Interaction of drought and ozone exposure on isoprene emission from extensively cultivated poplar

The combined effects of ozone (O₃) and drought on isoprene emission were studied for the first time. Young hybrid poplars (clone 546, Populus deltoides cv. 55/56 x P. deltoides cv. Imperial) were exposed to O₃ (charcoal‐filtered air, CF, and non‐filtered air +40 ppb, E‐O₃) and soil water stress (well‐watered, WW, and mild drought, MD, one‐third irrigation) for 96 days. Consistent with light‐saturated photosynthesis (A_sat), intercellular CO₂ concentration (C_i) and chlorophyll content, isoprene emission depended on drought, O₃, leaf position and sampling time. Drought stimulated emission (+38.4%), and O₃ decreased it (?40.4%). Ozone increased the carbon cost per unit of isoprene emission. Ozone and drought effects were stronger in middle leaves (13th–15th from the apex) than in upper leaves (6th–8th). Only A_sat showed a significant interaction between O₃ and drought. When the responses were up‐scaled to the entire‐plant level, however, drought effects on total leaf area translated into around twice higher emission from WW plants in clean air than in E‐O₃. Our results suggest that direct effects on plant emission rates and changes in total leaf area may affect isoprene emission from intensively cultivated hybrid poplar under combined MD and O₃ exposure, with important feedbacks for air quality.     

Impact of elevated ozone concentration on growth,physiology, and yield of wheat (Triticum aestivum L.): a meta‐analysis

We quantitatively evaluated the effects of elevated concentration of ozone (O₃) on growth, leaf chemistry, gas exchange, grain yield, and grain quality relative to carbon‐filtered air (CF) by means of meta‐analysis of published data. Our database consisted of 53 peer‐reviewed studies published between 1980 and 2007, taking into account wheat type, O₃ fumigation method, rooting environment, O₃ concentration ([O₃]), developmental stage, and additional treatments such as drought and elevated carbon dioxide concentration ([CO₂]). The results suggested that elevated [O₃] decreased wheat grain yield by 29% (CI: 24–34%) and aboveground biomass by 18% (CI: 13–24%), where CI is the 95% confidence interval. Even in studies where the [O₃] range was between 31 and 59 ppb (average 43 ppb), there was a significant decrease in the grain yield (18%) and biomass (16%) relative to CF. Despite the increase in the grain protein content (6.8%), elevated [O₃] significantly decreased the grain protein yield (?18%). Relative to CF, elevated [O₃] significantly decreased photosynthetic rates (?20%), Rubisco activity (?19%), stomatal conductance (?22%), and chlorophyll content (?40%). For the whole plant, rising [O₃] induced a larger decrease in belowground (?27%) biomass than in aboveground (?18%) biomass. There was no significant response difference between spring wheat and winter wheat. Wheat grown in the field showed larger decreases in leaf photosynthesis parameters than wheat grown in < 5 L pots. Open‐top chamber fumigation induced a larger reduction than indoor growth chambers, when plants were exposed to elevated [O₃]. The detrimental effect was progressively greater as the average daily [O₃] increased, with very few exceptions. The impact of O₃ increased with developmental stages, with the largest detrimental impact during grain filling. Both drought and elevated [CO₂] significantly ameliorated the detrimental effects of elevated [O₃], which could be explained by a significant decrease in O₃ uptake resulting from decreased stomatal conductance.     

The effect of elevated atmospheric CO2 and drought on sources and sinks of isoprene in a temperate and tropical rainforest mesocosm

Isoprene is the most abundant volatile hydrocarbon emitted by many tree species and has a major impact on tropospheric chemistry, leading to formation of pollutants and enhancing the lifetime of methane, a powerful greenhouse gas. Reliable estimates of global isoprene emission from different ecosystems demand a clear understanding of the processes of both production and consumption. Although the biochemistry of isoprene production has been studied extensively and environmental controls over its emission are relatively well known, the study of isoprene consumption in soil has been largely neglected. Here, we present results on the production and consumption of isoprene studied by measuring the following different components: (1) leaf and soil and (2) at the whole ecosystem level in two distinct enclosed ultraviolet light‐depleted mesocosms at the Biosphere 2 facility: a cottonwood plantation with trees grown at ambient and elevated atmospheric CO₂ concentrations and a tropical rainforest, under well watered and drought conditions. Consumption of isoprene by soil was observed in both systems. The isoprene sink capacity of litter‐free soil of the agriforest stands showed no significant response to different CO₂ treatments, while isoprene production was strongly depressed by elevated atmospheric CO₂ concentrations. In both mesocosms, drought suppressed the sink capacity, but the full sink capacity of dry soil was recovered within a few hours upon rewetting. We conclude that soil uptake of atmospheric isoprene is likely to be modest but significant and needs to be taken into account for a comprehensive estimate of the global isoprene budget. More studies investigating the capacity of soils to uptake isoprene in natural conditions are clearly needed.     

Importance of changing CO2, temperature, precipitation, and ozone on carbon and water cycles of an upland-oak forest: incorporating experimental results into model simulations

Observed responses of upland‐oak vegetation of the eastern deciduous hardwood forest to changing CO₂, temperature, precipitation and tropospheric ozone (O₃) were derived from field studies and interpreted with a stand‐level model for an 11‐year range of environmental variation upon which scenarios of future environmental change were imposed. Scenarios for the year 2100 included elevated [CO₂] and [O₃] (+385 ppm and +20 ppb, respectively), warming (+4°C), and increased winter precipitation (+20% November–March). Simulations were run with and without adjustments for experimentally observed physiological and biomass adjustments. Initial simplistic model runs for single‐factor changes in CO₂ and temperature predicted substantial increases (+191% or 508 g C m^?2 yr^?1) or decreases (?206% or ?549 g C m^?2 yr^?1), respectively, in mean annual net ecosystem carbon exchange (NEE_a≈266±23 g C m^?2 yr^?1 from 1993 to 2003). Conversely, single‐factor changes in precipitation or O₃ had comparatively small effects on NEE_a (0% and ?35%, respectively). The combined influence of all four environmental changes yielded a 29% reduction in mean annual NEE_a. These results suggested that future CO₂‐induced enhancements of gross photosynthesis would be largely offset by temperature‐induced increases in respiration, exacerbation of water deficits, and O₃‐induced reductions in photosynthesis. However, when experimentally observed physiological adjustments were included in the simulations (e.g. acclimation of leaf respiration to warming), the combined influence of the year 2100 scenario resulted in a 20% increase in NEE_a not a decrease. Consistent with the annual model's predictions, simulations with a forest succession model run for gradually changing conditions from 2000 to 2100 indicated an 11% increase in stand wood biomass in the future compared with current conditions. These model‐based analyses identify critical areas of uncertainty for multivariate predictions of future ecosystem response, and underscore the importance of long term field experiments for the evaluation of acclimation and growth under complex environmental scenarios.     

Effects of elevated O3, alone and in combination with elevated CO2, on tree leaf chemistry and insect herbivore performance: a meta-analysis

We reviewed the effects of elevated ozone (O₃), alone and in combination with elevated carbon dioxide (CO₂) on primary and secondary metabolites of trees and performance of insect herbivores by means of meta‐analysis. Our database consisted of 63 studies conducted on 22 species of trees and published between 1990 and 2005. Ozone alone had no overall effect on concentrations of carbohydrates or nutrients, whereas in combination with CO₂, elevated O₃ reduced nutrient concentrations and increased carbohydrate concentrations. In contrast to primary metabolites, concentrations of phenolics and terpenes were significantly increased by 16% and 8%, respectively, in response to elevated O₃. Effects of ozone in combination with elevated CO₂ were weaker than those of ozone alone on phenolics, but stronger than those of ozone alone on terpenes. The magnitude of secondary metabolite responses depended on the type of ozone exposure facility and increased in the following order: indoor growth chamber 3 than gymnosperms, as shifts in concentrations of carbohydrate and phenolics were observed in the former, but not in the latter. Elevated O₃ had positive effects on some indices of insect performance: pupal mass increased and larval development time shortened, but these effects were counteracted by elevated CO₂. Therefore, despite the observed increase in secondary metabolites, elevated O₃ tends to increase tree foliage quality for herbivores, but elevated CO₂ may alleviate these effects. Our meta‐analysis clearly demonstrated that effects of elevated O₃ alone on leaf chemistry and some indices of insect performance differed from those of O₃+CO₂, and therefore, it is important to study effects of several factors of global climate change simultaneously.     

Effects of elevated carbon dioxide, ozone and water availability on spring wheat growth and yield

Spring wheat (Triticum aestivum L. cv. Dragon) was exposed to elevated carbon dioxide (CO₂), alone (1995) or in combination with two levels of increased ozone (O₃) (1994) or increased irrigation (1996) during three successive growing seasons as part of the EU ESPACE‐wheat programme and conducted in open‐top chambers (OTCs) and ambient air (AA) plots at Östad, 50 km north‐east of Göteborg, Sweden. Doubling the CO₂ concentration had a positive effect on grain yield in all 3 years (+21, +7 and +11%, respectively), although only statistically significant in 1994. That year was characterised by a warm and dry summer in comparison with 1995 and 1996, in which the summers were more humid and typical for south‐west Sweden. In 1994, the CO₂‐induced increase in grain yield was associated with an increase in the duration of the green leaf area, a positive effect on straw yield and on the number of ears per square metre and a negative effect (?13%) on grain protein concentration. Harvest index was unaffected by the elevated CO₂ concentration. The only statistically significant effect of elevated CO₂ in 1995 was a decrease in the grain protein concentration (?11% in both CO₂ concentrations), and in 1996 an increase (+21%) in the straw yield. In 1996 the soil water potential was less negative in elevated CO₂, which is likely to reflect a lower water consumption of these plants. Addition of extra O₃ significantly affected the grain yield (?6 and ?10%, respectively) and the 1 000‐grain weight negatively (?3 and ?6%). Statistically significant interactions between CO₂ and O₃ were obtained for the number of ears per unit area and for the 1 000‐grain weight. The 1 000‐grain weight was negatively affected by O₃ in low CO₂, but remained unaffected in the high CO₂ treatment. There was a significant decrease (?6%) in the grain protein concentration induced by elevated irrigation. The chambers, compared with AA plots, had a positive effect on plant development and on grain yield in all 3 years.     

Accumulation of phenolic compounds in birch leaves is changed by elevated carbon dioxide and ozone

Atmospheric change may affect plant phenolic compounds, which play an important part in plant survival. Therefore, we studied the impacts of CO₂ and O₃ on the accumulation of 27 phenolic compounds in the short‐shoot leaves of two European silver birch (Betula pendula Roth) clones (clones 4 and 80). Seven‐year‐old soil‐grown trees were exposed in open‐top chambers over three growing seasons to ambient and twice ambient CO₂ and O₃ concentrations singly and in combination in central Finland. Elevated CO₂ increased the concentration of the phenolic acids (+25%), myricetin glycosides (+18%), catechin derivatives (+13%) and soluble condensed tannins (+19%) by increasing their accumulation in the leaves of the silver birch trees, but decreased the flavone aglycons (?7%) by growth dilution. Elevated O₃ increased the concentration of 3,4′‐dihydroxypropiophenone 3‐β‐d ‐glucoside (+22%), chlorogenic acid (+19%) and flavone aglycons (+4%) by inducing their accumulation possibly as a response to increased oxidative stress in the leaf cells. Nevertheless, this induction of antioxidant phenolic compounds did not seem to protect the birch leaves from detrimental O₃ effects on leaf weight and area, but may have even exacerbated them. On the other hand, elevated CO₂ did seem to protect the leaves from elevated O₃ because all the O₃‐derived effects on the leaf phenolics and traits were prevented by elevated CO₂. The effects of the chamber and elevated CO₂ on some compounds changed over time in response to the changes in the leaf traits, which implies that the trees were acclimatizing to the altered environmental conditions. Although the two clones used possessed different composition and concentrations of phenolic compounds, which could be related to their different latitudinal origin and physiological characteristics, they responded similarly to the treatments. However, in some cases the variation in phenolic concentrations caused by genotype or chamber environment was much larger than the changes caused by either elevated CO₂ or O₃.     

Physiological acclimation dampens initial effects of elevated temperature and atmospheric CO2 concentration in mature boreal Norway spruce

Physiological processes of terrestrial plants regulate the land–atmosphere exchange of carbon, water, and energy, yet few studies have explored the acclimation responses of mature boreal conifer trees to climate change. Here we explored the acclimation responses of photosynthesis, respiration, and stomatal conductance to elevated temperature and/or CO₂ concentration ([CO₂]) in a 3‐year field experiment with mature boreal Norway spruce. We found that elevated [CO₂] decreased photosynthetic carboxylation capacity (?23% at 25 °C) and increased shoot respiration (+64% at 15 °C), while warming had no significant effects. Shoot respiration, but not photosynthetic capacity, exhibited seasonal acclimation. Stomatal conductance at light saturation and a vapour pressure deficit of 1 kPa was unaffected by elevated [CO₂] but significantly decreased (?27%) by warming, and the ratio of intercellular to ambient [CO₂] was enhanced (+17%) by elevated [CO₂] and decreased (?12%) by warming. Many of these responses differ from those typically observed in temperate tree species. Our results show that long‐term physiological acclimation dampens the initial stimulation of plant net carbon assimilation to elevated [CO₂], and of plant water use to warming. Models that do not account for these responses may thus overestimate the impacts of climate change on future boreal vegetation–atmosphere interactions.     

Isoprenoid emission in trees of Quercus pubescens and Quercus ilex with lifetime exposure to naturally high CO2 environment†

The long‐term effect of elevated atmospheric CO₂ on isoprenoid emissions from adult trees of two Mediterranean oak species (the monoterpene‐emitting Quercus ilex L. and the isoprene‐emitting Quercus pubescens Willd.) native to a high‐CO₂ environment was investigated. During two consecutive years, isoprenoid emission was monitored both at branch level, measuring the actual emissions under natural conditions, and at leaf level, measuring the basal emissions under the standard conditions of 30 °C and at light intensity of 1000 µmol m^?2 s^?1. Long‐term exposure to high atmospheric levels of CO₂ did not significantly affect the actual isoprenoid emissions. However, when leaves of plants grown in the control site were exposed for a short period to an elevated CO₂ level by rapidly switching the CO₂ concentration in the gas‐exchange cuvette, both isoprene and monoterpene basal emissions were clearly inhibited. These results generally confirm the inhibitory effect of elevated CO₂ on isoprenoid emission. The absence of a CO₂ effect on actual emissions might indicate higher leaf temperature at elevated CO₂, or an interaction with multiple stresses some of which (e.g. recurrent droughts) may compensate for the CO₂ effect in Mediterranean ecosystems. Under elevated CO₂, isoprene emission by Q. pubescens was also uncoupled from the previous day's air temperature. In addition, pronounced daily and seasonal variations of basal emission were observed under elevated CO₂ underlining that correction factors may be necessary to improve the realistic estimation of isoprene emissions with empirical algorithms in the future. A positive linear correlation of isoprenoid emission with the photosynthetic electron transport and in particular with its calculated fraction used for isoprenoid synthesis was found. The slope of this relationship was different for isoprene and monoterpenes, but did not change when plants were grown in either ambient or elevated CO₂. This suggests that physiological algorithms may usefully predict isoprenoid emission also under rising CO₂ levels.     

Effects of elevated ozone concentration on CH4 and N2O emission from paddy soil under fully open‐air field conditions

We investigated the effects of elevated ozone concentration (E‐O₃) on CH₄ and N₂O emission from paddies with two rice cultivars: an inbred Indica cultivar Yangdao 6 (YD6) and a hybrid one II‐you 084 (IIY084), under fully open‐air field conditions in China. A mean 26.7% enhancement of ozone concentration above the ambient level (A‐O₃) significantly reduced CH₄ emission at tillering and flowering stages leading to a reduction of seasonal integral CH₄ emission by 29.6% on average across the two cultivars. The reduced CH₄ emission is associated with O₃‐induced reduction in the whole‐plant biomass (?13.2%), root biomass (?34.7%), and maximum tiller number (?10.3%), all of which curbed the carbon supply for belowground CH₄ production and its release from submerged soil to atmosphere. Although no significant difference was detected between the cultivars in the CH₄ emission response to E‐O₃, a larger decrease in CH₄ emission with IIY084 (?33.2%) than that with YD6 (?7.0%) was observed at tillering stage, which may be due to the larger reduction in tiller number in IIY084 by E‐O₃. Additionally, E‐O₃ reduced seasonal mean NO_x flux by 5.7% and 11.8% with IIY084 and YD6, respectively, but the effects were not significant statistically. We found that the relative response of CH₄ emission to E‐O₃ was not significantly different from those reported in open‐top chamber experiments. This study has thus confirmed that increasing ozone concentration would mitigate the global warming potential of CH₄ and suggested consideration of the feedback mechanism between ozone and its precursor emission into the projection of future ozone effects on terrestrial ecosystem.     

How does elevated carbon dioxide (CO2) affect plant–herbivore interactions? A field experiment and meta‐analysis of CO2‐mediated changes on plant chemistry and herbivore performance

First, we report the results of the longest‐known field study (9 years) to examine the effects of elevated carbon dioxide (CO₂) on leaf miner densities in a scrub‐oak community at Kennedy Space Center, Florida. Here, the densities of all leaf miner species (6) on all host species (3) were lower in every year in elevated CO₂ than they were in ambient CO₂. Second, meta‐analyses were used to review the effects of elevated CO₂ on both plants (n=59 studies) and herbivores (n=75 studies). The log of the response ratio was chosen as the metric to calculate effect sizes. Results showed that elevated CO₂ significantly decreased herbivore abundance (−21.6%), increased relative consumption rates (+16.5%), development time (+3.87%) and total consumption (+9.2%), and significantly decreased relative growth rate (−8.3%), conversion efficiency (−19.9%) and pupal weight (−5.03%). No significant differences were observed among herbivore guilds. Host plants growing under enriched CO₂ environments exhibited significantly larger biomass (+38.4%), increased C/N ratio (+26.57%), and decreased nitrogen concentration (−16.4%), as well as increased concentrations of tannins (+29.9%) and other phenolics. Effects of changes on plant primary and secondary chemistry due to elevated CO₂ and consequences for herbivore growth and development are discussed.     

30 years of free‐air carbon dioxide enrichment (FACE): What have we learned about future crop productivity and its potential for adaptation?

Free‐air CO₂ enrichment (FACE) allows open‐air elevation of [CO₂] without altering the microclimate. Its scale uniquely supports simultaneous study from physiology and yield to soil processes and disease. In 2005 we summarized results of then 28 published observations by meta‐analysis. Subsequent studies have combined FACE with temperature, drought, ozone, and nitrogen treatments. Here, we summarize the results of now almost 250 observations, spanning 14 sites and five continents. Across 186 independent studies of 18 C₃ crops, elevation of [CO₂] by ca. 200 ppm caused a ca. 18% increase in yield under non‐stress conditions. Legumes and root crops showed a greater increase and cereals less. Nitrogen deficiency reduced the average increase to 10%, as did warming by ca. 2°C. Two conclusions of the 2005 analysis were that C₄ crops would not be more productive in elevated [CO₂], except under drought, and that yield responses of C₃ crops were diminished by nitrogen deficiency and wet conditions. Both stand the test of time. Further studies of maize and sorghum showed no yield increase, except in drought, while soybean productivity was negatively affected by early growing season wet conditions. Subsequent study showed reduced levels of nutrients, notably Zn and Fe in most crops, and lower nitrogen and protein in the seeds of non‐leguminous crops. Testing across crop germplasm revealed sufficient variation to maintain nutrient content under rising [CO₂]. A strong correlation of yield response under elevated [CO₂] to genetic yield potential in both rice and soybean was observed. Rice cultivars with the highest yield potential showed a 35% yield increase in elevated [CO₂] compared to an average of 14%. Future FACE experiments have the potential to develop cultivars and management strategies for co‐promoting sustainability and productivity under future elevated [CO₂].     

Nitrification,denitrification, and related functional genes under elevated CO2: A meta-analysis in terrestrial ecosystems

Global change may have profound effects on soil nitrogen (N) cycling that can induce positive feedback to climate change through increased nitrous oxide (N₂O) emissions mediated by nitrification and denitrification. We conducted a meta-analysis of the effects of elevated CO₂ on nitrification and denitrification based on 879 observations from 58 publications and 46 independent elevated CO₂ experiments in terrestrial ecosystems. We investigated the effects of elevated CO₂ alone or combined with elevated temperature, increased precipitation, drought, and N addition. We assessed the response to elevated CO₂ of gross and potential nitrification, potential denitrification, and abundances of related functional genes (archaeal amoA, bacterial amoA, nirK, nirS, and nosZ). Elevated CO₂ increased potential nitrification (+28%) and the abundance of bacterial amoA functional gene (+62%) in cropland ecosystems. Elevated CO₂ increased potential denitrification when combined with N addition and higher precipitation (+116%). Elevated CO₂ also increased the abundance of nirK (+25%) and nirS (+27%) functional genes in terrestrial ecosystems and of nosZ (+32%) functional gene in cropland ecosystems. The increase in the abundance of nosZ under elevated CO₂ was larger at elevated temperature and high N (+62%). Four out of 14 two-way interactions tested between elevated CO₂ and elevated temperature, elevated CO₂ and increased precipitation, and elevated CO₂ and N addition were marginally significant and mostly synergistic. The effects of elevated CO₂ on potential nitrification and abundances of bacterial amoA and nirS functional genes increased with mean annual temperature and mean annual precipitation. Our meta-analysis thus suggests that warming and increased precipitation in large areas of the world could reinforce positive responses of nitrification and denitrification to elevated CO₂ and urges the need for more investigations in the tropical zone and on interactive effects among multiple global change factors, as we may largely underestimate the effects of global change on soil N₂O emissions.     

Increased temperatures may safeguard the nutritional quality of crops under future elevated CO2 concentrations

Iron (Fe) and zinc (Zn) deficiencies are a global human health problem that may worsen by the growth of crops at elevated atmospheric CO₂ concentration (eCO₂). However, climate change will also involve higher temperature, but it is unclear how the combined effect of eCO₂ and higher temperature will affect the nutritional quality of food crops. To begin to address this question, we grew soybean (Glycine max) in a Temperature by Free‐Air CO₂ Enrichment (T‐FACE) experiment in 2014 and 2015 under ambient (400 μmol mol^?1) and elevated (600 μmol mol^?1) CO₂ concentrations, and under ambient and elevated temperatures (+2.7°C day and +3.4°C at night). In our study, eCO₂ significantly decreased Fe concentration in soybean seeds in both seasons (?8.7 and ?7.7%) and Zn concentration in one season (?8.9%), while higher temperature (at ambient CO₂ concentration) had the opposite effect. The combination of eCO₂ with elevated temperature generally restored seed Fe and Zn concentrations to levels obtained under ambient CO₂ and temperature conditions, suggesting that the potential threat to human nutrition by increasing CO₂ concentration may not be realized. In general, seed Fe concentration was negatively correlated with yield, suggesting inherent limitations to increasing seed Fe. In addition, we confirm our previous report that the concentration of seed storage products and several minerals varies with node position at which the seeds developed. Overall, these results demonstrate the complexity of predicting climate change effects on food and nutritional security when various environmental parameters change in an interactive manner.     

Genotypic differences in leaf biochemical, physiological and growth responses to ozone in 20 winter wheat cultivars released over the past 60 years

Ozone (O₃) concentrations in periurban areas in East Asia are sufficiently high to decrease crop yield. However, little is known about the genotypic differences in O₃ sensitivity in winter wheat in relation to year of cultivar release. This paper reports genotypic variations in O₃ sensitivity in 20 winter wheat cultivars released over the past 60 years in China highlighting O₃‐induced mechanisms. Wheat plants were exposed to elevated O₃ (82 ppb O₃, 7 h day^?1) or charcoal‐filtered air (<5 ppb O₃) for 21 days in open top chambers. Responses to O₃ were assessed by the levels of antioxidative activities, protein alteration, membrane lipid peroxidation, gas exchange, leaf chlorophyll, dark respiration and growth. We found that O₃ significantly reduced foliar ascorbate (?14%) and soluble protein (?22%), but increased peroxidase activity (+46%) and malondialdehyde (+38%). Elevated O₃ depressed light saturated net photosynthetic rate (?24%), stomatal conductance (?8%) and total chlorophyll (?11%), while stimulated dark respiration (+28%) and intercellular CO₂ concentration (+39%). O₃ also reduced overall plant growth, but to a greater extent in root (?32%) than in shoot (?17%) biomass. There was significant genotypic variation in potential sensitivity to O₃ that did not correlate to observed O₃ tolerance. Sensitivity to O₃ in cultivars of winter wheat progressed with year of release and correlated with stomatal conductance and dark respiration in O₃‐exposed plants. O₃‐induced loss in photosynthetic rate was attributed primarily to impaired activity of mesophyll cells and loss of integrity of cellular membrane as evidenced by increased intercellular CO₂ concentration and lipid peroxidation. Our findings demonstrated that higher sensitivity to O₃ in the more recently released cultivars was induced by higher stomatal conductance, larger reduction in antioxidative capacity and lower levels of dark respiration leading to higher oxidative damage to proteins and integrity of cellular membranes.     

Profiles of isoprene emission and photosynthetic parameters in hybrid poplars exposed to free‐air CO2 enrichment†

Poplar (Populus × euroamericana) saplings were grown in the field to study the changes of photosynthesis and isoprene emission with leaf ontogeny in response to free air carbon dioxide enrichment (FACE) and soil nutrient availability. Plants growing in elevated [CO₂] produced more leaves than those in ambient [CO₂]. The rate of leaf expansion was measured by comparing leaves along the plant profile. Leaf expansion and nitrogen concentration per unit of leaf area was similar between nutrient treatment, and this led to similar source–sink functional balance. Consequently, soil nutrient availability did not cause downward acclimation of photosynthetic capacity in elevated [CO₂] and did not affect isoprene synthesis. Photosynthesis assessed in growth [CO₂] was higher in plants growing in elevated than in ambient [CO₂]. After normalizing for the different number of leaves over the profile, maximal photosynthesis was reached and started to decline earlier in elevated than in ambient [CO₂]. This may indicate a [CO₂]‐driven acceleration of leaf maturity and senescence. Isoprene emission was adversely affected by elevated [CO₂]. When measured on the different leaves of the profile, isoprene peak emission was higher and was reached earlier in ambient than in elevated [CO₂]. However, a larger number of leaves was emitting isoprene in plant growing in elevated [CO₂]. When integrating over the plant profile, emissions in the two [CO₂] levels were not different. Normalization as for photosynthesis showed that profiles of isoprene emission were remarkably similar in the two [CO₂] levels, with peak emissions at the centre of the profile. Only the rate of increase of the emission of young leaves may have been faster in elevated than in ambient [CO₂]. Our results indicate that elevated [CO₂] may overall have a limited effect on isoprene emission from young seedlings and that plants generally regulate the emission to reach the maximum at the centre of the leaf profile, irrespective of the total leaf number. In comparison with leaf expansion and photosynthesis, isoprene showed marked and repeatable differences among leaves of the profile and may therefore be a useful trait to accurately monitor changes of leaf ontogeny as a consequence of elevated [CO₂].     

Atmospheric CO2 and O3 alter competition for soil nitrogen in developing forests

Plant growth responses to rising atmospheric CO₂ and O₃ vary among genotypes and between species, which could plausibly influence the strength of competitive interactions for soil N. Ascribable to the size‐symmetric nature of belowground competition, we reasoned that differential growth responses to CO₂ and O₃ should shift as juvenile individuals mature, thereby altering competitive hierarchies and forest composition. In a 12‐year‐long forest FACE experiment, we used tracer ¹⁵N and whole‐plant N content to assess belowground competitive interactions among five Populus tremuloides genotypes, between a single P. tremuloides genotype and Betula papryrifera, as well as between the same single P. tremuloides genotype and Acer saccharum. Under elevated CO₂, the amount of soil N and ¹⁵N obtained by the P. tremuloides genotype common to each community was contingent on the nature of belowground competition. When this genotype competed with its congeners, it obtained equivalent amounts of soil N and tracer ¹⁵N under ambient and elevated CO₂; however, its acquisition of soil N under elevated CO₂ increased by a significant margin when grown in competition with B. papyrifera (+30%) and A. saccharum (+60%). In contrast, elevated O₃ had no effect on soil N and ¹⁵N acquisition by the P. tremuloides genotype common in each community, regardless of competitive interactions. Under elevated CO₂, the rank order of N acquisition among P. tremuloides genotypes shifted over time, indicating that growth responses to CO₂ change during ontogeny; this was not the case under elevated O₃. In the aspen‐birch community, the competitive advantage elevated CO₂ initially conveyed on birch diminished over time, whereas maple was a poor competitor for soil N in all regards. The extent to which elevated CO₂ and O₃ will shape the genetic structure and composition of future forests is, in part, contingent on the time‐dependent effects of belowground competition on plant growth response.     

Interactive effects of elevated CO2 and soil fertility on isoprene emissions from Quercus robur

The effects of global change on the emission rates of isoprene from plants are not clear. A factor that can influence the response of isoprene emission to elevated CO₂ concentrations is the availability of nutrients. Isoprene emission rate under standard conditions (leaf temperature: 30°C, photosynthetically active radiation (PAR): 1000 μmol photons m^?2 s^?1), photosynthesis, photosynthetic capacity, and leaf nitrogen (N) content were measured in Quercus robur grown in well‐ventilated greenhouses at ambient and elevated CO₂ (ambient plus 300 ppm) and two different soil fertilities. The results show that elevated CO₂ enhanced photosynthesis but leaf respiration rates were not affected by either the CO₂ or nutrient treatments. Isoprene emission rates and photosynthetic capacity were found to decrease with elevated CO₂, but an increase in nutrient availability had the converse effect. Leaf N content was significantly greater with increased nutrient availability, but unaffected by CO₂. Isoprene emission rates measured under these conditions were strongly correlated with photosynthetic capacity across the range of different treatments. This suggests that the effects of CO₂ and nutrient levels on allocation of carbon to isoprene production and emission under near‐saturating light largely depend on the effects on photosynthetic electron transport capacity.     

Enhanced isoprene emission capacity and altered light responsiveness in aspen grown under elevated atmospheric CO2 concentration

Controversial evidence of CO₂‐responsiveness of isoprene emission has been reported in the literature with the response ranging from inhibition to enhancement, but the reasons for such differences are not understood. We studied isoprene emission characteristics of hybrid aspen (Populus tremula x P. tremuloides) grown under ambient (380 μmol mol^?1) and elevated (780 μmol mol^?1) [CO₂] to test the hypothesis that growth [CO₂] effects on isoprene emission are driven by modifications in substrate pool size, reflecting altered light use efficiency for isoprene synthesis. A novel in vivo method for estimation of the pool size of the immediate isoprene precursor, dimethylallyldiphosphate (DMADP) and the activity of isoprene synthase was used. Growth at elevated [CO₂] resulted in greater leaf thickness, more advanced development of mesophyll and moderately increased photosynthetic capacity due to morphological “upregulation”, but isoprene emission rate under growth light and temperature was not significantly different among ambient‐ and elevated‐[CO₂]‐grown plants independent of whether measured at 380 μmol mol^?1 or 780 μmol mol^?1 CO₂. However, DMADP pool size was significantly less in elevated‐[CO₂]‐grown plants, but this was compensated by increased isoprene synthase activity. Analysis of CO₂ and light response curves of isoprene emission demonstrated that the [CO₂] for maximum isoprene emission was shifted to lower [CO₂] in elevated‐[CO₂]‐grown plants. The light‐saturated isoprene emission rate (I_max,Q) was greater, but the quantum efficiency at given I_max,Q was less in elevated‐[CO₂]‐grown plants, especially at higher CO₂ measurement concentration, reflecting stronger DMADP limitation at lower light and higher [CO₂]. These results collectively demonstrate important shifts in light and CO₂‐responsiveness of isoprene emission in elevated‐[CO₂]‐acclimated plants that need consideration in modeling isoprene emissions in future climates.     

Stress-induced changes in carbon sources for isoprene production in Populus deltoides

Isoprene is emitted from leaves of numerous plant species and has important implications for plant metabolism and atmospheric chemistry. The ability to use stored carbon (alternative carbon sources), as opposed to recently assimilated photosynthate, for isoprene production may be important as plants routinely experience photosynthetic depression in response to environmental stress. A CO₂‐labelling study was performed and stable isotopes of carbon were used to examine the role of alternative carbon sources in isoprene production in Populus deltoides during conditions of water stress and high leaf temperature. Isotopic fractionation during isoprene production was higher in heat‐ and water‐stressed leaves (?8.5 and ?9.3‰, respectively) than in unstressed controls (?2.5 to ?3.2‰). In unstressed plants, 84–88% of the carbon in isoprene was derived from recently assimilated photosynthate. A significant shift in the isoprene carbon composition from photosynthate to alternative carbon sources was observed only under severe photosynthetic limitation (stomatal conductance < 0.05 mol m^?2 s^?1). The contribution of photosynthate to isoprene production decreased to 77 and 61% in heat‐ and water‐stressed leaves, respectively. Across water‐ and heat‐stress experiments, allocation of photosynthate was negatively correlated to the ratio of isoprene emission to photosynthesis. In water‐stressed plants, the use of alternative carbon was also related to stomatal conductance. It has been proposed that isoprene emission may be regulated by substrate availability. Thus, understanding carbon partitioning to isoprene production from multiple sources is essential for building predictive models of isoprene emission.