Ultraviolet-absorbing compounds from the cell walls of an aquatic liverwort are more efficiently extracted by alkaline than by enzymatic digestion

Abstract

The accumulation of ultraviolet-absorbing compounds (UVACs) within the cell walls of bryophytes provides a spatially uniform filter of ultraviolet-B (UV-B) radiation, and thus represents an important protective mechanism against its adverse effects. Here, the abilities of several cell-wall-degrading enzymes and of alkali (NaOH) in extracting cell-wall-bound UVACs from the aquatic liverwort Jungermannia exsertifolia Steph. subsp. cordifolia (Dumort.) Váňa were compared, in order to select the most appropriate extraction method and to achieve the most reliable assessment of the degree of protection afforded by UVACs against UV-B. We analysed both the overall level of UVACs and the concentrations of two hydroxycinnamic acids (p-coumaric acid and ferulic acid). The most effective extraction (statistically significant) of cell-wall-bound UVACs, in terms both of their overall levels and of the concentrations of individual compounds, was achieved using alkaline digestion, which was more efficient than any of the enzymatic digestions trialled. This may be attributable to the ability of the alkali to break simultaneously both hydrogen bonds and covalent bonds within the cell wall, including ester linkages between phenolics and carbohydrates. In addition, alkaline digestion was more rapid than enzymatic digestions, and was not excessively aggressive, because the molecular integrity of the individual compounds analysed was preserved. Thus, alkaline digestion can be recommended for the extraction of cell-wall-bound UVACs in J. exsertifolia subsp. cordifolia, whether these compounds are to be evaluated overall or individually.     

Dynamic response of UV-absorbing compounds, quantum yield and the xanthophyll cycle to diel changes in UV-B and photosynthetic radiations in an aquatic liverwort

We studied the diel responses of the liverwort Jungermannia exsertifolia subsp. cordifolia to radiation changes under laboratory conditions. The samples were exposed to three radiation regimes: P (only PAR), PA (PAR + UV-A), and PAB (PAR + UV-A + UV-B). The day was divided in four periods: darkness, a first low-PAR period, the high-PAR plus UV period, and a second low-PAR period. After 15 days of culture, we measured photosynthetic pigments, chlorophyll fluorescence and UV-absorbing compounds in the four periods of the day on two consecutive days. With respect to UV-absorbing compounds, we analyzed their global amount (as the bulk UV absorbance of methanolic extracts) and the concentration of seven hydroxycinnamic acid derivatives, both in the soluble (mainly vacuolar) and insoluble (cell wall-bound) fractions of the plant extracts. PAB samples increased the bulk UV absorbance of the soluble and insoluble fractions, and the concentrations of p-coumaroylmalic acid in the soluble fraction and p-coumaric acid in the cell wall. Most of these variables showed significant diel changes and responded within a few hours to radiation changes (more strongly to UV-B), increasing at the end of the period of high-PAR plus UV. F_v/F_m, Φ_PSII, NPQ and the components of the xanthophyll cycle showed significant and quick diel changes in response to high PAR, UV-A and UV-B radiation, indicating dynamic photoinhibition and protection of PSII from excess radiation through the xanthophyll cycle. Thus, the liverwort showed a dynamic protection and acclimation capacity to the irradiance level and spectral characteristics of the radiation received.     

Response of cotton and sorghum to several levels of subambient solar UV-B radiation: a test of the saturation hypothesis

Some have proposed that plant responses to above-ambient or supplemented levels of solar ultraviolet-B radiation (UV-B; 280–315 nm) are typically subtle because targets or receptors in plants become saturated. If true, in solar UV-B filter exclusion experiments we would expect that plant responses would level off or 'saturate' as doses approached ambient levels. To test this so-called 'saturation hypothesis' we examined the response of Gossypium hirsutum (cotton) and Sorghum bicolor (sorghum) to filter exclusions that provided five levels of biologically effective UV-B, ranging from 36 to 91% of ambient solar levels in Arizona, USA. UV-B dose had no effect on biomass production of either species. As UV-B dose increased or approached ambient, individual leaves of S. bicolor were smaller, but plants produced more tillers and leaves. In G. hirsutum , individual leaves as well as total plant leaf area were smaller, but plants produced more branches. Bulk concentrations of soluble UV-B absorbing compounds increased with UV-B dose in both species. Leaf epidermal UV-B transmittance, assessed with the chlorophyll fluorescence technique, declined with increasing UV-B dose, and was well correlated with bulk concentrations of soluble UV-B screening compounds. Bulk concentrations of insoluble or wall-bound UV-B absorbing compounds were not affected by UV-B dose. The intensity of UV-induced blue fluorescence from leaf surfaces was strongly correlated with bulk concentrations of wall-bound UV-B absorbing compounds, and this signal has the potential to provide a rapid, non-invasive method to estimate concentrations of these compounds, which are time-consuming to extract. While both species were responsive to solar UV-B, responses did not appear to become saturated as doses approached ambient levels. Rather, responses required a threshold dose of >70% of solar ambient UV-B levels before they became apparent.     

Photosynthesis of two moss crusts from the Tengger Desert with contrasting sensitivity to supplementary UV-B radiation

Predicting the effects of increased ultraviolet-B (UV-B) radiation due to stratospheric ozone depletion on temperate desert ecosystems requires better knowledge of the ecophysiological response of common moss species. The aim of the current work was to determine whether elevated UV-B radiation affected photosynthetic performance and chloroplast ultrastructure of two moss crusts and whether response differences were observed between the crusts. In laboratory experiments, Bryum argenteum and Didymodon vinealis, which show microdistributions and are dominant in soil crusts at the Tengger Desert, Northern China, were subjected to four levels of UV-B radiation of 2.75 (control), 3.08, 3.25, and 3.41 W m^?2 for 10 days, simulating 0, 6, 9, and 12% of stratospheric ozone at the latitude of Shapotou, respectively. The results showed that chlorophyll a fluorescence parameters (i.e., the maximal quantum yield of PSII photochemistry, the effective quantum yield of PSII photochemistry, and photochemical quenching coefficient), pigment contents, soluble protein contents, and the ultrastructure were negatively influenced by elevated UV-B radiation and the degree of detrimental effects significantly increased with the intensity of UV-B radiation. Moreover, results indicated that B. argenteum was probably more sensitive to supplementary UV-B radiation than D. vinealis. Therefore, we propose the use of B. argenteum crusts as a bioindicator of responses to elevated UV-B radiation.     

Long-Term Dosimetry of Solar UV Radiation in Antarctica with Spores of Bacillus subtilis

The main objective was to assess the influence of the seasonal stratospheric ozone depletion on the UV climate in Antarctica by using a biological test system. This method is based on the UV sensitivity of a DNA repair-deficient strain of Bacillus subtilis (TKJ 6321). In our field experiment, dried layers of B. subtilis spores on quartz discs were exposed in different seasons in an exposure box open to solar radiation at the German Antarctic Georg von Neumayer Station (70°37′S, 8°22′W). The UV-induced loss of the colony-forming ability was chosen as the biological end point and taken as a measure for the absorbed biologically harmful UV radiation. Inactivation constants were calculated from the resulting dose-response curves. The results of field experiments performed in different seasons indicate a strongly season-dependent trend of the daily UV-B level. Exposures performed at extremely depleted ozone concentrations (October 1990) gave higher biologically harmful UV-B levels than expected from the calculated season-dependent trend, which was determined at normal ozone values. These values were similar to values which were measured during the Antarctic summer, indicating that the depleted ozone column thickness has an extreme influence on the biologically harmful UV climate on ground.     

Do effects of ultraviolet radiation on microbial films have indirect effects on larval attachment of the barnacle Balanus amphitrite?

We have examined the indirect effects of UV-A and UV-B on cypris attachment of the barnacle Balanus amphitrite Darwin through their effects on microbial films. Specifically, we tested the hypothesis that both UV-A and UV-B radiation can indirectly affect the larval attachment of barnacles by altering the microbial film bioactivity. Microbial films were developed from mid-intertidal region (∼1 m above Mean Low Water Level) for 6 days and subjected to ambient levels of ultraviolet radiation. Response of cyprids to untreated and UV-treated microbial films was investigated using double-dish still water choice bioassay. Results showed that both UV-A and UV-B caused a decrease in the percentage of respiring bacterial cells in microbial films and this effect increased with UV energy. With the same UV energy, UV-B caused a greater decrease in respiring bacterial cells than UV-A. However, despite strong UV radiation, the bioactivities of microbial films (i.e., stimulation of cypris attachment) remain unchanged. Results of this study suggest that increased UV radiation, which might occur due to ozone depletion, may not significantly affect the barnacle recruitment by means of affecting the inductive larval attachment cues of microbial films.     

Differential effects of elevated ultraviolet-B radiation on plant species of a dune grassland ecosystem

In a greenhouse study, plants of three monocotyledonous and five dicotyledonous species, which occur in a Dutch dune grassland, were exposed to four levels of ultraviolet-B (UV-B) radiation. UV-B levels simulated up to 30% reduction of the stratospheric ozone column during summertime in The Netherlands. Six of the plant species studied in the greenhouse were also exposed to enhanced UV-B irradiance in an experimental field study. In the field experiment plants either received the ambient UV-B irradiance (control) or an enhanced UV-B level simulating 15–20% ozone depletion during summertime in The Netherlands. The purpose of both experiments was to determine the response of the plant species to UV-B radiation and to compare results obtained in the greenhouse with results of the field experiment. Large intraspecific differences in UV-B sensitivity were observed in the greenhouse study. Total dry matter accumulation of monocotyledons was increased, while dry matter accumulation of dicotyledons remained unaffected or decreased. The increase in biomass production of monocotyledons at elevated UV-B was not related to the rate of photosynthesis but to alterations in leaf orientation. In the greenhouse study, UV-B radiation also affected morphological characteristics. Shoot height or maximum leaf length of five out of eight species was reduced. In the field study only one species showed a significantly decreased maximum leaf length at enhanced UV-B. Possible reasons for this discrepancy are discussed. The absorbance of methanolic leaf extracts also differed between species. UV absorbance of field-grown plants was higher than greenhouse-grown plants. In the greenhouse study, the highest UV-B level increased UV-B absorbance of some species. In the field study however, this stimulation of UV absorbance was not observed. In general, results obtained in the greenhouse study were similar to results obtained in the field study. Difficulties in extrapolating results of UV-B experiments conducted in the greenhouse to the field situation are discussed.     

Principal component analysis of intraspecific responses of tartary buckwheat to UV-B radiation under field conditions

Fifteen populations of tartary buckwheat (Fagopyrum tataricum Gaertn.) occurring in habitats with different natural UV-B levels were sampled, and the plants were exposed to enhanced UV-B radiation under field conditions simulating 25% depletion of the stratospheric ozone layer. The experimental design was a 2 × 15 factorial, with two levels of UV-B radiation (ambient and enhanced UV-B radiation) and plants from 15 populations. The responses of plants in growth, morphology, productivity and in the composition of photosynthetic pigments were measured. The results demonstrated that there were significant differences among populations in responses to UV-B radiation: some populations exhibited a positive effect while others were negatively affected. The UV-B effects on plant traits were correlated with the constitutive values. A principal component analysis (PCA) was used to evaluate the overall sensitivity of responses to UV-B radiation. Our results suggest that the sensitivity of plants to UV-B radiation is not only associated with the ambient UV-B level in natural habitats but also with the relative growth rate and other factors.     

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.     

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.     

Boreal peatland ecosystems under enhanced UV-B radiation and elevated tropospheric ozone concentration

Boreal peat-forming wetlands, mires, are globally important sources of methane and sinks for CO₂. As peatland vegetation plays a significant role regulating the exchange of these greenhouse gases, we have assessed the responses of the dominant plants and ecosystem functions to increasing tropospheric ozone concentration and enhanced ultraviolet-B (UV-B) radiation in long-term experiments, the results of which are summarized in this review. The dominant sedge, Eriophorum vaginatum, and especially the Sphagnum mosses common on peatlands, appear fairly tolerant to the future predicted ozone levels. Similarly, UV-B radiation only caused few alterations in the carbohydrates and pigments of the dominant sedge, Eriophorum russeolum, and had no effects on the dominant moss species of the experimental site, Warnstorfia exannulata. Surprisingly, there were alterations in organic acid concentrations in the peat pore water and peat microbial community composition in both experiments. Elevated ozone caused a transient decrease in ecosystem-level gross photosynthesis and methane (CH₄) emission, which shifted to a slight increase later on. Enhanced UV-B decreased dark ecosystem respiration and increased CH₄ emission in the course of the six measurement years. The emission of isoprene was increased by both ozone and UV-B during warm weather periods, suggesting interactive effects with temperature. All in all, we suggest that ozone and UV-B have limited effects on the carbon cycle in boreal peatlands, because other environmental factors, such as temperature, water level and photosynthetically active radiation more strongly regulate CO₂ and CH₄ exchange rates.     

Effects of supplemental UV-B radiation on primary photosynthetic carboxylating enzymes and soluble proteins in leaves of C3 and C4 crop plants

Effects of increased UV-B radiation on activities of primary photosynthetic carboxylating enzymes and on contents of soluble proteins were studied in soybean (Glycine max [L.] Merr. cv. Bragg), pea (Pisum sativum L. cv. Little Marvel), tomato (Lycopersicon esculentum L. cv. Rutgers), and sweet corn (Zea mays L. cv. Golden Cross Bantam). The purpose was to evaluate the responses of agronomic crops to increases in solar UV-B radiation. Plants were grown and exposed under greenhouse conditions for 6 h daily to supplemental UV-B radiation which was provided by Westinghouse FS-40 fluorescent sun lamps filtered with 0.127-mm film of cellulose acetate (UV-B treated) or Mylar S (Mylar control). Three UV-B levels were tested: 1.09 (treatment T₁), 1.36 (treatment T₂), and 1.83 (treatment T₃) UV-B_seu where 1 UV-B_seu equals 16.0 mW-m² weighted by EXP-[(λ-265)/21]². These UV-B levels corresponded to 6%,21%, and 36%, respectively, of decrease in stratospheric ozone content, based on the interpolations of UV-B irradiances at a solar elevation angle of 60°. Leaves of plants of soybean, pea, and tomato exposed to UV-B radiation were generally low in RuBP carboxylase activity. On a fresh weight basis, all three UV-B radiation levels significantly reduced the enzyme activity in soybean and pea, whereas tomato plants showed significant reduction in RuBP carboxylase activity only when exposed to 1.83 and 1.36 UV-B_seu. An apparent decrease in soluble proteins was observed in leaf extracts of soybean and pea plants exposed to 1.36 and 1.83 UV-B_seu whereas higher amounts of proteins were detected in leaves of tomato plants grown under UV-B radiation. Leaves of sweet corn plants grown under Mylar control were low in PEP carboxylase activity and proteins as compared with those of control plants receiving no supplemental UV and UV-B treatment. Activities of PEP carboxylase in crode extracts from leaves of sweet corn were significantly suppressed under 1.36 and 1.83 UV-B_seu as compared with the no UV control. Some stimulation of PEP carboxylase activity was observed in corn plants exposed to 1.09 UV-B_seu.     

Effect of enhanced UV-B radiation on pollen quantity,quality, and seed yield in Brassica rapa (Brassicaceae)

We conducted three experiments to examine the influence of ultraviolet-B radiation (UV-B; 280–320 nm) exposure on reproduction in Brassica rapa (Brassicaceae). Plants were grown in a greenhouse under three biologically effective UV-B levels that simulated either an ambient stratospheric ozone level (control), 16% (“low enhanced”), or 32% (“high enhanced”) ozone depletion levels at Morgantown, WV, USA in mid-March. In the first experiment, we examined whether UV-B level during plant growth influenced in vivo pollen production and viability, and flower production. Pollen production and viability per flower were reduced by ≈50% under both enhanced UV-B levels relative to ambient controls. While plants under high-enhanced UV-B produced over 40% more flowers than plants under the two lower UV-B treatments, whole-plant production of viable pollen was reduced under high-enhanced UV-B to 17% of that of ambient controls. Whole-plant production of viable pollen was reduced under low-enhanced UV-B to 34% of ambient controls. In the second experiment, we collected pollen from plants under the three UV-B levels and examined whether source-plant UV-B exposure influenced in vitro pollen germination and viability. Pollen from plants under both enhanced-UV-B treatments had initially lower germination and viability than pollen from the ambient level. After in vitro exposure to the high-enhanced UV-B levels for 6 h, viability of the pollen from plants grown under ambient UV-B was reduced from 65 to 18%. In contrast, viability of the pollen from plants grown under both enhanced UV-B treatments was reduced to a much lesser extent: only from ≈43 to 22%. Thus, ambient source-plant pollen was more sensitive to enhanced UV-B exposure. In the third experiment, we used pollen collected from source plants under the three UV-B levels to fertilize plants growing under ambient-UV-B levels, and assessed subsequent seed production and germination. Seed abortion rates were higher in plants pollinated with pollen from the enhanced UV-B treatments, than from ambient UV-B. Despite this, seed yield (number and mass) per plant was similar, regardless of the UV-B exposure of their pollen source. Our findings demonstrate that enhanced UV-B levels associated with springtime ozone depletion events have the capacity to substantially reduce viable pollen production, and could ultimately reduce reproductive success of B. rapa.     

Different response of two reindeer forage plants to enhanced UV-B radiation: modification of the phenolic composition

The long-term effects of enhanced UV-B radiation on the content and composition of leaf phenolics in Epilobium angustifolium L. and Eriophorum russeolum Fries ex Hartman were studied in northern Finland (68°N) using two UV-B enhancement experiments, both simulating UV-B_CIE radiation and corresponding to a 20% loss of ozone layer. High proportions of hydrolyzable tannins (69%) and condensed tannins (66%) characterized both Epilobium and Eriophorum leaves, respectively. No UV treatment effect was detected in the content or composition of Epilobium leaf soluble phenolics, whereas significant UV effects were detected in Eriophorum leaves in a developmental-specific manner. At the end of the growing season, the proportion of total soluble phenolics was higher in leaves exposed to enhanced UV-A and UV-B radiation than in the control leaves, but the phenolic composition was not significantly modified. This study introduces a new example on plants’ phenolic response to UV radiation being species-specific and detectable only at certain developmental stages. Possible consequences of increased phenolic content in forage plants for selection and digestibility by reindeer are, however, not yet known.     

Effects of UV-B radiation on near-surface zooplankton of puget sound

Summary An increase in incident solar ultraviolet irradiation, resulting from possible deterioration of the stratospheric ozone layer, would have important biological effects. Though the oceans are relatively opaque to UV radiation, compared to visible light, increases in incident UV may affect organisms living within the first few meters of the sea surface.Shrimp larvae, crab larvae, and euphausids were exposed to various low levels of simulated solar UV radiation (UV-B, 290–315 nm) under laboratory conditions. Comparisons between solar and artificial spectra were based on spectroradiometric measurements converted to erythemally effective irradiance. These zooplankton tolerated UV-B irradiance levels up to threshold levels with no significant reduction in survival or developmental rates compared to control organisms. Beyond the threshold levels, activity, development, and survival rapidly declined. The apparent UV thresholds are near present incident UV levels.Observed survival threshold levels for each experimental group were superimposed on seasonal solar incident UV levels at the experimental site. These threshold levels appeared to be exceeded by median ambient UV levels late in the season of surface occurrence of each species. UV increases resulting from ozone depletion may significantly shorten this season. Although the apparent impact would be lessened by the decrease in UV with depth, irreversible detrimental effects would probably occur before reported survival thresholds were exceeded.Contribution No. 1107 from the Department of Oceanography, University of Washington, Seattle, WA 98195, USA     

Changes in biologically-active ultraviolet radiation reaching the Earth's surface.

The Montreal Protocol is working. Concentrations of major ozone-depleting substances in the atmosphere are now decreasing, and the decline in total column amounts seen in the 1980s and 1990s at mid-latitudes has not continued. In polar regions, there is much greater natural variability. Each spring, large ozone holes continue to occur in Antarctica and less severe regions of depleted ozone continue to occur in the Arctic. There is evidence that some of these changes are driven by changes in atmospheric circulation rather than being solely attributable to reductions in ozone-depleting substances, which may indicate a linkage to climate change. Global ozone is still lower than in the 1970s and a return to that state is not expected for several decades. As changes in ozone impinge directly on UV radiation, elevated UV radiation due to reduced ozone is expected to continue over that period. Long-term changes in UV-B due to ozone depletion are difficult to verify through direct measurement, but there is strong evidence that UV-B irradiance increased over the period of ozone depletion. At unpolluted sites in the southern hemisphere, there is some evidence that UV-B irradiance has diminished since the late 1990s. The availability and temporal extent of UV data have improved, and we are now able to evaluate the changes in recent times compared with those estimated since the late 1920s, when ozone measurements first became available. The increases in UV-B irradiance over the latter part of the 20th century have been larger than the natural variability. There is increased evidence that aerosols have a larger effect on surface UV-B radiation than previously thought. At some sites in the Northern Hemisphere, UV-B irradiance may continue to increase because of continuing reductions in aerosol extinctions since the 1990s. Interactions between ozone depletion and climate change are complex and can be mediated through changes in chemistry, radiation, and atmospheric circulation patterns. The changes can be in both directions: ozone changes can affect climate, and climate change can affect ozone. The observational evidence suggests that stratospheric ozone (and therefore UV-B) has responded relatively quickly to changes in ozone-depleting substances, implying that climate interactions have not delayed this process. Model calculations predict that at mid-latitudes a return of ozone to pre-1980 levels is expected by the mid 21st century. However, it may take a decade or two longer in polar regions. Climate change can also affect UV radiation through changes in cloudiness and albedo, without involving ozone and since temperature changes over the 21st century are likely to be about 5 times greater than in the past century. This is likely to have significant effects on future cloud, aerosol and surface reflectivity. Consequently, unless strong mitigation measures are undertaken with respect to climate change, profound effects on the biosphere and on the solar UV radiation received at the Earth's surface can be anticipated. The future remains uncertain. Ozone is expected to increase slowly over the decades ahead, but it is not known whether ozone will return to higher levels, or lower levels, than those present prior to the onset of ozone depletion in the 1970s. There is even greater uncertainty about future UV radiation, since it will be additionally influenced by changes in aerosols and clouds.     

Influence of Temperature on the Effects of Artificially Enhanced UV-B Radiation on Aquatic Bryophytes Under Laboratory Conditions

We examined, under laboratory conditions, the influence of temperature (2 °C vs. 10 °C) on the physiological responses of two aquatic bryophytes from a mountain stream to artificially enhanced UV-B radiation for 82 d. These organisms may be exposed naturally to relatively low temperatures and high levels of UV-B radiation, and this combination is believed to increase the adverse effects of UV-B radiation. In the moss Fontinalis antipyretica, UV-B-treated samples showed severe physiological damages, including significant decreases in chlorophyll (Chl) and carotenoid (Car) contents, Chl a/b and Chl/phaeopigment ratios, Chl a fluorescence parameters F_v/F_m and _PS2, electron transport rate (ETR_max), and growth. In the liverwort Jungermannia cordifolia, UV-B radiation hardly caused any physiological change except for growth reduction. Thus, this liverwort seemed to be more tolerant to UV-B radiation than the moss under the specific experimental conditions used, maybe partly due to the accumulation of UV-B absorbing compounds. The influence of temperature on the effects of UV-B radiation depended on the species: the higher the UV-B tolerance, the lower the influence of temperature. Also, different physiological variables showed varied responses to this influence. Particularly, the lower temperature used in our study enhanced the adverse effects of UV-B radiation on important physiological variables such as F_v/F_m, growth, and Chl/phaeopigment ratios in the UV-B-sensitive F. antipyretica, but not in the more UV-B-tolerant J. cordifolia. Thus, the adverse effects of cold and UV-B radiation were apparently additive in the moss, but this additiveness was lacking in the liverwort. The Principal Components Analyses (PCA) conducted for both species with the physiological data obtained after 36 and 82 d of culture confirmed the above results. Under natural conditions, the relatively high water temperatures in summer might facilitate the acclimation of aquatic bryophytes from mountain streams to high levels of UV-B radiation. This may be relevant to predict the consequences of concomitant global warming and increasing UV-B radiation.     

UV-B辐射对马尾松凋落叶分解和养分释放的影响

由大气臭氧层减薄导致的UV-B辐射变化将直接影响到凋落物的分解。目前,有关UV-B辐射影响木本植物凋落物分解的研究还很少,在国内还没有开展。采用分解袋法开展了马尾松凋落叶在自然环境和UV-B辐射滤减两种辐射环境下的分解试验。结果表明：在UV-B辐射滤减环境下的马尾松凋落叶年分解速率比对照环境减慢了47.74%。UV-B辐射极显著(p<0.01)地加快了马尾松凋落叶的分解速率,促进了凋落叶中碳、磷、钾的释放和木质素的降解,对氮的释放无明显影响。研究结果意味着UV-B辐射将加快马尾松林的营养循环速度,降低马尾松林凋落物层的碳储量。     

IMPACTS OF SOLAR UV RADIATION ON THE PHOTOSYNTHESIS,GROWTH, AND UV‐ABSORBING COMPOUNDS IN GRACILARIA LEMANEIFORMIS (RHODOPHYTA) GROWN AT DIFFERENT NITRATE CONCENTRATIONS1

Solar ultraviolet radiation (UVR, 280–400 nm) is known to affect macroalgal physiology negatively, while nutrient availability may affect UV‐absorbing compounds (UVACs) and sensitivity to UVR. However, little is known about the interactive effects of UVR and nitrate availability on macroalgal growth and photosynthesis. We investigated the growth and photosynthesis of the red alga Gracilaria lemaneiformis (Bory) Grev. at different levels of nitrate (natural or enriched nitrate levels of 41 or 300 and 600 μM) under different solar radiation treatments with or without UVR. Nitrate‐enrichment enhanced the growth, resulted in higher concentrations of UVACs, and led to negligible photoinhibition of photosynthesis even at noon in the presence of UVR. Net photosynthesis during the noon period was severely inhibited by both ultraviolet‐A radiation (UVA) and ultraviolet‐B radiation (UVB) in the thalli grown in seawater without enriched nitrate. The absorptivity of UVACs changed in response to changes in the PAR dose when the thalli were shifted back and forth from solar radiation to indoor low light, and exposure to UVR significantly induced the synthesis of UVACs. The thalli exposed to PAR alone exhibited higher growth rates than those that received PAR + UVA or PAR + UVA + UVB at the ambient or enriched nitrate concentrations. UVR inhibited growth approximately five times as much as it inhibited photosynthesis within a range of 60–120 μg UVACs · g^?1 (fwt) when the thalli were grown under nitrate‐enriched conditions. Such differential inhibition implies that other metabolic processes are more sensitive to solar UVR than photosynthesis.