antioxidant Redox Signaling 5: 23–32
Kimura M., Yoshizumi T, Manabe K, Yamamoto YY and Matsui M (2001) Arabidopsis transcriptional regulation by light stress via hydrogen peroxide-dependent and -independent pathways. Genes Cells 6: 607–617
Kimura M, Yamamoto YY, Seki M, Sato M, Abe T, Yoshida S, Manabe K, Shinozaki K and Matsui M (2003) Identification of Arabidopsis genes regulated by light stress using cDNA microarray. Photochem. Photobiol 77: 226–233
Kinkema M, Fan WH and Dong XN (2000) Nuclear localization of NPR1 is required for activation of PR gene expression. Plant Cell 12: 2339–2350
Klatt P and Lamas S (2000) Regulation of protein function by S-glutathiolation in response to oxidative and nitrosative stress. Eur J Biochem 267: 4928–4944
Köhler B and Blatt MR (2002) Protein phosphorylation activates the guard cell Ca2+ channel and is a prerequisite for gating by abscisic acid. Plant J 32: 185–194
Köhler B, Hills A and Blatt MR (2003) Control of guard cell ion channels by hydrogen peroxide and abscisic acid indicates their action through alternate signaling pathways. Plant Physiol 131: 385–388
Kovalchuk I, Bojko V, Kovalchuk O, Gloeckler V, Filkowski J, Heinlein M and Hohn B (2003) Pathogen induced systemic plant signal triggers genome instability. Nature 423: 760– 762
Krieger A, Rutherford AW, Vass I and Hideg E (1998) Relationship between activity, D1 loss, and Mn binding in photoinhibition of photosystem II. Biochemistry 37: 16262–16269
Kruk J and Strzalka K (1995) Occurrence and function of alphatocopherol quinone in plants. J Plant Physiol 145: 405–409
Kuge S, Jones N and Nomoto A (1997) Regulation of YAP-1 nuclear localization in response to oxidative stress. EMBO J 16: 1710–1720
Kwak JM, Mori IC, Pei ZM, Leonhardt N, Torres MA, Dangl JL, Bloom RE, Bodde S, Jones JD and Schroeder JI (2003) NADPH oxidase AtrbohD and AtrbohF genes function in ROS-dependent ABA signaling in Arabidopsis. EMBO J 22: 2623–2633
Lamb C and Dixon RA (1997) The oxidative burst in plant disease resistance. Annu Rev Plant Physiol Plant Mol Biol 48: 251–275
Lee DL, PrisbyllaMP, McLean ProvanW, Frase T and Mutter LC (1997) The discovery and structural requirements of inhibitors of p-hydroxyphenylpyruvate dioxygenase. Weed Sci 45: 601– 609
LeeKP, Kim C, LeeDWand Apel K (2003) TIGRINAd, required for regulating the biosynthesis of tetrapyrroles in barley, is an ortholog of the FLU gene of Arabidopsis thaliana. FEBS Lett 553: 119–124
Lemaire SD (2004) The glutaredoxin family in oxygenic photosynthetic organisms. Photosynth Res 79: 305–318
Lichtenthaler HK (1968) Plastoglobuli and the fine structure of plastids. Endeavour XXVII: 144–149
Lichtenthaler HK (1998) The plants 1-deoxy-d-xylulose-5- phosphate pathway for biosynthesis of isoprenoids. Fett Lipid 100: 128–138
Liere K and Link G (1997) Chloroplast endoribonuclease p54 involved in RNA 3’-end processing is regulated by phosphorylation and redox state. Nucleic Acids Res 25: 2403–2408
Liu YB, Fiskum G and Schubert D (2002) Generation of reactive oxygen species by the mitochondrial electron transport chain. J Neurochem 80: 780–787
Logan BA, Barker DH, Demmig-Adams B and Adams WW III (1996) Acclimation of leaf carotenoid composition and ascorbate levels to gradients in the light environment within an Australian rainforest. Plant Cell Environ 19: 1083–1090
Macpherson AN, Telfer A, Barber J and Truscott TG (1993) Direct detection of singlet oxygen from isolated photosystem II reaction centers. Biochim Biophys Acta 1143: 301–309
Maddison J, Lyons TM, Plöchl M and Barnes JD (2002) Hydroponically-cultivated radish fed L-galactono-1, 4-lactone exhibit increased tolerance to ozone. Planta 214: 383–391
Mathis P, Butler WL and Satoh K (1979) Carotenoid triplet state and chlorophyll fluorescence quenching in chloroplasts and subchloroplast particles. Photochem Photobiol 30: 603–614
Matsumoto H, Mizutani M, Yamaguchi T and Kadotani J (2002) Herbicide pyrazolate causes cessation of carotenoids synthesis in early watergrass by inhibiting 4-hydroxyphenylpyruvate dioxygenase. Weed Biol Manag 2: 39–45
Mattoo AK, Marder JB and EdelmanM(1989) Dynamics of the photosystem II reaction center. Cell 56: 241–246
MayMJ,VernouxT, Leaver C, Van MontaguMand Inzé D (1998) Glutathione homeostasis in plants: implications for environmental sensing and plant development. J Exp Bot 49: 649–667
Mayer M, Beyer P and Kleinig H (1990) Quinone compounds are able to replace molecular oxygen as terminal electron acceptor in phytoene desaturation in chromoplasts of Narcissus pseudonarcissus L. Eur J Biochem 191: 359–363
Melis A (1999) Photosystem II damage and repair cycle in chloroplasts:what modulates the rate of photodamage in vivo? Trends Plant Sci 4: 130–135
Meskauskine R, Nater M, Goslings D, Kessler F, op den Camp R and Apel K (2001) FLU: A negative regulator of chlorophyll biosynthesis in Arabidopsis thaliana. Proc Natl Acad Sci USA 98: 12826–12831
Millar AH, Mittova V, Kiddle G, Heazlewood JL, Bartoli CG, TheodoulouFLand Foyer CH (2003) Control of ascorbate synthesis by respiration and its implications for stress responses. Plant Physiol 133: 443–447
Mittler R (2002) Oxidative stress, antioxidants and stress tolerance. Trends Plant Sci 7: 405–410
Mittler R and Zilinskas BA (1992) Molecular cloning and characterization of a gene encoding pea cytosolic ascorbate peroxidase. J Biol Chem 267: 21802–21807
Mittova V, Kiddle G, Theodoulou FL, Gomez L, Volokita M, Tal M, Foyer CH and Guy M (2003) Co-ordinate induction of glutathione biosynthesis and glutathione-metabolising enzymes is correlated with salt tolerance in tomato. FEBS Lett 554: 417–421
Moeder W, Barry CS, Tauriainen AA, Betz C, Tuomainen J, Utriainen M, Grierson D, Sandermann H, Langebartels C and Kangasjärvi J (2002) Ethylene sunthesis regulated by biphasic induction of 1-aminocyclopropane-1-carboxylic acid synthase and 1-aminocyclopropane-1-carboxylic acid oxidase is required for hydrogen peroxide accumulation and cell death in ozone-exposed tomato. Plant Physiol 130: 1918–1926
Møller IM (2001) Plant mitochondria and oxidative stress: electron transport, NADPH turnover, and metabolism of reactive oxygen species. Annu Rev Plant Physiol Plant Mol Biol 52: 561–591
Morimura Y, Iwamoto K, Ohya T, Igarashi T, Nakamura Y, Kubo A, Tanaka K and Ikawa T (1999) Light-enhanced induction of ascorbate peroxidase in Japanese radish roots during postgerminative growth. Plant Sci 142: 123–132
Mou Z, Fan WH and Dong XN (2003) Inducers of plant systemic acquired resistance regulate NPR1 function through redox changes. Cell 113: 935–944
Mullineaux PM, Karpinski S, Jimenez A, Cleary SP, Robinson C and Creissen GP (1998) Identification of cDNAS encoding plastid-targeted glutathione peroxidases. Plant J 13: 375– 379
Mullineaux PM, Karpinski S and Creissen GP (2005) Integration of signaling in antioxidant defenses. In: Demmig-Adams B, Adams WW III and Mattoo AK (eds) Photoprotection, Photoinhibition, Gene Regulation, and Environment, pp 223–239. Springer, Dordrecht
Munné-Bosch S and Alegre L (2002a) Plant aging increases oxidative stress in chloroplasts. Planta 214: 608–615
Munné-Bosch S and Alegre L (2002b) The function of tocopherols and tocotrienols in plants. Crit Rev Plant Sci 21: 31– 57
Munné-Bosch S and Falk J (2003) New insights into the function of tocopherols in plants. Planta 218: 323–326
MurataY, Pei ZM, Mori IC and Schroeder JI (2001) Abscisic acid activation of plasma membrane Ca2+ channels in guard cells requires cytosolicNAD(P)H and is differentially disrupted upstream and downstream of reactive oxygen species production in abi1-1 and abi2-1 protein phosphatase 2C mutants. Plant Cell 13: 2513–2523
Neely WC, Martin JM and Barker SA (1988) Products and relative reaction rates of the oxidation of tocopherols with singlet molecular oxygen. Photochem Photobiol 48: 423–428
Nishizawa AN and Buchanan BB (1981) Enzyme regulation in C4 photosynthesis. Purification and properties of thioredoxinlinked fructose bisphosphatase and sedoheptulose bisphosphatase from corn leaves. J Biol Chem 256: 6119–6126
Noctor G and Foyer CH (1998) Ascorbate and glutathione: keeping active oxygen under control. Annu Rev Plant Physiol Plant Mol Biol 49: 249–279
Noctor G, Arisi ACM, Jouanin L, Kunert KJ, Rennenberg H and Foyer CH (1998) Glutathione: biosynthesis, metabolism and relationship to stress resistance explored in transformed plants. J Exp Bot 49: 623–647
Noctor G, Veljovic-Jovanovic S and Foyer CH (2000) Peroxide processing in photosynthesis: antixoxidant coupling and redox signalling. Proc Roy Soc Lond B 355: 1465–1475
Noctor G, Gomez L, Vanacker H and Foyer CH (2002a) Interactions between biosynthesis, compartmentation and transport in the control of glutathione homeostasis and signalling. J Exp Bot 53: 1283–1304
Noctor G, Veljovic-Jovanovic SD, Driscoll S, Novitskaya L and Foyer CH (2002b) Drought and oxidative load in the leaves of C3 plants: a predominant role for photorespiration? Ann Bot 89: 841–850
Noctor G, Dutilleul C, De Paepe R and Foyer CH (2004) The use of mitochondrial mutants to evaluate the effects of redox state on photosynthesis, stress tolerance, and the integration of carbon and nitrogen metabolism. J Exp Bot 55: 49–57
Norris SR, Barette TR and DellaPenna D (1995) Genetic dissection of carotenoid synthesis in Arabidopsis defines plastoquinone as an essential component of phytoene desaturation. Plant Cell 7: 2139–2148
Oba K, Ishikawa S, Nishikawa M, Mizuno H and Yamamoto T (1995) Purification and properties of L-galactono-γ -lactone dehydrogenase, a key enzyme for ascorbic acid biosynthesis, from sweet potato roots. J Biochem 117: 120–124
Ogawa K, Hatano-Iwasaki A, Yanagida M and Iwabuchi M (2004) Level of glutathione is regulated by ATP-dependent ligation of glutamate and cysteine through photosynthesis in Arabidopsis thaliana: Mechanism of strong interaction of light intensity with flowering. Plant Cell Physiol 45: 1–8
Op den Camp RG, Przybyla D, Ochsenbein C, Laloi C, Kim C, Danon A, Wagner D, Hideg E, Gobel C, Feussner I, Nater M and Apel K(2003) Rapid induction of distinct stress responses after the release of singlet oxygen in Arabidopsis. Plant Cell 15: 2320–2332
Orozco-Cardenas ML, Narvaez-Vazquez J and Ryan CA (2001) Hydrogen peroxide acts as a second messenger for the induction of defense genes in tomato plants in response towounding, systemin, and methyl jasmonate. Plant Cell 13: 179–191
Ostergaard J, Persiau G, DaveyMW, BauwGandVanMontaguM (1997) Isolation of a cDNA coding for L-galactono-γ -lactone dehydrogenase, an enzyme involved in the biosynthesis of ascorbic acid in plants. Purification, characterization, cDNA cloning, and expression in yeast. J Biol Chem 272: 30009– 30016
Osmond CB (1994) What is photoinhibition? Some insights from comparisons of shade and sun plants. In: Baker NR and Bowyer JR (eds) Photoinhibition of Photosynthesis from Molecular Mechanisms to the Field, pp 1–24. Bios Scientific Publishers, Oxford, UK
Osmond CB and Förster B (2005) Photoinhibition: then and now. In: Demmig-Adams B, Adams WW III and Mattoo AK (eds) Photoprotection, Photoinhibition, Gene Regulation, and Environment, pp 11–22. Springer, Dordrecht
Pallanca JE and Smirnoff N (1999) Ascorbic acid metabolism in pea seedlings. A comparison of D-glucosone, L-sorbosone, and L-galactono-1, 4-lactone as ascorbate precursors. Plant Physiol 120: 453–461
Pallett K (2000) The mode of action of isoxaflutole: a case study of an emerging target site. In: Cobb AH and Kirkwood RC (eds) Herbicides and Their Mechanism of Action, pp 215– 238. CRC Press, Boca Raton, FL
Pallett KE, Little JP, Sheekey M and Veeasekaran P (1998) The mode of action of isoxaflutole. Pestic Biochem Physiol 62: 113–124
Pasternak TP, Prinsen E, Ayaydin F, Miskolczi P, Potters G, Asard H, Van Onckelen HA, Dudits D and Fehér A (2002) The role of auxin, pH, and stress in the activation of embryogenic cell division in leaf protoplast-derived cells of alfalfa. Plant Physiol 129: 1807–1819
Pastori GM and Foyer CH (2002) Common components, pathways and networks of cross tolerance to stress: the central role of “redox” and hormone-mediated controls. Plant Physiol 129: 460–468
Pastori GM, Kiddle G, Antoniw J, Bernard S,Veljovic-Jovanovic S, Verrier PJ, Noctor G and Foyer CH (2003) Leaf vitamin C contents modulate plant defense transcripts and regulate genes controlling development through hormone signaling. Plant Cell 15: 939–951
Pei ZM, MurataY, Benning G, Thomine S, Klüsener B, Allen GJ, Grill E and Schroeder JI (2000) Calcium channels activated by hydrogen peroxide mediate abscisic acid signalling in guard cells. Nature 406: 731–734
Pignocchi C (2000) Establishing the functional significance of ascorbate oxidase in Planta. PhD Thesis. University of Newcastle, U.K.
Pignocchi C and Foyer CH (2003) Apoplastic ascorbate metabolism and its role in the regulation of cell signalling. Curr Opin Plant Biol 6: 379–389
Pignocchi C, Fletcher JM, Wilkinson JE, Barnes JD and Foyer CH (2003) The function of ascorbate oxidase in tobacco. Plant Physiol 132: 1631–1641
Polle A (2001) Dissecting the superoxide-dismutase-ascorbateglutathione pathway in chloroplasts by metabolic modeling. Computer simulations as a step towards flux analysis. Plant Physiol 126: 445–462
Porfirova S, Bergmüller E, Tropf S, Lemke R and Dörmann P (2002) Isolation of an Arabidopsis mutant lacking vitamin E and identification of a cyclase essential for all tocopherol biosynthesis. Proc Natl Acad Sci USA 99: 12495–12500
Potters G, Horemans N, Caubergs RJ and Asard H (2000) Ascorbate and dehydroascorbate influence cell cycle progression in a tobacco cell suspension. Plant Physiol 124: 17–20
Potters G, De Gara L, AsardHand HoremansN(2002) Ascorbate and glutathione: guardians of the cell cycle, partners in crime? Plant Physiol Biochem 40: 537–548
Potters G, Horemans N, Bellone S, Caubergs R J, Trost P, Guisez Y and Asard H (2004) Dehydroascorbate influences the plant cell cycle through a glutathione-independent reduction mechanism. Plant Physiol 134: 1479–1487
Price AH, Taylor A, Ripley SJ, Griffiths A, Trewavas AJ and Knight M (1994) Oxidative signals in tobacco increase cytosolic calcium. Plant Cell 6: 1301–1310
Puntarulo S, Sanchez RA and Boveris A (1988) Hydrogen peroxide metabolism in soybean embryonic axes at the onset of germination. Plant Physiol 86: 626–630
Puntarulo S, Galleano M, Sanchez RA and Boveris A (1991) Superoxide anion and hydrogen peroxide metabolism in soybean embryonic axes during germination. Biochim Biophys Acta 1074: 277–283
Raha S and Robinson BH (2000) Mitochondria, oxygen free radicals, disease and ageing. Trends Biochem Sci 25: 502– 508
Rennenberg H (1997) Molecular approaches to glutathione biosynthesis. In: Cram WJ, DeKok LJ, Stulem I, Brunnold C and Rennenberg H (eds) Sulphur Metabolism in Higher Plants, pp 59–70. Backhuys Publishers, Leiden, The Netherlands
Rentel MC and Knight MR (2004) Oxidative stress-induced calcium signaling in Arabidopsis. Plant Physiol 135: 1471–1479
Rodermel S (2001) Pathways of plastid-to-nucleus signalling. Trends Plant Sci 6: 471–478
Rodriguez AA, Grunberg KA and Taleisnik EL (2002) Reactive oxygen species in the elongation zone of maize leaves are necessary for leaf extension. Plant Physiol 129: 1627–1632
Rouhier N, Gelhaye E and Jacquot JP (2002) Glutaredoxindependent peroxiredoxin from poplar. Protein-protein interaction and catalytic mechanism. J Biol Chem 277: 13609–13614
Rutherford AW and Krieger-Liszkay A (2001) Herbicideinduced oxidative stress in photosystem II. Trends Biochem Sci 26: 648–653
Sandermann H (2001) Active oxygen species as mediators of plant immunity: three case studies. Biol. Chem 381: 649– 653
Sanmartin M, Drogouti PD, Lyons T, Barnes J and Kanellis AK (2003) Overexpression of ascorbate oxidase in the apoplast of transgenic tobacco results in altered ascorbate and glutathione redox states and increased sensitivity to ozone. Planta 216: 918–928
Satoh K and Mathis P (1981) Photosystem II chlorophyll α- protein complex: a study by flash absorption spectroscopy. Photobiochem Photobiophys 2: 189–198
Sattler SE, Cahoon EB, Coughlan SJ and DellaPenna D (2003) Characterisation of tocopherol cyclases from higher plants and cyanobacteria. Evolutionary implications for tocopherol synthesis and function. Plant Physiol 132: 2184–2195
Schafer FQ and Buettner GR (2001) Redox environment of the cell as viewed through the redox state of the glutathione disulfide/ glutathione couple. Free Rad Biol Medic 30: 1191–1212
Schledz M, Seidler A, Beyer P and Neuhaus G (2001) A novel phytyltransferase from Synechocystis sp. PCC 6803 involved in tocopherol biosynthesis. FEBS Lett 499: 15–20
Schopfer P (2001) Hydroxyl radical-induced cell-wall loosening in vitro and in vivo: Implications for the control of elongation growth. Plant J 28: 679–688
Schopfer P, Liszkay A, Bechtold M, Frahry G andWagner A (2002) Evidence that hydroxyl radicals mediate auxin-induced extension growth. Planta 214: 821–828
Schroeder JI, Allen GJ, Hugouvieux V, Kwak JM and Waner D (2001a) Guard cell signal transduction. Annu Rev Plant Physiol Plant Mol Biol 52: 627–658
Schroeder JI,Kwak JM and Allen GJ (2001b) Guard cell abscisic acid signalling and engineering drought hardiness in plants. Nature 410: 327–330
Schulz A, Ot O, Beyer P and Kleinig H (1993) SC-0051, a benzoyl-cyclohexane-1, 3-dione bleaching herbicide, is a potent inhibitor of the enzyme p-hydroxyphenylpyruvate dioxygenase. FEBS Lett 318: 162–166
Schupp R and Rennenberg H (1990) Diurnal changes in the thiol composition of spruce needles. In: Rennenberg H, Brunold CH, de Kok LJ and Stulen I (eds) Sulfur Nutrition and Sulfur Assimilation in Higher Plants, pp 249–254. SPB Acad Publ, The Hague, Netherlands
Schürmann P and Jacquot J-P (2000) Plant thioredoxin systems revisited. Annu Rev Plant Physiol Plant Mol Biol 51: 371–400
Scott JW, Cort WM, Harley H, Parrish DR and Saucy G (1974) 6-Hydroxychroman-2-carboxylic acid: Novel antioxidants. J Am Oil Chem Soc 51: 200–203
Sen Gupta A, Alscher RG and McCune D (1991) Response of photosynthesis and cellular antioxidants to ozone in Populus leaves. Plant Physiol 96: 650–655
Shimaoka T, Yokota A and Miyake C (2000) Purification and characterization of chloroplast dehydroascorbate reductase from spinach leaves. Plant Cell Physiol 41: 1110–1118
Siendones E, Gonzalez-Reyes JA, Santos-Ocana C, Navas P and Cordoba F (1999) Biosynthesis of ascorbic acid in kidney bean. L-galactono-γ -lactone dehydrogenase is an intrinsic protein located at the mitochondrial inner membrane. Plant Physiol 120: 907–912
Smirnoff N (2000) Ascorbic acid: metabolism and functions of a multi-facetted molecule. Curr Opin Plant Biol 3: 229–235
Smirnoff N and Pallanca JE (1996) Ascorbate metabolism in relation to oxidative stress. Biochem Soc Trans 24: 472–478
Smirnoff N and Wheeler GL (2000) Ascorbic acid in plants. Biosynthesis and function. Crit Rev Biochem Mol Biol 35: 291–314
Smirnoff N, Running JA and Gatzek S (2004) Ascorbate metabolism in relation to oxidative stress. In: Asrad H, MayJ M and Smirnoff N (eds) Vitamin C Functions and Biochemistry in Animals and Plants, pp 1–30. Bios Scientific Publishers. London, UK
Smith IK, Kendall AC, Keys AJ, Turner JC and Lea PJ (1984) Increased levels of glutathione in a catalase-deficient mutant of barley (Hordeum vulgare L.). Plant Sci Lett 37: 29–33
Soll J, Schultz G, Joyard J, Douce R and BlockMA(1985) Localization and synthesis of prenylquinones in isolated outer and inner envelope membranes from spinach chloroplasts. Arch Biochem Biophys 238: 290–299
StarkeDW, ChockPBand Mieyal JJ (2003) Glutathione thiyl radical scavenging and transferase properties of human glutaredoxin (thioltransferase). Potential role in redox signal transduction. J Biol Chem 278: 14607–14613
Sticher L, MauchMani B and Metraux JP (1997) Systemic acquired resistance. Annu Rev Phytopath 35: 235–270
Streb P, Aubert S, Gout E and Bligny R (2003) Cold- and lightinduced changes of metabolite and antioxidant levels in two high mountain plant species Soldanella alpina and Ranunculus glacialis and a lowland species Pisum sativum. Physiol Plant 118: 96–104
Strand A, Asami T, Alonso J, Ecker JR and Chory J (2003) Chloroplast to nucleus communication triggered by accumulation of Mg-protoporphyrin IX. Nature 421: 79–83
Sweetlove L and Foyer CH (2004). Roles for reactive oxygen species and antioxidants in plant mitochondria. In: Day DA, Millar AH and Whelan J (eds) Plant Mitochondria: From Genome to Function. Advances in Photosynthesis and Respiration, Vol 17, pp 307–320. Kluwer Academic Publishers, Dordrecht, The Netherlands
Tabata K, Oba K, Suzuki K and Esaka M (2001) Generation and properties of ascorbic acid-deficient transgenic tobacco cells expressing antisense RNA for L-galactono-1, 4-lactone dehydrogenase. Plant J 27: 139–148
Tabata K, Takaoka T and Esaka M (2002) Gene expression of ascorbic acid-related enzymes in tobacco. Phytochemistry 61: 631–635
Tamaoki M, Mukai F, Asai N, Nakajima N, Kubo A, AonoMand Saji H (2003) Light-controlled expression of a gene encoding L-galactono-γ -lactone dehydrogenase which affects ascorbate pool size in Arabidopsis thaliana. Plant Sci 164: 1111– 1117
Telfer A (2002) What is β-carotene doing in the photosystem II reaction centre? Phil Trans Roy Soc Lond 357: 1431–1440
Telfer A and Barber J (1995) Role of carotenoid bound to the photosystem II reaction centre. In: Mathis P (ed) Photosynthesis: from Light to Biosphere, Vol IV, pp 15–20. Kluwer Academic Publishers, Dordrecht
Telfer A, Bishop SM, Philipps D and Barber J (1994a) Isolated photosynthetic reaction center of photosystem II as a sensitizer for the formation of singlet oxygen. Detection and quantum yield determination using a chemical trapping technique. J Biol Chem 269: 13244–13253
Telfer A, Dhami S, Bishop SM, Phillips D and Barber J (1994b) β-carotene quenches singlet oxygen formed in isolated photosystem II reaction center. Biochemistry 33: 14469– 14474
TeviniMand Lichtenthaler HK (1970) Untersuchungen über die Pigment- und Lipochinonausstattung der zwei photosynthetischen Pigmentsysteme. Z Pflanzenphysiol 62: 17–32
Trebst A (1999) Singlet oxygen in photosynthesis. In: Denke A and Dornisch K (eds) Different Pathways through Life, pp 125–142. Lincom Europe, München, Germany
Trebst A (2003) Function of β-carotene and tocopherol in photosystem II. Z Naturforsch 58c: 609–620
Trebst A and Depka B (1997) Role of carotene in the rapid turnover and assembly of photosystem II in Chlamydomonas reinhardtii. FEBS Lett 400: 359–362
Trebst A, Depka B and Holländer-Czytko H (2002) A specific role for tocopherol and of chemical singlet oxygen quenchers in the maintenance of photosystem II structure and function in Chlamydomonas reinhardtii. FEBS Lett 516: 156–160
Trebst A, Depka B, Jäger J and Oettmeier W (2004) Reversal of the inhibition of photosynthesis by herbicides affecting hydroxyphenylpyruvate dioxygenase by plastoquinone- and tocopheryl-derivatives in Chlamydomonas reinhardtii. Pest Manag Sci publication 60: 669–674.
Trumper S, Follmann H and Haberlein I (1994) A novel dehydroascorbate reductase from spinach chloroplasts homologous to plant trypsin inhibitor. FEBS Lett 352: 159–162
Trumpower BL (1990) The protomotive Q cycle. Energy transduction by coupling of proton translocation to electron transfer by the cytochrome bc1 complex. J Biol Chem 265: 11409– 11412
Tsegaye Y, Shintani DK and DellaPenna D (2002) Overexpression of the enzyme p-hydroxyphenolpyruvate dioxygenase in Arabidopsis and its relation to tocopherol biosynthesis. Plant Physiol Biochem 40: 913–920.
Tsukaguchi H, Tokui T, Mackenzie B, Berger UV, Chen XZ, Wang Y, Brubaker RF and Hediger MA (1999) A family of mammalian Na+-dependent L-ascorbic acid transporters. Nature 399: 70–75
Urano J, Nakagawa T, Maki Y, Masumura T, Tanaka K, Murata N and Ushimaru T (2000) Molecular cloning and characterization of a rice dehydroascorbate reductase. FEBS Lett 466: 107–111
Vanacker H, Carver TL and Foyer CH (1998) Pathogen-induced changes in the antioxidant status of the apoplast in barley leaves. Plant Physiol 117: 1103–1114
Vanlerberghe G C and Ordog SH (2002) Alternative oxidase: integrating carbon metabolism and electron transport in plant respiration. In: Foyer CH and Noctor G (eds) Photosynthetic Nitrogen Assimilation and Associated Carbon and Respiratory Metabolism, Advances in Photosynthesis and Respiration, Vol 12, pp 173–191. Kluwer Academic Publishers, Dordrecht
Van Gorkom HJ and Schelvi JPM (1993) Kok’s oxygen clock: what makes it tick? The structure of P680 and consequences of its oxidizing power. Photosynth Res 38: 297–301
Veljovic-Jovanovic SD, Pignocchi C, Noctor G and Foyer CH (2001) Low ascorbic acid in the vtc-1 mutant of Arabidopsis is associated with decreased growth and intracellular redistribution of the antioxidant system. Plant Physiol 127: 426–435
Vernoux T, Wilson RC, Seeley KA, Reichheld JP, Muroy S, Brown S, Maughan SC, Cobbett CS, Van Montagu M, Inzé D, May MJ and Sung ZR (2000) The ROOTMERISTEMLESS1/ CADMIUM SENSITIVE2 gene defines a glutathionedependent pathway involved in initiation and maintenance of cell division during post-embryonic root development. Plant Cell 12: 97–100
Vranova E, Inze D and Van Breusegem F (2002a) Signal transduction during oxidative stress. J Exp Bot 53: 1227–1236
Vranova E, Atichartpongkul S, Villarroel R., Van Montagu M., Inzé D and Van Camp W (2002b) Comprehensive analysis of gene expression in Nicotiana tabacum leaves acclimated to oxidative stress. Proc Natl Acad Sci USA 99: 10870–10875
Wagner D, Przybyla D, op den Camp R, Kim C, Landgraf F, Lee K P,Wursch M, Laloi C, Nater M, Higeg E and Apel K (2004) The genetic basis of singlet oxygen-induced stress reactions of Arabidopsis thaliana. Science 306: 1183–1185.
Wells WW, Xu DP, Yang Y and Rocque PA (1990) Mammalian thioltransferase (glutaredoxin) and protein disulfide isomerase have dehydroascorbate reductase activity. J Biol Chem 265: 15361–15364
Weber A, Servaites JC, Geiger DR, Kofler H, Hille D, Groner F, Hebbeker U and Flugge UI (2000) Identification, purification, and molecular cloning of a putative plastidic glucose translocator. Plant Cell 12: 787–802
Wheeler GL, Jones MA and Smirnoff N (1998) The biosynthetic pathway of vitamin C in higher plants. Nature 393: 365– 369
Willekens H, Chamnongpol S, Davey M, Schraudner M, Langebartels C, Van Montagu M, Inzé D and Van Camp W (1997) Catalase is a sink for H2O2 and is indispensable for stress defense in C3 plants. EMBO J 16: 4806–4816
Wingate VPM, Lawton MA and Lamb CJ (1988) Glutathione causes a massive and selective induction of plant defense genes. Plant Physiol 87: 206–210
Winkler BS (1992) Unequivocal evidence in support of the nonenzymatic redox coupling between glutathione/glutathione disulfide and ascorbic acid/dehydroascorbic acid. Biochim Biophys Acta 1117: 287–290
Witt HT (1996) Photosynthesis. Ber Bunsenge Phys Chem 100: 1923–1927
Xiang C and Oliver DJ (1998) Glutathione metabolic genes coordinately respond to heavy metals and jasmonic acid in Arabidopsis. Plant Cell 10: 1539–1550
Yabuta Y, Yoshimura K, Takeda T and Shigeoka S (2000) Molecular characterization of tobacco mitochondrial L-galactono-γ -lactone dehydrogenase and its expression in Escherichia coli. Plant Cell Physiol. 41: 666–675
Yamaguchi K, Mori H and Nishimura M(1995) A novel isozyme of ascorbate peroxidase localized on glyoxysomal and leaf peroxisomal membranes in pumpkin. Plant Cell Physiol 36: 1157–1162
Yoshimura K, Ishikawa T, NakamuraY, Tamoi M, Takeda T, Tada T, Nishimura K and Shigeoka S (1998) Comparative study on recombinant chloroplastic and cytosolic ascorbate peroxidase isozymes of spinach. Arch Biochem Biophys 353: 55–63
Zhang W, Zhang L, Dong F, Gao J, Galbraith DW and Song CP (2001) Hydrogen peroxide is involved in abscisic acid-induced stomatal closure in Vicia faba. Plant Physiol 126: 1438–1448
Author information
Authors and Affiliations
Editor information
Editors and Affiliations
Rights and permissions
Copyright information
© 2008 Springer Science+Business Media B.V.
About this chapter
Cite this chapter
Foyer, C.H., Trebst, A., Noctor, G. (2008). Signaling and Integration of Defense Functions of Tocopherol, Ascorbate and Glutathione. In: Demmig-Adams, B., Adams, W.W., Mattoo, A.K. (eds) Photoprotection, Photoinhibition, Gene Regulation, and Environment. Advances in Photosynthesis and Respiration, vol 21. Springer, Dordrecht. https://doi.org/10.1007/1-4020-3579-9_16
Download citation
DOI: https://doi.org/10.1007/1-4020-3579-9_16
Publisher Name: Springer, Dordrecht
Print ISBN: 978-1-4020-3564-7
Online ISBN: 978-1-4020-3579-1
eBook Packages: Biomedical and Life SciencesBiomedical and Life Sciences (R0)