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Ultraviolet radiation (UV) is a minor fraction of the solar spectrum reaching the ground surface. In this assessment we summarize the results of previous work on the effects of the UV-B component (280-315 nm) on terrestrial ecosystems, and draw attention to important knowledge gaps in our understanding of the interactive effects of UV radiation and climate change. We highlight the following points: (i) The effects of UV-B on the growth of terrestrial plants are relatively small and, because the Montreal Protocol has been successful in limiting ozone depletion, the reduction in plant growth caused by increased UV-B radiation in areas affected by ozone decline since 1980 is unlikely to have exceeded 6%. (ii) Solar UV-B radiation has large direct and indirect (plant-mediated) effects on canopy arthropods and microorganisms. Therefore, trophic interactions (herbivory, decomposition) in terrestrial ecosystems appear to be sensitive to variations in UV-B irradiance. (iii) Future variations in UV radiation resulting from changes in climate and land-use may have more important consequences on terrestrial ecosystems than the changes in UV caused by ozone depletion. This is because the resulting changes in UV radiation may affect a greater range of ecosystems, and will not be restricted solely to the UV-B component. (iv) Several ecosystem processes that are not particularly sensitive to UV-B radiation can be strongly affected by UV-A (315-400 nm) radiation. One example is the physical degradation of plant litter. Increased photodegradation (in response to reduced cloudiness or canopy cover) will lead to increased carbon release to the atmosphere via direct and indirect mechanisms.

Assuntos

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The southern part of Tierra del Fuego, in the southernmost tip of South America, is covered by dense Nothofagus spp. forests and Sphagnum-dominated peat bogs, which are subjected to the influence of ozone depletion and to increased levels of solar ultraviolet-B radiation (UV-B). Over the last 5 years we have studied some of the biological impacts of solar UV-B on natural ecosystems of this region. We have addressed two general problems: (i) do the fluctuations in UV-B levels under the influence of the Antarctic ozone 'hole' have any measurable biological impact, and (ii) what are the long-term effects of solar (ambient) UV-B on the Tierra del Fuego ecosystems? In this paper, we provide an overview of the progress made during the first 4 years of the project. We highlight and discuss the following results: (1) ambient UV-B has subtle but significant inhibitory effects on the growth of herbaceous and graminoid species of this region (growth reduction < or = 12%), whereas no consistent inhibitory effects could be detected in woody perennials; (2) in the species investigated in greatest detail, Gunnera magellanica, the inhibitory effect of solar UV-B is accompanied by increased levels of DNA damage in leaf tissue, and the DNA damage density in the early spring is clearly correlated with the dose of weighted UV-B measured at ground level; (3) the herbaceous species investigated thus far show little or no acclimation responses to ambient UV-B such as increased sunscreen levels and DNA repair capacity; and (4) ambient UV-B has significant effects on heterotrophic organisms, included marked inhibitory effects on insect herbivory. The results from the experiments summarized in this review clearly indicate that UV-B influences several potentially important processes and ecological interactions in the terrestrial ecosystems of Tierra del Fuego.

Assuntos

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Ultraviolet B radiation (UV-B, 290-315 nm) can cause damage and induce photomorphogenic responses in plants. The mechanisms that mediate the photomorphogenic effects of UV-B are unclear. In etiolated Arabidopsis seedlings, a daily exposure to 2.5 h of UV-B enhanced the cotyledon opening response induced by a subsequent red light (R) pulse. An R pulse alone, 2.5 h of UV-B terminated with a far-red pulse, or 2.5 h of continuous R caused very little cotyledon opening. The enhancing effect of UV-B increased with fluence rate up to approximately 7.58 micromol m(-2) s(-1); at higher fluence rates the response to UV-B was greatly reduced. The phyA, phyA cry1, and cry1 cry2 mutants behaved like the wild type when exposed to UV-B followed by an R pulse. In contrast, phyB, phyB cry1, and phyB phyA mutants failed to open the cotyledons. Thus, phytochrome B was required for the cotyledon opening response to UV-B --> R treatments, whereas phytochrome A and cryptochromes 1 and 2 were not necessary under the conditions of our experiments. The enhancing effect of low doses of UV-B on cotyledon opening in uvr1 uvr2 and uvr1 uvr3 mutants, deficient in DNA repair, was similar to that found in the wild type, suggesting that this effect of UV-B was not elicited by signals derived from UV-B-induced DNA lesions (cyclobutane pyrimidine dimers and 6-4 photoproducts). We conclude that low doses of UV-B, perceived by a receptor system different from phytochromes, cryptochromes, or DNA, enhance a de-etiolation response that is induced by active phytochrome B.

Assuntos

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Colorless phenylpropanoid derivatives are known to protect plants from ultraviolet (UV) radiation, but their photoregulation and physiological roles under field conditions have not been investigated in detail. Here we describe a fast method to estimate the degree of UV penetration into photosynthetic tissue, which is based on chlorophyll fluorescence imaging. In Arabidopsis this technique clearly separated the UV-hypersensitive transparent testa (tt) tt5 and tt6 mutants from the wild type (WT) and tt3, tt4, and tt7 mutants. In field-grown soybean (Glycine max), we found significant differences in UV penetration among cultivars with different levels of leaf phenolics, and between plants grown under contrasting levels of solar UV-B. The reduction in UV penetration induced by ambient UV-B had direct implications for DNA integrity in the underlying leaf tissue; thus, the number of cyclobutane pyrimidine dimers caused by a short exposure to solar UV-B was much larger in leaves with high UV transmittance than in leaves pretreated with solar UV-B to increase the content phenylpropanoids. Most of the phenylpropanoid response to solar UV in field-grown soybeans was induced by the UV-B component (lambda

Assuntos

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The primary motivation behind the considerable effort in studying stratospheric ozone depletion is the potential for biological consequences of increased solar UVB (280-315 nm) radiation. Yet, direct links between ozone depletion and biological impacts have been established only for organisms of Antarctic waters under the influence of the ozone "hole;" no direct evidence exists that ozone-related variations in UVB affect ecosystems of temperate latitudes. Indeed, calculations based on laboratory studies with plants suggest that the biological impact of ozone depletion (measured by the formation of cyclobutane pyrimidine dimers in DNA) is likely to be less marked than previously thought, because UVA quanta (315-400 nm) may also cause significant damage, and UVA is unaffected by ozone depletion. Herein, we show that the temperate ecosystems of southern South America have been subjected to increasingly high levels of ozone depletion during the last decade. We found that in the spring of 1997, despite frequent cloud cover, the passages of the ozone hole over Tierra del Fuego (55 degrees S) caused concomitant increases in solar UV and that the enhanced ground-level UV led to significant increases in DNA damage in the native plant Gunnera magellanica. The fluctuations in solar UV explained a large proportion of the variation in DNA damage (up to 68%), particularly when the solar UV was weighted for biological effectiveness according to action spectra that assume a sharp decline in quantum efficiency with increasing wavelength from the UVB into the UVA regions of the spectrum.

Assuntos

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Most of our present knowledge about the impacts of solar UVB radiation on terrestrial ecosystems comes from studies with plants. Recently, the effects of UVB on the growth and survival of consumer species have begun to receive attention, but very little is known about UVB impacts on animal behavior. Here we report that manipulations of the flux of solar UVB received by field-grown soybean crops had large and consistent effects on the density of the thrips (Caliothrips phaseoli, Thysanoptera: Thripidae) populations that invaded the canopies, as well as on the amount of leaf damage caused by the insects. Solar UVB strongly reduced thrips herbivory. Thrips not only preferred leaves from plants that were not exposed to solar UVB over leaves from UVB-exposed plants in laboratory and field choice experiments, but they also appeared to directly sense and avoid exposure to solar UVB. Additional choice experiments showed that soybean leaf consumption by the late-season soybean worm Anticarsia gemmatalis (Lepidoptera: Noctuidae) was much less intense in leaves with even slight symptoms of an early thrips attack than in undamaged leaves. These experiments suggest that phytophagous insects can present direct and indirect behavioral responses to solar UVB. The indirect responses are mediated by changes in the plant host that are induced by UVB and, possibly, by other insects whose behavior is affected by UVB.

Assuntos

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Transgenic tobacco plants that express an oat phytochrome gene (phyA) under control of the cauliflower mosaic virus (CaMV) 35S promoter and display altered photophysiology were used to test the role of light as a vehicle of information in dominance relationships between neighboring plants. Compared with the isogenic wild type, phyA-overexpressing plants showed dramatically reduced morphological responsivity to changes in the red/far red ratio of the incident light and to the proximity of neighboring plants in spacing experiments. In transgenic canopies an increase in stand density caused the small plants of the population to be rapidly suppressed by their neighbors. In wild-type canopies, plants responded to increased density with large morphological changes, and there appeared to be an inverse relationship between the magnitude of this morphological response and the ranking of the individual plant in the population size hierarchy. In these wild-type populations, size inequality increased only moderately with density within the time frame of the experiments. Our results suggest that, in crowded stands, the ability of individual plants to acquire information about their light environment via phytochrome plays a central role in driving architectural changes that, at the population level, delay the development of size differences between neighbors.

Assuntos

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Seedlings of shade-intolerant species react to alterations of the light climate caused by their neighbors with morphological changes that may influence the pattern of resource acquisition and utilization at the whole-canopy level. One such change, the increased stem elongation rate that is triggered by low red (R, 660 nm) to far-red (FR, 730 nm) ratios (R:FR) in dense canopies, might reduce the amount of assimilates available for leaf area expansion or root growth, and in that way affect resource capture by the canopy. We have tested this hypothesis by comparing the growth of both isolated individuals and canopies of the weed Amaranthus quitensis under conditions differing only in the spectral distribution of the incident light. When canopies received the full spectrum of sunlight, the stems were a large proportion (40-57%) of total biomass. Filtering the FR waveband (and hence raising the R:FR ratio to eliminate the neighbors' proximity-signal) resulted in shorter canopies with lighter stems. However, the growth of leaves and roots was not promoted by this treatment, indicating that the opportunity cost of the assimilates invested in the stems was nil or very small. Filtering the FR had no effect on biomass accumulation when plants were grown as isolated individuals. The higher growth of the canopics under full spectrum could be due to a higher light interception or to a higher efficiency of light conversion into biomass. The first possibility is weakened by the observation that filtering the FR had no effect on the dynamics of soil covering by the crops. The second is indirectly strengthened by results of an experiment with isolated plants showing that stem elongation, stem growth, and total plant biomass can be increased by reducing the flux of R light received by the stems without affecting the light climate of the leaves. Further work is needed to distinguish between these two possibilities; whatever the cause, our results show that the elongation responses to decreased R:FR may lead to a net increase in canopy productivity, and do not necessarily have a negative impact on the growth of resource-harvesting organs.

RESUMO

A fiber optic probe inserted into plant tissues was used to investigate the effects of canopy density on the light environment in different organs. The red:far-red ratio inside the stem of Datura ferox L. seedlings and the estimated phytochrome photoequilibrium were strongly reduced by the presence of neighbors forming canopies too sparse to cause any mutual shading at the level of the leaves. In such canopies, changes in plant density had little effects on the light regime inside the leaves of the succulent Aeonium haworthii (S.D.) Webb et Berth., particularly when the lamina was kept nearly normal to the direct rays of the sun. In field experiments using D. ferox and Sinapis alba L. seedlings, the elongation of the internodes responded to various types of localized light-quality treatments that simulated different plant densities in sparse canopies. The responses were quantitatively similar to those elicited by changes in plant density. The evidence supports the hypothesis that, in stands formed by plants of similar size, the red:far-red ratio of the light that impinges laterally on the stems is among the earliest environmental cues that allow plants to detect local canopy density and adjust axis extension accordingly.

RESUMO

We studied the effects of density on the dynamics of seedling growth and canopy microclimate within experimental stands composed of Datura ferox L. seedlings grown in individual pots. Interception of photosynthetically active radiation (PAR) by seedlings was evaluated either indirectly, by measuring leaf area, proportion of leaf area shaded by neighbouring individuals and laminar orientation with respect to sunlight, or directly, by measuring PAR at individual leaves at their natural angle of display. An integrating cylinder, with a geometry approximating that of a stem, was used within the canopies to measure the red:far-red (R:FR) ratio of the light flux from all compass points parallel to the soil surface. Seedlings responded rapidly (i.e. 1-2 weeks) to increased density by producing longer internodes and partitioning more dry matter to stems relative to leaves. These responses were observed before either PAR interception of growth rate were reduced by the presence of neighbours. Conversely, morphogenetic adjustment was preceded by a consistent effect of plant density on the R:FR ratio of the light received by the integrating cylinder. Air and soil temperature were not affected by density in these experiments. Differences in wind velocity within the canopy associated with plant density were avoided by the experimental procedure. The results support the idea that the drop in R:FR ratio of the light flux parallel to the ground - e.g. reflected sunlight - is an early signal that allows rapid adjustment of plant form to changes in canopy structure.