Influence of solar ultraviolet-B on pelagic fish embryos: osmolality, mortality and viable hatch

Eggs of dab (Limanda limanda) and plaice (Pleuronectes platessa) were experimentally exposed to ultraviolet-B (UV-B) radiation in a solar radiation simulator. The experimental design tried to simulate present and future conditions with reference to increased UV-B exposure due to northern hemisphere ozone loss, employing mainly two scenarios, a reduction to 270 (S1) and to 180 (S2) Dobson units (DU) in single or repetitive exposures of 2, 4 or 6 h. Depending on the total dose of UV-B irradiation and the developmental stage, exposed eggs displayed loss of buoyancy as a sublethal effect, as well as increased embryo mortality and reduced viable hatch. In the single exposure experiments only under conditions of 180 DU for 6 h were effects apparent. Double exposure under conditions of 270 DU did not lead to lasting effects. At the sublethal effect level, i.e. loss of buoyancy, considerable photorepair was observed. It was concluded, that under the present general weather conditions in spring and at the present levels of environmental ozone, allowing for a reduction to 180 DU, the embryonic development of North Sea spring spawning fish is not endangered by UV-B radiation. Received in revised form: 19 June 2000 Electronic Publication     

Short-term UV-B radiation and ozone exposure effects on aromatic secondary metabolite accumulation and shoot growth of flavonoid-deficient Arabidopsis mutants

The presence of UV-absorptive substances in the epidermal cells of leaves is thought to protect mesophyll tissues from the harmful effects of UV-B radiation. We examined the influence of short-term UV-B exposures on UV-absorptive (330 nm) sinapates and flavonols, and on shoot growth of the Arabidopsis wild type ecotype Landsberg erecta and two mutants. 114 deficient in chalcone synthase, and 115 , deficient in chalcone/flavonone isomerase. Sequential ozone exposures were used to determine the effects of oxidative stress The levels of sinapates and flavonols on a leaf fresh weight basis increased substantially in the wild type and sinapates increased in the 114 mutant in vegetative vegetative/reproductive transitional and reproductive stage plants in response to short-term (48h) UV-B radiation. When UV-B was discontinued the levels generally decreased lo pre-exposure levels after 48 h in vegetative/reproductive but not in reproductive plants. Exposure to ozone before or alter UV-B treatment did not consistently affect the levels of these UV-absorptive compounds. Dry matter accumulation was less affected by UV-B at the vegetative and reproductive stages than at the vegetative/reproductive stage. At the vegetative/reproductive stage, shoot growth of all 3 genotypes was retarded by UV-B. Growth was not retarded by short-term ozone exposure alone but when exposure to ozone followed UV-B exposure, growth was reduced in all genotypes. Leaf cupping appeared on 115 plants exposed to UV-B.     

The effect of enhanced ultraviolet-B radiation on growth,photosynthesis and stable carbon isotope composition (δ13C) of two soybean cultivars (Glycine max) under field conditions

Two Chinese cultivars of Glycine max, namely Heidou and Jindou, were exposed to ambient and supplemental levels of ultraviolet-B (UV-B) radiation simulating a 24% depletion in stratospheric ozone over a 9-week growing period at an outdoor experimental site. Enhanced UV-B irradiation significantly reduced leaf, stem and root biomass, and plant height in the Heidou cultivar. These changes were associated with a diminished photosynthetic (net CO₂) rate, stomatal conductance, transpiration rate and water use efficiency, and accompanied by decreased foliar chlorophyll a and b, and total carotenoid concentrations and elevated foliar flavonoid levels. In contrast, the Jindou cultivar displayed only a significantly reduced stem mass and stomatal conductance, but no changes in pigment composition under elevated UV-B. The greater tolerance of elevated UV-B exposures by the Jindou cultivar was attributed partly to its higher foliar flavonoid content, smaller leaf size, thicker leaf cuticle and scabrous (hairy) lamina. Nevertheless both the Heidou cultivar and the less UV-B sensitive Jindou cultivar displayed an altered carbon isotope composition (δ¹³C) in their tissues following exposure to elevated UV-B. Such carbon isotope composition changes in plant tissues suggested a means of early detection of photosynthetic disruption in plants with anticipated increase in UV-B due to stratospheric ozone depletion.     

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

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

The effects of ultraviolet-B radiation on loblolly pine

Summary Depletion of stratospheric ozone and the resulting increase in ultraviolet-B (UV-B) radiation may negatively impact the productivity of terrestrial ecosystems. This concern has led to a number of studies that report the influence of supplementing UV-B radiation on plant growth and development. However, only two of these field studies have included tree species and both were singleseason experiments. In this study, loblolly pine (Pinus taeda L.) from seven seed sources was grown under natural and supplemental levels of UV-B radiation. Irradiation treatments were continued for three seasons on plants from four of the seven groups and for 1 year only for three groups. The supplemental irradiances simulated those that would be anticipated with stratospheric ozone reductions of 16% and 25% over Beltsville, Md. The effects of UV-B radiation during the 1st year on plant growth varied among the seed sources. The growth of plants from two of the seven seed sources tested showed significant reductions following a single irradiation season and plants from one group tended to be larger under increased UV-B radiation. However, after 3 years of supplemental irradiation, plant biomass was reduced in all four groups by 12–20% at the highest simulated ozone depletion. These results suggest that the effects of UV-B radiation may accumulate in trees and that increased UV-B radiation could significantly reduce the growth of loblolly pine over its lifetime. However, they also point to a need for multiple season research in any analysis of potential consequences of global change on the long-term growth of trees.     

The effects of desiccation and root diffusates from potato and mustard on eggs of the white potato cyst nematode Globodera pallida

Following exposures to potato root diftusate of between 6 and 12 h, desiccation killed a proportion of juveniles in eggs of G. pallida and affected the hatching behaviour of survivors. In hatching tests of 9 wk duration, more juveniles hatched in the final wk from cysts, which were soahed and dried alternately for 9 wk than from cysts soaked in tap water for the final 2 wk only. Mustard root diffusate prevented eggs previously stimulated by potato root diffusate from hatching, but it did not alleviate the effects of desiccation.     

Isoprene emission from a subarctic peatland under enhanced UV-B radiation

Isoprene is a reactive hydrocarbon with an important role in atmospheric chemistry, and emissions from vegetation contribute to atmospheric carbon fluxes. The magnitude of isoprene emissions from arctic peatlands is not known, and it may be altered by increasing UV-B radiation. Isoprene emission was measured with the dynamic chamber method from a subarctic peatland under long-term enhancement of UV-B radiation targeted to correspond to a 20% loss in the stratospheric ozone layer. The site type of the peatland was a flark fen dominated by the moss Warnstorfia exannulata and sedges Eriophorum russeolum and Carex limosa. The relationship between species densities and the emission was also assessed. Isoprene emissions were significantly increased by enhanced UV-B radiation during the second (2004) and the fourth (2006) growing seasons under the UV-B exposure. Emissions were related to the density of E. russeolum. The dominant moss, W. exannulata, proved to emit small amounts of isoprene in a laboratory trial. Subarctic fens, even without Sphagnum moss, are a significant source of isoprene to the atmosphere, especially under periods of warm weather. Warming of the Arctic together with enhanced UV-B radiation may substantially increase the emissions.     

Review article. Ozone depletion and increased UV-B radiation: is there a real threat to photosynthesis?

This critical review of recent literature questions earlier predictions that photosynthetic productivity of higher plants is vulnerable to increased ultraviolet-B (UV-B) radiation as a result of stratospheric ozone (O3) depletion. Direct UV-B-induced inhibition of photosynthetic competence is observed only at high UV-B irradiances and primarily involves the loss of soluble Calvin cycle enzymes and adaxial stomatal closure in amphistomatous plants. However, even under these extreme UV-B exposures, acclimation (e.g. induction of UV-B absorbing flavonoids) can protect the photosynthetic processes. In plants irradiated with UV-B throughout development a reduction in productivity is usually associated with a reduced ability to intercept light (i.e. smaller leaf area) and not an inhibition of photosynthetic competence. Finally, a review of field experiments utilizing realistic UV-B enhancement is made to evaluate whether the mechanisms involved in UV-B-induced depressions of photosynthesis are likely to impact on the photosynthetic productivity of crops and natural vegetation in the future. Predictions of plant responses to O3 depletion are suspect from square-wave irradiance experiments due to the increased sensitivity of plants to UV-B at relatively low photosynthetically-active photon flux densities (PPFD) and ultraviolet-A (UV-A) irradiances. Realistic modulated UV-B irradiances in the field do not appear to have any significant effects on photosynthetic competence or light-interception. It is concluded that O3 depletion and the concurrent rise in UV-B irradiance is not a direct threat to photosynthetic productivity of crops and natural vegetation.Key words: Biomass, development, ozone depletion, photosynthesis, ultraviolet-B.      

The effects of ultraviolet-B radiation on plant competition in terrestrial ecosystems

Evidence regarding the interaction of ultraviolet-B (UV-B, 280-320 nm) radiation and plant competition in terrestrial ecosystems is examined. The competitive interactions of some species pairs were affected even by ambient solar UV-B radiation (as exists without ozone depletion), when compared to control pairs grown without UV-B. Also, the total shoot biomass of these species pairs was depressed under ambient UV-B. Relatively large increases in UV-B radiation (approximating a 40% ozone layer reduction when weighted with the generalized plant action spectrum) altered the competitive interactions of some species pairs grown in pots under field conditions, but did not affect the total shoot biomass production of those pairs. Recent field experiments have examined the competitive interactions of wheat ( Triticum aestivum L. cv. Bannock) and wild oat ( Avena fatua L.) under a simulated increased UV-B regime resulting from a 16% ozone layer reduction when weighted with the generalized plant action spectrum. This increase in UV-B altered the competitive interactions of these two species without affecting the total shoot biomass production of the species pair. The manner in which increased UV-B affected the relative competitive abilities of the two species was highly dependent upon the environmental conditions during the early life stages of the plants. The implications of these results for both agricultural and natural plant communities are discussed.     

Enhanced UV-B radiation alleviates the adverse effects of summer drought in two Mediterranean pines under field conditions

The effects of enhanced UV-B (290-320 nm) radiation on two native Mediterranean pines (Pinus pinea L., Pinus halepensis Mill.) were recorded during a one-year field study. Plants received ambient or ambient plus supplemental UV-B radiation (simulating a 15% stratospheric ozone depletion over Patras. Greece, 38.3°N. 29.1°E) and only natural precipitation, i.e. they were simultaneously exposed to other natural stresses. particularly water stress during summer. Supplemental UV-B irradiation started in early February, 1993 and up to late June, no effects were observed on growth and photochemical efficiency of photosystem II, as measured by chlorophy II fluorescence induction. Water stress during the summer was manifested in the control plants as a decline in the ratio of variable to maximum fluorescence (F_v/F_m), the apparent photon yield for oxygen evolution (φ_l) and the photosynthetic capacity at 5% CO₂ (P_m). In addition, a partial needle loss was evident. Under supplemental UV-B radiation, however, the decreases in F_v/F_m, φ_i, and P_m. as well as needle losses were significantly less. Soon after the first heavy autumn rains. photosynthetic parameters in both control and UV-B treated plants recovered to similar values. but the transient summer superiority of UV-B irradiated plants resulted in a significant increase in their dry weight measured at plant harvest. during late January. 1994. Plant height. UV-B absorbing compounds, photosynthetic pigments and relative water content measured at late spring. late summer and at plant harvest, were not significantly affected by supplemental UV-B radiation. The results indicate that enhanced UV-B radiation may be beneficial for Mediterranean pines through a partial alleviation of the adverse effects of summer drought.     

The effect of enhanced ultraviolet-B radiation on germination and seedling development of plant species occurring in a dune grassland ecosystem

The germination of seeds of seven plant species occurring in a dune grassland vegetation of the Netherlands, was studied at four levels of UV-B radiation simulating unto 45% stratospheric ozone reduction during April. With the exception of seeds of Senecio jacobaea, germination of the dune grassland species was not affected by enhanced UV-B irradiance. Although a clear UV-B fluence-response relationship was not observed, the germination rate of S. jacobaea seeds and maximal germination percentage were reduced at enhanced UV-B. Germination rate in the dark was higher than germination in the light for Oenothera biennis, Plantago lanceolata, Rumex obtusifolius and S. jacobaea. Total dry biomass accumulation of seedlings was not affected by increased UV-B radiation in any of the species tested. Clear-cut differences in UV-absorbance of methanolic extracts were observed between species. Enhanced UV-B irradiance stimulated UV-absorbance of seedling extracts of Holcus lanatus and Verbascum thapsus. A clear UV-B fluence-response relationship was observed for both species. The results indicate that germination of the studied plant species probably will not be adversely affected by the expected stratospheric ozone reduction in The Netherlands.     

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.     

Plant reaction to elevated ultraviolet irradiation: potential impacts of stratospheric ozone depletion

The ozone layer depletion evokes the increase of solar UV-B radiation intensity and corresponding reductions of growth (height, leaf area, fresh and dry weight), photosynthetic activity and flowering in higher plants. Competitive interactions also may be altered indirectly by differential growth responses. The UV-B-sensitivity of plants varies both among species and among cultivars of a given species. Photosynthetic activity may be reduced by direct effects on the photosynthetic process or metabolic pathways, or indirectly through effects on photosynthetic pigments or stomatal function. Plants may also respond by accumulating UV-absorbing compounds in their outer tissue layers, which presumably protect sensitive target from UV-damage. The key enzymes in the biosynthetic pathways of these compounds are specifically induced by UV-B irradiation via gene activation. The effects of UV-B radiation on plants can be modified by prevailing microclimatic conditions. Plants tend to be less sensitive to UV-B under drought or mineral deficiency, while sensitivity increases under low levels of visible light. Prognoses of agricultural yield reduction and economic loss for different scenarious of stratospheric ozone depletion are presented.     

Influence of UV-B Supplemental Radiation on Growth and Pigment Content in Suaeda Maritima L.

In a field experiment with a mangrove species Suaeda maritima L. grown under ambient and supplementary UV-B radiation corresponding to 20 % ozone depletion, changes in growth and contents of photosynthetic and UV-absorbing pigments were determined. Supplemental UV-B irradiation for 9 d significantly reduced the growth and concentration of photosynthetic pigments. However, anthocyanin and flavonoid contents were significantly increased in UV-treated plants and which could be reduce the UV-B penetration and damage to the underlying tissues.     

Differential effects of elevated ultraviolet-B radiation on plant species of a dune grassland ecosystem

In a greenhouse study, plants of three monocotyledonous and five dicotyledonous species, which occur in a Dutch dune grassland, were exposed to four levels of ultraviolet-B (UV-B) radiation. UV-B levels simulated up to 30% reduction of the stratospheric ozone column during summertime in The Netherlands. Six of the plant species studied in the greenhouse were also exposed to enhanced UV-B irradiance in an experimental field study. In the field experiment plants either received the ambient UV-B irradiance (control) or an enhanced UV-B level simulating 15–20% ozone depletion during summertime in The Netherlands. The purpose of both experiments was to determine the response of the plant species to UV-B radiation and to compare results obtained in the greenhouse with results of the field experiment. Large intraspecific differences in UV-B sensitivity were observed in the greenhouse study. Total dry matter accumulation of monocotyledons was increased, while dry matter accumulation of dicotyledons remained unaffected or decreased. The increase in biomass production of monocotyledons at elevated UV-B was not related to the rate of photosynthesis but to alterations in leaf orientation. In the greenhouse study, UV-B radiation also affected morphological characteristics. Shoot height or maximum leaf length of five out of eight species was reduced. In the field study only one species showed a significantly decreased maximum leaf length at enhanced UV-B. Possible reasons for this discrepancy are discussed. The absorbance of methanolic leaf extracts also differed between species. UV absorbance of field-grown plants was higher than greenhouse-grown plants. In the greenhouse study, the highest UV-B level increased UV-B absorbance of some species. In the field study however, this stimulation of UV absorbance was not observed. In general, results obtained in the greenhouse study were similar to results obtained in the field study. Difficulties in extrapolating results of UV-B experiments conducted in the greenhouse to the field situation are discussed.     

The impact of food quantity on UV-B tolerance and recovery from UV-B damage in Daphnia pulex

UV-B (290 nm) tolerance of Daphnia pulex, conditioned to four different food levels (Chlorophyta), was tested under standardized conditions with an artificial radiation source. Parameters measured were survival, percentage of egg bearing Daphnia and the number of juveniles produced after irradiation. UV-B tolerance of Daphnia pulex was found to be significantly improved with increasing food concentrations at all three parameters. The impact of the four different food concentrations on the photoreactivation system was tested with simultanous UV-B and white-light irradiation of Daphnia. Survival rate improved significantly with increasing food levels compared to solely UV-B irradiation. Photoreactivation had no effect on the reproductive parameters.     

A test of the effect of supplemental UV-B radiation on the common frog, Rana temporaria L., during embryonic development

The release of certain man-made chemicals has led to recurrent, seasonal destruction of ozone in the upper atmosphere, allowing more solar radiation in the UV-B waveband to reach the Earth. Consequently, many amphibians may suffer increased exposure to UV-B at various stages in their lives. Embryonic stages of species which spawn in the spring, in shallow, open water, are at high risk of increased exposure. We exposed newly fertilized eggs of one such species, Rana temporaria L., to sunlight with and without supplemental UV-B. We used outdoor arrays of lamps to simulate the increase in UV-B which might result from previously documented ozone depletion. From immediately after fertilization to when hatchlings began feeding, ambient solar UV-B, weighted for DNA-damaging potential, was supplemented by ≈ 81% in 1995 and 113% in 1996. These levels of supplementation approximated the increase in solar UV-B expected to result from losses of 21% and 25%, respectively, of the total amount of ozone in the atmospheric column, relative to pre-ozone-depletion values. We found no evidence that these additions of UV-B radiation increased the incidence of mortality or overt developmental abnormality among embryos. We stress the need for appropriate dosimetry in studies of effects of UV-B on organisms.     

Boreal peatland ecosystems under enhanced UV-B radiation and elevated tropospheric ozone concentration

Boreal peat-forming wetlands, mires, are globally important sources of methane and sinks for CO₂. As peatland vegetation plays a significant role regulating the exchange of these greenhouse gases, we have assessed the responses of the dominant plants and ecosystem functions to increasing tropospheric ozone concentration and enhanced ultraviolet-B (UV-B) radiation in long-term experiments, the results of which are summarized in this review. The dominant sedge, Eriophorum vaginatum, and especially the Sphagnum mosses common on peatlands, appear fairly tolerant to the future predicted ozone levels. Similarly, UV-B radiation only caused few alterations in the carbohydrates and pigments of the dominant sedge, Eriophorum russeolum, and had no effects on the dominant moss species of the experimental site, Warnstorfia exannulata. Surprisingly, there were alterations in organic acid concentrations in the peat pore water and peat microbial community composition in both experiments. Elevated ozone caused a transient decrease in ecosystem-level gross photosynthesis and methane (CH₄) emission, which shifted to a slight increase later on. Enhanced UV-B decreased dark ecosystem respiration and increased CH₄ emission in the course of the six measurement years. The emission of isoprene was increased by both ozone and UV-B during warm weather periods, suggesting interactive effects with temperature. All in all, we suggest that ozone and UV-B have limited effects on the carbon cycle in boreal peatlands, because other environmental factors, such as temperature, water level and photosynthetically active radiation more strongly regulate CO₂ and CH₄ exchange rates.     

Influence of UV-B radiation on bacterial activity in coastal waters

The impact of UV-B radiation (290–315 nm) on bacterialactivity and abundance in coastal water was studied in mesocosmexperiments in May 1994 and May 1995 at Kristineberg MarineResearch Station, Sweden. Mesocosms (6 m³) containing naturalpelagic communities were exposed either to ambient irradiation(AMB), ambient irradiation with enhanced UV-B (+UV) (0.7 W m^–24 h every day around noon), or ambient irradiation screenedfor UV-B (–UV). Bacterial activity in the mesocosms wasmeasured by means of thymidine incorporation in short-term testsduring incubations at ambient irradiation, at ambient irradiationwith enhanced UV-B, and at ambient irradiation screened forUV-B. In +UV mesocosms, bacterial activity was significantlystimulated when incubated at ambient radiation. The stimulatingeffect was suggested to be due to an increase in carbon or nutrientsupply through a photodegradation of recalcitrant dissolvedorganic material (DOM). Low attenuation coefficients for UV-Band PAR (400–700 nm) in the +UV mesocosms supported thishypothesis. The bacterial activity in +UV mesocosms, however,was inhibited when incubations were made at enhanced UV-B irradiation,implying that the bacteria had become more sensitive to UV-Bradiation. The increased sensitivity to UV-B exposure in bacterialassemblages that already had been exposed and stressed by UV-Bradiation is suggested to be due to an overburdening of theenergy-consuming DNA repair mechanism. The data suggest thatincreased UV-B radiation, which might occur with ozone depletion,may both stimulate and suppress bacterial activity in coastalwaters, implying that the net outcome of enhanced UV-B radiationcould be an unchanged bacterial activity.     

Effects of ultraviolet-B irradiation on the cuticular wax of cucumber (<Emphasis Type="Italic">Cucumis sativus</Emphasis>) cotyledons

Cucumber seedlings were grown under three doses of supplemental ultraviolet-B (UV-B) irradiation to examine the effects on the surface structure of the cotyledons. Medium and high doses of irradiation induced glazing (formation of translucent, glossy layers) on the adaxial surfaces of cotyledons, especially those exposed to a high dose of UV-B. Observation with a scanning electron microscope revealed that the adaxial surfaces of cotyledons exposed to a medium dose of UV-B and controls became rough in appearance, but unevenness of the surface was not apparent in cotyledons irradiated with a high dose of UV-B. UV-B irradiation affected the types and amounts of alkanes and primary alcohols, the main components of cucumber cuticular wax. Based on cotyledon area, the amounts of these components were significantly higher in cotyledons irradiated with a medium dose of UV-B than in controls. This effect could be a consequence of small cotyledon area and constant wax production in the cotyledons irradiated with a medium dose of UV-B. The distribution patterns of homologs within the alkane and primary alcohol fractions shifted during growth to longer alkyl chain length in the control cotyledons. UV-B irradiation repressed these changes, suggesting that UV-B acts on cuticular wax biosynthetic pathways.