UV-B effects on the nutritional chemistry of plants and the responses of a mammalian herbivore.

Stratospheric ozone depletion has caused ground-level ultraviolet-B (UV-B) radiation to rise in temperate latitudes of both hemispheres. Because the effects of enhanced UV-B radiation on the nutrition of food consumed by mammalian herbivores are unknown, we measured nutritional and chemical constituents of 18 forages and related changes to in vitro dry matter digestibility. We also measured intake and in vivo digestibility of Pacific willow (Salix lasiandra) and alfalfa (Medicago sativa L.) by blue duikers (Cephalophus monticola). Forages were irradiated for 3 months with ambient (1x) or supplemental (1.6 x) UV-B radiation representing a 15% ozone depletion for Pullman, Washington, USA. Enhanced UV-B radiation had minimal and inconsistent effects on the nutritional content, in vitro dry matter digestibility, and protein-binding capacity of forages. However, flavonoid compounds increased in seven of the 13 forbs and woody dicots that were evaluated. Flavonoids were found to decrease only in yarrow (Achillea millefolium). When offered simultaneously, blue duikers preferred 1x and 1.6 x UV-B irradiated plants of alfalfa equally, but ate 26% less willow grown under 1.6 x UV-B radiation. However, when fed to duikers in separate feeding experiments, total dry matter intake and in vivo digestibility of dry matter, fiber, protein, and apparent energy did not differ between alfalfa and willow grown under 1x and 1.6 x UV-B radiation. We conclude that expected increases in UV-B radiation from ozone depletion would have minimal effects on intake and digestion of ruminant herbivores.

Effects of enhanced UV-B radiation on plant chemistry: nutritional consequences for a specialist and generalist lagomorph.

Ultraviolet-B (UV-B) radiation has been increasing in temperate latitudes in recent decades and is expected to continue rising for some time. Enhanced UV-B radiation can change plant chemistry, yet the effects of these changes on mammalian herbivores are unknown. To examine the influence of enhanced UV-B radiation on nutrition of a specialist and generalist hindgut fermenter, we measured nutritional and chemical constituents of three common North American range plants, big sagebrush (Artemisia tridentata), yarrow (Achillea millefolium), and bluebunch wheatgrass (Pseudoregneria spicata), and how these changes influenced in vitro dry matter digestibility and in vivo digestibility by pygmy rabbits (Brachylagus idahoensis) and eastern cottontails (Sylvilagus floridanus). Forages were irradiated for 3 mo with ambient (1x) or supplemental (1.6x) UV-B radiation representing a 15% ozone depletion for Pullman, WA, USA. Enhanced UV-B radiation had minimal effects on the nutritional content and the tannin-binding capacity of forages. Similarly, the terpene concentration in sagebrush and yarrow was not affected by higher UV-B irradiances. Flavonoid compounds increased in sagebrush but decreased in yarrow. Rabbit preference and intake was not affected by treatment levels for any forage species and no differences were found between treatments for dry matter, fiber, protein digestibility, and apparent digestible energy.

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

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

Ecosystem consequences of enhanced solar ultraviolet radiation: secondary plant metabolites as mediators of multiple trophic interactions in terrestrial plant communities.

The potential role of ultraviolet-B (UV-B)-induced secondary plant metabolites as mediators of multiple trophic responses in terrestrial ecosystems is considered through review of the major classes of secondary metabolites, the pathways for their biosynthesis, interactions with primary and secondary consumers and known UV effects on their induction. Gross effects of UV-B radiation on plant growth and survival under realistic spectral balances in the field have been generally lacking, but subtle changes in carbon allocation and partitioning induced by UV-B, in particular production of secondary metabolites, can affect ecosystem-level processes. Secondary metabolites are important in plant-herbivore interactions and may affect pathogens. They act as feeding or oviposition deterrents to generalists and nonadapted specialists, but adapted specialists are stimulated to feed by these same compounds, which they detoxify and often sequester for use against their predators. This provides a route for tritrophic effects of enhanced UV-B radiation whereby herbivory may be increased while predation on the herbivore is simultaneously reduced. It is in this context that secondary metabolites may manifest their most important role. They can be the demonstrable mechanism establishing cause and effect at higher trophic levels because the consequences of their induction can be established at all trophic levels.

Rocky Mountain ecosystems: diversity, complexity and interactions.

The interior west of North America provides many opportunities to study ecosystem responses to climate change, biological diversity and management of disturbance regimes. These ecosystem responses are not unique to the Rocky Mountains, but they epitomize similar scientific problems throughout North America. Better management of these ecosystems depends on a thorough understanding of the underlying biology and ecological interactions of the species that occupy the diverse habitats of this region. This review highlights progress in research to understand aspects of this complex ecosystem.

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

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

Leaf chemical changes induced in Populus trichocarpa by enhanced UV-B radiation and concomitant effects on herbivory by Chrysomela scripta (Coleoptera: Chrysomelidae).

To assess the potential impact of enhanced ultraviolet-B (UV-B) radiation over two trophic levels, we monitored key leaf chemical constituents and related changes in their concentration to dietary preference and performance of a specialist insect herbivore. Ramets of Populus trichocarpa Torr. & Gray (black cottonwood) were subjected to near zero (0X), ambient (1X) or twice ambient (2X) doses of biologically effective UV-B radiation (UV-B(BE)) in a randomized block design using either a square-wave (greenhouse) or a modulated (field) lamp system. After a 3-month treatment period, apparent photosynthesis was determined in situ and plants were harvested for biomass determination. Leaf subsamples were analyzed for nitrogen, sulfur, chlorophylls, UV-absorbing compounds and protein-precipitable tannins. Effects of changes in these constituents on feeding by Chrysomela scripta Fab. (cottonwood leaf beetle) were determined by (1) adult feeding preference trials and (2) larval growth rate trials. Enhanced UV-B(BE) radiation had minimal effects on photosynthesis, growth, leaf area and biomass distribution. In the greenhouse study, concentrations of foliar nitrogen and chlorophylls increased, but tannins decreased slightly in young leaves exposed to enhanced UV-B(BE) radiation. There were no significant effects on these parameters in the field study. The concentration of methanol-extractable foliar phenolics increased in plants grown with enhanced UV-B(BE) radiation in both the greenhouse and field studies. In feeding preference trials, adult C. scripta chose 2X-treated tissue almost twice as often as 1X-treated tissue in both greenhouse and field studies, but differences were not statistically significant (P = 0.12). In the field study, first instar larvae grown to adult eclosion on 2X-treated leaves had a significant (P < 0.001) reduction in consumption efficiency compared with larvae grown on 1X-treated leaves. We conclude that effects of enhanced UV-B(BE) radiation at the molecular-photochemical level can elicit significant responses at higher trophic levels that may ultimately affect forest canopy structure, plant competitive interactions and ecosystem-level processes.

Alteration of foliar flavonoid chemistry induced by enhanced UV-B radiation in field-grown Pinus ponderosa, Quercus rubra and Pseudotsuga menziesii.

Chromatographic analyses of foliage from several tree species illustrate the species-specific effects of UV-B radiation on both quantity and composition of foliar flavonoids. Pinus ponderosa, Quercus rubra and Pseudotsuga menziesii were field-grown under modulated ambient (1x) and enhanced (2x) biologically effective UV-B radiation. Foliage was harvested seasonally over a 3-year period, extracted, purified and the flavonoid fraction applied to a mu Bondapak/C(18) column HPLC system sampling at 254 nm. Total flavonoid concentrations in Quercus rubra foliage were more than twice (leaf area basis) that of the other species; Pseudotsuga menziesii foliage had intermediate levels and P. ponderosa had the lowest concentrations of total flavonoids. No statistically significant UV-B radiation-induced effects were found in total foliar flavonoid concentrations for any species; however, concentrations of specific compounds within each species exhibited significant treatment effects. Higher (but statistically insignificant) levels of flavonoids were induced by UV-B irradiation in 1- and 2-year-old P. ponderosa foliage. Total flavonoid concentrations in 2-year-old needles increased by 50% (1x ambient UV-B radiation) or 70% (2x ambient UV-B radiation) from that of 1-year-old tissue. Foliar flavonoids of Q. rubra under enhanced UV-B radiation tended to shift from early-eluting compounds to less polar flavonoids eluting later. There were no clear patterns of UV-B radiation effects on 1-year-old P. menziesii foliage. However, 2-year-old tissue had slightly higher foliar flavonoids under the 2x UV-B radiation treatment compared to ambient levels. Results suggest that enhanced UV-B radiation will alter foliar flavonoid composition and concentrations in forest tree species, which could impact tissue protection, and ultimately, competition, herbivory or litter decomposition.