Ultraviolet-B radiation alters phenolic salicylate and flavonoid composition of Populus trichocarpa leaves

We investigated foliar phenolic composition of field- and greenhouse-grown Populus trichocarpa Torr. & A. Gray (black cottonwood) ramets subjected to near zero (0x), ambient (1x) or twice ambient (2x) concentrations of biologically effective ultraviolet-B (UV-B) radiation. After a 3-month treatment period, several age classes of foliage samples were harvested and the phenolic compounds extracted, separated by high performance liquid chromatography and identified and quantified by diode-array spectrometry and mass spectrometry. Foliar phenolic concentration was greater in 1x- and 2x-treated tissue than in 0x-treated tissue. Phenolic compounds that increased in response to UV-B radiation were predominantly flavonoids, primarily quercetin and kaempferol glycosides. Enhancement of UV-B radiation from 1x to 2x ambient concentration did not result in further flavonoid accumulation in either greenhouse or field ramets; however, a non-flavonoid phenolic glycoside, salicortin, increased in response to an increase in UV-B radiation from 1x to 2x ambient concentration. Increased salicortin concentrations accounted for at least 30-40% of the total (5%) increase in UV-absorption potential of 2x-treated tissue. Because salicortin and other salicylates are important in plant-herbivore-predator relationships, these increases are discussed in the context of collateral feeding studies. We conclude that enhanced solar UV-B radiation may significantly alter trophic structure in some ecosystems by stimulating specific phenolic compounds.     

Growth and gas exchange in field-grown and greenhouse-grown Quercus rubra following three years of exposure to enhanced UV-B radiation

Long-term effects of enhanced UV-B radiation were evaluated in field-grown and greenhouse-grown Quercus rubra L. (northern red oak), a species with a multiple flushing shoot growth habit. Seeds were germinated and grown in ambient, twice ambient (2x) or three times ambient (3x) biologically effective UV-B radiation from square-wave (greenhouse) or modulated (field) lamp systems for three growing seasons. Greenhouse plants in the 2x treatment had greater heights and diameters during the later part of the first year and into the second year, but by the third year there were no differences among treatments. There were no significant differences in growth among treatments for field plants. Enhanced UV-B radiation did not significantly reduce total biomass or distribution of biomass in either field or greenhouse plants. Net photosynthesis (3x), leaf conductance (2x and 3x) and water-use efficiency (3x) of greenhouse plants were greater in the enhanced UV-B radiation treatments in the second year but unaffected by the treatments in other years. None of the treatments affected these parameters in field plants. Dark respiration was increased by the 3x treatment in the first and third years in greenhouse plants, and by the 2x treatment during the second year in field plants. Enhanced UV-B had variable effects on apparent quantum yield and light compensation points. Chlorophylls were unaffected by enhanced UV-B radiation in both greenhouse and field conditions. Bulk methanol-extractable UV-absorbing compounds were increased only by the 3x treatment in greenhouse plants during the third year and by the 2x treatment in field plants during the second year. Overall, Q. rubra appears relatively resistant to potentially damaging enhanced UV-B radiation and is unlikely to be negatively impacted even in the predicted worst-case scenarios.     

Growth, leaf anatomy, and physiology of Populus clones in response to solar ultraviolet-B radiation

We compared the physiological and morphological responses of rooted cuttings of Populus trichocarpa Torr. & Gray and P. trichocarpa x P. deltoides Bartr. ex Marsh. grown in either near-ambient solar ultraviolet-B (UV-B; 280-320 nm) radiation (cellulose diacetate film) or subambient UV-B radiation (polyester film) for one growing season. Midday biologically effective UV-B radiation was 120.6 and 1.6 mJ m(-2) s(-1) under the cellulose diacetate and polyester films, respectively. Gas exchange, leaf chlorophyll, light harvesting efficiency of photosystem II, and foliar UV-B radiation-absorbing compounds (i.e., flavonoid derivatives) were measured in expanding (leaf plastochron index (LPI) 5), nearly expanded (LPI 10), and fully expanded mature (LPI 15) leaves of intact plants of plastochron index 30 to 35. Plants were then harvested and height, diameter, biomass allocation and leaf anatomical attributes determined. Net photosynthesis, transpiration, and stomatal conductance were significantly greater in mature leaves exposed to subambient UV-B radiation than in mature leaves exposed to near-ambient UV-B radiation. Concentrations of UV-B radiation-absorbing compounds (measured as absorbance of methanol-extracts at 300 nm) were significantly greater in mature leaves exposed to near-ambient UV-B radiation than in mature leaves exposed to subambient UV-B radiation. The UV-B radiation treatments had no effects on chlorophyll content or intrinsic light harvesting efficiency of photosystem II. Height, diameter, and biomass were not significantly affected by UV-B radiation regime in either clone. Leaf anatomical development was unaffected by UV-B radiation treatment in P. trichocarpa x P. deltoides. For P. trichocarpa, leaf anatomical development was complete by LPI 10 in the near-ambient UV-B radiation treatment, but continued through to LPI 15 in the subambient UV-B radiation treatment. Mature leaves of P. trichocarpa were thicker in the subambient UV-B radiation treatment than in the near-ambient UV-B radiation treament as a result of greater development of palisade parenchyma tissue. We conclude that exposure to near-ambient UV-B radiation for one growing season caused shifts in carbon allocation from leaf development to other pools, probably including but not limited to, UV-B absorbing compounds. This reallocation curtailed leaf development and reduced photosynthetic capacity of the plants compared with those in the subambient UV-B radiation treatment and may affect growth over longer periods of exposure.     

Accumulation of flavonoids and related compounds in birch induced by UV-B irradiance

A growth chamber experiment was conducted to examine the effects of UV-B exposure (4.9 kJ m(-2) day(-1) of biologically effective UV-B, 280-320 nm) on shoot growth and secondary metabolite production in Betula pendula (Roth) and B. resinifera (Britt.) seedlings originating from environments in Finland, Germany and Alaska differing in solar UV-B radiation and climate. Neither shoot growth nor the composition of secondary metabolites was affected by UV-B irradiance, but the treatment induced significant changes in the amounts of individual secondary metabolites in leaves. Leaves of seedlings exposed to UV-B radiation contained higher concentrations of several flavonoids, condensed tannins and some hydroxycinnamic acids than leaves of control seedlings that received no UV-B radiation. At the population level, there was considerable variation in secondary metabolite responses to UV-B radiation: among populations, the induced response was most prominent in Alaskan populations, which were adapted to the lowest ambient UV-B radiation environment. I conclude that solar UV-B radiation plays an important role in the formation of secondary chemical characteristics in birch trees.     

Enhanced tolerance of photosynthesis to high-light and drought stress in Pseudotsuga menziesii seedlings grown in ultraviolet-B radiation

We investigated the effects of an ambient dose of ultraviolet-B (UV-B) radiation on chamber-grown Pseudotsuga menziesii var. glauca (Beissn.) Franco (Douglas-fir) seedlings, to determine if the presence of UV-B radiation in the growth light regime induces tolerance to environmental stresses such as high light and drought. Douglas-fir seedlings were grown without UV-B radiation or with 6 kJ m-2 day-1 of biologically effective UV-B, which is ambient for the intermountain regions of Idaho. Non-stressed seedlings grown with UV-B radiation had 35% lower seedling dry mass, 36% higher concentrations of UV-B absorbing compounds per unit leaf area, 30% lower stomatal frequencies, 25% lower light-saturated photochemical efficiencies of Photosystem II and 45% lower light-saturated stomatal conductance than non-stressed seedlings grown without UV-B radiation. After 4 days of high-light stress, seedlings grown with UV-B radiation had 32% higher light-saturated carbon assimilation rates (A(CO2)) than seedlings grown without UV-B radiation. After water was withheld from the seedlings for up to 15 days, seedlings grown with UV-B radiation had 50% higher A(CO2) and 40% higher seedling water potentials than seedlings grown without UV-B radiation. The results support the hypothesis that UV-B radiation can act as an environmental signal to induce tolerance to high-light and drought stress in Douglas-fir seedlings. Possible mechanisms for the enhanced stress tolerance are discussed.     

Growth and physiological responses to supplemental UV-B radiation of two contrasting poplar species

We compared the growth and physiological responses of cuttings of Populus kangdingensis C. Wang et Tung and P. cathayana Rehder originating from altitudes of 3500 m and 1500 m, respectively, when exposed to three ultraviolet-B (UV-B) radiation regimes: zero UV-B, ambient (4.5 kJ m(-2) day(-1)) and twice-ambient (9 kJ m(-2) day(-1)) biologically effective UV-B radiation for one growing season, to determine if Populus trees that are adapted to contrasting UV-B habitats exhibit different tolerances to enhanced UV-B radiation. Compared with cuttings grown without UV-B radiation, twice-ambient UV-B radiation significantly decreased total biomass, total leaf area and internode length in both species, whereas root/shoot ratio, leaf number, amount of photosynthetic pigments and ascorbate peroxidase activity were unaffected. Differences in responses were also observed between the study species. Cutting height increment, total biomass, total leaf area, free proline concentration and membrane damage assessed by electrolyte leakage were significantly more affected by the twice-ambient UV-B radiation in P. cathayana than in P. kangdingensis. However, specific leaf mass, amount of UV-B absorbing compounds and superoxide dismutase and guaiacol peroxidase activities increased more with increasing UV-B radiation in P. kangdingensis than in P. cathayana, perhaps reflecting important characteristics of species with low to moderate tolerance to UV-B radiation. Overall, the results indicated that P. kangdingensis, which originates from altitudes of 3500 m, has greater tolerance to enhanced UV-B radiation than P. cathayana originating from altitudes of 1500 m.     

Responses to ultraviolet-B radiation by purely symbiotic and NO3-fed nodulated tree and shrub legumes indigenous to southern Africa

Purely symbiotic and NO3-fed nodulated seedlings of Virgilia oroboides (Bergius) T.M. Salter, Cyclopia maculata (L.) Vent and Podalyria calyptrata Willd. were exposed to biologically effective ultraviolet-B radiation (UV-B) to assess the effects of above- and below-ambient UV-B on growth, symbiotic function and metabolite concentrations. Seedlings were grown outdoors either on tables under ambient or 34 or 66% above-ambient UV-B conditions (UV-B100 control, UV-B134 and UV-B166, respectively), or in chambers providing below-ambient (22% of ambient) UV-B (UV-B22) along with a UV-A control and a photosynthetically active radiation (PAR) control. Exposure of seedlings to UV-B166 radiation reduced (P < or = 0.05) leaf and stem dry mass by 34 and 39%, respectively, in C. maculata, and reduced leaf nitrogen concentration (%N) by 12% in V. oroboides. Nodule %N in C. maculata and stem %N in P. calyptrata also decreased (P < or = 0.05) in response to UV-B22 radiation compared with the UV-A control, but not compared with the PAR control. Concentrations of flavonoids, soluble sugars and starch were unaltered by the UV-B treatments. Application of 1 mM NO3 to UV-B166-treated seedlings increased whole-plant dry mass of V. oroboides and P. calyptrata by 47 and 52%, respectively. Dry mass of organs, nodule %N and total N concentration of these species also increased with NO3 application. However, NO3 supply decreased (P < or = 0.05) nodule dry mass, stem %N and leaf %N as well as root and leaf anthocyanin concentrations in C. maculata. In terms of UV-B x N interactions, dry mass of stems, roots, nodules and total biomass of NO3-fed C. maculata seedlings were reduced, and nodule %N, total N and leaf anthocyanins were depressed by the UV-B134 and UV-B166 treatments relative to UV-B100-treated seedlings. Although we found that above-ambient UV-B had no effects on growth and symbiotic function of V. oroboides and P. calyptrata seedlings, feeding NO3 to these species increased (P < or = 0.05) seedling growth. In contrast, purely symbiotic C. maculata seedlings were sensitive to the UV-B166 radiation treatment, and adding NO3 further increased their sensitivity to both the UV-B134 and UV-B166 treatments.     

Different growth sensitivity to enhanced UV-B radiation between male and female Populus cathayana

We investigated sex-related morphological and physiological responses to enhanced UV-B radiation in the dioecious species Populus cathayana Rehd. Cuttings were subjected to two UV-B radiation regimes: ambient (4.5 kJ m?2 day?1) and enhanced (12.5 kJ m?2 day?1) biologically effective UV-B radiation for one growing season. Enhanced UV-B radiation was found to significantly decrease the shoot height and basal diameter and to reduce the leaf area, dry matter accumulation, net photosynthesis rate (P(n)), chlorophyll a/b ratio (Chl a/b) and anthocyanin content. Enhanced UV-B radiation also increased chlorophyll pigment, leaf nitrogen, malondialdehyde and abscisic acid (ABA) content, superoxide dismutase and peroxidase activities and UV-B-absorbing compounds. No significant effects of enhanced UV-B radiation were found on biomass allocation, gas exchange (except for P(n)), photochemical efficiency of photosystem II or water use efficiency. Moreover, different sensitivity to enhanced UV-B radiation between males and females was detected. Under enhanced UV-B radiation, males exhibited significantly higher basal diameter and leaf nitrogen, and lower Chl a/b, ABA content, UV-B-absorbing compounds, as well as less decrement of leaf area and dry matter accumulation than did females. However, no significant sexual differences in these traits were found under ambient UV-B radiation. Our results suggest that males may possess a greater UV-B resistance than do females, with males having a more efficient antioxidant system and higher anthocyanin content to alleviate UV-B penetration stress than females.     

Evaluating effects of nursery and post-planting nutrient regimes on leaf chemistry and browsing of eucalypt seedlings in plantations

We examined the effects of nursery and field nutrient treatments on leaf chemistry and browsing during establishment of Eucalyptus globulus (two trials; Cressy and Dunalley) and Eucalyptus nitens (one trial; Tyenna) seedlings. Seedlings had high- or low-fertilizer application treatments in the nursery, were fertilized or not in the field and were individually fenced (used for leaf chemistry sampling) or not (used for browsing assessment). The high-fertilizer nursery treatment resulted in relatively high foliar nitrogen (N) and low tannin content (all seedling types), and high total essential oils (seedling batch planted at Cressy only), but nursery fertilizer level did not affect dry matter, neutral detergent (NDF) or acid detergent fibre (ADF) or lignin. After planting, levels of N, tannins and total essential oils changed over time, but the degree differed between sites. Field fertilizing generally had no effect on leaf chemistry during 10 weeks after planting. Across sites, the pattern of browsing was the same. Seedlings with the high-fertilizer nursery treatment were browsed more than the low-fertilizer nursery treatment, though the difference was only significant at two of the three sites. Thus trends in browsing damage were consistent with differences in foliar N and tannins as a result of nursery- and not field fertilizing treatment. Manipulation of seedling chemistry, using particular fertilizer application regimes in the nursery, may be one tool for managers wishing to decrease seedling damage by browsers, particularly when browsing severity is not extreme.     

Autumnal changes of sulfur fractions and the ratio of organic sulfur to total nitrogen in leaves and adjacent bark of eastern cottonwood, white basswood and actinorhizal black alder

Autumnal changes in organic-S, sulfate-S, total-S and the ratios of organic-S to total-N and sulfate-S to organic-S were followed in leaves and adjacent bark of actinorhizal (Frankia-nodulated) black alder (Alnus glutinosa (L.) Gaertn.) and eastern cottonwood (Populus deltoides Bartr. ex Marsh.) trees growing on a minespoil site high in extractable soil sulfate, and in black alder and white basswood (Tilia heterophylla Venten.) trees growing on a prairie-derived soil in Illinois. Organic-S concentrations decreased significantly (P < 0.05) during autumn only in foliage of trees growing on the prairie-derived soil where losses of leaf organic-S were 65% for black alder and 100% for white basswood. Leaf sulfate concentrations were relatively stable throughout autumn in white basswood growing on prairie-derived soil and in black alder at both sites. Sulfate-S concentrations in leaves were significantly (P < 0.05) higher in trees at the minespoil site than in trees growing in the prairie-derived soil (5.1 mg g(-1) for the minespoil site and 1.2 mg g(-1) for the prairie-derived soil), and in the non-actinorhizal species during late summer. During the autumn, the ratio of organic-S to total-N doubled in leaves of eastern cottonwood at the minespoil site, but in black alder and white basswood growing on the prarie-derived soil, it decreased by 60 and 74%, respectively. Organic-S concentrations in bark increased more during autumn in species unable to fix atmospheric N(2), than in black alder. The results suggest that patterns of autumnal translocation of leaf S can be site-dependent and that leaf S and leaf N are, at least in part, translocated independently in the fall. Black alder and eastern cottonwood seemed to incorporate sulfate-S readily into organic substances in leaves when grown in soils with a high sulfate content.     

Effects of UV-B radiation on Olea europaea: comparisons between a greenhouse and a field experiment

We compared growth, morphology, anatomy and physiology of field-grown Olea europaea L. plants after one year with or without supplemental UV-B radiation equivalent to the increase at ground level resulting from a 15% depletion in stratospheric ozone concentration over Patras, Greece (38.3 degrees N, 29.1 degrees E). In a parallel greenhouse experiment, plants were exposed for four months to either zero UV-B radiation or UV-B radiation equivalent to that of the supplemented outdoor treatment. In the field, the only significant changes in response to supplemental UV-B radiation were an increase in adaxial epidermal thickness and a decrease in total protein concentration. In the greenhouse-grown plants, UV-B radiation caused significant increases in abaxial cuticle thickness and trichome UV-B absorbing compounds. We conclude that Olea europaea exhibits high UV-B tolerance and will not be affected by the predicted increases in UV-B radiation.     

Effects of enhanced ultraviolet-B radiation on water use efficiency, stomatal conductance, leaf nitrogen content and morphological characteristics of Spiraea pubescens in a warm-temperate deciduous broad-leaved forest

Spiraea pubescens, a common shrub in the warm-temperate deciduous forest zone which is distributed in the Dongling Mountain area of Beijing, was exposed to ambient and enhanced ultraviolet-B (UV-B, 280–320 nm) radiation by artificially supplying a daily dose of 9.4 kJ/m² for three growing seasons, a level that simulated a 17% depletion in stratospheric ozone. The objective of this study was to explore the effects of long-term UV-B enhancement on stomatal conductance, leaf tissue δ ¹³C, leaf water content, and leaf area. Particular attention was paid to the effects of UV-B radiation on water use efficiency (WUE) and leaf total nitrogen content. Enhanced UV-B radiation significantly reduced leaf area (50.1%) but increased leaf total nitrogen content (102%). These changes were associated with a decrease in stomatal conductance (16.1%) and intercellular CO₂ concentration/ air CO₂ concentration (C _i /C _a) (4.0%), and an increase in leaf tissue δ ¹³C (20.5‰), leaf water content (3.1%), specific leaf weight (SLW) (5.2%) and WUE (4.1%). The effects of UV-B on the plant were greatly affected by the water content of the deep soil (30–40 cm). During the dry season, differences in the stomatal conductance, δ ¹³C, and WUE between the control and UV-B treated shrubs were very small; whereas, differences became much greater when soil water stress disappeared. Furthermore, the effects of UV-B became much less significant as the treatment period progressed over the three growing seasons. Correlation analysis showed that enhanced UV-B radiation decreased the strength of the correlation between soil water content and leaf water content, δ ¹³C, C _i/C _a, stomatal conductance, with the exception of WUE that had a significant correlation coefficient with soil water content. These results suggest that WUE would become more sensitive to soil water variation due to UV-B radiation. Based on this experiment, it was found that enhanced UV-B radiation had much more significant effects on morphological traits and growth of S. pubescens than hydro-physiological characteristics. __________ Translated from Journal of Plant Ecology, 2006, 30(1): 47–56 [译自: 植物生态学报]     

Effects of long-term,elevated ultraviolet-B radiation on phytochemicals in the bark of silver birch (Betula pendula)

Long-term outdoor experiments were conducted to investigate the effects of elevated ultraviolet-B (UV-B, 280-320 nm) radiation on secondary metabolites (phenolics and terpenoids) and the main soluble sugars (sucrose, raffinose and glucose) in the bark of silver birch (Betula pendula Roth) saplings. Saplings were exposed to a constant 50% increase in erythemal UV irradiance (UV-B(CIE); based on the CIE (International Commission on Illumination) erythemal action spectrum) and a small increase in UV-A radiation (320-400 nm) for three growing seasons in an irradiation field in central Finland. Two control groups were used: saplings exposed to ambient radiation and saplings exposed to slightly increased UV-A radiation. Concentrations of sucrose, raffinose and glucose in bark were higher in UV-treated saplings than in saplings grown in ambient radiation, indicating that stem carbohydrate metabolism was changed by long-term elevated UV radiation. Saplings in the elevated UV-A + UV-B radiation treatment and the UV-A radiation control treatment had significantly increased concentrations of certain UV-absorbing phenolics, such as salidroside, 3,4'-dihydroxypropiophenone-3-glucoside, (+)-catechin and (-)-epicatechin compared with saplings in ambient radiation. In contrast, the radiation treatments had no effect on the non-UV-B-absorbing terpenoids, papyriferic acid and deacetylpapyriferic acid. We conclude that plant parts, in addition to leaves, accumulate specific phenolic UV-filters in response to UV radiation exposure.     

Foliar herbivory and leaf traits of five native tree species in a young plantation of Central Panama

This study examined foliar herbivory on 1 year-old tree saplings planted in previously abandoned fields in central Panama. Plots (15 × 15 trees) of Anacardium excelsum (Anacardiaceae), Dalbergia retusa (Fabaceae), Pachira quinata (Malvaceae), Tabebuia rosea (Bignoniaceae), and Terminalia amazonia (Combretaceae) were tested for herbivory using leaf counts and digital image analysis. Values of foliar carbon, foliar nitrogen, specific leaf area (SLA), and leaf toughness were analyzed to describe mechanical defenses and leaf nutrients on young and mature leaves of each of these species. For all five species, less than 10% of total leaf area was found to be damaged by arthropods. Significant (P-value < 0.001) differences in herbivory were found among both the tree species and the insect feeding guilds considered: chewing, skeletonizing, mining, and leaf-rolling. On mature leaves, Anacardium excelsum had the highest amount of leaf damage (3.53%) while Dalbergia retusa exhibited the lowest herbivore damage (1.72%). Tabebuia rosea had statistically significantly higher damage than other species for young leaves caused by leaf-rolling insects (4.21% rolling of 5.55% total damage). Leaf toughness was negatively correlated with SLA and foliar N. Linear regressions showed that herbivory was positively correlated with foliar N for young leaves and negatively correlated with foliar N for mature leaves. No statistically significant relationships were found between herbivory and the mechanical properties of toughness and SLA. Overall, results from this study indicate that, as young saplings, the species evaluated did not suffer high amounts of foliar herbivory in the plantation environment.     

Production physiology of three fast-growing hardwood species along a soil resource gradient

We determined how specific leaf area (SLA), specific leaf nitrogen (SLN), leaf area index (LAI), light-saturated photosynthesis (Amax) and aboveground net primary productivity (ANPP) of three commercially important hardwood species, eastern cottonwood (Populus deltoides Bartr.), American sycamore (Platanus occidentalis L.) and cherrybark oak (Quercus falcata var.pagodafolia Ell.), vary across a soil resource gradient. Five treatments were applied in a randomized block design (control, irrigation only (IRR), and irrigation plus fertilization with 56, 112 or 224 kg N ha-1 year-1 (N56, N112 and N224)) with four replications per species. When trees were 6 years old, Amax, SLA, SLN, LAI and ANPP were quantified during peak leaf production within a single growing season. In all species, Amax for sun leaves was significantly higher than for shade leaves (34, 32 and 29 micromol m2 s-1 versus 27, 23 and 23 micromol m2 s-1 for cottonwood, cherrybark oak and sycamore sun and shade leaves, respectively) and tended to plateau in the N112 treatment. The SLA was significantly lower in sun than in shade leaves and reached a plateau in IRR-treated cottonwood and sycamore, and in N56-treated oak. Values of SLN peaked in the N122 treatment for cottonwood sun leaves (1.73 g N m2) and in the N56 treatment for sycamore and oak (1.54 and 1.90 g N m2, respectively). In sun and shade leaves of all species, Amax increased with increasing SLN. Cherrybark oak LAI reached a plateau across the resource gradient in the N56 treatment, whereas cottonwood and sycamore LAI reached a plateau in the IRR treatment. All species exhibited significant curvilinear relationships between canopy Amax and ANPP. These findings indicate that nutrients and water regulate leaf-level traits such as SLA and SLN, which in turn influence LAI and canopy photosynthesis, thereby affecting ANPP at the tree and stand levels.     

The effects of ultraviolet-B radiation on photosynthesis of different aged needles in field-grown loblolly pine

We examined the effect of supplemental UV-B radiation (290-320 nm) on photosynthetic characteristics of different aged needles of 3-year-old, field-grown loblolly pine (Pinus taeda L.). Needles in four age classes were examined: I, most recently fully expanded, year 3; II, first flush, year 3; III, final flush, year 2; and IV, oldest needles still present, year 2. Enhanced UV-B radiation caused a statistically significant decrease (6%) in the ratio of variable to maximum fluorescence (F(v)/F(m)) following dark adaptation only in needles from the youngest age class, suggesting transient damage to photosynthesis. However, no effects of enhanced UV-B radiation on other instantaneous measures of photosynthesis, including maximum photosynthesis, apparent quantum yield and dark respiration, were seen for needles of any age. Foliar nitrogen concentration was unaffected by UV-B treatment. However, the (13)C/(12)C carbon isotope ratios (delta(13)C-a time integrated measure of photosynthetic function) of needles in age classes II and IV were 3% (P < 0.01) and 2% (P < 0.05) more negative, respectively, in treated plants than in control plants. Exposure to enhanced UV-B radiation caused a 20% decrease in total biomass and a 4% (P < 0.05), 25% (P < 0.01), and 9% (P < 0.01) decrease in needle length of needles in age classes I, II, and IV, respectively. The observed decreases in delta(13)C, and F(v)/F(m) of the needles in the youngest needle age class suggest subtle damage to photosynthesis, although overall growth reductions were probably a result of decreased total leaf surface rather than decreased photosynthetic capacity. Needles of age class IV had lower light- and CO(2)-saturated maximum photosynthetic rates (39%), lower dark respiration (34%), lower light saturation points (37%), lower foliar nitrogen concentration (28%), and lower delta(13)C (14%) values than needles of age class I. Apparent quantum yield and F(v)/F(m) did not change with needle age. The observed changes in photosynthesis and foliage chemical composition with needle age are consistent with previous studies of coniferous trees and may represent adaptations of older needles to shaded conditions within the canopy.     

Streamside trees: responses of male, female and hybrid cottonwoods to flooding

Cottonwoods, riparian poplars, are dioecious and prior studies have indicated that female poplars and willows can be more abundant than males in low-elevation zones, which are occasionally flooded. We investigated the response to flooding of clonal saplings of 12 male and 9 female narrowleaf cottonwoods (Populus angustifolia) grown for 15 weeks in a greenhouse, along with three females of a co-occurring native hybrid (Populus?×?jackii?=?Populus deltoides?×?Populus balsamifera). Three water-level treatments were provided, with substrate inundation as the flood treatment. In the non-flooded condition, the hybrids produced about four-fold more dry weight (DW) than the narrowleaf cottonwoods (P??P. angustifolia male?>?P.?×?jackii female. This indicates that narrowleaf cottonwoods are relatively flood tolerant and suggests that females are more flood tolerant than males. We propose the concept of 'strategic positioning', whereby the seed-producing females could be better adapted to naturally flooded, low-elevation streamside zones where seedling recruitment generally occurs.     

Relative efficacy of strobilurin formulations in the control of downy mildew of sunflower

The efficacy of three commercial formulations of strobilurins, viz., trifloxystrobin, kresoxim-methyl, and azoxystrobin was evaluated against sunflower downy mildew disease caused by Plasmopara halstedii under laboratory, greenhouse, and field conditions. Complete inhibition of sporangial sporulation, zoospore release, and motility was observed with 2 μg ml⁻¹ in trifloxystrobin, and 5 μg ml⁻¹ for azoxystrobin and kresoxim-methyl. Seed treatment with different concentrations of strobilurins enhanced seed germination and seedling vigor of sunflower to varying degrees compared to control. Highest seed germination was recorded at 10 μg ml⁻¹ in kresoxim-methyl, and maximum seedling vigor was noticed with trifloxystrobin at 30 μg ml⁻¹. The effect of strobilurins was tested as seed treatment, foliar application, and seed treatment followed by foliar application. Under greenhouse conditions none of the concentrations used, either as seed treatment and foliar application, were phytotoxic. For the three strobilurins, the seed treatment along with foliar application enhanced the protection of the plants as compared to only the treatment of seeds. Foliar spray treatments alone provided an intermediate control of the disease. Trifloxystrobin showed a better effect than kresoxim-methyl and azoxystrobin. Disease curative activity of trifloxystrobin was higher compared to kresoxim-methyl and azoxystrobin. Tested fungicides when applied on adaxial leaf surface showed partial translaminar activity, and disease inhibition was marginal. Loss of trifloxystrobin and azoxystrobin activity over time was low, indicating stable rainfastness residual activity. A trend in protection against downy mildew disease similar to greenhouse results was evident in the field trials. This is the first report of strobilurins exhibiting high activity against P. halstedii and is a promising fungicide for controlling sunflower downy mildew disease by seed treatment and foliar spray.     

Induction of flavonoid production by UV-B radiation in Passiflora quadrangularis callus cultures

Callus cultures from several species of Passiflora were initiated in vitro, and their capacity to produce four glycosyl flavonoids (orientin, isoorientin, vitexin, isovitexin) was analysed. The aim of the present work was to examine the possible role of UV-B irradiation and elicitation with methyl jasmonate (MJ) on the production of these compounds in callus cultures. All the species tested (P. incarnata, P. quadrangularis, P. edulis) formed friable callus from leaf explants after 4 weeks on medium supplemented with kinetin and 2,4-dichlorophenoxyacetic acid. Among them, P. quadrangularis turned out to have a faster growth rate and a more friable texture, and was therefore chosen for experiments with elicitors. In callus cultures only small amounts of isoorientin were found, while the concentration of the other flavonoids was below the detection limit. UV-B irradiation of calluses was able to increase the production of all four glycosyl flavonoids. After a 7-day exposure of cultures to UV-B light, the production of isoorientin reached concentrations similar to those found in fresh leaves from glasshouse-grown plants. Elicitation with methyl jasmonate also enhanced orientin, vitexin and isovitexin concentrations, even though the stimulation was about 6-fold weaker for orientin and vitexin and about 40-fold for isovitexin, than that exerted by UV-B treatment. Callus cultures treated with the UV-B dose which most enhanced flavonoid production showed a higher antioxidant activity compared to untreated calluses, with an increase ranging from 28% to 76%. Results show that the secondary metabolite biosynthetic capacity of Passiflora tissue cultures can be enhanced by appropriate forms of elicitation.     

UV-B辐射对亚热带森林凋落叶分解的影响

采用分解袋法研究自然和UV-B辐射滤减2种环境下6种亚热带代表性树种(杉木、马尾松、木荷、香樟、青冈和甜槠)凋落叶的分解情况。结果表明:除个别分解阶段外,各树种凋落叶在2种UV-B辐射环境下的干质量剩余率均存在显著差异,且随着分解时间延长,差异逐渐加大;与对照相比,UV-B辐射滤减显著降低了6个树种凋落叶的分解速率(P<0.01),降幅为33.3%～69.6%,对香樟凋落叶分解的影响最小,对杉木凋落叶分解的影响最大;UV-B辐射处理和凋落物类型对凋落叶的分解速率均有极显著影响(P<0.001),以UV-B辐射的影响更强烈;自然和UV-B辐射滤减环境下凋落叶的分解速率均与C∶N呈显著负相关(P<0.05)。