Yield and nutritive quality of sericea lespedeza (Lespedeza cuneata) and little bluestem (Schizachyrium scoparium) exposed to ground-level ozone

Sericea lespedeza (Lespedeza cuneata cv. Interstate 76) and little bluestem (Schizachyrium scoparium cv. Aldous) were raised from seed in a glasshouse, transplanted into 5.7-l pots and placed into open-top chambers (OTC) on 6 June 1999. Following a 7-day adjustment period, each of six OTCs (duplicate OTCs per treatment) was ventilated with either air that had been carbon-filtered (CF) to remove ambient ozone (O3); non-filtered (NF), representative of ambient air; or enriched to twice-ambient O3 concentration (2X). Primary-growth forage was harvested on days 7, 32, 46, 59 and 72 following the start of fumigation, and regrowth forage from the first primary-growth harvest was harvested on days 36, 54 and 72 following the start of fumigation. Dry matter (DM) yield of either forage species did not differ among treatments except in the final regrowth period when yield of sericea lespedeza was greater for the NF than 2X O3 treatment. In vitro DM digestibility (IVDMD) and concentrations of crude protein (CP), soluble phenolics (SP) and condensed tannins (CT) in primary-growth sericea lespedeza did not differ between treatments, but NF primary-growth forage had higher concentration of protein-precipitating tannins (PPT) than did 2X primary-growth forage. Concentrations of neutral detergent fiber (NDF), acid detergent fiber (ADF) and acid detergent lignin (ADL) were higher in NF and 2X than in CF primary-growth sericea lespedeza. Similarly, concentrations of NDF and ADL were higher, and IVDMD was lower for NF and 2X than for CF regrowth sericea lespedeza. Concentrations of ADF and ADL were lower, whereas IVDMD, in vitro NDF digestibility (IVNDFD) and concentrations of CP and SP were greater, in CF than in NF and 2X primary-growth little bluestem. Percentages IVDMD and IVNDFD and concentrations of CP and SP in NF primary-growth little bluestem were greater than those in forage exposed to 2X O3 treatment. No significant differences were observed among treatments in percentages IVDMD and IVNDFD, or concentrations of cell wall constituents or SP in little bluestem regrowth. Nutritive quality of little bluestem was decreased by < 2%, and that of sericea lespedeza by approximately 7% as a result of increased concentrations of cell wall constituents and decreased in vitro digestibility of NF and 2X compared with CF forages. Results indicate that existing and projected O3 levels can drive alterations in forage quality of select warm-season forages sufficient to have nutritional and economic implications for their utilization by ruminant herbivores.     

Sub-canopy deposition of ozone in a stand of cutleaf coneflower

Although there has been a great deal of research on ozone, interest in exposure of native, herbaceous species is relatively recent and it is still not clear what role the pollutant has in their ecological fitness. The ozone exposure of a plant is usually expressed in terms of the concentration above the canopy or as a time-weighted index. However, to understand the physiological effects of ozone it is necessary to quantify the ozone flux to individual leaves as they develop, which requires knowing the deposition velocity and concentration of the pollutant as a function of height throughout the plant canopy. We used a high-order closure model of sub-canopy turbulence to estimate ozone profiles in stands of cutleaf coneflower (Rudbeckia laciniata L.) located in the Great Smoky Mountains National Park, USA. The model was run for periods coinciding with a short field study, during which we measured vertical concentration profiles of ozone along with measurements of atmospheric turbulence and other meteorological and plant variables. Predictions of ozone profiles by the model are compared with observations throughout the canopy.     

Ambient ozone effects on gas exchange and total non-structural carbohydrate levels in cutleaf coneflower (Rudbeckia laciniata L.) growing in Great Smoky Mountains National Park

Ozone-sensitive and -tolerant individuals of cutleaf coneflower (Rudbeckia laciniata L.) were compared for their gas exchange characteristics and total non-structural carbohydrates at Purchase Knob, a high elevation site in Great Smoky Mountains National Park, USA. Photosynthesis and stomatal conductance decreased with increased foliar stipple. Sensitive plants had lower photosynthetic rates for all leaves, except the very youngest and oldest when compared to tolerant plants. Stomatal conductance decreased with increasing leaf age, but no ozone-sensitivity differences were found. Lower leaves had less starch than upper ones, while leaves on sensitive plants had less than those on tolerant plants. These results show that ambient levels of ozone in Great Smoky Mountains National Park can adversely affect gas exchange, water use efficiency and leaf starch content in sensitive coneflower plants. Persistence of sensitive genotypes in the Park may be due to physiological recovery in low ozone years.     

Seasonal development of ozone-induced foliar injury on tall milkweed (Asclepias exaltata) in Great Smoky Mountains National Park

The goals of this study were to document the development of ozone-induced foliar injury, on a leaf-by-leaf basis, and to develop ozone exposure relationships for leaf cohorts and individual tall milkweeds (Asclepias exaltata L.) in Great Smoky Mountains National Park. Plants were classified as either ozone-sensitive or insensitive based on the amount of foliar injury. Sensitive plants developed injury earlier in the season and to a greater extent than insensitive plants. Older leaf cohorts were more likely to belong to high injury classes by the end of each of the two growing seasons. In addition, leaf loss was more likely for older cohorts (2000) and lower leaf positions (2001) than younger cohorts and upper leaves, respectively. Most leaves abscised without prior ozone-like stippling or chlorosis. Failure to take this into account can result in underestimation of the effects of ozone on these plants.     

Ozone injury on cutleaf coneflower (Rudbeckia laciniata) and crown-beard (Verbesina occidentalis) in Great Smoky Mountains National Park

Incidence and severity of visible foliar ozone injury on cutleaf coneflower (Rudbeckia laciniata L.) and crown-beard (Verbesina occidentalis Walt.) were determined along selected trails at three locations in Great Smoky Mountains National Park during the summers of 2000 and 2001: Clingmans Dome, Cherokee Orchard Road and Purchase Knob. Cutleaf coneflower exhibited a greater amount of foliar injury than crown-beard each year of the 2-year study. Incidence and severity of injury was significantly greater for cutleaf coneflower growing near the edge of the Clingmans Dome trail than in the interior of the stand. Injury was greater at Clingmans Dome than Purchase Knob (70% vs. 40% ozone-injured plants, respectively), coincident with greater ozone exposures. In contrast to Clingmans Dome, there were no differences in injury between plants growing near- and off-trail at Purchase Knob. Differences in sensitivity to ozone were not observed for crown-beard growing near the edge compared with the interior of the stand adjacent to the Cherokee Orchard Road Loop. Ozone injury was greatest on the lower leaves for both species sampled with over 95% of the injured leaves occurring on the lower 50% of the plant. This is the first report of foliar ozone injury on these plant species in situ, in the Park, illustrating the great variability in symptom expression with time, and within and between populations.     

Effects of soil-applied lead on seedling growth and ectomycorrhizal colonization of loblolly pine

Six-month-old loblolly pine (Pinus taeda) were grown for 15 weeks in two native soils amended with 0, 30, 60, 120, 240 or 480 mg kg(-1) Pb as PbCl2. Ectomycorrhizae were quantified, by morphotype, as the total number of tips per centimeter, and as the number of tips for each morphotype and for Cenococcum geophilum. Total numbers of non-ectomycorrhizal short roots and necrotic tips were recorded. Total height and biomass exhibited a non-linear response to soil-applied lead. Growth generally was greatest in the controls and higher treatments, and least in the intermediate treatments. In both soils, Pb concentrations in roots increased linearly with increasing levels of soil-applied Pb. Neither foliage nor stems exhibited significant increases in Pb concentrations with increasing levels of Pb. Significant linear decreases in total numbers of ectomycorrhizal tips and significant linear increases in non-ectomycorrhizal short roots and necrotic tips occurred with increasing levels of Pb in the soil. The majority of individual morphotypes decreased with increasing Pb. However, the number of ectomycorrhizal tips formed by C. geophilium increased with increasing soil Pb levels after 15 weeks of treatment. These results indicate that short-term loblolly pine seedling growth is not inhibited by increasing Pb levels. Ectomycorrhizal formation decreased, and alterations in species composition occurred as a result of increasing concentrations of soil-applied Pb. These effects on ectomycorrhizae may cause long-term changes in nutrient and water balances, which could reduce tree vigor.     

Temporal patterns of foliar ozone symptoms on tall milkweed (Asclepias exaltata L.) in Great Smoky Mountains National Park

Incidence and severity of ozone-induced foliar symptoms on tall milkweed (Asclepias exaltata L.) along selected trails in Great Smoky Mountains National Park (GRSM) were determined by two surveys/season conducted from 1992 through 1996. Overall incidence was 73%, and was 84%, 44%, 90%, 58%, and 82% for 1992-1996, respectively for the same clusters. Average incidence was 61% and 84% for the 1st and 2nd surveys, respectively. Seasonal comparisons showed two distinct injury groupings regarding incidence and severity of injury: 1992, 1994 and 1996 (high injury); 1993 and 1995 (low injury). No discernible patterns were observed between symptomatic and asymptomatic plants regarding height, herbivory or flowering. Regression analyses indicated no differentiation in foliar symptoms regarding topographic position, aspect, slope or elevation over the 5-year study period. Our findings indicate other micro-site or genetic factors may control ozone sensitivity of tall milkweed in GRSM.     

Perspectives regarding 50 years of research on effects of tropospheric ozone air pollution on US forests

Tropospheric ozone (O(3)) was first determined to be phytotoxic to grapes in southern California in the 1950s. Investigations followed that showed O(3) to be the cause of foliar symptoms on tobacco and eastern white pine. In the 1960s, "X" disease of ponderosa pines within the San Bernardino Mountains was likewise determined to be due to O(3). Nearly 50 years of research have followed. Foliar O(3) symptoms have been verified under controlled chamber conditions. Studies have demonstrated negative growth effects on forest tree seedlings due to season-long O(3) exposures, but due to complex interactions within forest stands, evidence of similar losses within mature tree canopies remains elusive. Investigations on tree growth, O(3) flux, and stand productivity are being conducted along natural O(3) gradients and in open-air exposure systems to better understand O(3) effects on forest ecosystems. Given projected trends in demographics, economic output and climate, O(3) impacts on US forests will continue and are likely to increase.     

Impacts of climatic and atmospheric changes on carbon dynamics in the Great Smoky Mountains National Park

We used the Dynamic Land Ecosystem Model (DLEM) to estimate carbon (C) storage and to analyze the impacts of environmental changes on C dynamics from 1971 to 2001 in Great Smoky Mountain National Park (GRSM). Our simulation results indicate that forests in GRSM have a C density as high as 15.9kgm(-2), about twice the regional average. Total carbon storage in GRSM in 2001 was 62.2Tg (T=10(12)), 54% of which was in vegetation, the rest in the soil detritus pool. Higher precipitation and lower temperatures in the higher elevation forests result in larger total C pool sizes than in forests at lower elevations. During the study period, the CO(2) fertilization effect dominated ozone and climatic stresses (temperature and precipitation), and the combination of these multiple factors resulted in net accumulation of 0.9Tg C in this ecosystem.