Effect of Enhanced UV-B Radiation and Low-Energy N+ Ion Beam Radiation on the Response of Photosynthesis, Antioxidant Enzymes, and Lipid Peroxidation in Rice (Oryza sativa) Seedlings

To understand the effect of enhanced UV-B radiation and low-energy N⁺ ion beam radiation on the response of photosynthesis, antioxidant enzymes, and lipid peroxidation in rice seedlings, Oryza sativa was exposed to three different doses of low-energy N⁺ ion beam and enhanced UV-B alone and in combination. Enhanced UV-B caused a marked decline in some photosynthetic parameters (net photosynthetic rate, transpiration rate, and stomatal conductance) and photosynthetic pigments, whereas it induced an increase in hydrogen peroxide (H₂O₂) accumulation, the rate of superoxide radical production, and the content of malondialdehyde (MDA). Enhanced UV-B also induced an increase in the activity of antioxidant enzymes (superoxide dismutase [SOD], peroxidase (POD), and catalase [CAT]) and some nonenzymatic antioxidants such as proline. Under the combined treatment of enhanced UV-B and low-energy N⁺ ion beam at the dose of 3.0?×?10¹⁷ N⁺ cm^?2, the activity of antioxidant compounds (SOD, POD, CAT, proline, and glutathione), photosynthetic pigments, and some photosynthetic parameters (net photosynthetic rate, transpiration rate, and stomatal conductance) increased significantly; however, the MDA content, H₂O₂ accumulation, and rate of superoxide radical production showed a remarkable decrease compared with the enhanced UV-B treatment alone. These results implied that the appropriate dose of low-energy N⁺ ion beam treatment may alleviate the damage caused by the enhanced UV-B radiation on rice.     

Effect of ultraviolet-B radiation on salt marsh vegetation: Trends of the genus Salicornia along the Americas

The effects of natural UV-B radiation on growth, photosynthetic and photoprotective pigment composition of different Salicornia species were analyzed in salt marshes at three different sites along the Americas (Puerto Rico, southern Brazil and Patagonia, Argentina). Plants were exposed to different levels of UV-B radiation for 1-2 years in situ as well as in outdoor garden UV-B exclusion experiments. Different UV-B levels were obtained by covering plants with UV-B opaque (blocked 93-100% of ambient UV-B) and UV-B attenuating (near-ambient) filters (reduced 20-25% of UV-B). Unfiltered plants were exposed to natural irradiance. UV-B filters had significant effects on temperature and photosynthetic pigments (due to changes in PAR; 400-700 nm). The growth of Salicornia species was inhibited after 35 to 88 days of exposure to mean UV-B radiation dosages between 3.6 and 4.1 kJ m(-2) day(-1). The highest number of branches on the main shoot (S. bigelovii and S. gaudichaudiana) and longest total length of the branches (S. gaudichaudiana) were observed in the UV-B opaque treatment. Salicornia species responded to increasing levels of UV-B radiation by increasing the amount of UV-B absorbing pigments up to 330%. Chromatographic analyses of seedlings and adult S. bigelovii plants found seven different UV-B absorbing flavonoids that are likely to serve as UV-B filtering pigments. No evidence of differential sensitivity or resilience to UV-B radiation was found between Salicornia species from low-mid latitudes and a previously published study of a high-latitude population.     

EFFECTIVENESS OF UV-B RADIATION ON THE GROWTH AND PHYSIOLOGY OF FIELD-GROWN SOYBEAN MODIFIED BY WATER STRESS

Abstract— Soybeans [ Glycine max (L) Merr. cv Essex] were grown in field plots during May-October 1985 under ambient and an enhanced level of ultraviolet-B (UV-B) radiation (supplemental daily dose: 5.1 effective kJ m^-2). They were either subjected to water stress or supplementally irrigated, resulting in a 2.0 MPa lower soil water potential in stressed plots. Increased levels of UV-B radiation reduced leaf area, total plant dry weight and net photosynthesis under well-watered conditions, but no significant UV-B effects were detected in plants concurrently subjected to water stress. The insensitivity of growth and net photosynthesis to UV-B radiation in water-stressed plants may be related to anatomical and biochemical changes induced by water stress. These include an increase in the concentration of UV absorbing compounds in leaf tissues and leaf thickening.     

How do UV Photomorphogenic Responses Confer Water Stress Tolerance?¶†

Although ultraviolet-B (UV-B) radiation is potentially harmful, it is an important component of terrestrial radiation to which plants have been exposed since invading land. Since then, plants have evolved mechanisms to avoid and repair UV radiation damage; therefore, it is not surprising that photomorphogenic responses to UV-B are often assumed to be adaptations to harmful radiation. This presupposes that the function of the observed responses is to prevent UV damage. It has been hypothesized that, as with blue light, UV-B provides a signal important for normal plant development and might be perceived within developing plants through nondestructive processes, perhaps through UV-specific signal perception mechanisms. UV signal perception can lead to photomorphogenic responses that may confer adaptive advantages under conditions associated with high-light environments, such as water stress. Plant responses to UV radiation in this regard include changes in leaf area, leaf thickness, stomatal density, photosynthetic pigment production and altered stem elongation and branching patterns. Such responses may lead to altered transpiration rates and water-use efficiencies. For example, we found that the cumulative effect of ambient UV-B radiation upon stomatal density and conductance can lead to altered water-use efficiencies. In field settings, UV might more properly be viewed as a photomorphogenic signal than as a stressor. Hence, it might be insufficient to attempt to fully evaluate the adaptive roles of plant responses to UV-B cues upon stress tolerance by the simultaneous application of UV and drought stress during development. We propose that rather than examining a plant's response to combinations of stressors one might also examine how a plant's response to UV induces tolerance to subsequently applied stresses.     

Flavonoid concentrations in three grass species and a sedge grown in the field and under controlled environment conditions in response to enhanced UV-B radiation

An investigation was carried out to find whether enhanced levels of UV-B radiation induce increased concentrations of flavonoids in the leaves of the grass species Deschampsia antarctica, Deschampsia borealis and Calamagrostis epigeios and the sedge Carex arenaria. Whether the enhanced levels of UV-B influenced the proportions of the various flavonoids in the leaves was also studied. Increased flavonoid concentrations would improve the UV-B shielding of UV-B susceptible tissues. Using HPLC analysis the flavonoids orientin and luteolin were identified in D. antarctica, orientin in D. borealis and tricin in C. arenaria. Neither flavonoid concentrations nor the proportion of the various flavonoids in climate room grown D. antarctica and D. borealis plants differed between individuals grown under 0, ambient or elevated UV-B levels. After 12 weeks of growth biomass production and shoot-to-root ratios of D. antarctica were not affected by elevated UV-B radiation. Greenhouse grown C. epigeios plants contained higher concentrations and different proportions of flavonoids grown under elevated levels of UV-B than when grown under ambient or 0 UV-B. In C. epigeios plants grown in their natural habitat in the field under ambient or elevated levels of UV-B, flavonoid concentrations and proportions were the same in plants from both treatments. In the leaves of the sedge C. arenaria grown in a greenhouse flavonoid concentrations and proportions were not affected by UV-B radiation. Leaves were harvested four times during the growing season from C. arenaria plants grown in their natural habitat in the field under ambient or elevated levels of UV-B. Leaves harvested in January contained higher concentrations of flavonoids when grown under elevated UV-B than when grown under ambient UV-B radiation. In leaves harvested in May, September and December flavonoid concentrations were the same in plants grown under ambient or elevated UV-B. The proportion of the different flavonoids was the same for both treatments in all months. These results indicate that constitutive levels of flavonoids in these grass and sedge species are adequately high to protect them against ambient and elevated levels of solar UV-B radiation.     

INFLUENCE OF PHOTON FLUX DENSITY IN THE 400–700 nm WAVEBAND ON INHIBITION OF PHOTOSYNTHESIS BY UV-B (280–320 nm) IRRADIATION IN SOYBEAN LEAVES: SEPARATION OF INDIRECT AND IMMEDIATE EFFECTS

Abstract— Visible radiation can substantially influence the degree to which plant photosynthesis is inhibited by UV-B radiation. This study was designed to separate the immediate effects of visible radiation on UV-B photosynthetic inhibition from the indirect influence of visible irradiation on morphological and physiological properties of leaves during leaf development. Soybean plants were pretreated in growth chambers with either high or low visible irradiance (750 and 70 μmol m^-2s^-1 quantum flux in the 400–700 nm waveband, respectively) during the development of leaves used subsequently for UV irradiation. Test leaves still attached to the plant were exposed to 5 h of polychromatic UV-B irradiation and the photosynthetic capacity (net CO₂ exchange) was determined before and after the UV irradiation. During the UV irradiation, plants from both pretreatment groups received either high or low visible flux. Development of leaves in the high visible flux pretreatment conditions resulted in thicker leaves, higher chlorophyll a/b ratios, more UV-absorbing pigments, and reduced sensitivity to the UV-B irradiation. However, higher visible flux during the UV-B irradiation resulted in greater depression of photosynthesis by the UV-B irradiation. The relative magnitude of photosynthetic depression under these treatment combinations was the same when photosynthesis was measured under either light-limited or light-saturated conditions.     

The influence of enhanced UV-B radiation on Batrachium trichophyllum and Potamogeton alpinus -- aquatic macrophytes with amphibious character

The responses of two amphibious species, Batrachium trichophyllum and Potamogeton alpinus to different UV-B environments were studied. Plant material from natural environments, as well as from outdoor treatments was examined. In long-term outdoor experiments plants were grown under three different levels of UV-B radiation: reduced and ambient UV-B levels, and a UV-B level simulating 17% ozone depletion. The following parameters were monitored: contents of total methanol soluble UV-absorbing compounds and chlorophyll a, terminal electron transport system (ETS) activity and optimal and effective quantum yield of photosystem II. No effect of the different UV-B levels on the measured parameters was observed. The amount of UV-B absorbing compounds seems to be saturated, since no differences were observed between treatments and no increase was found in peak season, when natural UV-B levels were the highest. Physiological measurements revealed no harmful effects; neither on potential and actual photochemical efficiency, nor on terminal ETS activity. The contents of UV-B absorbing compounds were examined also in plant material sampled in low and high altitude environments during the growth season. Both species exhibited no seasonal dynamics of production of UV-absorbing compounds. The contents were variable and showed no significant differences between high and low altitude populations.     

Leaves of Citrus aurantifolia exhibit a different sensibility to solar UV-B radiation according to development stage in relation to photosynthetic pigments and UV-B absorbing compounds production

Plants of Citrus aurantifolia grown in a greenhouse without solar UV radiation (UVR) were transferred outdoors to evaluate the effect of solar UV-B radiation (UVBR, 280–315 nm) in prior-developed leaves, constituted by apical bud and those fully expanded before being taken outdoors, and post-developed leaves, formed by those expanded outdoors. Results demonstrated that over a 40 d outdoor period leaf chlorophyll content and distribution pattern were different with and without solar UVBR. Chlorophyll a, chlorophyll b and total chlorophyll contents in both treatments were higher in prior-developed leaves than in post-developed ones. However, highest values were observed in prior-developed leaves under solar UVBR, whereas in post-developed leaves an opposite trend was observed. Carotenoids content in prior-developed leaves was higher with solar UVBR, whereas in post-developed leaves there were no significant differences in both with and without solar UVBR. In addition, prior-developed leaves under solar UVBR accumulated flavonoids, but not anthocyanins. Growth parameters (e.g. DW, DW/FW ratio, LMA, plant height, length and width of foliar lamina) did not show significant differences between plants grown with and without solar UVBR. Thus, our results demonstrated that C. aurantifolia leaves exhibited a different sensibility to solar UVBR according to development stage in relation to photosynthetic pigments and UV-B absorbing compounds production. In addition, the solar UVBR was not necessary as inductor of photosynthetic protection mechanisms in a short-time growth period. On the other hand, our results also demonstrated that solar UVBR acted as an effective feeding deterrent against citrus leafminer.     

The role of UV-B radiation in aquatic and terrestrial ecosystems--an experimental and functional analysis of the evolution of UV-absorbing compounds

We analysed and compared the functioning of UV-B screening pigments in plants from marine, fresh water and terrestrial ecosystems, along the evolutionary line of cyanobacteria, unicellular algae, primitive multicellular algae, charophycean algae, lichens, mosses and higher plants, including amphibious macrophytes. Lichens were also included in the study. We were interested in the following key aspects: (a) does the water column function effectively as an 'external UV-B filter'?; (b) do aquatic plants need less 'internal UV-B screening' than terrestrial plants?; (c) what role does UV screening play in protecting the various plant groups from UV-B damage, such as the formation of thymine dimers?; and (d) since early land 'plants' (such as the predecessors of present-day cyanobacteria, lichens and mosses) experienced higher UV-B fluxes than higher plants, which evolved later, are primitive aquatic and land organisms (cyanobacteria, algae, lichens, mosses) better adapted to present-day levels of UV-B than higher plants? Furthermore, polychromatic action spectra for the induction of UV screening pigments of aquatic organisms have been determined. This is relevant for translating 'physical' radiation measurements of solar UV-B into 'biological' and 'ecological' effects. From the action spectra, radiation amplification factors (RAFs) have been calculated. These action spectra allow us to determine any mitigating or antagonistic effects in the ecosystems and therefore qualify the damage prediction for the ecosystems under study. We summarize and discuss the main results based on three years of research of four European research groups. The central theme of the work was the investigation of the effectiveness of the various screening compounds from the different species studied in order to gain some perspective of the evolutionary adaptations from lower to higher plant forms. The induction of mycosporine-like amino acids (MAAs) was studied in the marine dinoflagellate Gyrodinium dorsum, the green algal species Prasiola stipitata and in the cyanobacterium Anabaena sp. While visible (400-700 nm) and long wavelength UV-A (315-400 nm) showed only a slight effect, MAAs were effectively induced by UV-B (280-315 nm). The growth of the lower land organisms studied, i.e. the lichens Cladina portentosa, Cladina foliacaea and Cladonia arbuscula, and the club moss Lycopodiumannotinum, was not significantly reduced when grown under elevated UV-B radiation (simulating 15% ozone depletion). The growth in length of the moss Tortula ruralis was reduced under elevated UV-B. Of the aquatic plants investigated the charophytes Chara aspera showed decreased longitudinal growth under elevated UV-B. In the 'aquatic higher plants' studied, Ceratophyllum demersum, Batrachium trichophyllum and Potamogeton alpinus, there was no such depressed growth with enhanced UV-B. In Chara aspera, neither MAAs nor flavonoids could be detected. Of the terrestrial higher plants studied, Fagopyrum esculentum, Deschampsia antarctica, Vicia faba, Calamagrostis epigejos and Carex arenaria, the growth of the first species was depressed with enhanced UV-B, in the second species length growth was decreased, but the shoot number was increased, and in the latter two species of a dune grassland there was no reduced growth with enhanced UV-B. In the dune grassland species studied outdoors, at least five different flavonoids appeared in shoot tissue. Some of the flavonoids in the monocot species, which were identified and quantified with HPLC, included orientin, luteolin, tricin and apigenin. A greenhouse study with Vicia faba showed that two flavonoids (aglycones) respond particularly to enhanced UV-B. Of these, quercetin is UV-B inducible and mainly located in epidermal cells, while kaempferol occurs constitutively. In addition to its UV-screening function, quercetin may also act as an antioxidant. Polychromatic action spectra were determined for induction of the UV-absorbing pigments in three photosynthetic organisms, representing very different taxonomic groups and different habitats. In ultraviolet photobiology, action spectra mainly serve two purposes: (1) identification of the molecular species involved in light absorption; and (2) calculation of radiation amplification factors for assessing the effect of ozone depletion. Radiation amplification factors (RAFs) were calculated from the action spectra. In a somewhat simplified way, RAF can be defined as the percent increase of radiation damage for a 1% depletion of the ozone layer. Central European summer conditions were used in the calculations, but it has been shown that RAF values are not critically dependent on latitude or season. If only the ultraviolet spectral region is considered, the RAF values obtained are 0.7 for the green alga Prasiola stipitata, 0.4 for the dinoflagellate Gyrodinium dorsum, and 1.0 for the cyanobacterium Anabaena sp. In the case of P. stipitata, however, the effect of visible light (PAR, photosynthetically active radiation, 400-700 nm) is sufficient to lower the RAF to about 0.4, while the PAR effect for G. dorsum is negligible. RAFs for some damage processes, such as for DNA damage (RAF=2.1 if protective effects or photorepair are not considered [1]), are higher than those above. Our interpretation of this is that if the ozone layer is depleted, increased damaging radiation could overrule increased synthesis of protective pigments. In addition to investigating the functional effectiveness of the different screening compounds, direct UV effects on a number of key processes were also studied in order to gain further insight into the ability of the organisms to withstand enhanced UV-B radiation. To this end, the temperature-dependent repair of cyclobutane dimers (CPD) and (6-4) photoproducts induced by enhanced UV-B was studied in Nicotiana tabacum, and the UV-B induction of CPD was studied in the lichen Cladonia arbuscula. Also, photosynthesis and motility were monitored and the response related to the potential function of the screening compounds of the specific organism.     

UNDERSTANDING PHOTOSYNTHESIS, PIGMENT AND GROWTH RESPONSES INDUCED BY UV-B AND UV-A IRRADIANCES

Abstract—Plant response to UV-B (0.290–0.320 μm) irradiation in controlled environments has been difficult to assess, possibly because plants also respond to UV-A (0.320–0.400 μm) and visible radiation. Photosynthetic dysfunction is often reported, but effects on photosynthetic pigments have been equivocal. Because UV-A/blue radiation is involved in pigment synthesis, the experimental UV-A irradiation was controlled and this study was conducted under high ambient photosynthetic photon flux (mid-day PPF > 1400 pmol m ^–2 s^–1). Two biologically effective UV-B irradiances (10.7 and 14.1 kJ m^-2 day^-I) were utilized and the UV-A irradiances were matched in controls (˜5 and 9 kJ m^-2 day^-1). Normal and two mutant pigment isolines (chlorophyll-deficient, flavonoid-deficient) of soybean cultivar Clark were utilized for comparisons. Many pigmedgrowth variables exhibited a statistical interaction between spectral quality and quantity. UV-A/blue photoregulation was demonstrated in the UV-A controls. The pigmentlgrowth pattern observed at the lower UV-B irradiance was interpreted as a photosystem II response similar to shade adaptation, suggesting phytochrome involvement in UV-B irradiation responses. On the other hand, two variables most commonly observed to manifest UV-B-induced effects—decreased photosynthesis and increased leaf flavonoid content—exhibited no interactions due to UV exposure or spectral quality. In general, the observed response patterns indicated either moderation of UV-B-induced responses by UV-A/blue radiation, or coaction between them, and provides an explanation for the common failure to demonstrate fluence-related responses in UV-B experiments.     

Influence of solar ultraviolet-B radiation on Antarctic terrestrial plants: results from a 4-year field study.

We examined the influence of solar ultraviolet-B radiation (UV-B; 280-315 nm) on the performance of Antarctic vascular plants (Colobanthus quitensis and Deschampsia antarctica) by placing filters that either absorbed or transmitted most solar UV-B over tundra along the Antarctic Peninsula for four consecutive growing seasons. The difference in biologically effective UV-B levels between our treatments was 65%, which was similar to the enhancement in ambient UV-B levels that appeared attributable to ozone depletion during the first 2 months of the growing season (November and December) at our site (62%). In both species, exposure to UV-B reduced vegetative growth, primarily through slower leaf elongation rates that led to shorter fully expanded leaves. In C. quitensis, exposure to UV-B also led to reductions in leaf longevity, branch production, cushion diameter growth, aboveground biomass, and thickness of the non-green cushion base and litter layer. Exposure to UV-B accelerated the development of reproductive structures and increased the number of panicles (D. antarctica) and capsules (C. quitensis) that reached maturity per unit of ground surface area covered by mother plants. However, this effect was offset by a tendency for these panicles and capsules to produce fewer spikelets and seeds. Ultimately, UV-B exposure did not effect the numbers of spikelets or seeds produced per unit of ground surface area. While seeds from plants exposed to UV-B tended to be lighter, germination rates were similar between UV-B treatments. The relative reductions in leaf elongation rates in D. antarctica attributable to UV-B exposure increased from the first (23%) through the fourth (43%) growing season, and relative reductions in leaf longevity in C. quitensis tended to increase from the first (9%) through the fourth (19%) growing season, suggesting that UV-B growth responses tended to be cumulative over successive years.     

Laser pretreatment protects cells of broad bean from UV-B radiation damage

In order to determine the role of lasers in the stress resistance of broad bean (Vicia faba L.) to ultraviolet-B (UV-B) radiation, the embryos in seeds were exposed to He-Ne laser or CO2 laser radiation. Afterwards they were cultivated in Petri dishes in a constant temperature incubator until the lengths of epicotyls were nearly 3 cm. The epicotyls were then exposed to 1.02, 3.03 or 4.52 kJ m(-2) UV-B radiation, respectively, under 70 micromol m(-2) s(-1) photosynthetically active radiation (PAR) in a growth cabinet. Changes in the concentration of malondialdehyde (MDA), ascorbic acid (AsA) and UV-B absorbing compounds (absorbance at 300 nm) were measured to test the effects of laser pretreatment. The results showed that laser pretreatment of embryos enhanced UV-B stress resistance in the epicotyls of the broad bean by decreasing the MDA concentration and increasing the content of AsA and UV-B absorbing compounds. We suggest that those changes in MDA, AsA and UV-B absorbing compounds were responsible for the increase in stress resistance observed in the broad bean. This is the first investigation reporting the use of laser pretreatment to protect the cells of the broad bean from UV-B-induced damage.     

Photosynthetic,Biochemical and Secondary Metabolite Changes in a Medicinal Plant Chlorophytum borivillianum (Safed musli) against Low and High Doses of UV-B Radiation

Plants are inevitably grown in presence of sunlight, therefore bound to be exposed to natural UV-B radiation. Several studies have already been conducted with UV-B and medicinal plants and only few studies showed dose dependent variation. The present study aims to find out the variations and adaptation in Chlorophytum borivillianum under two different doses of UV-B radiation; ambient + low (3.2 kJm⁻² d⁻¹) and high (7.2 kJm⁻² d⁻¹) UV-B dose, denoted as LD and HD, respectively. Reduction in photosynthetic rate was higher at HD, while plants receiving LD displayed nonsignificant variation. During vegetative and reproductive stage, significant reduction (P ≤ 0.001) in stomatal conductance was obtained when exposed to HD-eUV-B. F_v/F_m showed more reductions in HD-eUV-B (12.6%) followed by LD-eUV-B (7.9%). Low and high doses of UV-B enhanced the anthocyanin content but the increase was significant in HD, indicates epidermal protection strategy by the plants. Under LD-eUV-B, the content of saponin, a major phytochemical constituent was enhanced by 26%. Phytochemical analysis of roots revealed reduction mostly in fatty acid components whereas the steroidal components (stigmasterol and sarsasapogenin) showed enhancement in response to LD. The study suggests the importance of LD-eUV-B in the stimulation of medicinal compounds in C. borivillianum.     

Alteration of foliar flavonoid chemistry induced by enhanced UV-B radiation in field-grown Pinus ponderosa, Quercus rubra and Pseudotsuga menziesii

Chromatographic analyses of foliage from several tree species illustrate the species-specific effects of UV-B radiation on both quantity and composition of foliar flavonoids. Pinus ponderosa, Quercus rubra and Pseudotsuga menziesii were field-grown under modulated ambient (1x) and enhanced (2x) biologically effective UV-B radiation. Foliage was harvested seasonally over a 3-year period, extracted, purified and the flavonoid fraction applied to a mu Bondapak/C(18) column HPLC system sampling at 254 nm. Total flavonoid concentrations in Quercus rubra foliage were more than twice (leaf area basis) that of the other species; Pseudotsuga menziesii foliage had intermediate levels and P. ponderosa had the lowest concentrations of total flavonoids. No statistically significant UV-B radiation-induced effects were found in total foliar flavonoid concentrations for any species; however, concentrations of specific compounds within each species exhibited significant treatment effects. Higher (but statistically insignificant) levels of flavonoids were induced by UV-B irradiation in 1- and 2-year-old P. ponderosa foliage. Total flavonoid concentrations in 2-year-old needles increased by 50% (1x ambient UV-B radiation) or 70% (2x ambient UV-B radiation) from that of 1-year-old tissue. Foliar flavonoids of Q. rubra under enhanced UV-B radiation tended to shift from early-eluting compounds to less polar flavonoids eluting later. There were no clear patterns of UV-B radiation effects on 1-year-old P. menziesii foliage. However, 2-year-old tissue had slightly higher foliar flavonoids under the 2x UV-B radiation treatment compared to ambient levels. Results suggest that enhanced UV-B radiation will alter foliar flavonoid composition and concentrations in forest tree species, which could impact tissue protection, and ultimately, competition, herbivory or litter decomposition.     

Epidermal lignin deposition in quinoa cotyledons in response to UV-B radiations

UV-B radiation (280-320 nm) is harmful to living organisms and has detrimental effects on plant growth, development and physiology. In this work we examined some mechanisms involved in plant responses to UV-B radiation. Seedlings of quinoa (Chenopodium quinoa Willd.) were exposed to variable numbers of UV-B radiation doses, and the effect on cotyledons was studied. We analyzed (1) cotyledons anatomy and chloroplasts ultrastructure; (2) peroxidase activity involved in the lignification processes; and (3) content of photosynthetic pigments, phenolic compounds and carbohydrates. Exposure to two UV-B doses induced an increase in the wall thickness of epidermal cells, which was associated with lignin deposition and higher activity of the peroxidase. The chloroplast ultrastructure showed an appearance typical of plants under shade conditions, likely in response to reduced light penetration into the mesophyll cells due to the screening effect of epidermal lignin deposition. Exposure to UV-B radiation also led to (1) enhancement in the level of phenolics, which may serve a protective function; (2) strong increase in the fructose content, a fact that might be related to higher requirement of erythrose-4P as a substrate for the synthesis of lignin and phenolics; and (3) reduction in the chlorophyll concentration, evidencing alteration in the photosynthetic system. We propose that the observed lignin deposition in epidermal tissues of quinoa is a resistance mechanism against UV-B radiation, which allows growing of this species in Andean highlands.     

AMELIORATION OF UV-B DAMAGE UNDER HIGH IRRADIANCE. I: ROLE OF PHOTOSYNTHESIS

Sensitivity to ultraviolet-B radiation (UV-B,280–315 nm) is generally reduced when background irradiance is high. We tested the involvement of photosynthesis in the amelioration of UV-B damage by treating plants at high PAR (photosynthetically-active radiation, 400–700 nm; 1000 μmol m-² s-¹) with supplemental UV-B at double ambient levels of biologically-effective radiation (18 kJ m-²d-¹) and either “ambient” (450 μmol mol-¹) or short term elevated (750 μmol mol-¹) CO₂ levels. Responses to UV-B were assessed by photosynthetic gas exchange, leaf expansion and production of UV-absorbing compounds (presumptive flavonoids) in cultivars of cucumber (Cucumis sativus L.) previously demonstrated to be relatively sensitive (cv. Poinsett) and insensitive (cv. Ashley) to UV-B. Except for marginal leaf interveinal chlorosis observed in Poinsett, both cultivars responded similarly. UV-B had little direct effect on leaf photosynthesis, but it did cause reductions in leaf area and corresponding increases in leaf dry matter per area. Increased CO, stimulated plant growth, counteracting the effect of UV-B on leaf growth and indicating an important role for photosynthesis. In contrast, the accumulation of UV-absorbing flavonoid compounds was enhanced by UV-B exposure but was not affected by COz enrichment.     

UV-B EFFECTS ON TERRESTRIAL PLANTS

The potential impacts of an increase in solar UV-B radiation reaching the Earth's surface due to stratospheric ozone depletion have been investigated by several research groups during the last 15 years. Much of this research has centered on the effects of plant growth and physiology under artificial UV-B irradiation supplied to plants in growth chambers or greenhouses. Since these artificial sources do not precisely match the solar spectrum and due to the wavelength dependency of photobiol-ogical processes, weighting functions, based on action spectra for specific responses, have been developed to assess the biological effectiveness of the irradiation sources and of predicted ozone depletion. Recent experiments have also utilized artificially produced ozone cuvettes to filter natural solar radiation and simulate an environment of reduced UV-B for comparative purposes. Overall, the effectiveness of UV-B varies both among species and among cultivars of a given species. Sensitive plants often exhibit reduced growth (plant height, dry weight, leaf area, etc.), photosynthetic activity and flowering. Competitive interactions may also be altered indirectly by differential growth responses. Photosynthetic activity may be reduced by direct effects on photosynthetic enzymes, metabolic pathways or indirectly through effects on photosynthetic pigments or stomatal function. The fluence response of these changes has yet to be clearly demonstrated in most cases. Plants sensitive to UV-B may also respond by accumulating UV-absorbing compounds in their outer tissue layers, which presumably protect sensitive targets from UV damage. Several key enzymes in the biosynthetic pathways of these compounds have been shown to be specifically induced by UV-B irradiation. Few studies have documented the effects of UV-B on total plant yield under field conditions. One notable exception is a 6-yr study with soybean demonstrating harvestable yield reductions under a simulated 25% ozone depletion. These effects are further modified by prevailing microclimatic conditions. Plants tend to be less sensitive to UV-B radiation under drought or mineral deficiency, while sensitivity increases under low levels of visible light. Further studies are needed to understand the mechanisms of UV-B effects and the interactions with present stresses and future projected changes in the environment.     

Serious complications in experiments in which UV doses are effected by using different lamp heights

Many experiments examining plant responses to enhanced ultraviolet-B radiation (280–315 nm) simply compare an enhanced UV-B treatment with ambient UV-B (or no UV-B radiation in most greenhouse and controlled-environment studies). Some more detailed experiments utilize multiple levels of UV-B radiation. A number of different techniques have been used to adjust the UV dose. One common technique is to place racks of fluorescent UV-emitting lamps at different heights above the plant canopy. However, the lamps and associated support structure cast shadows on the plant bed below. We calculated one example of the sequence of shade intervals for two common heights of lamp racks and show the patterns and duration of shade which the plants receive is distributed differently over the course of the day for different heights of the lamp racks. We also conducted a greenhouse experiment with plants (canola, sunflower and maize) grown under unenergized lamp racks suspended at the same two heights above the canopy. Growth characteristics differed in unpredictable ways between plants grown under the two heights of lamp racks. These differences could enhance or obscure potential UV-B effects. Also, differences in leaf mass per unit foliage area, which were observed in this experiment, could contribute to differences in plant UV-B sensitivity. We recommend the use of other techniques for achieving multiple doses of UV-B radiation. These range from simple and inexpensive approaches (e.g., wrapping individual fluorescent tubes in layers of a neutral-density filter such as cheese cloth) to more technical and expensive alternatives (e.g., electronically modulated lamp control systems). These choices should be determined according to the goals of the particular experiment.     

Increased UV-B Radiation Affects the Viability, Reactive Oxygen Species Accumulation and Antioxidant Enzyme Activities in Maize (Zea mays L.) Pollen

The increase in UV-B radiation reaching the earth's surface has prompted extensive studies on the effects of UV-B on plants. However, most of these studies have not addressed the close characteristics related to future survival of plant populations. The purpose of this study was to investigate the effects of UV-B radiation on reactive oxygen species (ROS) accumulation and antioxidant defense system in relation to germination, tube length and viability of maize pollen. Our results indicate that increased UV-B radiation decreased the pollen germination rate and tube length in vitro and also its fertilization ability in the field. Production of O₂^•− and H₂O₂ increased by UV-B radiation treatment, and their continuous accumulation resulted in lipid peroxidization. The activities of superoxide dismutase, catalase, peroxidase and DPPH-radical scavenging were decreased by increased UV-B radiation. The increased ROS and lipid peroxidization, and decreased activities of the antioxidants may be attributed to the effects of UV-B radiation on pollen germination, tube growth and fertilization ability.     

Ultraviolet-induced responses in two species of climax tropical marine macrophytes.

In tropical regions nominal reductions in stratospheric ozone could be detrimental to marine organisms that live near their upper tolerance levels of ultraviolet (UV) radiation and temperature. Well-known plant responses to UV include inhibition of photosynthesis, reductions in chlorophyll content, morphological changes and production of UV absorbing compounds such as flavonoids. An assessment of the effects and responses of two tropical marine macrophytes to full solar radiation and solar radiation depleted of UV were conducted in southwestern Puerto Rico. Changes in concentration of photosynthetic and photoprotective pigments, and in leaf optical properties of the red mangrove Rhizophora mangle and the seagrass Thalassia testudinum, were evaluated in field exclusion experiments. Rhizophora mangle exposed to full solar radiation showed lower leaf reflectance and a shift of 5 nm in the inflection point of the red edge. Thalassia testudinum samples excluded from UV had significant increases in total chlorophyll and carotenoid concentrations. These marine macrophytes showed increments in their concentration of UV-B absorbing compounds with exposure to UV radiation. Results indicate that even minor increases in UV radiation at low latitudes could have significant effects on the pigment composition of these climax species.