Stomatal characteristics and infection biology of Pyrenopeziza betulicola in Betula pendula trees grown under elevated CO2 and O3

Two silver birch clones were exposed to ambient and elevated concentrations of CO₂ and O₃, and their combination for 3 years, using open-top chambers. We evaluated the effects of elevated CO₂ and O₃ on stomatal conductance (g_s), density (SD) and index (SI), length of the guard cells, and epidermal cell size and number, with respect to crown position and leaf type. The relationship between the infection biology of the fungus (Pyrenopeziza betulicola) causing leaf spot disease and stomatal characteristics was also studied. Leaf type was an important determinant of O₃ response in silver birch, while crown position and clone played only a minor role. Elevated CO₂ reduced the g_s, but had otherwise no significant effect on the parameters studied. No significant interactions between elevated CO₂ and O₃ were found. The infection biology of P. betulicola was not correlated with SD or g_s, but it did occasionally correlate positively with the length of the guard cells.     

Diurnal changes in photosynthetic parameters of Populus tremuloides, modulated by elevated concentrations of CO2 and/or O3 and daily climatic variation

The diurnal changes in light-saturated photosynthesis (Pn) under elevated CO₂ and/or O₃ in relation to stomatal conductance (g_s), water potential, intercellular [CO₂], leaf temperature and vapour-pressure difference between leaf and air (VPD_L) were studied at the Aspen FACE site. Two aspen (Populus tremuloides Michx.) clones differing in their sensitivity to ozone were measured. The depression in Pn was found after 10:00 h. The midday decline in Pn corresponded with both decreased g_s and decreased Rubisco carboxylation efficiency, Vc_max. As a result of increasing VPD_L, g_s decreased. Elevated [CO₂] resulted in more pronounced midday decline in Pn compared to ambient concentrations. Moreover, this decline was more pronounced under combined treatment compared to elevated CO₂ treatment.The positive impact of CO₂ on Pn was relatively more pronounced in days with environmental stress but relatively less pronounced during midday depression. The negative impact of ozone tended to decrease in both cases.     

Anthropogenic NOx emissions alter the intrinsic water-use efficiency (WUEi) for Quercus cerris stands under Mediterranean climate conditions

We investigated the effect of N deposition (Ndep) on intrinsic water-use efficiency (WUE_i), the ratio of photosynthesis (A) to stomatal conductance (g_s), for two Quercus cerris stands at different distances to an oil refinery in Southern Italy. We used δ¹³C in tree rings for assessing changes in WUE_i; while the influence of climate and NO_x emission was explored through δ¹⁸O and δ¹⁵N, respectively. Differences in WUE_i between the two sites were significant, with trees exposed to different degrees of NO_x emissions showing an abrupt increase with the onset of pollution. Assuming similar g_s at the two sites, as inferred through δ¹⁸O, the higher N availability at the polluted site caused the shift of the A/g_s ratio in favour of A. Overall, our result suggests that an increase of Ndep may enhance tree WUE under a scenario of reduction of precipitation predicted for Mediterranean area.     

Stomatal uptake of O3 in aspen and aspen-birch forests under free-air CO2 and O3 enrichment

Rising atmospheric carbon dioxide (CO₂) may alleviate the toxicological impacts of concurrently rising tropospheric ozone (O₃) during the present century if higher CO₂ is accompanied by lower stomatal conductance (g_s), as assumed by many models. We investigated how elevated concentrations of CO₂ and O₃, alone and in combination, affected the accumulated stomatal flux of O₃ (AFst) by canopies and sun leaves in closed aspen and aspen-birch forests in the free-air CO₂-O₃ enrichment experiment near Rhinelander, Wisconsin. Stomatal conductance for O₃ was derived from sap flux data and AFst was estimated either neglecting or accounting for the potential influence of non-stomatal leaf surface O₃ deposition. Leaf-level AFst (AFst_l) was not reduced by elevated CO₂. Instead, there was a significant CO₂ × O₃ interaction on AFst_l, as a consequence of lower values of g_s in control plots and the combination treatment than in the two single-gas treatments. In addition, aspen leaves had higher AFst_l than birch leaves, and estimates of AFst_l were not very sensitive to non-stomatal leaf surface O₃ deposition. Our results suggest that model projections of large CO₂-induced reductions in g_s alleviating the adverse effect of rising tropospheric O₃ may not be reasonable for northern hardwood forests.     

Effects of sub-lethal glyphosate concentrations on growth and photosynthetic performance of non-target species Bolboschoenus maritimus

Glyphosate use has increased over the last decades for the control of invasive plant species in wetland ecosystems. Although glyphosate has been considered ‘environmentally’ safe, its repeated use could increase the toxicological risk derived from diffuse pollution of surface and groundwater on non-target vegetation. A glasshouse study was designed to determine the effect produced by the addition of different sub-lethal doses of glyphosate herbicides (5–30 mg L⁻¹) to the nutrient solution on the growth and photosynthetic apparatus of Bolboschoenus maritimus. Although B. maritimus plants were able to grow and survive after 20 d of exposure to glyphosate, the presence of this herbicide affected their growth, through a direct interaction with the root system. Particularly, at 30 mg L⁻¹ glyphosate, B. maritimus showed ca. 30% of biomass decrease. The reduction in B. maritimus growth was due to a decrease in net photosynthetic rate (A), which ranged between values ca. 11.5 and 5.5 μmol m⁻² s⁻¹ CO₂ for the control and the highest glyphosate treatment, respectively. The response of A to glyphosate could be largely accounted for by non-stomatal limitations, since stomatal conductance was similar in all glyphosate treatments. Thus, A decrease was prompted by the negative impact of herbicide on photochemical (PSII) apparatus, the reduction in the absorption of essential nutrients, the reduction of photosynthetic pigments and possibly the reduction in Rubisco carboxilation capacity. Moreover, glyphosate excess caused photoinhibitory damage. In conclusion, in this study we have shown that herbicide water pollution could be a source of indirect phytotoxicity for B. maritimus.     

The growth response of Alternanthera philoxeroides in a simulated post-combustion emission with ultrahigh [CO2] and acidic pollutants

Although post-combustion emissions from power plants are a major source of air pollution, they contain excess CO₂ that could be used to fertilize commercial greenhouses and stimulate plant growth. We addressed the combined effects of ultrahigh [CO₂] and acidic pollutants in flue gas on the growth of Alternanthera philoxeroides. When acidic pollutants were excluded, the biomass yield of A. philoxeroides saturated near 2000 μmol mol⁻¹ [CO₂] with doubled biomass accumulation relative to the ambient control. The growth enhancement was maintained at 5000 μmol mol⁻¹ [CO₂], but declined when [CO₂] rose above 1%, in association with a strong photosynthetic inhibition. Although acidic components (SO₂ and NO₂) significantly offset the CO₂ enhancement, the aboveground yield increased considerably when the concentration of pollutants was moderate (200 times dilution). Our results indicate that using excess CO₂ from the power plant emissions to optimize growth in commercial green house could be viable.     

Combined effects of elevated CO2 and natural climatic variation on leaf spot diseases of redbud and sweetgum trees

Atmospheric CO₂ concentrations are predicted to double within the next century and alter climate regimes, yet the extent that these changes will affect plant diseases remains unclear. In this study conducted over five years, we assessed how elevated CO₂ and interannual climatic variability affect Cercospora leaf spot diseases of two deciduous trees. Climatic data varied considerably between the five years and altered disease expression. Disease incidence and severity for both species were greater in years with above average rainfall. In years with above average temperatures, disease incidence for Liquidambar styraciflua was decreased significantly. When significant changes did occur, disease incidence and severity always increased under elevated CO₂. Chlorophyll fluorescence imaging of leaves revealed that any visible increase in disease severity induced by elevated CO₂ was mitigated by higher photosynthetic efficiency in the remaining undamaged leaf tissue and in a halo surrounding lesions.     

Modelling annual pasture dynamics: Application to stomatal ozone deposition

Modelling ozone (O₃) deposition for impact risk assessment is still poorly developed for herbaceous vegetation, particularly for Mediterranean annual pastures. High inter-annual climatic variability in the Mediterranean area makes it difficult to develop models characterizing gas exchange behaviour and air pollutant absorption suitable for risk assessment. This paper presents a new model to estimate stomatal conductance (g_s) of Trifolium subterraneum, a characteristic species of dehesa pastures. The MEDPAS (MEDiterranean PAStures) model couples 3 modules estimating soil water content (SWC), vegetation growth and g_s. The g_s module is a reparameterized version of the stomatal component of the EMEP DO₃SE O₃ deposition model. The MEDPAS model was applied to two contrasting years representing typical dry and humid springs respectively and with different O₃ exposures. The MEDPAS model reproduced realistically the g_s seasonal and inter-annual variations observed in the field. SWC was identified as the major driver of differences across years. Despite the higher O₃ exposure in the dry year, meteorological conditions favoured 2.1 times higher g_s and 56 day longer growing season in the humid year compared to the dry year. This resulted in higher ozone fluxes absorbed by T. subterraneum in the humid year. High inter-family variability was found in gas exchange rates, therefore limiting the relevance of single species O₃ deposition flux modelling for dehesa pastures. Stomatal conductance dynamics at the canopy level need to be considered for more accurate O₃ flux modelling for present and future climate scenarios in the Mediterranean area.     

Production of carbon dioxide in a fattening pig house under field conditions. II. Release from the manure

Carbon dioxide (CO₂) release from manure was investigated under field conditions in one of the compartments in a mechanically ventilated fattening pig house. The potential of CO₂ release was tested in the emptied compartment by imposing high ventilation rate and measuring CO₂ concentrations in the incoming and exhaust air. When the compartment was filled with pigs, a model of tranquil CO₂ exhalation rate (TCER) was used to estimate the part of CO₂ produced by animals, in addition to the measurement of CO₂ concentrations and ventilation rate. Useful data were extracted from 135 days of continuously measured field data obtained when the housing system was in steady-state. The CO₂ release per m² of pit surface was 18.4 and 25.8 g h⁻¹ m⁻² in the emptied pig compartment during the impulse and pulse ventilation rate tests, respectively. It ranged from 1.1 and 116.7 g h⁻¹ m⁻² and averaged 42.1 g h⁻¹ m⁻² during the 135-day continuous field measurement. In 128 of the 135 days when the compartment was filled with pigs, the mean CO₂ release from the manure reached an average of 37.5% of the TCER by pigs, instead of less than 5% believed by many researchers. The maximum CO₂ release rate in 2-h steady-state measurements was 94% of the TCER by pigs. This research demonstrated that, in the previous knowledge, there might be a serious underestimation of the CO₂ release from the manure in pig houses. The total pig weight, manure temperature and ventilation rate were found the most important variables related to the CO₂ release. A mathematical model was developed to estimate the CO₂ release as a function of these three variables with R²=0.525.     

Seasonal soil and leaf CO2 exchange rates in a Mediterranean holm oak forest and their responses to drought conditions

We measured the soil and leaf CO₂ exchange in Quercus ilex and Phillyrea latifolia seasonally throughout the year in a representative site of the Mediterranean region, a natural holm oak forest growing in the Prades Mountains in southeastern Catalonia. In the wet seasons (spring and autumn), we experimentally decreased soil moisture by 30%, by excluding rainfall and water runoff in 12 plots, 1×10 m, and left 12 further plots as controls. Our aim was to predict the response of these gas exchanges to the drought forecasted for the next decades for this region by GCM and ecophysiological models.Annual average soil CO₂ exchange rate was 2.27±0.27 μmol CO₂ m⁻² s⁻¹. Annual average leaf CO₂ exchange rates were 8±1 and 5±1 μmol m⁻² s⁻¹ in Q. ilex and P. latifolia, respectively. Soil respiration rates in control treatments followed a seasonal pattern similar to photosynthetic activity. They reached maximum values in spring and autumn (2.5–3.8 μmol m⁻² s⁻¹ soil CO₂ emission rates and 7–15 μmol m⁻² s⁻¹ net photosynthetic rates) and minimum values (almost 0 for both variables) in summer, showing that soil moisture was the most important factor driving the soil microbial activity and the photosynthetic activity of plants. In autumn, drought treatment strongly decreased net photosynthesis rates and stomatal conductance of Q. ilex by 44% and 53%, respectively. Soil respiration was also reduced by 43% under drought treatment in the wet seasons. In summer there were larger soil CO₂ emissions in drought plots than in control plots, probably driven by autotrophic (roots) metabolism. The results indicate that leaf and soil CO₂ exchange may be strongly reduced (by ca. 44%) by the predicted decreases of soil water availability in the next decades. Long-term studies are needed to confirm these predictions or to find out possible acclimation of those processes.     

Will photosynthetic capacity of aspen trees acclimate after long-term exposure to elevated CO2 and O3?

Photosynthetic acclimation under elevated carbon dioxide (CO₂) and/or ozone (O₃) has been the topic of discussion in many papers recently. We examined whether or not aspen plants grown under elevated CO₂ and/or O₃ will acclimate after 11 years of exposure at the Aspen Face site in Rhinelander, WI, USA. We studied diurnal patterns of instantaneous photosynthetic measurements as well as A/C_i measurements monthly during the 2004-2008 growing seasons. Our results suggest that the responses of two aspen clones differing in O₃ sensitivity showed no evidence of photosynthetic and stomatal acclimation under either elevated CO₂, O₃ or CO₂ + O₃. Both clones 42E and 271 did not show photosynthetic nor stomatal acclimation under elevated CO₂ and O₃ after a decade of exposure. We found that the degree of increase or decrease in the photosynthesis and stomatal conductance varied significantly from day to day and from one season to another.     

Effects of ammonia from livestock farming on lichen photosynthesis

This study investigated if atmospheric ammonia (NH₃) pollution around a sheep farm influences the photosynthetic performance of the lichens Evernia prunastri and Pseudevernia furfuracea. Thalli of both species were transplanted for up to 30 days in a semi-arid region (Crete, Greece), at sites with concentrations of atmospheric ammonia of ca. 60 μg/m³ (at a sheep farm), ca. 15 μg/m³ (60 m from the sheep farm) and ca. 2 μg/m³ (a remote area 5 km away). Lichen photosynthesis was analysed by the chlorophyll a fluorescence emission to identify targets of ammonia pollution. The results indicated that the photosystem II of the two lichens exposed to NH₃ is susceptible to this pollutant in the gas-phase. The parameter PI_ABS, a global index of photosynthetic performance that combines in a single expression the three functional steps of the photosynthetic activity (light absorption, excitation energy trapping, and conversion of excitation energy to electron transport) was much more sensitive to NH₃ than the F_V/F_M ratio, one of the most commonly used stress indicators.     

Gene expression responses of paper birch (Betula papyrifera) to elevated CO2 and O3 during leaf maturation and senescence

Gene expression responses of paper birch (Betula papyrifera) leaves to elevated concentrations of CO₂ and O₃ were studied with microarray analyses from three time points during the summer of 2004 at Aspen FACE. Microarray data were analyzed with clustering techniques, self-organizing maps, K-means clustering and Sammon's mappings, to detect similar gene expression patterns within sampling times and treatments. Most of the alterations in gene expression were caused by O₃, alone or in combination with CO₂. O₃ induced defensive reactions to oxidative stress and earlier leaf senescence, seen as decreased expression of photosynthesis- and carbon fixation-related genes, and increased expression of senescence-associated genes. The effects of elevated CO₂ reflected surplus of carbon that was directed to synthesis of secondary compounds. The combined CO₂ + O₃ treatment resulted in differential gene expression than with individual gas treatments or in changes similar to O₃ treatment, indicating that CO₂ cannot totally alleviate the harmful effects of O₃.     

The estimation of historical CO2 trajectories is indeterminate: Comment on “A new look at atmospheric carbon dioxide”

A paper by Hofmann et al. (2009, this journal) is critiqued. It is shown that their exponential model for characterizing CO₂ trajectories for historical data is not estimated properly. An exponential model is properly estimated and is shown to fit over the entire 51 year period of available data. Further, the entire problem of estimating models for the CO₂ historical data is shown to be ill-posed because alternate model forms fit the data equally well. To illustrate this point the past 51 years of CO₂ data were analyzed using three different time-dependent models that capture the historical pattern of CO₂ increase. All three fit with R² > 0.98, are visually indistinguishable when overlaid, and match each other during the calibration period with R² > 0.999. Projecting the models forward to 2100, the exponential model comes quite close to the Intergovernmental Panel on Climate Change (IPCC) best estimate of 836 ppmv. The other two models project values far below the IPCC low estimates. The problem of characterizing historical CO₂ levels is thus indeterminate, because multiple models fit the data equally well but forecast very different future trajectories.     

Belowground effects of enhanced tropospheric ozone and drought in a beech/spruce forest (Fagus sylvatica L./Picea abies [L.] Karst)

The effects of experimentally elevated O₃ on soil respiration rates, standing fine-root biomass, fine-root production and δ¹³C signature of newly produced fine roots were investigated in an adult European beech/Norway spruce forest in Germany during two subsequent years with contrasting rainfall patterns. During humid 2002, soil respiration rate was enhanced under elevated O₃ under beech and spruce, and was related to O₃-stimulated fine-root production only in beech. During dry 2003, the stimulating effect of O₃ on soil respiration rate vanished under spruce, which was correlated with decreased fine-root production in spruce under drought, irrespective of the O₃ regime. δ¹³C signature of newly formed fine-roots was consistent with the differing g_s of beech and spruce, and indicated stomatal limitation by O₃ in beech and by drought in spruce. Our study showed that drought can override the stimulating O₃ effects on fine-root dynamics and soil respiration in mature beech and spruce forests.     

Spring leaf flush in aspen (Populus tremuloides) clones is altered by long-term growth at elevated carbon dioxide and elevated ozone concentration

Early spring leaf out is important to the success of deciduous trees competing for light and space in dense forest plantation canopies. In this study, we investigated spring leaf flush and how long-term growth at elevated carbon dioxide concentration ([CO₂]) and elevated ozone concentration ([O₃]) altered leaf area index development in a closed Populus tremuloides (aspen) canopy. This work was done at the Aspen FACE experiment where aspen clones have been grown since 1997 in conditions simulating the [CO₂] and [O₃] predicted for ∼2050. The responses of two clones were compared during the first month of spring leaf out when CO₂ fumigation had begun, but O₃ fumigation had not. Trees in elevated [CO₂] plots showed a stimulation of leaf area index (36%), while trees in elevated [O₃] plots had lower leaf area index (−20%). While individual leaf area was not significantly affected by elevated [CO₂], the photosynthetic operating efficiency of aspen leaves was significantly improved (51%). There were no significant differences in the way that the two aspen clones responded to elevated [CO₂]; however, the two clones responded differently to long-term growth at elevated [O₃]. The O₃-sensitive clone, 42E, had reduced individual leaf area when grown at elevated [O₃] (−32%), while the tolerant clone, 216, had larger mature leaf area at elevated [O₃] (46%). These results indicate a clear difference between the two clones in their long-term response to elevated [O₃], which could affect competition between the clones, and result in altered genotypic composition in future atmospheric conditions.     

Elevated CO2 response of photosynthesis depends on ozone concentration in aspen

The effect of elevated CO₂ and O₃ on apparent quantum yield (?), maximum photosynthesis (P_max), carboxylation efficiency (V_cmax) and electron transport capacity (J_max) at different canopy locations was studied in two aspen (Populus tremuloides) clones of contrasting O₃ tolerance. Local light climate at every leaf was characterized as fraction of above-canopy photosynthetic photon flux density (%PPFD). Elevated CO₂ alone did not affect ? or P_max, and increased J_max in the O₃-sensitive, but not in the O₃-tolerant clone. Elevated O₃ decreased leaf chlorophyll content and all photosynthetic parameters, particularly in the lower canopy, and the negative impact of O₃ increased through time. Significant interaction effect, whereby the negative impact of elevated O₃ was exaggerated by elevated CO₂ was seen in Chl, N and J_max, and occurred in both O₃-tolerant and O₃-sensitive clones. The clonal differences in the level of CO₂ × O₃ interaction suggest a relationship between photosynthetic acclimation and background O₃ concentration.     

Evaluation of the UNC toluene-SOA mechanism with respect to other chamber studies and key model parameters

In a companion paper by Hu et al. [2007. A kinetic mechanism for predicting secondary organic aerosol formation from toluene oxidation in the presence of NO_x and natural sunlight. Atmospheric Environment, doi:10.1016/j.atmosenv.2007.04.025], a kinetic mechanism was developed from data generated in the University of North Carolina's (UNC) 270 m³ dual outdoor aerosol smog chamber, to predict secondary organic aerosol (SOA) formation from toluene oxidation in the atmosphere. In this paper, experimental data sets from European Photoreactor (EUPHORE), smog chambers at the California Institute of Technology (Caltech), and the UNC 300 m³ dual-outdoor gas phase chamber were used to evaluate the toluene mechanism. The model simulates SOA formation for the ‘low-NO_x’ and ‘mid-NO_x’ experiments from EUPHORE chambers reasonably well, but over-predicts SOA mass concentrations for the ‘high-NO_x’ run. The model well simulates the SOA mass concentrations observed from the Caltech chambers. Experiments with the three key toluene products, 1,4-butenedial, 4-oxo-2-pentenal and o-cresol in the presence of oxides of nitrogen (NO_x) are also simulated by the developed mechanism. The model well predicts the NO_x time–concentration profiles and the decay of these two carbonyls, but underestimates ozone (O₃) formation for 4-oxo-2-pentenal. It well simulates SOA formation from 1,4-butenedial but overestimates (possibly due to experimental problems) the measured aerosol mass concentrations from 4-oxo-2-pentenal. The model underestimates SOA production from o-cresol, mostly due to its under-prediction of o-cresol decay. The effects of varying temperature, relative humidity, glyoxal uptake, organic nitrate yields, and background seed aerosol concentrations, were also investigated.     

Differential response of aspen and birch trees to heat stress under elevated carbon dioxide

The effect of high temperature on photosynthesis of isoprene-emitting (aspen) and non-isoprene-emitting (birch) trees were measured under elevated CO₂ and ambient conditions. Aspen trees tolerated heat better than birch trees and elevated CO₂ protected photosynthesis of both species against moderate heat stress. Elevated CO₂ increased carboxylation capacity, photosynthetic electron transport capacity, and triose phosphate use in both birch and aspen trees. High temperature (36-39 °C) decreased all of these parameters in birch regardless of CO₂ treatment, but only photosynthetic electron transport and triose phosphate use at ambient CO₂ were reduced in aspen. Among the two aspen clones tested, 271 showed higher thermotolerance than 42E possibly because of the higher isoprene-emission, especially under elevated CO₂. Our results indicate that isoprene-emitting trees may have a competitive advantage over non-isoprene emitting ones as temperatures rise, indicating that biological diversity may be affected in some ecosystems because of heat tolerance mechanisms.     

Direct and indirect effects of ammonia, ammonium and nitrate on phosphatase activity and carbon fluxes from decomposing litter in peatland

Here we investigate the response of soils and litter to 5 years of experimental additions of ammonium (NH₄), nitrate (NO₃), and ammonia (NH₃) to an ombrotrophic peatland. We test the importance of direct (via soil) and indirect (via litter) effects on phosphatase activity and efflux of CO₂. We also determined how species representing different functional types responded to the nitrogen treatments. Our results demonstrate that additions of NO₃, NH₄ and NH₃ all stimulated phosphatase activity but the effects were dependent on species of litter and mechanism (direct or indirect). Deposition of NH₃ had no effect on efflux of CO₂ from Calluna vulgaris litter, despite it showing signs of stress in the field, whereas both NO₃ and NH₄ reduced CO₂ fluxes. Our results show that the collective impacts on peatlands of the three principal forms of nitrogen in atmospheric deposition are a result of differential effects and mechanisms on individual components.