花色苷（书籍文献）Anthocyanins，Kevin Gould · Kevin Davies · Chris Winefield，Biosynthesis, Functions, and Applications.pdf_P31-P35

Anthocvanin Function in vegetative Organs 17 Murray,J.R.and Hackett,W.P.(1991)Dihydroflavonol reductase activity in relation to in juvenile and mature phase Neill.S.O.an nd Gould K S (1999)Ontical of leaves in relation to anthocyanin concentration and distribution.Can.J.Bot.77.1777-1782. Neill,S.O.and Gould,K.S.(2003)Anthocyanins in leaves:light attenuators or antioxidants? Neill sO Gould Ks Kilmartin PA Mitchell K A and Markham K R (2002 Antioxidant activities of red versus green leaves of Elatostema rugosum.Plant Cell S.o Kilmartin, P.A.,Mitchell,K.A.and Markham K.R.(2002c An nct.Plant Biol.29 1437-1443 and cyanic leaves in th e sun species,Ountinia serrala NishioN.0Whyare hir pantsgEvolution of the higher plant photosynthetic pigment complement.Plant Cell Environ.23,539-548. Niyogi,K.K.()Safety valves for photosynthesis.Curr.Opin Plant Biol.,455460. nced UV-B radiati iol.Plant.107. PeltigrewW.and Vaughn.K.C(Physioloic structural,and immunoloical PfundelFENB Mever s and Cerovic G007 Investiti Pfunde IC, Z.G.(2007)Investigating UV screening rent typep of porta creening by th Re 3.reveals in vi Pietrini.F lannelli.MA and Massacei A.(2002)Anthocyanin accumulation in the ue whout urther my illuminated surface of maize leaves enhances protection from photo-inhibitory risks at Plant Cell Environ.25 Rao.M.V..Paliyath,G.,Murt DP and Fletcher,R.A.(1997)Cha of an and th induced chilling tolerance of maize seedlings.Plant Physiol.114.695-704. Pitzschke,A.,Forzani,C.and Hirt,H.(2006)Reactive oxygen species signaling in plants. Redox Signal.8,1757-1764 Polle,A.(1997)Defens Spring Harbor a Defenses.Ne Polle,A.(2001)Dissecting the s uperoxide dismutase-ascorbate peroxidase-glutathione pathway in chloroplasts by metabolic modelling.Computer simulations as a step towards Pos A. Post and Vesk.M.()Photosynthesis.pigments.and chloroplast ultra ucture of ar antarctic liverwort from sun-exposed and shaded sites.Can.J.Bot.70,2259-2264. 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

Anthocyanin Function in Vegetative Organs 17 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. 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. Neill, S.O. and Gould, K.S. (2003) Anthocyanins in leaves: light attenuators or antioxidants? Funct. Plant Biol. 30, 865–873. Neill, S., Desikan, R. and Hancock, J. (2002a) Hydrogen peroxide signalling. Curr. Opin, Plant Biol. 5, 388–395. 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. 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. Nishio, J.N. (2000) Why are higher plants green? Evolution of the higher plant photosynthetic pigment complement. Plant Cell Environ. 23, 539–548. Niyogi, K.K. (2000) Safety valves for photosynthesis. Curr. Opin. Plant Biol. 3, 455–460. Olsson, L.C., Veit, M. and Bornman, J.F. (1999) Epidermal transmittance and phenolic composition in leaves of atrazine-tolerant and atrazine-sensitive cultivars of Brassica napus grown under enhanced UV-B radiation. Physiol. Plant. 107, 259–266. Pettigrew, W.T. and Vaughn, K.C. (1998) Physiological, structural, and immunological characterization of leaf and chloroplast development in cotton. Protoplasma 202, 23–37. 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. 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. 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 paclobutrazolinduced chilling tolerance of maize seedlings. Plant Physiol. 114, 695–704. Pitzschke, A., Forzani, C. and Hirt, H. (2006) Reactive oxygen species signaling in plants. Antioxid. Redox Signal. 8, 1757–1764. 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. 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. Post, A. (1990) Photoprotective pigment as an adaptive strategy in the Antarctic moss Ceratodon purpureus. Polar Biol. 10, 241–245. 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. 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

18 J.-H. B. Hatier, K.S. Gould 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. 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. Rice-Evans, C.A., Miller, N. and Paganga, G. (1997) Antioxidant properties of phenolic compounds. Trends Plant Sci. 2, 152–159. 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. 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. Roitsch, T. (1999) Source-sink regulation by sugar and stress. Curr. Opin. Plant Biol. 2, 198–206. Ryan, K.G. and Hunt, J.E. (2005) The effects of UVB radiation on temperate southern hemisphere forests. Environ. Pollut. 137, 415–427. 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. Šesták, Z. and Šiffel, P. (1997) Leaf-age related differences in chlorophyll fluorescence. Photosynthetica 33, 347–369. Singh, A., Selvi, M. and Sharma, R. (1999) Sunlight-induced anthocyanin pigmentation in maize vegetative tissues. J. Exp. Bot. 50, 1619–1625. 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. 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. 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. 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. 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. Sun, J., Nishio, J.N. and Vogelmann, T.C. (1998) Green light drives CO2 fixation deep within leaves. Plant Cell Environ. 39, 1020–1026. Takahama, U. (2004) Oxidation of vacuolar and apoplastic phenolic substrates by peroxidases: physiological significance of the oxidation reactions. Phytochem. Rev. 3, 207–219. 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. 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. 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. 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

2 Role of Anthocyanins in Plant Defence Abstract. In addition to their well-documented beneficial effects on plant physiological processes, anthocyanins have also been proposed to function in a diverse array of plant/animal interactions. These include the attraction of pollinators and frugivores, as well as the repellence of herbivores and parasites. The optical properties of anthocyanins may serve as visual signals to potential herbivores, indicating a strong metabolic investment in toxic or unpalatable chemicals. Anthocyanins have also been implicated in the camouflage of plant parts against their backgrounds, in the undermining of insect crypsis, and in the mimicry of defensive structures. These hypotheses have in recent years attracted strong theoretical support and increasing experimental evidence. We emphasize that both the defensive and the physiological functions of anthocyanins may operate in plants simultaneously. 2.1 Introduction In their natural environments, plants run the risk of multiple attacks by many different species of herbivores and pathogens. Phytophagous species feature in all major groups of vertebrates; mammals and birds are undoubtedly the most injurious to plants, but reptiles, amphibians, and fish also inflict damage (Schulze et al. 2002). Invertebrates, too, have the potential to devastate plant communities. In a collection of German woodlands, for example, Reichelt and Wilmanns (1973) identified numerous species of leaf chewers, excavators, leaf miners, bark borers, wood borers, sap suckers, bud and shoot eaters, root eaters, and seedling eaters. Some herbivores feed preferentially at the canopy, others focus on lower branches or seedlings; some parasitize the sugars and nitrogen flowing through leaf veins, others will eat the lamina tissues surrounding the veins; some cause the plants to develop galls in which the animals live and feed, yet others employ chemical signals to redirect the flow of plant nutrients in their direction (Karban and Baldwin 1997). Little wonder, therefore, that plants have evolved elaborate strategies of avoidance and/or a sophisticated armoury of morphological devices to counteract herbivore attacks. Chemical weaponry, too, is known to play a significant role in plant defence. A vast K. Gould et al. (eds.), Anthocyanins, DOI: 10.1007/978-0-387-77335-3_2, © Springer Science+Business Media, LLC 2009 Simcha Lev-Yadun1 and Kevin S. Gould2 1 Department of Biology Education, Faculty of Science and Science Education, University of Haifa, Oranim, Tivon 36006, Israel, levyadun@research.haifa.ac.il 2 School of Biological Sciences, Victoria University of Wellington, P.O. Box 600, Wellington, New Zealand, kevin.gould@vuw.ac.nz

22 s Lev-Yadun KS Gould assortment of secondary metabolites has been demonstrated to act as antifeedants toxins,warning signals,or precursors to physical defence systems (Bennett and Wallsgrove 1994;Harborne 1997).Among them are the phenolics,a large group of structurally diverse compounds that includes terpenoids,cinnamic acids,catechols. coumarins,tannins,as well as certain flavonoids such as the anthocyanins,the subject of this chapter. There are several different ways anthocvanins might assist plants in their defence against other organisms.These include both direct roles as chemical repellents and more indirect roles as visual signals.In common with other flavonoids,certain anthocyanins have demonstrable antiviral,antibacterial,and fungicidal activities (Konczak and Zhang 2004:Wrolstad 2004,and references therein).They have the potential,therefore,to protect plants from infections by pathogenic micro organism In general,howe ver,the antimicrobial activities of anthocyanins area effective than those of other phenolic compo nds such as kev hydrox nnamic acids that also likely to be nt in the shoo (Padmavati et al 1997:Werlein et al.2005).Moreo ps have not been found to be toxic to any higher animal sp es (Lee et al.1987).Aphid survival ra s,for example,ar in their diet (Cost 2001).Thus chemical defence is unlikely to be functio of these pig s in plan ole stron the upport and owing emp g roles of hough of the echa involve associated chemica or mechanica com ccepted that the colo rs of flowe ers and fruits enhance su 1990,we (Ridley 193 30.commu and van der n pla eieo fer et al.2004),there is no a prior reason to assum tha urs annot in de nce from herbivory Thi colours(i)undermine an herbivorous invertebrate's camouf age,(I1 are apose an unpalatable plant or anim or (Iv)serve in pl camouflage (see Hinton 1973:Givnish 1990:Cole and Cole 2005;Lev-Yadun 2006) Of course,anthocyanins are not the only class of pigments that might contribute to defence in these ways.In several species,leaf variegation caused by pigments unrelated to anthocyanins has been shown to correlate to reduced herbivory (Cahn and Harper 1976;Smith 1986).Such examples may,however,help to understand the principles that operate when anthocyanins serve in defence. Moreover.it ha long been recognised that the non-photosynthetic plant pigments have the potentia to serve more than one function concurrently (Gould et al.2002;Lev-Yadun et al 2002 2004:Schaefer and Wilkinson 2004:Lev-Yadun 2006).The UV-absorbing dearomatized isoprenylated phloroglucinols.for example.serve a defensive role in the stamens and ovaries of Hypericum calycinum,but an attractive role in the petals of the same species (Gronguist et al 2001)Thus the various functional hypotheses concerning pigmentation in leaves and other plant parts need not contrast or exclude any other functional explanation for specific types of plant colouration,and those traits,such as colouration,that might have more than one type of benefit,may be

22 S. Lev-Yadun, K.S. Gould assortment of secondary metabolites has been demonstrated to act as antifeedants, toxins, warning signals, or precursors to physical defence systems (Bennett and Wallsgrove 1994; Harborne 1997). Among them are the phenolics, a large group of structurally diverse compounds that includes terpenoids, cinnamic acids, catechols, coumarins, tannins, as well as certain flavonoids such as the anthocyanins, the subject of this chapter. There are several different ways anthocyanins might assist plants in their defence against other organisms. These include both direct roles as chemical repellents and more indirect roles as visual signals. In common with other flavonoids, certain anthocyanins have demonstrable antiviral, antibacterial, and fungicidal activities (Konczak and Zhang 2004; Wrolstad 2004, and references therein). They have the potential, therefore, to protect plants from infections by pathogenic microorganisms. In general, however, the antimicrobial activities of anthocyanins are appreciably less effective than those of other phenolic compounds, such as key flavonols and hydroxycinnamic acids that are also likely to be present in the shoot (Padmavati et al. 1997; Werlein et al. 2005). Moreover, anthocyanins have not been found to be toxic to any higher animal species (Lee et al. 1987). Aphid survival rates, for example, are unaffected by anthocyanins in their diet (Costa-Arbulú et al. 2001). Thus, direct chemical defence is unlikely to be a major function of these pigments in plants. There is, in contrast, strong theoretical support and growing empirical evidence for a role of anthocyanins in the defence from “visually oriented” herbivores. This discussion focuses largely on the defensive roles of anthocyanins as visual cues, though some of these mechanisms also involve associated chemical or mechanical components such as poisons and thorns. Although it is generally accepted that the colours of flowers and fruits enhance reproductive success by facilitating communication between plants, their pollinators, and seed-dispersers (Ridley 1930; Faegri and van der Pijl 1979; Willson and Whelan 1990; Weiss 1995; Schaefer et al. 2004), there is no a priori reason to assume that flower, fruit and leaf colours cannot also serve in defence from herbivory. This is achievable if the colours (i) undermine an herbivorous invertebrate’s camouflage, (ii) are aposematic, (iii) mimic an unpalatable plant or animal, or (iv) serve in plant camouflage (see Hinton 1973; Givnish 1990; Cole and Cole 2005; Lev-Yadun 2006). Of course, anthocyanins are not the only class of pigments that might contribute to defence in these ways. In several species, leaf variegation caused by pigments unrelated to anthocyanins has been shown to correlate to reduced herbivory (Cahn and Harper 1976; Smith 1986). Such examples may, however, help to understand the principles that operate when anthocyanins serve in defence. Moreover, it has long been recognised that the non-photosynthetic plant pigments have the potential to serve more than one function concurrently (Gould et al. 2002; Lev-Yadun et al. 2002, 2004; Schaefer and Wilkinson 2004; Lev-Yadun 2006). The UV-absorbing dearomatized isoprenylated phloroglucinols, for example, serve a defensive role in the stamens and ovaries of Hypericum calycinum, but an attractive role in the petals of the same species (Gronquist et al. 2001). Thus, the various functional hypotheses concerning pigmentation in leaves and other plant parts need not contrast or exclude any other functional explanation for specific types of plant colouration, and those traits, such as colouration, that might have more than one type of benefit, may be