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.     

Effects of elevated CO2 and O3 on the rate and duration of grain growth and harvest index in spring wheat (Triticum aestivum L.)

Wheat (Triticum aestivum L.) cv. Minaret was grown in open-top chambers (OTCs) in 1995 and 1996 under three carbon dioxide (CO₂) and two ozone (O₃) levels. Plants were harvested regularly between anthesis and maturity to examine the rate of grain growth (dG/dt; mg d^–1) and the rate of increase in harvest index (dHI/dt;% d^–1). The duration of grain filling was not affected by elevated CO₂ or O₃, but was 12 days shorter in 1995, when the daily mean temperature was over 3 °C higher than in 1996. Season-long exposure to elevated CO₂ (680 μmol mol^–1) significantly increased the rate of grain growth in both years and mean grain weight at maturity (MGW) was up to 11% higher than in the chambered ambient air control (chAA; 383 μmol mol^–1). However, the increase in final yield obtained under elevated CO₂ relative to the chAA control in 1996 resulted primarily from a 27% increase in grain number per unit ground area. dG/dt was significantly reduced by elevated O₃ under ambient CO₂ conditions in 1995, but final grain yield was not affected because of a concurrent increase in grain number. Neither dG/dt nor dHI/dt were affected by the higher mean O₃ concentrations applied in 1996 (77 vs. 66 nmol mol^–1); the differing effects of O₃ on grain growth in 1995 and 1996 observed in both the ambient and elevated CO₂ treatments may reflect the contrasting temperature environments experienced. Grain yield was nevetheless reduced under elevated O₃ in 1996, primarily because of a substantial decrease in grain number. The data obtained show that, although exposure to elevated CO₂ and O₃ individually or in combination may affect both dG/dt and dHI/dt, the presence of elevated CO₂ does not protect against substantial O₃-induced yield losses resulting from its direct deleterious impact on reproductive processes. The implications of these results for food production under future climatic conditions are considered.     

Yield vs. Quality trade‐offs for wheat in response to carbon dioxide and ozone

Although it is established that there exist potential trade‐offs between grain yield and grain quality in wheat exposed to elevated carbon dioxide (CO₂) and ozone (O₃), their underlying causes remain poorly explored. To investigate the processes affecting grain quality under altered CO₂ and O₃, we analysed 57 experiments with CO₂ or O₃ exposure in different exposure systems. The study covered 24 cultivars studied in 112 experimental treatments from 11 countries. A significant growth dilution effect on grain protein was found: a change in grain yield of 10% by O₃ was associated with a change in grain protein yield of 8.1% (R² = 0.96), whereas a change in yield effect of 10% by CO₂ was linked to a change in grain protein yield effect of 7.5% (R² = 0.74). Superimposed on this effect, elevated CO₂, but not O₃, had a significant negative effect on grain protein yield also in the absence of effects on grain yield, indicating that there exists a process by which CO₂ restricts grain protein accumulation, which is absent for O₃. Grain mass, another quality trait, was more strongly affected by O₃ than grain number, whereas the opposite was true for CO₂. Harvest index was strongly and negatively influenced by O₃, but was unaffected by CO₂. We conclude that yield vs. protein trade‐offs for wheat in response to CO₂ and O₃ are constrained by close relationships between effects on grain biomass and less than proportional effects on grain protein. An important and novel finding was that elevated CO₂ has a direct negative effect on grain protein accumulation independent of the yield effect, supporting recent evidence of CO₂‐induced impairment of nitrate uptake/assimilation. Finally, our results demonstrated that processes underlying responses of grain yield vs. quality trade‐offs are very different in wheat exposed to elevated O₃ compared with elevated CO₂.     

Growth dynamics and genotypic variation in tropical, field-grown paddy rice (Oryza sativa L.) in response to increasing carbon dioxide and temperature

While previous studies have examined the growth and yield response of rice to continued increases in CO₂ concentration and potential increases in air temperature, little work has focused on the long-term response of tropical paddy rice (i.e. the bulk of world rice production) in situ, or genotypic differences among cultivars in response to increasing CO₂ and/or temperature. At the International Rice Research Institute, rice (cv IR72) was grown from germination until maturity for 4 field seasons, the 1994 and 1995 wet and the 1995 and 1996 dry seasons at three different CO₂ concentrations (ambient, ambient + 200 and ambient + 300 μL L^–1 CO₂) and two air temperatures (ambient and ambient + 4 °C) using open-top field chambers placed within a paddy site. Overall, enhanced levels of CO₂ alone resulted in significant increases in total biomass at maturity and increased seed yield with the relative degree of enhancement consistent over growing seasons across both temperatures. Enhanced levels of temperature alone resulted in decreases or no change in total biomass and decreased seed yield at maturity across both CO₂ levels. In general, simultaneous increases in air temperature as well as CO₂ concentration offset the stimulation of biomass and grain yield compared to the effect of CO₂ concentration alone. For either the 1995 wet and 1996 dry seasons, additional cultivars (N-22, NPT1 and NPT2) were grown in conjunction with IR72 at the same CO₂ and temperature treatments. Among the cultivars tested, N-22 showed the greatest relative response of both yield and biomass to increasing CO₂, while NPT2 showed no response and IR72 was intermediate. For all cultivars, however, the combination of increasing CO₂ concentration and air temperature resulted in reduced grain yield and declining harvest index compared to increased CO₂ alone. Data from these experiments indicate that (a) rice growth and yield can respond positively under tropical paddy conditions to elevated CO₂, but that simultaneous exposure to elevated temperature may negate the CO₂ response to grain yield; and, (b) sufficient intraspecific variation exists among cultivars for future selection of rice cultivars which may, potentially, convert greater amounts of CO₂ into harvestable yield.     

Elevated ozone reduces photosynthetic carbon gain by accelerating leaf senescence of inbred and hybrid maize in a genotype‐specific manner

Exposure to elevated tropospheric ozone concentration ([O₃]) accelerates leaf senescence in many C₃ crops. However, the effects of elevated [O₃] on C₄ crops including maize (Zea mays L.) are poorly understood in terms of physiological mechanism and genetic variation in sensitivity. Using free air gas concentration enrichment, we investigated the photosynthetic response of 18 diverse maize inbred and hybrid lines to season‐long exposure to elevated [O₃] (~100 nl L^?1) in the field. Gas exchange was measured on the leaf subtending the ear throughout the grain filling period. On average over the lifetime of the leaf, elevated [O₃] led to reductions in photosynthetic CO₂ assimilation of both inbred (?22%) and hybrid (?33%) genotypes. There was significant variation among both inbred and hybrid lines in the sensitivity of photosynthesis to elevated [O₃], with some lines showing no change in photosynthesis at elevated [O₃]. Based on analysis of inbred line B73, the reduced CO₂ assimilation at elevated [O₃] was associated with accelerated senescence decreasing photosynthetic capacity and not altered stomatal limitation. These findings across diverse maize genotypes could advance the development of more O₃ tolerant maize and provide experimental data for parameterization and validation of studies modeling how O₃ impacts crop performance.     

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.     

Water availability moderates N2 fixation benefit from elevated [CO2]: A 2‐year free‐air CO2 enrichment study on lentil (Lens culinaris MEDIK.) in a water limited agroecosystem

Increased biomass and yield of plants grown under elevated [CO₂] often corresponds to decreased grain N concentration ([N]), diminishing nutritional quality of crops. Legumes through their symbiotic N₂ fixation may be better able to maintain biomass [N] and grain [N] under elevated [CO₂], provided N₂ fixation is stimulated by elevated [CO₂] in line with growth and yield. In Mediterranean‐type agroecosystems, N₂ fixation may be impaired by drought, and it is unclear whether elevated [CO₂] stimulation of N₂ fixation can overcome this impact in dry years. To address this question, we grew lentil under two [CO₂] (ambient ~400 ppm and elevated ~550 ppm) levels in a free‐air CO₂ enrichment facility over two growing seasons sharply contrasting in rainfall. Elevated [CO₂] stimulated N₂ fixation through greater nodule number (+27%), mass (+18%), and specific fixation activity (+17%), and this stimulation was greater in the high than in the low rainfall/dry season. Elevated [CO₂] depressed grain [N] (?4%) in the dry season. In contrast, grain [N] increased (+3%) in the high rainfall season under elevated [CO₂], as a consequence of greater post‐flowering N₂ fixation. Our results suggest that the benefit for N₂ fixation from elevated [CO₂] is high as long as there is enough soil water to continue N₂ fixation during grain filling.     

Response of wheat restricted‐tillering and vigorous growth traits to variables of climate change

The response of wheat to the variables of climate change includes elevated CO₂, high temperature, and drought which vary according to the levels of each variable and genotype. Independently, elevated CO₂, high temperature, and terminal drought affect wheat biomass and grain yield, but the interactive effects of these three variables are not well known. The aim of this study was to determine the effects of elevated CO₂ when combined with high temperature and terminal drought on the high‐yielding traits of restricted‐tillering and vigorous growth. It was hypothesized that elevated CO₂ alone, rather than combined with high temperature, ameliorates the effects of terminal drought on wheat biomass and grain yield. It was also hypothesized that wheat genotypes with more sink capacity (e.g. high‐tillering capacity and leaf area) have more grain yield under combined elevated CO₂, high temperature, and terminal drought. Two pairs of sister lines with contrasting tillering and vigorous growth were grown in poly‐tunnels in a four‐factor completely randomized split‐plot design with elevated CO₂ (700 µL L^?1), high day time temperature (3 °C above ambient), and drought (induced from anthesis) in all combinations to test whether elevated CO₂ ameliorates the effects of high temperature and terminal drought on biomass accumulation and grain yield. For biomass and grain yield, only main effects for climate change variables were significant. Elevated CO₂ significantly increased grain yield by 24–35% in all four lines and terminal drought significantly reduced grain yield by 16–17% in all four lines, while high temperature (3 °C above the ambient) had no significant effect. A trade‐off between yield components limited grain yield in lines with greater sink capacity (free‐tillering lines). This response suggests that any positive response to predicted changes in climate will not overcome the limitations imposed by the trade‐off in yield components.     

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.     

CO2 enrichment reduces the relative contribution of latex and latex-related hydrocarbons to biomass in Euphorbia lathyris

The hypothesis that plants grown under elevated CO₂ allocate more carbon to the production of latex and C‐rich secondary compounds whereas nutrient addition counteracts this effect was tested. Two similar experiments were conducted in two different experimental facilities. In both facilities seedlings of Euphorbia lathyris were exposed to factorial combinations of two CO₂ concentrations and two levels of nutrient availability for 2 months. The CO₂ treatments and growth conditions differed substantially between these two experiments but treatment responses to elevated CO₂ and fertilizer addition were remarkably similar, underlining the robustness of our findings. Elevated CO₂ increased biomass to a greater extent in fertilized than in unfertilized plants and reduced the leaf biomass fraction by accelerating leaf senescence. Concentrations of non‐structural carbohydrates (NSC) increased in elevated CO₂. However, this apparent carbon surplus did not feed into the whole plant latex pool. The latex harvest per leaf (?25%) and the concentration of latex‐related hydrocarbons (?20%) even decreased under elevated CO₂ (both experiments P < 0.05). Fertilization reduced NSC concentrations (?25%) but neither affected latex yield per leaf nor the concentration of latex‐related hydrocarbons. It is concluded that latex and related hydrocarbons in CO₂‐enriched plants are a negligible sink for excess carbon irrespective of nutrient status and thus, vigour of growth.     

The influence of elevated CO2 and O3 on fine roots and mycorrhizas of naturally growing young Scots pine trees during three exposure years

Young Scots pine trees naturally established at a pine heath were exposed to two concentrations of CO₂ (ambient and doubled ambient) and two O₃ regimes (ambient and doubled ambient) and their combination in open-top field chambers during growing seasons 1994, 1995 and 1996 (late May to 15 September). Filtered ozone treatment and chamberless control trees were also included in the treatment comparisons. Root ingrowth cores were inserted to the undisturbed soil below the branch projection of each tree at the beginning of the fumigation period in 1994 and were harvested at the end of the fumigation periods in 1995 and 1996. Root biomasses were determined from different soil layers in the ingrowth cores, and the infection levels of different mycorrhizal types were calculated. Elevated O₃ and CO₂ did not have significant effects on the biomass production of Scots pine coarse (Ø > 2 mm) or fine roots (Ø < 2 mm) and roots of grasses and dwarf shrubs. Elevated O₃ caused a transient stimulation, observable in 1995, in the proportion of tuber-like mycorrhizas, total mycorrhizas and total short roots but this stimulation disappeared during the last study year. Elevated CO₂ did not enhance carbon allocation to root growth or mycorrhiza formation, although a diminishing trend in the mycorrhiza formation was observed. In the combination treatment increased CO₂ inhibited the transient stimulating effect of ozone, and a significant increase of old mycorrhizas was observed. Our conclusion is that doubled CO₂ is not able to increase carbon allocation to growth of fine roots or mycorrhizas in nutrient poor forest sites and realistically elevated ozone does not cause a measurable limitation to roots within a period of three exposure years.     

Ozone and density affect the response of biomass and seed yield to elevated CO2 in rice

Tropospheric O₃ reduces growth and yield of many crop species, whereas CO₂ ameliorates the negative effects of O₃. Thus, in a combined elevated CO₂ and O₃ atmosphere, seed yield is at least restored to that of charcoal‐filtered (CF) air at ambient CO₂. The CO₂‐induced yield increase in CF air is highly variable, suggesting other potential resource limitations. To understand such variability in response, we tested that (1) competition for resources precludes some of the CO₂ enhancement on biomass and yield; and (2) O₃ reduces competition in elevated CO₂. We grew rice (Oryza sativa L.) at five densities in CF and O₃‐fumigated (+O3) air at ambient (A) and elevated [CO₂] (+CO2) in 1997 and 1998. O₃ reduced biomass by 25% and seed yield by 13–20% in A, but had little effect in +CO2. A competition model of biomass and yield response to density based on resource availability without competition showed that fewer resources were used for biomass in +O3 than in CF (average 53% vs. 70%) in A, while in +CO2 85% of resources were used for biomass regardless of O₃ suggesting greater depletion of resources. The enhanced biomass response to CO₂ with O₃ is consistent with a 22% greater CO₂ enhancement ratio [mass in +CO2 air/mass in A air; enhancement ratio (ER)] in +O3 than in CF air. For seed yield, few resources were used (average 17% and 25% for CF in 1997 and 1998, respectively), and ER was 13% greater in +O3. With competition the rate of change of individual plant biomass to density was not affected by +CO2 in CF air in 1997 but was increased 19% with more nutrients in 1998, indicating resource limitations with +CO2. The rate of change of individual plant yield to density was reduced with CO₂ in 1997 and unchanged in 1998 showing a different response to resource limitation for reproductive biomass. The resource use in +O3‐A suggested that increased density and soil fertility might compensate for pollutant damage. Although ambient [O₃] can modulate the response to elevated CO₂, resource limitation precludes the CO₂ fertilization impact and both factors need consideration for better management and forecasts of future productivity.     

Isoprene is more affected by climate drivers than monoterpenes: A meta‐analytic review on plant isoprenoid emissions

Isoprene and monoterpenes (MTs) are among the most abundant and reactive volatile organic compounds produced by plants (biogenic volatile organic compounds). We conducted a meta‐analysis to quantify the mean effect of environmental factors associated to climate change (warming, drought, elevated CO₂, and O₃) on the emission of isoprene and MTs. Results indicated that all single factors except warming inhibited isoprene emission. When subsets of data collected in experiments run under similar change of a given environmental factor were compared, isoprene and photosynthesis responded negatively to elevated O₃ (?8% and ?10%, respectively) and drought (?15% and ?42%), and in opposite ways to elevated CO₂ (?23% and +55%) and warming (+53% and ?23%, respectively). Effects on MTs emission were usually not significant, with the exceptions of a significant stimulation caused by warming (+39%) and by elevated O₃ (limited to O₃‐insensitive plants, and evergreen species with storage organs). Our results clearly highlight individual effects of environmental factors on isoprene and MT emissions, and an overall uncoupling between these secondary metabolites produced by the same methylerythritol 4‐phosphate pathway. Future results from manipulative experiments and long‐term observations may help untangling the interactive effects of these factors and filling gaps featured in the current meta‐analysis.     

The influence of elevated CO2 and O3 concentrations on Scots pine needles: changes in starch and secondary metabolites over three exposure years

Scots pine (Pinus sylvestris L.) trees, aged about 20 years old, growing on a natural pine heath were exposed to two concentrations of CO₂ (ambient CO₂ and double-ambient CO₂) and two O₃ regimes (ambient O₃ and double-ambient O₃) and their combination in open-top chambers during growing seasons 1994, 1995 and 1996. Concentrations of foliar starch and secondary compounds are reported in this paper. Starch concentrations remained unaffected by elevated CO₂ and/or O₃ concentrations during the first 2 study years. But in the autumn of the last study year, a significantly higher concentration of starch was found in current-year needles of trees exposed to elevated CO₂ compared with ambient air. There were large differences in concentrations of starch and secondary compounds between individual trees. Elevated concentrations of CO₂ and/or O₃ did not have any significant effects on the concentrations of foliar total monoterpenes, total resin acids or total phenolics. Significantly higher concentrations of monoterpenes and resin acids and mostly lower concentrations of starch were found in trees growing without chambers than in those growing in open-top chambers, while there were no differences in concentrations of total phenolics between trees growing without or in chambers. The results suggest that elevated concentrations of CO₂ might increase foliar starch concentrations in Scots pine, while secondary metabolites remain unaffected. Realistically elevated O₃ concentrations do not have clear effects on carbon allocation to starch and secondary compounds even after 3 exposure years. Received: 2 June 1997 / Accepted: 12 December 1997     

Does elevated atmospheric carbon dioxide affect internal nitrogen allocation in the temperate trees Alnus glutinosa and Pinus sylvestris?

Nitrogen‐fixing plant species growing in elevated atmospheric carbon dioxide concentration ([CO₂]) should be able to maintain a high nutrient supply and thus grow better than other species. This could in turn engender changes in internal storage of nitrogen (N) and remobilisation during periods of growth. In order to investigate this one‐year‐old‐seedlings of Alnus glutinosa (L.) Gaertn and Pinus sylvestris (L.) were exposed to ambient [CO₂] (350 µ mol mol ^{? 1}) and elevated [CO₂] (700 µ mol mol ^{? 1}) in open top chambers (OTCs). This constituted a main comparison between a nitrogen‐fixing tree and a nonfixer, but also between an evergreen and a deciduous species. The trees were supplied with a full nutrient solution and in July 1994, the trees were given a pulse of ¹⁵N‐labelled fertiliser. The allocation of labelled N to different tissues (root, leaves, shoots) was followed from September 1994 to June 1995. While N allocation in P. sylvestris (Scots pine) showed no response to elevated [CO₂], A. glutinosa (common alder) responded in several ways. During the main nutrient uptake period of June–August, trees grown in elevated [CO₂] had a higher percentage of N derived from labelled fertiliser than trees grown in ambient [CO₂]. Remobilisation of labelled N for spring growth was significantly higher in A. glutinosa grown in elevated [CO₂] (9.09% contribution in ambient vs. 29.93% in elevated [CO₂] leaves). Exposure to elevated [CO₂] increased N allocation to shoots in the winter of 1994–1995 (12.66 mg in ambient vs. 43.42 mg in elevated 1993 shoots; 4.81 mg in ambient vs. 40.00 mg in elevated 1994 shoots). Subsequently significantly more labelled N was found in new leaves in April 1995. These significant increases in movement of labelled N between tissues could not be explained by associated increases in tissue biomass, and there was a significant shift in C‐biomass allocation away from the leaves towards the shoots (all above‐ground material except leaves) in A. glutinosa. This experiment provides the first evidence that not only are shifts in C allocation affected by elevated [CO₂], but also internal N resource utilisation in an N₂‐fixing tree.     

Elevated carbon dioxide and ozone alter productivity and ecosystem carbon content in northern temperate forests

Three young northern temperate forest communities in the north‐central United States were exposed to factorial combinations of elevated carbon dioxide (CO₂) and tropospheric ozone (O₃) for 11 years. Here, we report results from an extensive sampling of plant biomass and soil conducted at the conclusion of the experiment that enabled us to estimate ecosystem carbon (C) content and cumulative net primary productivity (NPP). Elevated CO₂ enhanced ecosystem C content by 11%, whereas elevated O₃ decreased ecosystem C content by 9%. There was little variation in treatment effects on C content across communities and no meaningful interactions between CO₂ and O₃. Treatment effects on ecosystem C content resulted primarily from changes in the near‐surface mineral soil and tree C, particularly differences in woody tissues. Excluding the mineral soil, cumulative NPP was a strong predictor of ecosystem C content (r² = 0.96). Elevated CO₂ enhanced cumulative NPP by 39%, a consequence of a 28% increase in canopy nitrogen (N) content (g N m^?2) and a 28% increase in N productivity (NPP/canopy N). In contrast, elevated O₃ lowered NPP by 10% because of a 21% decrease in canopy N, but did not impact N productivity. Consequently, as the marginal impact of canopy N on NPP (?NPP/?N) decreased through time with further canopy development, the O₃ effect on NPP dissipated. Within the mineral soil, there was less C in the top 0.1 m of soil under elevated O₃ and less soil C from 0.1 to 0.2 m in depth under elevated CO₂. Overall, these results suggest that elevated CO₂ may create a sustained increase in NPP, whereas the long‐term effect of elevated O₃ on NPP will be smaller than expected. However, changes in soil C are not well‐understood and limit our ability to predict changes in ecosystem C content.     

Growth, light interception and yield responses of spring wheat (Triticum aestivum L.) grown under elevated CO2 and O3 in open-top chambers

Spring wheat cv. Minaret was grown to maturity under three carbon dioxide (CO₂) and two ozone (O₃) concentrations in open-top chambers (OTC). Green leaf area index (LAI) was increased by elevated CO₂ under ambient O₃ conditions as a direct result of increases in tillering, rather than individual leaf areas. Yellow LAI was also greater in the 550 and 680 μmol mol^–1 CO₂ treatments than in the chambered ambient control; individual leaves on the main shoot senesced more rapidly under 550 μmol mol^–1 CO₂, but senescence was delayed at 680 μmol mol^–1 CO₂. Fractional light interception (f) during the vegetative period was up to 26% greater under 680 μmol mol^–1 CO₂ than in the control treatment, but seasonal accumulated intercepted radiation was only increased by 8%. As a result of greater carbon assimilation during canopy development, plants grown under elevated CO₂ were taller at anthesis and stem and ear biomass were 27 and 16% greater than in control plants. At maturity, yield was 30% greater in the 680 μmol mol^–1 CO₂ treatment, due to a combination of increases in the number of ears per m^–2, grain number per ear and individual grain weight (IGW). Exposure to a seasonal mean (7 h d^–1) of 84 nmol mol^–1 O₃ under ambient CO₂ decreased green LAI and increased yellow LAI, thereby reducing both f and accumulated intercepted radiation by ≈ 16%. Individual leaves senesced completely 7–28 days earlier than in control plants. At anthesis, the plants were shorter than controls and exhibited reductions in stem and ear biomass of 15 and 23%. Grain yield at maturity was decreased by 30% due to a combination of reductions in ear number m^–2, the numbers of grains per spikelet and per ear and IGW. The presence of elevated CO₂ reduced the rate of O₃-induced leaf senescence and resulted in the maintenance of a higher green LAI during vegetative growth under ambient CO₂ conditions. Grain yields at maturity were nevertheless lower than those obtained in the corresponding elevated CO₂ treatments in the absence of elevated O₃. Thus, although the presence of elevated CO₂ reduced the damaging impact of ozone on radiation interception and vegetative growth, substantial yield losses were nevertheless induced. These data suggest that spring wheat may be susceptible to O₃-induced injury during anthesis irrespective of the atmospheric CO₂ concentration. Possible deleterious mechanisms operating through effects on pollen viability, seed set and the duration of grain filling are discussed.     

Stem wood properties of Populus tremuloides, Betula papyrifera and Acer saccharum saplings after 3 years of treatments to elevated carbon dioxide and ozone

The aim of this study was to examine the effects of elevated carbon dioxide [CO₂] and ozone [O₃] and their interaction on wood chemistry and anatomy of five clones of 3‐year‐old trembling aspen (Populus tremuloides Michx.). Wood chemistry was studied also on paper birch (Betula papyrifera Marsh.) and sugar maple (Acer saccharum Marsh.) seedling‐origin saplings of the same age. Material for the study was collected from the Aspen Free‐Air CO₂ Enrichment (FACE) experiment in Rhinelander, WI, USA, where the saplings had been exposed to four treatments: control (C; ambient CO₂, ambient O₃), elevated CO₂ (560 ppm during daylight hours), elevated O₃ (1.5 × ambient during daylight hours) and their combination (CO₂+O₃) for three growing seasons (1998–2000). Wood chemistry responses to the elevated CO₂ and O₃ treatments differed between species. Aspen was most responsive, while maple was the least responsive of the three tree species. Aspen genotype affected the responses of wood chemistry and, to some extent, wood structure to the treatments. The lignin concentration increased under elevated O₃ in four clones of aspen and in birch. However, elevated CO₂ ameliorated the effect. In two aspen clones, nitrogen in wood samples decreased under combined exposure to CO₂ and O₃. Soluble sugar concentration in one aspen clone and starch concentration in two clones were increased by elevated CO₂. In aspen wood, α‐cellulose concentration changed under elevated CO₂, decreasing under ambient O₃ and slightly increasing under elevated O₃. Hemicellulose concentration in birch was decreased by elevated CO₂ and increased by elevated O₃. In aspen, elevated O₃ induced statistically significant reductions in distance from the pith to the bark and vessel lumen diameter, as well as increased wall thickness and wall percentage, and in one clone, decreased fibre lumen diameter. Our results show that juvenile wood properties of broadleaves, depending on species and genotype, were altered by atmospheric gas concentrations predicted for the year 2050 and that CO₂ ameliorates some adverse effects of elevated O₃ on wood chemistry.     

Influence of carbon dioxide enrichment, ozone and nitrogen fertilization on cotton (Gossypium hirsutum L.) leaf and root composition

The objective of this study was to test whether elevated [CO₂], [O₃] and nitrogen (N) fertility altered leaf mass per area (LMPA), non‐structural carbohydrate (TNC), N, lignin (LTGA) and proanthocyanidin (PA) concentrations in cotton (Gossypium hirsutum L.) leaves and roots. Cotton was grown in 14 dm³ pots with either sufficient (0·8 g N dm ^{? 3}) or deficient (0·4 and 0·2 g N dm ^{? 3}) N fertilization, and treated in open‐top chambers with either ambient or elevated ( + 175 and + 350 μ mol mol ^{? 1}) [CO₂] in combination with either charcoal‐filtered air (CF) or non‐filtered air plus 1·5 times ambient [O₃]. At about 50 d after planting, LMPA, starch and PA concentrations in canopy leaves were as much as 51–72% higher in plants treated with elevated [CO₂] compared with plants treated with ambient [CO₂], whereas leaf N concentration was 29% lower in elevated [CO₂]‐treated plants compared with controls. None of the treatments had a major effect on LTGA concentrations on a TNC‐free mass basis. LMPA and starch levels were up to 48% lower in plants treated with elevated [O₃] and ambient [CO₂] compared with CF controls, although the elevated [O₃] effect was diminished when plants were treated concurrently with elevated [CO₂]. On a total mass basis, leaf N and PA concentrations were higher in samples treated with elevated [O₃] in ambient [CO₂], but the difference was much reduced by elevated [CO₂]. On a TNC‐free basis, however, elevated [O₃] had little effect on tissue N and PA concentrations. Fertilization treatments resulted in higher PA and lower N concentrations in tissues from the deficient N fertility treatments. The experiment showed that suppression by elevated [O₃] of LMPA and starch was largely prevented by elevated [CO₂], and that interpretation of [CO₂] and [O₃] effects should include comparisons on a TNC‐free basis. Overall, the experiment indicated that allocation to starch and PA may be related to how environmental factors affect source–sink relationships in plants, although the effects of elevated [O₃] on secondary metabolites differed in this respect.     

Effects of elevated temperature and carbon dioxide on seed-set and yield of kidney bean (Phaseolus vulgaris L.)

It is important to quantify and understand the consequences of elevated temperature and carbon dioxide (CO₂) on reproductive processes and yield to develop suitable agronomic or genetic management for future climates. The objectives of this research work were (a) to quantify the effects of elevated temperature and CO₂ on photosynthesis, pollen production, pollen viability, seed‐set, seed number, seeds per pod, seed size, seed yield and dry matter production of kidney bean and (b) to determine if deleterious effects of high temperature on reproductive processes and yield could be compensated by enhanced photosynthesis at elevated CO₂ levels. Red kidney bean cv. Montcalm was grown in controlled environments at day/night temperatures ranging from 28/18 to 40/30 °C under ambient (350 µmol mol^?1) or elevated (700 µmol mol^?1) CO₂ levels. There were strong negative relations between temperature over a range of 28/18–40/30 °C and seed‐set (slope, ? 6.5% °C^?1) and seed number per pod (? 0.34 °C^?1) under both ambient and elevated CO₂ levels. Exposure to temperature > 28/18 °C also reduced photosynthesis (? 0.3 and ? 0.9 µmol m^?2 s^?1 °C^?1), seed number (? 2.3 and ? 3.3 °C^?1) and seed yield (? 1.1 and ? 1.5 g plant^?1 °C^?1), at both the CO₂ levels (ambient and elevated, respectively). Reduced seed‐set and seed number at high temperatures was primarily owing to decreased pollen production and pollen viability. Elevated CO₂ did not affect seed size but temperature > 31/21 °C linearly reduced seed size by 0.07 g °C^?1. Elevated CO₂ increased photosynthesis and seed yield by approximately 50 and 24%, respectively. There was no beneficial interaction of CO₂ and temperature, and CO₂ enrichment did not offset the negative effects of high temperatures on reproductive processes and yield. In conclusion, even with beneficial effects of CO₂ enrichment, yield losses owing to high temperature (> 34/24 °C) are likely to occur, particularly if high temperatures coincide with sensitive stages of reproductive development.