Short communication. Do anthocyanins play a role in UV protection of the red juvenile leaves of Syzygium?

The biological function of juvenile leaves pigmented with anthocyanin is poorly understood. The role anthocyanins play in UV protection was assessed in juvenile leaves of two Syzygium species (S. luehmannii and S. wilsonii) which contain high anthocyanin concentrations. HPLC was used to separate UV-absorbing anthocyanins from other soluble UV-absorbing phenolic compounds. The isolated anthocyanins (predominantly malvidin-3,5-diglucoside) contributed little to the total absorbance of UV-A and UV-8 radiation. This was because the non-acylated anthocyanins only effectively absorbed shortwave UV-B radiation and the strong absorbance by other compounds. These results suggest that the UV protection hypothesis is not valid for anthocyanins in juvenile Syzygium leaves.     

Higher amounts of anthocyanins and UV-absorbing compounds effectively lowered CPD photorepair in purple rice (Oryza sativa L.)

Growth of a near‐isogenic line (NIL) for the purple leaf gene Pl of rice with a genetic background of Taichung 65 (T‐65) rice was significantly retarded by supplementary ultraviolet‐B radiation (UV‐B), despite the fact that the amounts of UV‐absorbing compounds and anthocyanins in NIL were significantly higher than those in T‐65. In order to understand the role of flavonoids in UV‐B induced damage protection in T‐65 and the NIL, both the (1) relationships between changes in the steady state of cyclobutane pyrimidine dimer (CPD) levels and changes in accumulation of anthocyanins and UV‐absorbing compounds in leaves with leaf age, and (2) the susceptibility to CPD induction by UV‐B radiation and the ability to photorepair CPD were examined. Although supplementary UV‐B elevated the steady state of CPD levels in leaves in both strains, the level in the leaf of the NIL was higher than that in T‐65 at any time. The susceptibility to CPD induction by short‐term (challenge) UV‐B exposure was lower in the NIL than in T‐65. On the other hand, the CPD photorepair was also lower in the leaves of the NIL than in those of T‐65. The decrease in CPD‐photorepair in the NIL was due to a lowering of the leaf‐penetrating blue/UV‐A radiation, which is effective for photoreactivation by photolyase, by anthocyanins. Thus, accumulation of anthocyanins and UV‐absorbing compounds did not effectively function as screening against damage caused by elevated UV‐B radiation in the NIL, and the retardation of growth in the NIL resulted from its lower ability to photorepair CPD by higher amounts of anthocyanins.     

Investigating UV screening in leaves by two different types of portable UV fluorimeters reveals in vivo screening by anthocyanins and carotenoids

Two portable instruments, designed to evaluate epidermal UV screening in leaves, were compared: the Dualex and the UV-A-PAM fluorimeter. Both instruments excite chlorophyll fluorescence at the same UV wavelengths but reference excitation is in the red and the blue spectral range in the former and the latter fluorimeter, respectively. When analyzing green leaves, general agreement of the data is obtained with the two instruments. In the presence of anthocyanins, the UV-A-PAM fluorimeter provided higher estimates for epidermal UV transmittance than the Dualex fluorimeter, which was attributed to absorption of blue excitation light by anthocyanins. By comparing data from the instruments, anthocyanin-dependent transmittance of 50% was determined in abaxial sides of some autumn leaves, and also in abaxial sides of tropical shade plants. Further, with leaves of chlorophyll b-less mutants of H. vulgare, unusually high epidermal UV transmittance was detected but this was attributed to the lack of chlorophyll b absorption and, in addition, to absorption of blue radiation by xanthophylls which are not functionally connected to photosystems.     

The fascinating diatom frustule—can it play a role for attenuation of UV radiation?

Diatoms are ubiquitous organisms in aquatic environments and are estimated to be responsible for 20–25 % of the total global primary production. A unique feature of diatoms is the silica wall, called the frustule. The frustule is characterized by species-specific intricate nanopatterning in the same size range as wavelengths of visible and ultraviolet (UV) light. This has prompted research into the possible role of the frustule in mediating light for the diatoms’ photosynthesis as well as into possible photonic applications of the diatom frustule. One of the possible biological roles, as well as area of potential application, is UV protection. In this review, we explore the possible adaptive value of the silica frustule with focus on research on the effect of UV radiation on diatoms. We also explore the possible effect of the frustules on UV radiation, from a theoretical, biological, and applied perspective, including recent experimental data on UV transmission of diatom frustules.     

Flavonoids can protect maize DNA from the induction of ultraviolet radiation damage.

Diverse flavonoid compounds are widely distributed in angiosperm families. Flavonoids absorb radiation in the ultraviolet (UV) region of the spectrum, and it has been proposed that these compounds function as UV filters. We demonstrate that the DNA in Zea mays plants that contain flavonoids (primarily anthocyanins) is protected from the induction of damage caused by UV radiation relative to the DNA in plants that are genetically deficient in these compounds. DNA damage was measured with a sensitive and simple assay using individual monoclonal antibodies, one specific for cyclobutane pyrimidine dimer damage and the other specific for pyrimidine(6,4)pyrimidone damage.     

Immobile and tough versus mobile and weak: effects of ultraviolet B radiation on eggs and larvae of Manduca sexta

Although indirect effects of solar ultraviolet (UV) radiation on insects are well known (e.g. UV radiation can modify plant chemistry), direct effects of solar radiation on insects have received little attention. Radiation in the UVB range (300–320 nm) is damaging because it is absorbed directly by proteins and DNA. UVB should be toughest on immobile or small life stages, such as eggs or early larval instars. In the present study, the effects of UVB radiation on eggs and larvae of the tobacco hornworm Manduca sexta L. (Lepidoptera: Sphingidae) are examined. The present study aimed to address: what natural levels of UV do they experience; how does UVB affect the performance of eggs; and how does it affect the performance of larvae? In addition, do M. sexta larvae use behaviour to avoid UVB exposure and, consequently, are they physiologically less robust to UVB? In these experiments, eggs and late larval instars of M. sexta are found to be robust to natural levels of UV radiation. By contrast, young larvae are not only more susceptible to damage from UVB, but also they use behavioural means to avoid it. The strategy of using behaviour may relax selection pressures on morphological and physiological mechanisms for preventing (or recovering from) damage by environmental UV radiation.     

Photomorphogenic responses to UV radiation IV: a comparative study of UVB effects on growth and pigment accumulation in etiolated and de-etiolated hypocotyls of wild-type and aurea mutant of tomato Lycopersicon esculentum Mill)

ABSTRACT

The question of how de-etiolation of tomato seedling under continuous monochromatic yellow light exerts an influence on UV radiation-induced responses has been studied. Hypocotyl extension and the accumulation of anthocyanins and UV-absorbing compounds was compared in the aurea mutant of tomato and its isogenic wild type. The data of the present paper provide evidence that, during de-etiolation of tomato seedlings, yellow light exerts its effects over seedlings responsiveness to subsequent UV irradiation through several mechanisms: 1) a significant enhancement of shorter UVB wavelength efficiency in both the genotypes; 2) the abolition of UVA -blue light-induced accumulation of UV-absorbing compounds that does not involve pnyA; 3) the disappearance of UVA-blue light-induced hypocotyl growth inhibition that does not involve phyA; 4) higher anthocyanin accumulation rate in response to UV radiation mediated by phyA. Yellow light-induced growth inhibition and accumulation of UV-absorbing compound both mediated by phyA and present only in wild type tomato, appear to be completely separate from the action of UV radiation on the same responses.     

Differences in growth and physiological traits of two poplars originating from different altitudes as affected by UV‐B radiation and nutrient availability

Cuttings of Populus kangdingensis and Populus cathayana originating from altitudes of 3500 and 1500 m in southwestern China, respectively, were grown for one growing season in the field under ambient or ambient plus supplemental ultraviolet‐B (UV‐B) radiation with two levels of nutrients. In both species, enhanced UV‐B radiation significantly increased UV‐B absorbing compounds and guaiacol peroxidase (GPX) activity, while no significant effects were observed in photosynthetic pigments and proline content. On the other hand, cuttings grown with high‐nutrient availability had larger leaf area, higher total biomass and GPX activity as well as higher water use efficiency (WUE) (as measured by stable carbon isotope composition, δ¹³C) when compared with low‐nutrient conditions, while UV‐B absorbing compounds and ascorbic acid (AsA) content significantly decreased. Differences in responses to enhanced UV‐B radiation and nutrient availability were observed between the two species. Nutrient‐induced increases in chlorophyll a, chlorophyll b and total chlorophyll as well as in carotenoids were greater in P. kangdingensis than in P. cathayana. In P. cathayana, enhanced UV‐B radiation significantly decreased leaf area and total biomass, while it significantly increased WUE and ascorbate peroxidase (APX). In contrast, such changes were not observed in P. kangdingensis. In addition, the effects of enhanced UV‐B radiation on leaf area, total biomass and UV‐B absorbing compounds were closely related to the nutrient status. Our results indicated that P. kangdingensis, which originates from the altitude of 3500 m and is apparently adapted to low‐nutrient and high‐UV‐B habitats, exhibits better tolerance to enhanced UV‐B radiation and greater growth under low‐nutrient availability than does P. cathayana originating from the altitude of 1500 m.     

Inhibitory effects of ambient levels of solar UV‐A and UV‐B radiation on growth of cv. New Red Fire lettuce

The influence of solar UV‐A and UV‐B radiation at Beltsville, MD, USA, on growth of Lactuca sativa L. (cv. New Red Fire lettuce) was examined during early summer of 1996 and 1997. Plants were grown from seed in plastic window boxes covered with Llumar to exclude UV‐A and UV‐B, polyester to exclude UV‐B, or tefzel (1996) or teflon (1997) to transmit UV‐A and UV‐B radiation. After 31–34 days, plants grown in the absence of solar UV‐B radiation (polyester) had 63 and 57% greater fresh weight and dry weight of tops, respectively, and 57, 72 and 47% greater dry weight of leaves, stems and roots, respectively, as compared to those grown under ambient UV‐B (tefzel or teflon). Plants protected from UV‐A radiation as well (Llumar) showed an additional 43 and 35% increase, respectively, in fresh and dry weight of tops and a 33 and 33% increase, respectively, in dry weight of leaves and stems, but no difference in root biomass over those grown under polyester. Excluding ambient UV‐B (polyester) significantly reduced the UV absorbance of leaf extracts at 270, 300 and 330 nm (presumptive flavonoids) and the concentration of anthocyanins at 550 nm as compared to those of leaf extracts from plants grown under ambient UV‐A and UV‐B. Additional removal of ambient UV‐A (Llumar) reduced the concentration of anthocyanins, but had no further effect on UV absorbance at 270, 300 or 330 nm. These findings provide evidence that UV‐B radiation is more important than UV‐A radiation for flavonoid induction in this red‐pigmented lettuce cultivar. Although previous workers have obtained decreases in lettuce yield under enhanced UV‐B, this is the first evidence for inhibitory effects of solar UV‐A and UV‐B radiation on lettuce growth.     

UV-A screening in plants determined using a new portable fluorimeter

UV screening by plant surfaces can be determined by exposing plant organs to UV radiation and measuring the chlorophyll (Chl) fluorescence elicited. From this fluorescence, the UV transmittance can be derived: the more intense the screening the lower the reporter Chl fluorescence and the lower the UV transmittance. The relationships between UV screening at 375 nm, as determined in the field by a portable UV-A-PAM fluorimeter, and UV screening at 314 and 360 nm, measured in the laboratory with the non-portable XE-PAM fluorimeter, were investigated in leaves of grapevine (Vitis vinifera L. cv. Bacchus) and barley (Hordeum vulgare cv. Ricarda), as well as in white grape berries. With leaves, linear trends were observed between XE-PAM measurements at 314 nm and UV-A-PAM measurements at 375 nm but the relationship between transmittance at 360 and 375 nm in barley was curved: a simple model calculation suggests that this curvi-linearity arises from particularly weak absorbance of barley flavonoids at 375 nm relative to absorbance at 360 nm. Transmittance values at 314 nm plotted against 375 nm yielded a much smaller slope in grapevine leaves than in barley leaves, which was attributed to screening in the short-wavelength UV by hydroxycinnamic acids in the former but not in the latter species. With grape berries, a poor correlation was detected between transmittances at 314 and 375 nm which might arise from high scattering of UV radiation at the berry surface. Such artefacts appear to be confined to the UV-B region, as berry transmittance at 360 nm correlated very well with that at 375 nm. Thus, assessment of UV screening in the field at short UV wavelengths using 375 nm readings from a UV-A-PAM fluorimeter is possible provided that information is available on the relationship between the transmittance at the UV wavelength of interest and at 375 nm for the sample tissue being investigated.     

The influence of ultraviolet radiation on growth,photosynthesis and phenolic levels of green and red lettuce: potential for exploiting effects of ultraviolet radiation in a production system

Studies have shown that natural ultraviolet (UV) radiation increases secondary products such as phenolics but can significantly inhibit biomass accumulation in lettuce plants. In the work presented here, the effect of UV radiation on phenolic concentration and biomass accumulation was assessed in relation to photosynthetic performance in red and green lettuce types. Lettuce plants in polythene clad tunnels were exposed to either ambient (UV transparent film) or UV-free conditions (UV blocking film). The study tested whether growth reduction in lettuce plants exposed to natural UV radiation is because of inhibition of photosynthesis by direct damage to the photosynthetic apparatus or by internal shading by anthocyanins. Ambient levels of UV radiation did not limit the efficiency of photosynthesis suggesting that phenolic compounds may effectively protect the photosynthetic apparatus. Growth inhibition does, however, occur in red lettuce and could be explained by the high metabolic cost of phenolic compounds for UV protection. From a commercial perspective, UV transparent and UV blocking films offer opportunities because, in combination, they could increase plant quality as well as productivity. Growing plants continuously under a UV blocking film, and then 6 days before the final harvest transferring them to a UV transparent film, showed that high yields and high phytochemical content can be achieved complementarily.     

Asymmetric somatic cell hybridization in plants

Summary An investigation into the possible application of UV radiation as a pretreatment for the donor cells in asymmetric plant cell hybridization protocols has been carried out. A comparison was made between the effects of UV doses in the range 700–4200 J/m² and those of ⁶⁰Co gamma radiation over the range 0.15–1 kGy on Beta vulgaris suspension cell protoplasts. The investigation had two aspects. Firstly, alterations to cell physiology (cell wall resynthesis, viability, division and colony formation) in irradiated protoplasts were examined during a 4-week culture period. Results have indicated that a dose of 700 J/m² UV is necessary to prevent further cell division and colony formation in these cells. A dose of 0.15 kGy gamma radiation generally prevented colony formation, although some early cell division did occur (as was also observed even after 0.45 kGy had been applied). Membrane integrity, as measured after 6 days, using fluorescein diacetate staining, was not affected by either treatment within the dose ranges applied. Secondly, denaturing (alkaline) gel electrophoresis, in association with a pulsed field gel DNA preparation technique, was used to determine the degree of in vivo DNA damage following the radiation treatments. After UV radiation, considerable fragmentation of the DNA was observed, the extent of which was dose-dependent. Gamma radiation, however, appeared to result in fewer DNA lesions, with only the 1 kGy treatment revealing a pattern significantly altered from that of the control. These results augur well for the potential use of UV radiation in asymmetric fusion experiments.     

An enhancing effect of visible light and UV radiation on phenolic compounds and various antioxidants in broad bean seedlings

Exposure of dark- or ambient visible light-grown broad bean seedlings to low (LL) and high (HL) visible light intensities, UV-A or UV-C, either alone or in combination, induced significant increases in total phenolic compounds as well as in anthocyanins content, throughout the germination period, as compared with the respective levels in control seedlings. In general, as compared with control levels, exposure of both dark- or light-grown broad bean seedlings to LL, HL, UV-A or UV-C, induced significant increases in the contents of non-enzymatic antioxidants (total ascorbate; ASA-DASA and total glutathione; GSSG-GSH) and enzymatic antioxidant activities (superoxide dismutase; SOD, catalase; CAT, ascorbate peroxidase; APO and glutathione reductase; GR). The obtained results are discussed in relation to induced mechanisms of protection and repair from the inevitable exposure to damaging visible light and UV radiation.Key words: Vicia faba, anthocyanins, phenolic compounds, total ascorbate, total glutathione, SOD, APO, CAT, GR     

Effects of UV radiation on five marine microphytobenthic Wadden sea diatoms isolated from the Solthörn tidal flat (Lower Saxony,southern North Sea) – Part I: growth and antioxidative defence strategies

The unconsolidated sediment of intertidal mudflats constitutes a highly unstable environment, due to continuously changing water levels and currents as well as temporary exposure to the air. Therefore, diatoms inhabiting marine intertidal areas are subjected to strongly changing surface light and UV intensities due to exposure at low tide. Five marine intertidal diatoms (Achnanthes exigua, Cocconeis peltoides, Diploneis littoralis, Navicula digitoradiata and Amphora exigua) were isolated from the Solthörn tidal flat (Lower Saxony, southern North Sea). Semi-continuous cultures were used to determine the effect of UV radiation (photosynthetically active radiation only [PAR], PAR+UV-B, PAR+UV-A, PAR+UV-B+UV-A) during short- and long-term exposure (6 h or 30 days). Growth rates, chlorophyll a (chl a), antioxidant capacities, accumulation of phenolic compounds (e.g. flavonoids) and DMSP, and activities of antioxidant enzymes (superoxide dismutase, ascorbate peroxidase, monodehydroascorbate reductase and glutathione reductase) were assessed. UV-A had only minor effects on cells, while growth rate, chl a content and protein content were significantly reduced after long-term UV-B exposure. Achnanthes exigua extracts showed the highest antioxidant capacity. The highest activity of SOD, APX and MDHAR was found under long-term combined UV exposure (PAR+UV-B+UV-A). Overall, the antioxidative defence of the five isolates was stimulated during exposure to UV radiation, as may be found during emersion. Emersion induces oxidative stress and, as a result, growth of the five diatom taxa was inhibited to suit changing environmental conditions. All five taxa tested in the present study showed species-specific acclimatization potentials, providing possible explanations for variability in population, species composition and ecosystem structures in the face of climatic variations.     

Lichens as survivors in space and on Mars

Many lichens are able to live and photosynthesize under harsh conditions, characterized by low temperatures, aridity and high UV radiation fluxes. Some lichen species are even able to survive simulated and real space conditions. Many tests after space exposure on the satellite FOTON M3 and on the International Space Station have shown their capacity to maintain physiological and photosynthetic activity, and their capacity to germinate and grow after being exposed to space parameters. Further tests using simulated Martian atmospheres, temperatures, humidity profiles and UV radiation spectra and fluxes have shown maintenance of photosynthetic activity of Xanthoria elegans. Results from space and Mars simulation experiments on lichens such as X. elegans are valuable for determining the habitability of a planet and for the search for possible life-supporting habitats on planets like Mars.     

pEffects of UV radiation on the ultrastructure of several red algae

The effect of ultraviolet (UV) radiation on the ultrastructure of four red algae, the endemic Antarctic Palmaria decipiens (Reinsch) Ricker and Phycodrys austrogeorgica Skottsberg, the Arctic‐cold temperate Palmaria palmata (Linnaeus) O. Kuntze and the cosmopolitan Bangia atropurpurea (Roth) C. Agardh was studied. All four species showed a formation of ‘inside‐out’ vesicles from the chloroplast thylakoids upon exposure to artificial UV‐radiation. In P. decipiens, most vesicles were developed after 8 h and in P. palmata, after 48 h of UV exposure. In B. atropurpurea, vesi‐culation of thylakoids was observed after 72 h of UV irradiation. In Ph. austrogeorgica, the chloroplast envelope and thylakoid membranes were damaged and the phycobilisomes became detached from the thylakoids after 12 h of UV exposure. Ultraviolet‐induced changes in the membrane structure of mitochondria were observed in P. decipiens and P. palmata. However, in P. decipiens they were reversible as was the damage in chloroplast fine structure after 12 h of UV treatment. Protein crystals in Ph. austrogeorgica showed degradation after exposure to UV radiation. Different methods of fixation and embedding macroalgal material are discussed. These findings give insight into the fine structural changes which occur during and after UV exposure and indicate a relationship between the species dependent sensitivity to UV‐exposure and the depth distribution of the different species.     

Photoinhibition by visible and ultraviolet radiation in the red macroalga Porphyra umbilicalis grown in the laboratory

Photosynthetic oxygen production and PAM fluorescence measurements were used to follow photoinhibition in the red macroalga Porphyra umbilicalis. Exposure to simulated solar radiation caused inhibition of the effective photosynthetic quantum yield from which the thalli partially recovered in the shade in subsequent hours. There were no significant differences between samples exposed to unfiltered radiation and those exposed to radiation from which increasing portions of UV radiation had been removed indicating that the thalli are well adapted to current levels of solar PAR and UV radiation. This notion was supported by the finding of high concentrations of UV screening pigments which were even enhanced by exposure to increased UV radiation. However, when exposed to (only) UV radiation about 50% higher than that encountered by the organisms in their natural habitat, the photosynthetic yield decreased slowly and did not show any recovery even when the degree of inhibition did not exceed 10%.     

LACK OF DEFENSE OR PHLOROTANNIN INDUCTION BY UV RADIATION OR MESOGRAZERS IN DESMARESTIA ANCEPS AND D. MENZIESII (PHAEOPHYCEAE)1

Phlorotannins are polyphenoloic metabolites occurring only in the Phaeophyceae that have numerous putative primary roles (e.g. cell‐wall construction and storage) as well as secondary metabolic roles, which include herbivore feeding deterrence and protection from UV radiation. The proposed role of phlorotannins in the defense against UV radiation is of particular importance in the Antarctic due to depletion of the stratospheric ozone layer in that area. Several studies of brown algae have found evidence of an induction response (the production of defensive metabolites, including phlorotannins) after grazing by various mesograzers, after simulated grazing/wounding, and after exposure to increases in UV radiation. This study aimed to determine if phlorotannin production or other defenses in two dominant, endemic Antarctic species (Desmarestia menziesii Montagne and Desmarestia anceps J. Agardh) could be induced by an increase in exposure to UV radiation or by natural and artificial grazing. An in situ experiment failed to detect any effect of UV radiation on phlorotannin concentrations in either species or on subsequent palatability in feeding bioassays. A laboratory‐based experiment did not detect any effect of mesoherbivore grazing or simulated grazing (wounding) on palatability or the concentration of phlorotannins in D. menziesii. Instead, phlorotannin concentrations increased in all treatments in both experiments, consistent with an increase in overall resource availability due to an increase in available PAR compared with the in situ irradiance at the algal collection sites.     

Chemical profiling of mycosporine‐like amino acids in twenty‐three red algal species

Rhodophyta produce a variety of chemically different mycosporine‐like amino acids (MAAs), compounds that are known as some of the strongest ultraviolet (UV) absorbing molecules in nature. Accordingly, they primarily act as photoprotectants against harmful levels of solar ultraviolet radiation in the UV‐A and UV‐B range. In order to get a deeper understanding of the chemical diversity of MAAs in red algae, pure standards of eleven mycosporine‐like amino acids were isolated from three different species (Agarophyton chilense, Pyropia plicata and Champia novae‐zelandiae) using various chromatographic methods. Their structures were confirmed by nuclear magnetic resonance and mass spectrometry. Four out of the eleven MAAs are reported for the first time in algae. In addition, a new high‐performance liquid chromatography method was developed for the separation of all isolated MAAs and successfully applied for the analysis of twenty‐three red algal species of marine origin. All of them contained MAAs, the most abundant compounds were shinorine, palythine, asterina‐330 and porphyra‐334. For some samples, the direct assignment of MAAs based on their UV spectra was not possible; therefore, the target analytes were enriched by a simple concentration step, followed by liquid chromatography‐mass spectrometry analysis of the extracts. This approach enabled a deeper insight into the MAA pattern of red algae, indicating that not only the four dominant ones are synthesized but also many others, which were often described as unknown compounds in previous studies.