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 (O₃) 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 O₃ 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 O₃. Both VPD and N_L had a strong influence on leaf gas exchange. Foliar O₃-induced injury appeared when cumulative O₃ uptake reached 8-12 mmol m⁻², depending on soil water availability. The mechanistic assessment of O₃-induced injury is a valuable approach for a biologically relevant O₃ risk assessment for forest trees.     

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.     

Responses of hybrid poplar clones and red maple seedlings to ambient O(3) under differing light within a mixed hardwood forest

The responses of ramets of hybrid poplar (Populus spp.) (HP) clones NE388 and NE359, and seedlings of red maple (Acer rubrum, L.) to ambient ozone (O(3)) were studied during May-September of 2000 and 2001 under natural forest conditions and differing natural sunlight exposures (sun, partial shade and full shade). Ambient O(3) concentrations at the study site reached hourly peaks of 109 and 98 ppb in 2000 and 2001, respectively. Monthly 12-h average O(3) concentrations ranged from 32.3 to 52.9 ppb. Weekly 12-h average photosynthetically active radiation (PAR) within the sun, partial shade and full shade plots ranged from 200 to 750, 50 to 180, and 25 to 75 micromol m(-2) s(-1), respectively. Ambient O(3) exposure induced visible foliar symptoms on HP NE388 and NE359 in both growing seasons, with more severe injury observed on NE388 than on NE359. Slight foliar symptoms were observed on red maple seedlings during the 2001 growing season. Percentage of total leaf area affected (%LAA) was positively correlated with cumulative O(3) exposures. More severe foliar injury was observed on plants grown within the full shade and partial shade plots than those observed on plants grown within the sun plot. Lower light availability within the partial shade and full shade plots significantly decreased net photosynthetic rate (Pn) and stomatal conductance (g(wv)). The reductions in Pn were greater than reductions in g(wv), which resulted in greater O(3) uptake per unit Pn in plants grown within the partial shade and full shade plots. Greater O(3) uptake per unit Pn was consistently associated with more severe visible foliar injury in all species and/or clones regardless of differences in shade tolerance. These studies suggest that plant physiological responses to O(3) exposure are likely complicated due to multiple factors under natural forest conditions.     

Effects of acidic fog on seedlings of Pinus ponderosa and Abies concolor: foliar injury, physiological and biochemical responses

Seedlings of Pinus ponderosa (ponderosa pine) and Abies concolor (white fir) were exposed to acidic fog (pH 2.0, 3.0 or 4.0) in open-field plots for six weeks. The two species exhibited dissimilar injury responses; neither current year nor previous year needles of ponderosa pine were injured by pH 2.0 fog, but current year needles exhibited higher membrane permeability responses (i.e. needle extract conductivity, K+ concentration). In comparison, both needle age classes in white fir were significantly injured by pH 2.0 fog, but no significant effects on membrane permeability were observed. For both species, whole-study average rates of net photosynthesis in previous year needles were lower in plants exposed to pH 2.0 fog than in plants treated with pH 4.0 fog. While decreased process rates coincided with leaf necrosis in white fir, stomatal closure appeared to be the mechanism of inhibition in ponderosa pine with pH 2.0 fog (i.e. no visible injury). The findings of the present study provide evidence that frequent applications of highly acidic fog (i.e. pH 2.0-3.0) can cause temporal alterations in membrane permeability and gas exchange rates in western conifer seedlings, in the presence or absence of visible injury. However, because incipient effects on other measures of foliage health were species-specific (i.e. concentrations of starch, photosynthetic pigments, inorganic nutrients), a general mechanism of phytotoxicity could not be identified.     

Effect of chronic ozone fumigation on the photosynthetic process of poplar clones showing different sensitivity.

Rooted cuttings from two poplar clones (Populus x euramericana, I-214, and Populus deltoides x maximowiczii, Eridano) were exposed for 15 days to diurnal square-wave treatment with ozone (60 nL L-1 for 5 h day-1). Completely fully expanded leaves exposed to ozone showed a reduction in net CO2 assimilation rate as compared to the control leaves during whole exposure period in both the clones. The reduction was related to a strong stomatal closure in clone I-214, but also to an altered mesophyll activity ascribed to limitation of the dark reactions of photosynthetic process. The results obtained in leaves of I-214 subjected to long-term fumigation seem to support the view that the decrease in quantum yield of electron transport may be a mechanism to down-regulate photosynthetic electron transport so that production of ATP and NADPH would be in equilibrium with the decreased demand in the Calvin cycle. In Eridano the CO2 assimilation was reduced because of the exposure and any alteration in stomatal conductance was observed. Thus, chlorophyll fluorescence parameters showed that an inhibition of photosystem II had occurred (reduction in Fv/Fm ratio), while no alterations in quenching parameters were observed upon illumination. The results seem to indicate that an alternative sink for reducing equivalent, other than carbon metabolism is present.     

Growth responses of Populus tremuloides clones to interacting elevated carbon dioxide and tropospheric ozone.

The Intergovernmental Panel of Climate Change (IPCC) has concluded that the greenhouse gases carbon dioxide (CO2) and tropospheric ozone (O3) are increasing concomitantly globally. Little is known about the effect of these interacting gases on growth, survival, and productivity of forest ecosystems. In this study we assess the effects of three successive years of exposure to combinations of elevated CO2 and O3 on growth responses in a five trembling aspen (Populus tremuloides) clonal mixture in a regenerating stand. The experiment is located in Rhinelander, Wisconsin, USA (45 degrees N 89 degrees W) and employs free air carbon dioxide and ozone enrichment (FACE) technology. The aspen stand was exposed to a factorial combination of four treatments consisting of elevated CO2 (560 ppm), elevated O3 (episodic exposure-90 microl l(-1) hour(-1)), a combination of elevated CO2 and O3, and ambient control in 30 m treatment rings with three replications. Our overall results showed that our three growth parameters including height, diameter and volume were increased by elevated CO2, decreased by elevated O3, and were not significantly different from the ambient control under elevated CO2 + O3. However, there were significant clonal differences in the responses; all five clones exhibited increased growth with elevated CO2, one clone showed an increase with elevated O3, and two clones showed an increase over the control with elevated CO2 + O3, two clones showed a decrease, and one was not significantly different from the control. Notably. there was a significant increase in current terminal shoot dieback with elevated CO2 during the 1999-2000 dormant season. Dieback was especially prominent in two of the five clones, and was attributed to those clones growing longer into the autumnal season where they were subject to frost. Our results show that elevated O3 negates expected positive growth effects of elevated CO2 in Populus tremuloides in the field, and suggest that future climate model predictions should take into account the offsetting effects of elevated O3 on CO2 enrichment when estimating future growth of trembling aspen stands.     

Arbuscular mycorrhizal fungi differentially affect the response to high zinc concentrations of two registered poplar clones

The effects of a high concentration of zinc on two registered clones of poplar (Populus alba Villafranca and Populus nigra Jean Pourtet), inoculated or not with two arbuscular mycorrhizal fungi (Glomus mosseae or Glomus intraradices) before transplanting them into polluted soil, were investigated, with special regard to the extent of root colonization by the fungi, plant growth, metal accumulation in the different plant organs, and leaf polyamine concentration. Zinc accumulation was lower in Jean Pourtet than in Villafranca poplars, and it was mainly translocated to the leaves; the metal inhibited mycorrhizal colonization, compromised plant growth, and, in Villafranca, altered the putrescine profile in the leaves. Most of these effects were reversed or reduced in plants pre-inoculated with G. mosseae. Results indicate that poplars are suitable for phytoremediation purposes, confirming that mycorrhizal fungi can be useful for phytoremediation, and underscore the importance of appropriate combinations of plant genotypes and fungal symbionts.     

Joint action of ozone and hydrogen fluoride on foliar senescence in maize

Maize (Zea mays, L.) plants were exposed intermittently to O(3), HF or both pollutants and the progression of foliar senescence was followed by measuring chlorophyll loss, membrane breakdown and changes in stomatal conductance. At concentrations insufficient to cause foliar symptoms (0.06 microl O(3) litre(-1) and 1.0 microg Fm(-3)), exposures to HF had little or no effect, whereas O(3) exposures accelerated the rate of senescence. The rapid rate of senescence produced by O(3) was moderated if the plants were also exposed to HF. Topical application of 6-benzyladenine (BA) prior to pollutant exposures delayed senescence in all plants and completely prevented the O(3)-induced acceleration of senescence.     

Responses of two birch (Betula pendula Roth) clones to different ozone profiles with similar AOT40 exposure

Saplings of two clones of European white birch (Betula pendula Roth) were exposed to three different ozone profiles resulting in same AOT40 value of 13–14 ppm h in a chamber experiment. The sensitive clone 5 and the more tolerant clone 2 were growing (1) under filtered air (=control), or (2) were exposed to 70 ppb ozone for 24 h d⁻¹ (=profile 1), (3) to 100 ppb ozone for 12 h d⁻¹ at 8:00–20:00 (=profile 2), or (4) to 200 ppb ozone for 4.5 h d⁻¹ at 9:30–14:00 (=profile 3) for 20 d. The saplings were determined for growth, visible leaf injuries, stomatal conductance, and concentrations of Rubisco, chlorophyll and carotenoids. Growth responses and induction of visible foliar injuries under different ozone profiles were variable, resulting in 4–17% lower dry mass of shoot, 16–46% reduction in stem height increment and 11–43% increase in visible injuries in clone 5, which was accompanied by higher leaf turnover rate under profile 3 indicating compensation growth. In clone 2, ozone-induced responses ranged from slight stimulation in stem height growth to 13% decrease in dry mass of shoot and 2–16% increase in visible injuries. Daytime stomatal conductance rates were lowered by 14–54% in clone 5 and 9–74% in clone 2, depending on profile. The additional power-weighted analyses revealed that high peak concentrations and exposure shape were important for induction of visible injuries in both clones and reduction in stomatal conductance in clone 5, whereas growth reductions were rather related to total cumulative exposure. The results indicate that profile of ozone exposure, night-time stomatal conductance (24 h flux), and recovery time for defence and compensations reactions should not be ignored in plant response and ozone flux modelling.     

Visible injury, crown condition, and growth responses of selected Italian forests in relation to ozone exposure

The impact of ozone on forest ecosystems in Italy is monitored within the CONECOFOR programme. Ozone levels are measured in 30 plots using passive samplers. Response parameters used are: crown condition (transparency), BAI (basal area increment), and visible symptoms on spontaneous vegetation. Levels of AOT40 are above the concentration-based critical level of 5 ppmh in all sites, but the evidence of impact on forest vegetation remains limited. Ozone is a predictor of crown transparency residuals in beech sites over two consecutive years, but the variance explained amounts to less than 10%. The relation between BAI reduction and ozone is even less certain. Transparency and BAI are more readily explainable in terms of ecological conditions of the site and climate fluctuations. The interpretation of visible symptoms is doubtful, and is conditioned by the prevailing ecological factors in the areas.     

Physiological and growth responses of differentially irrigated cotton to ozone

This study was conducted to determine the physiological and growth responses of cotton (Gossypium hirsutum L.) to the interaction of ozonee (O3) and drought stress. Cotton (cv SJ-2) was grown in open-top chambers in the field at three levels of soil water and exposed to charcoal-filtered air (CF), nonfiltered air (NF), and NF x 1.25, and NF x 1.5 ambient O3 concentrations in Riverside, CA, from June to October 1986. Ozone reduced carbon fixation an average of 74.6% in optimally watered (OW) plots, 63.4% in suboptimal (SO) plots, but only 19.3% in severely water-stressed (SS) plots. Leaf and stem biomass in OW and SO plots showed similar linear reductions in mass response to increased O3 concentrations, but SS plots showed no response to O3 except at the highest O3 treatment (seasonal 12-h O3 mean of 0.111 ppm 218 microm(-3)). These results showed that moderately water-stressed cotton had similar physiological and growth responses to O3 as well-watered plants, but severely water-stressed cotton showed little response to O3 at ambient O3 concentrations.     

Influence of uncertain reaction rates on ozone sensitivity to emissions

Air quality models rely upon simplified photochemical mechanisms to efficiently represent the thousands of chemical species that interact to form air pollution. Uncertainties in the chemical reaction rate constants and photolysis frequencies that comprise those mechanisms can generate uncertainty in the estimation of pollutant concentrations and their responsiveness to emission controls. A high-order sensitivity analysis technique is applied to quantify the extent to which reaction rate uncertainties influence estimates of ozone concentrations and their sensitivities to precursor emissions during an air pollution episode in Houston, Texas. Several reactions were found to have much larger proportional effects on ozone’s sensitivities to emissions than on its concentrations. In particular, uncertainties in photolysis frequencies and in the rate of reaction between NO₂ and OH to form nitric acid can significantly influence the magnitude and sign of peak ozone sensitivity to nitrogen oxide (NO_x) emissions. Ozone sensitivity to VOCs exhibits a much more muted response to uncertainties in the reaction rate constants and photolysis frequencies considered here. The results indicate the importance of accurate reaction rate constants to predicting the ozone impacts resulting from NO_x emission controls.     

Morphological,physiological and biochemical responses of two Australian biotypes of Parthenium hysterophorus to different soil moisture regimes

Parthenium weed is a problematic invasive species in several countries around the world. Although it is considered to be a highly invasive species within Australia, not all biotypes of parthenium weed exhibit the same ability in regard to aggressive colonization and distribution. Differences among biotypes, particularly in regard to environmental ranges as a possible basis for this variation, have not always been elucidated. To determine whether drought tolerance could be a factor in biotype demographics, we quantified the biological responses of two Australian parthenium weed biotypes known to differ in invasive ability Clermont (“high”) and Toogoolawah (“low”) to 100, 75 and 50% of soil water holding capacity (WHC). The Clermont biotype had greater vegetative growth, seed production and chlorophyll content than Toogoolawah, across all moisture levels. Net photosynthesis, stomatal conductance, internal CO₂ concentration, seed production per plant, 1000 seed weight and subsequent germination percentage were also higher for Clermont than for Toogoolawah and were maximum at 75% WHC. Clermont plants also had higher total soluble sugar, phenolics and free proline content than Toogoolawah, and a significant increase in the levels of all of these biochemicals was observed at 50% WHC. In conclusion, Clermont grew and reproduced better than Toogoolawah across all moisture regimes consistent of enhanced invasive ability of this biotype. Overall, the ability of parthenium weed to maintain good growth, physiology and seed production under moisture stress may enable it to colonize a wide range of Australian environments.

     

Assessing the risk of foliar injury from ozone on vegetation in parks in the U.S. National Park Service's Vital Signs Network

The risk of ozone injury to plants was assessed in support of the National Park Service's Vital Signs Monitoring Network program. The assessment examined bioindicator species, evaluated levels of ozone exposure, and investigated soil moisture conditions during periods of exposure for a 5-year period in each park. The assessment assigned each park a risk rating of high, moderate, or low. For the 244 parks for which assessments were conducted, the risk of foliar injury was high in 65 parks, moderate in 46 parks, and low in 131 parks. Among the well-known parks with a high risk of ozone injury are Gettysburg, Valley Forge, Delaware Water Gap, Cape Cod, Fire Island, Antietam, Harpers Ferry, Manassas, Wolf Trap Farm Park, Mammoth Cave, Shiloh, Sleeping Bear Dunes, Great Smoky Mountains, Joshua Tree, Sequoia and Kings Canyon, and Yosemite.     

Ethylenediurea as a potential tool in evaluating ozone phytotoxicity: a review study on physiological,biochemical and morphological responses of plants

Present-day climate change scenario has intensified the problem of continuously increasing ground-level ozone (O₃), which is responsible for causing deleterious effects on growth and development of plants. Studies involving use of ethylenediurea (EDU), a chemical with antiozonant properties, have given some promising results in evaluating O₃ injury in plants. The use of EDU is especially advantageous in developing countries which face a more severe problem of ground-level O₃, and technical O₃-induced yield loss assessment techniques like open-top chambers cannot be used. Recent studies have detected a hormetic response of EDU on plants; i.e. treatment with higher EDU concentrations may or may not show any adverse effect on plants depending upon the experimental conditions. Although the mode of action of EDU is still debated, it is confirmed that EDU remains confined in the apoplastic regions. Certain studies indicate that EDU significantly affects the electron transport chain and has positive impact on the antioxidant defence machinery of the plants. However, the mechanism of protecting the yield of plants without significantly affecting photosynthesis is still questionable. This review discusses in details the probable mode of action of EDU on the basis of available data along with the impact of EDU on physiological, biochemical, growth and yield response of plants under O₃ stress. Data regarding the effect of EDU on plant ‘omics’ is highly insufficient and can form an important aspect of future EDU research.     

Ozone air pollution and foliar injury development on native plants of Switzerland

The objectives of this study were to examine the foliar sensitivity to ozone exposure of 12 tree, shrub, and herbaceous species native to southern Switzerland and determine the seasonal cumulative ozone exposures required to induce visible foliar injury. The study was conducted from the beginning of May through the end of August during 2000 and 2001 using an open-top chamber research facility located within the Lattecaldo Cantonal Forest Nursery in Canton Ticino, southern Switzerland (600 m asl). Plants were examined daily and dates of initial foliar injury were recorded in order to determine the cumulative AOT40 ppb h ozone exposure required to cause visible foliar injury. Plant responses to ozone varied significantly among species; 11 species exhibited visible symptoms typical of exposures to ambient ozone. The symptomatic species (from most to least sensitive) were Populus nigra, Viburnum lantana, Salix alba, Crataegus monogyna, Viburnum opulus, Tilia platyphyllos, Cornus alba, Prunus avium, Fraxinus excelsior, Ribes alpinum, and Tilia cordata; Clematis spp. did not show foliar symptoms. Of the 11 symptomatic species, five showed initial injury below the critical level AOT40 10 ppmh O3 in the 2001 season.     

Influence of two growing seasons of experimental ozone fumigation on photosynthetic characteristics of white oak seedlings

White oak (Quercus alba L.) seedlings were exposed to charcoal-filtered air or to above-ambient ozone concentrations for 19-20 weeks during each of two growing seasons in continuously stirred tank reactors in greenhouses. Ozone treatments were 0.15 ppm (300 microg m(-3)) for 8 h day(-1), 3 days week(-1) in 1988, and continuous 15% above ambient in 1989. The seedlings were grown in forest soil watered twice weekly with simulated rain of pH 5.2. Responses of net photosynthesis to photosynthetically active radiation and intercellular CO(2) concentration were measured three times each year. There were no significant differences in light-saturated net photosynthesis or stomatal conductance, dark respiration, quantum or carboxylation efficiencies, and light or CO(2) compensation points on any date between control and ozone-exposed seedlings.