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     

Growth enhancement of soybean (Glycine max) upon exclusion of UV-B and UV-B/A components of solar radiation: characterization of photosynthetic parameters in leaves

Exclusion of UV (280–380 nm) radiation from the solar spectrum can be an important tool to assess the impact of ambient UV radiation on plant growth and performance of crop plants. The effect of exclusion of UV-B and UV-A from solar radiation on the growth and photosynthetic components in soybean (Glycine max) leaves were investigated. Exclusion of solar UV-B and UV-B/A radiation, enhanced the fresh weight, dry weight, leaf area as well as induced a dramatic increase in plant height, which reflected a net increase in biomass. Dry weight increase per unit leaf area was quite significant upon both UV-B and UV-B/A exclusion from the solar spectrum. However, no changes in chlorophyll a and b contents were observed by exclusion of solar UV radiation but the content of carotenoids was significantly (34–46%) lowered. Analysis of chlorophyll (Chl) fluorescence transient parameters of leaf segments suggested no change in the F _v/F _m value due to UV-B or UV-B/A exclusion. Only a small reduction in photo-oxidized signal I (P700⁺)/unit Chl was noted. Interestingly the total soluble protein content per unit leaf area increased by 18% in UV-B/A and 40% in UV-B excluded samples, suggesting a unique upregulation of biosynthesis and accumulation of biomass. Solar UV radiation thus seems to primarily affect the photomorphogenic regulatory system that leads to an enhanced growth of leaves and an enhanced rate of net photosynthesis in soybean, a crop plant of economic importance. The presence of ultra-violet components in sunlight seems to arrest carbon sequestration in plants. An erratum to this article can be found at     

Quantitative changes in secondary metabolites of dark-leaved willow (Salix myrsinifolia) exposed to enhanced ultraviolet-B radiation

This is a study of the impact of increased ultraviolet-B (UV-B) radiation on the secondary chemistry of Salix myrsinifolia (dark-leaved willow). For nearly two decades, the loss of stratospheric ozone above the high latitudes of the Northern Hemisphere has increased UV-B radiation (280–320 nm) over the long-term mean. Willows (Salicaceae) are widely distributed in these northern regions. To determine the effects of increased UV-B radiation on willows, the plantlets of three clones of S. myrsinifolia were grown under ambient (3.6 kJ m⁻² day⁻¹) or enhanced (7.18 kJ m⁻² day⁻¹) UV-B irradiance. After the 2-week indoor experiment, the concentrations of UV-B-screening phenolics (flavonoids and phenolic acids) and low-UV-B-screening phenolics (salicylates and condensed tannins) in fresh leaves were investigated and the biomass of leaves, stems and roots was determined. As expected, the total amount of flavonoids in willow leaves clearly increased when plantlets were exposed to higher UV-B irradiation. However, the degree of increase of individual compounds varied: luteolin-7-glucoside, monomethyl-monocoumaryl-luteolin-7-glucoside and one myricetin derivative increased significantly, while the apigenin-7-glucuronide increased only slightly. The enhanced UV-B also increased the amount of p -hydroxycinnamic acid derivative. The UV-B effects on other phenolic acids and tannins were minor. In contrast to the other phenolics, the amounts of two salicylates, salicin and saligenin, decreased under enhanced UV-B irradiation. Our results indicate that the concentrations of both UV-B-screening and low-UV-B-screening phenolic compounds in leaves of S. myrsinifolia may vary in response to elevated UV-B radiation. However, while the UV-B protective flavonoids and phenolic acids accumulate during UV-B exposure, the concentrations of certain salicylates decrease.     

Enhanced UV-B radiation,artificial wounding and leaf chemical defensive potential in Phlomis fruticosa L.

The combined effects of additional UV-B radiation and artificial wounding on leaf phenolics were studied in a short term field experiment with the drought semi-deciduous Mediterranean shrub Phlomis fruticosa L. The seedlings were grown under ambient or ambient plus supplemental UV-B radiation (biologically equivalent to a 15% ozone depletion over Patras, 38.3° N, 29.1° E) for 7 months before wounding. Unexpectedly, supplemental UV-B radiation decreased leaf phenolics. Subsequently, wounding was effected by removing leaf discs from some of the plants, while the rest remained intact and served as controls. Wounding significantly increased phenolics of the wounded leaves and the increase was more pronounced under supplemental UV-B radiation. In addition, wounding had a significant positive effect on the phenolics of the opposite, intact leaf, but only under additional UV-B radiation. We conclude that UV-B radiation, wounding and their combination may affect the chemical defensive potential of Phlomis fruticosa. In addition, increased levels of phenolics after herbivore attack under field conditions may afford extra protection against enhanced UV-B radiation levels.     

The effect of solar UV-B radiation on terpenes and biomass production in Grindelia chiloensis (Asteraceae), a woody perennial of Patagonia,Argentina

UV-B radiation is absorbed effectively by nucleic acids and other sensitive targets, potentially causing harmful photochemical effects. Protection against UV-B radiation may be afforded by flavonoids and other phenolics, which absorb strongly in the UV region, but little is known about the role played by other compounds, such as terpenes. Grindelia chiloensis, native of Patagonia (Argentina), can accumulate as much as 25% resin (terpenes) in its leaves. The present investigation was carried out to test the effect of solar UV-B radiation on the allocation of photoassimilates to biomass and terpenes. Exposure to UV-B radiation reduced whole plant biomass, plant height and leaf area, and increased leaf thickness and resin accumulation in Grindelia chiloensis. Higher absorbance was found for refined resin in the UV-B waveband from plants grown under solar UV-B radiation than plants without UV-B radiation. These chemical and structural changes could protect the plant from UV radiation, and may help elucidate the importance of epicuticular resins for a species as G. chiloensis native to an environment with maximum daily integrated values of solar UV-B irradiance.     

Effects of UV-B radiation and additional irrigation on the Mediterranean evergreen sclerophyll Ceratonia siliqua L. under field conditions

Seedlings of Ceratonia siliqua L. were grown for 1 year in the field under ambient or ambient plus supplemental UV-B radiation (corresponding to 15% ozone depletion over Patras) and received two levels of additional irrigation during the summer dry period. The experiment was started during February 1998 and two major samplings were performed, the first at the end of the dry period (September 1998) and the second at the end of the experiment (January 1999). Plants receiving additional irrigation showed significantly higher leaf number, plant height and chlorophyll content at the end of the summer, but these differences were abolished at the final harvest. Plants growing under enhanced UV-B radiation had significantly fewer leaves and less nitrogen content at the end of the dry period, but these effects were also abolished at the final harvest, during which significant UV-B induced increases in stem dry mass were observed. None of the other measured parameters (mean leaf area, leaf dry mass, leaf thickness, UV-B absorbing compounds, phenolics, tannins and photochemical efficiency of PSII) were affected by either treatment. Combined UV-B / water effects were not significant. We may conclude that although some minor responses to enhanced UV-B radiation were evident, C. siliqua is resistant against UV-B radiation damage at the level applied.     

Stratospheric ozone reduction and ecosystem processes: enhanced UV-B radiation affects chemical quality and decomposition of leaves of the dune grassland species Calamagrostis epigeios

This study reports changes in the plant's chemical composition and the decomposition of this plant material under enhanced solar UV-B radiation. Calamagrostis epigeios, a dominant grass species in the dune grassland in The Netherlands, was grown outdoor on an experimental field under ambient and enhanced solar UV-B (5 and 7.5 kJ m^-2 day^-1 UV-B_BE, respectively), corresponding to about 15% stratospheric ozone depletion. After one growing season aerial plant parts were harvested. The decomposition of this harvested leaf material was studied in a dune grassland and on the above mentioned experimental field under ambient (5 kJ m^-2 day^-1 UV-B_BE) and enhanced (7.5 kJ m^-2 day^-1 UV-B_BE) radiation, using litter bags. The chemical quality of the leaves grown under enhanced solar UV-B changed. There was an increase in the leaf content of lignin, while no significant changes occurred for the content of -cellulose, hemicellulose and tannins under enhanced UV-B. In the field, the rate of decomposition of leaf material grown under enhanced UV-B (with an increased content of lignin) was reduced. The content of lignin of the decomposing leaf material increased, but less under exposure to enhanced UV-B. The latter may be explained by photodegradation of the lignin. The consequences of enhanced UV-B radiation for carbon fluxes in the dune grassland ecosystem are discussed.     

Contrasting strategies for UV-B screening in sub-Arctic dwarf shrubs

The content and distribution of UV-absorbing phenolic compounds was investigated in leaves of three species of Vaccinium co-existing at a site in north Sweden. Vaccinium myrtillus L., Vaccinium vitis-idaea L., and Vaccinium uliginosum L. exhibit markedly different strategies, in terms of localization and content of leaf phenolics and in their responses to UV-B enhancement. Plants were exposed to either ambient radiation or to enhancement of UV-B corresponding to 15% (clear sky) depletion of stratospheric ozone for approximately 10 years prior to commencement of this study. Vaccinium myrtillus contained the highest concentration of methanol-extractable UV-B-absorbing compounds, which was elevated in plants exposed to enhanced UV-B. Fluorescence and confocal laser scanning microscopy showed that these compounds were distributed throughout the leaf, and were particularly concentrated in chlorophyll-containing cells. In V. vitis-idaea, most phenolic compounds were cell wall-bound and concentrated in the walls of the epidermis; this pool increased in response to UV-B enhancement. It is suggested that these two plants represent extreme forms of two divergent strategies for UV-B screening, the different responses possibly being related to leaf longevity in the two species. The response of V. uliginosum was intermediate between the other two, with high concentrations of cell wall-bound phenolics in the epidermis but with this pool decreasing, and the methanol-soluble pool tending to increase, after exposure to enhanced UV-B. One explanation for this response is that this plant is deciduous, like V. myrtillus, but has leaves that are structurally similar to those of V. vitis-idaea.     

Phenolic compounds in seedlings of Betula pubescens and B. pendula are affected by enhanced UVB radiation and different nitrogen regimens during early ontogeny

We studied the ability of tree seedlings to respond to two environmental factors, elevated ultraviolet B (UVB) radiation and availability of nitrogen (N), at the beginning of their development. Seeds of two birch species, Betula pubescens Ehrh. (common white birch) and B. pendula Roth (silver birch), were germinated and the seedlings grown in an experimental field in eastern Finland. The experimental design consisted of a constant 50% increase in UVB radiation (including a slight increase in UVA), a UVA control (a slight increase in UVA) and a control. The seedlings were fertilized with three levels of N. The experiment lasted for 2 months; aboveground biomass was measured and the most mature leaf of each seedling was taken for the analyses of phenolics. Growth of the seedlings was not significantly affected by enhanced UVB, but was increased by increasing N. Elevated UVB induced significant changes in phenolic compounds. Quercetin glycosides were accumulated in the leaves of both species in response to UVB; this is considered to be a protective response. However, the direction of the responses of individual phenolics to different N regimens differed. In addition, concentration of soluble condensed tannins was lower at moderate N than that at lower levels of N in both species; on the contrary, in B. pubescens the concentration of insoluble condensed tannins was highest at moderate N. No significant interaction between UV and N was detected, and the responses of the two species were highly similar to UVB, while the responses to N regimens varied slightly more between species.     

Temperature is the key to altitudinal variation of phenolics in Arnica montana L. cv. ARBO

Plants in alpine habitats are exposed to many environmental stresses, in particular temperature and radiation extremes. Recent field experiments on Arnica montana L. cv. ARBO indicated pronounced altitudinal variation in plant phenolics. Ortho-diphenolics increased with altitude compared to other phenolic compounds, resulting in an increase in antioxidative capacity of the tissues involved. Factors causing these variations were investigated by climate chamber (CC) experiments focusing on temperature and ultraviolet (UV)-B radiation. Plants of A. montana L. cv. ARBO were grown in CCs under realistic climatic and radiation regimes. Key factors temperature and UV-B radiation were altered between different groups of plants. Subsequently, flowering heads were analyzed by HPLC for their contents of flavonoids and caffeic acid derivatives. Surprisingly, increased UV-B radiation did not trigger any change in phenolic metabolites in Arnica. In contrast, a pronounced increase in the ratio of B-ring ortho-diphenolic (quercetin) compared to B-ring monophenolic (kaempferol) flavonols resulted from a decrease in temperature by 5°C in the applied climate regime. In conclusion, enhanced UV-B radiation is probably not the key factor triggering shifts in the phenolic composition in Arnica grown at higher altitudes but rather temperature, which decreases with altitude.     

Seasonal fluctuations in leaf phenolic composition under UV manipulations reflect contrasting strategies of alder and birch trees

Seasonal variation in leaf phenolic composition may be important for acclimation of plants to seasonal changes in their biotic and abiotic environment. For a realistic assessment of how plants respond to solar UV‐B (280–315 nm) and UV‐A (315–400 nm) radiation, seasonal variation in both environment and plant responses needs to be taken into account. This also has implications for studies concerning stratospheric ozone depletion and resulting increased UV‐B radiation, as other environmental variables and/or plant phenology could interact with UV radiation. To elucidate this, we established a field experiment using plastic films attenuating different parts of the solar UV spectrum. The concentration of individual phenolic compounds was measured during one growing season in leaves of grey alder (Alnus incana) and white birch (Betula pubescens) trees. Our results showed changes in concentration of, e.g. hydrolyzable tannins in birch that suggest an effect of UV‐A alone and e.g. chlorogenic acids in alder indicate a quadratic effect of UV‐B irradiance and both linear and quadratic effect for UV‐A in second‐degree polynomial fits. Further, there was interaction between treatment and sampling time for some individual metabolites; hence, the UV response varied during the season. In addition to the UV effects, three temporal patterns emerged in the concentrations of particular groups of phenolics. Possible implications for both sampling methods and timing are discussed. Moreover, our results highlight differences in responses of the two tree species, which are taken to indicate differences in their ecological niche differentiation.     

Short-term impacts of enhanced UV-B radiation on photo-assimilate allocation and metabolism: a possible interpretation for time-dependent inhibition of growth

To test the hypothesis that plant source-sink relations are important in determining response to UV-B radiation, a short-term (45 d) field experiment was conducted at Abisko Scientific Research Station, Abisko, Sweden (68° N). Tillers of the grass Calamagrostis purpurea were grown outdoors at levels of UV-B radiation representing 25% ozone depletion. Growth, respiration, photo-assimilate allocation and UV-B protective compounds were subsequently measured.There were no significant effects of enhanced UV-B on total plant dry weight, leaf area, Shoot: Root ratio, leaf weight ratio, leaf area ratio, specific leaf area, tiller number per plant or blade thickness of this species. However, the amount of UV-B absorbing compounds and respiration rates were significantly increased in young and mature leaves. Increases in leaf respiration were accompanied by alterations in plant carbohydrate allocation at enhanced UV-B. The amount of soluble root carbohydrates was reduced following UV-B exposure. Enhanced UV-B also caused increases in the soluble sugar: starch ratio of young leaves, the stem and total aboveground biomass. The importance of source-sink relations and constitutive versus induced defense are discussed in relation to UV-B response.     

Functional significance and induction by solar radiation of ultraviolet-absorbing sunscreens in field-grown soybean crops

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

Partitioning of 14C-labeled photosynthate to allelochemicals and primary metabolites in source and sink leaves of aspen: evidence for secondary metabolite turnover

Theories on allelochemical concentrations in plants are often based upon the relative carbon costs and benefits of multiple metabolic fractions. Tests of these theories often rely on measuring metabolite concentrations, but frequently overlook priorities in carbon partitioning. We conducted a pulse-labeling experiment to follow the partitioning of ¹⁴CO₂-labeled photosynthate into ten metabolic pools representing growth and maintenance (amino acids, organic acids, lipids plus pigments, protein, residue), defense (phenolic glycosides, methanol:water and acetone-soluble tannins/phenolics), and transport and storage (sugars and starch) in source and importing sink leaves of quaking aspen (Populus tremuloides). The peak period of ¹⁴C incorporation into sink leaves occurred at 24 h. Within 48 h of labeling, the specific radioactivity (dpm/mg dry leaf weight) of phenolic glycosides declined by over one-third in source and sink leaves. In addition, the specific radioactivity in the tannin/phenolic fraction decreased by 53% and 28% in source and sink leaves, respectively. On a percent recovery basis, sink leaves partitioned 1.7 times as much labeled photosynthate into phenolic glycosides as source leaves at peak ¹⁴C incorporation. In contrast, source leaves partitioned 1.8 times as much ¹⁴C-labeled photosynthate into tannins/phenolics as importing sink leaves. At the end of the 7-day chase period, sink leaves retained 18%, 52%, and 30% of imported ¹⁴C photosynthate, and labeled source leaves retained 15%, 66%, and 19% of in situ photosynthate in metabolic fractions representing transport and storage, growth and maintenance, and defense, respectively. Analyses of the phenolic fractions showed that total phenolics were twice as great and condensed tannins were 1.7 times greater in sink than in source leaves. The concentration of total phenolics and condensed tannins did not change in source and sink leaves during the 7-day chase period. Received: 31 July 1998 / Accepted: 8 February 1999     

Photoperiodic control of phenolic metabolism in Kalanchoe blossfeldiana

When grown in conditions of long day length, the leaves of Kalanchoe blossfeldiana contain high levels of soluble phenolic compounds, mainly present as tannins. A decreasing concentration gradient is observed in the leaves from the apex to the base. When transferred to short day conditions, the ability of leaves of the same physiological age to accumulate phenolics decreases with time. The very low phenolic content after 25 short days indicates pronounced changes in the metabolism of the plant induced by new photoperiodic conditions. Moreover, during development in short days the amount of tannins per leaf reaches a maximum then decreases suggesting an over-polymerization or even a degradation of the substances. A similar lag time is required for the depressing effect of short days on phenolies and for their stimulating effect on CAM.     

Combined effects of enhanced UV-B radiation and additional nutrients on growth of two Mediterranean plant species

Seedlings of two Mediterranean plants, the slow-growing, evergreen sclerophyll Ceratonia siliqua L. and the fast growing drought semi-deciduous Phlomis fruticosa L., were grown for one year in the field at ambient or ambient plus supplemental UV-B radiation (equivalent to a 15% ozone depletion) and two levels of applied fertilizers (NPK). The effects on growth, morphological, anatomical and physiological parameters were measured at final plant harvest. Additional nutrients increased leaf nitrogen, improved growth and reduced the root/shoot ratio in both plants, yet these effects were more pronounced in the fast growing P. fruticosa. A nutrient-induced increase in chlorophyll content was also observed in this plant. The growth responses to UV-B radiation were different for the two species. Growth in C. siliqua was not affected by UV-B radiation at both nutrient levels and the same was true for P. fruticosa at low nutrients. However, at the high nutrient level, supplemental UV-B radiation improved growth in P. fruticosa, indicating a strong interaction between the treatments. Photosystem II (PSII) photochemical efficiency, methanol-extractable UV-B absorbing capacity, total phenolics and tannins were not affected by either treatment in both plants. It is concluded that nutrient levels can strongly modify the UV-B radiation effects on growth of P. fruticosa. We presume that this may be correlated to the fast growing habit of this species.     

Short communication. Do anthocyanins play a role in UV protection of the red juvenile leaves of Syzygium?

The biological function of juvenile leaves pigmented with anthocyanin is poorly understood. The role anthocyanins play in UV protection was assessed in juvenile leaves of two Syzygium species (S. luehmannii and S. wilsonii) which contain high anthocyanin concentrations. HPLC was used to separate UV-absorbing anthocyanins from other soluble UV-absorbing phenolic compounds. The isolated anthocyanins (predominantly malvidin-3,5-diglucoside) contributed little to the total absorbance of UV-A and UV-8 radiation. This was because the non-acylated anthocyanins only effectively absorbed shortwave UV-B radiation and the strong absorbance by other compounds. These results suggest that the UV protection hypothesis is not valid for anthocyanins in juvenile Syzygium leaves.     

Effects of enhanced ultraviolet-B radiation on plant nutrients and decomposition of spring wheat under field conditions

Spring wheat (Triticum aestivum) was grown in the field under ambient and supplemental levels of ultraviolet-B (UV-B, 280–315 nm) radiation to determine the potential for alteration in plant nutrients, decomposition, leaf quality and dry matter yield. Supplemental UV-B radiation simulating a 12, 20 and 25% stratospheric ozone depletion significantly decreased dry matter yield, but had no significant impact on harvest index. UV-B radiation resulted in an increase of the concentrations of N and K in all plant parts; changes of the concentrations of P, Mg, Fe and Zn varied in a tissue-dependent manner, as the decrease of P in leaves and stems, and its increase in spikes and grains. The mass of N, P, K, Mg, Fe and Zn in various plant parts and whole plant was generally decreased except leaf N mass was increased by enhanced UV-B radiation. Enhanced UV-B radiation decreased the concentrations of soluble carbohydrates in leaves and increased that of holocellulose and soluble proteins. After 60 and 100 days of decomposition of leaves and stems in the field, enhanced UV-B radiation stimulated the loss of organic C. As a consequence, the nutrient content of soils might be less diminished under enhanced UV-B radiation.     

Tannin Dynamics of Propagules and Leaves of Kandelia candel and Bruguiera gymnorrhiza in the Jiulong River Estuary, Fujian, China

Changes in the total phenolics, condensed tannins (CT), protein-precipitable phenolics content and protein precipitation capacity were determined on a series of mangrove leaves from two true viviparous mangrove species (Kandelia candel and Bruguiera gymnorrhiza) at various stages of development and decomposition in the Jiulong River Estuary, Fujian, China. Similar measurements were also done for the propagules at different developmental stages. The results showed that the total phenolics, extractable condensed tannins, total condensed tannins, protein-precipitable phenolics content and protein precipitation capacity in young leaves were higher than those in mature and senescent leaves. Tannin dynamics during leaf decomposition varied with species, and the rapid loss of phenolics observed during decomposition can be ascribed to leaching and degradation. Protein-bound CT and fibre-bound CT tended to increase with leaf decomposition, with CT binding more strongly to protein than to fibre. Protein-bound CT was higher than fibre-bound CT with the exception of mature leaves. Total phenolics, extractable CT and protein-precipitable phenolics contents in flower tissues were relatively lower than those in hypocotyls at different developmental stages. Protein precipitation capacity fluctuated with the development of propagules. Increases in nitrogen in decaying litter, and declines in contents of total phenolics and total condensed tannins of detritus support the general conclusion that decomposing mangrove detritus can be a more palatable heterotrophic substrate than living leaves.