Possible impacts of changes in UV-B radiation on North American trees and forests

Approximately 35 species representing 14 tree genera have been evaluated for responses to UV-B radiation in North America. The best representation has been in the conifers where some 20 species representing three genera have been studied. Overall, about 1/3 of these have demonstrated some deleterious response to UV-B. However, most negative impacts have been observed under controlled environment conditions where sensitivity may be enhanced. Therefore, it seems unlikely that expected levels of ozone depletion will result in direct losses in productivity. However, the role that ambient or enhanced levels of UV-B may play in forest ecosystem processes is more difficult to access. One possible indirect response of forests to changes in UV-B radiation levels could be via alterations in plant secondary metabolites. Increases in phenolics and flavonoids that enhance epidermal UV-screening effectiveness may also influence leaf development, water relations or ecosystem processes such as plant-herbivore interactions or decomposition.     

Assessment of the impact of rising carbon dioxide and other potential climate changes on vegetation

The projected doubling of current levels of atmospheric carbon dioxide concentration ([CO(2)]) during the next century along with increases in other radiatively active gases have led to predictions of increases in global air temperature and shifts in precipitation patterns. Additionally, stratospheric ozone depletion may result in increased ultraviolet-B (UV-B) radiation incident at the Earth's surface in some areas. Since these changes in the Earth's atmosphere may have profound effects on vegetation, the objectives of this paper are to summarize some of the recent research on plant responses to [CO(2)], temperature and UV-B radiation. Elevated [CO(2)] increases photosynthesis and usually results in increased biomass, and seed yield. The magnitude of these increases and the specific photosynthetic response depends on the plant species, and are strongly influenced by other environmental factors including temperature, light level, and the availability of water and nutrients. While elevated [CO(2)] reduces transpiration and increases photosynthetic water-use efficiency, increasing air temperature can result in greater water use, accelerated plant developmental rate, and shortened growth duration. Experiments on UV-B radiation exposure have demonstrated a wide range of photobiological responses among plants with decreases in photosynthesis and plant growth among more sensitive species. Although a few studies have addressed the interactive effects of [CO(2)] and temperature on plants, information on the effects of UV-B radiation at elevated [CO(2)] is scarce. Since [CO(2)], temperature and UV-B radiation may increase concurrently, more research is needed to determine plant responses to the interactive effects of these environmental variables.     

Effects of the ultraviolet-B radiation (UV-B) on conifers: a review

The current knowledge on conifer responses to enhanced ultraviolet-B (UV-B) radiation is mainly based on greenhouse or growth chamber experiments of one growing season in duration. However, the biomass losses observed in greenhouses do not occur in field-grown trees in their natural habitats. Moreover, the majority of the 20 conifer species studied have been 1-year-old seedlings, and no studies have been undertaken on mature trees. Fully grown needles, with their glaucous waxy surfaces and thick epidermal cells with both soluble and wall-bound UV-B screening metabolites, are well protected against UV-B radiation. However, it is not known whether these are sufficient protectants in young emerging needles or during the early spring period of high UV-B levels reflected from snow. In order to understand all the mechanisms that result in the protection of conifer needles against UV-B radiation, future research should focus on the epidermal layer, separating the waxes, cuticle and epidermal and hypodermal cells. Parallel studies should consist of wall-bound and soluble secondary metabolite analysis, antioxidant measurements and microscopic observations.     

The effect of reciprocal treatments with ozone and ultraviolet-B radiation on photosynthesis and growth of perennial grass Elymus athericus

The impact on plant growth of the simultaneously changing factors of the global climate, rising tropospheric O3 concentrations and increasing UV-B radiation fluxes, has been tested in a combined glasshouse and growth chamber experiment. The saltmarsh grass species Elymus athericus was sequentially fumigated for two weeks with O3 and for another two weeks irradiated with UV-B (vv). Exposure to elevated UV-B did not negatively affect photosynthesis or plant growth. Fumigation with O3 had a depressing effect on net photosynthesis, the number and biomass of flowers, the number of leaves and the number of shoots. O3-induced damage only was observed in plants which had been fumigated during the last two weeks of the experiment. Since interactive responses were not observed, results suggest different primary target sites for O3 and UV-B within the plant.     

The effect of elevated UV-B (280-320 nm) radiation levels on Silene vulgaris: a comparison between a highland and a lowland population

Highland (altitude 1600 m above sea level) and lowland (altitude -2 m below sea level) populations of the perennial herb Silene vulgaris (Moench) Garcke, were tested on their response to elevated levels of UV-B radiation. Highland populations typically receive high natural UV-B fluxes, whereas lowland populations receive a lower natural UV-B dose. Adaptation to high UV-B levels of the highland population is to be expected. Experimental comparison of growth rates, gas exchange rates, transpiration and biochemical parameters using adult plants as well as seedlings did not show a difference in the response to elevated UV-B levels between the two populations. Individuals of both populations were relatively insensitive to elevated UV-B radiation. The response of alpine and lowland populations of Silene vulgaris is discussed in relation to the dispersal of this species after the last ice age.     

Long-term exposure to enhanced UV-B radiation has no significant effects on growth or secondary compounds of outdoor-grown Scots pine and Norway spruce seedlings

The effects of long-term enhanced UV-B radiation on growth and secondary compounds of two conifer species were studied in an outdoor experiment. Scots pine (Pinus sylvestris) seedlings were exposed for two growing seasons and Norway spruce (Picea abies) seedlings for three growing seasons to supplemental UV-B radiation, corresponding to a 30% increase in ambient UV-B radiation. The experiment also included appropriate controls for ambient and increased UV-A radiation. Enhanced UV-B did not affect the growth of the conifer seedlings. In addition, neither the concentrations of terpenes and phenolics in the needles nor the concentrations of terpenes in the wood were affected. However, in the UV-A control treatment the concentrations of diterpenes in the wood of Scots pine decreased significantly compared to the ambient control. Apparently, a small increase in UV-B radiation has no significant effects on the secondary compounds and growth of Scots pine and Norway spruce seedlings.     

The effect of elevated UV-B (280–320 nm) radiation levels on Silene vulgaris: A comparison between a highland and a lowland population

Highland (altitude 1600 m above sea level) and lowland (altitude −2 m below sea level) populations of the perennial herb Silene vulgaris (Moench) Garcke, were tested on their response to elevated levels of UV-B radiation. Highland populations typically receive high natural UV-B fluxes, whereas lowland populations receive a lower natural UV-B dose. Adaptation to high UV-B levels of the highland population is to be expected. Experimental comparison of growth rates, gas exchange rates, transpiration and biochemical parameters using adult plants as well as seedlings did not show a difference in the response to elevated UV-B levels between the two populations. Individuals of both populations were relatively insensitive to elevated UV-B radiation. The response of alpine and lowland populations of Silene vulgaris is discussed in relation to the dispersal of this species after the last ice age.     

增强的UV-B辐射对陆地生态系统的影响

本文根据最近的文献,回顾了ＵＶ－ Ｂ 辐射对陆地生态系统的影响。ＵＶ－ Ｂ 辐射影响植物物候、形态和次生代谢,从而改变生态系统的物种结构、竞争性平衡、食物链、植物病原体、物质循环、真菌移殖与叶片分解。植物物候、形态和次生代谢的变化是ＵＶ－ Ｂ 辐射调控影响生态系统的重要途径,具有重要的生态学意义。虽然我们已经适当了解了ＵＶ－ Ｂ 辐射对植物作用的机理,但由于温室内植物个体水平的短期响应与野外条件下生态系统的长期响应具有明显的差异,到目前为止还不能预测ＵＶ－ Ｂ辐射影响生态系统的精确后果,有时甚至连变化方向也不能预测。因此,必须强调生态系统对ＵＶ－ Ｂ 辐射长期响应的野外研究的重要性。     

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.     

Effects of ultraviolet-B radiation on behaviour and growth of three species of amphibian larvae

Effects of ultraviolet-B (UV-B) radiation on amphibian embryos have been investigated in a number of studies, but the effects on larvae have received less attention. We investigated the effects of UV-B radiation on the behaviour and growth of larvae of three amphibians (Rana arvalis, Rana temporaria and Bufo bufo) in two different experiments. First, we tested whether larvae of the three species actively avoid UV-B exposure if given a choice. We found no evidence for active avoidance of UV-B or changes in activity in the presence of UV-B in any of the species. Second, we assessed the effects of natural (1.25 kJm(-2)) and enhanced (1.58 kJm(-2)) UV-B radiation on the survival and growth of the three species and found that the exposure to UV-B radiation did not have any effect on survival rates of any of the species. However, UV-B radiation had a positive effect on the growth of R. arvalis and R. temporaria, whereas the growth of B. bufo tadpoles was unaffected by the UV-B treatments. Our results suggest that a short-term exposure to UV-B radiation does not induce any UV-B avoidance behaviour in tadpoles of these three species. Furthermore, unlike some previous studies, the results suggest that the young tadpoles of these species are not negatively affected by UV-B radiation. In fact, our results demonstrate that a moderate amount of UV-B radiation enhance tadpole growth rates in two of the three species.     

UV-B radiation and acclimation in timberline plants

Research has shown that some plants respond to enhanced UV-B radiation by producing smaller and thicker leaves, by increasing the thickness of epidermis and concentration of UV-B absorbing compounds of their surface layers and activation of the antioxidant defence system. The response of high-altitude plants to UV-B radiation in controlled conditions is often less pronounced compared to low-altitude plants, which shows that the alpine timberline plants are adapted to UV-B. These plants may have a simultaneous co-tolerance for several stress factors: acclimation or adaptation to the harsh climate can also increase tolerance to UV-B radiation, and vice versa. On the other hand, alpine timberline plants of northern latitudes may be less protected against increasing UV-B radiation than plants from more southern latitudes and higher elevations due to harsh conditions and weaker preadaptation resulting from lower UV-B radiation exposure. It is evident that more long-term experimental field research is needed in order to study the interaction of climate, soil and UV-B irradiance on the timberline plants.     

The effects of solar UV-B radiation on embryonic mortality and development in three boreal anurans (Rana temporaria, Rana arvalis and Bufo bufo).

Many species of amphibians have experienced population and range reductions. It has been hypothesized that sensitivity to UV-B may contribute to the population declines of some amphibian species. We performed field experiments to measure the effects of solar UV-B on the hatching success of three Finnish anuran species, the common frog (Rana temporaria), moor frog (Rana arvalis) and common toad (Bufo bufo). Further, the effects of natural UV-B radiation on survival of the tadpoles of the same three species of anurans were tested. A significant percentage of R. temporaria and B. bufo embryos survived when exposed to and protected from solar UV-B and hatching success was not affected by solar radiation. Elimination of solar UV-B significantly increased the hatching success of R. arvalis, but embryonic mortality was high in both treatments. The data indicates that under natural conditions, solar UV-B radiation influences embryo survival in R. arvalis, but has no effect on R. temporaria and B. bufo. Solar UV-B radiation had no effect on R. temporaria and R. arvalis tadpoles, but elimination of UV-B significantly increased survival of B. bufo tadpoles. It seems that ambient UV-radiation levels have no effect on R. temporaria but may affect R. arvalis and B. bufo at different developmental stages.     

Depletion of stratospheric ozone over the Antarctic and Arctic: responses of plants of polar terrestrial ecosystems to enhanced UV-B, an overview

Depletion of stratospheric ozone over the Antarctic has been re-occurring yearly since 1974, leading to enhanced UV-B radiation. Arctic ozone depletion has been observed since 1990. Ozone recovery has been predicted by 2050, but no signs of recovery occur. Here we review responses of polar plants to experimentally varied UV-B through supplementation or exclusion. In supplementation studies comparing ambient and above ambient UV-B, no effect on growth occurred. UV-B-induced DNA damage, as measured in polar bryophytes, is repaired overnight by photoreactivation. With UV exclusion, growth at near ambient may be less than at below ambient UV-B levels, which relates to the UV response curve of polar plants. UV-B screening foils also alter PAR, humidity, and temperature and interactions of UV with environmental factors may occur. Plant phenolics induced by solar UV-B, as in pollen, spores and lignin, may serve as a climate proxy for past UV. Since the Antarctic and Arctic terrestrial ecosystems differ essentially, (e.g. higher species diversity and more trophic interactions in the Arctic), generalization of polar plant responses to UV-B needs caution.     

The impact of UV-B radiation and ozone on terrestrial vegetation

Although terrestrial vegetation has been exposed to UV-B radiation and ozone over the course of evolutionary history, it is essential to view the effects on vegetation of changing levels of these factors in the context of other features of climate change, such as increasing CO(2) levels and changes in temperature and precipitation patterns. Much of our understanding of the impacts of increased UV-B and ozone levels has come from studies of the effects of each individual factor. While such information may be relevant to a wider understanding of the roles that these factors may play in climate change, experience has shown that the interactions of environmental stresses on vegetation are rarely predictable. A further limitation on the applicability of such information results from the methodologies used for exposing plants to either factor. Much of our information comes from growth chamber, greenhouse or field studies using experimental protocols that made little or no provision for the stochastic nature of the changes in UV-B and ozone levels at the earth's surface, and hence excluded the roles of repair mechanisms. As a result, our knowledge of dose-response relationships under true field conditions is both limited and fragmentary, given the wide range of sensitivities among species and cultivars. Adverse effects of increased levels of either factor on vegetation are qualitatively well established, but the quantitative relationships are far from clear. In both cases, sensitivity varies with stage of plant development. At the population and community levels, differential responses of species to either factor has been shown to result in changes in competitiveness and community structure. At the mechanistic level, ozone generally inhibits photosynthetic gas exchange under both controlled and field conditions, and although UV-B is also inhibitory in some species under controlled conditions, others appear to be indifferent, particularly in the field. Both factors affect metabolism; a common response is increased secondary metabolism leading to the accumulation of phenolic compounds that, in the case of UV-B, offer the leaf cell some protection from radiation. Virtually no information is available about the effects of simultaneous or sequential exposures. Since both increased surface UV-B and ozone exposures have spatial and temporal components, it is important to evaluate the different scenarios that may occur, bearing in mind that elevated daytime ozone levels will attenuate the UV-B reaching the surface to some extent. The experimentation needed to acquire unequivocal effects data that are relevant to field situations must therefore be carried out using technologies and protocols that focus on quantification of the interactions of UV-B and ozone themselves and their interactions with other environmental factors.     

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.     

Growth and defense in deciduous trees and shrubs under UV-B

Reflection by waxy or resinous surface structures and hairs, repair reactions of biomolecules and induction of different sheltering components provide the means of plant protection from harmful solar UV-B radiation. Secondary products, especially flavonoids and phenolic acids as defense components are also important in plant tolerance to UV-B, fulfilling the dual role as screens that reduce UV-B penetration in plant tissues, and as antioxidants protecting from damage by reactive oxidant species. Plants are sensitive to UV-B radiation, and this sensitivity can be even more clone-specific than species-specific. The results available in the literature for deciduous trees and shrubs indicate that UV-B radiation may affect several directions in the interaction of woody species with biotic (herbivores) and abiotic (CO2 and nutrition) factors depending on the specific interaction in question. These multilevel interactions should have moderate ecological significance via the overall changed performance of woody species and shrubs.     

Air pollution, forest condition and forest decline in Southern Europe: an overview

Over the last decades much of the work on the impact of air pollution on forests in Europe has concentrated on central and northern countries. The southern part of Europe has received far less attention, although air pollutants-especially the photochemical ones-can reach concentrations likely to have adverse effects on forest vegetation. Although international forest condition surveys present serious problems where data consistency is concerned, they reveal considerable year-by-year species-specific fluctuations rather than a large-scale forest decline. Cases of obvious decline related to environmental factors are well circumscribed: (1) the deterioration of some coastal forests due to the action of polluted seaspray; (2) the deterioration of reforestation projects, especially conifers, mainly due to the poor ecological compatibility between species and site; and (3) the decline of deciduous oaks in southern Italy and of evergreen oaks in the Iberian peninsula apparently due to the interaction of climate stresses and pests and diseases. However, besides obvious deterioration, changes in environmental factors can provoke situations of more subtle stress. The most sensitive stands are Mediterranean conifer forests and mesophile forests of the Mediterranean-montane plane growing at the edges of the natural ecological distribution. Evergreen sclerophyllous forests appear less sensitive to variations in climatic parameters, since they can adapt quite well to both drought and the action of UV-B rays. Several experiments were carried out to test the sensitivity of Mediterranean forest species to air pollutants. Most of those experiments used seedlings of different species treated with pollutant concentrations too high to be realistic, so it is difficult to derive adequate information on the response of adult trees in field conditions. Ozone has been proved to cause foliar injury in a variety of native forest species in different Southern European countries, while the effects of other pollutants (e.g. nitrogen, sulphur, acidic deposition) are less obvious and likely to be very localized. In the case of ozone, visible symptoms were almost completely missed by large-scale surveys and-at the same time-non-visible symptoms are suspected to be even more widespread than the visible ones. Owing to this and to the complex relationships existing between species sensitivity, ozone exposure and doses, length of the vegetative periods, influence of climatic and edaphic condition on the tree's response, the impacted areas are yet to be identified. Therefore, the large-scale impact of air pollutants on the forests of Southern Europe remains largely unknown, until more specific investigations are carried out.     

Growth, phenology and reproduction of an arid-environment winter ephemeral Dimorphotheca pluvialis in response to combined increases in CO2 and UV-B radiation

The winter ephemeral Dimorphotheca pluvialis was grown in open-top chambers in ambient or elevated CO2 (350 or 650 micromol mol(-1)), combined with ambient (2.39 to 7.59 kJ m(-2) d(-1)) or increased (4.94 to 11.13 kJ m(-2) d(-1)) UV-B radiation. Net CO2 assimilation rate and leaf water use efficiency increased in elevated CO2, but increased UV-B did not affect gas exchange. Leaf biomass was greater under increased UV-B, but vegetative biomass was unaffected in elevated CO2. Initiation of reproduction was delayed, and proportional investment in reproductive biomass at harvest was reduced in elevated CO2. Increased UV-B stimulated reproduction, particularly in ambient CO2, but also in elevated CO2 at a later stage. Changes in reproductive phenology and prolonged development in elevated CO2 during the stressful late season could indirectly be detrimental to reproductive success of D. pluvialis, but stimulation of reproduction by enhanced UV-B may to some extent mitigate this.     

Phenogenetic response of silver birch populations and half-sib families to elevated ozone and ultraviolet-B radiation at juvenile age

Phenogenetic response of silver birch populations and half-sib families to separate and combined elevated ozone (O₃) concentrations and ultraviolet-B (UV-B) radiation dozes was studied at juvenile age in the climatic chambers. Significant population and family effects were found for seedling height, lamina width, and leaf damage. The exposure to UV-B radiation decreased genetic variation at the stage of seed germination. Complex exposure to UV-B and O₃ caused an increase of genetic variation at the stage of intensive seedling growth: seedling height genetic variation in separate treatments increased from 23.7–38.6 to 33.7–65.7%, the increase for lamina width was from 10.2–13.9 to 13.6–31.8%. Different populations and families demonstrated differing response to elevated complex UV-B and O₃ exposure. Changes of genetic intra-population variation were population-specific. Such changes in genetic variation under the impact of stressors can alter adaptation, stability, and competitive ability of regenerating populations in a hardly predictive way.     

Mycorrhizal mediation of plant response to atmospheric change: Air quality concepts and research considerations

The term 'global climate change' encompasses many physical and chemical changes in the atmosphere that have been induced by anthropogenic pollutants. Increases in concentrations of CO2 and CH4 enhance the 'greenhouse effect' of the atmosphere and may contribute to changes in temperature and precipitation patterns at the earth's surface. Nitrogen oxides and SO2 are phytotoxic and also react with other pollutants to produce other phytotoxins in the troposphere such as O3 and acidic substances. However, release of chlorofluorocarbons into the atmosphere may cause depletion of stratospheric O3, increasing the transmittance of ultraviolet-B (UV-B) radiation to the earth's surface. Increased intensities of UV-B could affect plants and enhance photochemical reactions that generate some phytotoxic pollutants. The role of mycorrhizae in plant responses to such stresses has received little attention. Although plans for several research programs have acknowledged the importance of drought tolerance and soil fertility in plant responses to atmospheric stresses, mycorrhizae are rarely targeted to receive specific investigation. Most vascular land plants form mycorrhizae, so the role of mycorrhizae in mediating plant responses to atmospheric change may be an important consideration in predicting effects of atmospheric changes on plants in managed and natural ecosystems.