Outdoor Studies on the Effects of Solar UV-B on bryophytes: Overview and Methodology

In this review all recent field studies on the effects of UV-B radiation on bryophytes are discussed. In most of the studies fluorescent UV-B tubes are used to expose the vegetation to enhanced levels of UV-B radiation to simulate stratospheric ozone depletion. Other studies use screens to filter the UV-B part of the solar spectrum, thereby comparing ambient levels of UV-B with reduced UV-B levels, or analyse effects of natural variations in UV-B arising from stratospheric ozone depletion. Nearly all studies show that mosses are well adapted to ambient levels of UV-B radiation since UV-B hardly affects growth parameters. In contrast with outdoor studies on higher plants, soluble UV-B absorbing compounds in bryophytes are typically not induced by enhanced levels of UV-B radiation. A few studies have demonstrated that UV-B radiation can influence plant morphology, photosynthetic capacity, photosynthetic pigments or levels of DNA damage. However, there is only a limited number of outdoor studies presented in the literature. More additional, especially long-term, experiments are needed to provide better data for statistical meta-analyses. A mini UV-B supplementation system is described, especially designed to study effects of UV-B radiation at remote field locations under harsh conditions, and which is therefore suited to perform long-term studies in the Arctic or Antarctic. The first results are presented from a long-term UV-B supplementation experiment at Signy Island in the Maritime Antarctic.     

青藏高原强UV-B辐射对美丽风毛菊光合作用和色素含量的影响

以青藏高原矮嵩草草甸的主要伴随种美丽风毛菊为材料,通过滤除太阳辐射光谱中UV-B成分的模拟试验,研究了强太阳UV-B辐射对高山植物光合作用、光合色素和紫外吸收物质的影响.结果表明：与对照相比,弱UV-B处理能促使美丽风毛菊叶片净光合速率增加和提高稳态PSⅡ光化学效率;对照中叶片厚度的相对增加能弥补单位叶面积光合色素的光氧化损失,是高山植物对强UV B辐射的一种适应方式.短期滤除UV-B辐射处理时紫外吸收物质含量几乎没有变化,说明高山植物叶表皮层中该类物质受环境波动的影响较小.强UV-B环境下光合色素的相对增加是一种表象,而青藏高原强太阳UV-B辐射对高山植物美丽风毛菊的光合生理过程仍具有潜在的负影响.     

Photosynthesis, Growth, and Ultraviolet Irradiance Absorbance of Cucurbita pepo L. Leaves Exposed to Ultraviolet-B Radiation (280-315 nm)

Net photosynthesis, growth, and ultraviolet (UV) radiation absorbance were determined for the first leaf of Cucurbita pepo L. exposed to two levels of UV-B irradiation and a UV-B radiation-free control treatment. Absorbance by extracted flavonoid pigments and other UV-B radiation-absorbing compounds from the first leaves increased with time and level of UV-B radiation impinging on leaf surfaces. Although absorbance of UV-B radiation by extracted pigments increased substantially, UV-B radiation attenuation apparently was insufficient to protect completely the photosynthetic apparatus or leaf growth processes. Leaf expansion was repressed by daily exposure to 1365 Joules per meter per day of biologically effective UV-B radiation but not by exposure to 660 Joules per meter per day. Photosynthesis measured through ontogenesis of the first leaf was depressed by both UV-B radiation treatments. Repression of photosynthesis by UV-B radiation was especially evident during the ontogenetic period of maximum photosynthetic activity.     

UV responses of Lolium perenne raised along a latitudinal gradient across Europe: a filtration study

Lolium perenne (cv. AberDart) was grown at 14 locations along a latitudinal gradient across Europe (37-68°N) to study the impact of ultraviolet radiation (UV) and climate on aboveground growth and foliar UV-B absorbing compounds. At each location, plants were grown outdoors for 5 weeks in a replicated UV-B filtration experiment consisting of open, UV-B transparent (cellulose diacetate) and UV-B opaque (polyester) environments. Fourier transform-infrared spectroscopy was used to compare plant metabolite profiles in relation to treatment and location. UV radiation and climatic parameters were determined for each location from online sources and the data were assessed using a combination of anova and multiple regression analyses. Most of the variation in growth between the locations was attributable to the combination of climatic parameters, with minimum temperature identified as an important growth constraint. However, no single environmental parameter could consistently account for the variability in plant growth. Concentrations of foliar UV-B absorbing compounds showed a positive trend with solar UV across the latitudinal gradient; however, this relationship was not consistent in all treatments. The most striking experimental outcome from this study was the effect of presence or absence of filtration frames on UV-absorbing compounds. Overall, the study demonstrates the value of an European approach in studying the impacts of natural UV across a large latitudinal gradient. We have shown the feasibility of coordinated UV filtration at multiple sites but have also highlighted the need for open controls and careful interpretation of plant responses.     

(Poly)phenolic compounds in pollen and spores of Antarctic plants as indicators of solar UV-B – A new proxy for the reconstruction of past solar UV-B?

The morphology, size and characteristics of the pollen of the plant species Antarctic hairgrass (Deschampsia antarctica, Poaceae) and Antarctic pearlwort (Colobanthus quitensis, Caryophyllaceae) are described by scanning electron microscopy and light microscopy. Based on the number of pores the pollen of Colobanthus quitensis is classified as periporate or polypantorate, while that of Deschampsia antarctica is monoporate.

Pollen of Vicia faba plants, exposed to enhanced UV-B (10.6 kJ m^?2 day^?1 UV-B_BE) in a greenhouse, showed an increased content of UV-B absorbing compounds. There was also an increase of UV-B absorbing compounds in response to exposure to UV-A. By sequential chemical extraction three `compartments' of UV-B absorbance of pollen can be distinguished: a cytoplasmic fraction consisting of, e.g., flavonoids (acid-methanol extractable), a wall-bound fraction, consisting of, e.g., ferulic acid (NaOH extractable) and aromatic groups in the bioresistant polymer sporopollenin possibly consisting of, e.g., para-coumaric acid monomers (fraction remaining after acetolysis). The sporopollenin fraction in the pollen of Helleborus foetidus showed considerable UV-B absorbance (280–320 nm). There is evidence that enhanced solar UV-B induces increased UV-B absorbance (of sporopollenin) in pollen and spores of mosses, which may be preserved in the fossil record. As there are no instrumental records of solar UV-B in the Antarctic before 1970 and no instrumental records of stratospheric ozone over the Antarctic before 1957, the use of UV-B absorbing polyphenolics in pollen (and spores) as bio-indicator, or proxy of solar UV-B, may allow reconstruction of pre-ozone hole and subrecent UV-B and stratospheric ozone levels. Pollen and spores from herbarium specimens and from frozen moss banks (about 5000–10?000 years old) in the Antarctic may, therefore, represent a valuable archive of historical UV-B levels.

     

Morphological and physiological responses of bean plants to supplemental UV radiation in a Mediterranean climate

During the last few decades many experiments have been performed to evaluate the responses of plants to enhanced solar UV-B radiation (280–320 nm) that may occur because of stratospheric ozone depletion; most of them were performed in controlled environment conditions where plants were exposed to low photosynthetically active radiation (PAR) levels and high UV-B irradiance. Since environmental radiative regimes can play a role in the response of plants to UV-B enhancement, it appears doubtful whether it is valid to extrapolate the results from these experiments to plants grown in natural conditions. The objective of this work was to evaluate the effects on physiology and morphology of a bean (Phaseolus vulgaris L.) cultivar Nano Bobis, exposed to supplemental UV radiation in the open-air. UV-B radiation was supplied by fluorescent lamps to simulate a 20% stratospheric ozone reduction. Three groups of plants were grown: control (no supplemental UV), UV-A treatment (supplementation in the UV-A band) and UV-B treatment (supplemental UV-B and UV-A radiation). Each group was replicated three times. After 33 days of treatment plants grown under UV-B treatment had lower biomass, leaf area and reduced leaf elongation compared to UV-A treatment. No significant differences were detected in photosynthetic parameters, photosynthetic pigments and UV-B absorbing compounds among the three groups of plants. However, plants exposed to UV-A treatment showed a sort of 'stimulation' of their growth when compared to the control. The results of this experiment showed that plants may be sensitive to UV-A radiation, thus it is difficult to evaluate the specific effects of UV-B (280–320 nm) radiation from fluorescent lamps and it is important to choose the appropriate control. Environmental conditions strongly affect plant response to UV radiation so further field studies are necessary to assess the interaction between UV-B exposure and meteorological variability.     

Solar ultraviolet radiation affects the activity of ribulose-1,5-bisphosphate carboxylase-oxygenase and the composition of photosynthetic and xanthophyll cycle pigments in the intertidal green alga Ulva lactuca L

The effect of solar UV radiation on the physiology of the intertidal green macroalga Ulva lactuca L. was investigated. A natural Ulva community at the shore of Helgoland was covered with screening foils, excluding UV-B or UV-B + UV-A from the solar spectrum. In the sampled material, changes in the activity and concentration of ribulose-1,5-bisphosphate carboxylase-oxygenase (Rubisco), and the concentration of photosynthetic and xanthophyll cycle pigments were determined. Exclusion of UV radiation from the natural solar spectrum resulted in an elevated overall activity of Rubisco, related to an increase in its cellular concentration. Among the photosynthetic pigments, lutein concentration was substantially elevated under UV exclusion. In addition, marked UV effects on the xanthophyll cycle were found: exclusion of solar UV radiation (and particularly UV-B) resulted in an increased ratio of zeaxanthin concentration to the total xanthophyll content, indicating adverse effects of UV-B on the efficiency of photoprotection under high irradiances of photosynthetically active radiation. The results confirm a marked impact of present UV-B levels on macroalgal physiology under field conditions.     

Abscisic acid is involved in the response of grape (Vitis vinifera L.) cv. Malbec leaf tissues to ultraviolet-B radiation by enhancing ultraviolet-absorbing compounds, antioxidant enzymes and membrane sterols

We investigated the interactions of abscisic acid (ABA) in the responses of grape leaf tissues to contrasting ultraviolet (UV)-B treatments. One-year-old field-grown plants of Vitis vinifera L. were exposed to photosynthetically active radiation (PAR) where solar UV-B was eliminated by using polyester filters, or where PAR was supplemented with UV-B irradiation. Treatments combinations included weekly foliar sprays of ABA or a water control. The levels of UV-B absorbing flavonols, quercetin and kaempferol were significantly decreased by filtering out UV-B, while applied ABA increased their content. Concentration of two hydroxycinnamic acids, caffeic and ferulic acids, were also increased by ABA, but not affected by plus UV-B (+UV-B) treatments. Levels of carotenoids and activities of the antioxidant enzymes, catalase, ascorbate peroxidase and peroxidase were elevated by +ABA treatments, but only if +UV-B was given. Cell membrane β -sitosterol was enhanced by ABA independently of +UV-B. Changes in photoprotective compounds, antioxidant enzymatic activities and sterols were correlated with lessened membrane harm by UV-B, as assessed by ion leakage. Oxidative damage expressed as malondialdehyde content was increased under +UV-B treatments. Our results suggest that the defence system of grape leaf tissues against UV-B is activated by UV-B irradiation with ABA acting downstream in the signalling pathway.     

Rapid determination of spore chemistry using thermochemolysis gas chromatography-mass spectrometry and micro-Fourier transform infrared spectroscopy.

Spore chemistry is at the centre of investigations aimed at producing a proxy record of harmful ultraviolet radiation (UV-B) through time. A biochemical proxy is essential owing to an absence of long-term (century or more) instrumental records. Spore cell material contains UV-B absorbing compounds that appear to be synthesised in variable amounts dependent on the ambient UV-B flux. To facilitate these investigations we have developed a rapid method for detecting variations in spore chemistry using combined thermochemolysis gas chromatography-mass spectrometry and micro-Fourier transform infrared spectroscopy. Our method was tested using spores obtained from five populations of the tropical lycopsid Lycopodium cernuum growing across an altitudinal gradient (650-1981 m a.s.l.) in S.E. Asia with the assumption that they experienced a range of UV-B radiation doses. Thermochemolysis and subsequent pyrolysis liberated UV-B pigments (ferulic and para-coumaric acid) from the spores. All of the aromatic compounds liberated from spores by thermochemolysis and pyrolysis were active in UV-B protection. The various functional groups associated with UV-B protecting pigments were rapidly detected by micro-FTIR and included the aromatic C[double bond, length as m-dash]C absorption band which was exclusive to the pigments. We show increases in micro-FTIR aromatic absorption (1510 cm(-1)) with altitude that may reflect a chemical response to higher UV-B flux. Our results indicate that rapid chemical analyses of historical spore samples could provide a record ideally suited to investigations of a proxy for stratospheric O3 layer variability and UV-B flux over historical (century to millennia) timescales.     

Living on the edge – plants and global change in continental and maritime Antarctica

Antarctic terrestrial ecosystems experience some of the most extreme growth conditions on Earth and are characterized by extreme aridity and subzero temperatures. Antarctic vegetation is therefore at the physiological limits of survival and, as a consequence, even slight changes to growth conditions are likely to have a large impact, rendering Antarctic terrestrial communities sensitive to climate change. Climate change is predicted to affect the high‐latitude regions first and most severely. In recent decades, the Antarctic has undergone significant environmental change, including the largest increases in ultraviolet‐B (UV‐B; 290–320 nm) radiation levels in the world and, in the maritime region at least, significant temperature increases. This review describes the current evidence for environmental change in Antarctica, and the impacts of this change on the terrestrial vegetation. This is largely restricted to cryptogams, such as bryophytes, lichens and algae; only two vascular plant species occur in the Antarctic, both restricted to the maritime region. We review the range of ecological and physiological consequences of increasing UV‐B radiation levels, and of changes in temperature, water relations and nutrient availability. It is clear that climate change is already affecting the Antarctic terrestrial vegetation, and significant impacts are likely to continue in the future. We conclude that, in order to gain a better understanding of the complex dynamics of this important system, there is a need for more manipulative, long‐term field experiments designed to address the impacts of changes in multiple abiotic factors on the Antarctic flora.     

Role of ultraviolet-B radiation on bacterioplankton and the availability of dissolved organic matter

Attenuation of ultraviolet (UV)-radiation into the water column is highly correlated with the concentration of the dissolved organic matter (DOM). Thus UV penetrates deeper into marine waters than into freshwater systems. DOM is efficiently cleaved by solar surface radiation levels consuming more oxygen than bacterial metabolism. This photolytically cleaved DOM exhibits higher absorbance ratios (250/365 nm) than untreated DOM. Natural bacterioplankton reach higher abundance if inoculated in previously solar-exposed DOM than in untreated DOM; during bacterial growth the absorbance ratio declines steadily indicating the utilization of the photolytically cleaved DOM. On the other hand, bacterioplankton are greatly reduced in their activity if exposed to surface solar radiation levels. Photoenzymatic repair of DNA induced by UV-A radiation, however, leads to an efficient recovery of bacterial activity once the UV-B stress is released. Turbulent mixing of the upper layers of the water column leads to a continuous alteration of the UV exposure regime. Close to the surface, bacteria and DOM are exposed to high levels of UV-B leading to a reduction in bacterial activity and to photolysis of DOM. Once mixed into deeper layers where UV-B is attenuated, but sufficient UV-A is remaining to allow photoenzymatic repair, the photolytically cleaved DOM is efficiently taken up by bacterioplankton leading to even higher bacterial activity than prior to the exposure. Thus, the overall effect of UV on bacterioplankton is actually an enhancement of bacterial activity despite their lack of protective pigments.     

Solar UV-B radiation inhibits the growth of Antarctic terrestrial fungi

We tested the effects of solar radiation, and UV-B in particular, on the growth of Antarctic terrestrial fungi. The growth responses to solar radiation of five fungi, Geomyces pannorum, Phoma herbarum, Pythium sp., Verticillium sp., and Mortierella parvispora, each isolated from Antarctic terrestrial habitats, were examined on an agar medium in the natural Antarctic environment. A 3-h exposure to solar radiation of >287 nm reduced the hyphal extension rates of all species relative to controls kept in the dark. Pythium sp. cultures exposed to solar radiation for 1.5 h on five consecutive days were most sensitive to radiation of >287 nm, but radiation of >313 nm also inhibited growth to a lesser extent. Radiation of >400 nm had no effect on hyphal growth relative to controls kept in the dark. Short-wave solar UV-B radiation of between 287 and 305 nm inhibited the growth of Pythium sp. hyphae on and below the surface of the agar medium after 24 h, but radiation of > or =345 nm only reduced the growth of surface hyphae. Similar detrimental effects of UV-B on surface and, to a lesser extent, submerged hyphae of all five fungi were shown in the laboratory by using artificial UV-B from fluorescent lamps. A comparison of growth responses to solar radiation and temperature showed that the species that were most resistant to UV radiation grew fastest at higher temperatures. These data suggest that solar UV-B reduces the growth of fungi on the soil surface in the Antarctic terrestrial environment.     

Effect of solar ultraviolet-B radiation during springtime ozone depletion on photosynthesis and biomass production of Antarctic vascular plants

We assessed the influence of springtime solar UV-B radiation that was naturally enhanced during several days due to ozone depletion on biomass production and photosynthesis of vascular plants along the Antarctic Peninsula. Naturally growing plants of Colobanthus quitensis (Kunth) Bartl. and Deschampsia antarctica Desv. were potted and grown under filters that absorbed or transmitted most solar UV-B. Plants exposed to solar UV-B from mid-October to early January produced 11% to 22% less total, as well as above ground biomass, and 24% to 31% less total leaf area. These growth reductions did not appear to be associated with reductions in photosynthesis per se: Although rates of photosynthetic O(2) evolution were reduced on a chlorophyll and a dry-mass basis, on a leaf area basis they were not affected by UV-B exposure. Leaves on plants exposed to UV-B were denser, probably thicker, and had higher concentrations of photosynthetic and UV-B absorbing pigments. We suspect that the development of thicker leaves containing more photosynthetic and screening pigments allowed these plants to maintain their photosynthetic rates per unit leaf area. Exposure to UV-B led to reductions in quantum yield of photosystem II, based on fluorescence measurements of adaxial leaf surfaces, and we suspect that UV-B impaired photosynthesis in the upper mesophyll of leaves. Because the ratio of variable to maximal fluorescence, as well as the initial slope of the photosynthetic light response, were unaffected by UV-B exposure, we suggest that impairments in photosynthesis in the upper mesophyll were associated with light-independent enzymatic, rather than photosystem II, limitations.     

Leaf expansion and development of PHOTOSYNTHETIC CAPACITY AND PIGMENTS IN Liquidambar styraciflua (Hamamelidaceae)—EFFECTS OF UV-B RADIATION

In order to perform their functions as photosynthetic organs, leaves must cope with excess heat and potentially damaging ultraviolet radiation. Possible increases in the UV-B portion of the solar spectrum may place an additional burden on leaves, and this could be particularly important for young expanding leaves with poorly developed UV-B defense mechanisms. We evaluated the effects of supplemental UV-B radiation on leaf expansion and the development of photosynthetic capacity and pigments in sweetgum (Liquidambar styraciflua L.) seedlings. Seedlings were grown in the field under either ambient or ambient plus 3 or 5.0 kJ of biologically effective supplemental UV-B radiation. Although final leaf size was unaffected, the rate of leaf elongation and accumulation of leaf area was slower in leaves exposed to the lower supplemental UV-B irradiance. In contrast, chlorophyll accumulation and the development of photosynthetic capacity was more rapid in plants exposed to the higher, compared to the lower supplemental UV-B irradiance. The accumulation of anthocyanins and other putative flavonoids or UV-absorbing compounds was scarcely affected by exposure to supplemental UV-B radiation. These results suggest that the UV-B portion of the solar spectrum may, in the absence of gross affects on biomass, exert subtle influences on leaf ontogeny and the development of photosynthetic pigments and capacity in sweetgum.     

Impact of changes in natural ultraviolet radiation on pigment composition, physiological and morphological characteristics of the Antarctic moss, Grimmia antarctici

The impact of ambient ultraviolet (UV)‐B radiation on the endemic bryophyte, Grimmia antarctici, was studied over 14 months in East Antarctica. Over recent decades, Antarctic plants have been exposed to the largest relative increase in UV‐B exposure as a result of ozone depletion. We investigated the effect of reduced UV and visible radiation on the pigment concentrations, surface reflectance and physiological and morphological parameters of this moss. Plexiglass screens were used to provide both reduced UV levels (77%) and a 50% decrease in total radiation. The screen combinations were used to separate UV photoprotective from visible photoprotective strategies, because these bryophytes are growing in relatively high light environments compared with many mosses. G. antarctici was affected negatively by ambient levels of UV radiation. Chlorophyll content was significantly lower in plants grown under near‐ambient UV, while the relative proportions of photoprotective carotenoids, especially β‐carotene and zeaxanthin, increased. However, no evidence for the accumulation of UV‐B‐absorbing pigments in response to UV radiation was observed. Although photosynthetic rates were not affected, there was evidence of UV effects on morphology. Plants that were shaded showed fewer treatment responses and these were similar to the natural variation observed between moss growing on exposed microtopographical ridges and in more sheltered valleys within the turf. Given that other Antarctic bryophytes possess UV‐B‐absorbing pigments which should offer better protection under ambient UV‐B radiation, these findings suggest that G. antarctici may be disadvantaged in some settings under a climate with continuing high levels of springtime UV‐B radiation.     

Impact of solar ultraviolet-B radiation (290-320 nm) upon marine microalgae

For years scientists and laymen alike have casually noted the impact of solar ultraviolet radiation upon the non-human component of the biosphere. It was not until recently, when human activities were thought to threaten the protective stratospheric ozone shield, that researchers undertook intensive studies into the biological stress caused by the previously neglected short-wavelength edge of the global solar spectrum. Stratospheric ozone functions effectively as an ultraviolet screen by filtering out solar radiation in the 220–320 nm waveband as it penetrates through the atmosphere, thus allowing only small amounts of the longer wavelengths of radiation in this waveband to leak through to the surface of the earth. Although this ultraviolet radiation (UV-B radiation, 290–320 nm) comprises only a small fraction (less than 1%) of the total solar spectrum, it can have a major impact on biological systems due to its actinic nature. Many organic molecules, most notably DNA and proteins, absorb UV-B radiation which can initiate photochemical reactions. It is life's ability, or lack thereof, to cope with enhanced levels of solar UV-B radiation that has generated the concern over the potential depletion of stratospheric ozone. The defense mechanisms that serve to protect both plants and animals from current levels of UV-B radiation are quite varied. Whether these mechanisms will suffice for marine microalgae under conditions of enhanced levels of UV-B radiation is the subject of this review.     

低剂量微波辐射对增强UV-B辐射损伤菘蓝幼苗抗氧化酶活性的影响

为了探讨低剂量微波对增强UV-B辐射损伤菘蓝(Isatis indigotica Fort.)的修复作用,将经过10.08 kJ·m-2·d-1辐射损伤(PAR=220 μmol·m-2·s-1)的菘蓝幼苗分别经0、3、6、9和12 s等不同时间的微波辐照(126 mW·cm-2,2 450 MHz),然后测定其幼苗MDA含量、紫外吸收物质含量、抗坏血酸含量以及3种抗氧化酶SOD、CAT和POD活性.结果表明,增强UV-B辐射损伤菘蓝在微波的作用下其菘蓝幼苗中SOD、CAT和POD活性及紫外吸收物质含量、抗坏血酸含量提高,MDA含量得到显著的降低,说明微波对增强UV-B辐射伤害菘蓝幼苗具有修复作用.但是,随着微波剂量的增加,这种修复效应减弱,甚至消失.上述参数的变化说明适量时间的微波处理可以提高菘蓝对增强UV-B辐射的抵抗能力,并在此基础上初步探讨了微波的修复机理.     

Effects of solar ultraviolet radiation on the potential efficiency of photosystem II in leaves of tropical plants

The effects of solar ultraviolet (UV)-B and UV-A radiation on the potential efficiency of photosystem II (PSII) in leaves of tropical plants were investigated in Panama (9°N). Shade-grown tree seedlings or detached sun leaves from the outer crown of mature trees were exposed for short periods (up to 75 min) to direct sunlight filtered through plastic or glass filters that absorbed either UV-B or UV-A+B radiation, or transmitted the complete solar spectrum. Persistent changes in potential PSII efficiency were monitored by means of the dark-adapted ratio of variable to maximum chlorophyll a fluorescence. In leaves of shade-grown tree seedlings, exposure to the complete solar spectrum resulted in a strong decrease in potential PSII efficiency, probably involving protein damage. A substantially smaller decline in the dark-adapted ratio of variable to maximum chlorophyll a fluorescence was observed when UV-B irradiation was excluded. The loss in PSII efficiency was further reduced by excluding both UV-B and UV-A light. The photoinactivation of PSII was reversible under shade conditions, but restoration of nearly full activity required at least 10 d. Repeated exposure to direct sunlight induced an increase in the pool size of xanthophyll cycle pigments and in the content of UV-absorbing vacuolar compounds. In sun leaves of mature trees, which contained high levels of UV-absorbing compounds, effects of UV-B on PSII efficiency were observed in several cases and varied with developmental age and acclimation state of the leaves. The results show that natural UV-B and UV-A radiation in the tropics may significantly contribute to photoinhibition of PSII during sun exposure in situ, particularly in shade leaves exposed to full sunlight.