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1.
The impact of ambient ultraviolet (UV)‐B radiation on the endemic bryophyte, Grimmia antarctici, was studied over 14 months in East Antarctica. Over recent decades, Antarctic plants have been exposed to the largest relative increase in UV‐B exposure as a result of ozone depletion. We investigated the effect of reduced UV and visible radiation on the pigment concentrations, surface reflectance and physiological and morphological parameters of this moss. Plexiglass screens were used to provide both reduced UV levels (77%) and a 50% decrease in total radiation. The screen combinations were used to separate UV photoprotective from visible photoprotective strategies, because these bryophytes are growing in relatively high light environments compared with many mosses. G. antarctici was affected negatively by ambient levels of UV radiation. Chlorophyll content was significantly lower in plants grown under near‐ambient UV, while the relative proportions of photoprotective carotenoids, especially β‐carotene and zeaxanthin, increased. However, no evidence for the accumulation of UV‐B‐absorbing pigments in response to UV radiation was observed. Although photosynthetic rates were not affected, there was evidence of UV effects on morphology. Plants that were shaded showed fewer treatment responses and these were similar to the natural variation observed between moss growing on exposed microtopographical ridges and in more sheltered valleys within the turf. Given that other Antarctic bryophytes possess UV‐B‐absorbing pigments which should offer better protection under ambient UV‐B radiation, these findings suggest that G. antarctici may be disadvantaged in some settings under a climate with continuing high levels of springtime UV‐B radiation.  相似文献   

2.
Using steel pins inserted into growing moss colonies near Casey Station, Wilkes Land, continental Antarctica, we have measured the growth rate of three moss species: Bryum pseudotriquetrum and Schistidium antarctici over 20 years and Ceratodon purpureus over 10 years. This has provided the first long-term growth measurements for plants in Antarctica, confirming that moss shoots grow extremely slowly in Antarctica, elongating between 1 and 5 mm per year. Moss growth rates are dependent on availability of water. Antheridia were observed on some stems of B. pseudotriquetrum; no archegonia or sporophytes were observed. Stems bearing antheridia elongated much more slowly than vegetative stems in the same habitat. Two other methods of growth rate measurement were tested, and gave similar rates of elongation over shorter periods of time. However, for long-term measurements, the steel pin measurements proved remarkably reproducible and reliable.  相似文献   

3.
Adele Post 《Polar Biology》1990,10(4):241-245
Summary Variation in leaf pigmentation from green to ginger is observed for Ceratodon purpureus (Hedw.) Brid. in Antarctica. Electron microscopy of ginger and green leaves reveals less thylakoid stacking, a response to greater light exposure, in the ginger leaves. In extremely exposed sites C. purpureus has low chlorophyll a/b ratios which correlate with decreased 77K chlorophyll fluorescence, indicating damage to chlorophyll a. Pigment analysis of ginger moss shows that even when the chlorophyll a/b ratio has not decreased the pigment composition differs from green moss. The increase in anthocyanin and decrease in chlorophyll concentrations largely account for the visual change from green to ginger. The ratio of total carotenoid to chlorophyll varies from 0.35 in green moss to 0.55 in the ginger moss, with violaxanthin increased preferentially. Since these changes in pigmentation are consistent with photoprotection and they are linked to light dependent variations in chloroplast structure, it appears that photoprotective pigments are a useful adaptation for the bright Antarctic environment.  相似文献   

4.
This study presents the effect of water content in moss turves from the Windmill Islands, Wilkes Land, continental Antarctica on the composition of associated filamentous fungi. The two most prevalent mosses were sampled: Grimmia antarctici and Bryum pseudotriquetrum. Results were expressed as dry weights. No relationship was found between water content of the mosses or the surrounding soil and fungal diversity. Ten species of fungi were recovered and all species belong to cosmopolitan taxa. Accepted: 21 June 1999  相似文献   

5.
Responses of plants in polar regions to UVB exposure: a meta-analysis   总被引:1,自引:0,他引:1  
We report a meta‐analysis of data from 34 field studies into the effects of ultraviolet B (UVB) radiation on Arctic and Antarctic bryophytes and angiosperms. The studies measured plant responses to decreases in UVB radiation under screens, natural fluctuations in UVB irradiance or increases in UVB radiation applied from fluorescent UV lamps. Exposure to UVB radiation was found to increase the concentrations of UVB absorbing compounds in leaves or thalli by 7% and 25% (expressed on a mass or area basis, respectively). UVB exposure also reduced aboveground biomass and plant height by 15% and 10%, respectively, and increased DNA damage by 90%. No effects of UVB exposure were found on carotenoid or chlorophyll concentrations, net photosynthesis, Fv/Fm or ΦPSII, belowground or total biomass, leaf mass, leaf area or specific leaf area (SLA). The methodology adopted influenced the concentration of UVB absorbing compounds, with screens and natural fluctuations promoting significant changes in the concentrations of these pigments, but lamps failing to elicit a response. Greater reductions in leaf area and SLA, and greater increases in concentrations of carotenoids, were found in experiments based in Antarctica than in those in the Arctic. Bryophytes typically responded in the same way as angiosperms to UVB exposure. Regression analyses indicated that the percentage difference in UVB dose between treatment and control plots was positively associated with concentrations of UVB absorbing compounds and carotenoids, and negatively so with aboveground biomass and leaf area. We conclude that, despite being dominated by bryophytes, the vegetation of polar regions responds to UVB exposure in a similar way to higher plant‐dominated vegetation at lower latitudes. In broad terms, the exposure of plants in these regions to UVB radiation elicits the synthesis of UVB absorbing compounds, reduces aboveground biomass and height, and increases DNA damage.  相似文献   

6.
Depletion of stratospheric ozone since the mid 1970s has led to significant increases in ultraviolet B (UVB) irradiation over Antarctica. The detrimental effects of UVB on plants are many, but plants produce photoprotective flavonoids that reduce cellular damage. We used herbarium samples of the moss Bryum argenteum collected in Antarctica to compare the levels of flavone aglycones in plants collected before and after the formation of the ozone hole. The interpretation of historical data is difficult, because environmental conditions immediately before sample collection are unknown. Factors such as cloud cover can have a significant influence on UVB dose at ground level, modifying the flavonoid content of the specimen and adding considerable variability to the results. Nevertheless, our results revealed significant relationships between total flavone concentration and pooled year classes ( P = 0.001). Furthermore, regression analysis showed a significant negative relationship of total flavone concentration and the level of ozone immediately before the time of collection ( P = 0.016). In addition, the ratio of luteolin (an ortho -dihydroxylated flavone) to apigenin (a monohydroxylated flavone) increased significantly with several environmental parameters. These included (a) increasing modelled midday UVB radiation ( P = 0.002), (b) increasing modelled midday UVB/PAR ratio ( P < 0.001), and (c) decreasing ozone concentration ( P < 0.001). We emphasise the utility of this ratio in interpreting the historical ozone trends rather than relying on changes in total flavone concentrations alone. These results illustrate that herbarium specimens may reveal historical levels of UVB radiation.  相似文献   

7.
Abstract. Bryum argenteum, B. pseudotriquetrum and Ceratodon purpureus are the predominant mosses in Victoria Land, continental Antarctica. All have cosmopolitan distributions and are widespread throughout Antarctica with wide ecological amplitudes resulting in considerable morphological variation. They are well adapted to tolerate the physiological stresses imposed by the severe environment. This study investigates aspects of their growth, physiology and survival in response to habitat constraints, especially hydrology. Their distribution is controlled almost exclusively by moisture availability. Each species tends to predominate in a specific zone along hydrological gradients, with B. pseudotriquetrum on moist soil, C. purpureus on drier soil, and B. argenteum on unstable stream margins, fluvial deposits and the marginal capillary zone. Where conditions are optimal, each species can form a turf 6–10 cm thick. Nutrient status of the soil does not appear to be an important determinant in the distribution pattern within communities. The thermal regime of the moss turf varies according to its moisture content; for a period of ca. six weeks during the summer, with the frequent long spells of 24-h sunshine, temperatures remain above 0 °C for much of the time even though air temperatures are frequently below the freezing point. This allows growth and metabolic activity to proceed continuously at a relatively rapid rate for quite long periods. Annual shoot incremental growth can exceed 3.5 mm in each species. Growth of B. argenteum may be inhibited by UV- B radiation. The optimal temperature for photosynthesis in each species is around 15 °C, but significant carbon fixation occurs at 5 °C. Photosynthetic rates at 5, 10 and 20 °C were B. argenteum > B. pseudotriquetrum > C. purpureus.  相似文献   

8.
We investigated the responses of ultraviolet (UV)‐absorbing compounds, chlorophylls a and b, carotenoids and the growth responses of the pleurocarpous moss Pleurozium schreberi (Britt.) Mitt. to enhanced UV radiation in situ. The moss was exposed to a 52% elevation above the ambient level of erythemally weighted UV‐B radiation, simulating an approximate 20% reduction in the ozone column, in a dry pine forest in Sodankylä, Finland (67 °22′N, 26 °38′E), under arrays of lamps filtered with cellulose diacetate, which transmitted both UV‐B and UV‐A radiation. The moss was also exposed to elevated UV‐A radiation under control arrays of lamps filtered with Melinex polyester and to ambient radiation under arrays with no lamps in them. Effects of enhanced UV radiation on P. schreberi were recorded during the first 3 years of exposure. Enhanced UV‐B radiation did not affect the segment height growth of the moss. The annual dry mass after the second growing season was higher in the UV‐A control than in the other treatments, and dry mass decreased significantly during the third treatment year in both UV treatments compared with the ambient. The specific leaf area of the UV‐B‐treated mosses was significantly higher than the ambient control mosses during the first 2 years. An increase of UV‐absorbing compounds was found in the mosses under enhanced UV‐B radiation compared with the UV‐A control mosses during the first year. Even though the treatment effect on UV‐absorbing compounds was transient, the concentrations of these compounds correlated with the amount of UV‐A and UV‐B radiation received under the elevated UV‐B treatment. A correlation with the irradiation of previous days and preceding month of the sampling day was found. A seasonal reduction occurred in the amount of UV‐absorbing compounds from the beginning of the summer to late summer. The amount of photosynthetic pigments correlated with the amount of photosynthetically active radiation. The moss P. schreberi was thus found to tolerate increasing UV‐B radiation. Our data indicate that P. schreberi tolerates a 52% increase in erythemally weighted UV‐B radiation above ambient, responding during the first few years of exposure by increasing UV‐absorbing compounds and specific leaf area, and decreasing annual dry mass, and then acclimating to its altered radiation environment.  相似文献   

9.
In a field experiment with a mangrove species Suaeda maritima L. grown under ambient and supplementary UV-B radiation corresponding to 20 % ozone depletion, changes in growth and contents of photosynthetic and UV-absorbing pigments were determined. Supplemental UV-B irradiation for 9 d significantly reduced the growth and concentration of photosynthetic pigments. However, anthocyanin and flavonoid contents were significantly increased in UV-treated plants and which could be reduce the UV-B penetration and damage to the underlying tissues.  相似文献   

10.
Long‐term growth response to natural solar radiation with enhanced ultraviolet‐B (UVB) exposure was examined in two species of dinoflagellates [Alexandrium tamarense (M. Lebour) Balech, At, and Heterocapsa triquetra (Ehrenb.) F. Stein, Ht], including two strains of A. tamarense, one from Spain and another from UK, and one diatom species (Thalassiosira pseudonana Hasle et Heimdal). We examined whether variable photoprotection (mycosporine‐like amino acids [MAAs] and xanthophyll‐cycle pigments) affected photosynthetic performance, phytoplankton light absorption, and growth. Growth rate was significantly reduced under enhanced UVB for the UK strain of At and for Ht (both grew very little) as well as for the diatom (that maintained high growth rates), but there was no effect for the Spanish strain of At. MAA concentration was high in the dinoflagellates, but undetectable in the diatom, which instead used the xanthophyll cycle for photoprotection. The highest cell concentrations of MAAs and photoprotective pigments were observed in the UK strain of At, along with lowest growth rates and Fv/Fm, indicating high stress levels. In contrast, the Spanish strain showed progressive acclimation to the experimental conditions, with no significant difference in growth between treatments. Increase in total MAAs followed linearly the cumulative UVB of the preceding day, and both total and primary MAAs were maintained at higher constitutive levels in this strain. Acclimation to enhanced UVB in the diatom resulted in an increase in PSII activity and reduction in nonphotochemical quenching, indicating an increased resistance to photoinhibition after a few weeks. All four species showed increased phytoplankton light absorption under enhanced UVB. Large intrastrain differences suggest a need to consider more closely intraspecific variability in UV studies.  相似文献   

11.
Depending on geographical location, plants are exposed to variable amounts of UVB radiation and herbivore attack. Because the role(s) of UVB in the priming and/or accumulation of plant defence metabolites against herbivores are not well understood, we used field‐grown Nicotiana attenuata plants to explore the effects of UVB on herbivore performance. Consistent with previous reports, UVB‐exposed plants accumulated higher levels of ultraviolet (UV)‐absorbing compounds (rutin, chlorogenic acid, crypto‐chlorogenic acid and dicaffeoylspermidine). Furthermore, UVB increased the accumulation of jasmonic acid, jasmonoyl‐L‐isoleucine and abscisic acid, all phytohormones which regulate plant defence against biotic and abiotic stress. In herbivore bioassays, N. attenuata plants experimentally protected from UVB were more infested by mirids in three consecutive field seasons. Among defence metabolites measured, 17‐hydroxygeranyllinalool diterpene glycosides (HGL‐DTGs) showed strongly altered accumulation patterns. While constitutive HGL‐DTGs levels were higher under UVB, N. attenuata plants exposed to mirid bugs (Tupiocoris notatus) had still more HGL‐DTGs under UVB, and mirids preferred to feed on HGL‐DTGs‐silenced plants when other UVB protecting factors were eliminated by UVB filters. We conclude that UVB exposure not only stimulates UV protective screens but also affects plant defence mechanisms, such as HGL‐DTGs accumulation, and modulates ecological interactions of N. attenuata with its herbivores in nature.  相似文献   

12.
This study aims to investigate the photoprotective properties of a Lomentaria hakodatensis ethanol extract (LHE) against ultraviolet B (UVB) radiation-induced cellular damage in human HaCaT keratinocytes. LHE exhibited scavenging activity against intracellular reactive oxygen species (ROS), which were generated by either hydrogen peroxide (H2O2) or UVB radiation. Moreover, LHE scavenged superoxide anion generated by the xanthine/xanthine oxidase system and hydroxyl radical generated by the Fenton reaction (FeSO4 + H2O2). Furthermore, LHE exhibited UVB absorptive properties and attenuated injury to cellular components (e.g., lipids, proteins and DNA), resulting from UVB-induced oxidative stress. In addition, LHE reduced apoptosis in response to UVB, as shown by decreased DNA fragmentation and the formation of apoptotic bodies. These results suggest that LHE protects human keratinocytes against UVB-induced oxidative stress by scavenging ROS and absorbing UVB rays; thereby reducing damage to biological components.  相似文献   

13.
The simple structure of mosses, a major component of Antarctic terrestrial vegetation, has led to suggestions that they might be exceptionally sensitive to enhanced UV from the ozone hole. The results presented here show that the mosses Ceratodon purpureus and Bryum subrotundifolium are resistant to UV-A and that the latter species can rapidly change its protection to suit the UV environment. The studies were made using a UV-A PAM chlorophyll fluorescence fluorometer that allowed absorption of UV-A before arrival at the chloroplast (i.e. UV-A shielding) to be estimated. Both C. purpureus and B. subrotundifolium have sun and shade forms that differ markedly in colour and their protection from UV-A. Shade forms of B. subrotundifolium, initially low in UV-A protection, achieve full, sun-form levels in about 6 days when exposed to ambient sunlight. These results, taken with other recent studies, suggest that not only are Antarctic mosses well protected from ambient UV, but are also as adaptable to incident UV as higher plants.  相似文献   

14.
UVB radiation is biologically active; in plants, it can induce a range of molecular, biochemical, morphological and developmental responses. Although much progress has been made in elucidating UVB perception and signalling pathways under controlled laboratory conditions, understanding of the adaptive, ecological role of UVB responses is still very limited. In this study, we looked at the functional role of UVR8 under outdoor light conditions, by studying growth, photosynthetic competence and accumulation of UV absorbing pigments in a mutant lacking functional UVR8 protein. It was found that the influence of UVB on morphology is restricted to summer and is independent of UVR8. In contrast, UVB had an effect on the content of UV‐absorbing pigments and the maximal efficiency of photosystem II of photosynthesis in the uvr8‐1 mutant throughout the year. It is concluded that the UVR8 photoreceptor plays a role throughout the year, in the temperate climate zone, even when UVB levels are relatively low.  相似文献   

15.
Net photosynthesis and dark respiration (CO2 flux) of Antarctic mosses were measured at Langhovde, East Antarctica, from 9 to 17 January 1988. Moss blocks were taken from communities in the Yukidori Valley (69°14′30″S, 39°46′00″E) at Langhovde. Each block was composed ofCeratodon purpureus andBryum pseudotriquetrum, orB. pseudotriquetrum. The upper part of the block was used to measure net photosynthesis and dark respiration. The net photosynthesis of each sample was measured in the field for one or three days with two infrared CO2 gas analyzers and an assimilation chamber. The relationships of net photosynthetic rate and dark respiration rate, to the water content of the sample, the intensity of solar radiation and the moss temperature were estimated from the field data. The maximum rate of net photosynthesis was about 4 μmol CO2 m−2s−1 at saturating radiation intensity and at optimum temperature, about 10°C. Environmental features of moss habitats in the Yukidori Valley are discussed in relation to these results.  相似文献   

16.
To assess the effects of UV radiation and its interaction with water availability on Mediterranean plants, we performed an experiment with seedlings of six Mediterranean species (three mesophytes vs three xerophytes) grown in a glasshouse from May to October under three UV conditions (without UV, with UVA and with UVA+UVB) and two irrigation levels (watered to saturation and low watered). Morphological, physiological and biochemical measures were taken. Exposure to UVA+UVB increased the overall leaf mass per area (LMA) and the leaf carotenoids/chlorophyll a + b ratio of plants in relation to plants grown without UV or with UVA, respectively. In contrast, we did not find a general effect of UV on the leaf content of phenols or UVB‐absorbing compounds of the studied species. Regarding plant growth, UV inhibited the above‐ground biomass production of well‐watered plants of Pistacia lentiscus. Conversely, under low irrigation, UVA tended to abolish the reduction in growth experienced by P. lentiscus plants growing in a UV‐free environment, in accordance with UVA‐enhanced apparent electron transport rate (ETR) values under drought in this species. UVA also induced an overall increase in root biomass when plants of the studied species were grown under a low water supply. In conclusion, while plant exposition to UVA favored root growth under water shortage, UVB addition only gave rise to photoprotective responses, such as the increase in LMA or in the leaf carotenoids/chlorophyll a + b ratio of plants. Species‐specific responses to UV were not related with the xerophytic or mesophytic character of the studied species.  相似文献   

17.
Recent studies suggest that complex interacting processes are driving global amphibian declines. Increased ultraviolet B (UVB) radiation in the solar spectrum associated with ozone depletion has been implicated in declines, and evidence suggests that the effects of UVB radiation on amphibians may be greater at cooler temperatures. We tested the thermal sensitivity of UVB effects on amphibians in a controlled factorial experiment using the striped marsh frog, Limnodynastes peronii as a model species. We compared survival, growth and locomotor performance of embryonic and larval L. peronii reared under low and high UVB exposures at both 20 and 30 °C. Embryonic and larval L. peronii proved extremely sensitive to UVB damage and exhibited greater sensitivity at 20 °C compared with 30 °C. Embryonic survival to Gosner stage 25 was unaffected by UVB exposure at 30 °C, but at 20 °C survival was reduced to 52% under high UVB. Larval survival exhibited a similar trend. At 20 °C, all tadpoles survived under low UVB, whereas under high UVB there was 100% mortality after 15 days of exposure. At 30 °C, 86% survived under low UVB, but only 46% survived under high UVB. Sublethal effects such as, embryonic malformation, retarded larval growth and reduced larval swimming performance were also greater at 20 °C compared with 30 °C. Our results strongly indicate that UVB damage in amphibians is markedly increased at cooler temperatures. Thus, populations of UVB sensitive species occurring at cold climates may be at greater risk of declines due to increased solar UVB radiation.  相似文献   

18.
The sensitivity of different life stages of the eulittoral green alga Urospora penicilliformis (Roth) Aresch. to ultraviolet radiation (UVR) was examined in the laboratory. Gametophytic filaments and propagules (zoospores and gametes) released from filaments were separately exposed to different fluence of radiation treatments consisting of PAR (P = 400–700 nm), PAR + ultraviolet A (UVA) (PA, UVA = 320–400 nm), and PAR + UVA + ultraviolet B (UVB) (PAB, UVB = 280–320 nm). Photophysiological indices (ETRmax, Ek, and α) derived from rapid light curves were measured in controls, while photosynthetic efficiency and amount of DNA lesions in terms of cyclobutane pyrimidine dimers (CPDs) were measured after exposure to radiation treatments and after recovery in low PAR; pigments of propagules were quantified after exposure treatment only. The photosynthetic conversion efficiency (α) and photosynthetic capacity (rETRmax) were higher in gametophytes compared with the propagules. The propagules were slightly more sensitive to UVB‐induced DNA damage; however, both life stages of the eulittoral inhabiting turf alga were not severely affected by the negative impacts of UVR. Exposure to a maximum of 8 h UVR caused mild effects on the photochemical efficiency of PSII and induced minimal DNA lesions in both the gametophytes and propagules. Pigment concentrations were not significantly different between PAR‐exposed and PAR + UVR–exposed propagules. Our data showed that U. penicilliformis from the Antarctic is rather insensitive to the applied UVR. This amphi‐equatorial species possesses different protective mechanisms that can cope with high UVR in cold‐temperate waters of both hemispheres and in polar regions under conditions of increasing UVR as a consequence of further reduction of stratospheric ozone.  相似文献   

19.
Bloom‐forming dinoflagellates often synthesize high concentrations of mycosporine‐like amino acids (MAAs) as a protection from high PAR and UV radiation, which may afford them a competitive advantage over other species. Photoprotection and long‐term acclimation to natural and enhanced UVB radiation were examined during 4 weeks in three dinoflagellates and one diatom. Semi‐continuous, nutrient‐replete cultures were maintained outdoors under natural light, with or without additional UVB (from lamps). The photosynthetic performance (XE‐PAM fluorometry) and the concentration and composition of MAAs and photoprotective carotenoids (PPCs; comprising xanthophylls) were followed weekly. A great variety of MAAs was found in the dinoflagellates (up to 10, identified by cochromatography and LC–MS). Large differences between sp. (and between two strains of the same sp.) were observed. In most cases, the composition and concentration of MAAs and PPCs increased under enhanced UVB. Growth rate and photosynthetic performance were sometime lower under enhanced UVB, while in other cases photoacclimation seemed to occur. The least affected sp. was Alexandrium tamarense who responded most rapidly to enhanced UVB. The algae also responded to changes in natural incident radiation. Some methodological aspects of MAA quantification were also investigated (with four species of dinoflagellates). Our results indicate how freezing (liquid‐N) likely breaks the cells, mobilizing the water‐soluble MAAs in the filter moist (otherwise highly packaged in fresh cells), while the organelles containing the pigments remain relatively intact. The implications of standard sample handling on the evaluation of MAA concentration in algae and on the estimation of the absorption of suspended material in water from in vivo spectroscopy (filter pad technique) will also be discussed.  相似文献   

20.
This study reports UV screening pigments in the upper cortices of two widespread lichens collected in three sun-exposed locations along a latitudinal gradient from the Arctic lowland to alpine sites of the Central European Alps. Populations from the Alps receive 3–5 times higher UV-B irradiance than their Arctic counterparts from Svalbard because of latitudinal and altitudinal gradients in UV-B irradiance. In Cetraria islandica, the screening capacity of melanin in the upper cortices was assessed by direct measurements of cortical transmittance (250–1,000 nm). A comparison of cortical transmittances in brown sun-exposed and pale shade-adapted forest C. islandica thalli showed that fungal melanins strongly absorb both UV-B and photosynthetically active radiation (PAR). For Xanthoria elegans cortical UV-B absorbing pigments, mainly the orange parietin, were extracted and quantified. Field experiments with extracted, parietin-deficient X. elegans thalli cultivated under various filters showed that UV-B was essential for the induction of parietin synthesis. The parietin resynthesis in these parietin-deficient samples increased with decreasing latitude of their location in which they had been sampled, which may imply that the synthesis of pigments is habitat specific. However, no latitudinal gradient in cortical screening capacity was detected for any of the two species investigated in the field. This implies that Arctic populations maintain a high level of screening pigments in spite of low ambient UV-B, and that the studied lichen species presumably may tolerate an increase in UV-B radiation due to the predicted thinning of the ozone layer over polar areas  相似文献   

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