Linkages between stratospheric ozone, UV radiation and climate change and their implications for terrestrial ecosystems.

Exposure of plants and animals to ultraviolet-B radiation (UV-B; 280-315 nm) is modified by stratospheric ozone dynamics and climate change. Even though stabilisation and projected recovery of stratospheric ozone is expected to curtail future increases in UV-B radiation at the Earth's surface, on-going changes in climate are increasingly exposing plants and animals to novel combinations of UV-B radiation and other climate change factors (e.g., ultraviolet-A and visible radiation, water availability, temperature and elevated carbon dioxide). Climate change is also shifting vegetation cover, geographic ranges of species, and seasonal timing of development, which further modifies exposure to UV-B radiation. Since our last assessment, there has been increased understanding of the underlying mechanisms by which plants perceive UV-B radiation, eliciting changes in growth, development and tolerances of abiotic and biotic factors. However, major questions remain on how UV-B radiation is interacting with other climate change factors to modify the production and quality of crops, as well as important ecosystem processes such as plant and animal competition, pest-pathogen interactions, and the decomposition of dead plant matter (litter). In addition, stratospheric ozone depletion is directly contributing to climate change in the southern hemisphere, such that terrestrial ecosystems in this region are being exposed to altered patterns of precipitation, temperature and fire regimes as well as UV-B radiation. These ozone-driven changes in climate have been implicated in both increases and reductions in the growth, survival and reproduction of plants and animals in Antarctica, South America and New Zealand. In this assessment, we summarise advances in our knowledge of these and other linkages and effects, and identify uncertainties and knowledge gaps that limit our ability to fully evaluate the ecological consequences of these environmental changes on terrestrial ecosystems.

Rapid modulation of ultraviolet shielding in plants is influenced by solar ultraviolet radiation and linked to alterations in flavonoids.

The accumulation of ultraviolet (UV)-absorbing compounds (flavonoids and related phenylpropanoids) and the resultant decrease in epidermal UV transmittance (TUV ) are primary protective mechanisms employed by plants against potentially damaging solar UV radiation and are critical components of the overall acclimation response of plants to changing solar UV environments. Whether plants can adjust this UV sunscreen protection in response to rapid changes in UV, as occurs on a diurnal basis, is largely unexplored. Here, we use a combination of approaches to demonstrate that plants can modulate their UV-screening properties within minutes to hours, and these changes are driven, in part, by UV radiation. For the cultivated species Abelmoschus esculentus, large (30-50%) and reversible changes in TUV occurred on a diurnal basis, and these adjustments were associated with changes in the concentrations of whole-leaf UV-absorbing compounds and several quercetin glycosides. Similar results were found for two other species (Vicia faba and Solanum lycopersicum), but no such changes were detected in Zea mays. These findings reveal a much more dynamic UV-protection mechanism than previously recognized, raise important questions concerning the costs and benefits of UV-protection strategies in plants and have practical implications for employing UV to enhance crop vigor and quality in controlled environments.

Molecular and physiological effects of environmental UV radiation on fungal conidia.

Conidia are specialized structures produced at the end of the asexual life cycle of most filamentous fungi. They are responsible for fungal dispersal and environmental persistence. In pathogenic species, they are also involved in host recognition and infection. Conidial production, survival, dispersal, germination, pathogenicity and virulence can be strongly influenced by exposure to solar radiation, although its effects are diverse and often species dependent. UV radiation is the most harmful and mutagenic waveband of the solar spectrum. Direct exposure to solar radiation for a few hours can kill conidia of most fungal species. Conidia are killed both by solar UV-A and UV-B radiation. In addition to killing conidia, which limits the size of the fungal population and its dispersion, exposures to sublethal doses of UV radiation can reduce conidial germination speed and virulence. The focus of this review is to provide an overview of the effects of solar radiation on conidia and on the major systems involved in protection from and repair of damage induced by solar UV radiation. The efforts that have been made to obtain strains of fungi of interest such as entomopathogens more tolerant to solar radiation will also be reviewed.

Rediscovering leaf optical properties: New insights into plant acclimation to solar UV radiation.

The accumulation of UV-absorbing compounds (flavonoids and other phenylpropanoid derivatives) and resultant decrease in the UV transmittance of the epidermis in leaves (TUV), is a primary protective mechanism against the potentially deleterious effects of UV radiation and is a critical component of the overall acclimation response of plants to changing UV environments. Traditional measurements of TUV were laborious, time-consuming and destructive or invasive, thus limiting their ability to efficiently make multiple measurements of the optical properties of plants in the field. The development of rapid, nondestructive optical methods of determining TUV has permitted the examination of UV optical properties of leaves with increased replication, on a finer time scale, and enabled repeated sampling of the same leaf over time. This technology has therefore allowed for studies examining acclimation responses to UV in plants in ways not previously possible. Here we provide a brief review of these earlier studies examining leaf UV optical properties and some of their important contributions, describe the principles by which the newer non-invasive measurements of epidermal UV transmittance are made, and highlight several case studies that reveal how this technique is providing new insights into this UV acclimation response in plants, which is far more plastic and dynamic than previously thought.

Growth under visible light increases conidia and mucilage production and tolerance to UV-B radiation in the plant pathogenic fungus Colletotrichum acutatum.

Light conditions can influence fungal development. Some spectral wavebands can induce conidial production, whereas others can kill the conidia, reducing the population size and limiting dispersal. The plant pathogenic fungus Colletotrichum acutatum causes anthracnose in several crops. During the asexual stage on the host plant, Colletototrichum produces acervuli with abundant mucilage-embedded conidia. These conidia are responsible for fungal dispersal and host infection. This study examined the effect of visible light during C. acutatum growth on the production of conidia and mucilage and also on the UV tolerance of these conidia. Conidial tolerance to an environmentally realistic UV irradiance was determined both in conidia surrounded by mucilage on sporulating colonies and in conidial suspension. Exposures to visible light during fungal growth increased production of conidia and mucilage as well as conidial tolerance to UV. Colonies exposed to light produced 1.7 times more conidia than colonies grown in continuous darkness. The UV tolerances of conidia produced under light were at least two times higher than conidia produced in the dark. Conidia embedded in the mucilage on sporulating colonies were more tolerant of UV than conidia in suspension that were washed free of mucilage. Conidial tolerance to UV radiation varied among five selected isolates.

Conidial pigmentation is important to tolerance against solar-simulated radiation in the entomopathogenic fungus Metarhizium anisopliae.

The importance of conidial pigmentation to solar UV radiation tolerance in the entomopathogenic fungus Metarhizium anisopliae var. anisopliae, was estimated by comparing the effects of exposure to simulated solar UV radiation on the wild-type parent strain U.S. Department of Agriculture (USDA)-Agricultural Research Service (ARS) Collection of Entomopathogenic Fungal Cultures (ARSEF) 23, which has dark green conidia, and three groups of color mutants with yellow, purple and white conidia. The comparisons included inactivation levels and the kinetics of germination of conidia exposed or not exposed to simulated solar UV radiation. In addition to significantly inactivating the conidia of different mutants, exposure to radiation delayed for several hours the germination of surviving conidia of the wild type and all mutants. In general, mutants with white conidia were more sensitive to simulated solar UV radiation than mutants with purple conidia, which were more sensitive than mutants with yellow conidia, which in turn were more sensitive than the green wild strain. A significant variation in tolerance to simulated solar radiation was observed among mutants within each color group, particularly among mutants with yellow conidia. Revertants with green conidia, DWR 179 and DWR 176, were obtained from the very sensitive UV mutants DWR 148 (yellow conidia) and DWR 149 (purple conidia), respectively. These revertants had levels of tolerance to simulated solar UV radiation similar to those of the wild-type ARSEF 23. This observation is strong evidence of the importance of green conidial pigmentation for tolerance to simulated solar UV radiation, a factor that could be manipulated to produce M. anisopliae strains with more tolerance to solar UV radiation.

Plant responses to current solar ultraviolet-B radiation and to supplemented solar ultraviolet-B radiation simulating ozone depletion: an experimental comparison.

Field experiments assessing UV-B effects on plants have been conducted using two contrasting techniques: supplementation of solar UV-B with radiation from fluorescent UV lamps and the exclusion of solar UV-B with filters. We compared these two approaches by growing lettuce and oat simultaneously under three conditions: UV-B exclusion, near-ambient UV-B (control) and UV-B supplementation (simulating a 30% ozone depletion). This permitted computation of "solar UV-B" and "supplemental UV-B" effects. Microclimate and photosynthetically active radiation were the same under the two treatments and the control. Excluding UV-B changed total UV-B radiation more than did supplementing UV-B, but the UV-B supplementation contained more "biologically effective" shortwave radiation. For oat, solar UV-B had a greater effect than supplemental UV-B on main shoot leaf area and main shoot mass, but supplemental UV-B had a greater effect on leaf and tiller number and UV-B-absorbing compounds. For lettuce, growth and stomatal density generally responded similarly to both solar UV-B and supplemented UV-B radiation, but UV-absorbing compounds responded more to supplemental UV-B, as in oat. Because of the marked spectral differences between the techniques, experiments using UV-B exclusion are most suited to assessing effects of present-day UV-B radiation, whereas UV-B supplementation experiments are most appropriate for addressing the ozone depletion issue.

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.