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.     

Physiological and foliar symptom response in the crowns of Prunus serotina, Fraxinus americana and Acer rubrum canopy trees to ambient ozone under forest conditions

The crowns of five canopy dominant black cherry (Prunus serotina Ehrh.), five white ash (Fraxinus americana L.), and six red maple (Acer rubrum L.) trees on naturally differing environmental conditions were accessed with scaffold towers within a mixed hardwood forest stand in central Pennsylvania. Ambient ozone concentrations, meteorological parameters, leaf gas exchange and leaf water potential were measured at the sites during the growing seasons of 1998 and 1999. Visible ozone-induced foliar injury was assessed on leaves within the upper and lower crown branches of each tree. Ambient ozone exposures were sufficient to induce typical symptoms on cherry (0-5% total affected leaf area, LAA), whereas foliar injury was not observed on ash or maple. There was a positive correlation between increasing cumulative ozone uptake (U) and increasing percent of LAA for cherry grown under drier site conditions. The lower crown leaves of cherry showed more severe foliar injury than the upper crown leaves. No significant differences in predawn leaf water potential (psi(L)) were detected for all three species indicating no differing soil moisture conditions across the sites. Significant variation in stomatal conductance for water vapor (g(wv)) was found among species, soil moisture, time of day and sample date. When comparing cumulative ozone uptake and decreased photosynthetic activity (P(n)), red maple was the only species to show higher gas exchange under mesic vs. drier soil conditions (P < 0.05). The inconsistent differences in gas exchange response within the same crowns of ash and the uncoupling relationship between g(wv) and P(n) demonstrate the strong influence of heterogeneous environmental conditions within forest canopies.     

Air pollution, precipitation chemistry and forest health in the Retezat Mountains, Southern Carpathians, Romania

In the Retezat Mountains concentrations of O3, NO2 and SO2 in summer season 2000-2002 were low and below toxicity levels for forest trees. While NH3 concentrations were low in 2000, the 2001 and 2002 concentrations were elevated indicating possibility for increased N deposition to forest stands. More than 90% of the rain events were acidic with pH values <5.5, contributing to increased acidity of soils. Crown condition of Norway spruce (Picea abies) and European beech (Fagus sylvatica) was good, however, defoliation described as >25% of foliage injured increased from 9.1% in 2000 to 16.1% in 2002. Drought that occurred in the southern Carpathians between fall 2000 and summer 2002 and frequent acidic rainfalls could cause the observed decline of forest condition. Both Norway spruce and European beech with higher defoliation had lower annual radial increments compared to the trees with low defoliation. Ambient O3 levels found in the Retezat did not affect crown condition of Norway spruce or European beech.     

An evaluation of ozone exposure metrics for a seasonally drought-stressed ponderosa pine ecosystem

Ozone stress has become an increasingly significant factor in cases of forest decline reported throughout the world. Current metrics to estimate ozone exposure for forest trees are derived from atmospheric concentrations and assume that the forest is physiologically active at all times of the growing season. This may be inaccurate in regions with a Mediterranean climate, such as California and the Pacific Northwest, where peak physiological activity occurs early in the season to take advantage of high soil moisture and does not correspond to peak ozone concentrations. It may also misrepresent ecosystems experiencing non-average climate conditions such as drought years. We compared direct measurements of ozone flux into a ponderosa pine canopy with a suite of the most common ozone exposure metrics to determine which best correlated with actual ozone uptake by the forest. Of the metrics we assessed, SUM0 (the sum of all daytime ozone concentrations > 0) best corresponded to ozone uptake by ponderosa pine, however the correlation was only strong at times when the stomata were unconstrained by site moisture conditions. In the early growing season (May and June). SUM0 was an adequate metric for forest ozone exposure. Later in the season, when stomatal conductance was limited by drought. SUM0 overestimated ozone uptake. A better metric for seasonally drought-stressed forests would be one that incorporates forest physiological activity, either through mechanistic modeling, by weighting ozone concentrations by stomatal conductance, or by weighting concentrations by site moisture conditions.     

Forest health status in the Carpathian Mountains over the period 1997-2001

The results of forest health status assessments in the Carpathian Mountains from the monitoring networks developed by the European Union Scheme on the Protection of Forest Against Atmospheric Pollution (EU Scheme) and International Co-operative Programme on Assessment and Monitoring of Air Pollution Effects on Forests (ICP-Forests), have led to a better understanding of the impact of air pollution and other stressors on forests at the regional scale. During the period 1997-2001, forests in the Carpathian Mountains were severely affected by air pollution and natural stresses with 29.7-34.9% of the trees included in defoliation classes 2-4. The broadleaves were slightly healthier than the conifers, and European beech (Fagus sylvatica) was the least affected species. Norway spruce (Picea abies) has poor health status, with 42.9-46.6% of the trees damaged (2-4% defoliation classes). Silver fir (Abies alba) damage was also high, with 46.0-50.9% in defoliation classes 2-4. Pines (primarily Pinus sylvestris) were the least affected of the conifers, with 24.9-33.8% in defoliation classes 2-4. The results from the transnational networks (16 x 16 km) show that the Carpathian forests are slightly more damaged than the average for the entire Europe. The correlative studies performed in individual European countries show the relationships between air pollution stressors with trends in defoliation and a possible effect of natural stresses at each site. More specific, effects of tree age, drought, ozone and acid deposition critical level exceedances were demonstrated to affect crown condition.     

Meteorological factors and air pollution in Lithuanian forests: possible effects on tree condition

This study investigates changes in tree condition and environmental factors in Lithuania during the active growing season in 1991-2001. The average crown defoliation and the proportion of healthy trees of Pinus sylvestris, Picea abies, Betula sp., Fraxinus excelsior, Alnus incana, Alnus glutinosa, Populus tremula, and Quercus robur, meteorological (average temperature, amount of precipitation, hydrothermal coefficient) and air pollution data (acidity of precipitation, concentrations of SO2, NO2 and exposure of O3) were analysed. During the period 1991-2001 the condition of Pinus sylvestris, Populus tremula showed a tendency of improvement, while defoliation of Fraxinus excelsior significantly increased. The proportion of healthy trees correlated well with the average temperature and O3 (AOT40), while defoliation correlated well with the acidity of precipitation and the concentrations of SO2 and NO2. Deciduous species appeared to be more sensitive to O3 exposure and conifers to the concentrations of SO2 and NO2.     

Ozone and forests in South-Western Europe--What have we learned?

Monitoring of forest condition and ozone (O3) at 83 sites in France, Italy, Luxenbourg, Spain and Switzerland resulted in a number of findings in relation to the knowledge of O3 exposure (concentration and cumulative AOT40), feasibility of the assessment of stomatal O3 flux and relationships between O3 and crown defoliation of beech and visible symptoms on native vegetation. However, the project provides evidence of issues to be addressed within the current monitoring system (data quality, validation sites and response indicators) and indications as to how the monitoring of O3 risk in the context of an effect-oriented monitoring program can be improved.     

Size-mediated foliar response to ozone in black cherry trees

Local ozone concentration and visible foliar injury were measured over the 1994 growing season on open-grown black cherry (Prunus serotina Ehrh.) trees of varying size (age) within forest stands and adjacent openings at a site in north-central Pennsylvania. Relationships were determined between visible ozone injury and ozone exposure, as well as calculated between injury and ozone uptake expressed as the product of stomatal conductance and ozone concentration. In addition, simultaneous measurements of visible symptoms and leaf gas exchange were also conducted to determine the correlation between visible and physiological injury and ozone exposure. By September, the amount of leaf area affected by visible foliar ozone injury was greatest for seedlings (46%), followed by canopy trees (20%) and saplings (15%). A large amount of variability in foliar ozone symptom expression was observed among trees within a size class. Sum40 and Sum60 (ozone concentration > 40 and > 60 nl liter(-1)) cumulative exposure statistics were the most meaningful indices for interpretation of foliar injury response. Seedlings were apparently more sensitive to ozone injury than larger trees because their higher rates of stomatal conductance resulted in higher rates of ozone uptake. Seedlings also had higher rates of early leaf abscission than larger trees with an average of nearly 30% of the leaves on a shoot abscised by 1 September compared to approximately 5% for larger trees. However, per unit ozone uptake into the leaf, larger trees exhibited larger amounts of foliar injury. The amount of visible foliar injury was negatively correlated (r(2) = 0.82) with net photosynthetic rates, but was not related to stomatal conductance. Net photosynthesis and stomatal conductance thus became uncoupled at high levels of visible foliar injury.     

Scots pine (Pinus sylvestris L.) growth and condition in a polluted environment: from decline to recovery

The results of long-term investigations of Scots pine (Pinus sylvestris L.) growth and condition in the impact zone of one of the biggest air pollution sources in Lithuania--mineral fertilizers plant "Achema" are presented. The main attention is laid to the recovery of damaged stands since annual emissions to air were reduced essentially. The investigations indicated, that the recovery of tree increment was mostly caused by distinct reduction of emissions of nitrogen and sulphur oxides and dust of mineral fertilizers. Despite reduced pollution, crown defoliation of investigated stands has continued to increase for a certain period. After the crown recovery of damaged stands has started, the recovery of most damaged survived trees was most intensive and convergence of defoliated to a different extend stands and trees is characteristic feature of this period. No defoliation threshold has been determined beyond of which recovery of trees would be impossible. Recovery of more than a quarter of damaged trees was registered even in the case of 90% of defoliation. Recovery of dominant trees occurs to be faster of that for suppressed trees within the same level of defoliation. The impact of stand density on the crown recovery rate is negative, the higher density (more intensive competition), the slower recovery of damaged trees. The dependence of growth rate on defoliation was found to be of logistic character: while crown defoliation consists up to 25-30%, tree increment losses are rather inconsiderable, further increase of defoliation leads to the essentially higher increment losses, however having achieved 65-70% defoliation, further increase of defoliation does not result such intensive decrease of radial increment.     

Ozone fluxes above and within a pine forest canopy in dry and wet conditions

The physiological and physical processes controlling ozone dry deposition to vegetated surfaces are still not fully understood. In particular, the role of the understorey and the possible action of dew on ozone deposition have not received much attention so far. This paper presents the results of an experiment aimed at quantifying ozone dry deposition to a maritime pine forest in the “Les Landes” area in France. Ozone deposition fluxes were measured using the eddy-covariance technique above and within the canopy. We investigate the factors acting on ozone deposition in both dew-wetted and dry conditions. The values obtained for the ozone deposition velocity are well in the range of previously published measurements over coniferous forests. For the present forest, ozone uptake by the understorey is a significant portion of ozone deposition to the whole pine stand. The understorey contributes more to the overall ozone flux than to the other measured scalar fluxes (sensible heat and water vapour). During dry nights the surface conductance for ozone and the friction velocity are strongly correlated, showing that ozone deposition is largely controlled by dynamical processes. During the day, in dry conditions, the canopy stomatal conductance is the major parameter controlling ozone deposition. However, in winter, when the stomatal conductance is low, the influence of dynamical processes persists during day-time. It is also found that surface wetness associated with dew significantly enhances ozone deposition, during the night as well as in the morning.     

Contribution of ambient ozone to Scots pine defoliation and reduced growth in the Central European forests: a Lithuanian case study

The study aimed to explore if changes in crown defoliation and stem growth of Scots pines (Pinus sylvestris L.) could be related to changes in ambient ozone (O₃) concentration in central Europe. To meet this objective the study was performed in 3 Lithuanian national parks, close to the ICP integrated monitoring stations from which data on meteorology and pollution were provided. Contribution of peak O₃ concentrations to the integrated impact of acidifying compounds and meteorological parameters on pine stem growth was found to be more significant than its contribution to the integrated impact of acidifying compounds and meteorological parameters on pine defoliation. Findings of the study provide statistical evidence that peak concentrations of ambient O₃ can have a negative impact on pine tree crown defoliation and stem growth reduction under field conditions in central and northeastern Europe where the AOT40 values for forests are commonly below their phytotoxic levels.     

Ozone exposure, defoliation of beech (Fagus sylvatica L.) and visible foliar symptoms on native plants in selected plots of South-Western Europe

The relationships between crown defoliation of beech, visible foliar symptoms on native vegetation and ozone exposure were investigated on permanent monitoring sites in South-Western Europe in the years 2000-2002. Relationships between defoliation of beech and O3 (seasonal mean, 2-week maximum, AOT40) were investigated by means of multiple regression models (11 plots, 1-3 years of data each) and a model based on temporal autocorrelation of defoliation data (14 plots, 1-3 years of data each). Different multiple regression techniques were used. The four models generated (R2=0.71-0.85, explained variance in cross-validation 61-78%) identified several significant predictors of defoliation, with AOT40 (p=0.008) and foliar content of phosphorous (p=0.0002-0.0004) being common to all models. The autocorrelation model (R2=0.55; p<0.0001) was used to calculate expected defoliation on the basis of the previous year's defoliation, and model predictions were used as an estimate of expected defoliation under constant site and environmental condition. Residuals (predicted-measured) plotted against current AOT40 shows that a possible effect of ozone occurs only at very high AOT40 (>35,000 ppbh). O3-like visible foliar symptoms were recorded on 65 species at 47% of the common monitoring sites in 2001 and 38% in 2002. No relationship was found between O3 exposure, frequency of symptomatic sites and frequency of species with symptoms (R2=0.11; p>0.05). A number of questions related to the ecological and methodological basis of the survey were identified. Inherent sampling and non-sampling errors and multicollinearity of the data suggest great caution when examining results obtained from mensurational, correlative studies.     

Limitations and perspectives about scaling ozone impacts in trees.

We review the need for scaling effects of ozone (O3) from juvenile to mature forest trees, identify the knowledge presently available, and discuss limitations in scaling efforts. Recent findings on O3/soil nutrient and O3/CO2 interactions from controlled experiments suggest consistent scaling patterns for physiological responses of individual leaves to whole-plant growth, carbon allocation, and water use efficiency of juvenile trees. These findings on juvenile trees are used to develop hypotheses that are relevant to scaling O3 effects to mature trees, and these hypotheses are examined with respect to existing research on differences in response to O3 between juvenile and mature trees. Scaling patterns of leaf-level physiological response to O3 have not been consistent in previous comparisons between juvenile and mature trees. We review and synthesize current understanding of factors that may cause such inconsistent scaling patterns, including tree-size related changes in environment, stomatal conductance, O3 uptake and exposure. carbon allocation to defense, repair, and compensation mechanisms, and leaf production phenology. These factors should be considered in efforts to scale O3 responses during tree ontogeny. Free-air O3 fumigation experiments of forest canopies allow direct assessments of O3 impacts on physiological processes of mature trees, and provide the opportunity to test current hypotheses about ontogenetic variation in O3 sensitivity by comparing O3 responses across tree-internal scales and ontogeny.     

Crown condition of Norway spruce in relation to sulphur and nitrogen deposition and soil properties in southeast Norway

Atmospheric deposition of sulphur and nitrogen compounds may lead to enhanced leaching of base cations, accumulation of nitrogen in organic matter, lowered pH and increased concentration of toxic aluminium in soil, which in turn may affect the vitality of forest trees. A general monitoring of forest condition has been initiated in many European countries, partly in order to reveal stresses caused by acidification. However, forest condition is also affected by many other factors. This paper examines a seven-year series of crown-condition data from Local County Monitoring Plots in Norway spruce stands in Norway. Average, time trend and lability variables were calculated for crown density and crown colour for each plot. Wet deposition of sulphate, ammonium and nitrate for each plot were estimated using data from the national air and precipitation monitoring programmes. Soil data are based on soil sampling within the plots. The analysis gave no evident support for the hypothesized negative effect on crown condition from sulphur and nitrogen deposition and related alterations in soil.     

Modelling stomatal ozone flux across Europe

A model has been developed to estimate stomatal ozone flux across Europe for a number of important species. An initial application of this model is illustrated for two species, wheat and beech. The model calculates ozone flux using European Monitoring and Evaluation Programme (EMEP) model ozone concentrations in combination with estimates of the atmospheric, boundary layer and stomatal resistances to ozone transfer. The model simulates the effect of phenology, irradiance, temperature, vapour pressure deficit and soil moisture deficit on stomatal conductance. These species-specific microclimatic parameters are derived from meteorological data provided by the Norwegian Meteorological Institute (DNMI), together with detailed land-use and soil type maps assembled at the Stockholm Environment Institute (SEI). Modelled fluxes are presented as mean monthly flux maps and compared with maps describing equivalent values of AOT40 (accumulated exposure over threshold of 40 ppb or nl l(-1)), highlighting the spatial differences between these two indices. In many cases high ozone fluxes were modelled in association with only moderate AOT40 values. The factors most important in limiting ozone uptake under the model assumptions were vapour pressure deficit (VPD), soil moisture deficit (for Mediterranean regions in particular) and phenology. The limiting effect of VPD on ozone uptake was especially apparent, since high VPDs resulting in stomatal closure tended to co-occur with high ozone concentrations. Although further work is needed to link the ozone uptake and deposition model components, and to validate the model with field measurements, the present results give a clear indication of the possible implications of adopting a flux-based approach for future policy evaluation.     

Defense and avoidance of ozone under global change

The level II approach of the critical loads concept adopted by the UNECE aims at a flux based evaluation and takes into account environmental factors governing stomatal conductance. These factors will probably be affected by global change. The flux concept predicts that a decrease in stomatal conductance would protect trees from air pollution effects by decreasing uptake. However, experimental evidence is inconclusive. Numerous results suggest that pollutants and factors subject to global change (drought, CO(2)) may interact and even exacerbate effects, probably because antioxidative defense systems are involved in both, defense against pollutant effects and protection from natural stress. An effective pollutant dose, which is weighted by physiological defense capacity, would better predict such effects. In this review paper we argue that the flux-based approach is imperfect, because global change effects may also modify the physiological susceptibility to ozone. Instead, a flux concept weighted by defense capacity should be tested.     

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.     

Ozone uptake (flux) as it relates to ozone-induced foliar symptoms of Prunus serotina and Populus maximowiziixtrichocarpa

Field studies were conducted during 2003 and 2004 from early June to the end of August, at 20 sites of lower or higher elevation within north-central Pennsylvania, using seedlings of black cherry (Prunus serotina, Ehrh.) and ramets of hybrid poplar (Populus maximowiziixtrichocarpa). A linear model was developed to estimate the influence of local environmental conditions on stomatal conductance. The most significant factors explaining stomatal variance were tree species, air temperature, leaf vapor pressure deficit, elevation, and time of day. Overall, environmental factors explained less than 35% of the variation in stomatal conductance. Ozone did not affect gas exchange rates in either poplar or cherry. Ozone-induced foliar injury was positively correlated with cumulative ozone exposures, expressed as SUM40. Overall, the amount of foliar injury was better correlated to a flux-based approach rather than to an exposure-based approach. More severe foliar injuries were observed on plants growing at higher elevations.     

Simulations of stomatal conductance and ozone uptake to Norway spruce saplings in open-top chambers

A simulation model was developed to estimate the stomatal conductance and ozone flux to Norway spruce saplings in open-top chambers. The model was parameterized against needle conductance measurements that were made on 4-6-year-old spruce saplings, grown in open-top chambers, in July-September during three different seasons. The spruce saplings were either maintained well watered or subject to a 7-8 week drought period in July-September each year. The simulated conductance showed a good agreement with the measured conductance for the well-watered as well as the drought stress-treated saplings. The simulations were significantly improved when different vapour pressure deficit (VPD) functions were applied for well-watered and drought-stressed spruce saplings. The cumulated ozone uptake which was calculated from the conductance simulations showed less variation between years, compared to the cumulative ozone exposure index AOT40 (accumulated exposure over a threshold of 40 ppb or nl l(-1)) for the corresponding time periods. Measurements in May 1995 demonstrated the occurrence of long-term 'memory-effects' from the drought stress treatments on the conductance. Memory-effects need to be considered when simulation models for stomatal conductance are to be applied to long-lived forest trees under a multiple stress situation.     

Seasonal differences and within-canopy variations of antioxidants in mature spruce (Picea abies) trees under elevated ozone in a free-air exposure system

The effect of free-air ozone fumigation and crown position on antioxidants were determined in old-growth spruce (Picea abies) trees in the seasonal course of two consecutive years (2003 and 2004). Levels of total ascorbate and its redox state in the apoplastic washing fluid (AWF) were increased under double ambient ozone concentrations (2xO3), whilst ascorbate concentrations in needle extracts were unchanged. Concentrations of apoplastic and symplastic ascorbate were significantly higher in 2003 compared to 2004 indicating a combined effect of the drought conditions in 2003 with enhanced ozone exposure. Elevated ozone had only weak effects on total glutathione levels in needle extracts, phloem exudates and xylem saps. Total and oxidised glutathione concentrations were higher in 2004 compared to 2003 and seemed to be more affected by enhanced ozone influx in the more humid year 2004 compared to the combined effect of elevated ozone and drought in 2003 as observed for ascorbate.