Interactive effects of changes in UV radiation and climate on terrestrial ecosystems, biogeochemical cycles, and feedbacks to the climate system.

Terrestrial organisms and ecosystems are being exposed to new and rapidly changing combinations of solar UV radiation and other environmental factors because of ongoing changes in stratospheric ozone and climate. In this Quadrennial Assessment, we examine the interactive effects of changes in stratospheric ozone, UV radiation and climate on terrestrial ecosystems and biogeochemical cycles in the context of the Montreal Protocol. We specifically assess effects on terrestrial organisms, agriculture and food supply, biodiversity, ecosystem services and feedbacks to the climate system. Emphasis is placed on the role of extreme climate events in altering the exposure to UV radiation of organisms and ecosystems and the potential effects on biodiversity. We also address the responses of plants to increased temporal variability in solar UV radiation, the interactive effects of UV radiation and other climate change factors (e.g. drought, temperature) on crops, and the role of UV radiation in driving the breakdown of organic matter from dead plant material (i.e. litter) and biocides (pesticides and herbicides). Our assessment indicates that UV radiation and climate interact in various ways to affect the structure and function of terrestrial ecosystems, and that by protecting the ozone layer, the Montreal Protocol continues to play a vital role in maintaining healthy, diverse ecosystems on land that sustain life on Earth. Furthermore, the Montreal Protocol and its Kigali Amendment are mitigating some of the negative environmental consequences of climate change by limiting the emissions of greenhouse gases and protecting the carbon sequestration potential of vegetation and the terrestrial carbon pool.

Environmental effects of stratospheric ozone depletion, UV radiation, and interactions with climate change: UNEP Environmental Effects Assessment Panel, Update 2021.

The Environmental Effects Assessment Panel of the Montreal Protocol under the United Nations Environment Programme evaluates effects on the environment and human health that arise from changes in the stratospheric ozone layer and concomitant variations in ultraviolet (UV) radiation at the Earth's surface. The current update is based on scientific advances that have accumulated since our last assessment (Photochem and Photobiol Sci 20(1):1-67, 2021). We also discuss how climate change affects stratospheric ozone depletion and ultraviolet radiation, and how stratospheric ozone depletion affects climate change. The resulting interlinking effects of stratospheric ozone depletion, UV radiation, and climate change are assessed in terms of air quality, carbon sinks, ecosystems, human health, and natural and synthetic materials. We further highlight potential impacts on the biosphere from extreme climate events that are occurring with increasing frequency as a consequence of climate change. These and other interactive effects are examined with respect to the benefits that the Montreal Protocol and its Amendments are providing to life on Earth by controlling the production of various substances that contribute to both stratospheric ozone depletion and climate change.

Environmental effects of stratospheric ozone depletion, UV radiation and interactions with climate change: UNEP Environmental Effects Assessment Panel, update 2019.

This assessment, by the United Nations Environment Programme (UNEP) Environmental Effects Assessment Panel (EEAP), one of three Panels informing the Parties to the Montreal Protocol, provides an update, since our previous extensive assessment (Photochem. Photobiol. Sci., 2019, 18, 595-828), of recent findings of current and projected interactive environmental effects of ultraviolet (UV) radiation, stratospheric ozone, and climate change. These effects include those on human health, air quality, terrestrial and aquatic ecosystems, biogeochemical cycles, and materials used in construction and other services. The present update evaluates further evidence of the consequences of human activity on climate change that are altering the exposure of organisms and ecosystems to UV radiation. This in turn reveals the interactive effects of many climate change factors with UV radiation that have implications for the atmosphere, feedbacks, contaminant fate and transport, organismal responses, and many outdoor materials including plastics, wood, and fabrics. The universal ratification of the Montreal Protocol, signed by 197 countries, has led to the regulation and phase-out of chemicals that deplete the stratospheric ozone layer. Although this treaty has had unprecedented success in protecting the ozone layer, and hence all life on Earth from damaging UV radiation, it is also making a substantial contribution to reducing climate warming because many of the chemicals under this treaty are greenhouse gases.

Solar UV radiation in a changing world: roles of cryosphere-land-water-atmosphere interfaces in global biogeochemical cycles.

Global change influences biogeochemical cycles within and between environmental compartments (i.e., the cryosphere, terrestrial and aquatic ecosystems, and the atmosphere). A major effect of global change on carbon cycling is altered exposure of natural organic matter (NOM) to solar radiation, particularly solar UV radiation. In terrestrial and aquatic ecosystems, NOM is degraded by UV and visible radiation, resulting in the emission of carbon dioxide (CO2) and carbon monoxide, as well as a range of products that can be more easily degraded by microbes (photofacilitation). On land, droughts and land-use change can reduce plant cover causing an increase in exposure of plant litter to solar radiation. The altered transport of soil organic matter from terrestrial to aquatic ecosystems also can enhance exposure of NOM to solar radiation. An increase in emission of CO2 from terrestrial and aquatic ecosystems due to the effects of global warming, such as droughts and thawing of permafrost soils, fuels a positive feedback on global warming. This is also the case for greenhouse gases other than CO2, including methane and nitrous oxide, that are emitted from terrestrial and aquatic ecosystems. These trace gases also have indirect or direct impacts on stratospheric ozone concentrations. The interactive effects of UV radiation and climate change greatly alter the fate of synthetic and biological contaminants. Contaminants are degraded or inactivated by direct and indirect photochemical reactions. The balance between direct and indirect photodegradation or photoinactivation of contaminants is likely to change with future changes in stratospheric ozone, and with changes in runoff of coloured dissolved organic matter due to climate and land-use changes.

Environmental effects of ozone depletion, UV radiation and interactions with climate change: UNEP Environmental Effects Assessment Panel, update 2017.

The Environmental Effects Assessment Panel (EEAP) is one of three Panels of experts that inform the Parties to the Montreal Protocol. The EEAP focuses on the effects of UV radiation on human health, terrestrial and aquatic ecosystems, air quality, and materials, as well as on the interactive effects of UV radiation and global climate change. When considering the effects of climate change, it has become clear that processes resulting in changes in stratospheric ozone are more complex than previously held. Because of the Montreal Protocol, there are now indications of the beginnings of a recovery of stratospheric ozone, although the time required to reach levels like those before the 1960s is still uncertain, particularly as the effects of stratospheric ozone on climate change and vice versa, are not yet fully understood. Some regions will likely receive enhanced levels of UV radiation, while other areas will likely experience a reduction in UV radiation as ozone- and climate-driven changes affect the amounts of UV radiation reaching the Earth's surface. Like the other Panels, the EEAP produces detailed Quadrennial Reports every four years; the most recent was published as a series of seven papers in 2015 (Photochem. Photobiol. Sci., 2015, 14, 1-184). In the years in between, the EEAP produces less detailed and shorter Update Reports of recent and relevant scientific findings. The most recent of these was for 2016 (Photochem. Photobiol. Sci., 2017, 16, 107-145). The present 2017 Update Report assesses some of the highlights and new insights about the interactive nature of the direct and indirect effects of UV radiation, atmospheric processes, and climate change. A full 2018 Quadrennial Assessment, will be made available in 2018/2019.

Effects of solar UV radiation and climate change on biogeochemical cycling: interactions and feedbacks.

Solar UV radiation, climate and other drivers of global change are undergoing significant changes and models forecast that these changes will continue for the remainder of this century. Here we assess the effects of solar UV radiation on biogeochemical cycles and the interactions of these effects with climate change, including feedbacks on climate. Such interactions occur in both terrestrial and aquatic ecosystems. While there is significant uncertainty in the quantification of these effects, they could accelerate the rate of atmospheric CO(2) increase and subsequent climate change beyond current predictions. The effects of predicted changes in climate and solar UV radiation on carbon cycling in terrestrial and aquatic ecosystems are expected to vary significantly between regions. The balance of positive and negative effects on terrestrial carbon cycling remains uncertain, but the interactions between UV radiation and climate change are likely to contribute to decreasing sink strength in many oceanic regions. Interactions between climate and solar UV radiation will affect cycling of elements other than carbon, and so will influence the concentration of greenhouse and ozone-depleting gases. For example, increases in oxygen-deficient regions of the ocean caused by climate change are projected to enhance the emissions of nitrous oxide, an important greenhouse and ozone-depleting gas. Future changes in UV-induced transformations of aquatic and terrestrial contaminants could have both beneficial and adverse effects. Taken in total, it is clear that the future changes in UV radiation coupled with human-caused global change will have large impacts on biogeochemical cycles at local, regional and global scales.

Ecological impact of solar ultraviolet-B (UV-B: 320-290 nm) radiation on Corynebacterium aquaticum and Xanthomonas sp. colonization on tea phyllosphere in relation to blister blight disease incidence in the field.

AIMS: To assess the effects of solar UV-B radiation on phyllosphere bacteria of tea leaves in relation to blister blight disease in the field. METHODS AND RESULTS: The effects of UV-B radiation on the phyllosphere microbiology of tea (Camellia sinensis) were studied in contrasting wet and dry seasons at a tropical site. Wavelength-selective filters were used to separate the effects of UV-B from those of other factors. Bacterial populations were quantified in relation to the incidence of blister blight disease. Attenuation of UV-B increased the survival of Xanthomonas sp. when populations were not water limited, and increased the incidence of blister blight, but had no effect on Corynebacterium aquaticum. CONCLUSIONS: The effects of solar UV-B on phyllosphere bacteria were substantial but depended on both species and interactions with other environmental variables. Xanthomonas sp. was more sensitive to UV-B than C. aquaticum, but this did not result in differences in population density under high radiation conditions (dry season), but only in the wet season when other factors were not limiting. SIGNIFICANCE AND IMPACT OF THE STUDY: The role of UV-B on leaf surface microbiology in the tropics is marked but depends on other conditions, and the contrasting UV-B responses of different organisms can be masked by other limiting factors.

Interactive effects of solar UV radiation and climate change on biogeochemical cycling.

This report assesses research on the interactions of UV radiation (280-400 nm) and global climate change with global biogeochemical cycles at the Earth's surface. The effects of UV-B (280-315 nm), which are dependent on the stratospheric ozone layer, on biogeochemical cycles are often linked to concurrent exposure to UV-A radiation (315-400 nm), which is influenced by global climate change. These interactions involving UV radiation (the combination of UV-B and UV-A) are central to the prediction and evaluation of future Earth environmental conditions. There is increasing evidence that elevated UV-B radiation has significant effects on the terrestrial biosphere with implications for the cycling of carbon, nitrogen and other elements. The cycling of carbon and inorganic nutrients such as nitrogen can be affected by UV-B-mediated changes in communities of soil organisms, probably due to the effects of UV-B radiation on plant root exudation and/or the chemistry of dead plant material falling to the soil. In arid environments direct photodegradation can play a major role in the decay of plant litter, and UV-B radiation is responsible for a significant part of this photodegradation. UV-B radiation strongly influences aquatic carbon, nitrogen, sulfur and metals cycling that affect a wide range of life processes. UV-B radiation changes the biological availability of dissolved organic matter to microorganisms, and accelerates its transformation into dissolved inorganic carbon and nitrogen, including carbon dioxide and ammonium. The coloured part of dissolved organic matter (CDOM) controls the penetration of UV radiation into water bodies, but CDOM is also photodegraded by solar UV radiation. Changes in CDOM influence the penetration of UV radiation into water bodies with major consequences for aquatic biogeochemical processes. Changes in aquatic primary productivity and decomposition due to climate-related changes in circulation and nutrient supply occur concurrently with exposure to increased UV-B radiation, and have synergistic effects on the penetration of light into aquatic ecosystems. Future changes in climate will enhance stratification of lakes and the ocean, which will intensify photodegradation of CDOM by UV radiation. The resultant increase in the transparency of water bodies may increase UV-B effects on aquatic biogeochemistry in the surface layer. Changing solar UV radiation and climate also interact to influence exchanges of trace gases, such as halocarbons (e.g., methyl bromide) which influence ozone depletion, and sulfur gases (e.g., dimethylsulfide) that oxidize to produce sulfate aerosols that cool the marine atmosphere. UV radiation affects the biological availability of iron, copper and other trace metals in aquatic environments thus potentially affecting metal toxicity and the growth of phytoplankton and other microorganisms that are involved in carbon and nitrogen cycling. Future changes in ecosystem distribution due to alterations in the physical and chemical climate interact with ozone-modulated changes in UV-B radiation. These interactions between the effects of climate change and UV-B radiation on biogeochemical cycles in terrestrial and aquatic systems may partially offset the beneficial effects of an ozone recovery.

Tolerance of Senecio vulgaris to Infection and Disease Caused by Native and Alien Rust Fungi.

ABSTRACT Plant defense strategies against pathogen attack can be divided into either resistance or tolerance. Variation in tolerance is expressed as differences in the relationship between host fitness (or yield) and the degree of infection. Plant tolerance of pathogen attack remains poorly understood both in terms of its specific mechanisms and in terms of the evolutionary processes by which it has arisen. Theoretical models predict that it is the result of coevolution between host and pathogen, suggesting greater tolerance in interactions with native as opposed to introduced pathogens. Therefore, we quantified and compared the degree of tolerance expressed in the interaction of Senecio vulgaris with the rust fungus Coleosporium tussilginis, which is native to the UK, and the introduced rust fungus Puccinia lagenophorae. We used the reaction norm approach to quantify tolerance and its components. The S. vulgaris-C. tussilaginis interaction expressed a significantly greater degree of tolerance, as reductions in host growth and fitness per unit infection were significantly less than with P. lagenophorae. The key mechanism for this greater tolerance to C. tussilaginis was a significantly smaller reduction in photosynthesis per unit infection than with P. lagenophorae, at both leaf and whole plant scales. There was no significant difference in the relationship between whole plant photosynthesis and host reproduction. We discuss these responses in the context of coevolution for tolerance in host-pathogen interactions.

Role for non-homologous end-joining in the repair of UVA-induced DNA damage.

PURPOSE: The biological significance of long-wavelength ultraviolet (UV) light, UVA, is increasingly realized, but the precise nature of the cellular damage responsible for the effects of this radiation is still not clear. It has been reported that UVA can induce double-strand breaks in DNA, but the biological significance of these is not known. We have therefore examined the UVA sensitivity of a cell line deficient in non-homologous end-joining, the major pathway for the repair of DNA double-strand breaks in mammalian cells in order to determine the biological importance of UVA-induced DSB. MATERIALS AND METHODS: Xrs-6, a Chinese hamster ovary cell line mutant for XRCC5 (Ku80) was compared with its parental CHO-K1 cell line for its sensitivity to UVA radiation (365 nm) using both a clonogenic assay and the micronucleus assay. RESULTS: Xrs-6 cells were sensitive to the cytotoxic effects of UVA. This resulted in the formation of chromosome damage, as measured by the micronucleus assay, which this cell line was unable to repair. CONCLUSIONS: Owing to the nature of the repair defect in these cells, these results imply that DNA double-strand breaks are produced in cells following UVA irradiation, that the non-homologous end-joining repair pathway is involved in their repair and that they are produced with sufficient frequency to have biological significance.

Coping with multiple enemies: an integration of molecular and ecological perspectives.

How plants respond to attack by the range of herbivores and pathogens that confront them in the field is the subject of considerable research by both molecular biologists and ecologists. However, in spite of the shared focus of these two bodies of research, there has been little integration between them. We consider the scope for such integration, and how greater dialogue between molecular biologists and ecologists could advance understanding of plant responses to multiple enemies.

Beetle grazing reduces natural infection of Rumex obtusifolius by fungal pathogens.

In previously reported laboratory experiments, infection of Rumex obtusifolius by the rust fungus Uromyces rumicis was decreased on leaves which had prior herbivory by the beetle Gastrophysa viridula. In this paper we investigate whether this interaction is robust for natural infection by a variety of fungi in field experiments carried out in spring and autumn with plants given different levels of nitrogen fertilization. Grazing by G. viridula led to a decrease in lesion density of Ramularia rubella and Venturia rumicis in the spring and V. rumicis and U. rumicis in the autumn experiment. For V. rumicis and U. rumicis significant reductions in lesion density occurred on the undamaged leaves of damaged plants, compared with similar leaves on undamaged plants, suggesting systemic induced resistance. This induced resistance was usually independent of the amount of nitrogen fertilization, although the inhibitory effect of grazing on R. rubella in the spring and V. rumicis in the autumn experiment was enhanced by increasing nitrogen fertilization and was inhibited by increasing nitrogen fertilization for V. rumicis in the spring. In both experiments, the lesion density of V. rumicis was greater on leaves on which R. rubella was also present, and the presence of U. rumicis in the autumn experiment was linked to a similar but greater effect on V. rumicis lesion density. We found no evidence of induced resistance by fungi against fungi in these experiments. We highlight the complex interactions between inhibitory and facilitatory processes acting on leaf fungal infection. These results are compared with the proposed molecular mechanisms of induced resistance(s) and we consider the benefits of closer integration between molecular and ecological investigations of induced resistances that occur in the field.

Stratospheric ozone depletion, UV-B radiation and crop disease.

Ultraviolet-B radiation (UV-B: 290-315 nm) is expected to increase as the result of stratospheric ozone depletion. Within the environmental range, UV-B effects on host plants appear to be largely a function of photomorphogenic responses, while effects on fungal pathogens may include both photomorphogenesis and damage. The effects of increased UV-B on plant-pathogen interactions has been studied in only a few pathosystems, and have used a wide range of techniques, making generalisations difficult. Increased UV-B after inoculation tends to reduce disease, perhaps due to direct damage to the pathogen, although responses vary markedly between and within pathogen species. Using Septoria tritici infection of wheat as a model system, it is suggested that even in a species that is inherently sensitive to UV-B, the effects of ozone depletion in the field are likely to be small compared with the effects of variation in UV-B due to season and varying cloud. Increased UV-B before inoculation causes a range of effects in different systems, but an increase in subsequent disease is a common response, perhaps due to changes in host surface properties or chemical composition. Although it seems unlikely that most crop diseases will be greatly affected by stratospheric ozone depletion within the limits currently expected, the lack of a detailed understanding of the mechanisms by which UV-B influences plant-pathogen interactions in most pathosystems is a significant limit to such predictions.

The effect of an insect herbivore and a rust fungus individually, and combined in sequence, on the growth of two Rumex species.

The chrysomelid beetle Gastrophysa viridula and the rust fungus Urmnyces rumicis both occur on leaves of Rumex crispus and R. obtusifolius. We investigated the effect of beetle grazing or rust infection individually and when combined in sequence on the growth of their hosts in the field. Singly, beetle or rust reduced leaf area and plant biomass; the effect was greater on R. crispus, and rust caused greater damage than the beetle. Beetle grazing with subsequent rust infection caused damage no greater than that caused by rust alone, although on R. obtusifolius damage was greater than that from beetle grazing alone. Rust infection of R. obtusifolius with subsequent beetle grazing produced damage similar to that from other treatments; involving rust infection. In R. crispus this treatment produced the greatest reduction in biomass, The reductions in root and total plant weight from rust infection with subsequent beetle grazing were accurately predicted by a model including the damage produced by beetle and rust alone and the length of time each was present on the plant. This model also predicted accurately the damage to R. obtusifolius from the beetle followed by rust treatment, but over-estimated by up to 40% the damage to R. crispus. This can be explained mainly by an inhibition of rust infection by beetle grazing.

A small-scale controlled environment chamber for the investigation of the effects of pollutant gases on plants growing at cool or sub-zero temperature.

A new, small-scale controlled environment plant growth chamber is described, which permits accurately controlled temperatures from ambient (c. 20 degrees C) down to -5 degrees C in the day or -15 degrees C at night. The system also allows controlled injection of pollutant gases. The chamber is based on a modified commercial chest freezer with temperature control achieved through by-passing the cooling coils in the inner walls of the freezer. This by-pass is controlled by a BBC microcomputer which gives nominal temperature control to 0.1 degrees C over definable diurnal temperature profiles. Photoperiod is also controlled by the microcomputer. The chamber forms a virtually closed system to which pollutant gases are added by injection from low concentration cylinders, regulated using mass flow controllers. The accuracy with which pollutant concentrations are controlled is increased by filtration of the air-stream after passing over the experimental plants: i.e. a 'single-pass' system is adopted. In operation air temperatures across the chambers are controlled to within 1-1.5 degrees C of the pre-set value over the range 25 to -10 degrees C. Leaf temperatures are 2.5-3.0 degrees C above air temperatures during the day, with a total energy input of c. 180 Wm(-2) from two 400 W metal halide lamps. The lamps provide a photon fluence rate of 270+/-30 micromol quanta PAR m(-2) s(-1) at plant height. During the dark, leaf temperatures deviate less than 1.0 degrees C from air temperature. Soil temperatures are controlled separately from air temperature and are held above zero at all times. To date, fumigation, with NO(2) and SO(2) has been confined to the daytime. At pre-set concentrations of 20 nl litre(-1) variations during fumigation were less than 2.0-2.5 nl litre(-1) and 0.5 nl litre(-1), respectively.

Botrytis cinerea kills groundsel (Senecio vulgaris) infected by rust (Puccinia lagenophorae).

Inoculation of healthy groundsel (Senecio vulgaris L.) with Botrytis cinerea Pers at 1.24 × 106 conidia ml -1 caused 10% mortality, and only 40%, mortality when plants were abiotically wounded before inoculation. However, all plants previously infected by rust (Puccinia lagenophorae Cooke) died after inoculation with B. cinerea. Mortality was most rapid it plants were inoculated with B. cinerea as rust colonies first erupted through the host's epidermis. A reduction in the concentration of conidial inoculum increased time to death, and concentrations below 015 × 103 conidia ml-1 failed to cause 100 % mortality within the 32 d of the experiment. Death of plants was associated with the growth of B. cinerea into stem bases; a reduction in the density of rust lesions on leaves had no effect on the time from inoculation to the appearance of foliar symptoms, but increased time-to-kill. There was little variation m virulence between isolates of B. cinerea made from different groundsel or ragwort (Senecio jacobaea L.) populations. Similar interactions between biotrophic fungi and opportunistic pathogens have occasionally been noted previously; their importance in natural vegetation processes and possible relevance to the biocontrol of weeds is discussed.