The effects of elevated ozone and CO2 on growth and defense of native,exotic and invader trees

Aims Elevated ozone and CO₂ can differentially affect the performance of plant species. Variation among native, exotic and invader species in their growth and defense responses to CO₂ and ozone may shape CO₂ and ozone effects on invasions, perhaps in part also due to variation between native and invasive populations of invaders.Methods We manipulated ozone (control or 100 ppb) and CO₂ (ambient or 800 ppm) in a factorial greenhouse experiment in replicated chambers. We investigated growth and defense (tannins) of seedlings of Triadica sebifera from invasive (USA) and native (China) populations and pairs of US and China tree species within three genera (Celtis, Liquidambar and Platanus).Important findings Overall, ozone reduced growth in ambient CO₂ but elevated CO₂ limited this effect. T. sebifera plants from invasive populations had higher growth than those from native populations in control conditions or the combination of elevated CO₂ and ozone in which invasive populations had greater increases in growth. Their performances were similar in elevated CO₂ because native populations were more responsive and their performances were similar with elevated ozone because invasive populations were more susceptible. Compared to other species, T. sebifera had high growth rates but low levels of tannin production that were insensitive to variation in CO₂ or ozone. Both China and US Platanus plants reduced tannins with increased CO₂ and/or ozone and US Liquidambar plants increased tannins with the combination of elevated CO₂ and ozone. The growth results suggest that intraspecific variation in T. sebifera will reduce the effects of CO₂ or ozone alone on invasions but increase their combined effects. The tannin results suggest that defense responses to CO₂ and ozone will be variable across native and exotic species. The effects of CO₂ and ozone on growth and defense of native and exotic species indicate that the benefit or harm to species from these global change drivers is an idiosyncratic combination of species origin and genus.     

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.     

Effects of enhanced UV-B radiation and nitrogen deposition on the growth of invasive plant Triadica sebifera

Aims Exotic plant invasions are important components of global change, threatening both the stability and function of invaded ecosystems. Shifts in competitive ability of invasive plants versus their native congeners have been documented. Enhanced UV-B radiation and nitrogen (N) deposition might interact with soil biota communities impacting the invasion process of exotic plant species. To understand the potential effects by UV-B and N with soil biota on plant growth would enhance our understanding of the mechanisms in plant invasions in the context of global change.
Methods We conducted a full-factorial pot experiment in the native range (China) of Triadica sebifera invading US to investigate how UV-B radiation, N and soil biota together determined their seedling growth.
Important findings The results showed that UV-B radiation, N and soil sterilization together impacted the growth of T. sebifera seedlings. UV-B radiation induced changes in biomass allocation with larger leaf biomass observed in response to UV-B radiation. In addition, N increased aboveground biomass and decreased root biomass simultaneously. Soil biota imposed positive effects on growth of T. sebifera, and the addition of N amplified these positive effects. The negative effects by UV-B radiation on growth of T. sebifera showed no response to N addition. Plant height, leaf biomass and total biomass of the invasive T. sebifera populations out- performed those of the native ones. In addition, invasive T. sebifera populations weakened the dependence of root/shoot ratio and root biomass on local soil microorganisms than native populations, but enhanced that of leaf area ratio.     

Principal component analysis of intraspecific responses of tartary buckwheat to UV-B radiation under field conditions

Fifteen populations of tartary buckwheat (Fagopyrum tataricum Gaertn.) occurring in habitats with different natural UV-B levels were sampled, and the plants were exposed to enhanced UV-B radiation under field conditions simulating 25% depletion of the stratospheric ozone layer. The experimental design was a 2 × 15 factorial, with two levels of UV-B radiation (ambient and enhanced UV-B radiation) and plants from 15 populations. The responses of plants in growth, morphology, productivity and in the composition of photosynthetic pigments were measured. The results demonstrated that there were significant differences among populations in responses to UV-B radiation: some populations exhibited a positive effect while others were negatively affected. The UV-B effects on plant traits were correlated with the constitutive values. A principal component analysis (PCA) was used to evaluate the overall sensitivity of responses to UV-B radiation. Our results suggest that the sensitivity of plants to UV-B radiation is not only associated with the ambient UV-B level in natural habitats but also with the relative growth rate and other factors.     

Evolutionary increases in defense during a biological invasion

Invasive plants generally escape from specialist herbivores of their native ranges but may experience serious damage from generalists. As a result, invasive plants may evolve increased resistance to generalists and tolerance to damage. To test these hypotheses, we carried out a common garden experiment comparing 15 invasive populations with 13 native populations of Chromolaena odorata, including putative source populations identified with molecular methods and binary choice feeding experiments using three generalist herbivores. Plants from invasive populations of C. odorata had both higher resistance to three generalists and higher tolerance to simulated herbivory (shoot removal) than plants from native populations. The higher resistance of plants from invasive populations was associated with higher leaf C content and densities of leaf trichomes and glandular scales, and lower leaf N and water contents. Growth costs were detected for tolerance but not for resistance, and plants from invasive populations of C. odorata showed lower growth costs of tolerance. Our results suggest that invasive plants may evolve to increase both resistance to generalists and tolerance to damage in introduced ranges, especially when the defense traits have low or no fitness costs. Greater defenses in invasive populations may facilitate invasion by C. odorata by reducing generalist impacts and increasing compensatory growth after damage has occurred.     

入侵植物的繁殖策略以及对本土植物繁殖的影响

理解入侵生物的繁殖策略是阐明生物入侵机制的一个重要方面。入侵植物常表现出一些共同的繁殖特征, 如以两性花为主的性系统、自动自交为主的繁育系统或不依赖传粉媒介的无融合生殖和无性繁殖以及高生殖投资的资源配置策略等。成功入侵的外来植物通过影响本土的传粉者, 在种群和群落水平上影响本土植物的有性繁殖, 甚至促使某些本土植物在繁殖对策和表型性状上发生快速转变。目前, 入侵植物繁殖策略及其生态效应的研究多侧重于入侵种的快速演化, 而有关外来植物与本土植物间的相互影响及其可能存在的协同适应研究还较为缺乏。探讨本土植物在外来种入侵压力下的繁殖对策和响应机制, 将丰富人们对物种间竞争、共存及群落构建等机制的深入了解。从繁殖和适应的角度探求入侵植物与本土植物之间的复杂关系, 将有助于解析生物入侵的机制及人类干扰下的物种演化规律, 也为预测和防控入侵植物提供科学依据。     

Adaptive rather than non-adaptive evolution of Mimulus guttatus in its invasive range

Adaptive and non-adaptive evolutionary processes are likely to play important roles in biological invasions but their relative importance has hardly ever been quantified. Moreover, although genetic differences between populations in their native versus invasive ranges may simply reflect different positions along a genetic latitudinal cline, this has rarely been controlled for. To study non-adaptive evolutionary processes in invasion of Mimulus guttatus, we used allozyme analyses on offspring of seven native populations from western North America, and three and four invasive populations from Scotland and New Zealand, respectively. To study quantitative genetic differentiation, we grew 2474 plants representing 17 native populations and the seven invasive populations in a common greenhouse environment under temporarily and permanently wet soil conditions. The absence of allozyme differentiation between the invasive and native range indicates that multiple genotypes had been introduced to Scotland and New Zealand, and suggests that founder effects and genetic drift played small, if any, roles in shaping genetic structure of invasive M. guttatus populations. Plants from the invasive and native range did not differ in phenology, floral traits and sexual and vegetative reproduction, and also not in plastic responses to the watering treatments. However, plants from the invasive range produced twice as many flower-bearing upright side branches than the ones from the native populations. Further, with increasing latitude of collection, vegetative reproduction of our experimental plants increased while sexual reproduction decreased. Plants from the invasive and native range shared these latitudinal clines. Because allozymes showed that the relatedness between native and invasive populations did not depend on latitude, this suggests that plants in the invasive regions have adapted to the local latitude. Overall, our study indicates that quantitative genetic variation of M. guttatus in its two invasive regions is shaped by adaptive evolutionary processes rather than by non-adaptive ones.     

Competitive interactions between native and invasive exotic plant species are altered under elevated carbon dioxide

We hypothesized that the greater competitive ability of invasive exotic plants relative to native plants would increase under elevated CO₂ because they typically have traits that confer the ability for fast growth when resources are not limiting and thus are likely to be more responsive to elevated CO₂. A series of competition experiments under ambient and elevated CO₂ glasshouse conditions were conducted to determine an index of relative competition intensity for 14 native-invasive exotic species-pairs. Traits including specific leaf area, leaf mass ratio, leaf area ratio, relative growth rate, net assimilation rate and root weight ratio were measured. Competitive rankings within species-pairs were not affected by CO₂ concentration: invasive exotic species were more competitive in 9 of the 14 species-pairs and native species were more competitive in the remaining 5 species-pairs, regardless of CO₂ concentration. However, there was a significant interaction between plant type and CO₂ treatment due to reduced competitive response of native species under elevated compared with ambient CO₂ conditions. Native species had significantly lower specific leaf area and leaf area ratio under elevated compared with ambient CO₂. We also compared traits of more-competitive with less-competitive species, regardless of plant type, under both CO₂ treatments. More-competitive species had smaller leaf weight ratio and leaf area ratio, and larger relative growth rate and net assimilation rate under both ambient and elevated CO₂ conditions. These results suggest that growth and allocation traits can be useful predictors of the outcome of competitive interactions under both ambient and elevated CO₂ conditions. Under predicted future atmospheric CO₂ conditions, competitive rankings among species may not change substantially, but the relative success of invasive exotic species may be increased. Thus, under future atmospheric CO₂ conditions, the ecological and economic impact of some invasive exotic plants may be even greater than under current conditions.     

Introduced Brassica nigra populations exhibit greater growth and herbivore resistance but less tolerance than native populations in the native range

Rapid post-introduction evolution has been found in many invasive plant species, and includes changes in defence (resistance and tolerance) and competitive ability traits. Here, we explored the post-introduction evolution of a trade-off between resistance to and tolerance of herbivory, which has received little attention. In a common garden experiment in a native range, nine invasive and 16 native populations of Brassica nigra were compared for growth and defence traits. Invasive populations had higher resistance to, but lower tolerance of, herbivore damage than native populations. Invasive populations survived better and produced more seeds than native ones when released from herbivores; but fitness was equivalent between the regions under ambient herbivory. The invasive populations grew taller, and produced more biomass and lighter seeds than natives, irrespective of insecticide treatment. In addition to supporting the idea of post-introduction rapid evolution of plant traits, our results also contribute to an emerging pattern of both increasing resistance and growth in invasive populations, contrary to the predictions of earlier theories of resistance-growth trade-offs.     

The role of interactions between trophic levels in determining the effects of UV-B on terrestrial ecosystems

Understanding the potential impact of ozone depletion on terrestrial ecosystems is constrained by lack of information on the effects of environmentally realistic UV-B doses on terrestrial organisms other than higher plants. Increasing UV-B may alter interactions between plants and consumers through direct effects on consumer organisms (herbivores, phytopathogens, decomposers, etc.). The effects of increasing UV-B on arthropods are not known. Significant UV-B effects on fungi have been reported, and may be either negative (inhibition of spore germination and mycelial growth) or positive (increased growth, induction of reproductive development and sporulation). However, in many cases consumers are unlikely to be directly exposed to UV-B in the field. In addition, UV action spectra for fungi suggest that this major group may be less sensitive to the effects of ozone depletion than higher plants. Host mediated effects of UV-B on consumers may include alterations in plant chemistry. While secondary metabolites such as phenolics may increase under increased UV-B, this is not invariably the case and evidence that such changes have significant effects on consumers is limited. In particular, there is no evidence that increased UV-B increases resistance of higher plants to fungal pathogens. Indeed, increased UV-B prior to inoculation results in no significant effect or increased disease. Such responses may be attributable to UV-B effects on host surface properties or on compounds other than phenolics. However, such changes are poorly known, and their potential effects on phytopathogens, herbivores or decomposers cannot be assessed. Understanding the effects of UV-B on terrestrial ecosystems is further limited since virtually nothing is known of possible impacts on higher trophic levels, i.e. predators, parasites or pathogens.     

Phenotypes of Two Generations of Sporobolus airoides Seedlings Derived from Acroptilon repens‐invaded and Non‐invaded Grass Populations

Although the ecological impacts of invasive species are well known, the evolutionary impacts on recipient native grass communities are not. We suggest that remnant native plants may provide desirable seed sources for restoration and native plant production. Native populations exposed to the selective pressures associated with exotic invasion may retain traits that increase their ability to coexist with invasive species. Two generations of Sporobolus airoides Torr. (Alkali sacaton) plants derived from lineages collected from within long‐term invaded areas of Acroptilon repens (L.) DC (Russian knapweed) and from adjacent non‐invaded areas were propagated in a greenhouse to evaluate generational changes in phenotypic traits from the production environment. Given the difference in invasion history of the two populations, we hypothesized that invaded and non‐invaded subpopulations would differ phenotypically. Phenotypic measurements revealed that invaded subpopulations had greater vegetative growth, whereas non‐invaded subpopulations had increased sexual reproduction. Phenotypic expression changed from the first to the second generation, predominantly in the invaded subpopulation. Generational phenotypic shifts are disadvantageous for native seed production which requires a standard product to sell commercially. However, phenotypic variation may improve field seed survival. This research demonstrates the potential value of targeting post‐invasion remnant grass populations for restoration.     

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.     

Effect of enhanced UV-B radiation on pollen quantity,quality, and seed yield in Brassica rapa (Brassicaceae)

We conducted three experiments to examine the influence of ultraviolet-B radiation (UV-B; 280–320 nm) exposure on reproduction in Brassica rapa (Brassicaceae). Plants were grown in a greenhouse under three biologically effective UV-B levels that simulated either an ambient stratospheric ozone level (control), 16% (“low enhanced”), or 32% (“high enhanced”) ozone depletion levels at Morgantown, WV, USA in mid-March. In the first experiment, we examined whether UV-B level during plant growth influenced in vivo pollen production and viability, and flower production. Pollen production and viability per flower were reduced by ≈50% under both enhanced UV-B levels relative to ambient controls. While plants under high-enhanced UV-B produced over 40% more flowers than plants under the two lower UV-B treatments, whole-plant production of viable pollen was reduced under high-enhanced UV-B to 17% of that of ambient controls. Whole-plant production of viable pollen was reduced under low-enhanced UV-B to 34% of ambient controls. In the second experiment, we collected pollen from plants under the three UV-B levels and examined whether source-plant UV-B exposure influenced in vitro pollen germination and viability. Pollen from plants under both enhanced-UV-B treatments had initially lower germination and viability than pollen from the ambient level. After in vitro exposure to the high-enhanced UV-B levels for 6 h, viability of the pollen from plants grown under ambient UV-B was reduced from 65 to 18%. In contrast, viability of the pollen from plants grown under both enhanced UV-B treatments was reduced to a much lesser extent: only from ≈43 to 22%. Thus, ambient source-plant pollen was more sensitive to enhanced UV-B exposure. In the third experiment, we used pollen collected from source plants under the three UV-B levels to fertilize plants growing under ambient-UV-B levels, and assessed subsequent seed production and germination. Seed abortion rates were higher in plants pollinated with pollen from the enhanced UV-B treatments, than from ambient UV-B. Despite this, seed yield (number and mass) per plant was similar, regardless of the UV-B exposure of their pollen source. Our findings demonstrate that enhanced UV-B levels associated with springtime ozone depletion events have the capacity to substantially reduce viable pollen production, and could ultimately reduce reproductive success of B. rapa.     

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.     

Impact of drought on plant populations of native and invasive origins

Invasive populations often shift phenotypically during introduction. Moreover, they are postulated to show an increased phenotypic plasticity compared with their native counterparts, which could be advantageous. However, less is known about trait selection across populations along the invasion gradient in response to environmental factors, such as increasing drought caused by climate change. In this study, we investigated the impacts of drought on growth, regrowth, and various leaf traits in plants of different origin. Therefore, seeds of 18 populations of the perennial Tanacetum vulgare were collected along the invasion gradient (North America, invasive; West Europe, archaeophyte; East Europe, native) and grown in competition with the grass Poa pratensis under control or dry conditions in a common garden. Above-ground biomass was cut once and the regrowth was measured as an indicator for tolerance over a second growth period. Initially, drought had little effects on growth of T. vulgare, but after cutting, plants grew more vigorously. Against expectations, phenotypic plasticity was not higher in invasive populations, but even reduced in one trait, which may be attributable to ecological constraints imposed by multiple stress conditions. Trait responses reflected the range expansion and invasion gradient and were influenced by the latitudinal origin of populations. Populations of invaded ranges may be subject to faster and more extensive genetic mixing or had less time to undergo and reflect selective processes.     

Elevated ozone reduces methane emissions from peatland mesocosms

Although the effects of elevated ozone on aboveground carbon (C) assimilation are well understood, its effects on soil C fluxes are less certain. Mesocosms taken from a lowland raised bog in northern England were exposed in open‐top chambers for 2 years to ambient air or ambient air plus ozone elevated for 8 h day^?1 by an average of 49 ppb in summer and 10 ppb in winter. The effects of elevated ozone on methane emission and ecosystem dark respiration were measured throughout this period, along with soil and plant variables. Methane emissions were significantly reduced, by about 25%, by elevated ozone during midsummer periods of both years, but no significant effect of ozone was found during the winter periods. Dark ecosystem respiration was not significantly affected by elevated ozone. There was no evidence that effects of elevated ozone on methane emissions were mediated through changes in aboveground plant biomass or soil water dissolved organic C concentrations. Our results imply that the increased northern hemisphere background ozone concentrations over the 21st century that are predicted by most models may reduce the rate of increase in methane emissions as the region warms.     

Ozone suppresses soil drying- and abscisic acid (ABA)-induced stomatal closure via an ethylene-dependent mechanism

Elevated atmospheric ozone concentrations (70 ppb) reduced the sensitivity of stomatal closure to abscisic acid (ABA) in Leontodon hispidus after at least 24 h exposure (1) when detached leaves were fed ABA, and (2) when intact plants were sprayed or injected with ABA. They also reduced the sensitivity of stomatal closure to soil drying around the roots. Such effects could already be occurring under current northern hemisphere peak ambient ozone concentrations. Leaves detached from plants which had been exposed to elevated ozone concentrations generated higher concentrations of ethylene, although leaf tissue ABA concentrations were unaffected. When intact plants were pretreated with the ethylene receptor binding antagonist 1-methylcyclopropene, the stomatal response to both applied ABA and soil drying was fully restored in the presence of elevated ozone. Implications of ethylene's antagonism of the stomatal response to ABA under oxidative stress are discussed. We suggest that this may be one mechanism whereby elevated ozone induces visible injury in sensitive species. We emphasize that drought linked to climate change and tropospheric ozone pollution, are both escalating problems. Ozone will exacerbate the deleterious effects of drought on the many plant species including valuable crops that respond to this pollutant by emitting more ethylene.     

Response of purely symbiotic and NO3-fed nodulated plants of Lupinus luteus and Vicia atropurpurea to ultraviolet-B radiation

The effects of elevated UV-B radiation on growth, symbiotic function and concentration of metabolites were assessed in purely symbiotic and NO3-fed nodulated plants of Lupinus luteus and Vicia atropurpurea grown outdoors either on tables under supplemental UV-B radiation or in chambers covered with different types of plexi-glass to attenuate solar ultraviolet radiation. Moderately and highly elevated UV-B exposures simulating 15% and 25% ozone depletion as well as sub- ambient UV-B did not alter organ growth, plant total dry matter and N content per plant in both L. luteus and V. atropurpurea. In contrast, elevated UV-B increased (P <0.05) flavonoid and anthocyanin concentrations in roots and leaves of L. luteus, but not of V. atropurpurea. Feeding nodulated plants of L. luteus under elevated UV-B radiation with 2 mM NO3 increased (P <0.05) nodule, leaf and total dry matter, and whole plant N content. With V. atropurpurea, NO3 reduced (P <0.05) nodule activity, root %N and concentrations of flavonoids, anthocyanins in roots and leaves and soluble sugars in roots, in contrast to an observed increase (P <0.05) in nodule dry matter per plant. Similarly, supplying 2 mM NO3 to L. luteus plants exposed to sub-ambient UV-B radiation significantly reduced individual organ growth, plant total biomass, nodule dry matter, nodule %N, and whole plant N content, as well as root concentrations of flavonoids, anthocyanins, soluble sugars, and starch of L. luteus, but not V. atropurpurea plants. These results show no adverse effect of elevated UV-B radiation on growth and symbiotic function of L. luteus and V. atropurpurea plants. However, NO3 supply promoted growth in L. luteus plants exposed to the highly elevated UV-B radiation.     

Interactions between willows and insect herbivores under enhanced ultraviolet-B radiation

We studied the effects of elevated ultraviolet-B radiation on interactions between insect herbivores and their host plants by exposing two species of phytochemically different willows, Salix myrsinifolia and S. phylicifolia, to a modulated increase in ultraviolet radiation in an outdoor experiment and monitoring the colonisation of insect herbivores on these willows. We examined the effect of increased ultraviolet-B (UV-B) radiation on (1) the quality of willow leaves, (2) the distribution and abundance of insect herbivores feeding on these willows, (3) the resulting amount of damage, and (4) the performance of insect larvae feeding on the exposed plant tissue. Six clones of each of the two willow species were grown in eight blocks for 12 weeks in the UV-B irradiation field. The clones were exposed to a constant 50% increase in UV-B radiation (simulating 20-25% ozone depletion), to a small increase in UV-A radiation or to ambient solar irradiation. We allowed colonisation on the willows by naturally occurring insects, but also introduced adults of a leaf beetle, Phratora vitellinae, a specialist herbivore on S. myrsinifolia. Increased UV-B radiation did not affect any of the measured indices of plant quality. However, numbers of P. vitellinae on S. myrsinifolia were higher in plants with UV-B treatment compared with UV-A and shade controls. In laboratory tests, growth of the second-instar larva of P. vitellinae was not affected by UV-B treatment of S. myrsinifolia, but was retarded on UV-B treated leaves of S. phylicifolia. In addition, naturally occurring insect herbivores were more abundant on willows exposed to elevated UV-B radiation compared to those grown under control treatments. In spite of the increased abundance of insect herbivores, willows treated with elevated UV-B did not suffer more herbivore damage than willows exposed to ambient solar radiation (shade control). The observed effects of UV-B on herbivore abundance, feeding and growth varied significantly due to spatial variation in environment quality, as indicated by the UV-treatment x block interaction. The results suggest that (1) environmental variation modifies the effects of UV-B radiation on plant-insect interactions and (2) specialist herbivores might be more sensitive to chemical changes in their secondary host plants (S. phylicifolia) than to changes in their primary hosts (S. myrsinifolia).