Anthocyanin Function in Vegetative Organs



Possible functions of anthocyanins in leaves, stems, roots and other vegetative organs have long attracted scientific debate. Key functional hypotheses include: (i) protection of chloroplasts from the adverse effects of excess light; (ii) attenuation of UV-B radiation; and (iii) antioxidant activity. However, recent data indicate that the degree to which each of these processes is affected by anthocyanins varies greatly across plant species. Indeed, none of the hypotheses adequately explains variation in spatial and temporal patterns of anthocyanin production. We suggest instead that anthocyanins may have a more indirect role, as modulators of reactive oxygen signalling cascades involved in plant growth and development, responses to stress, and gene expression.


Reactive Oxygen Species Chlorophyll Fluorescence Green Leaf Methyl Viologen Vegetative Organ 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.


  1. Adir, N., Zer, H., Scholat, S. and Ohad, I. (2003) Photoinhibition – a historical review. Photosynth. Res. 76, 343–370.PubMedGoogle Scholar
  2. Agati, G., Matteini, P., Goti, A. and Tattini, M. (2007) Chloroplast-located flavonoids can scavenge singlet oxygen. New Phytol. 174, 77–89.PubMedGoogle Scholar
  3. Ålenius, C.M., Vogelmann, T.C. and Bornman, J.F. (1995) A three-dimensional representation of the relationship between penetration of U.V.-B radiation and U.V.-screening pigments in leaves of Brassica napus. New Phytol. 131, 297–302.Google Scholar
  4. Allan, A.C. and Fluhr, R. (1997) Two distinct sources of elicited reactive oxygen species in tobacco epidermal cells. Plant Cell 9, 1559–1572.PubMedGoogle Scholar
  5. Alscher, R.G., Donahue, J.L. and Cramer, C.L. (1997) Reactive oxygen species and antioxidants: relationships in green cells. Physiol. Plant. 100, 224–233.Google Scholar
  6. Anderson, M.D., Prasad, T.K. and Stewart, C.R. (1995) Changes in isozyme profiles of catalase, peroxidase, and glutathione reductase during acclimation to chilling in mesocotyls of maize seedlings. Plant Physiol. 109, 1247–1257.PubMedGoogle Scholar
  7. Apel, K. and Hirt, H. (2004) Reactive oxygen species: metabolism, oxidative stress, and signal transduction. Annu. Rev. Plant Biol. 55, 373–399.PubMedGoogle Scholar
  8. 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.PubMedGoogle Scholar
  9. Asada, K. (2000) The water-water cycle as alternative photon and electron sinks. Phil. Trans. R. Soc. Lond. B 355, 1419–1431.Google Scholar
  10. Bowler, C. and Fluhr, R. (2000) The role of calcium and activated oxygens as signals for controlling cross-tolerance. Trends Plant Sci. 5, 241–246.PubMedGoogle Scholar
  11. Bowler, C., van Montagu, M. and Inzé, D. (1992) Superoxide dismutase and stress tolerance. Annu. Rev. Plant Physiol. Plant Mol. Biol. 43, 83–116.Google Scholar
  12. Brown, J.E., Khodr, H., Hider, R.C. and Rice-Evans, C.A. (1998) Structural dependence of flavonoid interactions with Cu2 + ions: implications for their antioxidant properties. Biochem. J. 330, 1173–1178.PubMedGoogle Scholar
  13. Buchholz, G., Elmann, B. and Wellmann, E. (1995) Ultraviolet light inhibition of phytochrome-induced flavonoid biosynthesis and DNA photolyase formation in mustard cotyledons (Sinapis alba L.). Plant Physiol. 108, 227–234.PubMedGoogle Scholar
  14. Burger, J. and Edwards, G. (1996) Photosynthetic efficiency, and photodamage by UV and visible radiation, in red versus green leaf Coleus varieties. Plant Cell Physiol. 37, 395–399.Google Scholar
  15. Cai, Z.-Q., Slot, M. and Fan, Z.-X. (2005) Leaf development and photosynthetic properties of three tropical tree species with delayed greening. Photosynthetica 43, 91–98.Google Scholar
  16. Caldwell, M.M., Robberecht, R. and Flint, S.D. (1983) Internal filters: prospects for UV-acclimation in higher plants. Physiol. Plant. 58, 445–450.Google Scholar
  17. Casano, L.M., Gómez, L.D., Lascano, H.R., González, C.A. and Trippi, V.S. (1997) Inactivation and degradation of CuZn-SOD by active oxygen species in wheat chloroplasts exposed to photooxidative stress. Plant Cell Physiol. 38, 433–440.PubMedGoogle Scholar
  18. Chalker-Scott, L. (1999) Environmental significance of anthocyanins in plant stress responses. Photochem. Photobiol. 70, 1–9.Google Scholar
  19. Chalker-Scott, L. (2002) Do anthocyanins function as osmoregulators in leaf tissues? Adv. Bot. Res. 37, 103–127.Google Scholar
  20. Choinski, J.S. Jr., Ralph, P. and Eamus, D. (2003) Changes in photosynthesis during leaf expansion in Corymbia gummifera. Aust. J. Bot. 51, 111–118.Google Scholar
  21. Close, D.C. and Beadle, C.L. (2003) The ecophysiology of foliar anthocyanin. Bot. Rev. 69, 149–161.Google Scholar
  22. Corpas, F.J., Barroso, J.B. and del Río, L.A. (2001) Peroxisomes as a source of reactive oxygen species and nitric oxide signal molecules in plant cells. Trends Plant Sci. 6, 145–150.PubMedGoogle Scholar
  23. Couée, I., Sulmon, C., Gouesbet, G. and El Amrani, A. (2006) Involvement of soluble sugars in reactive oxygen species balance and responses to oxidative stress in plants. J. Exp. Bot. 57, 449–459.PubMedGoogle Scholar
  24. Dat, J., Vandenabeele, S., Vranová, E., Van Montagu, M., Inzé, D. and Van Breusegem, F. (2000) Dual action of the active oxygen species during plant stress responses. Cell. Mol. Life Sci. 57, 779–795.PubMedGoogle Scholar
  25. Day, T.A., Vogelmann, T.C. and DeLucia, E.H. (1992) Are some plant life forms more effective than others in screening out ultraviolet-B radiation? Oecologia 92, 513–519.Google Scholar
  26. Demmig-Adams, B. and Adams III, W.W. (2006) Photoprotection in an ecological context: the remarkable complexity of thermal energy dissipation. New Phytol. 172, 11–21.PubMedGoogle Scholar
  27. Dröge, W. (2002) Free radicals in the physiological control of cell function. Physiol. Rev. 82, 47–95.PubMedGoogle Scholar
  28. Feild, T.S., Lee, D.W. and Holbrook, N.M. (2001) Why leaves turn red in Autumn. The role of anthocyanins in senescing leaves of red-osier dogwood. Plant Physiol. 127, 566–574.PubMedGoogle Scholar
  29. Foyer, C.H. and Noctor, G. (2005) Oxidant and antioxidant signalling in plants: a re-evaluation of the concept of oxidative stress in a physiological context. Plant Cell Environ. 28, 1056–1071.Google Scholar
  30. Foyer, C.H., Lelandais, M. and Kunert, K.J. (1994) Photooxidative stress in plants. Physiol. Plant. 92, 696–717.Google Scholar
  31. Fry, S.C. (1998) Oxidative scission in plant cell wall polysaccharides by ascorbate-induced hydroxyl radicals. Biochem. J. 332, 507–515.PubMedGoogle Scholar
  32. Genty, B., Briantais, J.-M. and Baker, N.R. (1989) The relationship between quantum yield of photosynthetic electron transport and quenching of chlorophyll fluorescence. Biochim. Biophys. Acta 990, 87–92.Google Scholar
  33. Gibson, S. (2005) Control of plant development and gene expression by sugar signaling. Curr. Opin. Plant Biol. 8, 93–102.PubMedGoogle Scholar
  34. Gitelson, A.A., Merzylak, M.N. and Chivkunova, O.B. (2001) Optical properties and non-destructive estimation of anthocyanin content in plant leaves. Photochem. Photobiol. 74, 38–45.PubMedGoogle Scholar
  35. Giusti, M., Rodriguez-Saona, L. and Wrolstad, R. (1999) Molar absorptivity and color characteristics of acylated and non-acylated pelargonidin-based anthocyanins. J. Agric. Food Chem. 47, 4631–4637.PubMedGoogle Scholar
  36. Gorton, H. and Vogelmann, T.C. (1996) Effects of epidermal cell shape and pigmentation on optical properties of Antirrhinum petals at visible and ultraviolet wavelengths. Plant Physiol. 112, 879–888.PubMedGoogle Scholar
  37. Gould, K.S. (2003) Free radicals, oxidative stress and antioxidants. In: Thomas, B., Murphy, D.J. and Murray, B.G. (Eds), Encyclopedia of Applied Plant Sciences. Elsevier, Amsterdam, pp. 9–16.Google Scholar
  38. Gould, K.S. (2004) Nature’s Swiss army knife: the diverse protective roles of anthocyanins in leaves. J. Biomed. Biotechnol. 2004, 314–320.PubMedGoogle Scholar
  39. Gould, K.S. and Lister, C. (2005) Flavonoid functions in plants. In: Andersen, Ø.M. and Markham, K.R. (Eds), Flavonoids: Chemistry, Biochemistry, and Applications. CRC Press, Boca Raton, pp. 397–441.Google Scholar
  40. Gould, K.S. and Quinn, B.D. (1999) Do anthocyanins protect leaves of New Zealand native species from UV-B? New Zeal. J. Bot. 37, 175–178.Google Scholar
  41. Gould, K.S., Kuhn, D.N., Lee, D.W. and Oberbauer, S.F. (1995) Why leaves are sometimes red. Nature 378, 241–242.Google Scholar
  42. Gould, K.S., Markham, K.R., Smith, R.G. and Goris, J.J. (2000) Functional role of anthocyanins in the leaves of Quintinia serrata A Cunn. J. Exp. Bot. 51, 1107–1115.PubMedGoogle Scholar
  43. Gould, K.S., McKelvie, J. and Markham, K.R. (2002a) Do anthocyanins function as antioxidants in leaves? Imaging of H2O2 in red and green leaves after mechanical injury. Plant Cell Environ. 25, 1261–1269.Google Scholar
  44. Gould, K.S., Neill, S.O. and Vogelmann, T.C. (2002b) A unified explanation for anthocyanins in leaves? Adv. Bot. Res. 37, 167–192.Google Scholar
  45. Gould, K.S., Vogelmann, T.C., Han, T. and Clearwater, M.J. (2002c) Profiles of photosynthesis of red and green leaves of Quintinia serrata. Physiol. Plant. 116, 127–133.Google Scholar
  46. Grant, J.J. and Loake, G.J. (2000) Role of reactive oxygen intermediates and cognate redox signalling in disease resistance. Plant Physiol. 124, 21–29.PubMedGoogle Scholar
  47. Hada, H., Hidema, J., Maekawa, M. and Kumagai, T. (2003) Higher amounts of anthocyanins and UV-absorbing compounds effectively lowered CPD photorepair in purple rice (Oryza sativa L.). Plant Cell Environ. 26, 1691–1701.Google Scholar
  48. Halliwell, B. and Gutteridge, J.M.C. (1999) Free Radicals in Biology and Medicine. Oxford University Press, Oxford.Google Scholar
  49. Hara, M., Oki, K., Hoshino, K. and Kuboi, T. (2003) Enhancement of anthocyanin biosynthesis by sugar in radish (Raphanus sativus) hypocotyl. Plant Sci. 164, 259–265.Google Scholar
  50. Havaux, M. and Kloppstech, K. (2001) The protective functions of carotenoid and flavonoid pigments against excess visible radiation at chilling temperature investigated in Arabidopsis npq and tt mutants. Planta 213, 953–966.Google Scholar
  51. Hoch, W.A., Zeldin, E.L. and McCown, B.H. (2001) Physiological significance of anthocyanins during autumnal leaf senescence. Tree Physiol. 21, 1–8.PubMedGoogle Scholar
  52. Hoch, W.A., Singaas, E.L. and McCown, B.H. (2003) Resorption protection. Anthocyanins facilitate nutrient recovery in Autumn by shielding leaves from potentially damaging light levels. Plant Physiol. 133, 1–10.Google Scholar
  53. Hooijmaijers, C.A.M. and Gould, K.S. (2007) Photoprotective pigments in red and green gametophytes of two New Zealand liverworts. New Zeal. J. Bot. 45, 451–461.Google Scholar
  54. Hoque, E. and Remus, G. (1999). Natural UV-screening mechanisms of Norway spruce (Picea abies [L.] Karst.) needles. Photochem. Photobiol. 69, 177–192.Google Scholar
  55. Hughes, N.M. and Smith, W.K. (2007) Attenuation of incident light in Galax urceolata (Diapensiaceae): concerted influence of adaxial and abaxial anthocyanic layers on photoprotection. Am. J. Bot. 94, 784–790.Google Scholar
  56. Hughes, N.M., Neufeld, H.S. and Burkey, K.O. (2005) Functional role of anthocyanins in high-light winter leaves of the evergreen herb Galax urceolata. New Phytol. 168, 575–587.PubMedGoogle Scholar
  57. Hughes, N.M., Morley, C.B. and Smith, W.K. (2007) Coordination of anthocyanin decline and photosynthetic maturation in juvenile leaves of three deciduous tree species. New Phytol. 175, 675–685.PubMedGoogle Scholar
  58. Jahnke, L.S., Hull, M.R. and Long, S.P. (1991) Chilling stress and oxygen metabolizing enzymes in Zea mays and Zea diploperennis. Plant Cell Environ. 14, 97–104.Google Scholar
  59. Jordan, B.R., James, P., Strid, Å. and Anthony, R. (1994) The effect of ultraviolet-B radiation on gene expression and pigment composition in etiolated and green pea leaf tissue: UV-B-induced changes are gene-specific and dependent upon the developmental stage. Plant Cell Environ. 17, 45–54.Google Scholar
  60. Karageorgou, P. and Manetas, Y. (2006) The importance of being red when young: anthocyanins and the protection of young leaves of Quercus coccifera from insect herbivory and excess light. Tree Physiol. 26, 613–621.PubMedGoogle Scholar
  61. Kato, M. and Shimizu, S. (1985) Chlorophyll metabolism in higher plants VI. Involvement of peroxidase in chlorophyll degradation. Plant Cell Physiol. 26, 1291–1301.Google Scholar
  62. Kawano, T. (2003) Roles of the reactive oxygen species-generating peroxidase reactions in plant defense and growth induction. Plant Cell Rep. 21, 829–837.PubMedGoogle Scholar
  63. Koostra, A. (1994) Protection from UV-B induced DNA damage by flavonoids. Plant Mol. Biol. 26, 771–774.Google Scholar
  64. Krause, G.H. and Weis, E. (1991) Chlorophyll fluorescence and photosynthesis: the basics. Annu. Rev. Plant Physiol. Plant Mol. Biol. 42, 331–349.Google Scholar
  65. Krol, M., Gray, G.R., Hurry, V.M., Oquist, G., Malek, L. and Huner, N.P.A. (1995) Low-temperature stress and photoperiod affect an increased tolerance to photoinhibition inPinus banksiana seedlings. Can. J. Bot. 73, 1119–1127.Google Scholar
  66. Kuk, Y.I., Shin, J.S., Burgos, N.R., Hwang, T.E., Han, O., Cho, B.H., Jung, S. and Guh J.O. (2003) Antioxidative enzymes offer protection from chilling damage in rice plants. Crop Sci. 43, 2109–2117.Google Scholar
  67. Kunz, S. and Becker, H. (1995) Cell wall pigment formation of in vitro cultures of the liverwort Ricciocarpos natans. Z. Naturforsch. 50, 235–240.Google Scholar
  68. Kunz, S., Burkhardt, G. and Becker, H. (1994) Riccionidins A and B, anthocyanidins from the cell walls of the liverwort Ricciocarpos natans. Phytochemistry 35: 233–235.Google Scholar
  69. Kyparissis, A., Grammatikopoulos, G. and Manetas, Y. (2007) Leaf morphological and physiological adjustments to the spectrally selective shade imposed by anthocyanins in Prunus cerasifera. Tree Physiol. 27, 849–857.PubMedGoogle Scholar
  70. Kytridis, V.-P. and Manetas, Y. (2006) Mesophyll versus epidermal anthocyanins as potentialin vivo antioxidants: evidence linking the putative antioxidant role to the proximity of oxy-radical source. J. Exp. Bot. 57, 2203–2210.PubMedGoogle Scholar
  71. Laloi, C., Apel, K. and Danon A. (2004) Reactive oxygen signalling: the latest news. Curr. Opin. Plant Biol. 7, 323–328.PubMedGoogle Scholar
  72. Lee, D.W. and Collins, T.M. (2001) Phylogenetic and ontogenetic influences on the distribution of anthocyanins and betacyanins in leaves of tropical plants. Int. J. Plant Sci. 162, 1141–1153.Google Scholar
  73. Lee, D.W. and Gould, K.S. (2002a) Anthocyanins in leaves and other vegetative organs: an introduction. Adv. Bot. Res. 37, 2–16.Google Scholar
  74. Lee, D.W. and Gould, K.S. (2002b) Why leaves turn red. Am. Sci. 90, 524–531.Google Scholar
  75. Li, J., Ou-Lee, T.M., Raba, R., Amundson, R.G. and Last, R.L. (1993) Arabidopsis flavonoid mutants are hypersensitive to UV-B irradiation. Plant Cell 5, 171–179.PubMedGoogle Scholar
  76. Liakopoulos, G., Nikolopoulos, D., Klouvatou, A., Vekkos, K.-A., Manetas, Y. and Karabourniotis, G. (2006) The photoprotective role of epidermal anthocyanins and surface pubescence in young leaves of grapevine (Vitis vinifera). Ann. Bot. 98, 257–265.PubMedGoogle Scholar
  77. Logan, B.A., Adams III, W.W. and Demmig-Adams, B. (2007) Avoiding common pitfalls of chlorophyll fluorescence analysis under field conditions. Funct. Plant Biol. 3, 853–859.Google Scholar
  78. Long, S.P., Humphries, S. and Falkowski, P.G. (1994) Photoinhibition of photosynthesis in nature. Annu. Rev. Plant Physiol. Plant Mol. Biol. 45, 633–662.Google Scholar
  79. Mahalingam, R. and Fedoroff, N. (2003) Stress response, cell death and signalling: the many faces of reactive oxygen species. Physiol. Plant. 119, 56–68.Google Scholar
  80. Manetas, Y. (2006) Why some leaves are anthocyanic, and why most anthocyanic leaves are red. Flora 201, 163–177.Google Scholar
  81. Manetas, Y., Drinia, A. and Petropoulou, Y. (2002) High contents of anthocyanins in young leaves are correlated with low pools of xanthophyll cycle components and low risk of photoinhibition. Photosynthetica 40, 349–354.Google Scholar
  82. Manetas, Y., Petropoulou, Y., Psaras, G.K. and Drinia, A. (2003). Exposed red (anthocyanic) leaves of Quercus coccifera display shade characteristics. Funct. Plant Biol. 30, 265–270.Google Scholar
  83. Markham, K.R. (1982) Techniques of Flavonoid Identification. Academic Press, London.Google Scholar
  84. Maxwell, K. and Johnson, G.N. (2000) Chlorophyll fluorescence – a practical guide. J. Exp. Bot. 51, 659–668.PubMedGoogle Scholar
  85. McClure, J.W. (1975) Physiology and functions of flavonoids. In: Harborne, J.B. (Ed.), The Flavonoids. Chapman and Hall, London, pp. 970–1055.Google Scholar
  86. Meiers, S., Kemény, M., Weyand, U., Gastpar, R., Von Angerer E. and Marko, D. (2001) The anthocyanidins cyanidin and delphinidin are potent inhibitors of the epidermal growth-factor receptor. J. Agric. Food Chem. 49, 958–962.PubMedGoogle Scholar
  87. Mendez, M., Jones, D.G. and Manetas, Y. (1999) Enhanced UV-B radiation under field conditions increases anthocyanin and reduces the risk of photoinhibition but does not affect growth in the carnivorous plant Pinguicula vulgaris. New Phytol. 144, 275–282.Google Scholar
  88. Mittler, R. (2002) Oxidative stress, antioxidants and stress tolerance. Trends Plant Sci. 7, 495–410.Google Scholar
  89. Mittler, R., Vanderauwera, S., Gollery, M. and van Breusegem, F. (2004) Reactive oxygen gene network of plants. Trends Plant Sci. 9, 490–498.PubMedGoogle Scholar
  90. Murray, J.R. and Hackett, W.P. (1991) Dihydroflavonol reductase activity in relation to differential anthocyanin accumulation in juvenile and mature phase Hedera helix L. Plant Physiol. 97, 343–351.PubMedGoogle Scholar
  91. Neill, S.O. and Gould, K.S. (1999) Optical properties of leaves in relation to anthocyanin concentration and distribution. Can. J. Bot. 77, 1777–1782.Google Scholar
  92. Neill, S.O. and Gould, K.S. (2003) Anthocyanins in leaves: light attenuators or antioxidants? Funct. Plant Biol. 30, 865–873.Google Scholar
  93. Neill, S., Desikan, R. and Hancock, J. (2002a) Hydrogen peroxide signalling. Curr. Opin, Plant Biol. 5, 388–395.Google Scholar
  94. Neill, S.O., Gould, K.S., Kilmartin, P.A., Mitchell, K.A. and Markham, K.R. (2002b) Antioxidant activities of red versus green leaves of Elatostema rugosum. Plant Cell Environ. 25, 539–547.Google Scholar
  95. Neill, S.O., Gould, K.S., Kilmartin, P.A., Mitchell, K.A. and Markham, K.R. (2002c) Antioxidant capacities of green and cyanic leaves in the sun species, Quintinia serrata. Funct. Plant Biol. 29, 1437–1443.Google Scholar
  96. Nishio, J.N. (2000) Why are higher plants green? Evolution of the higher plant photosynthetic pigment complement. Plant Cell Environ. 23, 539–548.Google Scholar
  97. Niyogi, K.K. (2000) Safety valves for photosynthesis. Curr. Opin. Plant Biol. 3, 455–460.PubMedGoogle Scholar
  98. Olsson, L.C., Veit, M. and Bornman, J.F. (1999) Epidermal transmittance and phenolic composition in leaves of atrazine-tolerant and atrazine-sensitive cultivars ofBrassica napus grown under enhanced UV-B radiation. Physiol. Plant. 107, 259–266.Google Scholar
  99. Pettigrew, W.T. and Vaughn, K.C. (1998) Physiological, structural, and immunological characterization of leaf and chloroplast development in cotton. Protoplasma 202, 23–37.Google Scholar
  100. Pfündel, E.E., Ghozlen, N.B., Meyer, S. and Cerovic, Z.G. (2007) Investigating UV screening in leaves by two different types of portable UV fluorimeters reveals in vivo screening by anthocyanins and carotenoids. Photosynth. Res. 93, 205–221.PubMedGoogle Scholar
  101. Pietrini, F., Iannelli, M.A. and Massacci, A. (2002) Anthocyanin accumulation in the illuminated surface of maize leaves enhances protection from photo-inhibitory risks at low temperature, without further limitation to photosynthesis. Plant Cell Environ. 25, 1251–1259.Google Scholar
  102. Pinhero, R.G., Rao, M.V., Paliyath, G., Murr, D.P. and Fletcher, R.A. (1997) Changes in activities of antioxidant enzymes and their relationship to genetic and paclobutrazol-induced chilling tolerance of maize seedlings. Plant Physiol. 114, 695–704.PubMedGoogle Scholar
  103. Pitzschke, A., Forzani, C. and Hirt, H. (2006) Reactive oxygen species signaling in plants. Antioxid. Redox Signal. 8, 1757–1764.PubMedGoogle Scholar
  104. Polle, A. (1997) Defense against photooxidative damage in plants. In: Scandalios, J.G. (Ed.), Oxidative Stress and the Molecular Biology of Antioxidant Defenses. New York, Cold Spring Harbor Laboratory Press, pp. 623–666.Google Scholar
  105. Polle, A. (2001) Dissecting the superoxide dismutase-ascorbate peroxidase-glutathione pathway in chloroplasts by metabolic modelling. Computer simulations as a step towards flux analysis. Plant Physiol. 126, 445–462.PubMedGoogle Scholar
  106. Post, A. (1990) Photoprotective pigment as an adaptive strategy in the Antarctic moss Ceratodon purpureus. Polar Biol. 10, 241–245.Google Scholar
  107. Post, A. and Vesk, M. (1992) Photosynthesis, pigments, and chloroplast ultrastucture of an antarctic liverwort from sun-exposed and shaded sites. Can. J. Bot. 70, 2259–2264.Google Scholar
  108. Poustka, F., Irani, N.G., Feller, A., Lu, Y., Pourcel, L., Frame, K. and Grotewold, E. (2007) Trafficking pathway for anthocyanins overlaps with the endoplasmic reticulum-to-vacuole protein-sorting route in Arabidopsis and contributes to the formation of vacuolar inclusions. Plant Physiol. 145, 1323–1335.PubMedGoogle Scholar
  109. Rhoads, D., Umbach, A.L., Subbaiah C.C. and Siedow J.N. (2006) Mitochondrial reactive oxygen species. Contribution to oxidative stress and interorganellar signaling. Plant Physiol. 141, 357–366.PubMedGoogle Scholar
  110. Rice-Evans, C.A., Miller, N. and Paganga, G. (1996) Structure-antioxidant activity relationships of flavonoids and phenolic acids. Free Radical Biol. Med. 20, 933–956.Google Scholar
  111. Rice-Evans, C.A., Miller, N. and Paganga, G. (1997) Antioxidant properties of phenolic compounds. Trends Plant Sci. 2, 152–159.Google Scholar
  112. Rizhsky, L., Liang, H.J. and Mittler, R. (2003) The water-water cycle is essential for chloroplast protection in the absence of stress. J. Biol. Chem. 278, 38921–38925.PubMedGoogle Scholar
  113. Rodríguez, A.A., Grunberg, K.A. and Taleisnik, E.L. (2002). Reactive oxygen species in the elongation zone of maize leaves are necessary for leaf extension. Plant Physiol. 129, 1627–1632.PubMedGoogle Scholar
  114. Roitsch, T. (1999) Source-sink regulation by sugar and stress. Curr. Opin. Plant Biol. 2, 198–206.PubMedGoogle Scholar
  115. Ryan, K.G. and Hunt, J.E. (2005) The effects of UVB radiation on temperate southern hemisphere forests. Environ. Pollut. 137, 415–427.PubMedGoogle Scholar
  116. Scebba, F., Sebustiani, L. and Vitagliano, C. (1999) Protective enzymes against activated oxygen species in wheat (Triticum aestivum L.) seedlings: responses to cold acclimation. J. Plant Physiol. 155, 762–768.Google Scholar
  117. Šesták, Z. and Šiffel, P. (1997) Leaf-age related differences in chlorophyll fluorescence. Photosynthetica 33, 347–369.Google Scholar
  118. Singh, A., Selvi, M. and Sharma, R. (1999) Sunlight-induced anthocyanin pigmentation in maize vegetative tissues. J. Exp. Bot. 50, 1619–1625.Google Scholar
  119. Solfanelli, C., Poggi, A., Loreii, E., Alpi, A. and Perata, P. (2006) Sucrose-specific induction of the anthocyanin biosynthetic pathway in Arabidopsis. Plant Physiol. 140, 637–646.PubMedGoogle Scholar
  120. Solovchenko, A. and Merzlyak, M. (2003) Optical properties and contribution of cuticle to UV protection in plants: experiments with apple fruit. Photochem. Photobiol. Sci. 2, 861–866.PubMedGoogle Scholar
  121. Steyn, W.J., Wand, S.J.E., Holcroft, D.M. and Jacobs, G. (2002) Anthocyanins in vegetative tissues: a proposed unified function in photoprotection. New Phytol. 155, 349–361.Google Scholar
  122. Stintzing, F.C. and Carle, F. (2005) Functional properties of anthocyanins and betalains in plants, food and human nutrition. Trends Food Sci. Technol. 15, 19–38.Google Scholar
  123. Streb, P.F., Feierabend, J. and Bligney, R. (1997) Resistance to photoinhibition of photosystem II and catalase and antioxidative protection in high mountain plants. Plant Cell Environ. 20, 1030–1040.Google Scholar
  124. Sun, J., Nishio, J.N. and Vogelmann, T.C. (1998) Green light drives CO2 fixation deep within leaves. Plant Cell Environ. 39, 1020–1026.Google Scholar
  125. Takahama, U. (2004) Oxidation of vacuolar and apoplastic phenolic substrates by peroxidases: physiological significance of the oxidation reactions. Phytochem. Rev. 3, 207–219.Google Scholar
  126. Takahashi, A., Takeda, K. and Ohnishi, T. (1991) Light-induced anthocyanin reduces the extent of damge to DNA in UV-irradiated Centaurea cyanus cells in culture. Plant Cell Physiol. 32, 541–547.Google Scholar
  127. Tanyolaç, D., Ekmekçi, Y. and Ünalan, Ş. (2007) Changes in photochemical and antioxidant enzyme activities in maize (Zea mays L.) leaves exposed to excess copper. Chemosphere 67, 89–98.PubMedGoogle Scholar
  128. van Acker, S.A.B.E., van den Berg, D.-J., Tromp, M.N.J.L., Griffioen, D.H., van Bennekom, W.P., van Der Vijgh, W.J.F. and Bast, A. (1996) Structural aspects of antioxidant activity of flavonoids. Free Radical Biol. Med. 20, 331–342.Google Scholar
  129. van den Berg, A. and Perkins, T.D. (2007) Contribution of anthocyanins to the antioxidant capacity of juvenile and senescing sugar maple (Acer saccharum) leaves. Funct. Plant Biol. 34, 714–719.Google Scholar
  130. Vranová, E., Inzé, D. and van Breusegem, F. (2002) Signal transduction during oxidative stress. J. Exp. Bot. 53, 1227–1236.PubMedGoogle Scholar
  131. Wagner, D., Przybyla, D., op den Camp, R., Kim, C., Landgraf, F., Pyo Lee, K., Würsch, M., Laloi, C., Nater, M., Hideg, E. and Apel, K. (2004) The genetic basis of singlet oxygen-induced stress responses of Arabidopsis thaliana. Science 306, 1183–1185.PubMedGoogle Scholar
  132. Wang, H., Cao, G. and Prior, R.L. (1997) Oxygen radical absorbing capacity of anthocyanins. J. Agric. Food Chem. 45, 304–309.Google Scholar
  133. Wheldale, M. (1916) The Anthocyanin Pigments of Plants. Cambridge University Press, Cambridge.Google Scholar
  134. Wise, R.R. (1995) Chilling-enhanced photooxidation: the production, action and study of reactive oxygen species produced during chilling in the light. Photosynth. Res. 45, 79–97.Google Scholar
  135. Woodall, G.S. and Stewart, G.R. (1998) Do anthocyanins play a role in UV protection of the red juvenile leaves of Syzygium? J. Exp. Bot. 49, 1447–1450.Google Scholar
  136. Yamasaki, H., Sakihama, Y. and Ikehara, N. (1997) Flavonoid-peroxidase reaction as a detoxification mechanism of plant cells against H2O2. Plant Physiol. 115, 1405–1412.PubMedGoogle Scholar

Copyright information

© Springer Science+Business Media, LLC 2008

Authors and Affiliations

  1. 1.School of Biological SciencesUniversity of AucklandAuckland
  2. 2.AgResearch Limited, Grasslands Research CentreTennent Drive, Private Bag 11008 Palmerston NorthNew Zealand
  3. 3.School of Biological SciencesVictoria University of WellingtonWellingtonNew Zealand

Personalised recommendations