Distribution Patterns of Lentic-Breeding Amphibians in Relation to Ultraviolet Radiation Exposure in Western North America

An increase in ultraviolet-B (UV-B) radiation has been posited to be a potential factor in the decline of some amphibian population. This hypothesis has received support from laboratory and field experiments showing that current levels of UV-B can cause embryo mortality in some species, but little research has addressed whether UV-B is influencing the distribution of amphibian populations. We compared patterns of amphibian presence to site-specific estimates of UV-B dose at 683 ponds and lakes in Glacier, Olympic, and Sequoia–Kings Canyon National Parks. All three parks are located in western North America, a region with a concentration of documented amphibian declines. Site-specific daily UV-B dose was estimated using modeled and field-collected data to incorporate the effects of elevation, landscape, and water-column dissolved organic carbon. Of the eight species we examined (Ambystoma gracile, Ambystoma macrodactylum, Bufo boreas, Pseudacris regilla, Rana cascadae, Rana leuteiventris, Rana muscosa, Taricha granulosa), two species (T. granulosa and A. macrodactylum) had quadratic relationships with UV-B that could have resulted from negative UV-B effects. Both species were most likely to occur at moderate UV-B levels. Ambystoma macrodactylum showed this pattern only in Glacier National Park. Occurrence of A. macrodactylum increased as UV-B increased in Olympic National Park despite UV-B levels similar to those recorded in Glacier. We also found marginal support for a negative association with UV-B for P. regilla in one of the two parks where it occurred. We did not find evidence of a negative UV-B effect for any other species. Much more work is still needed to determine whether UV-B, either alone or in concert with other factors, is causing widespread population losses in amphibians.     

Exposure of red-legged frog embryos to ambient UV-B radiation in the field negatively affects larval growth and development

Exposure to ultraviolet-B radiation (UV-B; 280-320 nm) has a wide array of effects on aquatic organisms, including amphibians, and has been implicated as a possible factor contributing to global declines and range reductions in amphibian populations. Both lethal and sublethal effects of UV-B exposure have been documented for many amphibian species at various life-history stages. Some species, such as red legged frogs, Rana aurora, appear to be resistant to current ambient levels of UV-B, at least at the embryonic and larval stages, despite the fact that they have experienced range reductions in the Willamette Valley of Oregon, USA. However, UV-B is lethal to embryonic and larval R. aurora at levels slightly above those currently experienced during development. Therefore, we predicted that exposure of embryos to ambient UV-B radiation would result in sublethal effects on larval growth and development. We tested this by exposing R. aurora embryos to ambient UV-B in the field and then raising individuals in the laboratory for 1 month after hatching. Larvae that were exposed to UV-B as embryos were smaller and less developed than the non-exposed individuals 1 month post-hatching. These types of sublethal effects of UV-B exposure indicate that current levels of UV-B could already be influencing amphibian development.     

Sub-alpine amphibian distributions related to species palatability to non-native salmonids in the Klamath mountains of northern California

The goal of this study was to examine how introduced trout influence the distributions and abundances of a sub‐alpine amphibian assemblage whose members display a variety of different life‐history and defence strategies. Our study was conducted in the sub‐alpine lentic habitats of three wilderness areas that form the core of the Klamath‐Siskiyou Bioregion of northern California, a biodiversity ‘hotspot’ that supports the highest diversity of sub‐alpine, lentic‐breeding amphibians in the western USA. These wilderness areas contain no native fishes, but all have been populated with non‐native trout for recreational fishing. Five of the eight amphibian species that occur in this region were sufficiently common to use in our study; these included one that breeds in both temporary and permanent waters and is palatable to fish (Pacific treefrog, Pseudacris regilla), two that breed primarily in permanent waters and are unpalatable to fish (western toad, Bufo boreas, and rough‐skinned newt, Taricha granulosa), and two that breed primarily in permanent waters and are palatable to fish (Cascades frog, Rana cascadae, and long‐toed salamander, Ambystoma macrodactylum). Based on life histories and predator defence strategies (i.e. palatable or not), we predicted that the three palatable species would likely be negatively correlated with introduced trout, but with P. regilla less impacted because of its use of both temporary and permanent waters. We predicted that B. boreas and T. granulosa would not be significantly correlated with introduced trout due to the lack of any predator/prey interactions between them. We surveyed 728 pond, lake, or wet meadow sites during the summers of 1999–2002, using timed gill‐net sets to measure trout occurrence and relative density, and visual encounter surveys to determine amphibian presence and abundance. We used semiparametric logistic regression models to quantify the effect of trout presence/absence and density on the probability of finding amphibian species in a water body while accounting for variation within and among the various lentic habitats sampled. The distributions of P. regilla, A. macrodactylum and R. cascadae were strongly negatively correlated with trout presence across all three wilderness areas. Ambystoma macrodactylum was 44 times more likely to be found in lakes without fish than in lakes with fish. Rana cascadae and P. regilla were 3.7 and 3.0 times more likely, respectively, to be found in fishless than fish‐containing waters. In contrast, the two unpalatable species were either uncorrelated (T. granulosa) or positively correlated (B. boreas) with fish presence. We found that the relative density of fish (catch per unit effort) was negatively correlated with the combined abundances of the three palatable amphibians, and also with both the length and the condition of the fish themselves. Our results are consistent with a compelling body of evidence that introduced fishes greatly alter the aquatic community structure of mountain lakes, ponds, and wet meadows.     

Seasonal and diurnal variations in ultraviolet-B and ultraviolet-A irradiances at and below the sea surface at Helgoland (North Sea) over a 6-year period

Ultraviolet (UV) radiation at four wavelengths (305, 320, 340 and 380 nm) and photosynthetically active radiation (PAR) were measured from May 1994 to October 1999 using Biospherical UV radiometers. A surface reference sensor located on the roof of the Marine Station at Helgoland recorded values every 5 min, and an equivalent profiling underwater sensor was used for measurements in the sea at approximately monthly intervals. The ratio of 305-nm radiation to PAR varied seasonally, with a 14-fold increase from winter to summer. A much weaker seasonal trend (ca. 1.5-fold) was apparent in the ratio of 320-nm radiation to PAR, but there was no seasonal trend in the ratios of 340- or 380-nm radiation to PAR. The year-to-year variations in 305-nm radiation were also much greater relative to PAR than for the other UV wavelengths, but there was no evidence of a change in the 305 nm:PAR ratio over the study period. The ratios of both 305- and 320-nm radiation to PAR increased from dawn to midday, but those of 340- and 380-nm radiation were almost constant through the day, except shortly before sunrise and after sunset when the proportions of 340- and 380-nm radiation increased. Underwater measurements of PAR and UV suggest that the 1% depth for 305-nm radiation was little more than 1 m, but this estimate is valid only for summer and autumn because, in other seasons, few reliable readings for 305-nm radiation could be obtained underwater, and no attenuation coefficient could be calculated. The 1% depths recorded for the other UV wavelengths in the middle 6 months of the year were 2.0 m for 320 nm, 2.6 m for 340 nm and 4.6 m for 380 nm, compared with 12 m for PAR, but the attenuation of all wavebands increased sharply in October and remained higher until March. An analysis of the influence of sun angle, total column ozone concentration, the proportion of skylight, and cloud cover on the ratio of UV wavelengths to PAR in surface irradiance demonstrated that solar angle has a greater influence than ozone concentration on the irradiance at 305 nm, and that the typical occurrence of ozone "holes" in spring may not result in higher UV-B irradiances than occur under higher ozone concentrations in summer. The implications of the data for attempts to model the biological effects of natural UV radiation on marine organisms are considered. Received in revised form: 3 July 2000 Electronic Publication     

Temporal dynamics of ROS biogenesis under simulated solar radiation in the cyanobacterium Anabaena variabilis PCC 7937

We studied the temporal generation of reactive oxygen species (ROS) in the cyanobacterium Anabaena variabilis PCC 7937 under simulated solar radiation using WG 280, WG 295, WG 305, WG 320, WG 335, WG 345, and GG 400 nm cut-off filters to find out the minimum exposure time and most effective region of the solar spectrum inducing highest level of ROS. There was no significant generation of ROS in all treatments in comparison to the samples kept in the dark during the first 8 h of exposure; however, after 12 h of exposure, ROS were significantly generated in samples covered with 305, 295, or 280 nm cut-off filters. In contrast with ROS, the fragmentation of filaments was predominantly seen in 280 nm cut-off filter covered samples after 12 h of exposure. After 24 h of exposure, ROS levels were significantly higher in all samples than in the dark; however, the ROS signals were more pronounced in 320, 305, 295, or 280 nm cut-off filter covered samples. In contrast, the length of filaments was reduced in 305, 295, or 280 nm cut-off filter covered samples after 24 h of exposure. Thus, fragmentation of the filament was induced by all wavelengths of the UV-B region contrary to the UV-A region where only shorter wavelengths were able to induce the fragmentation. In contrast, ROS were generated by all wavelengths of the solar spectrum after 24 h of exposure; however, shorter wavelengths of both the UV-A and the UV-B regions were more effective in generating ROS in comparison to their higher wavelengths and photosynthetic active radiation (PAR). Moreover, lower wavelengths of UV-B were more efficient than the lower wavelengths of the UV-A radiation. Findings from this study suggest that certain threshold levels of ROS are required to induce the fragmentation of filaments.     

Risposte Fotomorfogeniche Indotte Dalla Radiazione UV-B in Brassica Oleracea var. Capitata

Abstract

Photomorphogenic responses induced by UV-B radiation in Brassica oleracea var. capitata.—Ultraviolet radiation can induce a plethora of “damaging” and “non damaging” effects in higher plants. We analyzed two possible photomorphogenic responses to UV-B radiation, the anthocyanin accumulation and the inhibition of hypocotyl elongation by modifying the UV spectral range with specific cut-off filters, under two levels of photon fluence rate. Experimental results suggest that detrimental effects are due to shorter wavelenghts of UV-B region (less than 305 nm); in contrast some adaptative responses may be induced by longer wavelenghts of UV-B region (between 305 and 320 nm). We attempted to determine the involvment of endogenous anthocyanin content in the UV-B photoprotection. Our experiments suggest a secondary role of anthocyanin accumulation in UV-B plant adaptation.     

Studies on the effects of UV-B radiation on phytoplankton of sub-antarctic lakes and ponds

Summary Experiments were performed to determine the effects of UV-B (ultraviolet, 280–320 nm) radiation on motility and growth of phytoplankton from lakes and ponds in South Georgia. After 4 h of solar radiation and 4h artificial radiation (UV-B_BE 11.6 kJ m^-2 day^-1, UV-B lamps) the swimming velocity of Cryptomonas sp. decreased. The growth rate of Botryococcus, Lyngbya sp. and Stauraslrum sp. did not show any significant variations between the different light conditions. The UV-B component was reduced by filtering solar radiation through glass bottles und cellulose acetate. Cloudy days had only 30% of the radiation of clear days in both the PAR (photosynthetic active radiation) and UV-B regions. The ponds contained large amounts of humic substances, which are responsible for the absorbance in the UV region.     

Spatial and temporal niche partitioning in grassland ants

We examined whether the species composition of a community influences the persistence of larval Ambystoma maculatum in assemblages composed of two larger intraguild predators (A. opacum and A. jeffersonianum) and an alternative prey species (tadpoles of Rana sylvatica). We predicted a priori that A. maculatum would have higher survival in more diverse communities containing alternative species of prey and top predators (A. opacum), the latter of which may lower the abundance of intermediate predators (A. jeffersonianum) via intraguild predation. In a factorial experiment, we manipulated the presence of larval A. opacum, A. jeffersonianum, and R. sylvatica in replicated artificial ponds containing larval A. maculatum. The presence of all three species significantly depressed biomass production in A. maculatum: biomass was highest in ponds lacking the other species and was zero in ponds initially containing all four species. Tadpoles severely reduced the growth of filamentous algae in the ponds. This, in turn, may have affected the abundance of some herbivorous prey of larval salamanders, although this possibility was not tested. The presence of congeneric predators severely restricted the presence in the water column of larval A. maculatum, which otherwise exhibited significant diel patterns of activity in the absence of predators. Together, the presence of tadpoles and a predator-mediated reduction in activity patterns may have limited foraging opportunities for A. maculatum, thus exacerbating the direct impact of predation on survival in this species. These results suggest that diverse assemblages consisting of these particular species may actually inhibit, rather than promote, inclusion of A. maculatum in some communities of pond-breeding amphibians.     

Estimated Ultraviolet Radiation Doses in Wetlands in Six National Parks

Ultraviolet-B radiation (UV-B, 280–320-nm wavelengths) doses were estimated for 1024 wetlands in six national parks: Acadia (Acadia), Glacier (Glacier), Great Smoky Mountains (Smoky), Olympic (Olympic), Rocky Mountain (Rocky), and Sequoia/Kings Canyon (Sequoia). Estimates were made using ground-based UV-B data (Brewer spectrophotometers), solar radiation models, GIS tools, field characterization of vegetative features, and quantification of DOC concentration and spectral absorbance. UV-B dose estimates were made for the summer solstice, at a depth of 1 cm in each wetland. The mean dose across all wetlands and parks was 19.3 W-h m⁻² (range of 3.4–32.1 W-h m⁻²). The mean dose was lowest in Acadia (13.7 W-h m⁻²) and highest in Rocky (24.4 W-h m⁻²). Doses were significantly different among all parks. These wetland doses correspond to UV-B flux of 125.0 μW cm⁻² (range 21.4–194.7 μW cm⁻²) based on a day length, averaged among all parks, of 15.5 h. Dissolved organic carbon (DOC), a key determinant of water-column UV-B flux, ranged from 0.6 (analytical detection limit) to 36.7 mg C L⁻¹ over all wetlands and parks, and reduced potential maximal UV-B doses at 1-cm depth by 1%–87 %. DOC concentration, as well as its effect on dose, was lowest in Sequoia and highest in Acadia (DOC was equivalent in Acadia, Glacier, and Rocky). Landscape reduction of potential maximal UV-B doses ranged from zero to 77% and was lowest in Sequoia. These regional differences in UV-B wetland dose illustrate the importance of considering all aspects of exposure in evaluating the potential impact of UV-B on aquatic organisms.     

Effects of UV-B Radiation on Anti-predator Behavior in Three Species of Amphibians

Ultraviolet radiation has been suggested as a possible contributing cause of amphibian declines around the world. Both laboratory and field studies have demonstrated that exposure to ultraviolet radiation can lead to increased mortality of developing amphibians. Virtually no studies have examined the sub‐lethal effects of ultraviolet on amphibian behavior. In this study, we examine the anti‐predator behavior of three species of amphibians after short‐term exposure to ultraviolet‐B radiation. Toad (Bufo boreas) juveniles that had been exposed to ultraviolet radiation did not respond to chemical extracts from conspecifics and heterospecifics as much as juveniles that had not been exposed. Both newt larvae (Taricha granulosa) that had been exposed to ultraviolet radiation and those that had not been exposed responded to chemical cues from conspecific predators by increasing the amount of time spent in shelter. Frog tadpoles (Rana cascadae) that had been exposed to ultraviolet radiation did not reduce their movement in response to chemical cues from predators as much as tadpoles that had not been exposed. These results indicate that ultraviolet exposure may have important sub‐lethal effects in amphibians that could adversely effect their fitness.     

The influence of UV-B radiation on light-dependent photosynthetic performance in <Emphasis Type="Italic">Sanionia uncinata </Emphasis>(Hedw.) Loeske in Antarctica

Photosynthetic activity of the moss Sanionia uncinata (Hedw.) Loeske was investigated on Léonie Island (67°35'S, 68°20'W, Antarctic Peninsula) in response to short-term changes of UV-B radiation. The UV-environment of natural mat formations dominated by S. uncinata was altered using filter screens. Two filter experiments were conducted in the Antarctic summers 1998 and 1999. A third filter experiment was conducted during springtime ozone depletion in October 1998. Photosynthetic activity of S. uncinata was mainly determined by photosynthetically active photon flux density (PPFD). Light response of relative electron transport rate through photosystem II (rel ETR=jF/Fm'2PPFD) remained unaffected by ambient summer levels of UV-B radiation. The same was found for net photosynthesis and dark respiration. In October 1998, S. uncinata was mainly metabolically inactive due to low temperatures. No significant levels of DNA-damage measured as cyclobutane pyrimidine dimers (CPDs) were induced by ambient summer levels of UV-B. Artificially enhanced UV-B radiation supplying a Setlow-DNA-dose of 8.7 kJ m^ф day^у UV-B led to formation of 7Dž CPD (10⁶ nucleotides)^-1. It is concluded that current ambient summer levels of UV-B radiation do not affect photosynthetic activity in S. uncinata.     

Experimental examination of the effects of ultraviolet-B radiation in combination with other stressors on frog larvae

Ultraviolet-B radiation (UVB) is a ubiquitous stressor with negative effects on many aquatic organisms. In amphibians, ambient levels of UVB can result in impaired growth, slowed development, malformations, altered behavior and mortality. UVB can also interact with other environmental stressors to amplify these negative effects on individuals. In outdoor mesocosm and laboratory experiments we studied potential synergistic effects of UVB, a pathogenic fungus, Batrachochytrium dendrobatidis (Bd), and varying temperatures on larval Cascades frogs (Rana cascadae). First, we compared survivorship, growth and development in two mesocosm experiments with UVB- and Bd-exposure treatments. We then investigated the effects of UVB on larvae in the laboratory under two temperature regimes, monitoring survival and behavior. We found reduced survival of R. cascadae larvae with exposure to UVB radiation in all experiments. In the mesocosm experiments, growth and development were not affected in either treatment, and no effect of Bd was found. In the laboratory experiment, larvae exposed to UVB demonstrated decreased activity levels. We also found a trend towards reduced survival when UVB and cold temperatures were combined. Our results show that amphibian larvae can suffer both lethal and sublethal effects when exposed to UVB radiation.     

Elevational differences in trait response to UV-B radiation by long-toed salamander populations

Amphibian species capable of optimizing trait response to environmental stressors may develop complex strategies for defending against rapid environmental change. Trait responses may differ between populations, particularly if stressor strength varies across spatial or temporal gradients. Ultraviolet-B (UV-B) radiation is one such stressor that poses a significant threat to amphibian species. We examined the ability of long-toed salamanders (Ambystoma macrodactylum) at high- and low-elevation breeding sites to cooperatively employ behavioral and physiological trait responses to mediate UV-B damage. We performed a microhabitat survey to examine differences in oviposition behavior and UV-B conditions among breeding populations at high- (n = 3; >1,500 m) and low-elevation (n = 3; <100 m) sites. We found significant differences in oviposition behavior across populations, with females at high-elevation sites selecting oviposition substrates in UV-B protected microhabitats. We also collected eggs (n = 633) from each of the breeding sites for analysis of photolyase activity, a photoreactivating enzyme that repairs UV-B damage to the DNA, using a photoproduct immunoassay. Our results revealed no significant differences in photolyase activity between long-toed salamander populations at high and low elevations. For high-elevation salamander populations, relatively low physiological repair capabilities in embryos appear to be buffered by extensive behavioral modifications to reduce UV-B exposure and standardize developmental temperatures. This study provides valuable insight into environmental stress responses via the assessment of multiple traits in allowing sensitive species to persist in rapidly changing landscapes.     

Inhibition of photosynthetic activity by UV-B radiation in radish seedlings

Inhibition of primary photosynthetic reactions by UV-B radiation (280 nm-320 nm) was demonstrated in radish leaves ( Raphanus sativus cv. Saxa Treib). Detached radish cotyledons from 10-day-old seedlings were irradiated with continuous white light and increasing UV-B irradiances using cut-off filters with increasing transmission for shorter wavelengths (WG 360, WG 345, WG 320, WG 305, WG 295, WG 280). Photosynthetic activity measured in terms of chlorophyll fluorescence induction (Kautsky effect) after 2, 4, 6, 8 and 24 h irradiation decreased in a wavelength dependent way with increasing UV-B irradiance and irradiation time.
Radish seedlings grown for 10 days from the time of germination under the same UV-B irradiation conditions exhibited similar reductions of the variable fluorescence as detached cotyledons irradiated for short time periods. They additionally had lower initial fluorescence at high UV-B radiation levels, although the chlorophyll content per leaf area increased. In contrast to short term experiments, the plastoquinone and flavonoid content increased with increasing UV-B irradiance when based on leaf area.     

Interspecific Variability in Sensitivity to UV Radiation and Subsequent Recovery in Selected Isolates of Marine Bacteria

The interspecific variability in the sensitivity of marine bacterial isolates to UV-B (295- to 320-nm) radiation and their ability to recover from previous UV-B stress were examined. Isolates originating from different microenvironments of the northern Adriatic Sea were transferred to aged seawater and exposed to artificial UV-B radiation for 4 h and subsequently to different radiation regimens excluding UV-B to determine the recovery from UV-B stress. Bacterial activity was assessed by thymidine and leucine incorporation measurements prior to and immediately after the exposure to UV-B and after the subsequent exposure to the different radiation regimens. Large interspecific differences among the 11 bacterial isolates were found in the sensitivity to UV-B, ranging from 21 to 92% inhibition of leucine incorporation compared to the bacterial activity measured in dark controls and from 14 to 84% for thymidine incorporation. Interspecific differences in the recovery from the UV stress were also large. An inverse relation was detectable between the ability to recover under dark conditions and the recovery under photosynthetic active radiation (400 to 700 nm). The observed large interspecific differences in the sensitivity to UV-B radiation and even more so in the subsequent recovery from UV-B stress are not related to the prevailing radiation conditions of the microhabitats from which the bacterial isolates originate. Based on our investigations on the 11 marine isolates, we conclude that there are large interspecific differences in the sensitivity to UV-B radiation and even larger differences in the mechanisms of recovery from previous UV stress. This might lead to UV-mediated shifts in the bacterioplankton community composition in marine surface waters.     

Amphibian defenses against ultraviolet-B radiation

As part of an overall decline in biodiversity, amphibian populations throughout the world are disappearing. There are a number of potential causes for these declines, including those related to environmental changes such as increasing ultraviolet-B (UV-B) radiation due to stratospheric ozone depletion. UV-B radiation can kill amphibian embryos or can cause sublethal effects that can harm amphibians in later life stages. However, amphibians have defenses against UV-B damage that can limit damage or repair it after exposure to UV-B radiation. These include behavioral, physiological, and molecular defenses. These defenses differ interspecifically, with some species more able to cope with exposure to UV-B than others. Unfortunately, the defense mechanisms of many species may not be effective against increasing persistent levels of UV-B radiation that have only been present for the past several decades due to human-induced environmental damage. Moreover, we predict that persistent UV-B-induced mortality and sublethal damage in species without adequate defenses could lead to changes in community structure. In this article we review the effects of UV-B radiation on amphibians and the defenses they use to avoid solar radiation and make some predictions regarding community structure in light of interspecific differences in UV-B tolerance.     

The interactive effects of UV-B radiation and a herbicide on uptake and allocation of carbon in two strains of the green algaScenedesmus

We examined the effects of UV B radiation on¹⁴C-uptake rates and carbon assimilation into the major end-products of photosynthesis of the green algaScenedesmus in the presence and absence of the triazine herbicide simetryn. Experiments were conducted using both a herbicide-susceptible and herbicide-tolerant strains ofScenedesmus. Three different UV-B dose rates were used as well as a light control. The lowest dose rate was almost the same level as in subsurface of ponds and lakes, while the other two were slightly lower and higher than natural sunlight on the surface of ponds and lakes, respectively. Total uptake rates of¹⁴C were not reduced by the UV B irradiation alone even at the highest dose rate. However, in the presence of the herbicide, uptake rates were clearly reduced by the highest dose rate of UV-B concomitant with increasing herbicide concentrations in the herbicide-susceptible strain. On the other hand, the proportion of lipid fraction was slightly reduced by all the UV-B treatments in the herbicide-susceptible strain even in the absence of the herbicide. In the herbicide-tolerant strain, uptake rates were not affected by UV-B radiation or by the herbicide. These facts indicated that UV-B effects could be smaller than predicted. It may be important to examine combined effects of UV-B and other anthropogenic and/or natural stresses for assessing actual UV-B effects in the field.     

Effect of UV-B Radiation on Leaf Optical Properties Measured with Fibre Optics

Changes in the internal light microenvironment in leaves ofplants of Brassica campestris L. cv. Emma, B. carinata L., andMedicago saliva L. cv. Armour in response to exposure to UV-B(UV-B, 280–320 nm) radiation were measured using a fibreopticmicroprobe. Plants were exposed for 2 weeks either to high visiblelight or to supplemental ultraviolet-B radiation. The spectral regime (400–700 nm; PAR) was measured eithermidway through the leaf palisade or the spongy mesophyll. Afterexposure to UV-B radiation leaves of Brassica campesiris attenuatedtransmitted light more than the controls. At the same time bothforward and back scattered light increased in the palisade andspongy mesophylls. In contrast, UV-treatment of Medicago salivaleaves increased light transmission into the palisade, whilethe back scattered component showed little change. Leaves ofcariiwla showed little change in response to UV. Other responsesto UV-B radiation included increases in leaf thickness, decreasedtotal chlorophyll content, and changes in UV-B screening pigmentsand chlorophyll fluorescence induction kinetics. Brassica campestriswas most sensitive to exposure to enhanced levels of UV-B radiation,whereas leaves of B. carinata were the least sensitive. Ourdata indicate that exposure to UV-B radiation altered the lightmicroenvironment within leaves of the species different ways.These changes appeared to be caused by alterations in pigmentcontent and leaf anatomy. In turn, the altered distributionof PAR within the leaf could influence photosynthesis. Key words: Brassica campestris, Brassica carinata, fibre optics, light scattering, Medicago saliva, optical properties, ozone depletion, photosynthesis, ultraviolet radiation     

Spatial and Temporal Variability in the Amount and Source of Dissolved Organic Carbon: Implications for Ultraviolet Exposure in Amphibian Habitats

The amount, chemical composition, and source of dissolved organic carbon (DOC), together with in situ ultraviolet (UV-B) attenuation, were measured at 1–2 week intervals throughout the summers of 1999, 2000, and 2001 at four sites in Rocky Mountain National Park (Colorado). Eight additional sites, four in Sequoia and Kings Canyon National Park/John Muir Wilderness (California) and four in Glacier National Park (Montana), were sampled during the summer of 2000. Attenuation of UV-B was significantly related to DOC concentrations over the three years in Rocky Mountain (R² = 0.39, F = 25.71, P < 0.0001) and across all parks in 2000 (R² = 0.44, F = 38.25, P < 0.0001). The relatively low R² values, however, reflect significant temporal and spatial variability in the specific attenuation per unit DOC. Fluorescence analysis of the fulvic acid DOC fraction (roughly 600–2,000 Daltons) indicated that the source of DOC significantly affected the attenuation of UV-B. Sites in Sequoia–Kings Canyon were characterized by DOC derived primarily from algal sources and showed much deeper UV-B penetration, whereas sites in Glacier and Rocky Mountain contained a mix of algal and terrestrial DOC-dominated sites, with more terrestrially dominated sites characterized by greater UV-B attenuation per unit DOC. In general, site characteristics that promoted the accumulation of terrestrially derived DOC showed greater attenuation of UV-B per unit DOC; however, catchment vegetation and soil characteristics, precipitation, and local hydrology interacted to make it difficult to predict potential exposure from DOC concentrations.     

Growth, structure, stomatal responses and secondary metabolites of birch seedlings (Betula pendula) under elevated UV-B radiation in the field

Three-year-old birch (Betula pendula Roth.) seedlings were exposed, in the field, to supplemental levels of UV-B radiation. Control seedlings were exposed to ambient levels of UV radiation, using arrays of unenergized lamps. A control for UV-A radiation was also included in the experiment. Enhanced UV-B radiation had no significant effects on height growth, and shoot and root biomass of birch seedlings. Leaf expansion rate increased transiently in the middle of the growing period in enhanced UV-B- and UV-A-exposed plants; however, final leaf size and relative growth rate remained unaffected. Leaf thickness and spongy intercellular spaces were increased in UV-B-exposed seedlings along with increased density of glandular trichomes. At the ultrastructural level, enhanced UV-B increased the number of cytoplasmic lipid bodies, and abnormal membrane whorls were found. Both enhanced UV-B and UV-A radiation induced swelling of chloroplast thylakoids. Stomatal density and conductance were significantly increased by elevated UV-B radiation. UV-A radiation increased the length and width of stomata, whereas UV-B radiation had only a marginal effect on stomatal size. UV-A and enhanced UV-B radiation attenuated the appearance of necrotic spots in autumn, probably caused by the fungus Pyrenopeziza betulicola, suggesting a direct harmful effect of UV on pathogens or reduced susceptibility to pathogens in UV-exposed seedlings. Secondary metabolite analysis showed increases in (+)-catechin, quercetin, cinnamic acid derivative, apigenin and pentagalloylglucose in birch leaves under enhanced UV-B radiation. Negative correlations between apigenin, and particularly quercetin concentrations and lipid peroxidation levels indicated an antioxidant role of secondary metabolites in birch leaves exposed to UV-B radiation.