Better estimation of evapotranspiration and transpiration using an improved modified Priestly-Taylor model based on a new parameter of leaf senescence in a rice field.

Evapotranspiration (ET) is at the heart of the global water, energy, and carbon cycles. As ET is difficult and expensive to measure, it is crucial to develop estimation models that can be widely applied. Currently, an improved Priestley-Taylor (PT) model considers soil moisture stress, temperature constraints, and leaf senescence; however, its parameter (fs) for simulating crop senescence is based on empirical values, making it difficult to apply to different varieties and complex external conditions and thus challenging to generalize. We improved the parameters fs in the original model based on the chlorophyll decomposition that accompanies crop senescence through easily observable SPAD values (Soil-Plant Analysis Development readings) in the field. We validated the improved model by obtaining ET of different rice varieties in 2022 and 2023 using the energy balance residual method at the Free Air Concentration Enrichment Experimental (FACE) Facility located in Yangzhou City, China. The results showed that the simulation of leaf senescence using SPAD values was feasible and could be extended to different varieties. The new model using improved leaf senescence parameter for estimating ET and transpiration (T) in three plots (2022 and 2023) exhibited slightly enhanced accuracy, particularly at the later stages of crop growth. Moreover, the higher the T/ET ratio of the cropland, the more significant the improvement. This new development enhances the ability of PT models to estimate ET and T using readily available field observations and provides some suggestions for wider application in the field for other crop species.

Drought mitigates the adverse effects of O3 on plant photosynthesis rather than growth: A global meta-analysis considering plant functional types.

Tropospheric ozone (O3 ) is a phytotoxic air pollutant adversely affecting plant growth. High O3 exposures are often concurrent with summer drought. The effects of both stresses on plants are complex, and their interactions are not yet well understood. Here, we investigate whether drought can mitigate the negative effects of O3 on plant physiology and growth based on a meta-analysis. We found that drought mitigated the negative effects of O3 on plant photosynthesis, but the modification of the O3 effect on the whole-plant biomass by drought was not significant. This is explained by a compensatory response of water-deficient plants that leads to increased metabolic costs. Relative to water control condition, reduced water treatment decreased the effects of O3 on photosynthetic traits, and leaf and root biomass in deciduous broadleaf species, while all traits in evergreen coniferous species showed no significant response. This suggested that the mitigating effects of drought on the negative impacts of O3 on the deciduous broadleaf species were more extensive than on the evergreen coniferous ones. Therefore, to avoid over- or underestimations when assessing the impact of O3 on vegetation growth, soil moisture should be considered. These results contribute to a better understanding of terrestrial ecosystem responses under global change.

Climate gradient and leaf carbon investment influence the effects of climate change on water use efficiency of forests: A meta-analysis.

Forest ecosystems cover a large area of the global land surface and are important carbon sinks. The water-carbon cycles of forests are prone to climate change, but uncertainties remain regarding the magnitude of water use efficiency (WUE) response to climate change and the underpinning mechanism driving WUE variation. We conducted a meta-analysis of the effects of elevated CO2 concentration (eCO2 ), drought and elevated temperature (eT) on the leaf- to plant-level WUE, covering 80 field studies and 95 tree species. The results showed that eCO2 increased leaf intrinsic and instantaneous WUE (WUEi, WUEt), whereas drought enhanced both leaf- and plant-level WUEs. eT increased WUEi but decreased carbon isotope-based WUE, possibly due to the influence of mesophyll conductance. Stimulated leaf-level WUE by drought showed a progressing trend with increasing latitude, while eCO2 -induced WUE enhancement showed decreasing trends after >40° N. These latitudinal gradients might influence the spatial pattern of climate and further drove WUE variation. Moreover, high leaf-level WUE under eCO2 and drought was accompanied by low leaf carbon contents. Such a trade-off between growth efficiency and defence suggests a potentially compromised tolerance to diseases and pests. These findings add important ecophysiological parameters into climate models to predict carbon-water cycles of forests.

Adapting crop production to climate change and air pollution at different scales.

Air pollution and climate change are tightly interconnected and jointly affect field crop production and agroecosystem health. Although our understanding of the individual and combined impacts of air pollution and climate change factors is improving, the adaptation of crop production to concurrent air pollution and climate change remains challenging to resolve. Here we evaluate recent advances in the adaptation of crop production to climate change and air pollution at the plant, field and ecosystem scales. The main approaches at the plant level include the integration of genetic variation, molecular breeding and phenotyping. Field-level techniques include optimizing cultivation practices, promoting mixed cropping and diversification, and applying technologies such as antiozonants, nanotechnology and robot-assisted farming. Plant- and field-level techniques would be further facilitated by enhancing soil resilience, incorporating precision agriculture and modifying the hydrology and microclimate of agricultural landscapes at the ecosystem level. Strategies and opportunities for crop production under climate change and air pollution are discussed.

Physiological and molecular responses of different rose (Rosa hybrida L.) cultivars to elevated ozone levels.

The increasing ground-level ozone (O3) pollution resulting from rapid global urbanization and industrialization has negative effects on many plants. Nonetheless, many gaps remain in our knowledge of how ornamental plants respond to O3. Rose (Rosa hybrida L.) is a commercially important ornamental plant worldwide. In this study, we exposed four rose cultivars ("Schloss Mannheim," "Iceberg," "Lüye," and "Spectra") to either unfiltered ambient air (NF), unfiltered ambient air plus 40 ppb O3 (NF40), or unfiltered ambient air plus 80 ppb O3 (NF80). Only the cultivar "Schloss Mannheim" showed significant O3-related effects, including foliar injury, reduced chlorophyll content, reduced net photosynthetic rate, reduced stomatal conductance, and reduced stomatal apertures. In "Schloss Mannheim," several transcription factor genes-HSF, WRKY, and MYB genes-were upregulated by O3 exposure, and their expression was correlated with that of NCED1, PP2Cs, PYR/PYL, and UGTs, which are related to ABA biosynthesis and signaling. These results suggest that HSF, WRKY, and MYB transcription factors and ABA are important components of the plant response to O3 stress, suggesting a possible strategy for cultivating O3-tolerant rose varieties.

Variations in leaf anatomical characteristics drive the decrease of mesophyll conductance in poplar under elevated ozone.

Decline in mesophyll conductance (gm ) plays a key role in limiting photosynthesis in plants exposed to elevated ozone (O3 ). Leaf anatomical traits are known to influence gm , but the potential effects of O3 -induced changes in leaf anatomy on gm have not yet been clarified. Here, two poplar clones were exposed to elevated O3 . The effects of O3 on the photosynthetic capacity and anatomical characteristics were assessed to investigate the leaf anatomical properties that potentially affect gm . We also conducted global meta-analysis to explore the general response patterns of gm and leaf anatomy to O3 exposure. We found that the O3 -induced reduction in gm was critical in limiting leaf photosynthesis. Changes in liquid-phase conductance rather than gas-phase conductance drive the decline in gm under elevated O3, and this effect was associated with thicker cell walls and smaller chloroplast sizes. The effects of O3 on palisade and spongy mesophyll cell traits and their contributions to gm were highly genotype-dependent. Our results suggest that, while anatomical adjustments under elevated O3 may contribute to defense against O3 stress, they also cause declines in gm and photosynthesis. These results provide the first evidence of anatomical constraints on gm under elevated O3 .

Effects of elevated ozone on the uptake and allocation of macronutrients in poplar saplings above- and belowground.

Ground-level ozone (O3) is a secondary air pollutant and affects the roots and soil processes of trees. Therefore, O3 can affect the uptake and allocation of nutrients in trees, which merits further clarification. A fumigation experiment with five O3 levels was conducted in 15 open top chambers for two poplar clones, and the concentrations of six macronutrients (N, P, K, S, Ca, Mg) in different organs and leaf positions were determined. Under all O3 levels, the concentration of mobile nutrients (N and P) was higher in upper leaves than in lower leaves, while the non-mobile nutrients (Ca and S) concentration was the opposite. Relative to charcoal filtered ambient air (CF), high O3 treatment (NF60) significantly increased the concentration of mobile nutrients K and Mg in upper leaves by 38 % and 33 %, in lower leaves by 142 % and 65 %, respectively, which suggested the effect of O3 on their concentrations was greater at the lower leaf position than at the upper leaf position. Elevated O3 significantly increased the macronutrient concentrations in most organs. The effects of O3 on nutrient concentrations were attributed using graphical vector analysis, suggested that the increase of nutrient concentration in the shoots was attributed to excessive nutrient stocks, while their increase in root was attributed to the "concentration" effect. Compared to CF, NF60 also reduced the root-to-shoot ratio of N, P, S, K, Ca and Mg stocks by 34 %, 39 %, 37 %, 64 %, 46 % and 42 %, respectively, indicating the allocation of increased nutrients to shoots in response to O3 stress. Changes in the allocation pattern of nutrients in different leaf positions and organs of poplar were primarily in response to O3 stress since these nutrients play important roles in some physiological processes. These results will help improve the plantation nutrient utilization by optimizing fertilizer management regimes under O3 pollution.

Elevated ozone decreases the activity of Rubisco in poplar but not its activation under fluctuating light.

Increasing tropospheric ozone (O3) is well-known to decrease leaf photosynthesis under steady-state light through reductions in biochemical capacity. However, the effects of O3 on photosynthetic induction and its biochemical limitations in response to fluctuating light remain unclear, despite the rapid fluctuations of light intensity occurring under field conditions. In this study, two hybrid poplar clones with different O3 sensitivities were exposed to elevated O3. Dynamic photosynthetic CO2 response measurements were conducted to quantify the impact of elevated O3 and exposure duration on biochemical limitations during photosynthetic induction. We found that elevated O3 significantly reduced the steady-state light-saturated photosynthetic rate, the maximum rate of carboxylation (Vcmax) and Rubisco content. In addition, elevated O3 significantly decreased the time constants for slow phases and weighting of the fast phase of the Vcmax induction in poplar clone '546' but not in clone '107'. However, elevated O3 did not affect the time, it took to reach a given percentage of full Vcmax activation or photosynthetic induction in either clone. Overall, photosynthetic induction was primarily limited by the activity of Rubisco rather than the regeneration of ribulose-1,5-biphosphate regardless of O3 concentration and exposure duration. The lack of O3-induced effects on the activation of Rubisco observed here would simplify the simulation of impacts of O3 on nonsteady-state photosynthesis in dynamic photosynthetic models.

Ozone pollution threatens the production of major staple crops in East Asia.

East Asia is a hotspot of surface ozone (O3) pollution, which hinders crop growth and reduces yields. Here, we assess the relative yield loss in rice, wheat and maize due to O3 by combining O3 elevation experiments across Asia and air monitoring at about 3,000 locations in China, Japan and Korea. China shows the highest relative yield loss at 33%, 23% and 9% for wheat, rice and maize, respectively. The relative yield loss is much greater in hybrid than inbred rice, being close to that for wheat. Total O3-induced annual loss of crop production is estimated at US$63 billion. The large impact of O3 on crop production urges us to take mitigation action for O3 emission control and adaptive agronomic measures against the rising surface O3 levels across East Asia.

[Spatiotemporal Distribution and Health Impacts of PM2.5 and O3 in Beijing, from 2014 to 2020].

In China, fine particulate matter (PM2.5) and tropospheric ozone (O3) have become major air pollutants that threaten human health. Since 2013, the government has strengthened air pollution controls in Beijing and achieved significant effects. A spatial-temporal analysis was conducted of the distribution and health impacts of PM2.5 and O3 in Beijing, using data collected from 34 air quality monitoring sites between 2014 and 2020. In 2014, the annual average PM2.5 and seasonal (April to September) average of daily one-hourly maximum O3 concentrations (O3_max) were 92.0 µg·m-3 and 81.9 nmol·mol-1, respectively. From 2014 to 2020, annual average PM2.5 decreased at a rate of 7.5 µg·m-3. However, there was no significant difference in O3_max over the years. The concentrations of PM2.5 were highest in December and January (in winter) and lowest in August (in summer). On the contrary, O3_max was highest in June. The diurnal variations of PM2.5 were affected by meteorological conditions and emission sources, and maximum concentrations occurred between 22:00 to 00:00, while minimum concentrations occurred between 14:00 to 16:00. The concentration of O3_max showed an opposite pattern, with minimum vales occurring at 07:00 and maximum values occurring in the afternoon. The spatial distribution of PM2.5 showed similar patterns in 2014 and 2019, with the south of Beijing exhibiting the highest concentrations, and the north the lowest. The concentration of O3_max was higher in suburban areas than in traffic areas. In terms of health impacts, 1580 cases of cardiovascular disease and 821 of respiratory disease were attributed to PM2.5 in 2014, while 2180 cases of respiratory disease were attributable to O3 in 2014. In 2019, mortalities attributable to PM2.5 had decreased by 50% compared to 2014. While the number of disease cases attributable to O3 were similar in 2014 and 2019. the results indicate that PM2.5 pollution in Beijing has been successfully controlled, while O3 pollution has become more severe, and was the primary air pollutant threatening human health in 2019. Therefore, the synchronous control of PM2.5 and O3 should be implemented in the future.

Uptake of nitrogen forms by diploid and triploid white poplar depends on seasonal carbon use strategy and elevated summer ozone.

The ability of plants to acquire soil nitrogen (N) sources is plastic in response to abiotic and biotic factors. However, information about how plant preferences among N forms changes in response to internal plant N demand through growth phases, or to environmental stress such as ozone (O3), is scarce. Diploid and triploid Chinese white poplar were used to investigate N form preferences at two key developmental periods (spring, summer) and in response to summer O3 (ambient, 60 ppb above ambient). We used stable isotopes to quantify NH4+, NO3- and glycine N-uptake rates. Carbon acquisition was recorded simultaneously. Both ploidy levels differed in growth, N form preferences, and N and C use strategies. Diploid white poplars grew faster in spring but slower in summer compared with triploids. Diploid white poplars also showed plasticity among N form preferences through the season, with no preferences in spring, and NO3- preferred in summer, while triploids showed an overall preference for NO3-. Carbon acquisition and NO3- uptake were inhibited in both ploidy levels of poplar at elevated O3, which also reduced diploid total N uptake. However, triploid white poplars alleviated N uptake reduction, switching to similar preferences among N forms. We conclude that N form preferences by white poplar are driven by internal C and N use in response to nutrient demands, and external factors such as O3.

Performances of a system for free-air ozone concentration elevation with poplar plantation under increased nitrogen deposition.

The increasing emission of nitrogen oxides exerts large impacts on vegetation by raising surface ozone (O3) concentrations and enhancing atmospheric nitrogen (N) deposition. We established a free-air O3 concentration elevation and enhanced N deposition system (O3-N-FACE) in Beijing, China, to investigate long-term effects of elevated O3 and N deposition on poplar plantation. Eight square plots with a side length of 16 m were randomly allocated to elevated O3 (E-O3) and ambient air (AA) treatments. Ozone generated by electric discharge in pure oxygen is mixed with clean and dry air, and released from small holes on the tubes installed above the plant canopy at a rate controlled to keep O3 concentration in E-O3 plots by 50% higher than that in AA plots. Each O3 treatment plot consisted of four subplots with a factorial combination of 2 lines of poplar clones and 2 levels of N deposition rate. In enhanced N deposition subplots, we sprayed urea solution on the plantation floor at a rate of 60 kg ha-1 year-1. We hereby present the system performances during the growing seasons of 2018 and 2019: the first 2 years of experiment. The mean daytime O3 concentrations of E-O3 plots were 38% and 31% higher than AA plots in 2018 and 2019, respectively. And, in 2019, the accumulated O3 exposure over 40 ppb (AOT40) in E-O3 plots was 70% higher than that in AA plots. The hourly mean O3 concentrations in E-O3 plots were within 20% of the target for 83% of time on average across the four E-O3 plots. Within the E-O3 plots, spatial distribution of the hourly O3 concentration exhibited the maximum deviation at 24% in 2019. We concluded that performance of this system is better than other similar facilities for trees and suitable for a long-term experiment of enhanced O3 and N.

Reduced photosynthetic thermal acclimation capacity under elevated ozone in poplar (Populus tremula) saplings.

The sensitivity of photosynthesis to temperature has been identified as a key uncertainty for projecting the magnitude of the terrestrial carbon cycle response to future climate change. Although thermal acclimation of photosynthesis under rising temperature has been reported in many tree species, whether tropospheric ozone (O3 ) affects the acclimation capacity remains unknown. In this study, temperature responses of photosynthesis (light-saturated rate of photosynthesis (Asat ), maximum rates of RuBP carboxylation (Vcmax ), and electron transport (Jmax ) and dark respiration (Rdark ) of Populus tremula exposed to ambient O3 (AO3 , maximum of 30 ppb) or elevated O3 (EO3 , maximum of 110 ppb) and ambient or elevated temperature (ambient +5°C) were investigated in solardomes. We found that the optimum temperature of Asat (ToptA ) significantly increased in response to warming. However, the thermal acclimation capacity was reduced by O3 exposure, as indicated by decreased ToptA , and temperature optima of Vcmax (ToptV ) and Jmax (ToptJ ) under EO3 . Changes in both stomatal conductance (gs ) and photosynthetic capacity (Vcmax and Jmax ) contributed to the shift of ToptA by warming and EO3 . Neither Rdark measured at 25°C ( R dark 25 ) nor the temperature response of Rdark was affected by warming, EO3 , or their combination. The responses of Asat , Vcmax , and Jmax to warming and EO3 were closely correlated with changes in leaf nitrogen (N) content and N use efficiency. Overall, warming stimulated growth (leaf biomass and tree height), whereas EO3 reduced growth (leaf and woody biomass). The findings indicate that thermal acclimation of Asat may be overestimated if the impact of O3 pollution is not taken into account.

Nonlinear responses of foliar phenylpropanoids to increasing O3 exposure: Ecological implications in a Populus model system.

Plant phenolic compounds (phenylpropanoids) act as defense chemicals against herbivores and can mediate ecosystem processes. Tropospheric ozone (O3) pollution alters concentrations of plant phenolics; however, little is known about how these phytochemicals respond to different levels of O3 exposure. Here, we investigated the effects of five different O3 exposure levels on foliar concentrations of phenylpropanoids (53 compounds in total) and antioxidative capacity in hybrid Populus (Populus euramericana cv. '74/76') saplings grown in the presence of high or low soil nitrogen (N) load. Increasing O3 exposure initially increased and then decreased total concentrations of phenolic compounds, revealing a biphasic exposure-response profile (hormetic zone: 1.1-36.3 ppm h AOT40). This biphasic response pattern was driven by changes in a subset of phenylpropanoids with high antioxidative capacity (e.g. condensed tannins) but not in phenolics with low antioxidative capacity (e.g. salicinoids). The O3 exposure-response relationships of some phenylpropanoids (e.g. flavonoids and chlorogenic acids) varied in response to soil N, with hormesis occurring in high N soil but not in low N soil. Collectively, our findings indicated that plant phenolic compounds exhibit nonlinear responses to increasing O3 exposure, and that the responses vary in relation to phenolic compound class, antioxidative capacity, and soil nitrogen conditions. Our findings further suggest that the impact of O3 on ecological processes mediated by phenolics will be concentration-dependent, highlighting the complexity of the ecological effects of ground-level O3 pollution.

Effects of elevated ozone on maize under varying soil nitrogen levels: Biomass, nitrogen and carbon, and their allocation to kernel.

Effects of ozone (O3) on maize have been increasingly studied, but only few studies have focused on the combined impacts of O3 and nitrogen (N) on this important crop with C4 carbon (C) fixation. In this study, a maize cultivar with the largest acreage in China was exposed to two O3 treatments (NF: ambient air O3 concentration; NF60: NF plus 60 ppb O3) and four N levels (farmers' N practice: 240 kg N ha-1 yr-1; 150%, 50% and 25% of farmers' N practice). Generally, O3 and N significantly influenced biomass, N and C, but did not change their allocation to kernel. There were significant interactions between O3 and N in stem biomass, C concentration and uptake, and leaf biomass and C uptake, with significant O3 effects mainly occurring at N120 and N240. Based on the coefficient of determination (R2), root C:N ratio rather than the most commonly used leaf C:N ratio was the best trait to indicate maize productivity. Furthermore, O3 significantly increased the regression slopes between root C:N ratio and kernel N uptake, kernel C uptake and plant N uptake, strengthened the correlation of C:N ratio and kernel C uptake, and weakened the correlation of C:N ratio and hundred-kernels weight. These suggest that O3 pollution can change the relationship of C:N ratio and productivity in maize. The weak correlation between kernel harvest index (HI) and N harvest index (NHI) indicated that future breeding researches should consider how to improve the coupling between biomass and N-related nutrition allocations in crop edible parts. Our results not only are helpful to accurately estimate O3 impacts on maize with consideration of N but also provide a new insight into the relationship between plant traits and its productivity under O3 pollution.

High nitrogen addition decreases the ozone flux by reducing the maximum stomatal conductance in poplar saplings.

Ground-level ozone (O3) and nitrogen (N) deposition are major environmental pollutants, often occurring concurrently. Ozone exposure- and flux-response relationships for tree biomass are used for regional O3 risk assessment. In order to investigate whether soil N addition affects stomatal O3 uptake of poplar, poplar saplings were exposed to treatment combinations of five O3 levels and four N addition levels. High N addition treatment reduced the accumulated stomatal O3 uptake in the leaf due to reduced maximum stomatal conductance (gs). Nitrogen addition also significantly reduced the steady-state light-saturated gs in August and September. Elevated O3 significantly reduced and N addition increased total plant biomass; however, there were no significant O3 × N interactions. The slopes of biomass-based O3 exposure- and flux-response relationships did not differ significantly among N treatments. The critical levels for a 5% biomass reduction were estimated at 15.4 ppm h and 17.1 mmol O3 m-2 projected leaf area (PLA) for Accumulated O3 exposure Over an hourly Threshold of 40 ppb (AOT40) and Phytotoxic Ozone Dose above a threshold 1 nmol O3 m-2 PLA s-1 (POD1). These results can facilitate the evaluations of O3 effect on the carbon-sink capacity and productivity of forest.

Stomatal response drives between-species difference in predicted leaf water-use efficiency under elevated ozone.

Ozone-induced changes in the relationship between photosynthesis (An) and stomatal conductance (gs) vary among species, leading to inconsistent water use efficiency (WUE) responses to elevated ozone (O3). Thus, few vegetation models can accurately simulate the effects of O3 on WUE. Here, we conducted an experiment exposing two differently O3-sensitive species (Cotinus coggygria and Magnolia denudata) to five O3 concentrations and investigated the impact of O3 exposure on predicted WUE using a coupled An-gs model. We found that increases in stomatal O3 uptake caused linear reductions in the maximum rates of Rubisco carboxylation (Vcmax) and electron transport (Jmax) in both species. In addition, a negative linear correlation between O3-induced changes in the minimal gs of the stomatal model (g0) derived from the theory of optimal stomatal behavior and light-saturated photosynthesis was found in the O3-sensitive M. denudata. When the O3 dose-based responses of Vcmax and Jmax were included in a coupled An-gs model, simulated An under elevated O3 were in good agreement with observations in both species. For M. denudata, incorporating the O3 response of g0 into the coupled model further improved the accuracy of the simulated gs and WUE. In conclusion, the modified Vcmax, Jmax and g0 method presented here provides a foundation for improving the prediction for O3-induced changes in An, gs and WUE.

Impact of ozone pollution on nitrogen fertilization management during maize (Zea mays L.) production.

The impacts of ozone (O3) on crops have been extensively studied and are well understood. However, little information is available on the response of crops (especially maize) to the interactive effects of O3 and nitrogen (N) fertilizer. To this end, a maize cultivar (Zheng dan 958, ZD958) that is common in China was exposed to two O3 treatments and four N levels. We found that (1) the interactions between O3 and N were non-significant for grain yield, plant biomass, C and N, although N addition significantly increased all parameters except C concentrations in grain and plant; (2) compared to NF (non-filtered ambient air O3 concentration), NF60 (NF plus an extra 60 ppb O3) increased the optimum N application rates (Nopt) in grain yield and plant biomass, but not grain yield and plant biomass potentials, thus resulting in lower N use efficiencies (NUE) and a larger risk of N-related environmental pollution (e.g., increased N2O emission) under Nopt in NF60; (3) because of higher optimum plant N uptake (PNopt) in NF60, relative to NF, plant N-saturated conditions for grain yield potential can be gradually turned into N-limited conditions as O3 pollution increases. These findings manifest that O3 is a vital global change factor impacting the management of N fertilization. If current O3 pollution is substantially reduced, maize yield and biomass potentials can be increased under reductions in N input and N-related environmental pollution. In addition, these results can also contribute in developing and verifying Nopt model considering O3 pollution in the future.

Response of isoprene emission from poplar saplings to ozone pollution and nitrogen deposition depends on leaf position along the vertical canopy profile.

We investigated isoprene (ISO) emission and gas exchange in leaves from different positions along the vertical canopy profile of poplar saplings (Populus euramericana cv. '74/76'). For a growing season, plants were subjected to four N treatments, control (NC, no N addition), low N (LN, 50 kg N ha-1year-1), middle N (MN, 100 kg N ha-1year-1), high N (HN, 200 kg N ha-1year-1) and three O3 treatments (CF, charcoal-filtered ambient air; NF, non-filtered ambient air; NF + O3, NF + 40 ppb O3). Our results showed the effects of O3 and/or N on standardized ISO rate (ISOrate) and photosynthetic parameters differed along with the leaf position, with larger negative effects of O3 and positive effects of N on ISOrate and photosynthetic parameters in the older leaves. Expanded young leaves were insensitive to both treatments even at very high O3 concentration (67 ppb as 10-h average) and HN treatment. Significant O3 × N interactions were only found in middle and lower leaves, where ISOrate declined by O3 just when N was limited (NC and LN). With increasing light-saturated photosynthesis and chlorophyll content, ISOrate was reduced in the upper leaves but on the contrary increased in middle and lower leaves. The responses of ISOrate to AOT40 (accumulated exposure to hourly O3 concentrations > 40 ppb) and PODY (accumulative stomatal uptake of O3 > Y nmol O3 m-2 PLA s-1) were not significant in upper leaves, but ISOrate significantly decreased with increasing AOT40 or PODY under limited N supply in middle leaves but at all N levels in lower leaves. Overall, ISOrate changed along the vertical canopy profile in response to combined O3 and N exposure, a behavior that should be incorporated into multi-layer canopy models. Our results are relevant for modelling regional isoprene emissions under current and future O3 pollution and N deposition scenarios.

Effect of elevated ozone, nitrogen availability and mesophyll conductance on the temperature responses of leaf photosynthetic parameters in poplar.

Although ozone (O3) concentration and nitrogen (N) availability are well known to affect plant physiology, their impacts on the photosynthetic temperature response are poorly understood. We addressed this knowledge gap by exposing seedlings of hybrid poplar clone '107' (Populous euramericana cv. '74/76') to elevated O3 (E-O3) and N availability variation in a factorial experiment. E-O3 decreased light-saturated net photosynthesis (Asat), mesophyll conductance (gm) and apparent maximum rate of carboxylation (Vcmax, based on intercellular CO2 concentration) but not actual Vcmax (based on chloroplast CO2 concentration) and increased respiration in light (Rd) at 25 °C. Nitrogen fertilization increased Asat, gm, Vcmax and the maximum rate of electron transport (Jmax) and reduced Rd at 25 °C and the activation energy of actual Vcmax. No E-O3 or E-O3 x N interaction effects on the temperature response parameters were detected, simplifying the inclusion of O3 impacts on photosynthesis in vegetation models. gm peaked at 30 °C, apparent Vcmax and Jmax at 32-33 °C, while the optimum temperatures of actual Vcmax and Jmax exceeded the measured temperature range (15-35 °C). Ignoring gm would, thus, have resulted in mistakenly attributing the decrease in Asat at high temperatures to reduced biochemical capacity rather than to greater diffusion limitation.