花色苷（书籍文献）Anthocyanins，Kevin Gould · Kevin Davies · Chris Winefield，Biosynthesis, Functions, and Applications.pdf_P26-P30

12 J.-H.B.Hatier,K.S.Gould (Bowler and Fluhr 2000).Despite its efficier roplasts,mitoc ria,and pero somes, may ere stress (Yamasak et a ole typically occupy more than 70%of the mature plant cell volume, and as a s of all the majo anthocyanins are likely to have a crucial role,especially in preventing the symplastic movement of ROS from one cell to another(Mittler et al.2004). Finally,anthocyanins may interact with secondary messengers downstream of the ROS signalling pathway,or else be involved in the crosstalk with other response pathways.An intriguing possibility is the interaction between anthocyanins and sucrose.The anthocyanin biosynthetic pathway is strongly upregulated by sucrose in plants as diverse as radish (Raphanus sativus)English ivy (Hedera helix).and Arabidopsis thaliana (Murray and Hackett 1991;Hara et al.2003;Solfanelli et al. 2006).Soluble sugars,especially sucrose,glucose,and fructose,are now known to play central roles in the control of plant development.stress responses.and gene expression (Gibson 2005).Sugar accumulation has been associated with improved tolerance to diverse stressors including drought,salinity,high light,cold,anoxia and herbicides(Roitsch 1999:Couee et al.2006).It has been sugg sted that these roles relate to the regulation of the pro-oxidant and antioxidant balance in plant cells sucrose is known to be involved in both ROS-producing and ROS-scavenging metabolic However,the mechanism by which established A sinalline role for anthocvanins is attractive hecause it potentially also explains why anthocyanins often accumulate in c tosynthesise,or else for photosynthetic carbon assimilation is not the prim such as stem petioles and adventitious roots It can also explain why anthoc species ntially produced at certain developr ental stag s such as seed leaf i or leat h Establish g po sible relatio ellula sents the e of an ex new line investigat o this plant pigments References Adir,N.,Zer,H,Scholat,S.and Ohad,1.(2003)Photoinhibition-a historical perspective. p63 and Tattini,M.(2007)Chloroplast-located flavonoids car 4770 Alenius CM and 995)A three-dimensional representation of the relationship between penetration of U.V.-B radiation and U.V.-screening pigments in leaves o ica napu w Phytol.131,297-302 nA.C.and Alscher.R.G..Donahue. and Cr C.L(1997)Reactive oxygen species and antioxidants:relationships in green cells.Physiol.Plant.100,224-233

12 J.-H. B. Hatier, K.S. Gould (Bowler and Fluhr 2000). Despite its efficient scavenging by enzymes in the chloroplasts, mitochondria, and peroxisomes, H2O2 may leak into the cytosol and possibly the vacuole during periods of severe stress (Yamasaki et al. 1997). Vacuoles typically occupy more than 70% of the mature plant cell volume, and as a consequence of their size, vacuoles are one of the closest neighbours of all the major sources of organelle-derived ROS. For this reason, vacuolar LMWAs such as the anthocyanins are likely to have a crucial role, especially in preventing the symplastic movement of ROS from one cell to another (Mittler et al. 2004). Finally, anthocyanins may interact with secondary messengers downstream of the ROS signalling pathway, or else be involved in the crosstalk with other response pathways. An intriguing possibility is the interaction between anthocyanins and sucrose. The anthocyanin biosynthetic pathway is strongly upregulated by sucrose in plants as diverse as radish (Raphanus sativus), English ivy (Hedera helix), and Arabidopsis thaliana (Murray and Hackett 1991; Hara et al. 2003; Solfanelli et al. 2006). Soluble sugars, especially sucrose, glucose, and fructose, are now known to play central roles in the control of plant development, stress responses, and gene expression (Gibson 2005). Sugar accumulation has been associated with improved tolerance to diverse stressors including drought, salinity, high light, cold, anoxia and herbicides (Roitsch 1999; Couée et al. 2006). It has been suggested that these roles relate to the regulation of the pro-oxidant and antioxidant balance in plant cells; sucrose is known to be involved in both ROS-producing and ROS-scavenging metabolic pathways (Couée et al. 2006). However, the mechanism by which sucrose, anthocyanins, and ROS might contribute to plant function remains to be established. A signalling role for anthocyanins is attractive because it potentially also explains why anthocyanins often accumulate in organs that do not photosynthesise, or else for which photosynthetic carbon assimilation is not the primary function, such as stems petioles, and adventitious roots. It can also explain why anthocyanins are in some species preferentially produced at certain developmental stages, such as seed dormancy, leaf initiation or leaf senescence, or in certain seasons such as autumn or spring. Establishing possible relationships between the cellular redox balance and anthocyanin function presents the promise of an exciting, new line of investigation into this intriguing class of plant pigments. References Photosynth. Res. 76, 343–370. Agati, G., Matteini, P., Goti, A. and Tattini, M. (2007) Chloroplast-located flavonoids can scavenge singlet oxygen. New Phytol. 174, 77–89. 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. Allan, A.C. and Fluhr, R. (1997) Two distinct sources of elicited reactive oxygen species in tobacco epidermal cells. Plant Cell 9, 1559–1572. 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. Adir, N., Zer, H., Scholat, S. and Ohad, I. (2003) Photoinhibition – a historical perspective

Anthocyanin Function in Vegetative Organs 13 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. Apel, K. and Hirt, H. (2004) Reactive oxygen species: metabolism, oxidative stress, and signal transduction. Annu. Rev. Plant Biol. 55, 373–399. 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. Asada, K. (2000) The water-water cycle as alternative photon and electron sinks. Phil. Trans. R. Soc. Lond. B 355, 1419–1431. 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. Bowler, C., van Montagu, M. and Inzé, D. (1992) Superoxide dismutase and stress tolerance. Annu. Rev. Plant Physiol. Plant Mol. Biol. 43, 83–116. 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. Buchholz, G., Ehmann, 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. 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. 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. Caldwell, M.M., Robberecht, R. and Flint, S.D. (1983) Internal filters: prospects for UVacclimation in higher plants. Physiol. Plant. 58, 445–450. 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. Chalker-Scott, L. (1999) Environmental significance of anthocyanins in plant stress responses. Photochem. Photobiol. 70, 1–9. Chalker-Scott, L. (2002) Do anthocyanins function as osmoregulators in leaf tissues? Adv. Bot. Res. 37, 103–127. 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. Close, D.C. and Beadle, C.L. (2003) The ecophysiology of foliar anthocyanin. Bot. Rev. 69, 149–161. 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. 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. 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. 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. 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

14 J.-H. B. Hatier, K.S. Gould Dröge, W. (2002) Free radicals in the physiological control of cell function. Physiol. Rev. 82, 47–95. 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. Foyer, C.H. and Noctor, G. (2005) Oxidant and antioxidant signalling in plants: a reevaluation of the concept of oxidative stress in a physiological context. Plant Cell Environ. 28, 1056–1071. Foyer, C.H., Lelandais, M. and Kunert, K.J. (1994) Photooxidative stress in plants. Physiol. Plant. 92, 696–717. Fry, S.C. (1998) Oxidative scission in plant cell wall polysaccharides by ascorbate-induced hydroxyl radicals. Biochem. J. 332, 507–515. 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. Gibson, S. (2005) Control of plant development and gene expression by sugar signaling. Curr. Opin. Plant Biol. 8, 93–102. Gitelson, A.A., Merzylak, M.N. and Chivkunova, O.B. (2001) Optical properties and nondestructive estimation of anthocyanin content in plant leaves. Photochem. Photobiol. 74, 38–45. 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. 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. 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. Gould, K.S. (2004) Nature’s Swiss army knife: the diverse protective roles of anthocyanins in leaves. J. Biomed. Biotechnol. 2004, 314–320. 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. 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. Gould, K.S., Kuhn, D.N., Lee, D.W. and Oberbauer, S.F. (1995) Why leaves are sometimes red. Nature 378, 241–242. 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. 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. Gould, K.S., Neill, S.O. and Vogelmann, T.C. (2002b) A unified explanation for anthocyanins in leaves? Adv. Bot. Res. 37, 167–192. 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. 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. 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

Anthocvanin Function in vegetative Organs 15 Halliwell,B.and Gutteridge,JM.C.(1)Free Radicals in Biologyand Medicine.Oxford .K.and Kuboi.T.(200)Enhar Havaux.M.and Kloppstech.K.(2001)The protective functions of carotenoid and flavonoid pigments against excess visible radiation at chilling temperature investigated in Planta 213,9 Zeldin,E.L.an I significance of Hoch,W.A..Singsaas.EL.and McCown.B.H.(2003)Reso ion protection Anthocvanin facilitate nutrient recovery in Autumn by shielding leaves from potentially damaging light 13 C.A and Gou .(2007)Photopro red and green 0 .G.1999.Nat ral Uvns.New Ze Photochem Photobiol 6 177-19 s of Norway spruce (Pice 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 Bot.9 K O (2005)Fun -light winter 57587 Hughes,N.M.,Morley,C.B.and Smith,W.K.(2007)Coordination of anthocyanin decline and c maturation in juvenile leaves of three deciduous tree species.New Phytol I S Hull MR and Lo S.P.(1991)Chi metabolizing s in Zea m anden月 Jordan,B.R James,P Strid,A.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. E virange ne-specinic and dependent upon the developmental stage.Plant P and Man Y (2006)The of heir herbivory and excess light.Tree Physiol.26,613-621. Kato,M.and Shimiz lism in higher plants VI.Involvement of er ophyll egrada 6.1291-130 Kaw T.(2003 and o wth induct Koostra.A.()Protection from UV-B induced DNA damage by flavonoids.Plant Mol Biol.26,771-774 Krause,G.H and photosynthes the basics NPA (1995)Lo temperature stress and photop increased tolerance to photoinhibition in Pinus banksiana seedlings.Can.J.Bot.73,1119-1127. Kuk,Y.I,Shin,J.S.,Burgos,N.R,Hwang,T.E.,Han,O.,Cho,B.H.,Jung.S.and Guh J.O 0.2109-2 ve enzymes offer protection from chilling damage in rice plants.Crop Kunz,S.and Bec H.(1995) t f liverwort Ricciocarpos natans.Z.Naturfor tion of in vitro cultures of the

Anthocyanin Function in Vegetative Organs 15 Halliwell, B. and Gutteridge, J.M.C. (1999) Free Radicals in Biology and Medicine. Oxford University Press, Oxford. 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. 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. Hoch, W.A., Zeldin, E.L. and McCown, B.H. (2001) Physiological significance of anthocyanins during autumnal leaf senescence. Tree Physiol. 21, 1–8. Hoch, W.A., Singsaas, 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, 1296–1305. 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. Hoque, E. and Remus, G. (1999). Natural UV-screening mechanisms of Norway spruce (Picea abies [L.] Karst.) needles. Photochem. Photobiol. 69, 177–192. 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. 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. 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. 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. 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-Binduced changes are gene-specific and dependent upon the developmental stage. Plant Cell Environ. 17, 45–54. 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. Kato, M. and Shimizu, S. (1985) Chlorophyll metabolism in higher plants VI. Involvement of peroxidase in chlorophyll degradation. Plant Cell Physiol. 26, 1291–1301. Kawano, T. (2003) Roles of the reactive oxygen species-generating peroxidase reactions in plant defense and growth induction. Plant Cell Rep. 21, 829–837. Koostra, A. (1994) Protection from UV-B induced DNA damage by flavonoids. Plant Mol. Biol. 26, 771–774. Krause, G.H. and Weis, E. (1991) Chlorophyll fluorescence and photosynthesis: the basics. Annu. Rev. Plant Physiol. Plant Mol. Biol. 42, 313–349. Krol, M., Gray, G.R., Hurry, V.M., Oquist, G., Malek, L. and Huner, N.P.A. (1995) Lowtemperature stress and photoperiod affect an increased tolerance to photoinhibition in Pinus banksiana seedlings. Can. J. Bot. 73, 1119–1127. 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. Kunz, S. and Becker, H. (1995) Cell wall pigment formation of in vitro cultures of the liverwort Ricciocarpos natans. Z. Naturforsch. 50, 235–240

16 J.-H. B. Hatier, K.S. Gould 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. 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. Kytridis, V.-P. and Manetas, Y. (2006) Mesophyll versus epidermal anthocyanins as potential in vivo antioxidants: evidence linking the putative antioxidant role to the proximity of oxyradical source. J. Exp. Bot. 57, 2203–2210. Laloi, C., Apel, K. and Danon A. (2004) Reactive oxygen signalling: the latest news. Curr. Opin. Plant Biol. 7, 323–328. 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. Lee, D.W. and Gould, K.S. (2002a) Anthocyanins in leaves and other vegetative organs: an introduction. Adv. Bot. Res. 37, 2–16. Lee, D.W. and Gould, K.S. (2002b) Why leaves turn red. Am. Sci. 90, 524–531. 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. 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. 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. 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. Mahalingam, R. and Fedoroff, N. (2003) Stress response, cell death and signalling: the many faces of reactive oxygen species. Physiol. Plant. 119, 56–68. Manetas, Y. (2006) Why some leaves are anthocyanic, and why most anthocyanic leaves are red. Flora 201, 163–177. 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. 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. Markham, K.R. (1982) Techniques of Flavonoid Identification. Academic Press, London. Maxwell, K. and Johnson, G.N. (2000) Chlorophyll fluorescence – a practical guide. J. Exp. Bot. 51, 659–668. 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 growthfactor receptor. J. Agric. Food Chem. 49, 958–962. 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. Mittler, R. (2002) Oxidative stress, antioxidants and stress tolerance. Trends Plant Sci. 7, Mittler, R., Vanderauwera, S., Gollery, M. and van Breusegem, F. (2004) Reactive oxygen gene network of plants. Trends Plant Sci. 9, 490–498. 405–410. McClure, J.W. (1975) Physiology and functions of flavonoids. In: Harborne, J.B., Mabry, T.J. and Mabry, H. (Eds.), The Flavonoids. Chapman and Hall, London, pp. 970–1055