J.E. Savage

Observation and Confirmation of Foliar Ozone Symptoms of Native Plant Species of Switzerland and Southern Spain

Tropospheric ozone is considered as the major pollutant of concern to the health and productivity of forests in the eastern United States and has more recently become of increasing concern within the forests of southern Europe. Recent observations have clearly demonstrated foliar injury symptoms to be occurring on many tree and native plant species within remote forested areas. Several plant genera (and a few species within genera) found in both the forests of Switzerland and the southern coastal region of Spain exhibit field symptoms typical of ambient ozone exposures. Ozone exposures for many species have been conducted under controlled CSTR conditions and within open-top chambers within the study areas. Results have confirmed that the O3-like foliar symptoms as observed under natural forest and open grown conditions for many native tree, shrub, and herbaceous species in Spain and Switzerland are caused by exposures to ambient O3.

Physiology, morphology, and ozone uptake of leaves of black cherry seedlings, saplings, and canopy trees.

Patterns of ozone uptake were related to physiological, morphological, and phenological characteristics of different-sized black cherry trees (Prunus serotina Ehrh.) at a site in central Pennsylvania. Calculated ozone uptake differed among open-grown seedlings, forest gap saplings, and canopy trees and between leaves in the upper and lower crown of saplings and canopy trees. On an instantaneous basis, seedling leaves had the greatest ozone uptake rates of all tree size classes due to greater stomatal conductance and higher concentrations of ozone in their local environment. A pattern of higher stomatal conductance of seedlings was consistent with higher incident photosynthetically-active radiation, stomatal density, and predawn xylem water potentials for seedlings relative to larger trees. However, seedlings displayed an indeterminate pattern of shoot growth, with the majority of their leaves produced after shoot growth had ceased for canopy and sapling trees. Full leaf expansion occurred by mid-June for sapling and canopy trees. Because many of their leaves were exposed to ozone for only part of the growing season, seedlings had a lower relative exposure over the course of the growing season, and subsequently lower cumulative uptake, of ozone than canopy trees and a level of uptake similar to upper canopy leaves of saplings. Visible injury symptoms were not always correlated with patterns in ozone uptake. Visible symptoms were more apparent on seedling leaves in concurrence with their high instantaneous uptake rates. However, visible injury was more prevalent on leaves in the lower versus upper crown of canopy trees and saplings, even though lower crown leaves had less ozone uptake. Lower crown leaves may be more sensitive to ozone per unit uptake than upper crown leaves because of their morphology. In addition, the lower net carbon uptake of lower crown leaves may limit repair and anti-oxidant defense processes.

Response of native plants of northeastern United States and southern Spain to ozone exposures; determining exposure/response relationships

Tropospheric ozone has been identified as the most important regional scale air pollutant across much of eastern United States of America and many areas of Mediterranean climes in southern Europe. Recent field surveys in the northeastern USA and in southeastern Spain have revealed many additional plant species that exhibit symptoms typical of ozone-induced injuries. Objectives of this study were to confirm ozone as the cause of the observed foliar symptoms, determine ozone induced exposure/response relationships, and identify possible bio-indicator species. Thirteen native species of northeastern USA and 27 native species of southeastern Spain were selected for study. Plant species were exposed to ozone within 16 CSTR chambers in a greenhouse during the summer seasons of 2000 and 2001; ozone exposures of 30, 60, 90, and 120 ppb were delivered for 7 h/day, 5 days/week. Results have confirmed that with few exceptions, symptoms observed in the field were induced by exposures to ambient ozone. Species differed significantly in terms of the exposures required for the initiation of visible symptoms and subsequent injury progression.

Physiological and foliar injury responses of Prunus serotina, Fraxinus americana, and Acer rubrum seedlings to varying soil moisture and ozone

Sixteen black cherry (Prunus serotina, Ehrh.), 10 white ash (Fraxinus americana, L.) and 10 red maple (Acer rubrum, L.) 1-year old seedlings were planted per plot in 1997 on a former nursery bed within 12 open-top chambers and six open plots. Seedlings were exposed to three different ozone scenarios (ambient air: 100% O3; non-filtered air: 98% ambient O3; charcoal-filtered air: 50% ambient O3) within each of two different water regimes (nine plots irrigated, nine plots non-irrigated) during three growing seasons. During the 1998 and 1999 growing season, leaf gas exchange, plant water relations, and foliar injury were measured. Climatic data,ambient- and chamber-ozone-concentrations were monitored. We found that seedlings grown under irrigated conditions had similar (in 1998) but significantly higher gas exchange rates (in 1999) than seedlings grown within non-irrigated plots among similar ozone exposures. Cherry and ash had similar ozone uptake but cherry developed more ozone-induced injury (< 34% affected leaf area, LAA) than ash (<5% LAA), while maple rarely showed foliar injury, indicating the species differed in ozone sensitivity. Significantly more severe injury on seedlings grown under irrigated conditions than seedlings grown under non-irrigated conditions demonstrated that soil moisture altered seedling responses to ambient ozone exposures.

Use of bioindicators and passive sampling devices to evaluate ambient ozone concentrations in north central Pennsylvania

Ambient concentrations of tropospheric ozone and ozone-induced injury to black cherry (Prunus serotina) and common milkweed (Asclepias syriaca) were determined in north central Pennsylvania from 29 May to 5 September 2000 and from 28 May to 18 September 2001. Ogawa passive ozone samplers were utilized within openings at 15 forested sites of which six were co-located with TECO model 49 continuous ozone monitors. A significant positive correlation was observed between the Ogawa passive samplers and the TECO model 49 continuous ozone monitors for the 2000 (r=0.959) and 2001 (r=0.979) seasons. In addition, a significant positive correlation existed in 2000 and 2001 between ozone concentration and elevation (r=0.720) and (r=0.802), respectively. Classic ozone-induced symptoms were observed on black cherry and common milkweed. In 2000, initial injury was observed in early June, whereas for the 2001 season, initial injury was initially observed in late June. During both seasons, injury was noted at most sites by mid- to late-July. Soil moisture potential was measured for the 2001 season and a significant positive relationship (P<0.001) showed that injury to black cherry was a function of cumulative ozone concentrations and available soil moisture.

Foliar response of black cherry (Prunus serotina) clones to ambient ozone exposure in central Pennsylvania

During late summer of 1996 and 1997 we examined ozone-induced foliar injury in a plantation of 111 black cherry trees (ramets) comprising 15 clones originating from wild ortets growing in the Allegheny National Forest, Pennsylvania, and the Monongahela National Forest, West Virginia. The experimental plantation was a clonal seed orchard in Centre County, Pennsylvania, started in 1971 using ortet buds grafted onto seedling rootstocks of mixed origin. Clones differed significantly in severity of foliar injury symptoms (F=31.83, p<0.001). One clone (R-12) had significantly more foliar injury with >50% leaf area affected than other clones during both years. In contrast, clone R-14, which is from the same area in northcentral Pennsylvania as R-12, exhibited significantly less injury (LAA<6%). Although ambient O(3) concentrations were similar in both years, foliar injury was significantly greater (15.7%) in 1996 than in 1997 (9.9%). This is probably explained by lower stomatal conductance in 1997 caused by drier and hotter weather patterns in June and July of that year. Despite very different weather patterns and overall levels of injury in 1996 and 1997, mean clonal injury was significantly correlated between both years of assessment (r=0.92, p<0.001). Within tree crowns, foliage in lower and inner crown positions was significantly more injured than foliage in upper and exterior crown positions. There was no evidence of geographically based population differences in sensitivity to foliar O(3) injury. On the contrary, results demonstrate that wild genotypes of proximal geographic origin may differ greatly in sensitivity.

Observations of ozone-induced foliar injury on black cherry (Prunus serotina, var. Capuli) within the Desierto de los Leones National Park, Mexico City

A survey for ozone-induced foliar injury of black cherry was conducted in mid-June 1995 within the Desierto de Los Leones National Park located southwest of Mexico City. Evaluations of the upper and lower tree crowns of 18 trees revealed evidence of significant upper surface stipple, leaf reddening and premature senescence on 72% of the trees. A general survey of an additional 169 trees disclosed that 41% exhibited similar symptoms. A gradient of increasing symptoms with increasing elevation was also evident. For the most part, asymptomatic trees were observed to be situated within well-shaded coves at the lower elevations with very few symptomatic trees present in these areas.

Vertical gradients of ozone and carbon dioxide within a deciduous forest in Central Pennsylvania.

Ambient concentrations of ozone (O(3)) and carbon dioxide (CO(2)) were measured at locations from the forest floor to the top of the canopy in a deciduous forest at the Moshannon State Forest in northcentral Pennsylvania. O(3) concentrations were measured from May-September for three years (1993-1995) while CO(2) concentrations were measured only during July and August of 1994. O(3) concentrations increased steadily during the day at all locations, peaking during the middle to late afternoon hours. O(3) concentrations then steadily declined to their lowest point, just before dawn. Vertical O(3) concentration gradients varied seasonally and among years. However, O(3) concentrations were highest within the forest canopy and lowest at the forest floor, with an average difference of approximately 13%. Differences in O(3) concentrations between the canopy and forest floor were greatest at night. O(3) concentrations were slightly higher at locations within the canopy than above the canopy. CO(2) concentrations were consistenly higher near the forest floor and were higher above the canopy than within the canopy. CO(2) concentrations were higher at night than during the day at all locations, especially near the forest floor.

Leaf age affects the responses of foliar injury and gas exchange to tropospheric ozone in Prunus serotina seedlings

We investigated the effect of leaf age on the response of net photosynthesis (A), stomatal conductance (g(wv)), foliar injury, and leaf nitrogen concentration (N(L)) to tropospheric ozone (O(3)) on Prunus serotina seedlings grown in open-plots (AA) and open-top chambers, supplied with either carbon-filtered or non-filtered air. We found significant variation in A, g(wv), foliar injury, and N(L) (P < 0.05) among O(3) treatments. Seedlings in AA showed the highest A and g(wv) due to relatively low vapor pressure deficit (VPD). Older leaves showed significantly lower A, g(wv), N(L), and higher foliar injury (P < 0.001) than younger leaves. Leaf age affected the response of A, g(wv), and foliar injury to O(3). Both VPD and N(L) had a strong influence on leaf gas exchange. Foliar O(3)-induced injury appeared when cumulative O(3) uptake reached 8-12 mmol m(-2), depending on soil water availability. The mechanistic assessment of O(3)-induced injury is a valuable approach for a biologically relevant O(3) risk assessment for forest trees.

Ozone Sensitivity of 28 Plant Selections Exposed to Ozone Under Controlled Conditions

Abstract Ambient, ground-level ozone is the most important air pollutant affecting vegetation in the US. However, ozone-sensitive bioindicators need to be identified for use in field surveys to detect ozone-induced symptoms. To identify such bioindicators, 28 plant selections were exposed to ozone within greenhouse chambers during 2003 and 2004. Plants most sensitive to ozone included Asclepias incarnata (swamp milkweed), Asclepias syriaca (common milkweed), Cephalanthus occidentalis (buttonbush), Platanus occidentalis (American sycamore), Salix × cotteti (Bankers dwarf willow), Salix lucida (shining willow), Salix nigra (black willow), Salix sericea (silky willow), and Symphoricarpos albus (snowberry). Plants moderately sensitive included Aster novaeangliae (New England aster), Monarda didyma (bee-balm), Rhus aromatica (aromatic sumac), Salix discolor (pussy willow), Salix exigua (sandbar willow), Salix purpurea (basket willow), Sambucus ebulus (European dwarf elderberry), and Symphoricarpos spp. (mixture of “snowberries”). Plants more tolerant to ozone included Aster macrophyllus (bigleaf aster), Aster novibelgii (New York aster), Cercis canadensis (redbud), Populus maximowizii × trichocarpa (hybrid poplar), Rudbeckia laciniata (cutleaf, or tall green-headed coneflower), Salix amygdaloides (peach-leaved willow), Salix eriocephala (diamond willow), Sambucus canadensis (American elder), Sambucus nigra (European elder), Symphoricarpos orbiculatus (coralberry), and Viburnum trilobum (highbush-cranberry). The incidence and severity of ozone-induced symptoms varied with species, ozone concentration, and length of exposure. Pigmented adaxial leaf surface stipple, bifacial necrosis, and premature defoliation were common foliar symptoms induced by ozone; leaf reddening, bronzing, and chlorosis occurred less frequently. Species that produced classic ozone-induced stipple in this study may serve as useful bioindicators in field surveys to confirm that ozone injury occurs in many parts of the country. Such findings may have a positive impact on establishing more stringent air-quality standards to protect vegetation and the environment.