UV‐B effect on Quercus robur leaf litter decomposition persists over four years

The effects of elevated UV‐B (280–315 nm) radiation on the long‐term decomposition of Quercus robur leaf litter were assessed at an outdoor facility in the UK by exposing saplings to elevated UV‐B radiation (corresponding to a 30% increase above the ambient level of erythemally weighted UV‐B, equivalent to that resulting from a c. 18% reduction in ozone column) under arrays of cellulose diacetate‐filtered fluorescent UV‐B lamps that also produced UV‐A radiation (315–400 nm). Saplings were also exposed to elevated UV‐A radiation alone under arrays of polyester‐filtered fluorescent lamps and to ambient solar radiation under arrays of nonenergized lamps. After 8 months of irradiation, abscised leaves were placed into litter bags and allowed to decompose in the litter layer of a mixed deciduous woodland for 4.08 years. The dry weight loss of leaf litter from saplings irradiated with elevated UV‐B and UV‐A radiation during growth was 17% greater than that of leaf litter irradiated with elevated UV‐A radiation alone. Annual fractional weight loss of litter (k), and the estimated time taken for 95% of material to decay (3/k) were respectively increased and decreased by 27% for leaf litter exposed during growth to elevated UV‐B and UV‐A radiation, relative to that exposed to UV‐A alone. The present data corroborate those from a previous study indicating that UV‐B radiation applied during growth accelerates the subsequent decomposition of Q. robur leaf litter in soil, but indicate that this effect persists for over four years after abscission.     

Impact of changes in natural ultraviolet radiation on pigment composition, physiological and morphological characteristics of the Antarctic moss, Grimmia antarctici

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.     

Elevated UV-B radiation alters fluxes of methane and carbon dioxide in peatland microcosms

Increases in solar ultraviolet‐B radiation (UV‐B; 280–320 nm) reaching the earth have been estimated to continue until 2050s in the boreal and subarctic regions with an abundant peatland cover. Peatlands are significant sinks for carbon dioxide (CO₂) and sources for methane (CH₄). To assess whether the future increases in UV‐B could affect the fluxes of CO₂ and CH₄ in peatlands via an impact on vegetation, we exposed peatland microcosms to modulated 30% supplementation of erythemally weighted UV‐B at an outdoor facility for one growing season. The experimental design included appropriate controls for UV‐A and ambient radiation. The UV‐B caused a significant reduction in gross photosynthesis, net ecosystem CO₂ exchange, and CH₄ emission of the peatland microcosms. These changes in the carbon gas cycling can be partly explained by UV‐B‐induced morphological changes in Eriophorum vaginatum which acts as a conduit for CH₄. Leaf cross section and the percentage of CH₄‐conducting aerenchymatous tissue in E. vaginatum were significantly reduced by UV‐B. Methanol‐extractable UV‐B absorbing compounds decreased under both UV‐B and UV‐A in Sphagnum angustifolium, and tended to accumulate under UV‐B in S. papillosum. Membrane permeability to magnesium (Mg) and calcium (Ca) ions was higher in UV‐B exposed S. angustifolium. Amount of chlorophyll and carotenoid pigments was increased by UV‐A in S. magellanicum. The observed changes in Sphagnum mosses did not coincide with those in carbon gas fluxes but occurred at the time of the highest UV intensity in the mid summer. Our findings indicate that increasing UV‐B may have more substantial effects on gas exchange in peatlands than previously thought.     

Differences in growth and physiological traits of two poplars originating from different altitudes as affected by UV‐B radiation and nutrient availability

Cuttings of Populus kangdingensis and Populus cathayana originating from altitudes of 3500 and 1500 m in southwestern China, respectively, were grown for one growing season in the field under ambient or ambient plus supplemental ultraviolet‐B (UV‐B) radiation with two levels of nutrients. In both species, enhanced UV‐B radiation significantly increased UV‐B absorbing compounds and guaiacol peroxidase (GPX) activity, while no significant effects were observed in photosynthetic pigments and proline content. On the other hand, cuttings grown with high‐nutrient availability had larger leaf area, higher total biomass and GPX activity as well as higher water use efficiency (WUE) (as measured by stable carbon isotope composition, δ¹³C) when compared with low‐nutrient conditions, while UV‐B absorbing compounds and ascorbic acid (AsA) content significantly decreased. Differences in responses to enhanced UV‐B radiation and nutrient availability were observed between the two species. Nutrient‐induced increases in chlorophyll a, chlorophyll b and total chlorophyll as well as in carotenoids were greater in P. kangdingensis than in P. cathayana. In P. cathayana, enhanced UV‐B radiation significantly decreased leaf area and total biomass, while it significantly increased WUE and ascorbate peroxidase (APX). In contrast, such changes were not observed in P. kangdingensis. In addition, the effects of enhanced UV‐B radiation on leaf area, total biomass and UV‐B absorbing compounds were closely related to the nutrient status. Our results indicated that P. kangdingensis, which originates from the altitude of 3500 m and is apparently adapted to low‐nutrient and high‐UV‐B habitats, exhibits better tolerance to enhanced UV‐B radiation and greater growth under low‐nutrient availability than does P. cathayana originating from the altitude of 1500 m.     

Elevated UV-B radiation incident on Quercus robur leaf canopies enhances decomposition of resulting leaf litter in soil

We examined whether the exposure of Quercus robur L. to elevated UV-B radiation (280–315 nm) during growth would influence leaf decomposition rate through effects on litter quality. Saplings were exposed for eight months at an outdoor facility in the UK to a 30% elevation above the ambient level of erythemally weighted UV-B radiation under UV-B treatment arrays of fluorescent lamps filtered with cellulose diacetate, which transmitted both UV-B and UV-A (315–400 nm) radiation. Saplings were exposed to elevated UV-A alone under control arrays of lamps filtered with polyester and to ambient radiation under unenergised arrays of lamps. Abscised leaves from saplings were enclosed in 1 mm² mesh nylon bags, placed in a Quercus–Fraxinus woodland and were sampled at 0.11, 0.53, 1.10 and 1.33 years for dry weight loss, chemical composition and saprotrophic fungal colonization. At abscission, litters from UV-A control arrays had ≈ 7.5% higher lignin/nitrogen ratios than those from UV-B treatment and ambient arrays (P < 0.06). Dry weight loss of leaves treated with elevated UV-B radiation during growth was 2.5% and 5% greater than that of leaves from UV-A control arrays at 0.53 and 1.33 years, respectively. Litter samples from UV-B treatment arrays lost more nitrogen and phosphorus than samples from ambient arrays and more carbon than samples from UV-A control arrays. The annual fractional weight loss of litter from UV-B treatment arrays was 8% and 6% greater than that of litter from UV-A control and ambient arrays, respectively. Regression analyses indicated that the increased decomposition rate of UV-B treated litters was associated with enhanced colonization of leaves by basidiomycete fungi, the most active members of the soil fungal community, and that the frequency of these fungi was negatively associated with the initial lignin/nitrogen ratio of leaves.     

Nitric oxide is involved in integration of UV‐B absorbing compounds among parts of clonal plants under a heterogeneous UV‐B environment

In nature, ultraviolet‐B (UV‐B) radiation is highly heterogeneous, both spatially and temporally. Plants exposed to UV‐B radiation produce UV‐B absorbing compounds that function as a protective filter. For clonal plants under heterogeneous UV‐B radiation conditions, integration among ramets can allow irradiated ramets to benefit un‐irradiated ramets by causing them to increase their UV‐B absorbing compounds content. In this study, we evaluated integration between pairs of clonal ramets of Glechoma longituba under heterogeneous or homogeneous UV‐B conditions. We determined the levels of UV‐B absorbing compounds, nitric oxide (NO) and hydrogen peroxide (H₂O₂) and measured the activity of phenylalanine ammonia‐lyase (PAL) in connected ramet pairs under homogeneous or heterogeneous UV‐B conditions. Under heterogeneous UV‐B conditions, the UV‐B absorbing compounds content increased in leaves of irradiated and un‐irradiated ramets, but not in the connecting stolons. The NO content increased in irradiated and un‐irradiated leaves and stolons, but the H₂O₂ content did not. Application of NO synthesis inhibitors and an NO blocker to irradiated ramets blocked the increase in UV‐B absorbing compounds and PAL activity in un‐irradiated ramets. These results suggested that NO is involved in the integration process for UV‐B absorbing compounds among ramets. Our findings suggested that a UV‐B‐induced increase in NO transmits a signal to un‐irradiated ramets via the stolon, leading to an increase in PAL activity and UV‐B absorbing compounds content. The internal translocation of signal enables members of clonal networks to function as a whole unit and to mount an efficient defensive response to localized UV‐B radiation.     

UV‐B INDUCTION OF UV‐B PROTECTION IN ULVA PERTUSA (CHLOROPHYTA)1

The green macroalga Ulva pertusa Kjellman produced UV‐B absorbing compounds with a prominent absorption maximum at 294 nm in response only to UV‐B, and the amounts induced were proportional to the UV‐B doses. Under a 12:12‐h light:dark regime, the production of UV‐absorbing compounds occurred only during the exposure periods with little turnover in the dark. There was significant reduction in growth in parallel with the production of UV‐B absorbing compounds. The polychromatic action spectrum for the induction of UV‐B absorbing compounds in U. pertusa exhibits a major peak at 292 nm with a smaller peak at 311.5 nm. No significant induction was detected above 354.5 nm, and radiation below 285 nm caused significant reduction in the levels of UV‐B absorbing compounds. After UV‐B irradiation at 1.0 W·m^?2 for 9 h, the optimal photosynthetic quantum yield of the samples with UV‐B absorbing compounds slightly increased relative to the initial value, whereas that of thalli lacking the compounds declined to 30%–34% of the initial followed by subsequent recovery in dim light of up to 84%–85% of the initial value. There was a positive and significant relationship between the amount of UV‐B absorbing compounds with antioxidant activity as determined by the α,α‐diphenyl‐β‐picrylhydrazyl scavenging assay. In addition to mat‐forming characteristics and light‐driven photorepair, the existence and antioxidant capacity of UV‐B absorbing compounds may confer U. pertusa a greater selective advantage over other macroalgae, thereby enabling them to thrive in the presence of intense UV‐B radiation.     

Effects of elevated ultraviolet radiation on Quercus robur and its insect and ectomycorrhizal associates

Saplings of pedunculate oak (Quercus robur L.) were exposed at an outdoor facility to modulated levels of elevated UV-B radiation (280–315 nm) under treatment arrays of cellulose diacetate-filtered fluorescent lamps which also produced UV-A radiation (315–400 nm). Saplings were also exposed to UV-A radiation alone under control arrays of polyester-filtered lamps and to ambient levels of solar radiation under arrays of unenergized lamps. The UV-B treatment corresponded to a 30% elevation above the ambient level of erythemally weighted UV-B radiation. Sapling growth and the occurrence of associated organisms were examined over two years. In both years, leaves of saplings exposed to UV-B treatment were thicker and smaller in area relative to leaves exposed to ambient and control levels of radiation. UV-B treatment also retarded bud burst at one sampling in the first year of the study. Some responses were recorded which were common to both treatment and control arrays, implying that UV-A radiation, or some other factor associated with energized lamps, was responsible for the observed effects. Saplings under treatment and control arrays were taller in the first year of the study, suffered greater herbivory from chewing insects, and had lower root dry weights and greater insertion heights of secondary branches than saplings exposed to ambient levels of radiation. Exposure of saplings to elevated UV-A radiation alone under control arrays increased estimated leaf volumes in the second year of the study and reduced the number of secondary branches and the total number of branches per sapling after two years, relative to both treatment and ambient arrays. There were no effects of elevated ultraviolet radiation on shoot or total plant weight, root/shoot ratios, stem diameter, the numbers or insertion heights of primary or tertiary branches, total leaf number, timing of leaf fall or frequency of ectomycorrhizas. Our study suggests that any increases in UV-B radiation as a result of stratospheric ozone depletion will influence the growth of Q. robur primarily through effects on leaf morphology.     

DNA damage and photosynthesis in Antarctic and Arctic Sanionia uncinata (Hedw.) Loeske under ambient and enhanced levels of UV-B radiation

The response of the bipolar moss Sanionia uncinata (Hedw.) Loeske to ambient and enhanced UV‐B radiation was investigated at an Antarctic (Léonie Island, 67°35′ S, 68°20′ W) and an Arctic (Ny‐Alesund, 78°55′ N, 11°56′ E) site, which differed in ambient UV‐B radiation (UV‐BR: 280–320 nm) levels. The UV‐BR effects on DNA damage and photosynthesis were investigated in two types of outdoor experiments. First of all, sections of turf of S. uncinata were collected in an Arctic and Antarctic field site and exposed outdoors to ambient and enhanced UV‐BR for 2 d using UV‐B Mini‐lamps. During these experiments, chlorophyll a fluorescence, chlorophyll concentration and cyclobutyl pyrimidine dimer (CPD) formation were measured. Secondly, at the Antarctic site, a long‐term filter experiment was conducted to study the effect of ambient UV‐BR on growth and biomass production. Additionally, sections of moss turf collected at both the Antarctic and the Arctic site were exposed to UV‐BR in a growth chamber to study induction and repair of CPDs under controlled conditions. At the Antarctic site, a summer midday maximum of 2·1 W m^?2 of UV‐BR did not significantly affect effective quantum yield (ΔF/F_m′) and the ratio of variable to maximal fluorescence (F_v/F_m). The same was found for samples of S. uncinata exposed at the Arctic site, where summer midday maxima of UV‐BR were about 50% lower than at the Antarctic site. Exposure to natural UV‐BR in summer did not increase CPD values significantly at both sites. Although the photosynthetic activity remained largely unaffected by UV‐B enhancement, DNA damage clearly increased as a result of UV‐B enhancement at both sites. However, DNA damage induced during the day by UV‐B enhancement was repaired overnight at both sites. Results from the long‐term filter experiment at the Antarctic site indicated that branching of S. uncinata was reduced by reduction of ambient summer levels of UV‐BR, whereas biomass production was not affected. Exposure of specimens collected from both sites to UV‐BR in a growth chamber indicated that Antarctic and Arctic S. uncinata did not differ in UV‐BR‐induced DNA damage. It was concluded that S. uncinata from both the Antarctic and the Arctic site is well adapted to ambient levels of UV‐BR.     

Functional acclimation to solar UV-B radiation in Gunnera magellanica, a native plant species of southernmost Patagonia

The ecosystems of Tierra del Fuego (in southern Patagonia, Argentina) are seasonally exposed to elevated levels of ultraviolet‐B radiation (UV‐B: 280–315 nm), due to the passage of the ‘ozone hole’ over this region. In the experiments reported in this article the effects of solar UV‐B and UV‐A (315–400 nm) on two UV‐B defence‐related processes: the accumulation of protective UV‐absorbing compounds and DNA repair, were tested. It was found that the accumulation of UV‐absorbing sunscreens in Gunnera magellanica leaves was not affected by plant exposure to ambient UV radiation. Photorepair was the predominant mechanism of cyclobutane‐pyrimidine dimer (CPD) removal in G. magellanica. Plants exposed to solar UV had higher CPD repair capacity under optimal conditions of temperature (25 °C) than plants grown under attenuated UV. There was no measurable repair at 8 °C. The rates of CPD repair in G. magellanica plants were modest in comparison with other species and, under equivalent conditions, were about 50% lower than the repair rates of Arabidopsis thaliana (Ler ecotype). Collectively our results suggest that the susceptibility of G. magellanica plants to current ambient levels of solar UV‐B in southern Patagonia may be related to a low DNA repair capacity.     

Non-invasive measurements of leaf epidermal transmittance of UV radiation using chlorophyll fluorescence: field and laboratory studies

Ratios of chlorophyll fluorescence induced by ultraviolet (UV) and bluegreen (BG) radiation [F(UV)/F(BG)] were determined with a Xe‐PAM fluorometer to test the utility of this technique as a means of non‐intrusively assessing changes in the pigmentation and optical properties of leaves exposed to varying UV exposures under laboratory and field conditions. For plants of Vicia faba and Brassica campestris, grown under controlled‐environmental conditions, F(UV‐B)/F(BG) was negatively correlated with whole‐leaf UV‐B‐absorbing pigment concentrations. Fluorescence ratios of V. faba were similar to, and positively correlated with (r²=0.77 [UV‐B]; 0.85 [UV‐A]), direct measurements of epidermal transmittance made with an integrating sphere. Leaves of 2 of 4 cultivars of field‐grown Glycine max exposed to near‐ambient solar UV‐B at a mid‐latitude site (Buenos Aires, Argentina, 34° S) showed significantly lower abaxial F(UV‐B)/F(BG) values (i.e., lower UV‐B epidermal transmittance) than those exposed to attenuated UV‐B, but solar UV‐B reduction had a minimal effect on F(UV‐B)/F(BG) in plants growing at a high‐latitude site (Tierra del Fuego, Argentina, 55° S). Similarly, the exotic Taraxacum officinale did not show significant changes in F(UV‐B)/F(BG) when exposed to very high supplemental UV‐B (biologically effective UV‐B=14–15 kJ m^?2 day^?1) in the field in Tierra del Fuego, whereas a native species, Gunnera magellanica, showed significant increases in F(UV‐B)/F(BG) relative to those receiving ambient UV‐B. These anomalous fluorescence changes were associated with increases in BG‐absorbing pigments (anthocyanins), but not UV‐B‐absorbing pigments. These results indicate that non‐invasive estimates of epidermal transmittance of UV radiation using chlorophyll fluorescence can detect changes in pigmentation and leaf optical properties induced by UV‐B radiation under both field and laboratory conditions. However, this technique may be of limited utility in cold environments where UV and low temperatures can stimulate the production of BG‐absorbing pigments that interfere with these indirect measurements of UV‐transmittance.     

Responses to UV-B radiation in Trifolium repens L. – physiological links to plant productivity and water availability

This study used comparisons across nine populations of Trifolium repens (white clover) in conjunction with drought to examine physiological responses to ultraviolet‐B radiation (UV‐B). Plants were exposed for 12 weeks to supplementation with 13.3 kJ m^?2 d^?1 UV‐B, accompanied by 4 weeks of drought under controlled environmental conditions. UV‐B increased the levels of UV‐B‐absorbing compounds and of flavonol glycosides and this effect was synergistically enhanced by water stress. These changes were more pronounced for the ortho‐dihydroxylated quercetin, rather than the monohydroxylated kaempferol glycosides. UV‐B increased leaf water potential (ψ_L) by 16% under drought and proline levels by 23% under well‐watered conditions. The intraspecific comparisons showed that higher UV‐B‐induced levels of UV‐B‐absorbing compounds, of quercetin glycosides and of ψ_L were linked to lower plant productivity and to higher UV‐B tolerance under well‐watered conditions. These findings suggest that: (1) slow‐growing T. repens ecotypes adapted to other stresses have higher capacity for physiological acclimation to UV‐B; and (2) that these attributes also contribute to decreased UV‐B sensitivity under drought.     

Response of feather moss associated N2 fixation and litter decomposition to variations in simulated rainfall intensity and frequency

Feather mosses in boreal forests form a dense ground‐cover that is an important driver of both nutrient and carbon cycling. While moss growth is highly sensitive to moisture availability, little is known about how moss effects on nutrient and carbon cycling are affected by the dynamics of moisture input to the ecosystem. We experimentally investigated how rainfall regimes affected ecosystem processes driven by the dominant boreal feather moss Pleurozium schreberi by manipulating total moisture amount, frequency of moisture addition and moss presence/absence. Moisture treatments represented the range of rainfall conditions that occur in Swedish boreal forests as well as shifts in rainfall expected through climate change. We found that nitrogen (N) fixation by cyanobacteria in feather mosses (the main biological N input to boreal forests) was strongly influenced by both moisture amount and frequency, and their interaction; increased frequency had greater effects when amounts were higher. Within a given moisture amount, N fixation varied up to seven‐fold depending on how that amount was distributed temporally. We also found that mosses promoted vascular litter decomposition rates, concentrations of litter nutrients, and active soil microbial biomass, and reduced N release into soil solution. These effects were usually strongest under low moisture amount and/or frequency, and revealed a buffering effect of mosses on the decomposer subsystem under moisture limitation. These results highlight that both the amount and temporal distribution of rainfall, determine the effect of feather mosses on ecosystem N input and the decomposer subsystem. They also emphasize the role of feather mosses in mediating moisture effects on decomposer processes. Finally, our results suggest that projected shifts in precipitation in the Swedish boreal forest through climate change will result in increased moss growth and N₂ fixation but a reduced dependency of the decomposer subsystem on feather moss cover for moisture retention.     

Changes in photosynthesis and antioxidant defenses of Picea asperata seedlings to enhanced ultraviolet-B and to nitrogen supply

The paper mainly studied the effects of ultraviolet‐B (UV‐B) radiation, nitrogen, and their combination on photosynthesis and antioxidant defenses of Picea asperata seedlings. The experimental design included two levels of UV‐B treatments (ambient UV‐B, 11.02 KJ m⁻² day⁻¹; enhanced UV‐B, 14.33 KJ m⁻² day⁻¹) and two nitrogen levels (0; 20 g m⁻² a⁻¹ N) – to determine whether the adverse effects of UV‐B are eased by supplemental nitrogen. Enhanced UV‐B significantly inhibited plant growth, net photosynthetic rate (A), stomatal conductance to water vapor (Gs), transpiration rate and photosynthetic pigment, and increased intercellular CO₂ concentration, UV‐B absorbing compounds, proline content, malondialdehyde (MDA) content, and activity of antioxidant enzymes (peroxidase (POD), superoxide dimutase, and glutathione reductase). Enhanced UV‐B also reduced needle DW and increased hydrogen peroxide (H₂O₂) content and the rate of superoxide radical (O₂⁻) production only under supplemental nitrogen. On the other hand, supplemental nitrogen increased plant growth, A, Gs, chlorophyll content and activity of antioxidant enzymes (POD, ascorbate peroxidase, and catalase), and reduced MDA content, H₂O₂ content, and the rate of O₂⁻ production only under ambient UV‐B, whereas supplemental nitrogen reduced activity of antioxidant enzymes under enhanced UV‐B. Carotenoids content, proline content, and UV‐B absorbing compounds increased under supplemental nitrogen. Moreover, significant UV‐B × nitrogen interaction was found on plant height, basal diameter, A, chlorophyll a, activity of antioxidant enzymes, H₂O₂, MDA, and proline content. These results implied that supplemental nitrogen was favorable for photosynthesis and antioxidant defenses of P.asperata seedlings under ambient UV‐B. However, supplemental nitrogen made the plants more sensitive to enhanced UV‐B, although some antioxidant indexes increased.     

Effects of ultraviolet-B radiation on leaf elongation, production and phenylpropanoid concentrations of Deschampsia antarctica and Colobanthus quitensis in Antarctica

Stratospheric ozone depletion by anthropogenic chlorofluorocarbons has lead to increases in ultraviolet‐B radiation (UV‐B; 280–320 nm) along the Antarctic Peninsula during the austral spring. We manipulated UV‐B levels around plants of Antarctic hair grass (Deschampsia antarctica; Poaceae) and Antarctic pearlwort (Colobanthus quitensis; Caryophyllaceae) for one field season near Palmer Station along the west coast of the Antarctic Peninsula. Treatments involved placing frames over naturally growing plants that either (1) held filters that absorbed most biologically effective radiation (UV‐B_BE; ‘reduced UV‐B’, 22% of ambient UV‐B_BE levels), (2) held filters that transmitted most UV‐B_BE (‘near‐ambient UV‐B’, 87% of ambient UV‐B_BE levels), or (3) lacked filters (‘ambient UV‐B’). Leaves on D. antarctica exposed to near‐ambient and ambient UV‐B were 16–17% shorter than those exposed to reduced UV‐B, and this was associated with shorter epidermal cells at the leaf base and tip. Leaves on C. quitensis exposed to near‐ambient and ambient UV‐B tended to be shorter (P=0.18) and epidermal cells at the leaf base tended to be smaller than those under reduced UV‐B (P<0.10). In order to further explain reductions in leaf length, we examined leaf concentrations of insoluble (cell‐wall bound) phenylpropanoids, since it has been proposed that wall‐bound phenylpropanoids such as ferulic acid may constrain cell expansion and leaf elongation. In both species, HPLC analysis revealed that ferulic and p‐coumaric acid were major components of both insoluble and soluble phenylpropanoids. Although there were no significant differences in concentrations between UV‐B treatments, concentrations of insoluble ferulic acid in D. antarctica tended to be higher under ambient and near‐ambient UV‐B than under reduced UV‐B (P=0.17). We also examined bulk‐leaf concentrations of soluble (methanol extractable) UV‐B‐absorbing compounds and found that concentrations were higher in plants exposed to near‐ambient and ambient UV‐B than in plants exposed to reduced UV‐B. We also assessed the UV‐B‐screening effectiveness of leaves that had developed on plants at the field site with a fiber‐optic microprobe. Leaf epidermal transmittance of 300‐nm UV‐B was 4.0 and 0.6% for D. antarctica and C. quitensis, respectively, which is low compared to grasses and herbaceous dicotyledonous plants found in more temperate climates. While the leaves of Antarctic vascular plants are relatively effective at screening UV‐B, levels of UV‐B in Antarctica are sufficient to reduce leaf epidermal cell size and leaf elongation in these species, although the mechanisms for these reductions remain unclear.     

The ability of abaxial and adaxial epidermis of sun and shade leaves to attenuate UV-A and UV-B radiation in relation to the UV absorbing capacity of the whole leaf methanolic extracts

The UV‐absorbing capacity (measured as A₃₁₀ cm^?2 and A₃₆₅ cm^?2 or A_UVR cm^?2) of the shade leaves of four representative evergreen sclerophylls of the Mediterranean region (Quercus coccifera, Q. ilex, Arbutus andrachne and A. unedo) was considerably lower than the corresponding one of sun leaves of the same species. However, fibre optic microprobe measurements showed that adaxial as well as abaxial epidermis of shade leaves of all examined plants, except abaxial epidermis of A. andrachne, were almost as effective as the corresponding ones of the sun leaves in screening out most of the incident UV‐B radiation. There is probably a threshold, under which the concentration of the UV‐B absorbing compounds in the protective tissues is not furthermore reduced, in spite of the low levels of the stress factor (UV‐B radiation) in the environment. On the other hand, the ability of both abaxial and adaxial epidermis to attenuate UV‐A radiation, except of adaxial leaf epidermis of Quercus species, depended on the UV absorbing capacity of the whole‐leaf extracts, with different correlation patterns between the two Quercus species and the two Arbutus species. This could be explained by the fact that shade leaves showed not only quantitative, but also qualitative differences (higher A₃₁₀/A₃₆₅ ratio) in the absorbance of their methanolic extracts compared to these of sun leaves. The results of the present study showed that we should not always correlate the depth of penetration of UV radiation into sun and shade leaves according to the corresponding UV absorbing capacity of the whole leaf methanolic extracts, without taking into account all the anatomical, developmental and biochemical (such as different composition and distribution of the UV‐absorbing compounds among the different protective tissues) peculiarities of the leaves of each species.     

Appropriate controls in outdoor UV-B supplementation experiments

Quercus robur L. saplings were exposed in an outdoor experiment to supplemental levels of UV-8 (280–315 nm) radiation using treatment arrays of cellulose diacetate-filtered fluorescent lamps that also produce UV-A (315–400 nm) radiation. Saplings were also exposed to UV-A radiation alone using control arrays of the same lamps filtered with polyester and to ambient levels of radiation, using arrays of unenergized lamps. The UV-B treatment was modulated to maintain a 30% elevation above the ambient level of UV-B radiation, measured by a broad-band sensor weighted with an erythemal action spectrum. Saplings exposed to UV-B radiation beneath treatment arrays developed thicker leaves than those beneath ambient and control arrays. Despite the fact that supplemental levels of UV-A radiation were only a small percentage of ambient levels, apparent UV-A effects were also recorded. Significant increases in sapling height, lammas shoot length and herbivory by chewing insects were observed under treatment and control arrays, relative to ambient, but there were no differences between the responses of saplings under treatment and control. These data imply that supplemental UV-A radiation or other effects associated with energised lamps can significantly affect plant growth parameters and herbivory in outdoor studies. We conclude that the results from current outdoor UV-B supplementation experiments that lack control exposures using polyester-filtered lamps need to be interpreted with caution and that future supplementation experiments should include appropriate controls.     

Metabolite specific effects of solar UV-A and UV-B on alder and birch leaf phenolics

We measured the concentrations of ultraviolet (UV)‐absorbing phenolics varying in response to exclusion of either solar UV‐B or both solar UV‐A and UV‐B radiations in leaves of grey alder (Alnus incana) and white birch (Betula pubescens) trees under field conditions. In alder leaves 20 and in birch leaves 13 different phenolic metabolites were identified. The response to UV exclusion varied between and within groups of phenolics in both tree species. The changes in concentration for some metabolites suggest effects of only UV‐A or UV‐B, which band being effective depending on the metabolite. For some other metabolites, the results indicate that UV‐A and UV‐B affect concentrations in the same direction, while for a few compounds there was evidence suggesting opposite effects of UV‐A and UV‐B radiation. Finally, the concentration of some phenolics did not significantly respond to solar UV. We observed only minor effects on the summed concentration of all determined phenolic metabolites in alder and birch leaves, thus indicating that measuring only total phenolics concentration may not reveal the effects of radiation. Here, we show that the appropriate biological spectral weighting functions for plant‐protective responses against solar UV radiation extend in most cases – but not always – into the UV‐A region and more importantly that accumulation of different phenolic metabolites follows different action spectra. This demonstrates under field conditions that some of the implicit assumptions of earlier research simulating ozone depletion and studying the effects of UV radiation on plant secondary metabolites need to be reassessed.     

Bryophyte use by an insect herbivore: does the crane‐fly Tipula montana select food to maximise growth?

1. Growth rates of immature invertebrates can be of profound importance to subsequent reproductive success in the adult stage. If diet selection in herbivores is a principal means of maximising growth and rate of development, individuals should select food of the highest quality. 2. If disparities are found between performance and preference, these may be the result of individuals balancing nutritional gain against other demands on their behaviour, such as a need for shelter. The responses of the crane‐fly herbivore Tipula montana Curtis (Diptera: Tipulidae) to a range of moss genera from an upland environment were investigated to determine whether diet selection was determined by food quality alone, as measured by its effect on larval growth and survival. 3. Larvae were reared individually on single moss genus diets, and their growth, development, and survival were measured. Each of the mosses supported growth and development although the mean weights achieved differed by a factor of two between food genera. In two‐choice preference experiments, individuals broadly chose bryophyte foods as expected from their quality. One notable exception was the moss Pleurozium schreberi, which gave the best growth performance but was among the least preferred. 4. The two methods of assessing preference gave different results: observations of larvae showed relatively greater selection for dense mosses than faecal pellet analysis. A strong dietary preference for an angiosperm, the sedge Carex bigelowii, over all mosses supported the interpretation that crane‐fly larvae benefit from bryophytes as a refuge, and that this factor may override dietary quality.     

Responses of plants in polar regions to UVB exposure: a meta-analysis

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, F_v/F_m or Φ_PSII, 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.