Environmental effects of ozone depletion and its interactions with climate change: progress report, 2004.
The complexity of the linkages between ozone depletion, UV-B radiation and climate change has become more apparent.
Atmosphere↗
Biomedical subjects
Lars Olof Björn
Publications and source records attributed to Lars Olof Björn.
Key findings and extended summaries.
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Animals↗
Rationale, concepts and approach to the assessment.
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Arctic Regions↗
Past changes in Arctic terrestrial ecosystems, climate and UV radiation.
At the last glacial maximum, vast ice sheets covered many continental areas. The beds of some shallow seas were exposed thereby connecting previously separated landmasses. Although some areas were ice-free and supported a flora and fauna, mean annual temperatures were 10-13 degrees C colder than during the Holocene. Within a few millennia of the glacial maximum, deglaciation started, characterized by a series of climatic fluctuations between about 18,000 and 11,400 years ago. Following the general thermal maximum in the Holocene, there has been a modest overall cooling trend, superimposed upon which have been a series of millennial and centennial fluctuations in climate such as the "Little Ice Age spanning approximately the late 13th to early 19th centuries. Throughout the climatic fluctuations of the last 150,000 years, Arctic ecosystems and biota have been close to their minimum extent within the most recent 10,000 years. They suffered loss of diversity as a result of extinctions during the most recent large-magnitude rapid global warming at the end of the last glacial stage. Consequently, Arctic ecosystems and biota such as large vertebrates are already under pressure and are particularly vulnerable to current and projected future global warming. Evidence from the past indicates that the treeline will very probably advance, perhaps rapidly, into tundra areas, as it did during the early Holocene, reducing the extent of tundra and increasing the risk of species extinction. Species will very probably extend their ranges northwards, displacing Arctic species as in the past. However, unlike the early Holocene, when lower relative sea level allowed a belt of tundra to persist around at least some parts of the Arctic basin when treelines advanced to the present coast, sea level is very likely to rise in future, further restricting the area of tundra and other treeless Arctic ecosystems. The negative response of current Arctic ecosystems to global climatic conditions that are apparently without precedent during the Pleistocene is likely to be considerable, particularly as their exposure to co-occurring environmental changes (such as enhanced levels of UV-B, deposition of nitrogen compounds from the atmosphere, heavy metal and acidic pollution, radioactive contamination, increased habitat fragmentation) is also without precedent.
Animals↗
Biodiversity, distributions and adaptations of Arctic species in the context of environmental change.
The individual of a species is the basic unit which responds to climate and UV-B changes, and it responds over a wide range of time scales. The diversity of animal, plant and microbial species appears to be low in the Arctic, and decreases from the boreal forests to the polar deserts of the extreme North but primitive species are particularly abundant. This latitudinal decline is associated with an increase in super-dominant species that occupy a wide range of habitats. Climate warming is expected to reduce the abundance and restrict the ranges of such species and to affect species at their northern range boundaries more than in the South: some Arctic animal and plant specialists could face extinction. Species most likely to expand into tundra are boreal species that currently exist as outlier populations in the Arctic. Many plant species have characteristics that allow them to survive short snow-free growing seasons, low solar angles, permafrost and low soil temperatures, low nutrient availability and physical disturbance. Many of these characteristics are likely to limit species' responses to climate warming, but mainly because of poor competitive ability compared with potential immigrant species. Terrestrial Arctic animals possess many adaptations that enable them to persist under a wide range of temperatures in the Arctic. Many escape unfavorable weather and resource shortage by winter dormancy or by migration. The biotic environment of Arctic animal species is relatively simple with few enemies, competitors, diseases, parasites and available food resources. Terrestrial Arctic animals are likely to be most vulnerable to warmer and drier summers, climatic changes that interfere with migration routes and staging areas, altered snow conditions and freeze-thaw cycles in winter, climate-induced disruption of the seasonal timing of reproduction and development, and influx of new competitors, predators, parasites and diseases. Arctic microorganisms are also well adapted to the Arctic's climate: some can metabolize at temperatures down to -39 degrees C. Cyanobacteria and algae have a wide range of adaptive strategies that allow them to avoid, or at least minimize UV injury. Microorganisms can tolerate most environmental conditions and they have short generation times which can facilitate rapid adaptation to new environments. In contrast, Arctic plant and animal species are very likely to change their distributions rather than evolve significantly in response to warming.
Adaptation, Physiological↗
Responses to projected changes in climate and UV-B at the species level.
Environmental manipulation experiments showed that species respond individualistically to each environmental-change variable. The greatest responses of plants were generally to nutrient, particularly nitrogen, addition. Summer warming experiments showed that woody plant responses were dominant and that mosses and lichens became less abundant. Responses to warming were controlled by moisture availability and snow cover. Many invertebrates increased population growth in response to summer warming, as long as desiccation was not induced. CO2 and UV-B enrichment experiments showed that plant and animal responses were small. However, some microorganisms and species of fungi were sensitive to increased UV-B and some intensive mutagenic actions could, perhaps, lead to unexpected epidemic outbreaks. Tundra soil heating, CO2 enrichment and amendment with mineral nutrients generally accelerated microbial activity. Algae are likely to dominate cyanobacteria in milder climates. Expected increases in winter freeze-thaw cycles leading to ice-crust formation are likely to severely reduce winter survival rate and disrupt the population dynamics of many terrestrial animals. A deeper snow cover is likely to restrict access to winter pastures by reindeer/caribou and their ability to flee from predators while any earlier onset of the snow-free period is likely to stimulate increased plant growth. Initial species responses to climate change might occur at the sub-species level: an Arctic plant or animal species with high genetic/racial diversity has proved an ability to adapt to different environmental conditions in the past and is likely to do so also in the future. Indigenous knowledge, air photographs, satellite images and monitoring show that changes in the distributions of some species are already occurring: Arctic vegetation is becoming more shrubby and more productive, there have been recent changes in the ranges of caribou, and "new" species of insects and birds previously associated with areas south of the treeline have been recorded. In contrast, almost all Arctic breeding bird species are declining and models predict further quite dramatic reductions of the populations of tundra birds due to warming. Species-climate response surface models predict potential future ranges of current Arctic species that are often markedly reduced and displaced northwards in response to warming. In contrast, invertebrates and microorganisms are very likely to quickly expand their ranges northwards into the Arctic.
Adaptation, Physiological↗
Effects on the structure of Arctic ecosystems in the short- and long-term perspectives.
Species individualistic responses to warming and increased UV-B radiation are moderated by the responses of neighbors within communities, and trophic interactions within ecosystems. All of these responses lead to changes in ecosystem structure. Experimental manipulation of environmental factors expected to change at high latitudes showed that summer warming of tundra vegetation has generally led to smaller changes than fertilizer addition. Some of the factors manipulated have strong effects on the structure of Arctic ecosystems but the effects vary regionally, with the greatest response of plant and invertebrate communities being observed at the coldest locations. Arctic invertebrate communities are very likely to respond rapidly to warming whereas microbial biomass and nutrient stocks are more stable. Experimentally enhanced UV-B radiation altered the community composition of gram-negative bacteria and fungi, but not that of plants. Increased plant productivity due to warmer summers may dominate food-web dynamics. Trophic interactions of tundra and sub-Arctic forest plant-based food webs are centered on a few dominant animal species which often have cyclic population fluctuations that lead to extremely high peak abundances in some years. Population cycles of small rodents and insect defoliators such as the autumn moth affect the structure and diversity of tundra and forest-tundra vegetation and the viability of a number of specialist predators and parasites. Ice crusting in warmer winters is likely to reduce the accessibility of plant food to lemmings, while deep snow may protect them from snow-surface predators. In Fennoscandia, there is evidence already for a pronounced shift in small rodent community structure and dynamics that have resulted in a decline of predators that specialize in feeding on small rodents. Climate is also likely to alter the role of insect pests in the birch forest system: warmer winters may increase survival of eggs and expand the range of the insects. Insects that harass reindeer in the summer are also likely to become more widespread, abundant and active during warmer summers while refuges for reindeer/caribou on glaciers and late snow patches will probably disappear.
Animals↗
Effects on the function of Arctic ecosystems in the short- and long-term perspectives.
Historically, the function of Arctic ecosystems in terms of cycles of nutrients and carbon has led to low levels of primary production and exchanges of energy, water and greenhouse gases have led to low local and regional cooling. Sequestration of carbon from atmospheric CO2, in extensive, cold organic soils and the high albedo from low, snow-covered vegetation have had impacts on regional climate. However, many aspects of the functioning of Arctic ecosystems are sensitive to changes in climate and its impacts on biodiversity. The current Arctic climate results in slow rates of organic matter decomposition. Arctic ecosystems therefore tend to accumulate organic matter and elements despite low inputs. As a result, soil-available elements like nitrogen and phosphorus are key limitations to increases in carbon fixation and further biomass and organic matter accumulation. Climate warming is expected to increase carbon and element turnover, particularly in soils, which may lead to initial losses of elements but eventual, slow recovery. Individual species and species diversity have clear impacts on element inputs and retention in Arctic ecosystems. Effects of increased CO2 and UV-B on whole ecosystems, on the other hand, are likely to be small although effects on plant tissue chemisty, decomposition and nitrogen fixation may become important in the long-term. Cycling of carbon in trace gas form is mainly as CO2 and CH4. Most carbon loss is in the form of CO2, produced by both plants and soil biota. Carbon emissions as methane from wet and moist tundra ecosystems are about 5% of emissions as CO2 and are responsive to warming in the absence of any other changes. Winter processes and vegetation type also affect CH4 emissions as well as exchanges of energy between biosphere and atmosphere. Arctic ecosystems exhibit the largest seasonal changes in energy exchange of any terrestrial ecosystem because of the large changes in albedo from late winter, when snow reflects most incoming radiation, to summer when the ecosystem absorbs most incoming radiation. Vegetation profoundly influences the water and energy exchange of Arctic ecosystems. Albedo during the period of snow cover declines from tundra to forest tundra to deciduous forest to evergreen forest. Shrubs and trees increase snow depth which in turn increases winter soil temperatures. Future changes in vegetation driven by climate change are therefore, very likely to profoundly alter regional climate.
Arctic Regions↗
Effects of changes in climate on landscape and regional processes, and feedbacks to the climate system.
Biological and physical processes in the Arctic system operate at various temporal and spatial scales to impact large-scale feedbacks and interactions with the earth system. There are four main potential feedback mechanisms between the impacts of climate change on the Arctic and the global climate system: albedo, greenhouse gas emissions or uptake by ecosystems, greenhouse gas emissions from methane hydrates, and increased freshwater fluxes that could affect the thermohaline circulation. All these feedbacks are controlled to some extent by changes in ecosystem distribution and character and particularly by large-scale movement of vegetation zones. Indications from a few, full annual measurements of CO2 fluxes are that currently the source areas exceed sink areas in geographical distribution. The little available information on CH4 sources indicates that emissions at the landscape level are of great importance for the total greenhouse balance of the circumpolar North. Energy and water balances of Arctic landscapes are also important feedback mechanisms in a changing climate. Increasing density and spatial expansion of vegetation will cause a lowering of the albedo and more energy to be absorbed on the ground. This effect is likely to exceed the negative feedback of increased C sequestration in greater primary productivity resulting from the displacements of areas of polar desert by tundra, and areas of tundra by forest. The degradation of permafrost has complex consequences for trace gas dynamics. In areas of discontinuous permafrost, warming, will lead to a complete loss of the permafrost. Depending on local hydrological conditions this may in turn lead to a wetting or drying of the environment with subsequent implications for greenhouse gas fluxes. Overall, the complex interactions between processes contributing to feedbacks, variability over time and space in these processes, and insufficient data have generated considerable uncertainties in estimating the net effects of climate change on terrestrial feedbacks to the climate system. This uncertainty applies to magnitude, and even direction of some of the feedbacks.
Arctic Regions↗
Synthesis of effects in four Arctic subregions.
An assessment of impacts on Arctic terrestrial ecosystems has emphasized geographical variability in responses of species and ecosystems to environmental change. This variability is usually associated with north-south gradients in climate, biodiversity, vegetation zones, and ecosystem structure and function. It is clear, however, that significant east-west variability in environment, ecosystem structure and function, environmental history, and recent climate variability is also important. Some areas have cooled while others have become warmer. Also, east-west differences between geographical barriers of oceans, archipelagos and mountains have contributed significantly in the past to the ability of species and vegetation zones to relocate in response to climate changes, and they have created the isolation necessary for genetic differentiation of populations and biodiversity hot-spots to occur. These barriers will also affect the ability of species to relocate during projected future warming. To include this east-west variability and also to strike a balance between overgeneralization and overspecialization, the ACIA identified four major sub regions based on large-scale differences in weather and climate-shaping factors. Drawing on information, mostly model output that can be related to the four ACIA subregions, it is evident that geographical barriers to species re-location, particularly the distribution of landmasses and separation by seas, will affect the northwards shift in vegetation zones. The geographical constraints--or facilitation--of northward movement of vegetation zones will affect the future storage and release of carbon, and the exchange of energy and water between biosphere and atmosphere. In addition, differences in the ability of vegetation zones to re-locate will affect the biodiversity associated with each zone while the number of species threatened by climate change varies greatly between subregions with a significant hot-spot in Beringia. Overall, the subregional synthesis demonstrates the difficulty of generalizing projections of responses of ecosystem structure and function, species loss, and biospheric feedbacks to the climate system for the whole Arctic region and implies a need for a far greater understanding of the spatial variability in the responses of terrestrial arctic ecosystems to climate change.
Animals↗
Uncertainties and recommendations.
An assessment of the impacts of changes in climate and UV-B radiation on Arctic terrestrial ecosystems, made within the Arctic Climate Impacts Assessment (ACIA), highlighted the profound implications of projected warming in particular for future ecosystem services, biodiversity and feedbacks to climate. However, although our current understanding of ecological processes and changes driven by climate and UV-B is strong in some geographical areas and in some disciplines, it is weak in others. Even though recently the strength of our predictions has increased dramatically with increased research effort in the Arctic and the introduction of new technologies, our current understanding is still constrained by various uncertainties. The assessment is based on a range of approaches that each have uncertainties, and on data sets that are often far from complete. Uncertainties arise from methodologies and conceptual frameworks, from unpredictable surprises, from lack of validation of models, and from the use of particular scenarios, rather than predictions, of future greenhouse gas emissions and climates. Recommendations to reduce the uncertainties are wide-ranging and relate to all disciplines within the assessment. However, a repeated theme is the critical importance of achieving an adequate spatial and long-term coverage of experiments, observations and monitoring of environmental changes and their impacts throughout the sparsely populated and remote region that is the Arctic.
Animals↗
Effects of temperature on UV-B-induced DNA damage and photorepair in Arabidopsis thaliana.
DNA damage in the form of cyclobutane pyrimidine dimers(CPDs) and (6-4) photoproducts(6-4PPs) induced by UV-B radiation in Arabidopsis thaliana at different temperatures was investigated using ELISA with specific monoclonal antibodies. CPDs and 6-4PPs increased during 3 h UV-B exposure, but further exposure led to decreases. Contrary to the commonly accepted view that DNA damage induced by UV-B radiation is temperature-independent because of its photochemical nature, we found UV-B-induction of CPDs and 6-4PPs in Arabidopsis to be slower at a low than at a high temperature. Photorepair of CPDs at 24 degrees C was much faster than that at 0 degrees C and 12 degrees C, with 50% CPDs removal during 1 h exposure to white light. Photorepair of 6-4PPs at 12 degrees C was very slow as compared with that at 24 degrees C, and almost no removal of 6-4PPs was detected after 4 h exposure to white light at 0 degrees C. There was evidence to suggest that temperature-dependent DNA damage and photorepair could have important ecological implications.
Arabidopsis↗
Changes in biologically active ultraviolet radiation reaching the Earth's surface.
Since publication of the 1998 UNEP Assessment, there has been continued rapid expansion of the literature on UV-B radiation. Many measurements have demonstrated the inverse relationship between column ozone amount and UV radiation, and in a few cases long-term increases due to ozone decreases have been identified. The quantity, quality and availability of ground-based UV measurements relevant to assessing the environmental impacts of ozone changes continue to improve. Recent studies have contributed to delineating regional and temporal differences due to aerosols, clouds, and ozone. Improvements in radiative transfer modelling capability now enable more accurate characterization of clouds, snow-cover, and topographical effects. A standardized scale for reporting UV to the public has gained wide acceptance. There has been increased use of satellite data to estimate geographic variability and trends in UV. Progress has been made in assessing the utility of satellite retrievals of UV radiation by comparison with measurements at the Earth's surface. Global climatologies of UV radiation are now available on the Internet. Anthropogenic aerosols play a more important role in attenuating UV irradiances than has been assumed previously, and this will have implications for the accuracy of UV retrievals from satellite data. Progress has been made inferring historical levels of UV radiation using measurements of ozone (from satellites or from ground-based networks) in conjunction with measurements of total solar radiation obtained from extensive meteorological networks. We cannot yet be sure whether global ozone has reached a minimum. Atmospheric chlorine concentrations are beginning to decrease. However, bromine concentrations are still increasing. While these halogen concentrations remain high, the ozone layer remains vulnerable to further depletion from events such as volcanic eruptions that inject material into the stratosphere. Interactions between global warming and ozone depletion could delay ozone recovery by several years, and this topic remains an area of intense research interest. Future changes in greenhouse gases will affect the future evolution of ozone through chemical, radiative, and dynamic processes In this highly coupled system, an evaluation of the relative importance of these processes is difficult: studies are ongoing. A reliable assessment of these effects on total column ozone is limited by uncertainties in lower stratospheric response to these changes. At several sites, changes in UV differ from those expected from ozone changes alone, possibly as a result of long-term changes in aerosols, snow cover, or clouds. This indicates a possible interaction between climate change and UV radiation. Cloud reflectance measured by satellite has shown a long-term increase at some locations, especially in the Antarctic region, but also in Central Europe, which would tend to reduce the UV radiation. Even with the expected decreases in atmospheric chlorine, it will be several years before the beginning of an ozone recovery can be unambiguously identified at individual locations. Because UV-B is more variable than ozone, any identification of its recovery would be further delayed.
Earth, Planet↗
Terrestrial ecosystems, increased solar ultraviolet radiation and interactions with other climatic change factors.
Based on research to date, we can state some expectations about terrestrial ecosystem response as several elements of global climate change develop in coming decades. Higher plant species will vary considerably in their response to elevated UV-B radiation, but the most common general effects are reductions in height of plants, decreased shoot mass if ozone reduction is severe, increased quantities of some phenolics in plant tissues and, perhaps, reductions in foliage area. In some cases, the common growth responses may be lessened by increasing CO2 concentrations. However, changes in chemistry of plant tissues will generally not be reversed by elevated CO2. Among other things, changes in plant tissue chemistry induced by enhanced UV-B may reduce consumption of plant tissues by insects and other herbivores, although occasionally consumption may be increased. Pathogen attack on plants may be increased or decreased as a consequence of elevated UV-B, in combination with other climatic changes. This may be affected both by alterations in plant chemistry and direct damage to some pathogens. Water limitation may decrease the sensitivity of some agricultural plants to UV-B, but for vegetation in other habitats, this may not apply. With global warming, the repair of some types of UV damage may be improved, but several other interactions between warming and enhanced UV-B may occur. For example, even though warming may lead to fewer killing frosts, with enhanced UV-B and elevated CO2 levels, some plant species may have increased sensitivity to frost damage.
Climate↗
Paleoclimate. Toward solving the UV puzzle.
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Atmosphere↗
UV-induced changes in pigment content and light penetration in the fruticose lichen Cladonia arbuscula ssp. mitis.
The response of the lichen, Cladonia arbuscula (Wallr.) Flot. ssp. mitis (Sandst.) Ruoss to enhanced UV-B (280-315 nm) radiation was investigated with respect to: (a) changes in phenolic content; (b) differential pigment accumulation under visible and UV radiation with increasing distance from thallus apices; and (c) the internal distribution of UV-B radiation within the thallus measured with quartz optical fibres. In a short-term experiment, lichens were exposed for 7 days in a growth chamber to visible light with or without additional UV-B radiation. For a longer term experiment, lichens were grown outdoors under both natural UV radiation, and supplemental UV-A (315-400 nm)+UV-B provided by lamps. Controls were placed under filters that removed the radiation below 290 nm from the natural sunlight. The concentration of total phenolic compounds was measured spectrophotometrically at the termination of the experiments, in different parts of the lichen podetia. UV-exposed lichens showed increased accumulation of phenolics compared to those not grown under UV. At the termination of the long-term experiment, fibre optic measurements of the penetration of radiation into lichen thallus reflected the influence of growth under UV radiation, whereby UV was more strongly attenuated as compared to that in lichens not exposed to enhanced levels of UV-B radiation. Results indicated that in Cladonia, UV-B radiation induces the accumulation of phenolic compounds that may have a protective role. In addition, the morphological distribution of phenolic compounds was different under visible and supplemental UV-B radiation. Internal radiation measurements served to visualise the attenuation of radiation with thallus depth for different wavelengths in the UV-B waveband.
Lichens↗
Growth and production of buckwheat (Fagopyrum esculentum) treated with reduced, ambient, and enhanced UV-B radiation.
The effect of enhanced UV-B radiation on buckwheat (Fagopyrum esculentum Moench. variety 'Darja'), an important high elevation crop, was studied in order to estimate its vulnerability in changing UV-B environment. Plants were grown in outdoor experiments from July to October under reduced and ambient UV-B levels, and an UV-B level simulating 17% ozone depletion in Ljubljana. During the development the following parameters were monitored: light saturated photosynthetic activity, transpiration, potential and effective photochemical efficiencies of photosystem II, the contents of photosynthetic pigments and methanol soluble UV-B absorbing compounds. At the end of the experiment, growth rate and production of seeds were estimated. In the following growth season the seeds collected from plants exposed to different UV-B treatments were tested for germination capacity. Total UV-B absorbing compounds during plant development were increased by UV-B radiation, photosynthetic pigments (chlorophyll a and b and carotenoids) decreased. Photosynthetic rate was lowered in an early stage of development. UV-B treatment resulted in the increase in the transpiration rate and consequently the decrease in water use efficiency (WUE). The disturbances in water economy and in photosynthesis affected the reproduction potential negatively; the production of seeds in plants cultivated under ambient and enhanced UV-B was 57 and 39% of the production of specimens treated with reduced UV-B, respectively. The germination of seeds collected from treated plants revealed on average about 95% success, independently of the treatment, but the time needed for germination was the shortest for seeds developed under enhanced UV-B level treatment. Enhanced UV-B radiation affected water relations and production of buckwheat, but not the potential of seeds for germination.
Chlorophyll↗
Temperature-dependent formation and photorepair of DNA damage induced by UV-B radiation in suspension-cultured tobacco cells.
Two photoproducts of DNA damage, i.e. cyclobutane pyrimidine dimers (CPDs) and 6-4 photoproducts (6-4PPs), induced by UV-B radiation in suspension-cultured tobacco cells were quantified by enzyme-linked immunosorbent assay (ELISA) with monoclonal antibodies. CPDs and 6-4PPs were induced in tobacco cells by UV-B radiation. Photorepair of CPDs was faster than that of 6-4PPs. UV-B radiation induces formation of CPDs and 6-4PPs even at 0 degrees C, but low temperature significantly decreases the UV-B-induced (in contrast to UV-C-induced) formation of CPDs and 6-4PPs. Low temperature also retarded the removal of CPDs and 6-4PPs under white light, and almost no photorepair of CPDs and 6-4PPs was detected at 0 degrees C. When purified DNA from tobacco cells grown in darkness was irradiated with UV-B, formation of CPDs and 6-4PPs took place at the same speed at different temperatures. It indicated that formation of CPDs and 6-4PPs induced by UV-B was temperature-independent in a non-cellular system. Based on our results for suspension-cultured tobacco cells, not only the photorepair but also UV-B-induced formation of CPDs and 6-4PPs are temperature-dependent.
Cells, Cultured↗