Anthocyanins function as a light attenuator to compensate for insufficient photoprotection mediated by nonphotochemical quenching in young leaves of Acmena acuminatissima in winter

Ainsworth E.A., Gillespie K.M.: Estimation of total phenolic content and other oxidation substrates in plant tissues using Folin–Ciocalteu reagent.–Nat. Protoc. 2: 875–877, 2007. Albert N.W., Lewis D.H., Zhang H. et al.: Light-induced vegetative anthocyanin pigmentation in petunia.–J. Exp. Bot. 60: 2191–2202, 2009. Bilger W., Björkman O.: Role of the xanthophyll cycle in photoprotection elucidated by measurements of light-induced absorbance changes, fluorescence and photosynthesis in leaves of Hedera canariensis.–Photosynth. Res. 25: 173–185, 19 Bradford M.M.: A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding.–Anal. Biochem. 72: 248–254, 1976. Feild T.S., Lee D.W., Holbrook N.M.: Why leaves turn red in autumn. The role of anthocyanins in senescing leaves of redosier dogwood.–Plant Physiol. 127: 566–574, 2001. Genty B., Briantais J.M., Baker N.R.: The relationship between the quantum yield of photosynthetic electron transport and quenching of chlorophyll fluorescence.–Biochim. Biophys. Acta 990: 87–92 1989. Gould K.S., Kuhn D.N., Lee D.W. et al.: Why leaves are sometimes red.–Nature 378: 241–242, 1995. Gould K.S., Markham K.R., Smith R.H. et al.: Functional role of anthocyanins in the leaves of Quintinia serrata A. Cunn.–J. Exp. Bot. 51: 1107–1115, 2000. Guo Y.-H., Cao K.-F.: Effect of night chilling on photosynthesis of two coffee species grown under different irradiances.–J. Hortic. Sci. Biotech. 79: 713–716, 2004. Hakala M., Tuominen I., Keränen M. et al.: Evidence for the role of the oxygen-evolving manganese complex in photoinhibition of photosystem II.–BBA-Bioenergetics 1706: 68–80, 2005. Havaux M., Kloppstech K.: 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, 2001. He J., Yang W., Qin L. et al. Photoinactivation of photosystem II in wild type and chlorophyll b-less barley leaves: Which mechanism dominates depends on experimental circumstances.–Photosynth. Res. 126: 399–407, 2015. Heimler D., Vignolini P., Dini M.G. et al.: Rapid tests to assess the antioxidant activity of Phaseolus vulgaris L. Dry beans.–J. Agr. Food Chem. 53: 3053–3056, 2005. Hendrickson L., Ball M.C., Osmond C.B. et al.: Assessment of photoprotection mechanisms of grapevines at low temperature.–Funct. Plant Biol. 30: 631–642 2003. Hipskind J., Wood K., Nicholson R.L.: Localized stimulation of anthocyanin accumulation and delineation of pathogen ingress in maize genetically resistant Tobipolaris maydisrace O.–Physiol. Mol. Plant Pathol. 49: 247–256, 1996. Hirotsu N., Makino A., Ushio A. et al.: Changes in the thermal dissipation and the electron flow in the water–water cycle in rice grown under conditions of physiologically low temperature.–Plant Cell Physiol. 45: 635–644, 2004. Hu W., Song X., Shi K. et al.: Changes in electron transport, superoxide dismutase and ascorbate peroxidase isoenzymes in chloroplasts and mitochondria of cucumber leaves as influenced by chilling.–Photosynthetica 46: 581–588, 2008. Hughes N.M., Neufeld H.S., Burkey K.O.: Functional role of anthocyanins in high-light winter leaves of the evergreen herb Galax urceolata.–New Phytol. 168: 575–587, 20 Hughes N.M., Morley C.B., Smith W.K.: Coordination of anthocyanin decline and photosynthetic maturation in juvenile leaves of three deciduous tree species.–New Phytol. 175: 675–685, 2007. Hughes N.M., Smith W.K.: Attenuation of incident light in Galax urceolata (Diapensiaceae): Concerted influence of adaxial and abaxial anthocyanic layers on photoprotection.–Am. J. Bot. 94: 784–790, 2007. Karageorgou P., Manetas Y.: 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, 2006. Koćcielniak J., Biesaga-Koćcielniak J.: Photosynthesis and nonphotochemical excitation quenching components of chlorophyll excitation in maize and field bean during chilling at different photon flux density.–Photosynthetica 44: 174–180, 2006. Krall J.P., Edwards G.E.: Relationship between photosystem II activity and CO2 fixation in leaves.–Physiol. Plantarum 86: 180–187, 1992. Kratsch H.A., Wise R.R.: The ultrastructure of chilling stress.–Plant Cell Environ. 23: 337–350, 2000. Kumar V., Sharma S.S.: Nutrient deficiency-dependent anthocyanin development in Spirodela polyrhiza L. Schleid.–Biol. Plantarum 42: 621–624, 1999. Laine P.L., Bigot J., Ourry A. et al.: napus L.–New Phytol. 127: 675–683, 1994. Lea U.S., Slimestad R., Smedvig P. et al: Nitrogen deficiency enhances expression of specific MYB and BHLH transcription factors and accumulation of end products in the flavonoid pathway.–Planta 225: 1245–1253, 2007. Liakopoulos G., Nikolopoulos D., Klouvatou A. et al.: The photoprotective role of epidermal anthocyanins and surface pubescence in young leaves of grapevine (Vitis vinifera).–Ann. Bot.-London 98: 257–265, 2006. Liu X., Ardo S., Bunning M. et al.: Total phenolic content and DPPH radical scavenging activity of lettuce (Lactuca sativa L.) grown in Colorado.–LWT-Food Sci. Technol. 40: 552–557, 2007. Mai J., Herbette S., Vandame M. et al.: Effect of chilling on photosynthesis and antioxidant enzymes in Hevea brasiliensis Muell. Arg.–Trees 23: 863–874, 2009. Mai J., Herbette S., Vandame M. et al.: Contrasting strategies to cope with chilling stress among clones of a tropical tree, Hevea brasiliensis.–Tree Physiol. 30: 1391–1402, 20 Müller P., Li X.-P., Niyogi K.K.: Non-photochemical quenching. A response to excess light energy.–Plant Physiol. 125: 1558–1566, 2001. Müller-Moulé P., Conklin P.L., Niyogi K.K.: Ascorbate deficiency can limit violaxanthin de-epoxidase activity in vivo.–Plant Physiol. 128: 970–977, 20 Neill S.O., Gould K.S., Kilmartin P.A. et al.: Antioxidant activities of red versus green leaves in Elatostema rugosum.–Plant Cell Environ. 25: 539–547 20 Neill S.O., Gould K.S.: Anthocyanins in leaves: Light attenuators or antioxidants?–Funct. Plant Biol. 30: 865–873, 2003. Niyogi K.K.: Photoprotection revisited: Genetic and molecular approaches.–Annu. Rev. Plant Phys. 50: 333–359, 1999. Ohnishi N., Allakhverdiev S.I., Takahashi S. et al.: Two-step mechanism of photodamage to photosystem II: Step 1 occurs at the oxygen-evolving complex and step 2 occurs at the photochemical reaction center.–Biochemistry 44: 8494–8499, 2005. Oxborough K., Baker N.R.: Resolving chlorophyll a fluorescence images of photosynthetic efficiency into photochemical and non-photochemical components–calculation of qP and Fv’/Fm’; without measuring Fo’.–Photosynth. Res. 54: 135–142 1997. Peng S., Wang B.: Forest succession of Dinghushan, Guangdong, China.–Chin. J. Bot. 7: 75–80, 1994. Pietrini F., Iannelli M., Massacci A.: 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, 2002. Saha M.R, Hasan S.M.R., Akter R. et al.: In vitro free radical scavenging activity of methanol extract of the leaves of Mimusops elengi Linn.–Bangl. J. Vet. Med. 6: 197–202, 2008. Steyn W.J., Wand S.J., Jacobs G. et al.: Evidence for a photoprotective function of low-temperature-induced anthocyanin accumulation in apple and pear peel.–Physiol. Plantarum 136: 461–472, 2009. Tucić B., Vuleta A., Jovanović S.M.: Protective function of foliar anthocyanins: In situ experiments on a sun-exposed population of Iris pumila L. (Iridaceae).–Pol. J. Ecol. 57: 779–783, 2009. van den Berg A.K., Perkins T.D.: Contribution of anthocyanins to the antioxidant capacity of juvenile and senescing sugar maple (Acer saccharum) leaves.–Funct. Plant Biol. 34: 714–719, 2007. Wang L.-Z., Wang L.-M., Xiang H.-T. et al.: Relationship of photosynthetic efficiency and seed-setting rate in two contrasting rice cultivars under chilling stress.–Photosynthetica 54: 581–588, 2016. Wellburn A.R.: The spectral determination of chlorophylls a and b, as well as total carotenoids, using various solvents with spectrophotometers of different resolution.–J. Plant Physiol. 144: 307–313, 1994. Zeng X.Q., Chow W.S., Su L.J. et al.: Protective effect of supplemental anthocyanins on Arabidopsis leaves under high light.–Physiol. Plantarum 138: 215–225, 2010. Zhang K.-M., Yu H.-J., Shi K. et al.: Photoprotective roles of anthocyanins in Begonia semperflorens.–Plant Sci. 179: 202–208, 20 Zhang Q., Zhang T.-J., Chow W.S. et al.: Photosynthetic characteristics and light energy conversions under different light environments in five tree species occupying dominant status at different stages of subtropical forest succession.–Funct. Plant Biol. 42: 609–619, 2015. Zhang T.-J., Chow W.S., Liu X.-T. et al: A magic red coat on the surface of young leaves: Anthocyanins distributed in trichome layer protect Castanopsis fissa leaves from photoinhibition.–Tree Physiol. 36: 1296–1306, 2016.