Greg L. Somers

Evaluation of ozone injury on foliage of black cherry (Prunus serotina) and tall milkweed (Asclepias exaltata) in Great Smoky Mountains National Park

The incidence and severity of visible foliar ozone injury on black cherry (Prunus serotina) seedlings and saplings and tall milkweed (Asclepias exaltata) plants in Great Smoky Mountains National Park (GRSM) were determined by surveys along selected trails conducted during late summer 1992. The incidence (% injured plants) of ozone injury on black cherry was 47% and the percent injured leaves/injured plant and average leaf area injured were 43 and 6%, respectively. Maximum severity (avg. leaf area of the most severely injured leaf) was 12%. Black cherry seedlings and saplings exhibiting ozone injury were taller than non-injured plants. When insect feeding was present, it occurred 96% of the time on plants with ozone injury. Significantly more injury (p=0.007) on black cherry (% injured leaves/injured black cherry) occurred in the NW section of GRSM compared with the other Park sections. Regression analyses showed no relationships in ozone injury with respect to aspect, slope or elevation. Tall milkweed was evaluated twice during August for ozone injury. The incidence (% injured plants) of ozone injury was 74 and 79% for the first and second survey, respectively. The percentage of injured leaves per plant from the first to second survey was 63 to 79%, respectively. Tall milkweeds showing ozone injury were taller than the non-injured plants. The percentage of insect-damaged plants was 50% among plants without ozone injury and 60% among ozone-injured plants. Non-injured tall milkweed had fewer flowers and/or pods than the injured plants. Mean leaf area injured increased over time, and mean maximum leaf area injured increased from 8 to 11% during the same period. Regression analyses showed no differences in ozone injury regarding aspect, slope or elevation. Our findings indicate that ozone injury is widespread throughout the Park on sensitive vegetation.

Visible foliar injury caused by ozone alters the relationship between SPAD meter readings and chlorophyll concentrations in cutleaf coneflower

The ability of the SPAD-502 chlorophyll meter to quantify chlorophyll amounts in ozone-affected leaves of cutleaf coneflower (Rudbeckia laciniata var. digitata) was assessed in this study. When relatively uninjured leaves were measured (percent leaf area affected by stipple less than 6%), SPAD meter readings were linearly related to total chlorophyll with an adjusted R2 of 0.84. However, when leaves with foliar injury (characterized as a purple to brownish stipple on the upper leaf surface affecting more than 6% of the leaf area) were added, likelihood ratio tests showed that it was no longer possible to use the same equation to obtain chlorophyll estimations for both classes of leaves. Either an equation with a common slope or a common intercept was necessary. We suspect several factors are involved in altering the calibration of the SPAD meter for measuring chlorophyll amounts in visibly ozone-injured leaves, with the most likely being changes in either light absorption or scattering resulting from tissue necrosis.

Tree shelters and weed control: Effects on protection, survival and growth of cherrybark oak seedlings planted on a cutover site

In the southern USA oaks (Quercusspp.) are often favored by forest owners havingmultiple objectives for forest ownership as oaksprovide mast for wildlife, are consideredaesthetically pleasing, and are valuable for timberproducts. Regeneration and early seedling growth isa concern to those forest owners interested insustaining oaks as a component of their forests. Theeffects of tree shelters and herbaceous weed controlon second-year seedling survival, browse by deer andrabbits, and seedling growth of hand-plantedcherrybark oak (Quercus pagoda Raf.) wereexamined. The study was established on a cutovermixed pine-hardwood forest in Alabama. Fourtreatments were: weed control only, tree shelter only,tree shelter with weed control, and a controlconsisting of a seedling without a tree shelter orweed control. No significant difference in seedlingsurvival was found among the treatments after twoyears. Tree shelters were effective in preventingbrowsing. No seedlings in the tree-shelter-onlytreatment were browsed. There was no significant,difference, however, in the percentage of seedlingsbrowsed between the control treatment and the weedcontrol treatment. The use of tree shelters with weedcontrol was the most effective treatment for promoting2-year ground-line diameter, height, and stem volumegrowth.

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.

Empirical evidence of growth decline related to visible ozone injury

Differences in radial growth at breast height of yellow-poplar (Liriodendron tulipifera L.) and black cherry (Prunus serotina Ehrh.) were tested between individual trees with a history of visible foliar ozone injury and those not expressing foliar injury to ozone at three sites in Great Smoky Mountains National Park, USA. No significant differences in growth for black cherry were found although there was a 12% reduction in radial growth over 5 yrs (1990–1994) (p-level 0.4) and 8% over 10 yrs (1985–1994) (p-level 0.6). There was a significant difference in radial growth for yellow-poplar of 43% over 5 yrs (p-level 0.001) and 30% over 10 yrs (p-level 0.005). Even though the trees of both species were selected to balance the diameter distribution of each species between the two groups at each site, it was still possible that the differences were due to some other factors than sensitivity to ozone exposure. Therefore, a series of multiple linear regressions were used to identify the most explanatory model based on principal components derived from the following independent variables: Diameter at breast height, total height, height to the live crown, percent slope, and a number of competition indices based on the diameter and distance to competitors. These regressions were then tested for different intercepts and slopes between the sensitive and nonsensitive trees. Once again, no significant differences occurred for black cherry (p-levels of 0.4 and 0.7 for five-year and ten-year radial growth, respectively) and some differences for yellow-poplar (p-levels of 0.04 and 0.1 for five-year and ten-year radial growth, respectively). Although the conclusions did not change, the importance of proper balancing of the diameter distribution and accounting for the effects of uncontrollable independent variables are discussed.

Mature Black Cherry Used as a Bioindicator of Ozone Injury

Incidence and severity of foliar symptoms due to ambient ozone exposures were documented on mature black cherry (Prunus serotina) in two National Parks [Great Smoky Mountains National Park (GRSM) and Shenandoah National Park (SHEN)] in the Appalachian Mountains of the eastern USA during the summer of 1991-1993. Three plots in each park containing 30 trees each (Big Meadows in SHEN had 60 trees) with 90 and 120 trees total trees were evaluated in GRSM and SHEN, respectively. Plots were established at different elevations adjacent to ozone monitoring stations. Samples of foliage were collected and three exposed branches from the upper- crown and three branches from the mid-to-lower crown were examined for symptoms of foliar ozone injury. Incidence was greatest in 1991 at both locations; 60% and 45% for GRSM and SHEN, respectively. In 1992 and 1993, incidence was very similar in both parks, with approximately 33% of the trees affected. Black cherry at the highest elevations exhibited the greatest amount of symptoms in both parks all three years of the study. These sites also exhibited the highest ozone concentrations. In addition, the percent of trees injured by ozone was positively correlated with SUM06 and W126. These results along with forest surveys and open-top chamber studies indicate that black cherry may be a reliable bioindicator of foliar injury due to ambient ozone.

Visible Ozone Injury on Forest Trees in Great Smoky Mountains National Park, USA

During the summer of 1991 ozone injury trend plots in Great Smoky Mountains National Park, USA, consisting of mature black cherry, sassafras and yellow-poplar were established near three ozone monitors, ranging in elevation from 597-1265 m. Beginning in mid-August 1991-1993, three exposed branches each from the upper- and mid- to lower-crown of each tree were collected and evaluated for ozone injury. Of the trees examined, 63%, 52% and 36% exhibited some amount of foliar injury in 1991, 1992 and 1993, respectively. Ozone injury across species was the greatest at Cove Mountain in all three years of the study. Overall, across sites and years, 11, 12 and 11% of all leaves examined exhibited visible injury for black cherry, sassfras and yellow-poplar, respectively. The percentage of injured leaves per branch was greater in the mid- to lower-canopy for black cherry, across all sites. Trees for each species that exhibited the greatest or least amounts of visible injury did so in all three years of the study, indicating a differential sensitivity within each species population. No significant ozone exposure-tree response relationships were observed with any variable tested. These data indicate that ozone concentrations are high enough to cause visible symptoms to selected trees within Great Smoky Mountains National Park, USA.

Cutleaf coneflower (Rudbeckia laciniata L.) response to ozone and ethylenediurea (EDU)

Cutleaf coneflower (Rudbeckia laciniata L.) seedlings were placed into open-top chambers in May, 2004 and fumigated for 12 wks. Nine chambers were fumigated with either carbon-filtered air (CF), non-filtered air (NF) or twice-ambient (2x) ozone (O(3)). Ethylenediurea (EDU) was applied as a foliar spray weekly at 0 (control), 200, 400 or 600 ppm. Foliar injury occurred at ambient (30%) and elevated O(3) (100%). Elevated O(3) resulted in significant decreases in biomass and nutritive quality. Ethylenediurea reduced percent of leaves injured, but decreased root and total biomass. Foliar concentrations of cell-wall constituents were not affected by EDU alone; however, EDUxO(3) interactions were observed for total cell-wall constituents and lignocellulose fraction. Our results demonstrated that O(3) altered the physiology and productivity of cutleaf coneflower, and although reducing visible injury EDU may be phytotoxic at higher concentrations.

Ozone impacts on loblolly pine (Pinus taeda L.) grown in a competitive environment.

One-year old, bare-root, loblolly pine (Pinus taeda L.) seedlings grown in an early successional plant community, were exposed to four ozone treatments, 12 h day(-1) (09:00-21:59 h CST), 7 day week(-1) in east-central Alabama for 2 years (1994-1995). Treatments included AA=ambient air (open plot; no chamber), CF = carbon-filtered air (approximately 0.5xAA), 1X = non-filtered air, and 2X = twice ambient (AA) air. Pine height and diameter were measured monthly for 2 years and total above-ground biomass determined at termination of the experiment. In addition, height, diameter and volume (d2h) growth was modeled for the 2-year study. Trees growing in the CF chambers exhibited the least growth for any variable measured. The predicted height growth rate (r) was the least for the CF- and 2X-treated pines when compared with the AA and 1X treatments. In addition, percent total vegetative cover was significantly negatively correlated with height. diameter and volume growth in 1994, but not 1995. Our findings regarding the trees growing in the CF treatments, are contrary to previous reports regarding ozone impacts to loblolly pine grown without competition from other vegetation.

Environmental control of stomatal conductance in forest trees of the Great Smoky Mountains National Park

To determine if stomatal conductance (g(s)) of forest trees could be predicted from measures of leaf microclimate, diurnal variability in in situ g(s) was measured in black cherry (Prunus serotina), red maple (Acer rubrum), and northern red oak (Quercus rubra). Relative to overstory trees, understory saplings exhibited little diurnal variability in g(s) and ozone uptake. Depending on species and site, up to 30% of diurnal and seasonal variation in g(s )of overstory trees was explained by photosynthetically active radiation and vapor pressure deficit. Daily maximum g(s) was significantly related to soil moisture in overstory northern red oak and black cherry (R(2) ranged from 33 to 65%). Although g(s) was not fully predicted using instantaneous measures of leaf microclimate, ozone uptake of large forest trees was reduced by low soil moisture.