Comparative Transcriptomics of Multi-Stress Responses in Pachycladon cheesemanii and Arabidopsis thaliana.

The environment is seldom optimal for plant growth and changes in abiotic and biotic signals, including temperature, water availability, radiation and pests, induce plant responses to optimise survival. The New Zealand native plant species and close relative to Arabidopsis thaliana, Pachycladon cheesemanii, grows under environmental conditions that are unsustainable for many plant species. Here, we compare the responses of both species to different stressors (low temperature, salt and UV-B radiation) to help understand how P. cheesemanii can grow in such harsh environments. The stress transcriptomes were determined and comparative transcriptome and network analyses discovered similar and unique responses within species, and between the two plant species. A number of widely studied plant stress processes were highly conserved in A. thaliana and P. cheesemanii. However, in response to cold stress, Gene Ontology terms related to glycosinolate metabolism were only enriched in P. cheesemanii. Salt stress was associated with alteration of the cuticle and proline biosynthesis in A. thaliana and P. cheesemanii, respectively. Anthocyanin production may be a more important strategy to contribute to the UV-B radiation tolerance in P. cheesemanii. These results allowed us to define broad stress response pathways in A. thaliana and P. cheesemanii and suggested that regulation of glycosinolate, proline and anthocyanin metabolism are strategies that help mitigate environmental stress.

UV-B Induced Flavonoids Contribute to Reduced Biotrophic Disease Susceptibility in Lettuce Seedlings.

Biotrophic disease is one of the largest causes of decreased yield in agriculture. While exposure to ultraviolet B (UV-B) light (280-320 nm) has been previously observed to reduce plant susceptibility to disease, there is still a paucity of information regarding underlying biological mechanisms. In addition, recent advances in UV-LED technology raise the prospect of UV light treatments in agriculture which are practical and efficient. Here, we characterized the capability of UV-B LED pre-treatments to reduce susceptibility of a range of lettuce (Lactuca sativa) cultivars to downy mildew disease caused by the obligate biotroph Bremia lactucae. Innate cultivar susceptibility level did not seem to influence the benefit of a UV-B induced disease reduction with similar reductions as a percentage of the control observed (54-62% decrease in conidia count) across all susceptible cultivars. UV-B-induced reductions to conidia counts were sufficient to significantly reduce the infectivity of the diseased plant. Secondary infections caused by UV-B pre-treated plants exhibited yet further (67%) reduced disease severity. UV-B-induced flavonoids may in part mediate this reduced disease severity phenotype, as B. lactucae conidia counts of lettuce plants negatively correlated with flavonoid levels in a UV-B-dependent manner (r = -0.81). Liquid chromatography-mass spectrometry (LC-MS) was used to identify metabolic features which contribute to this correlation and, of these, quercetin 3-O-(6"-O-malonyl)-b-D-glucoside had the strongest negative correlation with B. lactucae conidia count (r = -0.68). When quercetin 3-O-(6"-O-malonyl)-b-D-glucoside was directly infiltrated into lettuce leaves, with those leaves subsequently infected, the B. lactucae conidia count was reduced (25-39%) in two susceptible lettuce cultivars. We conclude that UV-B induced phenolics, in particular quercetin flavonoids, may act as phytoanticipins to limit the establishment of biotrophic pathogens thus delaying or reducing their sporulation as measured by conidia count. These findings highlight the opportunity for UV-B morphogenesis to be exploited through the application of UV-LED technology, as part of the development of next-generation, sustainable disease control tools.

Herbivory and Attenuated UV Radiation Affect Volatile Emissions of the Invasive Weed Calluna vulgaris.

Calluna vulgaris (heather) is an aggressive invasive weed on the Central Plateau, North Is., New Zealand (NZ), where it encounters different environmental factors compared to its native range in Europe, such as high ultraviolet radiation (UV) and a lack of specialist herbivores. The specialist herbivore Lochmaea suturalis (heather beetle) was introduced from the United Kingdom (UK) in 1996 as a biocontrol agent to manage this invasive weed. Like other plant invaders, a novel environment may be challenging for heather as it adjusts to its new conditions. This process of "adjustment" involves morphological and physiological changes often linked to phenotypic plasticity. The biochemical responses of exotic plants to environmental variables in their invaded range is poorly understood. The production and release of volatile organic compounds (VOCs) is essential to plant communication and highly susceptible to environmental change. This study therefore aimed to explore the VOC emissions of heather in response to different levels of UV exposure, and to feeding damage by L. suturalis. Using tunnel houses clad with UV-selective filters, we measured VOCs produced by heather under NZ ambient, 20% attenuated, and 95% attenuated solar UV treatments. We also compared VOC emissions in the field at adjacent sites where L. suturalis was present or absent. Volatiles produced by the same target heather plants were measured at four different times in the spring and summer of 2018-2019, reflecting variations in beetle's abundance, feeding stage and plant phenology. Heather plants under 95% attenuated UV produced significantly higher amounts of (E)-ß-farnesene, decanal, benzaldehyde, and benzeneacetaldehyde compared to 25% attenuated and ambient UV radiation. We also found significant differences in volatiles produced by heather plants in beetle-present versus beetle-absent sites on most sampling occasions. We also recorded a lower number of generalist herbivores on heather at sites where L. suturalis was present. Interactions between invasive plants, a novel environment, and the native communities they invade, are discussed.

Genome draft of the Arabidopsis relative Pachycladon cheesemanii reveals novel strategies to tolerate New Zealand's high ultraviolet B radiation environment.

BACKGROUND: Pachycladon cheesemanii is a close relative of Arabidopsis thaliana and is an allotetraploid perennial herb which is widespread in the South Island of New Zealand. It grows at altitudes of up to 1000 m where it is subject to relatively high levels of ultraviolet (UV)-B radiation. To gain first insights into how Pachycladon copes with UV-B stress, we sequenced its genome and compared the UV-B tolerance of two Pachycladon accessions with those of two A. thaliana accessions from different altitudes. RESULTS: A high-quality draft genome of P. cheesemanii was assembled with a high percentage of conserved single-copy plant orthologs. Synteny analysis with genomes from other species of the Brassicaceae family found a close phylogenetic relationship of P. cheesemanii with Boechera stricta from Brassicaceae lineage I. While UV-B radiation caused a greater growth reduction in the A. thaliana accessions than in the P. cheesemanii accessions, growth was not reduced in one P. cheesemanii accession. The homologues of A. thaliana UV-B radiation response genes were duplicated in P. cheesemanii, and an expression analysis of those genes indicated that the tolerance mechanism in P. cheesemanii appears to differ from that in A. thaliana. CONCLUSION: Although the P. cheesemanii genome shows close similarity with that of A. thaliana, it appears to have evolved novel strategies allowing the plant to tolerate relatively high UV-B radiation.

Solar UV light regulates flavonoid metabolism in apple (Malus x domestica).

Ultraviolet-B light (UV-B) is one environmental signal perceived by plants that affects the flavonoid pathway and influences the levels of anthocyanins, flavonols, and proanthocyanidins. To understand the mechanisms underlying UV exposure, apple trees were grown under spectral filters that altered transmission of solar UV light. Fruit analysis showed that UV induced changes in physiology, metabolism, and gene expression levels during development over a season. These changes were sustained after storage. Under low UV, ripening was delayed, fruit size decreased, and anthocyanin and flavonols were reduced. Expression analysis showed changes in response to UV light levels for genes in the regulation and biosynthesis of anthocyanin and flavonols. Transcription of flavonol synthase (FLS), ELONGATED HYPOCOTYL 5 (HY5), MYB10, and MYB22 were down-regulated throughout fruit development under reduced UV. Functional testing showed that the FLS promoter was activated by HY5, and this response was enhanced by the presence of MYB22. The MYB22 promoter can also be activated by the anthocyanin regulator, MYB10. As ambient levels of UV light vary around the globe, this study has implications for future crop production, the quality of which can be determined by the response to UV.

Fungal endophyte infection of ryegrass reprograms host metabolism and alters development.

Beneficial associations between plants and microbes play an important role in both natural and agricultural ecosystems. For example, associations between fungi of the genus Epichloë, and cool-season grasses are known for their ability to increase resistance to insect pests, fungal pathogens and drought. However, little is known about the molecular changes induced by endophyte infection. To study the impact of endophyte infection, we compared the expression profiles, based on RNA sequencing, of perennial ryegrass infected with Epichloë festucae with noninfected plants. We show that infection causes dramatic changes in the expression of over one third of host genes. This is in stark contrast to mycorrhizal associations, where substantially fewer changes in host gene expression are observed, and is more similar to pathogenic interactions. We reveal that endophyte infection triggers reprogramming of host metabolism, favouring secondary metabolism at a cost to primary metabolism. Infection also induces changes in host development, particularly trichome formation and cell wall biogenesis. Importantly, this work sheds light on the mechanisms underlying enhanced resistance to drought and super-infection by fungal pathogens provided by fungal endophyte infection. Finally, our study reveals that not all beneficial plant-microbe associations behave the same in terms of their effects on the host.

Rediscovering leaf optical properties: New insights into plant acclimation to solar UV radiation.

The accumulation of UV-absorbing compounds (flavonoids and other phenylpropanoid derivatives) and resultant decrease in the UV transmittance of the epidermis in leaves (TUV), is a primary protective mechanism against the potentially deleterious effects of UV radiation and is a critical component of the overall acclimation response of plants to changing UV environments. Traditional measurements of TUV were laborious, time-consuming and destructive or invasive, thus limiting their ability to efficiently make multiple measurements of the optical properties of plants in the field. The development of rapid, nondestructive optical methods of determining TUV has permitted the examination of UV optical properties of leaves with increased replication, on a finer time scale, and enabled repeated sampling of the same leaf over time. This technology has therefore allowed for studies examining acclimation responses to UV in plants in ways not previously possible. Here we provide a brief review of these earlier studies examining leaf UV optical properties and some of their important contributions, describe the principles by which the newer non-invasive measurements of epidermal UV transmittance are made, and highlight several case studies that reveal how this technique is providing new insights into this UV acclimation response in plants, which is far more plastic and dynamic than previously thought.

Increased occurrence of pesticide residues on crops grown in protected environments compared to crops grown in open field conditions.

Crops grown under plastic-clad structures or in greenhouses may be prone to an increased frequency of pesticide residue detections and higher concentrations of pesticides relative to equivalent crops grown in the open field. To test this we examined pesticide data for crops selected from the quarterly reports (2004-2009) of the UK's Pesticide Residue Committee. Five comparison crop pairs were identified whereby one crop of each pair was assumed to have been grown primarily under some form of physical protection ('protected') and the other grown primarily in open field conditions ('open'). For each pair, the number of detectable pesticide residues and the proportion of crop samples containing pesticides were statistically compared (n=100 s samples for each crop). The mean concentrations of selected photolabile pesticides were also compared. For the crop pairings of cabbage ('open') vs. lettuce ('protected') and 'berries' ('open') vs. strawberries ('protected') there was a significantly higher number of pesticides and proportion of samples with multiple residues for the protected crops. Statistically higher concentrations of pesticides, including cypermethrin, cyprodinil, fenhexamid, boscalid and iprodione were also found in the protected crops compared to the open crops. The evidence here demonstrates that, in general, the protected crops possess a higher number of detectable pesticides compared to analogous crops grown in the open. This may be due to different pesticide-use regimes, but also due to slower rates of pesticide removal in protected systems. The findings of this study raise implications for pesticide management in protected-crop systems.

Endopolyploidy as a potential alternative adaptive strategy for Arabidopsis leaf size variation in response to UV-B.

The extent of endoreduplication in leaf growth is group- or even species-specific, and its adaptive role is still unclear. A survey of Arabidopsis accessions for variation at the level of endopolyploidy, cell number, and cell size in leaves revealed extensive genetic variation in endopolyploidy level. High endopolyploidy is associated with increased leaf size, both in natural and in genetically unstructured (mapping) populations. The underlying genes were identified as quantitative trait loci that control endopolyploidy in nature by modulating the progression of successive endocycles during organ development. This complex genetic architecture indicates an adaptive mechanism that allows differential organ growth over a broad geographic range and under stressful environmental conditions. UV-B radiation was identified as a significant positive climatic predictor for high endopolyploidy. Arabidopsis accessions carrying the increasing alleles for endopolyploidy also have enhanced tolerance to UV-B radiation. UV-absorbing secondary metabolites provide an additional protective strategy in accessions that display low endopolyploidy. Taken together, these results demonstrate that high constitutive endopolyploidy is a significant predictor for organ size in natural populations and is likely to contribute to sustaining plant growth under high incident UV radiation. Endopolyploidy may therefore form part of the range of UV-B tolerance mechanisms that exist in natural populations.

Reduction of photosynthetic sensitivity in response to abiotic stress in tomato is mediated by a new generation plant activator.

BACKGROUND: Yield losses as a result of abiotic stress factors present a significant challenge for the future of global food production. While breeding technologies provide potential to combat negative stress-mediated outcomes over time, interventions which act to prime plant tolerance to stress, via the use of phytohormone-based elicitors for example, could act as a valuable tool for crop protection. However, the translation of fundamental biology into functioning solution is often constrained by knowledge-gaps. RESULTS: Photosynthetic and transcriptomic responses were characterised in young tomato (Solanum lycopersicum L.) seedlings in response to pre-treatment with a new plant health activator technology, 'Alethea', followed by a subsequent 100 mM salinity stress. Alethea is a novel proprietary technology composed of three key constituent compounds; the hitherto unexplored compound potassium dihydrojasmonate, an analogue of jasmonic acid; sodium benzoate, a carboxylic acid precursor to salicylic acid, and the α-amino acid L-arginine. Salinity treatment led to a maximal 47% reduction in net photosynthetic rate 8 d following NaCl treatment, yet in Alethea pre-treated seedlings, sensitivity to salinity stress was markedly reduced during the experimental period. Microarray analysis of leaf transcriptional responses showed that while salinity stress and Alethea individually impacted on largely non-overlapping, distinct groups of genes, Alethea pre-treatment substantially modified the response to salinity. Alethea affected the expression of genes related to biotic stress, ethylene signalling, cell wall synthesis, redox signalling and photosynthetic processes. Since Alethea had clear effects on photosynthesis/chloroplastic function at the physiological and molecular levels, we also investigated the ability of Alethea to protect various crop species against methyl viologen, a potent generator of oxidative stress in chloroplasts. Alethea pre-treatment produced dramatic reductions in visible foliar necrosis caused by methyl viologen compared with non-primed controls. CONCLUSIONS: 'Alethea' technology mediates positive recovery of abiotic stress-induced photosynthetic and foliar loss of performance, which is accompanied by altered transcriptional responses to stress.

From ozone depletion to agriculture: understanding the role of UV radiation in sustainable crop production.

Largely because of concerns regarding global climate change, there is a burgeoning interest in the application of fundamental scientific knowledge in order to better exploit environmental cues in the achievement of desirable endpoints in crop production. Ultraviolet (UV) radiation is an energetic driver of a diverse range of plant responses and, despite historical concerns regarding the damaging consequences of UV-B radiation for global plant productivity as related to stratospheric ozone depletion, current developments representative of a range of organizational scales suggest that key plant responses to UV-B radiation may be exploitable in the context of a sustainable contribution towards the strengthening of global crop production, including alterations in secondary metabolism, enhanced photoprotection, up-regulation of the antioxidative response and modified resistance to pest and disease attack. Here, we discuss the prospect of this paradigm shift in photobiology, and consider the linkages between fundamental plant biology and crop-level outcomes that can be applied to the plant UV-B response, in addition to the consequences for related biota and many other facets of agro-ecosystem processes.

Multiple roles for UV RESISTANCE LOCUS8 in regulating gene expression and metabolite accumulation in Arabidopsis under solar ultraviolet radiation.

Photomorphogenic responses triggered by low fluence rates of ultraviolet B radiation (UV-B; 280-315 nm) are mediated by the UV-B photoreceptor UV RESISTANCE LOCUS8 (UVR8). Beyond our understanding of the molecular mechanisms of UV-B perception by UVR8, there is still limited information on how the UVR8 pathway functions under natural sunlight. Here, wild-type Arabidopsis (Arabidopsis thaliana) and the uvr8-2 mutant were used in an experiment outdoors where UV-A (315-400 nm) and UV-B irradiances were attenuated using plastic films. Gene expression, PYRIDOXINE BIOSYNTHESIS1 (PDX1) accumulation, and leaf metabolite signatures were analyzed. The results show that UVR8 is required for transcript accumulation of genes involved in UV protection, oxidative stress, hormone signal transduction, and defense against herbivores under solar UV. Under natural UV-A irradiance, UVR8 is likely to interact with UV-A/blue light signaling pathways to moderate UV-B-driven transcript and PDX1 accumulation. UVR8 both positively and negatively affects UV-A-regulated gene expression and metabolite accumulation but is required for the UV-B induction of phenolics. Moreover, UVR8-dependent UV-B acclimation during the early stages of plant development may enhance normal growth under long-term exposure to solar UV.

The UV-B photoreceptor UVR8 promotes photosynthetic efficiency in Arabidopsis thaliana exposed to elevated levels of UV-B.

The UV-B photoreceptor UVR8 regulates expression of genes in response to UV-B, some encoding chloroplast proteins, but the importance of UVR8 in maintaining photosynthetic competence is unknown. The maximum quantum yield of PSII (F (v)/F(m)) and the operating efficiency of PSII (Φ(PSII)) were measured in wild-type and uvr8 mutant Arabidopsis thaliana. The importance of specific UVR8-regulated genes in maintaining photosynthetic competence was examined using mutants. Both F (v)/F(m) and Φ(PSII) decreased when plants were exposed to elevated UV-B, in general more so in uvr8 mutant plants than wild-type. UV-B increased the level of psbD-BLRP (blue light responsive promoter) transcripts, encoding the PSII D2 protein. This increase was mediated by the UVR8-regulated chloroplast RNA polymerase sigma factor SIG5, but SIG5 was not required to maintain photosynthetic efficiency at elevated UV-B. Levels of the D1 protein of PSII decreased markedly when plants were exposed to elevated UV-B, but there was no significant difference between wild-type and uvr8 under conditions where the mutant showed increased photoinhibition. The results show that UVR8 promotes photosynthetic efficiency at elevated levels of UV-B. Loss of the DI polypeptide is probably important in causing photoinhibition, but does not entirely explain the reduced photosynthetic efficiency of the uvr8 mutant compared to wild-type.

Ecological responses to UV radiation: interactions between the biological effects of UV on plants and on associated organisms.

Solar ultraviolet (UV)-B radiation (280-315 nm) has a wide range of effects on terrestrial ecosystems, yet our understanding of how UV-B influences the complex interactions of plants with pest, pathogen and related microorganisms remains limited. Here, we report the results of a series of experiments in Lactuca sativa which aimed to characterize not only key plant responses to UV radiation in a field environment but also consequential effects for plant interactions with a sap-feeding insect, two model plant pathogens and phylloplane microorganism populations. Three spectrally modifying filters with contrasting UV transmissions were used to filter ambient sunlight, and when compared with our UV-inclusive filter, L. sativa plants grown in a zero UV-B environment showed significantly increased shoot fresh weight, reduced foliar pigment concentrations and suppressed population growth of green peach aphid (Myzus persicae). Plants grown under a filter which allowed partial transmission of UV-A radiation and negligible UV-B transmission showed increased density of leaf surface phylloplane microbes compared with the UV-inclusive treatment. Effects of UV treatment on the severity of two plant pathogens, Bremia lactucae and Botrytis cinerea, were complex as both the UV-inclusive and zero UV-B filters reduced the severity of pathogen persistence. These results are discussed with reference to known spectral responses of plants, insects and microorganisms, and contrasted with established fundamental responses of plants and other organisms to solar UV radiation, with particular emphasis on the need for future integration between different experimental approaches when investigating the effects of solar UV radiation.

Increased exposure to UV-B radiation during early development leads to enhanced photoprotection and improved long-term performance in Lactuca sativa.

Plant responses to solar UV radiation are numerous and have often been considered from a perspective of negative outcomes for plant productivity. In this study, we used two experimental approaches consisting of: (1) field-based spectrally modifying filters in addition to (2) controlled indoor exposure to UV-B, to examine the effects of UV radiation on growth and photosynthetic performance of lettuce (Lactuca sativa L.) seedlings. Various aspects of growth were affected in plants grown under a UV-inclusive environment compared to a UV-depleted environment, including reductions in leaf expansion, increases in leaf thickness and the rate of net photosynthesis. After transplantation to a uniform field environment, lettuce plants initially propagated under the UV-inclusive environment exhibited higher harvestable yields than those from a UV-depleted environment. In controlled conditions, photosynthetic rates were higher in plants grown in the presence of UV-B radiation, and relative growth of plants pre-acclimatized to UV-B was also increased, in addition to higher maximum photochemical efficiency of photosystem II (PSII) (F(v) /F(m) ) following subsequent exposure to high photosynthetically active radiation (PAR) and temperature stress. Our findings are discussed within the context of sustainability in agriculture and the paradigm shift in photobiology which such beneficial responses to UV radiation could represent.

UVR8 in Arabidopsis thaliana regulates multiple aspects of cellular differentiation during leaf development in response to ultraviolet B radiation.

Responses specific to ultraviolet B (UV-B) wavelengths are still poorly understood, both in terms of initial signalling and effects on morphogenesis. Arabidopsis thaliana UV RESISTANCE LOCUS8 (UVR8) is the only known UV-B specific signalling component, but the role of UVR8 in leaf morphogenesis is unknown. The regulatory effects of UVR8 on leaf morphogenesis at a range of supplementary UV-B doses were characterized, revealing both UVR8-dependent and independent responses to UV irradiation. Inhibition of epidermal cell division in response to UV-B is largely independent of UVR8. However, overall leaf growth under UV-B irradiation in wild-type plants is enhanced compared with a uvr8 mutant because of a UVR8-dependent compensatory increase of cell area in wild-type plants. UVR8 was also required for the regulation of endopolyploidy in response to UV-B, and the uvr8 mutant also has a lower density of stomata than the wild type in the presence of UV-B, indicating that UVR8 has a regulatory role in other developmental events. Our findings show that, in addition to regulating UV-protective gene expression responses, UVR8 is involved in controlling aspects of leaf growth and morphogenesis. This work extends our understanding of how UV-B response is orchestrated at the whole-plant level.

A comparative study on the aqueous photodegradation of two organophosphorus pesticides under simulated and natural sunlight.

Aqueous solutions of fenitrothion and methyl parathion were photochemically degraded in an Atlas Suntest solar simulator (500 W m(-2)) as well as under ambient sunlight at Lancaster University (June & August 2007, 54 degrees N) and the degradation kinetics and disappearance quantum yields are reported. Fenitrothion degradation confirmed to first order kinetics (r(2) = 0.90-0.99) with a half-life range of 4.9 h-5.3 h, shorter than previously reported studies. In contrast, methyl parathion did not show significant degradation over the duration of these experiments. Light irradiances were monitored with a spectroradiometer during the Suntest simulator and outdoor experiments. The filtered-xenon arc lamp of the Suntest yielded spectral irradiances comparable to natural sunlight in the UVB and UVA wavelength range (280-400 nm), but with higher irradiances in the visible region (400-750 nm). Nonetheless, as both compounds have light absorption spectra at wavelengths < 400 nm, then the half-lives and disappearance quantum yields were similar between the Suntest and natural sunlight, and demonstrated that the Suntest is suitable for environmentally-relevant photochemical degradation experiments.

The aqueous photodegradation of fenitrothion under various agricultural plastics: implications for pesticide longevity in agricultural 'micro-environments'.

Plastic cladding is increasingly used in agriculture to create micro-environments to improve crop yield/growth. Many of these plastics can alter the solar light spectrum by inhibiting or reducing the transmittance of certain parts of the solar spectrum. In this study, we investigated the aqueous photolysis of fenitrothion, under a selection of different plastic films commonly used in agriculture. Three different grades of polyethylene film were used: 'standard', 'transparent' and 'opaque'. The transmittance of light wavelengths in the UV region (290-400 nm), measured with a spectroradiometer, was found to decrease in the order of transparent>standard>opaque. Fenitrothion, an organophosphorothioate insecticide possesses molar absorptivity in the solar wavelength range of 290-400 nm. Aqueous first order degradation rate constants for fenitrothion determined over a 12 h period were found to be considerably less for those experiments conducted under the standard and opaque plastic films, compared to the transparent film and no-film control. The experiments were conducted in an Atlas Suntest solar simulator using a UV-filtered Xenon arc lamp to simulate sunlight. The first order half-life for fenitrothion was 100 and 250 h under the standard and opaque films, respectively, compared to approximately 10 h for the transparent film and no-film experiments. Our results suggest that pesticide longevity could be greatly extended within these plastic micro-environments, especially for those chemicals which may degrade/transform via photolytic or photochemical pathways.

Ultraviolet radiation as a limiting factor in leaf expansion and development.

Reductions in leaf growth are a commonly observed response to ultraviolet radiation, but the underlying mechanisms remain poorly defined. This study examined the response of leaves exposed to a UV environment across a range of organizational scales, including leaf expansion rate, epidermal cell size and number, biomechanical properties, leaf-water relations and activity of cell-wall peroxidases. Two experimental approaches were used; Lettuce (Lactuca sativa L.) plants were propagated under (a) supplementary UV-B (9 kJ m(-2) day(-1)) in controlled environment (CE) conditions, and (b) field conditions, where plants were placed under three horticultural films with differing UV transmissions. In both experiments, UV-B caused the greatest reductions in leaf expansion and final leaf size, with some reductions attributable to UV-A wavelengths. In supplementary UV-B conditions, adaxial cell size was reduced, while in field plants, both cell size and cell number were lower in an increased UV environment, as was the case with abaxial cells in CE plants. Although leaf turgor and leaf extensibility were not affected by UV wavelengths, breaking strain of leaf tissue was decreased under supplementary UV-B. Cell-wall peroxidase activity was increased in both supplementary UV conditions and in the field, where only a zero UV environment showed no upregulation of cell-wall peroxidase.

Tri-trophic consequences of UV-B exposure: plants, herbivores and parasitoids.

In this paper we demonstrate a UV-B-mediated link between host plants, herbivores and their parasitoids, using a model system consisting of a host plant Brassica oleracea, a herbivore Plutella xylostella and its parasitoid Cotesia plutellae. Ultraviolet-B radiation (UV-B) is a potent elicitor of a variety of changes in the chemistry, morphology and physiology of plants and animals. Recent studies have demonstrated that common signals, such as jasmonic acid (JA), play important roles in the mechanisms by which plants respond to UV-B and to damage by herbivores. Plant responses elicited by UV-B radiation can affect the choices of ovipositing female insects and the fitness of their offspring. This leads to the prediction that, in plants, the changes induced as a consequence of UV damage will be similar to those elicited in response to insect damage, including knock-on effects upon the next trophic level, predators. In our trials female P. xylostella oviposited preferentially on host plants grown in depleted UV-B conditions, while their larvae preferred to feed on tissues from UV-depleted regimes over those from UV-supplemented ones. Larval feeding patterns on UV-supplemented tissues met the predictions of models which propose that induced defences in plants should disperse herbivory; feeding scars were significantly smaller and more numerous--though not significantly so--than those on host plant leaves grown in UV-depleted conditions. Most importantly, female parasitoids also showed a clear pattern of preference when given the choice between host plants and attendant larvae from the different UV regimes; however, in the case of the female parasitoids, the choice was in favour of potential hosts foraging on UV-supplemented tissues. This study demonstrates the potential for UV-B to elicit a variety of interactions between trophic levels, most likely mediated through effects upon host plant chemistry.