Grape (Vitis vinifera cv Silvaner) vine plants were cultivated under shaded conditions in the absence of ultraviolet (UV) radiation in a greenhouse, and subsequently placed outdoors under three different light regimes for 7 d. Different light regimes were produced by filters transmitting natural radiation, or screening out the UV-B (280-315 nm), or screening out the UV-A (315-400 nm) and the UV-B spectral range. During exposure, synthesis of UV-screening phenolics in leaves was quantified using HPLC: All treatments increased concentrations of hydroxycinnamic acids but the rise was highest, reaching 230% of the initial value, when UV radiation was absent. In contrast, UV-B radiation specifically increased flavonoid concentrations resulting in more than a 10-fold increase. Transmittance in the UV of all extracted phenolics was lower than epidermal UV transmittance determined fluorimetrically, and the two parameters were curvilinearly related. It is suggested that curvilinearity results from different absorption properties of the homogeneously dissolved phenolics in extracts and of the non-homogeneous distribution of phenolics in the epidermis. UV-B-dependent inhibition of maximum photochemical yield of photosystem II (PSII), measured as variable fluorescence of dark-adapted leaves, recovered in parallel to the buildup of epidermal screening for UV-B radiation, suggesting that PSII is protected against UV-B damage by epidermal screening. However, UV-B inhibition of CO 2 assimilation rates was not diminished by efficient UV-B screening. We propose that protection of UV-B inactivation of PSII is observed because preceding damage is efficiently repaired while those factors determining UV-B inhibition of CO 2 assimilation recover more slowly.Photosynthetic organisms form energy-rich compounds using the energy of the sun's visible radiation. When harvesting light, photosynthetic organs are inevitably exposed to the UV region of natural radiation. In general, UV radiation damages lipids, nucleic acids, and proteins in leaves of higher plants, and specifically targets the photosystem II (PSII) reaction center, Rubisco, chloroplast ATPase, and violaxanthin deepoxidase (Jordan, 1996;Vass, 1997).To cope with UV radiation damage, plants have evolved a variety of mechanisms including: screening out UV radiation by accumulating UV-absorbing phenolic compounds in the leaf epidermis, repairing UV-induced DNA damage, and formation of antioxidants to scavenge peroxides and oxygen radicals (Bornman and Teramura, 1993; Jordan 1996). Increases in natural UV radiation due to decreased stratospheric ozone concentrations have stimulated research on mechanisms and maximum capacities for protection against UV exposure (Caldwell et al., 1998).Studies with Arabidopsis mutants deficient in synthesis of phenolic sunscreens have demonstrated the essential role of epidermal screening in UV protection (Li et al., 1993; Lois and Buchanan, 1994; Landry et al., 1995; Booij-James et al., 2000; Mazza et al., 2000). In many higher plants, two classes...
The role of phenolics in UV-screening was investigated in berries of a white grape cultivar (Vitis vinifera L. cv. Bacchus). Fluorescence microscopy revealed accumulation of phenolics in the skin of berries and, by high performance liquid chromatography and mass spectrometry, flavonols and hydroxycinnamic acids were identified as the main groups of UV-absorbing phenolics. Relationships between natural radiation and the synthesis of phenolics were studied in plants that were cultivated in the absence of UV radiation in a greenhouse before outdoor exposure to three different light regimes: the entire solar spectrum, the solar spectrum minus UV-B radiation and only visible radiation. During six days of exposure, flavonol synthesis was significantly stimulated by natural UV, in particular UV-B, but concentrations of hydroxycinnamic acids decreased under all conditions. Direct comparison of fluorimetrically-determined skin absorbance with absorbance of extracted flavonols or hydroxycinnamic acids suggested that acclimation of UV screening depends almost exclusively on flavonol synthesis. While increased flavonol levels resulted in efficient UV-A shielding, UV-B shielding was incomplete, probably due to decreased levels of the UV-B-absorbing hydroxycinnamic acids during exposure.
A recent review of climate patterns in Southern Germany has suggested significant increases in ultraviolet (UV) radiation due to decreases in cloud coverage and in cloud frequency which compound the effects of stratospheric ozone depletion. Whether such UV radiation increases result in UV damage of higher plant leaves depends partly on the capacity of UV-absorbing hydroxycinnamic acids and flavonoids located in the plant epidermis to screen out UV radiation. Epidermal UV screening is most often assessed from UV absorbance of whole-leaf extracts but in the present work, this method is critically examined. In grapevine ( Vitis vinifera L.), hydroxycinnamic acid as well as mono-hydroxylated and ortho-dihydroxylated flavonoid concentrations increased in parallel with fluorometrically detected adaxial epidermal UV absorbance but only the latter class of flavonoids was associated with epidermal UV absorbance in barley ( Hordeum vulgare L). For both species, curvilinear relationships between epidermal and total phenolic UV absorbance were established: initial slopes of the curves differed markedly between species. Modelling suggested that curvilinearity arises from UV-transparent epidermal areas located between vacuoles which are particularly UV-absorbing due to high levels of phenolics. The species-dependent differences were related to allocation of high amounts of phenolics in the mesophyll and abaxial epidermis in barley but not in grapevine. Both factors, optical heterogeneity and variable distribution of phenolics, severely restrict the use of phenolic absorbance to estimate true epidermal screening.
Grape (Vitis vinifera cv Silvaner) vine plants were cultivated under shaded conditions in the absence of ultraviolet (UV) radiation in a greenhouse, and subsequently placed outdoors under three different light regimes for 7 d. Different light regimes were produced by filters transmitting natural radiation, or screening out the UV-B (280-315 nm), or screening out the UV-A (315-400 nm) and the UV-B spectral range. During exposure, synthesis of UV-screening phenolics in leaves was quantified using HPLC: All treatments increased concentrations of hydroxycinnamic acids but the rise was highest, reaching 230% of the initial value, when UV radiation was absent. In contrast, UV-B radiation specifically increased flavonoid concentrations resulting in more than a 10-fold increase. Transmittance in the UV of all extracted phenolics was lower than epidermal UV transmittance determined fluorimetrically, and the two parameters were curvilinearly related. It is suggested that curvilinearity results from different absorption properties of the homogeneously dissolved phenolics in extracts and of the non-homogeneous distribution of phenolics in the epidermis. UV-B-dependent inhibition of maximum photochemical yield of photosystem II (PSII), measured as variable fluorescence of dark-adapted leaves, recovered in parallel to the buildup of epidermal screening for UV-B radiation, suggesting that PSII is protected against UV-B damage by epidermal screening. However, UV-B inhibition of CO(2) assimilation rates was not diminished by efficient UV-B screening. We propose that protection of UV-B inactivation of PSII is observed because preceding damage is efficiently repaired while those factors determining UV-B inhibition of CO(2) assimilation recover more slowly.
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