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。     

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.     

UV-B and Mediterranean forest species: direct effects and ecological consequences

Experimental results from plants receiving elevated doses of UV-B radiation generally show that Mediterranean forest species are well protected against increases in UV-B radiation. Natural adaptations to water stress and excess light (elevated concentrations of UV-B screening compounds, leaf hairs, thick cuticle and epidermis), and UV-B responses (thickening of the cuticle, increase in carotenoids) may avoid or counter-balance UV-B radiation damage. This response confirms that Mediterranean forest vegetation is adapted to face oxidative stress factors, such as elevated tropospheric ozone concentrations, drought and high radiation, including UV-B. Nevertheless, in the long term, species-specific and season-specific differential responses in growth, physiology, phenology and reproductive behaviour may alter the interactions between species and lead to slow but important changes in ecosystem structure and function.     

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.     

Effects of ultraviolet-B radiation (UV-B) on growth and physiology of the dune grassland species Calamagrostis epigeios

Seedlings of Calamagrostis epigeios were exposed to four levels of UV-B radiation (280-320 nm), simulating up to 44% reduction of stratospheric ozone concentration during summertime in The Netherlands, to determine the response of this plant species to UV-B irradiation. After six weeks of UV-B treatment, total biomass of all UV-B treated plants was higher, compared to plants that had received no UV-B radiation. The increase of biomass did not appear to be the result of a stimulation of net photosynthesis. Also, transpiration rate and water use efficiency were not altered by UV-B at any exposure level. Pigment analysis of leaf extracts showed no effect of enhanced UV-B radiation on chlorophyll content and accumulation of UV absorbing pigments. UV-B irradiance, however, did reduce the transmittance of visible light (400-700 nm) of intact attached leaves, suggesting a change in anatomical characteristics of the leaves. Additionally, the importance of including an ambient UV-B treatment in indoor experiments is discussed.     

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.     

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.     

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.     

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.     

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.     

Climate change: potential effects of increased atmospheric carbon dioxide (CO2), ozone (O3), and ultraviolet-B (UV-B) radiation on plant diseases

Continued world population growth results in increased emission of gases from agriculture, combustion of fossil fuels, and industrial processes. This causes changes in the chemical composition of the atmosphere. Evidence is emerging that increased solar ultraviolet-B (UV-B) radiation is reaching the earth's atmosphere, due to stratospheric ozone depletion. Carbon dioxide (CO(2)), ozone (O(3)) and UV-B are individual climate change factors that have direct biological effects on plants. Such effects may directly or indirectly affect the incidence and severity of plant diseases, caused by biotic agents. Carbon dioxide may increase plant canopy size and density, resulting in a greater biomass of high nutritional quality, combined with a much higher microclimate relative humidity. This would be likely to promote plant diseases such as rusts, powdery mildews, leaf spots and blights. Inoculum potential from greater overwintering crop debris would also be increased. Ozone is likely to have adverse effects on plant growth. Necrotrophic pathogens may colonize plants weakened by O(3) at an accelerated rate, while obligate biotroph infections may be lessened. Ozone is unlikely to have direct adverse effects on fungal pathogens. Ozone effects on plant diseases are host plant mediated. The principal effects of increased UV-B on plant diseases would be via alterations in host plants. Increased flavonoids could lead to increased diseased resistance. Reduced net photosynthesis and premature ripening and senescence could result in a decrease in diseases caused by biotrophs and an increase in those caused by necrotrophs. Microbial plant pathogens are less likely to be adversely affected by CO(2), O(3) and UV-B than are their corresponding host plants. Changes in host plants may result in expectable alterations of disease incidence, depending on host plant growth stages and type of pathogen. Given the importance of plant diseases in world food and fiber production, it is essential to begin studying the effects of increased CO(2), O(3) and UV-B (and other climate change factors) on plant diseases. We know very little about the actual impacts of climate change factors on disease epidemiology. Epidemiologists should be encouraged to consider CO(2), O(3) and UV-B as factors in their field studies.     

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.     

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.     

Effect of UV-B exposure on eggs and embryos of patagonian anurans and evidence of photoprotection

In Patagonia, enhanced levels of ultraviolet radiation (UVR) are affecting not only the southernmost territories but the northern as well, and there is a growing body of evidence indicating effects on terrestrial and aquatic communities. Anurans occur along a wide geographical gradient in Patagonia and are common in shallow waters, usually breeding during spring when seasonal UV-B radiation is at its highest level. In this paper we analyse the effective irradiances (I) experienced in nature by eggs and embryos of the patagonian anurans Pleurodema bufoninum, Pleurodema thaul and Bufo spinulosus papillosus, considering water and egg jelly optical properties. Besides, we determine photoprotective pigments in eggs and embryos of the different species, and ontogenetic shifts in pigment concentration. Finally, we studied the effects of natural and enhanced levels of UV-B on eggs of P. bufoninum aiming to evaluate the role of the jelly envelope in screening the damaging radiation. The evidence gathered in this work shows that natural and enhanced UV-B levels did not induce significant changes on the survivorship but incremented the occurrence of malformations. Also, the species experiencing high levels of exposure presented high concentrations of melanin suggesting that exposure to sunlight triggers photoprotection by pigments. Collectively our results suggest that the studied species bear certain levels of adaptation to cope with high ambient UVR conferred by environmental and biological factors.     

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.     

Damage to DNA caused by UV-B radiation in the desert cyanobacterium Scytonema javanicum and the effects of exogenous chemicals on the process

Radiation with UV-B increased the damage to DNA in Scytonema javanicum, a desert-dwelling soil microorganism, and the level of damage varied with the intensity of UV-B radiation and duration of exposure. Production of reactive oxygen species (ROS) also increased because of the radiation. Different exogenous chemicals (ascorbate acid, ASC; N-acetylcysteine, NAC; glyphosate, GPS; and 2-methyl-4-chlorophenoxyacetic acid, MCPA-Na) differed in their effect on the extent of DNA damage and ROS production: whereas NAC and ASC protected the DNA from damage and resulted in reduced ROS production, the herbicides (GPS and MCPA-Na) increased the extent of damage, lowered the rate of photosynthesis, and differed in their effect on ROS production. The chemicals probably have different mechanisms to exercise their effects: NAC and ASC probably function as antioxidant agents or as precursors of other antioxidant molecules that protect the DNA and photosynthetic apparatus directly from the ROS produced as a result of UV-B radiation, and GPS and MCPA-Na probably disrupt the normal metabolism in S. javanicum to induce the leaking of ROS into the photosynthetic electron transfer pathway following UV-B radiation, and thereby damage the DNA. Such mechanisms have serious implications for the use of environment-friendly herbicides, which, because they can destroy DNA, may prove harmful to soil microorganisms.     

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.     

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.     

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.