Ultraviolet radiation,ozone depletion,and marine photosynthesis

Concerns about stratospheric ozone depletion have stimulated interest in the effects of UVB radiation (280–320 nm) on marine phytoplankton. Research has shown that phytoplankton photosynthesis can be severely inhibited by surface irradiance and that much of the effect is due to UV radiation. Quantitative generalization of these results requires a biological weighting function (BWF) to quantify UV exposure appropriately. Different methods have been employed to infer the general shape of the BWF for photoinhibition in natural phytoplankton, and recently, detailed BWFs have been determined for phytoplankton cultures and natural samples. Results show that although UVB photons are more damaging than UVA (320–400 nm), the greater fluxes of UVA in the ocean cause more UV inhibition. Models can be used to analyze the sensitivity of water column productivity to UVB and ozone depletion. Assumptions about linearity and time-dependence strongly influence the extrapolation of results. Laboratory measurements suggest that UV inhibition can reach a steady-state consistent with a balance between damage and recovery processes, leading to a non-linear relationship between weighted fluence rate and inhibition. More testing for natural phytoplankton is required, however. The relationship between photoinhibition of photosynthesis and decreases in growth rate is poorly understood, so long-term effects of ozone depletion are hard to predict. However, the wide variety of sensitivities between species suggests that some changes in species composition are likely. Predicted effects of ozone depletion on marine photosynthesis cannot be equated to changes in carbon flux between the atmosphere and ocean. Nonetheless, properly designed studies on the effects of UVB can help identify which physiological and ecological processes are most likely to dominate the responses of marine ecosystems to ozone depletion.Abbreviations BWF biological weighting function - BWF/P-I photosynthesis versus photosynthetically available irradiance as influenced by biologically-weighted UV - Chl chlorophyll a - DOM dissolved organic matter - E _PAR irradiance in energy units (PAR) - E _s saturation parameter for PAR in the BWF/P-I model - E _inh ^* biologically-weighted dimensionless fluence rate for photoinhibition of photosynthesis by UV and PAR - biological weighting coefficient - % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGqiVu0Je9sqqrpepC0xbbL8F4rqqrFfpeea0xe9Lq-Jc9% vqaqpepm0xbba9pwe9Q8fs0-yqaqpepae9pg0FirpepeKkFr0xfr-x% fr-xb9adbaqaaeGaciGaaiaabeqaamaabaabaaGcbaGafqyTduMbae% baaaa!37AC!\[\bar \varepsilon \]_PAR biological weighting coefficient for damage to photosynthesis by E _PAR - k() diffuse attenuation coefficient for wavelength - MAAs mycosporine-like amino acids - PAR photosynthetically available radiation - P ^B rate of photosynthesis normalized to Chl - P _s ^B maximum attainable rate of photosynthesis in the absence of photoinhibition - UVA ultraviolet A (320–400 nm) - UVB ultraviolet B (280–320 nm)     

Effect of short-term exposure to UVA and UVB on potential phytoplankton production in UK coastal waters

The influence of vertical mixing on phytoplankton sensitivityto UV light has been assessed over an annual cycle. Photosynthesisrates of natural assemblages were compared in samples that wereincubated at fixed position in a light gradient and with duplicatesamples that simulated vertical mixing by movement in the samegradient with a periodicity of 4 h. This is the typical time-scaleof vertical mixing in coastal waters in the English Channel.There were clear seasonal differences in the short-term responseof phytoplankton to enhanced UVA+UVB. For most of the year,there was no detectable effect of UV on photosynthetic carbonfixation. But natural assemblages in late winter/early spring,when high UV light may sporadically occur at this latitude,were sensitive to UVA+UVB. In some samples, primary productionwas 40% of that measured in the absence of UV light. At thetime of maximum sensitivity to UV, the phytoplankton assemblagewas dominated by diatoms. Simulated vertical mixing resultedin more inhibition of photosynthesis by UVA+UVB light than whensamples were at constant light with the same time-integratedirradiance. Transient increases in UVA+UVB due to ozone depletion,such as have been observed over Northern Europe, could havea serious impact on coastal phytoplankton production in latewinter/early spring.     

Ultraviolet radiation and consumer effects on a field-grown intertidal macroalgal assemblage in Antarctica

Ultraviolet radiation (UVR) research on marine macroalgae has hithero focussed on physiological effects at the organism level, while little is known on the impact of UV radiation on macroalgal assemblages and even less on interactive effects with other community drivers, e.g. consumers. Field experiments on macrobenthos are scarce, particularly in the Antarctic region. Therefore, the effects of UVR and consumers (mainly limpets were excluded) on early successional stages of a hard bottom macroalgal community on King George Island, Antarctica, were studied. In a two‐factorial design experimental units [(1) ambient radiation, 280–700 nm; (2) ambient minus UVB, 320–700 nm and (3) ambient minus UVR, 400–700 nm vs. consumer–no consumer] were installed between November 2004 and March 2005 (n= 4 plus controls). Dry mass, species richness, diversity and composition of macroalgal assemblages developing on ceramic tiles were followed. Consumers significantly suppressed green algal recruits and total algal biomass but increased macroalgal richness and diversity. Both UVA and UVB radiation negatively affected macroalgal succession. UVR decreased the density of Monostroma hariotii germlings in the first 10 weeks of the experiment, whereas the density of red algal recruits was significantly depressed by UVR at the end of the study. After 106 days macroalgal diversity was significantly higher in UV depleted than in UV‐exposed assemblages. Furthermore, species richness was significantly lower in the UV treatments and species composition differed significantly between the UV‐depleted and the UV‐exposed treatment. Marine macroalgae are very important primary producers in coastal ecosystems, serving as food for herbivores and as habitat for many organisms. Both, UVR and consumers significantly shape macroalgal succession in the Antarctic intertidal. Consumers, particularly limpets can mediate negative effects of ambient UVR on richness and diversity till a certain level. UVB radiation in general and an increase of this short wavelength due to stratospheric ozone depletion in particular may have the potential to affect the zonation, composition and diversity of Antarctic intertidal seaweeds altering trophic interactions in this system.     

Variability of Primary Production in an Antarctic Marine Ecosystem as Estimated Using a Multi-scale Sampling Strategy

A major objective of the multidisciplinary Palmer Long TermEcological Research (LTER) program is to obtain a comprehensiveunderstanding of various components of the Antarctic marineecosystem—the assemblage of plants, animals, ocean, seaice, and island components south of the Antarctic Convergence.Phytoplankton production plays a key role in this polar ecosystem,and factors that regulate production include those that controlcell growth (light, temperature, nutrients) and those that controlcell accumulation rate and hence population growth (water columnstability, advection, grazing, and sinking). Several of thesefactors are mediated by the annual advance and retreat of seaice. In this study, we examine the results from nearly a decade(1991–2000) of ecological research in the western AntarcticPeninsula region. We evaluate the spatial and temporal variabilityof phytoplankton biomass (estimated as chlorophyll-a concentration)and primary production (determined in-situ aboard ship as wellas estimated from ocean color satellite data). We also presentthe spatial and temporal variability of sea ice extent (estimatedfrom passive microwave satellite data). While the data recordis relatively short from a long-term perspective, evidence isaccumulating that statistically links the variability in seaice to the variability in primary production. Even though thismarine ecosystem displays extreme interannual variability inboth phytoplankton biomass and primary production, persistentspatial patterns have been observed over the many years of study(e.g., an on to offshore gradient in biomass and a growing seasoncharacterized by episodic phytoplankton blooms). This high interannualvariability at the base of the food chain influences organismsat all trophic levels.     

Not just about sunburn – the ozone hole's profound effect on climate has significant implications for Southern Hemisphere ecosystems

Climate scientists have concluded that stratospheric ozone depletion has been a major driver of Southern Hemisphere climate processes since about 1980. The implications of these observed and modelled changes in climate are likely to be far more pervasive for both terrestrial and marine ecosystems than the increase in ultraviolet‐B radiation due to ozone depletion; however, they have been largely overlooked in the biological literature. Here, we synthesize the current understanding of how ozone depletion has impacted Southern Hemisphere climate and highlight the relatively few documented impacts on terrestrial and marine ecosystems. Reviewing the climate literature, we present examples of how ozone depletion changes atmospheric and oceanic circulation, with an emphasis on how these alterations in the physical climate system affect Southern Hemisphere weather, especially over the summer season (December–February). These potentially include increased incidence of extreme events, resulting in costly floods, drought, wildfires and serious environmental damage. The ecosystem impacts documented so far include changes to growth rates of South American and New Zealand trees, decreased growth of Antarctic mosses and changing biodiversity in Antarctic lakes. The objective of this synthesis was to stimulate the ecological community to look beyond ultraviolet‐B radiation when considering the impacts of ozone depletion. Such widespread changes in Southern Hemisphere climate are likely to have had as much or more impact on natural ecosystems and food production over the past few decades, than the increased ultraviolet radiation due to ozone depletion.     

THE EFFECTS OF ULTRAVIOLET‐B RADIATION ON ANTARCTIC SEA‐ICE ALGAE1

The impacts of ultraviolet‐B radiation (UVB) on polar sea‐ice algal communities have not yet been demonstrated. We assess the impacts of UV on these communities using both laboratory experiments on algal isolates and by modification of the in situ spectral distribution of the under‐ice irradiance. In the latter experiment, filters were attached to the upper surface of the ice so that the algae were exposed in situ to treatments of ambient levels of PAR and UV radiation, ambient radiation minus UVB, and ambient radiation minus all UV. After 16 d, significant increases in chl a and cell numbers were recorded for all treatments, but there were no significant differences among the different treatments. Bottom‐ice algae exposed in vitro were considerably less tolerant to UVB than those in situ, but this tolerance improved when algae were retained within a solid block of ice. In addition, algae extracted from brine channels in the upper meter of sea ice and exposed to PAR and UVB in the laboratory were much more tolerant of high UVB doses than were any bottom‐ice isolates. This finding indicates that brine algae may be better adapted to high PAR and UVB than are bottom‐ice algae. The data indicate that the impact of increased levels of UVB resulting from springtime ozone depletion on Antarctic bottom‐ice communities is likely to be minimal. These algae are likely protected by strong UVB attenuation by the overlying ice and snow, by other inorganic and organic substances in the ice matrix, and by algal cells closer to the surface.     

Ultraviolet B screening potential is higher in two cosmopolitan moss species than in a co-occurring Antarctic endemic moss: implications of continuing ozone depletion

Concentrations of UVB (ultraviolet B) absorbing pigments and anthocyanins were measured in three moss species, over a summer growing season in Antarctica. Pigment concentrations were compared with a range of climatic variables to determine if there was evidence that pigments were induced by UVB radiation, or other environmental parameters, and secondly if there were differences between species in their pigment responses. Significant seasonal differences in the potential UVB screening pigments were found, with the two cosmopolitan species Bryum pseudotriquetrum and Ceratodon purpureus appearing better protected from the potentially damaging effects of ozone depletion than the Antarctic endemic Schistidium antarctici. B. pseudotriquetrum accumulated the highest concentration of UVB screening pigments and showed positive associations between UVB radiation and both UVB absorbing and anthocyanin pigments. The negative associations between water availability measures and UVB absorbing and anthocyanin pigments also suggest that B. pseudotriquetrum is well protected in the desiccated state. This could offer B. pseudotriquetrum an advantage over the other species when high UVB radiation coincides with low temperatures and low water availability, thus limiting physiological activity and consequently, active photoprotective and repair mechanisms. As these pigments could act as either direct UVB screens or antioxidants, the results suggest that B. pseudotriquetrum is best equipped to deal with the negative effects of increased exposure to UVB radiation due to ozone depletion. The most exposed species, C. purpureus, has intermediate and stable concentrations of UVB absorbing pigments suggesting it may rely on constitutive UVB screens. Anthocyanin pigments were more responsive in this species and could offer increased antioxidant protection during periods of high UVB radiation. S. antarctici appears poorly protected and showed no evidence of any UV photoprotective response, providing additional evidence that this endemic is more vulnerable to climate change.     

History and evolution of primary productivity studies of the Southern Ocean

The main objective of the article is to give an overview of the history and evolution of phytoplankton research in the Southern Ocean during the past century and a half. It traces the evolution of phytoplankton investigations as it went through several phases commencing with intensive collecting and cataloging and leading to the Discovery investigations with its extensive and detailed studies of Southern Ocean phytoplankton. Following, in the footsteps of the Discovery was the USNS Eltanin with its specialized cruises that centered around the study of the ecology of the primary producers and the dynamics of the lower trophic levels of the food chain. Spurred by the findings of the Eltanin cruises and with the growing concern over the impending exploitation of the Antarctic marine living resources, in particular krill, the BIOMASS program was initiated. The program was the first international collaborative effort to study the dynamic functioning of the Antarctic marine ecosystem. The success of the program has rekindled great international interest in the Southern Ocean which resulted in an explosion of programs such as SO-JGOFS, SO-GLOBEC, EPOS, and several others. In recent years, there has been a major shift in phytoplankton research in the Southern Ocean. This was in response to worldwide concern over the effects of global warming and stratospheric ozone depletion on species diversity, primary production and ecosystem function. This has led to process-oriented programs to study phenomena and processes of global significance in which Antarctica and its surrounding seas play key roles.In an effort to avoid any confusion in the strict usage of terminology, the author has opted to use the Antarctic Ocean and the Southern Ocean interchangeably in this review.     

Phytoplankton and zooplankton response to ultraviolet radiation in a high-altitude Andean lake: short- versus long-term effects

Exclusion experiments on global UV (A and B) radiation and globalUVB were performed in 460 I mesocosms with plankton communitiesfrom the oligotrophic Andean lake Laguna Negra (33°35'S–70°04'W;2700 m a.s.l.). The experiments were run for 30 days duringthe summers of 1991–1992 and 1992–1993, and for48 days in 1993–1994. When UVB radiation was allowed toenter into the mesocosms (full sun), the population of Ankyrajudayi (Chlorophyta) reached the highest density, suggestingthat this species can endure high levels of UV radiation. Concurrently,an increase in chlorophyll a concentration was observed in thistreatment. The cladoceran Chydorus sphaericus and the rotiferLepadella ovallts were strongly inhibited by UVB. Conversely,UVB radiation had no effect on the survival of the differentlife stages of the calanoid copepod Boeckela gractlipes, suggestinga species-specific difference in the sensitivity to solar UVBradiation. Moreover, no reduction in the number of copepod eggsper female and the number of nauplii produced was observed.Apparently, herbivory does not strongly affect phytoplanktonabundance. Moreover, the phytoplankton species composition changedin the different treatments over the time. Fragilaria construensand Fragilaria crotonensis were dominant in those mesocosmswhere UVB was excluded. Populations fluc tuated depending ontheir life cycles and the period of time they were exposed toUVB radiation. It is important to define the time scale of exclusionexperiments, because different conclusions about the influenceof UVB irradiance result from short-, medium- or long-term exposures.     

Comparison of the impacts of consumers, ambient UV, and future UVB irradiance on mid-latitudinal macroepibenthic assemblages

Future levels in ultraviolet-B (UVB) radiation are expected to increase directly due to stratospheric ozone depletion and under water indirectly by, for example, global warming effects on DOC concentrations, altered trophic interactions in the plankton, or reduced eutrophication. While detrimental UV effects have been reported at the cellular level, little to nothing is known about community-wide effects of ambient and future UVB radiation. In a 4-month field experiment, the ambient UV regime was (i) reduced by cut-off filters which removed either UVB or total UV from the solar spectrum or (ii) increased to predicted future levels by UVB lamps. To allow relating the effects of present and future UV regimes to another important ecological control of community structure and diversity in subtidal marine habitats, consumer effects were quantified by an exclusion treatment under ambient light regimes. Ambient UV regimes did not affect community structure, biomass accrual, and diversity. In contrast, under enhanced UVB levels, the dominance of the competitively superior blue mussels increased and species richness and biomass accrual decreased. Species composition of the assemblages differed between the two UV regimes. Effects of enhanced UVB radiation and of consumption on biomass accrual, diversity, and structure of the community were comparable in magnitude and timing, but of opposite direction. In contrast, the effects of enhanced UVB radiation on growth and abundance of mussels were in the same direction, but shorter and weaker than consumer effects. Most UV effects were transitory and vanished within the first 2 months of succession. Our results indicate that present and future UVB levels may be of limited importance and not stronger in effect size than other ecological controls in structuring the shallow-water low-diversity macrobenthic communities in temperate regions.     

Antarctic terrestrial ecosystem and role of pigments in enhanced UV-B radiations

The terrestrial ecosystem of Antarctica are among the most extreme on earth, challenging the communities and making their existence difficult by rapidly increasing annual summer influx of solar ultraviolet radiations (UV-R), extremely cold conditions and lesser availability of nutrients. Spring time ozone depletion is due to release of chlorofluorocarbons in the earth atmosphere and is a serious cause of concern among environmentalists. Antarctic continent is mostly dominated by cryptogamic plants with limited distribution in different parts of the icy continent however; their distribution is mostly confined to Sub-Antarctic region. By the virtue of light requirement, cryptogams are exposed to extreme seasonal fluctuation in photosynthetically active radiation (PAR), and ultraviolet (UV) radiation which are closely associated with photosynthetic pigments in photoautotrophic organisms. Antarctic cryptogams cope up the stress imposed by UV radiation by the development of efficient systems for repairing damage by synthesis of screening compounds such as UV-B absorbing pigments and anthocyanin compounds. A major part of the UV absorbing compounds are appeared to be constitutive in lichens which are usnic acid, perlatolic acid and fumarphotocetraric acid which is particularly induced by UV-B. Secondary metabolites such as phenolics, atranorin, parietin and melanin also enhance the plant defense, by different molecular targets in specific solar irradiance and potential for increased antioxidative protection to UV induced vulnerability.     

Ultraviolet-B effects on photosystem II efficiency of natural phytoplankton communities from Antarctica

The impact of UVB on the Antarctic phytoplankton photosystem II repair cycle, involving the rapidly cycled D1 protein, was studied during summer 2002. On sunny and overcast days, phytoplankton (from 1-m depth) were exposed to natural light (+UVB) and Mylar-screened (–UVB) conditions. Half of the samples from each treatment were inoculated with lincomycin, an inhibitor of synthesis of chloroplast-encoded proteins including the D1 protein. Blocking D1 repair caused significant F_v/F_m depressions on sunny days but had not effect on the overcast day. Most of the F_v/F_m depression was caused by PAR and UVA with a non-significant contribution from UVB. In the presence of D1 repair, suppressing UVB had no effect on F_v/F_m when the samples originated from a weakly stratified water column with no defined upper mixed layer (UML) while it alleviated F_v/F_m depression when the phytoplankton samples originated from within an UML deeper than the depth of UVB penetration. These results suggest that UVB had more effect on the D1 repair process than on the damage process itself but that phytoplankton sensitivity to surface UVB exposure was influenced by their previous light history, partly determined by the vertical structure of the water column.     

Interannual variability in the distribution of the phytoplankton standing stock across the seasonal sea-ice zone west of the Antarctic Peninsula

The spatial distribution of phytoplankton cell abundance, carbon(C) biomass and chlorophyll a (Chl a) concentration was analysedduring three summers (1996, 1997 and 1999) in a seasonal sea-icearea, west of the Antarctic Peninsula. The objective of thestudy was to assess interannual variability in phytoplanktonspatial distribution and the mechanisms that regulate phytoplanktonaccumulation in the water column. Phytoplankton C biomass andChl a distributions were consistent from year to year, exhibitinga negative on/offshore gradient. The variations in C concentrationhad a close and non-linear relationship with the upper mixedlayer depth, suggesting that the vertical mixing of the watercolumn is the main factor regulating phytoplankton stock. Themagnitude of C gradients was 5-fold higher during 1996 thanduring 1997 and 1999. This was ascribed to interannual variationsin the concentration of diatom blooms in the region influencedby sea-ice melting. Vertical distribution of the phytoplankton,as estimated from Chl a profiles, also varied along an on/offshoregradient: Chl a was distributed homogeneously in the upper mixedlayer in coastal and mid-shelf stations and concentrated inthe deep layer (40–100 m) occupied by the winter waters(WW, remnants of the Antarctic surface waters during summer)in more offshore stations. The region with a deep Chl a maximumlayer (DCM layer) was dominated by a phytoplankton assemblagecharacterized by a relatively high concentration of diatoms.The extent of this region varied from year to year: it was restrictedto pelagic waters during 1996, extended to the shelf slope during1997 and occupied a major portion of the area during 1999. Itis hypothesized that iron depletion in near surface waters dueto phytoplankton consumption, and a higher concentration inWW, regulated this vertical phytoplankton distribution pattern.Furthermore, we postulate that year-to-year variations in thespatial distribution of the DCM layer were related to interannualvariations in the timing of the sea-ice retreat. The similaritybetween our results and those reported in literature for otherareas of the Southern Ocean allows us to suggest that the mechanismsproposed here as regulating phytoplankton stock in our areamay be applicable elsewhere.     

Effects of UV radiation on grazing by two marine heterotrophic nanoflagellates on autotrophic picoplankton

Phytoplankton <4-3 µm in diameter, or autotrophic picoplankton,can constitute the majority of the biomass and productivityof photosynthetic organisms in marine and freshwater systems.Indirect evidence has indicated that mortality of autotrophicpicoplankton occurs principally at night in the open ocean,but continuously in coastal water. The predominant view of thefate of autotrophic picoplankton production in the ocean isthat they are consumed by heterotrophic nanoflagellates. A possiblemechanism to explain these observations is that grazing of heterotrophicnanoflagellates on autotrophic picoplankton is inhibited byultraviolet radiation (W), at least in clear open-ocean environments.A series of laboratory experiments was conducted to examinethe effects of UV radiation on the grazing impact of two heterotrophicnanoflagellates on Synechococcus spp., a commonly occurringgenus of autotrophic picoplankton. The two nanoflagellates usedwere Paraphysomonas bandaieensis and Paraphysomonas imperforata.For both nanoflagellates, there was an inverse relationshipbetween the grazing mortality of Synechococcus and UV irradiance.The grazing mortality of Synechococcus was reduced less withP.imperforata than with P.bandaiensis. In some experiments,the effect of UV on the grazing impact of the nanoflagellatepopulations was caused in part by UV-related reductions in nanoflagellatesurvival. However, UV reduced the grazing impact of nanoflagellatesprimarily by reducing the rates of consumption of Synechococcusby individual nanoflagellates, to a degree directly relatedto UV irradiance. The results suggat that UV radiation may bean important selection factor in clear open-ocean water, andthat in order to predict the effect of increasing UV radiationon marine microbial plankton communities, we must consider interactionsbetween trophic levels as well as effects on single trophiclevels.     

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.     

Effects of UVB radiation on freshwater biota: a meta‐analysis

Aim Stratospheric ozone depletion and simultaneous increases in UVB radiation due to human activities have the potential to affect freshwater biota. The goal of our study is to summarize the impacts of UVB on freshwater biota by comparing the differences in the general patterns, including the directions and the magnitudes of the impacts of UVB on four major freshwater taxa (phytoplankton, zooplankton, fish and amphibians). The potential driving forces for these differences are also explored. Location Global. Methods We performed a meta‐analysis on a database consisting of 146 studies including 127 species from four taxonomic groups. We tested for the effects of taxonomic group, experimental venue, developmental stage, UVB dosage and the latitude of organism provenance. Results UVB had significant negative effects on freshwater biota from the molecular–cellular to individual–population levels. However, these effects were highly variable among the taxonomic groups. In general, zooplankton was the most negatively affected group, whereas fish and amphibians were less affected. As direct fitness components, survival and reproduction were the two responses most affected by UVB. The sensitivities of individuals to UVB at different developmental stages were different for the same taxon, while the stage‐dependent sensitivity patterns also differed among different taxa. Additionally, effects of different experimental venues, UVB dosages and latitudes of organism provenance on the effects of UVB were detected. Main conclusions Our results suggest that UVB has significant negative effects on freshwater biota. We found that the effects of UVB varied among taxonomic groups, developmental stages, experimental venues, UVB dosages and latitudes of organism provenance. The variation in sensitivity among the different taxa has important implications for ecosystem responses. Given that stratospheric ozone is unlikely to recover to the levels of the 1980s in the upcoming decades, more conservation efforts should be taken to protect freshwater habitats from further damage by UVB.     

Solar ultraviolet-B radiation affects plant-insect interactions in a natural ecosystem of Tierra del Fuego (southern Argentina)

We examined the effects of solar ultraviolet-B radiation (UVB) on plant-herbivore interactions in native ecosystems of the Tierra del Fuego National Park (southern Argentina), an area of the globe that is frequently under the Antarctic “ozone hole” in early spring. We found that filtering out solar UVB from the sunlight received by naturally-occurring plants of Gunnera magellanica, a creeping perennial herb, significantly increased the number of leaf lesions caused by chewing insects. Field surveys suggested that early-season herbivory was principally due to the activity of moth larvae (Lepidoptera: Noctuidae). Manipulative field experiments showed that exposure to solar UVB changes the attractiveness of G. magellanica leaf tissue to natural grazers. In a laboratory experiment, locally caught moth caterpillars tended to eat more tissue from leaves grown without UVB than from leaves exposed to natural UVB during development; however, the difference between treatments was not significant. Leaves grown under solar UVB had slightly higher N levels than leaves not exposed to UVB; no differences between UVB treatments in specific leaf mass, relative water content, and total methanol-soluble phenolics were detected. Our results show that insect herbivory in a natural ecosystem is influenced by solar UVB, and that this influence could not be predicted from crude measurements of leaf physical and chemical characteristics and a common laboratory bioassay. Received: 4 December 1997 / Accepted: 12 May 1998     

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.     

Interaction of nitrogen status and UVB sensitivity in a temperate phytoplankton assemblage

The influence of nitrate supplementation and ultraviolet-B (UVB; 280–320 nm) enhancement was tested on a coastal phytoplankton assemblage from eastern Canada exposed to ambient or supplemental UVB irradiance, equivalent to a local 60% ozone depletion. During a 10 d-long mesocosm experiment, 24-h surface bag incubations were repeated three times with half the bags supplemented with nitrate, phosphate and silicate while the other half received only phosphate and silicate. At beginning and the end of these 24-h surface incubations, chlorophyll fluorescence measurements were performed and the abundance of photoprotective pigments diadinoxanthin and diatoxanhtin, of the PSII reaction center D1 protein and of the large subunit of the ribulose-1,5-biphosphate carboxylase/oxygenase enzyme (RuBisCO LSU) was determined. Results showed that as long as nutrients were abundant inside the mesocosms, phytoplankton exposed to supplemental UVB at the surface were able to maintain their maximal quantum yield of PSII fluorescence. As nutrients became limiting inside the mesosocoms, however, phytoplankton showed an increased sensitivity to supplemental UVB and suffered more net photodamage to the PSII reaction centers (seen as decreases in D1 protein abundance). Supplemental UVB also resulted in low abundance of RuBisCO LSU and exacerbated photoinhibition compared to the phytoplankton exposed to surface ambient irradiance. Supplementing nitrate during this nutrient deficient period limited the inactivation of the PSII reaction centers and lowered photoinhibition. Nitrate supplementation had no clear effect on the abundance of the D1 protein but it helped the community to maintain a greater abundance of RuBisCO LSU. Overall, results from this study suggest that the sensitivity of the RuBisCO enzyme to the combined effects of supplemental UVB and nitrate limitation can influence the tolerance of PSII to UVB stress.     

EFFECTS OF INCREASING UV‐B RADIATION DUE TO OZONE DEPLETION ON PHOTOSYNTHESIS OF ANTARCTIC CYANOBACTERIA

Ground level ultraviolet‐B (UV‐B; 290–320 nm) fluxes in Antarctica have been increasing due to stratospheric ozone depletion. Although mat‐forming cyanobacteria are major component of freshwater algal biomass in Antarctica, little is known about their response to increasing ultraviolet radiation (UVR). The present study evaluated the sensitivity to UVR of two strains of mat‐forming cyanobacteria with different cell size, Phormidium murrayi (6.0 x 3.2 μm) and Schizothrix calcicola (2.2 x 2.3 μm). Cyanobacterial photosynthesis was measured under different UV spectral quality and quantity achieved by polychromatic filters with different cutoff wavelengths and neutral density screens. The productivity and irradiance data were used to generate biological weighting functions (BWF) for the assessment of UV inhibition on photosynthesis. The kinetics of UV inhibition, as determined by PAM fluorometry, differed between the two species so that inhibition of P. murrayi and S. calcicola were modeled based on UV‐irradiance and cumulative exposure, respectively. After a one hour exposure, BWF's did not differ between the two isolates of cyanobacteria despite their differences in cell size. To evaluate the negative impact of increased UV‐B exposure due to ozone depletion on cyanobacteria, the BWF's were applied to two solar spectra obtained from McMurdo Station, one on a day when the ozone hole was prominent (O₃ = 170 Dobson units; DU = 10‐3 cm O₃), and the other on a day with high ozone concentration (O₃ = 328 DU). The decrease in ozone level would reduce productivity by 3–8%. Seasonal variation of UVR has a bigger impact on cyanobacterial productivity than ozone depletion.