Effect of acute gamma radiation on some physiological features of lichens

The influence of acute gamma radiation on four physiological characteristics of lichens was investigated. Membrane permeability in irradiated thalli of Cladonia arbuscula, Cetraria islandica and Hypogymnia physodes increased considerably compared to controls. The nitrogen content in Peltigera aphthosa appeared to be stable to acute gamma radiation. Up to 1000 Gy there was no essential decrease in respiration intensity in H. physodes and P. aphthosa or of peroxidase activity in P. aphthosa.     

蓝藻对UV-B增强的响应与适应

平流层臭氧破坏导致地球表面紫外辐射(主要是UV-B)增强逐渐受到人们重视。由于蓝藻在生态系统中的重要性和在生物进化过程中的特殊性,用于研究UV-B对生物体的影响具有诸多优势。目前国内关于UV-B与蓝藻的研究报道较少,所以本文介绍了近年来国外该领域的相关研究,主要包括UV-B对蓝藻生物量、光合机构以及固氮等方面的影响,同时着重介绍了蓝藻对UV-B的适应策略。     

Effects of elevated ultraviolet-B radiation on a plant–herbivore interaction

Enhanced ultraviolet-B (UV-B) radiation may have multiple effects on both plants and animals and affect plant–herbivore interactions directly and indirectly by inducing changes in host plant quality. In this study, we examined combined effects of UV-B and herbivory on the defence of the mountain birch (Betula pubescens ssp. czerepanovii) and also the effects of enhanced UV-B radiation on a geometrid with an outbreak cycle: the autumnal moth (Epirrita autumnata). We established an experiment mimicking ozone depletion of 30% (a relevant level when simulating ozone depletion above Northern Lapland). Both arctic species responded only slightly to the enhanced level of UV-B radiation, which may indicate that these species are already adapted to a broader range of UV-B radiation. UV-B exposure slightly induced the accumulation of myricetin glycosides but had no significant effect on the contents of quercetin or kaempferol derivatives. Mountain birch seedlings responded more efficiently to herbivory wounding than to enhanced UV-B exposure. Herbivory induced the activities of foliar oxidases that had earlier been shown to impair both feeding and growth of moth larvae. In contrast, the contents of foliar phenolics did not show the same response in different clones, except for a decrease in the contents of tannin precursors. The induction of foliar phenoloxidase activities is a specific defence response of mountain birches against insect herbivory. To conclude, our results do not support the hypothesis that the outbreak cycle of the autumnal moth can be explained by the cycles of solar activity and UV-B.     

Stratospheric Ozone Depletion: High Arctic Tundra Plant Growth on Svalbard is not Affected by Enhanced UV-B after 7 years of UV-B Supplementation in the Field

The response of tundra plants to enhanced UV-B radiation simulating 15 and 30% ozone depletion was studied at two high arctic sites (Isdammen and Adventdalen, 78° N, Svalbard).The set-up of the UV-B supplementation systems is described, consisting of large and small UV lamp arrays, installed in 1996 and 2002. After 7 years of exposure to enhanced UV-B radiation, plant cover, density, morphological (leaf fresh and dry weight, leaf thickness, leaf area, reproductive and ecophysiological parameters leaf UV-B absorbance, leaf phenolic content, leaf water content) were not affected by enhanced UV-B radiation. DNA damage in the leaves was not increased with enhanced UV-B in Salix polaris and Cassiope tetragona. DNA damage in Salix polaris leaves was higher than in leaves of C. tetragona. The length of male gametophyte moss plants of Polytrichum hyperboreum was reduced with elevated UV-B as well as the number of Pedicularis hirsuta plants per plot, but the inflorescence length of Bistorta vivipara was not significantly affected. We discuss the possible causes of tolerance of tundra plants to UV-B (absence of response to enhanced UV-B) in terms of methodology (supplementation versus exclusion), ecophysiological adaptations to UV-B and the biogeographical history of polar plants     

Boreal peatland ecosystems under enhanced UV-B radiation and elevated tropospheric ozone concentration

Boreal peat-forming wetlands, mires, are globally important sources of methane and sinks for CO₂. As peatland vegetation plays a significant role regulating the exchange of these greenhouse gases, we have assessed the responses of the dominant plants and ecosystem functions to increasing tropospheric ozone concentration and enhanced ultraviolet-B (UV-B) radiation in long-term experiments, the results of which are summarized in this review. The dominant sedge, Eriophorum vaginatum, and especially the Sphagnum mosses common on peatlands, appear fairly tolerant to the future predicted ozone levels. Similarly, UV-B radiation only caused few alterations in the carbohydrates and pigments of the dominant sedge, Eriophorum russeolum, and had no effects on the dominant moss species of the experimental site, Warnstorfia exannulata. Surprisingly, there were alterations in organic acid concentrations in the peat pore water and peat microbial community composition in both experiments. Elevated ozone caused a transient decrease in ecosystem-level gross photosynthesis and methane (CH₄) emission, which shifted to a slight increase later on. Enhanced UV-B decreased dark ecosystem respiration and increased CH₄ emission in the course of the six measurement years. The emission of isoprene was increased by both ozone and UV-B during warm weather periods, suggesting interactive effects with temperature. All in all, we suggest that ozone and UV-B have limited effects on the carbon cycle in boreal peatlands, because other environmental factors, such as temperature, water level and photosynthetically active radiation more strongly regulate CO₂ and CH₄ exchange rates.     

Artificial cultures of the cyanobacterial lichenPeltigera didactyla (Peltigeraceae) in the natural environment

Soredia of the cyanobacterial lichenPeltigera didactyla were cultivated on soil substratum in the natural environment. Various developmental stages (germination, formation of a soredia field, differentiation of thallus lobes with soralia and fruiting bodies) were attained during a cultivation period of 6–7 months. Our field experiments revealed that the ephemeral pioneer lichenPeltigera didactyla has a short life cycle. Thus this lichen is a well-suited object for further investigations on thallus morphogenesis of lichens with cyanobionts.Dedicated to Prof. DrLothar Geitler on the occasion of his 90th birthday.     

Chitin-glucan complex in cell walls of the Peltigera aphthosa lichen

Cell walls and chitin-glucan complexes isolated from uneven-aged components of the thallus of the Peltigera aphthosa lichen were studied. The mass fraction of the cell wall and chitin-glucan complexes increased with age, but the content of nitrogen in these structures decreased with age. The basal area of the thallus was characterized by the largest mass fraction of the chitin-glucan complex from the dry mass of the thallus; the apical area, by the largest mass fraction of chitin in the complex. It was demonstrated that in P. aphthosa, the degree of deacetylation of chitin in the complex (depending on the age) was 33 and 54% in the apical and basal areas, respectively. The suggested method of functional analysis of chitin-glucan complexes for the presence of free amino groups in them can be used for studying other lichenified fungi.     

UV-B-tolerance of alpine and arctic Daphnia

Survival of transparent and pigmented Daphnia from alpine as well as arctic habitats, exposed to solar radiation with total or reduced UV-B, was measured. In the alpine pond, survivorship and reproduction of transparent juvenile as well as adult Daphnia were significantly reduced due to UV-B. Transparent adult arctic Daphnia also showed significant reduction in survivorship when exposed to total, when compared with controlled solar radiation. In contrast, heavily melanized adult arctic Daphnia showed no increase in mortality when exposed to natural solar radiation. There appeared to be differences both in tolerance and behavior in the two arctic forms when exposed to solar radiation.     

Solar ultraviolet-B radiation influence on Sphagnum bog and Carex fen ecosystems: first field season findings in Tierra del Fuego, Argentina

Stratospheric ozone depletion occurs over Tierra del Fuego, southern Argentina and Chile, in the austral spring and summer due to the precession of the Antarctic ‘ozone hole’ and the general erosion of the ozone layer. Plots receiving either near-ambient or reduced UV-B radiation were established using different louvered plastic film filters over Sphagnum bog and Carex fen ecosystems in October 1996. In the Sphagnum bog system, growth measurements during the late spring and summer showed no significant differences in the moss Sphagnum magellanicum, or the vascular plants (Empetrum rubrum, Nothofagus antarctica, and Tetroncium magellanicum) between near-ambient and attenuated UV-B radiation treatments. In the Carex fen system, leaf length and spike height did not differ in the two dominant species, Carex decidua and C. curta, between UV-B radiation treatments. The length of individual spikelets of C. curta under near-ambient UV-B radiation was less than under the reduced UV-B radiation treatment, but this was not evident in C. decidua. No differences in seed number, seed mass, or viability were seen in either Carex species between the UV-B treatments. Two important constituents of the microfauna that inhabit the Sphagnum bog are testate amoebae and rotifers. These both appeared to be more numerous under near-ambient UV-B radiation than under reduced UV-B radiation. The subtle responses of the Sphagnum and Carex ecosystems may become more apparent in subsequent years as the treatments are continued. Trophic-level changes, such as the differences in number of amoebae and rotifers, may be more sensitive to solar UV-B radiation than growth and productivity of the vegetation.     

Ion-exchange characteristics of the cell walls isolated from the thallus of the lichen <Emphasis Type="Italic">Peltigera aphthosa</Emphasis> (L.) Willd

Ion-exchange characteristics of the cell walls isolated from different zones of the foliose lichen Peltigera aphthosa (L.) Willd were determined. Four types of ionogenic groups were revealed in the thallus cell walls of P. aphthosa, namely amino groups, carboxylic groups of uronic acids, carboxylic groups of phenolic acids, and phenolic OH groups. They may participate in the ion-exchange reactions with the ions of the environment. The amount of ionogenic groups in P. aphthosa cell walls was found to depend on the zone and age of the thallus.     

Rapid photosynthetic recovery of a snow-covered feather moss and Peltigera lichen during sub-Arctic midwinter warming

Background: Arctic lichens and mosses are covered by snow for more than half the year and are generally considered as being dormant for most of this period. However, enhanced frequency of winter warming events due to climate change can cause increased disturbance of their protective subnivean environment.

Aim: To further understand cryptogamic responses to midwinter warming we compared the ecophysiological performance of one lichen and one moss species during a simulated warming event.

Methods: We measured photosynthesis and dark respiration in samples of the moss Hylocomium splendens and the lichen Peltigera aphthosa removed from under snow, and on natural refreezing after the warming event, which was simulated by using infrared heaters suspended above the ground.

Results: The moss exposed to light at +5 °C immediately after removal from their subnivean environment and from warmed plots showed positive net gas exchange within 332 s; the lichen required 1238 s. Photosynthesis and nitrogen fixation rates were equal to that, or higher than, during the preceding growing season. Upon refreezing after the event, moss photosynthesis declined considerably.

Conclusions: The moss, and to a lesser extent the lichen, may contribute to subnivean midwinter ecosystem respiration, and both are opportunistic, and can take advantage of warmer winter phases for photosynthesis and growth. This ought to be taken into account in vegetation change projections of cryptogam-rich ecosystems.     

Isoprene emission from a subarctic peatland under enhanced UV-B radiation

Isoprene is a reactive hydrocarbon with an important role in atmospheric chemistry, and emissions from vegetation contribute to atmospheric carbon fluxes. The magnitude of isoprene emissions from arctic peatlands is not known, and it may be altered by increasing UV-B radiation. Isoprene emission was measured with the dynamic chamber method from a subarctic peatland under long-term enhancement of UV-B radiation targeted to correspond to a 20% loss in the stratospheric ozone layer. The site type of the peatland was a flark fen dominated by the moss Warnstorfia exannulata and sedges Eriophorum russeolum and Carex limosa. The relationship between species densities and the emission was also assessed. Isoprene emissions were significantly increased by enhanced UV-B radiation during the second (2004) and the fourth (2006) growing seasons under the UV-B exposure. Emissions were related to the density of E. russeolum. The dominant moss, W. exannulata, proved to emit small amounts of isoprene in a laboratory trial. Subarctic fens, even without Sphagnum moss, are a significant source of isoprene to the atmosphere, especially under periods of warm weather. Warming of the Arctic together with enhanced UV-B radiation may substantially increase the emissions.     

Differential stimulation of an arid-environment winter ephemeral Dimorphotheca pluvialis (L.) Moench by ultraviolet-B radiation under nutrient limitation

A South African winter ephemeral D. pluvialis was exposed, under low and high nutrient conditions, to four different daily doses of biologically effective UV-B radiation. These simulated different depletions (range 0–30%) in the ozone layer at the southerly distribution limit (33° 56′S) of this species, and included daily UV-B doses received at the northerly distribution limit (26° 38′S) without ozone depletion. Growth inhibition by increased UV-B radiation was observed during early vegetative stages, but only under low nutrient conditions. Thereafter, net CO₂ assimilation rate, growth and reproduction were stimulated by an increase in UV-B radiation, though doses above those approximating a 20% ozone depletion appeared to be inhibitory. Differential stimulation occurred in the two nutrient treatments. Under high nutrient conditions, photosynthesis (specifically carboxylation efficiency), and numbers of leaves, inflorescences and diaspores per plant, and leaf areas increased, but leaf thickness decreased with increased UV-B radiation. Under low nutrient conditions, dry masses of leaves, stems, inflorescences and diaspores, and total above-ground dry masses increased with increased UV-B radiation. Foliar organic carbon and nitrogen concentrations and foliar concentrations of UV-B absorbing compounds were unaffected by increased UV-B radiation, but foliar P concentrations declined. Diaspore viability declined with increased UV-B radiation. The net effect was a 35 to 43% reduction in viable diaspore production under high nutrient conditions at UV-B doses equivalent to those currently received at the northerly distribution limit during the reproductive phase. It is concluded that anticipated increases in UV-B radiation could reduce regeneration success, and seedling survival in areas of low soil fertility, particularly at lower latitudes, and consequently increase the risk of localized population extinctions from stochastic causes.     

The response of five tropical dicotyledon species to solar ultraviolet-B radiation

Tropical regions currently receive the highest levels of global solar ultraviolet-B radiation (UV-B, 280–320 nm) even without ozone depletion. The influence of natural, present-day UV-B irradiance in the tropics was examined for five tropical species including three native rain forest tree species (Cecropia obtusifolia, Tetragastris panamensis, Calophyilum longifolium) and two economically important species (Swietenia macrophylla, Manihot esculenta). Solar UV-B radiation conditions in a small clearing on Barro Colorado Island, Panama (9° N), were obtained using either a UV-B-excluding plastic film or a film that transmits most of the solar UV-B. Significant differences between UV-B-excluded and near-ambient UV-B plants were often exhibited as increased foliar UV-B absorbing compounds and, in several cases, as reduced plant height with exposure to solar UV-B. Increases in leaf mass per area and reductions in leaf blade length under solar UV-B occurred less frequently. Biomass and photosystem II function using chlorophyll a fluorescence were generally unaffected. The results of this study provide evidence that tropical vegetation, including native rain forest species, responds to the present level of natural solar UV-B radiation. This suggests that even minor ozone depletion in the tropics may have biological implications.     

Direct and indirect effects of solar ultraviolet-B radiation on long-term decomposition

As a result of stratospheric ozone depletion, more solar ultraviolet-B radiation (UV-B, 280–315 nm) is reaching the Earth's surface. Enhanced levels of UV-B may, in turn, alter ecosystem processes such as decomposition. Solar UV-B radiation could affect decomposition both indirectly, by changes in the chemical composition of leaves during growth, or directly by photochemical breakdown of litter and through changes in decomposer communities exposed to sunlight. In this experiment, we studied indirect and direct effects of solar UV-B radiation on decomposition of barley (Hordeum vulgare). We used barley straw and leaf litter grown under reduced UV-B (20% of ambient UV-B) or under near-ambient UV-B (90% of ambient UV-B) in Buenos Aires, Argentina, and decomposed the litter under reduced or near-ambient solar UV-B for 29 months in Tierra del Fuego, Argentina. We found that the UV-B treatment applied during growth decreased the decay rate. On the other hand, there was a marginally significant direct effect of elevated UV-B during the early stages of decomposition, suggesting increased mass loss. The effect of UV-B during growth on decomposition was likely the result of changes in plant litter chemical composition. Near-ambient UV-B received during plant growth decreased the concentrations of nitrogen, soluble carbohydrates, and N/P ratio, and increased the concentrations of phosphorus, cellulose, UV-B-absorbing compounds, and lignin/N ratio. Thus, solar UV-B radiation affects the decomposition of barley litter directly and indirectly, and indirect effects are persistent for the whole decomposition period.     

Photosynthetic CO2-use efficiency in lichens and their isolated photobionts: the possible role of a CO2-concentrating mechanism

The CO₂ dependence of net CO₂ assimilation was examined in a number of green algal and cyanobacterial lichens with the aim of screening for the algal/cyanobacterial CO₂-concentrating mechanism (CCM) in these symbiotic organisms. For the lichens Peltigera aphthosa (L.) Willd., P. canina (L.) Willd. and P. neopolydactyla (Gyeln.) Gyeln., the photosynthetic performance was also compared between intact thalli and their respective photobionts, the green alga Coccomyxa PA, isolated from Peltigera aphthosa and the cyanobacterium Nostoc PC, isolated from Peltigera canina. More direct evidence for the operation of a CCM was obtained by monitoring the effects of the carbonic-anhydrase inhibitors acetazolamide and ethoxyzolamide on the photosynthetic CO₂use efficiency of the photobionts. The results strongly indicate the operation of a CCM in all cyanobacterial lichens investigated and in cultured cells of Nostoc PC, similar to that described for free-living species of cyanobacteria. The green algal lichens were divided into two groups, one with a low and the other with a higher CO₂-use efficiency, indicative of the absence of a CCM in the former. The absence of a CCM in the low-affinity lichens was related to the photobiont, because free-living cells of Coccomyxa PA also apparently lacked a CCM. As a result of the postulated CCM, cyanobacterial Peltigera lichens have higher rates of net photosynthesis at normal CO₂ compared with Peltigera aphthosa. It is proposed that this increased photosynthetic capacity may result in a higher production potential, provided that photosynthesis is limited by CO₂ under natural conditions.     

Solar ultraviolet-B radiation can affect slug feeding preference for some plant species native to a fen ecosystem in Tierra del Fuego,Argentina

The objectives of this study were to test potential effects of solar ultraviolet-B (UV-B) radiation on (i) foliage nutritional quality and foliage decomposition rates of six plant species of this fen ecosystem (Nothofagus antarctica, Carex curta, C. decidua and C. magellanica; Acaena magellanica and Gunnera magellanica) and (ii) feeding preferences for these plant species of the slug Deroceras reticulatum prevalent in this ecosystem. In a mixed-diet selection slugs were offered leaves of the six species that had been grown for three years in experimental field plots under either near-ambient or reduced solar ultraviolet-B (UV-B) radiation. The chosen characteristics of leaf quality (nitrogen concentration, carbon:nitrogen ratio, specific leaf area) and leaf decomposition rates of the six species varied significantly among species but were not affected by the UV-B treatments. However, there were UV-B treatment effects on slug feeding preference for two plant species. For the tree species, Nothofagus, slugs had consumed only one-third as much foliage grown under near-ambient UV-B radiation as of foliage grown under reduced UV-B by the end of the feeding experiment. In contrast, leaves of the sedge C. decidua that had been grown under near-ambient UV-B were consumed twice as much as leaves grown under reduced UV-B radiation. Consumption of foliage for the other four species was similar for the two UV-B treatments. Additionally, diet selection of the slugs was also significantly affected by prior UV-B conditions under which foliage had been grown. Nothofagus leaves were consumed proportionately less and C. decidua proportionately more if the foliage had been grown under near-ambient UV-B radiation. This revised version was published online in August 2006 with corrections to the Cover Date.     

TWO NEW SOREDIATE TAXA OFPELTIGERA

Examination of specimens ofPeltigera didactyla, from Africa, Asia, Europe and North America revealed that this sorediate species includes three entities that can be separated on morphological characters. A new species,P. lambinonii, is described from East Africa, and a new combinationP. didactylavar.extenuata, is proposed to accommodate morphs from mesic forest habitats in Asia, Europe and North America. Despite this taxonomic reduction,P. didactylavar.didactylaremains a ubiquitous taxon. The former two taxa often produce methyl gyrophorate, which can co-occur with traces of gyrophoric acid. These tridepsides were only rarely detected in var.didactyla; their occasional presence seems to be best explained by hybridization. The taxonomic and ecological significance of these substances is discussed.     

Symbiosis constraints: Strong mycobiont control limits nutrient response in lichens

Symbioses such as lichens are potentially threatened by drastic environmental changes. We used the lichen Peltigera aphthosa—a symbiosis between a fungus (mycobiont), a green alga (Coccomyxa sp.), and N₂‐fixing cyanobacteria (Nostoc sp.)—as a model organism to assess the effects of environmental perturbations in nitrogen (N) or phosphorus (P). Growth, carbon (C) and N stable isotopes, CNP concentrations, and specific markers were analyzed in whole thalli and the partners after 4 months of daily nutrient additions in the field. Thallus N was 40% higher in N‐fertilized thalli, amino acid concentrations were twice as high, while fungal chitin but not ergosterol was lower. Nitrogen also resulted in a thicker algal layer and density, and a higher δ¹³C abundance in all three partners. Photosynthesis was not affected by either N or P. Thallus growth increased with light dose independent of fertilization regime. We conclude that faster algal growth compared to fungal lead to increased competition for light and CO₂ among the Coccomyxa cells, and for C between alga and fungus, resulting in neither photosynthesis nor thallus growth responded to N fertilization. This suggests that the symbiotic lifestyle of lichens may prevent them from utilizing nutrient abundance to increase C assimilation and growth.     

The reduction of aboveground Calamagrostis epigeios mass and tiller number by enhanced UV-B in a dune-grassland ecosystem

Since early May 1997 dune-grassland vegetation in the Netherlands has been exposed to enhanced levels of ultraviolet-B (UV-B) radiation. Expected increases in the amount of biologically effective UV-B (UV-B_BE) upon a reduction of the stratospheric ozone layer with 15% were calculated and artificially supplemented.In June and September 1998, above- and belowground vegetation samples were taken. Of the dominant species Calamagrostis epigeios and Carex arenaria aboveground mass accumulation, leaf weight (LW), leaf area (LA), specific leaf area (SLA) and tiller number were assessed separate from the remaining vegetation.The results of our study indicate alterations in the vegetation structure due to UV-B supplementation. In June, a significant reduction due to UV-B supplementation in number of tillers and aboveground dry weight per soil area unit was found for C. epigeios. As C. epigeios is the most dominant species of the dune-grassland, these effects indicate a change in vegetation structure due to UV-B enhancement. Indications of UV-B effects on other parameters, such as the number of tillers of C. arenaria and aboveground plant dry weight of the group of species other than C. epigeios and C. arenaria, may also represent changes in vegetation structure. The LA and LW data show the same patterns as the mass accumulation trends. No significant UV-B effects on the SLA of the species or of the total vegetation could be found.Trends in patterns of species dry weight accumulation and partitioning of dry weight between species groups are different in June and September. This may indicate seasonal dependence of UV-B responses. Also, the consistently reducing trend in total and aboveground plant dry weight may indicate deleterious effects of UV-B on total plant matter accumulation. Possible causes of observed trends and effects are discussed.