Changes in leaf expansion and epidermal screening effectiveness in Liquidambar styraciflua and Pinus taeda in response to UV-B radiation

Absorption or screening of ultraviolet-B (UV-B) radiation by the epidermis may be an important protective method by which plants avoid damage upon exposure to potentially harmful UV-B radiation. In the present study we examined the relationships among epidermal screening effectiveness, concentration of UV-absorbing compounds, epidermal anatomy and growth responses in seedlings of loblolly pine (Pinus taeda L.) and sweetgum (Liquidambar styraciflua L.). Seedlings of each species were grown in a greenhouse at the University of Maryland under either no UV-B radiation or daily supplemental UV-B radiation levels of 4, 8 or 11 kJ m^?2 of biologically effective UV-B (UV-B_BE) radiation. Loblolly pine seedlings were subsequently grown in the field under either ambient or supplemental levels of UV-B radiation. At the conclusion of the growing season, measurements of epidermal UV-B screening effectiveness were made with a fiber-optic microprobe. In loblolly pine, less than 0.5% of incident UV-B radiation was transmitted through the epidermis of fascicle needles and about 1% was transmitted in primary needles. In contrast, epidermal transmittance in sweetgum ranged from about 20% in leaves not preconditioned to UV-B exposure, to about 10% in leaves grown under UV-B radiation. The concentration of UV-absorbing compounds was unaffected by UV-B exposure, but generally increased with leaf age. Increases in epidermal thickness were observed in response to UV-B treatment in loblolly pine, and this accounted for over half of the variability in UV-B screening effectiveness. In spite of the low levels of UV-B penetration into the mesophyll, delays in leaf development (both species) and final needle size (loblolly pine) were observed. Seedling biomass was reduced by supplemental UV-B radiation in loblolly pine. We hypothesize that the UV-induced growth reductions were manifested by changes in either epidermal anatomy or epidermal secondary chemistry that might negatively impact cell elongation.     

Penetration of UV-B radiation in foliage: evidence that the epidermis behaves as a non-uniform filter

In some plants, particularly herbaceous species, a considerable proportion of incident ultraviolet-B radiation (UV-B, 280-320 nm) penetrates into the leaf mesophyll where it is potentially damaging to nucleic acids and the photosyn-thetic machinery. We used optical techniques to look at the spatial variation in UV-B penetration through the epidermis of foliage of two herbaceous species (Chenopodium album and Smilacina stellata)and a conifer (Picea pun-gens). Measurements of UV-B penetration in intact foliage with a fibre-optic microprobe revealed that 300 nm radiation reached 161±36μm (mean±SD) into leaves of C. album, 154±40μm in S. stellata and 17±2μm in P. pungens, with epidermal transmittance being 39±14%, 55±19% and 0%, respectively. A thin polymer film was developed which fluoresced blue when irradiated by UV-B. Fresh epidermal leaf peels were placed over the film and irradiated with UV-B, and microscopic examination of the film from below allowed us to determine the spatial pattern of UV-B penetration through the epidermis. In herbaceous species, film fluorescence below cell walls, but not epidermal and guard cell protoplasts indicated that UV-B transmittance was much greater through anticlinal cell wall regions than protoplasts. Ultraviolet-B transmittance through large areas of epidermal cells could be induced by plasmolysis. Epidermal transmittance was also relatively high through stomal pores (and what appear to be nuclei in Smilacina), but relatively low through stomatal guard cells. Results from the fluorescing film technique were substantiated by direct measurements of UV-B transmittance through epidermal peels with a fibre-optic microprobe run paradermally along the bottom or inner side of irradiated peels. In Smilacina, we estimate that UV-B epidermal transmittance was up to 90% through anticlinal cell wall regions, but <10% through protoplast areas. In contrast to herbaceous species, we did not detect any UV-B transmittance through the epidermis of P. pungens with either the fluorescing film or the fibre-optic microprobe technique. The epidermis appears to be a much more spatially uniform UV-B filter in conifers than in these herbaceous species.     

Are some plant life forms more effective than others in screening out ultraviolet-B radiation?

Summary The unprecedented rate of depletion of the stratospheric ozone layer will likely lead to appreciable increases in the amount of ultraviolet-B radiation (UV-B, 280–320 nm) reaching the earth's surface. In plants, photosynthetic reactions and nucleic acids in the mesophyll of leaves are deleteriously affected by UV-B. We used a fiber-optic microprobe to make direct measurements of the amount of UV-B reaching these potential targets in the mesophyll of intact foliage. A comparison of foliage from a diverse group of Rocky Mountain plants enabled us to assess whether the foliage of some plant life forms appeared more effective at screening UV-B radiation. The leaf epidermis of herbaceous dicots was particularly ineffective at attenuating UV-B; epidermal transmittance ranged from 18–41% and UV-B reached 40–145 m into the mesophyll or photosynthetic tissue. In contrast to herbaceous dicots, the epidermis of 1-year old conifer needles attenuated essentially all incident UV-B and virtually none of this radiation reached the mesophyll. Although the epidermal layer was appreciably thinner in older needles (7 y) at high elevations (Krumholtz), essentially all incident UV-B was attenuated by the epidermis in these needles. The same epidermal screening effectiveness was observed after removal of epicuticular waxes with chloroform. Leaves of woody dicots and grasses appeared intermediate between herbaceous dicots and conifers in their UV-B screening abilities with 3–12% of the incident UV-B reaching the mesophyll. These large differences in UV-B screening effectiveness suggest that certain plant life forms may be more predisposed than others to meet the challenge of higher UV-B levels resulting from stratospheric ozone depletion.     

Site-dependent differences in transmittance and UV-B-absorbing capacity of isolated leaf epidermes and mesophyll in Urginea maritima (L.) Baker

The spectral transmittance of isolated 'intact' upper and lower epidermes as well as the extractable UV-B-absorbing capacity of epidermes and mesophyll were studied in the leaves of exposed and deeply shaded, field-grown plants of Urginea maritima (L.) Baker. Epidermal transmittance in the visible part of the spectrum was high (>80%) in all cases. Transmittance in the UV-B (280-320 nm) was comparatively high (c. 14%) in both the upper and lower epidermes of shaded plants, but more than an order of magnitude lower in exposed plants, with the lowest values observed on the upper leaf epidermis. UV-B transmittance was negatively correlated with the methanol extractable UV-B-absorbing capacity of the epidermes, but was independent of epidermal thickness. The UV-B-absorbing capacity of the mesophyll, when expressed on an area basis, was not affected by exposure. However, it was significantly higher in shaded plants, when expressed on a dry mass basis. The results indicate that although the concentrations of the UV-B-absorbing components of the whole leaf or its epidermis fluctuate according to the site-dependent radiation stress, the opposite is evident for the mesophyll. Therefore, high irradiance in U. maritima, apart from inducing an increase in UV-B-absorbing compounds on a whole leaf basis, also caused a change in the distribution of these compounds between epidermis and mesophyll.     

Epidermal transmittance and phenolic composition in leaves of atrazine-tolerant and atrazine-sensitive cultivars of Brassica napus grown under enhanced UV-B radiation

Experiments were conducted on the atrazine-tolerant mutant Stallion and the atrazine-sensitive cv. Paroll of Brassica napus L., which were grown under either visible light or with the addition of UV-B radiation (280–320 nm) for 15 days. The mutant has been shown to be sensitive to high levels of visible light as compared to the atrazine-sensitive cultivar and therefore we wished to determine plant response to UV-B radiation with respect to potential pigment changes, certain anatomical features, radiation penetration and partial photosynthesis. With regard to pigment changes, we were particularly interested in whether the compositional shift in flavonol pigments under enhanced UV-B radiation, previously suggested to favour increased antioxidant activity, is confined to the adaxial epidermis, which generally receives most UV-B radiation or whether the pigment shift is also inducible in the abaxial epidermis.As was to be expected, the penetration of UV-B radiation (310 nm) was lower in the UV-B-exposed plants, which was correlated with an increased amount of UV-screening pigments in the adaxial and abaxial epidermal layers. The main flavonoid glycosides showed the largest shift from kaempferol to quercetin as aglycone moiety in the adaxial epidermal layer. However, in the abaxial epidermal layer the hydroxycinnamic acid (HCA) derivatives and kaempferol glycosides were predominant. Penetration of 430 nm light was higher after UV-B exposure, and probably contributed to the fact that photosynthetic efficiency of photosystem II was unchanged or higher after UV-B exposure. UV-B radiation decreased leaf area in the atrazine-tolerant mutant only. Both cultivars showed an increased leaf thickness after UV-B exposure due to cell elongation mainly of the palisade tissue. This was especially evident in the mutant.     

Ultraviolet-B and visible light penetration into needles of two species of subalpine conifers during foliar development

The depth of penetration of Ultraviolet-B (UV-B, 300 and 320 nm) and visible (680 nm) light was measured in foliage of Abies lasiocarpa and Picea engelmannii using a fibre-optic microprobe. Measurements were made on foliage at four times during development: needles were sampled from within expanding buds (in bud); within 72 h of emergence from the bud scales (emergent); from elongating branches (elongating); and from foliage that emerged the previous summer (mature). Light attenuation in pre-emergent needles of both species was steep and showed strong wavelength dependence. Short wavelength 300-nm light was attenuated strongly in the developing epidermal layer, but a significant proportion of this potentially damaging UV-B radiation penetrated into the mesophyll. For A. lasiocarpa and P. engelmannii, 99% attenuation of 300-nm light occurred at 51 and 96 μm, respectively, well within the mesophyll. At this stage, however, the bud scales were opaque to light below 400nm. As the epidermal cell walls and cuticle continued to develop and chlorophyll accumulated following emergence from the bud scales, light attenuation, particularly of UV-B radiation, increased. Although no UV-B is transmitted through the epidermis-hypodermis of mature needles, small but measurable quantities of 300- and 320-nm light were measured in the photosynthetic mesophyll of post-emergent and elongating needles. Thus, shortly after emergence from the bud scales in mid-June to mid-July, when incident UV doses are highest, absorption of UV-B radiation by potentially sensitive chromophores in the mesophyll may disrupt physiological and developmental processes in these species. Soluble UV-absorbing pigments accumulated during needle maturation for P. engelmannii but not A. lasiocarpa, suggesting that, for A. lasiocarpa at least, the development of effective UV screening properties in the epidermis may not be related to the induction of soluble flavonoids.     

Effects of Natural Intensities of Visible and Ultraviolet Radiation on Epidermal Ultraviolet Screening and Photosynthesis in Grape Leaves

Grape (Vitis vinifera cv Silvaner) vine plants were cultivated under shaded conditions in the absence of ultraviolet (UV) radiation in a greenhouse, and subsequently placed outdoors under three different light regimes for 7 d. Different light regimes were produced by filters transmitting natural radiation, or screening out the UV-B (280-315 nm), or screening out the UV-A (315-400 nm) and the UV-B spectral range. During exposure, synthesis of UV-screening phenolics in leaves was quantified using HPLC: All treatments increased concentrations of hydroxycinnamic acids but the rise was highest, reaching 230% of the initial value, when UV radiation was absent. In contrast, UV-B radiation specifically increased flavonoid concentrations resulting in more than a 10-fold increase. Transmittance in the UV of all extracted phenolics was lower than epidermal UV transmittance determined fluorimetrically, and the two parameters were curvilinearly related. It is suggested that curvilinearity results from different absorption properties of the homogeneously dissolved phenolics in extracts and of the non-homogeneous distribution of phenolics in the epidermis. UV-B-dependent inhibition of maximum photochemical yield of photosystem II (PSII), measured as variable fluorescence of dark-adapted leaves, recovered in parallel to the buildup of epidermal screening for UV-B radiation, suggesting that PSII is protected against UV-B damage by epidermal screening. However, UV-B inhibition of CO(2) assimilation rates was not diminished by efficient UV-B screening. We propose that protection of UV-B inactivation of PSII is observed because preceding damage is efficiently repaired while those factors determining UV-B inhibition of CO(2) assimilation recover more slowly.     

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

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

Changes in UV-B radiation screening effectiveness with leaf age in Rhododendron maximum

We examined how ultraviolet-B radiation (UV-B; 300 nm) screening effectiveness changes with leaf age in Rhododendron maximum growing in a shaded understory by measuring depth of penetration and epidermal transmittance with a fibre-optic microprobe. Depth of penetration (and epidermal transmittance) of UV-B decreased with leaf age in 1- to 4-year-old leaves, averaging 62 (32), 52 (22), 45 (16) and 48 μm (13%), respectively. Epidermal thickness increased with age in 1- to 4-year-old leaves due to a thickening of the cuticle from an average of 20 to 29μm. Ultraviolet-B-absorbing compound concentrations increased with age from 1–3 to 1–5 A₃₀₀ cm^?2 leaf area. Concentrations of UV-B-absorbing compounds (area basis) were a strong predictor of depth of penetration (r²= 0.82) and epidermal transmittance (r²= 0.95) of UV-B in mature (1–4 year-old) foliage. Chlorophyll concentrations (area basis) increased in leaves up to 3 years of age. Current-year leaves (30 d old) were exceptional in that while they were particularly effective at screening UV-B (depth of penetration and epidermal transmittance averaged 39μm and 5%, respectively) they had relatively low concentrations of UV-B-absorbing compounds (1.3 A₃₀₀ cm^?2). Our findings show that UV-B-screening effectiveness is not necessarily related to absorbing compound concentrations on a whole-leaf basis, possibly due to anatomical changes within the epidermis that occur with leaf age.     

Effects of Epidermal Cell Shape and Pigmentation on Optical Properties of Antirrhinum Petals at Visible and Ultraviolet Wavelengths

We used the Mixta+ and mixta- lines of Antirrhinum majus as a model system to investigate the effects of epidermal cell shape and pigmentation on tissue optical properties in the visible and ultraviolet (UV) spectral regions. Adaxial epidermal cells of Mixta+ flowers have a conical-papillate shape; in the mixta- line the cells are slightly domed. Mixta+ cells contained significantly more anthocyanin and other flavonoids than mixta- cells when plants were grown under either high- or low-UV conditions. Mixta+ cells focused light (3.5-4.7 times incident) within their pigmented interiors, whereas mixta- cells focused light (2.1-2.7 times incident) in the unpigmented mesophyll. UV light penetrated the epidermis (commonly 20-50% transmittance at 312 nm) mainly through the unpigmented peripheral regions of the cells that were similar for the two lines, so that overall penetration through Mixta+ and mixta- epidermises was equal. However, maximum UV absorption in the central region of epidermal cells was slightly greater in Mixta+ than mixta-, and intact Mixta+ flowers reflected less light in the spectral regions with intermediate flavonoid absorbance. In both cases, about 50 to 75% of the difference could be attributed to cell shape and resulting changes in the optical pathlength or focusing.     

Epidermal UV-screening in vascular plants from Svalbard (Norwegian Arctic)

Stratospheric ozone depletion is most pronounced at high latitudes, and the concurring increased UV-B radiation might adversely affect plants from polar areas. However, vascular plants may protect themselves against UV-B radiation by UV-absorbing compounds located in the epidermis. In this 3-year study, epidermal UV-B (_max 314 nm) and UV-A (_max 366 nm) screening was assessed using a fluorescence method in 12 vascular species growing in their natural environment at Svalbard. The potential for acclimation to increased radiation was studied with artificially increased UV-B, simulating 11% ozone depletion. Open-top chambers simulated an increase in temperature of 2–3°C in addition to the UV-B manipulation. Adaxial epidermal UV-B transmittance varied between 1.6 and 11.4%. Artificially increased UV-B radiation and temperature did not consistently influence the epidermal UV-B transmittance in any of the measured species, suggesting that they may not have the potential to increase their epidermal screening, or that the screening is already high enough at the applied UV-B level. We propose that environmental factors other than UV-B radiation may influence epidermal UV-B screening.     

Impact of solar ultraviolet-B radiation on the antioxidant defense system in soybean lines differing in flavonoid contents

Exposure to ultraviolet-B (UV-B) radiation can lead to oxidative damage in plants. However, plants possess a number of UV-protection mechanisms including screening of potentially damaging UV-B and increased production or activities of antioxidants. The balance or trade-off between these two mechanisms has rarely been studied and is poorly understood. Two isolines of soybean (Glycine max [L.] Merr.) Clark cultivar, the normal line with moderate levels of flavonoids and the magenta line with reduced flavonoids levels, were grown in the field with or without natural levels of UV-B. Leaflet blades of the first trifoliate leaf were harvested after 4–12 days of exposure to the experimental conditions for analysis of active oxygen species (AOS) and antioxidant levels. Solar UV-B radiation caused oxidative stress in both lines and altered AOS metabolism primarily by decreasing superoxide dismutase activity and increasing the activities of ascorbate peroxidase, catalase and glutathione reductase. This resulted in decreased ascorbic acid content and increased dehydroascorbate content. The magenta line had greater oxidative stress than the normal line in spite of its enhanced oxidative defense capacity as compared to the normal line, even under UV-B exclusion. These results indicate enhanced sensitivity in the magenta line, especially under UV-B exclusion that was likely due to the absence of flavonoid epidermal screening compounds and subsequent increased penetration of solar ultraviolet radiation into the leaf.     

Effects of ultraviolet-B radiation on cotton (Gossypium hirsutum L.) morphology and anatomy

Cotton (Gossypium hirsutum L.) crop, cultivated between 40 degrees N and 40 degrees S, is currently experiencing 2-11 kJ m-2 d-1 of UV-B radiation. This is predicted to increase in the near future. An experiment was conducted to study the effect of enhanced UV-B radiation on vegetative and reproductive morphology and leaf anatomy of cotton in sunlit, controlled environment chambers. From emergence to harvest, cotton plants were exposed to 0, 8 or 16 kJ m-2 d-1 of UV-B in a square wave approach for 8 h from 0800 to 1600 h. Changes in plant height, internode and branch length, mainstem node number, leaf area, length and area of petals and bracts, and anther number per flower were recorded. Epidermal cell and stomatal density, stomatal index, leaf thickness, and epidermal, palisade and mesophyll tissue thickness were also measured. Initial chlorotic symptoms on leaves turned into necrotic patches on continued exposure to enhanced UV-B. Exposure to high UV-B reduced both vegetative and reproductive parameters and resulted in a smaller canopy indicating sensitivity of cotton to UV-B radiation. Enhanced UV-B radiation increased epicuticular wax content on adaxial leaf surfaces, and stomatal index on both adaxial and abaxial leaf surfaces. Leaf thickness was reduced following exposure to UV-B owing to a decrease in thickness of both the palisade and mesophyll tissue, while the epidermal thickness remained unchanged. The vegetative parameters studied were affected only by high levels of UV-B (16 kJ m-2 d-1), whereas the reproductive parameters were reduced at both ambient (8 kJ m-2 d-1) and high UV-B levels. The study shows that cotton plants are sensitive to UV-B at both the whole plant and anatomical level.     

Internal filters: Prospects for UV-acclimation in higher plants

Wavelength-selective absorption of solar radiation within plant leaves allows penetration of visible radiation (400–700 nm) to the chloroplasts, while removing much of the damaging ultraviolet-B (UV-B, 280–320 nm) radiation. Flavonoids are important in this wavelength-selective absorption. Induction of flavonoid synthesis by solar radiation, and specifically by UV-B radiation, is discussed as this relates to the potential acclimation of plants to enhanced solar UV-B radiation that would result from stratospheric ozone reduction.     

Measurement of leaf epidermal transmittance of UV radiation by chlorophyll fluorescence

In higher plants one of the important functions of the leaf epidermis is the effective screening of ultraviolet-B (280–320 nm, UV-B) radiation, due mostly to phenolic compounds. The assessment of the contribution of this function is necessary for an evaluation of the impact of increasing UV-B radiation. A method is proposed to estimate epidermal transmittance on the basis of chlorophyll fluorescence measurements. Fluorescence of chlorophyll induced by UV-A (320–400 nm, measuring beam centered at 366 nm, half band width 32 nm) or UV-B (measuring beam centered at 314 nm, half band width 18 nm) is compared to that induced by a blue-green measuring light (475 nm, half band width 140 nm). It is shown that the ratios of UV-and blue-green (BG)-induced fluorescence, F(UV-A)/F(BG) and F(UV-B)/F(BG), are relatively constant among leaf samples of various species ( Vicia faba, Spinacia oleracea, Rumex scutatus ) from which the epidermis was removed. In epidermis-free leaves no significant differences were found between adaxial and abaxial leaf sides, suggesting that leaf structure has negligible influence on the F(UV)/F(BG) ratios. On the other hand, fluorescence excitation ratios varied over a vast range when intact leaves from different species and habitats were investigated. Ratios were low in sun leaves and relatively high in shade- and greenhouse-grown leaves. By relating these results to those obtained with epidermis-free leaves, epidermal transmittances for UV-B radiation could be estimated, with values ranging between 1 and 45%. The data demonstrate a large adaptability of epidermal UV-A and UV-B transmittance in higher plants. The proposed method may prove a versatile and relatively simple tool for investigating epidermal UV transmittance complementing established methods.     

Penetration of UV-A, UV-B and blue light through the leaf trichome layers of two xeromorphic plants, olive and oak, measured by optical fibre microprobes

Quartz fibre-optic microprobes were used to monitor the light microenvironment beneath trichome layers of the xeromorphic leaves of two Mediterranean evergreen sclerophylls, Olea europaea and Quercus ilex . Young developing leaves of both plants were densely pubescent on both surfaces of the lamina, whereas the mature leaves were pubescent only on the abaxial side. Trichome layers of young as well as of mature leaves of both plants attenuated almost all incident ultraviolet (UV)-B (310 nm) and UV-A (360 nm) radiation and a considerable portion of blue light (430 nm). Abaxial trichome layers of young leaves were more effective in screening out the incident radiation compared to the adaxial ones of the same leaves and also compared to the abaxial layer of the mature leaves. The abaxial epidermis of dehaired mature leaves of O. europaea was ineffective in absorbing most of the incident UV-B and UV-A radiation. UV and visible spectra beneath trichome layers of O. europaea in mature leaves confirmed that the light microenvironment on the epidermis was deprived in the UV-B, UV-A and partly in the blue spectral regions. It is proposed that the occurrence of a dense trichome layer, especially in young leaves, may play a protective role against not only UV-B radiation damage, but also against high visible irradiance. This function is performed irrespective of the differing anatomy of individual hairs of both plants. The protection provided by the trichomes could afford advantages under stress conditions, especially during leaf development.     

Epidermal transmittance of leaves of Vicia faba for UV radiation as determined by two different methods

Leaves of Vicia faba were collected from the field and the greenhouse and transmittance of epidermal peels from adaxial and abaxial sides was determined in the wavelength range from 250 to 800 nm using a spectrophotometer equipped for the measurement of turbid samples. From the same leaves, epidermal transmittance was estimated by a recently developed fluorometric method. Both methods gave highly correlated results with a slope of the regression line between both methods close to 1 and an intercept close to 0. Transmittances at around 310 nm as low as 3% were detected in the adaxial epidermis of field-grown leaves, while transmittance could be as high as 70% in the abaxial epidermis of greenhouse-grown leaves. There was a strong correlation between UV-A (ca. 366 nm) and UV-B (ca. 310 nm) transmittance detected by both methods which could be explained by the pigment composition in methanolic extracts where flavonols accounted for 90% of the absorption at 310 nm in the extract, while hydroxycinnamic acid derivatives which absorb only at the shorter wavelength constituted about 5%. It is concluded that the fluorescence method which allows rapid measurements on intact leaves can provide a quantitative estimate of epidermal transmittance for UV-B (280–320 nm) and UV-A (320–400 nm) radiation.This revised version was published online in October 2005 with corrections to the Cover Date.     

SOME EFFECTS OF ULTRA-VIOLET RADIATION ON LEAVES OF FRENCH BEAN (PHASEOLUS VULGARIS L.)

Exposing the primary leaves of French bean to ultra-violet light produces a variety of effects, their magnitude depending on the extent of the exposure and the subsequent treatment of the leaves. Effects include some that are externally obvious, such as glazing and bronzing; others are detectable by changes in susceptibility to virus infection, by increased sensitivity to damage by ultra-violet, and by an increased tendency for cells to collapse when the leaves are kept in darkness. Some of the effects can be counteracted by exposing leaves to visible light. Effects of radiation are not confined to epidermal cells, and measurements with detached epidermis indicate that from a quarter to a half of incident radiation of wave-length 2536 A. may be transmitted by the epidermis.     

Giant clams in shallow reefs: UV-resistance mechanisms of Tridacninae in the Red Sea

The photosymbiosis of tropical giant clams (subfamily Tridacninae) with unicellular algae (Symbiodiniaceae) restricts their distribution to the sunlit, shallow waters of the euphotic zone where organisms are additionally exposed to potentially damaging levels of solar UV radiation. Metabolic and physiological responses of Red Sea Tridacna maxima clams, including net calcification and primary production, as well as valvometry (i.e., shell gaping behavior) were assessed when exposed to simulated high radiation levels received at 3 and 5 m underwater. The two levels of radiation included exposure treatments to photosynthetically active radiation (PAR; 400–700 nm) alone and to both, PAR and ultraviolet-B radiation (UV-B; 280–315 nm). The valvometry data obtained using flexible magnetic sensors indicated that specimens under PAR + UV-B exposure significantly reduced the proportion of their exposed mantle area, a potential photo-protective mechanism which, however, reduces the overall amount of PAR received by the algal symbionts. Consequently, specimens under PAR + UV-B displayed a slight, although non-significant, reduction in primary production rates but no signs of additional oxidative stress, changes in symbiont densities, chlorophyll content, or levels of mycosporine-like amino acids. Net calcification rates of T. maxima were not affected by exposure to UV-B; however, calcification was positively correlated with incident PAR levels. UV-B exposure changes the valvometry, reducing the exposed mantle area which consequently diminishes the available PAR for the photosymbionts. Still, T. maxima maintains high rates of primary production and net calcification, even under high levels of UV-B. This provides experimental support for a recently described, effective UV-defensive mechanism in Tridacninae, in which the photonic cooperation of the associated algal symbionts and giant clam iridocytes is assumed to establish optimal conditions for the photosynthetic performance of the clams’ symbionts.

     

Relating UV-B radiation screening effectiveness of foliage to absorbing-compound concentration and anatomical characteristics in a diverse group of plants

The ultraviolet-B radiation (UV-B, 300 nm) screening effectiveness of foliage of a diverse group of plants was examined by measuring epidermal transmittance and depth of penetration of UV-B with a fiberoptic microprobe. In addition, the concentration of UV-B-absorbing compounds and various anatomical characteristics were measured to assess whether they were useful predictors of UV-B screening. Sun foliage of naturally growing individuals of seven species were sampled in each of six life forms comprising two evergreen groups (gymnosperms and angiosperms) and four deciduous angiosperm groups (trees, shrubs and vines, herbaceous dicotyledons, and grasses). There was significant life-form variation in epidermal transmittance and depth of penetration of UV-B, concentration of UV-B-absorbing compounds (leaf-area basis), epidermal (including cuticle and hypodermis) thickness, and specific leaf area. Values of these parameters tended to be related to leaf longevity, with the most notable differences apparent between evergreen and deciduous species. The mean epidermal transmittance and depth of penetration of UV-B in foliage averaged 4% and 32 m in evergreens, compared to 28% and 75 m in deciduous species. These values are conservative estimates since the microprobe was oriented in foliage such that much of the side- and backscattered UV-B was ignored. The strongest predictors of epidermal transmittance and depth of penetration were epidermal thickness and the concentration of absorbing compounds, which averaged 32 m and 1.50 A cm^–2 in evergreens, but only 19 m and 0.99 A cm^–2 in deciduous foliage. However, the variation found in these relationships implies that additional factors warrant consideration in assessing UV-B-screening effectiveness. The relatively ineffective screening of UV-B by foliage of many deciduous plants suggests they may be more responsive to enhanced UV-B than evergreen species.