Micro-site performance of evergreen and deciduous dwarf shrubs in a subarctic heath in relation to nitrogen status

The relationship between micro-site nitrogen status (total soil N conc., total vegetation N, and leaf N conc.) and performance (biomass) for two evergreen and two deciduous dwarf shrubs was studied in a subarctic heath. One deciduous species, Vaccinium uliginosum , dominated on sites having the highest vegetation nitrogen content and highest leaf nitrogen concentrations. The evergreen Empetrum hermaphroditum dominated on sites with a low total vegetation nitrogen content and low leaf nitrogen concentrations. Two other species, V. myrtillus (deciduous) and V. vitis-idaea (evergreen), showed intermediate patterns that were not clearly separated from either V. uliginosum or Empetrum . Soil nitrogen content showed no significant differences between micro-sites. Possible physiological reasons for the observed patterns are discussed.     

Effects of enhanced UV-B radiation and elevated carbon dioxide concentrations on a sub-arctic forest heath ecosystem

An experiment is described which studies the effects of enhanced UV-B radiation (simulating a 15% reduction in the Ozone layer) and elevated atmospheric concentrations of CO₂ (600 ppm) on the dwarf shrub layer of a sub-arctic forest heath ecosystem at Abisko, North Sweden. The experimental treatments were first applied in 1993, and have covered most of the snow-free season (late May to early September) 1993–1995. Effects of the treatments on the four dwarf shrub species have been recorded largely using non-destructive measures (Vaccinium uliginosum, Vaccinium myrtillus – deciduous species and Vaccinium vitis-idaea and Empetrum hermaphroditum – evergreen species). Effects of the treatments on stem growth and leaf thickness have so far been small, although CO₂ treatments initially stimulated stem extension in Vaccinium myrtillus 1993 and depressed growth in V. vitis idaea in 1994 and E. hermaphroditum during 1995. UV-B treatments stimulated fruit production in V. myrtillus in both 1994 and 1995, but there was no effect on reproductive phenology. There were also marked effects of UV-B treatments on insect herbivory in the deciduous dwarf shrubs; with leaf area loss being greater than the control in the UV-B treatment in V. myrtillus and less in V. uliginosum. The results point to the possibility of important effects of the treatments on physiological and chemical processes within the plants. The ecological results of such effects may not be immediately apparent, but may be far reaching, pointing to the need for long-term in situ experimentation in predicting the effects of these global change variables.     

The functional quality of decomposing litter outputs from an Arctic plant community is affected by long-term exposure to enhanced UV-B

Plant exposure to enhanced UV-B radiation typically induces changes in leaf secondary metabolite profiles which will be inherited in litter, affecting litter breakdown and the carbon (C) dynamics of sensitive plant communities. A key enzyme in the decomposition process is phenol oxidase which is influenced by litter quality and, hence, a decomposition bioindicator. Here we investigated dwarf shrub litter decomposition following experimental community exposure to enhanced UV-B over two decades in the Swedish sub-Arctic. We examined the hypothesis that foliar UV-B exposure would alter litter quality to elevate phenol oxidase activity. This was tested in the field by measuring phenol oxidase activity in freshly collected mixed-community litter from under our experimental vegetation. A laboratory mesocosm was next used in a decomposition assay to investigate individual species responses over eight weeks, with an emphasis on the quality of leachate outputs from decomposing litter (from Empetrum hermaphroditum, Vaccinium vitis-idaea, Vaccinium uliginosum). In the assay bi-weekly collections of leachate were analysed for phenol oxidase activity, together with total phenolics and dissolved organic C (DOC). At the end of the assay litter mass loss and respired C were also determined. The initial assessment on field mixed-community litter found no enhanced UV-B treatment (henceforth: ‘UV-B treatment’) effect on phenol oxidase activity. However, in the controlled laboratory mesocosm assay, significant species-specific effects of the UV-B treatment were evident, with increased phenol oxidase activity in V. vitis-idaea leachate (P < 0.001) and a significant reduction (P = 0.05) in respired C. Leachate DOC release from the UV-B treatment was greater in both Vaccinium species and the effect on V. uliginosum was significant (P < 0.05). The UV-B treatment had no effect on the total phenolic concentration of litter or leachates for any species, but there were significant differences in leachate total phenolics, both over time and between species. Also the initial phenolic concentration in leachates from the decomposing litter of E. hermaphroditum was greater than both Vaccinum species. Results suggest a species specific role for UV-B in influencing enzyme function and decomposition, dependent on individual traits. This has implications for decomposition dynamics in this system and more widely. Our study highlights the value of using a laboratory assay to gain a mechanistic understanding the species level impacts of a global change factor (UV-B) on decomposition, which are otherwise obscured by community-level responses and difficult to determine under field conditions.     

The effects of seasons, exposure, enhanced UV-B radiation, and water stress on leaf epicuticular and internal UV-B absorbing capacity of Cistus creticus: a Mediterranean field study

The seasonal variation in leaf epicuticular and internal (cellular)UV-B absorbing compounds was followed in field-grown shrubsof Cistus creticus L. Fully exposed and permanently shaded (undera canopy of evergreen sclerophylls) individuals were used. Nearmonthly measurements were performed for three years. In addition,a 2-year field experiment was established, in which fully exposedplants received ambient or ambient plus supplemental UV-B radiation,simulating a 15% ozone depletion over Patras (38.3 N, 29.1E). A second, six-month field experiment investigated the effectsof additional summer irrigation on UV-B absorbing compounds.It was found that: (a) exposed plants had considerably higherconcentrations of UV-B absorbing compounds (both epicuticularand internal), irrespective of season, (b) exposed and shadedplants exhibited dramatic and yearly consistent seasonal fluctuationsin epicuticular material with a c. 18-fold difference betweenthe summer maxima and winter minima. Internal UV-B absorbingcompounds showed the same seasonal trends, but with far lesspronounced differences, (c) In the 2-year field experiment withenhanced UV-B radiation, an increase in both epicuticular andinternal UV-B absorbing capacity was evident only during thesummer dry period of the first year. Later on, however, andup to the termination of the experiment, the differences wereabolished, (d) Well-watered and water-stressed plants showedthe same levels of both epicuticular and internal UV-B absorbingcompounds. Although the results indicate that concentrations of epicuticularand, to a lesser extent, internal UVB absorbing compounds fluctuateaccording to the seasonally and site-dependent environmentalstresses, no indications for an effect of water availabilitywere obtained. Therefore it may be concluded that the main determinantof the levels of UVB absorbing compounds is the intensity ofsolar radiation. The relative influence of the various spectralregions of solar radiation cannot be assessed from the presentresults. It was predicted, however, that the effects of increasedsolar UV-B radiation, resulting from the expected ozone depletionover the next decades, will be negligible and probably maskedby the much larger seasonal and site-dependent changes. Key words: Epicuticular, exposure, seasons, UVB absorbing compounds, UVB radiation     

Increased exposure to UV-B radiation during early development leads to enhanced photoprotection and improved long-term performance in Lactuca sativa

Plant responses to solar UV radiation are numerous and have often been considered from a perspective of negative outcomes for plant productivity. In this study, we used two experimental approaches consisting of: (1) field-based spectrally modifying filters in addition to (2) controlled indoor exposure to UV-B, to examine the effects of UV radiation on growth and photosynthetic performance of lettuce (Lactuca sativa L.) seedlings. Various aspects of growth were affected in plants grown under a UV-inclusive environment compared to a UV-depleted environment, including reductions in leaf expansion, increases in leaf thickness and the rate of net photosynthesis. After transplantation to a uniform field environment, lettuce plants initially propagated under the UV-inclusive environment exhibited higher harvestable yields than those from a UV-depleted environment. In controlled conditions, photosynthetic rates were higher in plants grown in the presence of UV-B radiation, and relative growth of plants pre-acclimatized to UV-B was also increased, in addition to higher maximum photochemical efficiency of photosystem II (PSII) (F(v) /F(m) ) following subsequent exposure to high photosynthetically active radiation (PAR) and temperature stress. Our findings are discussed within the context of sustainability in agriculture and the paradigm shift in photobiology which such beneficial responses to UV radiation could represent.     

Few long-term effects of simulated climate change on volatile organic compound emissions and leaf chemistry of three subarctic dwarf shrubs

Climate change is exposing arctic ecosystems to higher temperature, increased nutrient availability and shading due to the increasing cloud cover and the expanding forests. In this work, we assessed how these factors affect the emissions of biogenic volatile organic compounds (BVOCs) from three subarctic dwarf shrub species in a field experiment after 18 treatment years. Of the studied species the willow Salix phylicifolia L. was the only isoprene-emitter with an emission potential of 16.1 ± 8.4 μg g⁻¹ dw h⁻¹ (at 30 °C and photosynthetic photon flux density of 1000 μmol m⁻² s⁻¹). The dwarf birch Betula nana L. had significant emissions of various reactive BVOCs, including monoterpenes and sesquiterpenes. The evergreen Cassiope tetragona (L.) D. Don emitted high amounts of mono- and sesquiterpenes. Due to chance, the temperature in the warming treatment (employing open-top plastic tents) and the unwarmed treatments was similar at the time of the measurements, and therefore long-term indirect effects of warming could be assessed without interference of temperature differences at the time of measurement. The long-term warming had not altered foliar N, P or condensed tannin concentrations, but it had led to other chemical changes detected in the near-infrared reflectance spectra of the leaves. Nevertheless, there were no significant differences in the BVOC emissions per unit leaf mass measured by the dynamic enclosure method and gas chromatography-mass spectrometry. Annual additions of NPK fertilizer, which mimicked increased nutrient availability, had accumulated P in the leaves of all species. In addition, fertilization marginally increased the leaf N concentration of B. nana. The only significant fertilization effect on BVOC emissions was a stimulation of emission of the sesquiterpene β-selinene from S. phylicifolia. The shading treatment obtained by dome-shaped hessian tents did not cause clear long-term changes in leaf chemistry or BVOC emissions. The only observed change was a marginally significant increase in sesquiterpene emissions from B. nana. When the treatment effects on long-term biomass changes in the different treatments were taken into account by proportioning the BVOC emissions to the biomass of each species in the field treatments, warming led to a significant increase and shading to a decrease in the total BVOC emissions per unit ground area from B. nana. These results highlight the importance of an integrated approach for realistic assessment of responses to climate change.     

Repeated exposure to enhanced UV-B radiation in successive generations increases developmental instability (leaf fluctuating asymmetry) in a desert annual

Populations of the desert annual Dimorphotheca sinuata , derived from a common seed stock, were exposed concurrently over four successive generations to either ambient (representing no stratospheric ozone depletion) or elevated (representing 20% stratospheric ozone depletion) UV-B levels during their complete life cycle. Leaf fluctuating asymmetry (FA) was measured in populations of plants grown from seeds of selected generations which had experienced different UV-B exposure histories, and from seeds collected from a wild population of this species which grows in a naturally enhanced UV-B environment. These measured plants had been grown in a greenhouse under essentially UV-B-free conditions. Leaf FA was significantly increased by greater numbers of enhanced UV-B exposures in the parentage of the seed. There was a linear to exponential dose–response relationship between number of UV-B exposure iterations in seed parentage and leaf FA, suggesting that damage to DNA caused by UV-B exposure during plant development may not be fully repaired, and thus be inherited by offspring and accumulated over successive generations in this species. Leaf FA of plants grown from seed from the wild population was not significantly greater than that of control plants whose parentage experienced only ambient UV-B exposures, although this negative result may have been due to low sampling intensity and measurement resolution, and the relatively low UV-B enhancement experienced by the wild population. We conclude that leaf FA may constitute a relatively sensitive yet inexpensive means of quantifying UV-B damage to plants.     

Leafing patterns and leaf traits of four evergreen shrubs in the Patagonian Monte, Argentina

We assessed leafing patterns (rate, timing, and duration of leafing) and leaf traits (leaf longevity, leaf mass per area and leaf-chemistry) in four co-occurring evergreen shrubs of the genus Larrea and Chuquiraga (each having two species) in the arid Patagonian Monte of Argentina. We asked whether species with leaves well-defended against water shortage (high LMA, leaf longevity, and lignin concentration, and low N concentration) have lower leaf production, duration of the leafing period, and inter-annual variation of leafing than species with the opposite traits. We observed two distinctive leafing patterns each related to one genus. Chuquiraga species produced new leaves concentrated in a massive short leafing event (5–48 days) while new leaves of Larrea species emerged gradually (128–258 days). Observed leafing patterns were consistent with simultaneous and successive leafing types previously described for woody plants. The peak of leaf production occurred earlier in Chuquiraga species (mid September) than in Larrea species (mid October–late November). Moreover, Chuquiraga species displayed leaves with the longest leaf lifespan, while leaves of Larrea species had the lowest LMA and the highest N and soluble phenolics concentrations. We also observed that only the leaf production of Larrea species increased in humid years. We concluded that co-occurring evergreen species in the Patagonian Monte displayed different leafing patterns, which were associated with some relevant leaf traits acting as plant defenses against water stress and herbivores. Differences in leafing patterns could provide evidence of ecological differentiation among coexisting species of the same life form.     

增强紫外—B对反枝苋形态、生理及异速生长的影响

在田间条件下,模拟西安地区21.6%的臭氧层减薄,研究增强紫外-B辐射 280～320nm,3.18kJ·m-2·d-1 对双子叶阔叶杂草反枝苋 Amaranthusretroflexus 生理、形态及异速生长的影响.结果表明: 1 与对照相比,处理组的叶绿素、类胡萝卜素含量降低,但叶片紫外吸收物质的含量增加; 2 处理组的株高、叶数及单株重有明显降低; 3 株高与单株重的线性关系有较大的偏离,表现在同等株高下处理组的生物量低于对照.这些表明在补充的紫外-B条件下,反枝苋的形态有较大的可塑性,并进一步会影响该植物在群落中的竞争能力.     

Effects of nutrient addition on leaf chemistry,morphology, and photosynthetic capacity of three bog shrubs

Plants in nutrient-poor environments typically have low foliar nitrogen (N) concentrations, long-lived tissues with leaf traits designed to use nutrients efficiently, and low rates of photosynthesis. We postulated that increasing N availability due to atmospheric deposition would increase photosynthetic capacity, foliar N, and specific leaf area (SLA) of bog shrubs. We measured photosynthesis, foliar chemistry and leaf morphology in three ericaceous shrubs (Vaccinium myrtilloides, Ledum groenlandicum and Chamaedaphne calyculata) in a long-term fertilization experiment at Mer Bleue bog, Ontario, Canada, with a background deposition of 0.8 g N m⁻² a⁻¹. While biomass and chlorophyll concentrations increased in the highest nutrient treatment for C. calyculata, we found no change in the rates of light-saturated photosynthesis (A _max), carboxylation (V _cmax)_, or SLA with nutrient (N with and without PK) addition, with the exception of a weak positive correlation between foliar N and A _max for C. calyculata, and higher V _cmax in L. groenlandicum with low nutrient addition. We found negative correlations between photosynthetic N use efficiency (PNUE) and foliar N, accompanied by a species-specific increase in one or more amino acids, which may be a sign of excess N availability and/or a mechanism to reduce ammonium (NH₄) toxicity. We also observed a decrease in foliar soluble Ca and Mg concentrations, essential minerals for plant growth, but no change in polyamines, indicators of physiological stress under conditions of high N accumulation. These results suggest that plants adapted to low-nutrient environments do not shift their resource allocation to photosynthetic processes, even after reaching N sufficiency, but instead store the excess N in organic compounds for future use. In the long term, bog species may not be able to take advantage of elevated nutrients, resulting in them being replaced by species that are better adapted to a higher nutrient environment.     

Maintenance of C sinks sustains enhanced C assimilation during long-term exposure to elevated [CO2] in Mojave Desert shrubs

During the first few years of elevated atmospheric [CO(2)] treatment at the Nevada Desert FACE Facility, photosynthetic downregulation was observed in desert shrubs grown under elevated [CO(2)], especially under relatively wet environmental conditions. Nonetheless, those plants maintained increased A (sat) (photosynthetic performance at saturating light and treatment [CO(2)]) under wet conditions, but to a much lesser extent under dry conditions. To determine if plants continued to downregulate during long-term exposure to elevated [CO(2)], responses of photosynthesis to elevated [CO(2)] were examined in two dominant Mojave Desert shrubs, the evergreen Larrea tridentata and the drought-deciduous Ambrosia dumosa, during the eighth full growing season of elevated [CO(2)] treatment at the NDFF. A comprehensive suite of physiological processes were collected. Furthermore, we used C labeling of air to assess carbon allocation and partitioning as measures of C sink activity. Results show that elevated [CO(2)] enhanced photosynthetic performance and plant water status in Larrea, especially during periods of environmental stress, but not in Ambrosia. δ(13)C analyses indicate that Larrea under elevated [CO(2)] allocated a greater proportion of newly assimilated C to C sinks than Ambrosia. Maintenance by Larrea of C sinks during the dry season partially explained the reduced [CO(2)] effect on leaf carbohydrate content during summer, which in turn lessened carbohydrate build-up and feedback inhibition of photosynthesis. δ(13)C results also showed that in a year when plant growth reached the highest rates in 5 years, 4% (Larrea) and 7% (Ambrosia) of C in newly emerging organs were remobilized from C that was assimilated and stored for at least 2 years prior to the current study. Thus, after 8 years of continuous exposure to elevated [CO(2)], both desert perennials maintained their photosynthetic capacities under elevated [CO(2)]. We conclude that C storage, remobilization, and partitioning influence the responsiveness of these desert shrubs during long-term exposure to elevated [CO(2)].     

UV-B辐射增强对反枝苋形态、生理及化学成分的影响

通过大田盆栽实验,研究了2种增强的UV-B辐射对双子叶C4植物反枝苋(Amaranthus retroflexus)形态、生理、生物量分配及化学成分的影响.结果表明:增强UV-B辐射处理下分枝数、叶片数、高度和基径均没有显著变化.高、低剂量的UV-B处理均导致过氧化氢酶(CAT)活性显著高于对照组(P＜0.05),但对超氧化物歧化酶(SOD)活性无影响,低剂量辐射下过氧化物酶(POD)活性显著高于对照而高剂量辐射下无显著变化,同时丙二醛(MDA)含量随辐射剂量的增加而显著增加(P＜0.05).收获期低剂量辐射处理下的资源分配模式发生变化,分配到叶片的生物量比例显著增加,而分配到根的比例显著减小,但高剂量辐射对生物量分配无影响.UV-B辐射处理下根、茎、叶部所检测的大多数化学成分与对照相比均有显著变化:叶片中可溶性蛋白、木质素和脂肪,根中木质素,茎中纤维素、木质素含量在辐射处理下显著高于对照;叶片中淀粉,茎中可溶性糖、淀粉和脂肪含量在辐射处理下显著低于对照(P＜0.05).     

Microgravity effects on leaf morphology, cell structure, carbon metabolism and mRNA expression of dwarf wheat

The use of higher plants as the basis for a biological life support system that regenerates the atmosphere, purifies water, and produces food has been proposed for long duration space missions. The objective of these experiments was to determine what effects microgravity (μg) had on chloroplast development, carbohydrate metabolism and gene expression in developing leaves of Triticum aestivum L. cv. USU Apogee. Gravity naive wheat plants were sampled from a series of seven 21-day experiments conducted during Increment IV of the International Space Station. These samples were fixed in either 3% glutaraldehyde or RNAlater^™ or frozen at −25°C for subsequent analysis. In addition, leaf samples were collected from 24- and 14-day-old plants during the mission that were returned to Earth for analysis. Plants grown under identical light, temperature, relative humidity, photoperiod, CO₂, and planting density were used as ground controls. At the morphological level, there was little difference in the development of cells of wheat under μg conditions. Leaves developed in μg have thinner cross-sectional area than the 1 g grown plants. Ultrastructurally, the chloroplasts of μg grown plants were more ovoid than those developed at 1 g, and the thylakoid membranes had a trend to greater packing density. No differences were observed in the starch, soluble sugar, or lignin content of the leaves grown in μg or 1 g conditions. Furthermore, no differences in gene expression were detected leaf samples collected at μg from 24-day-old leaves, suggesting that the spaceflight environment had minimal impact on wheat metabolism.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Acetylcholinesterase activation and enhanced lipid peroxidation after long-term exposure to low levels of aluminum on different mouse brain regions

Aluminum (Al), oxidative stress and impaired cholinergic functions have all been related to Alzheimer's disease (AD). The present study evaluates the effect of aluminum on acetylcholinesterase (AChE) and lipid peroxidation in the mouse brain. Mice were loaded by gavage with Al 0.1 mmol/kg/day 5 days per week during 12 weeks. The mice were divided into four groups: (1) control; (2) 10 mg/mL of citrate solution; (3) 0.1 mmol/kg of Al solution; (4) 0.1 mmol/kg of Al plus 10 mg/mL of citrate solution. AChE activity was determined in the hippocampus, striatum, cortex, hypothalamus and cerebellum and lipid peroxidation was determined in the hippocampus, striatum and cortex. An increase of AChE activity was observed in the fourth group (Al + Ci) in the hippocampus (36%), striatum (54%), cortex (44%) and hypothalamus (22%) (p<0.01). The third group (Al) presented a decrease of AChE activity in the hypothalamus (20%) and an enhancement in the striatum (27%). Lipid peroxidation, measured by TBARS (thiobarbituric acid reactive substances), was elevated in the hippocampus and cerebral cortex when compared with the control (p < 0.01). The effect of aluminum on AChE activity may be due to a direct neurotoxic effect of the metal or perhaps a disarrangement of the plasmatic membrane caused by increased lipid peroxidation.     

The effect of exposure to enhanced UV-B radiation on the penetration of monochromatic and polychromatic UV-B radiation in leaves of Brassica napus

Using quartz optical fibres, penetration of both monochromatic (310 nm) and polychromatic UV-B (280–320 nm) radiation in leaves of Brassica napus L. (cv. Ceres) was measured. Plants were grown under either visible light (750 μmol m⁻² s⁻¹ photosynthetically active radiation) or with the addition of 8. 9 KJ m⁻² day⁻¹ biologically effective UV-B (UV-B_BE) radiation. Results showed that of the 310 nm radiation that penetreated the leaf, 90% was within the intial one third of the leaf with high attenuation in the leaf epidermis, especially in UV-treated plants. Polychromatic UV-B radiation, relative to incident radiation, showed a relatively uniform spectral distribution within the leaf, except for collimated radiation. Over 30% of the UV-screening pigments in the leaf, including flavonoids, were found in the adaxial epidermal layer, making this layer less transparent to UV-B radiation than the abaxial epidermis, which contained less than 12% of the UV-screening pigments. UV-screening pigments increased by 20% in UV-treated leaves relative to control leaves. Densely arranged epicuticular wax on the adaxial leaf surface of UV-treated plants may have further decreased penetration of UV-B radiation by reflectance. An increased leaf thickness, and decreases in leaf area and leaf dry weight were also found for UV-treated plants.     

Increased prothrombin time and metabolic changes with high serum aluminum levels following long-term exposure to Bayer-process alumina

Serum aluminum levels twice to three times those in controls were found in 30 of 36 workers in a factory in an atmosphere of alumina dust. Inorganic phosphate and total alkaline phosphatase levels were within normal range, with no clinical evidence of phosphate depletion syndrome. Serum intestinal alkaline phosphatase, acid phosphatase and adenosine triphosphate levels were significantly reduced in the group with high levels of aluminum. The finding that the mean prothrombin time was significantly prolonged is of particular interest with respect to beneficial antithrombogenic effect.