Climatic conditions and concentrations of carbon dioxide and air pollutants during ‘ESPACE—wheat’ experiments

A major objective of the ESPACE—wheat programme was to perform by means of open-top chambers (OTCs) ‘standardised’ experimental investigations of spring wheat responses to increased atmospheric CO₂ and O₃ concentrations and to other environmental stresses at different locations in Europe, representing a broad range of different climatic conditions. From 1994 to 1996 a total number of 25 OTC experiments were carried out. In addition, four growth chamber experiments focusing on key physiological processes of wheat growth in CO₂-enriched air were performed. According to the specific needs for subsequent modelling purposes, environmental data were collected during experiments, i.e. air temperature, global radiation, humidity and trace gas concentrations. In the present paper results of these measurements are summarised. It was shown, that the OTC-experiments covered a considerable range of growing season mean-air-temperatures (13.0–23.4°C) and global irradiances (10.8–18.1 MJ m⁻² d⁻¹), the most important driving variables for crop growth simulation models. Mean concentrations of CO₂ and O₃ in ambient air and in different treatments illustrated the observed variability of trace gas exposures between different experiments. Implications for subsequent analyses of biological response data are discussed.     

The effects of climatic variation in Europe on the yield response of spring wheat cv. Minaret to elevated CO2 and O3: an analysis of open-top chamber experiments by means of two crop growth simulation models

In the ESPACE-Wheat programme, 25 open-top chamber experiments were carried out in 1994, 1995 and 1996, on nine locations, divided over eight European countries. In most experiments, spring wheat cv. Minaret was subjected to two levels of atmospheric CO₂ and two levels of ozone. Grain yields in the control treatments (ambient levels of CO₂ and O₃) varied strongly between sites. Also, yield response to elevated CO₂ and O₃ showed great variation. The present study was conducted to determine whether climatic differences between sites could account for the observed variation.

Two simulation models were used for the analysis: AFRCWHEAT2-O3 and LINTULCC. AFRCWHEAT2-O3 simulates phenology, canopy development and photosynthesis in greater detail than LINTULCC. Both models account for the effects of radiation and temperature on crop growth. New algorithms were developed to simulate the effects of CO₂ and O₃. Weather data that were measured in the experiments were used as input, and simulated growth responses to CO₂ and O₃ were compared with measurements. No attempt was made to merge the two models. Thus two independent tools for analysis of data related to climate change were developed and applied.

The average measured grain yield in the control treatment, across all 25 experiments, was 5.9 tons per hectare (t ha⁻¹), with a standard deviation (SD) of 1.9 t  ha⁻¹. The models predicted similar average yields (5.5 and 5.8 t ha⁻¹ for AFRCWHEAT2-O3 and LINTULCC, respectively), but smaller variation (SD for both models: 1.2 t ha⁻¹). Average measured yield increase due to CO₂-doubling was 30% (SD 22%). AFRCWHEAT2-O3 expected a slightly lower value (24%, SD 9%), whereas LINTULCC overestimated the response (42%, SD 11%). The average measured yield decrease due to nearly-doubled O₃ levels was 9% (SD 11%). Both models showed similar results, albeit at lower variation (7% yield decrease at SDs of 6 and 4%). Simulations accounted well for the observation that, at elevated CO₂, the percentage yield loss due to O₃ was lower than at ambient CO₂.

The models predicted lower variation among sites and years than was measured. Yield response to CO₂ and O₃ was predicted to depend on the climate, with a predominant effect of temperature on the response to CO₂. In the measurements, these climatic effects were indeed observed, but a greater part of the variation was not related to light intensity, temperature, CO₂, or O₃. This unexplained variability in the measured dataset was probably caused by factors not accounted for in the models, possibly related to soil characteristics.

We therefore conclude that even perfect information on the climate variables examined in ESPACE-Wheat, i.e. light intensity and temperature, by itself would be insufficient for accurate prediction of the response of spring wheat to future elevated levels of CO₂ and O₃.     

Phenological development, leaf emergence, tillering and leaf area index, and duration of spring wheat across Europe in response to CO2 and ozone

Phenological development, leaf emergence, tillering and leaf area index (LAI), and duration (LAD) of spring wheat cv. Minaret, grown in open-top chambers at different sites throughout Europe for up to 3 years at each site, were investigated in response to elevated CO₂ (ambient CO₂×2) and ozone (ambient ozone ×1.5) concentrations.

Phenological development varied among experiments and was partly explained by differences in temperature among sites and years. There was a weak positive relationship between the thermal rate of development and the mean daylength for the period from emergence to anthesis. Main stems produced on average 7.7 leaves with little variation among experiments. Variation was higher for the thermal rate of leaf emergence, which was partly explained by differences in the rate of change of daylength at plant emergence among seasons. Phenological development, rate of leaf emergence and final leaf number were not affected by CO₂ and ozone exposure. Responses of tillering and LAI to CO₂ and ozone exposure were significant only in some experiments. However, the direction of responses was consistent for most experiments. The number of tillers and ears per plant, respectively, was increased as a result of CO₂ enrichment by about 13% at the beginning of stem elongation (DC31), at anthesis and at maturity. Exposure to ozone had no effect on tillering. LAI was increased as a result of CO₂ elevation by about 11% at DC31 and by about 14% at anthesis. Ozone exposure reduced LAI at anthesis by about 9%. No such effect was observed at DC31. There were very few interactive effects of CO₂ and ozone on tillering and LAI. Variations in tillering and LAI, and their responses to CO₂ and ozone exposure, were partly explained by single linear relationships considering differences in plant density, tiller density and the duration of developmental phases among experiments. Consideration of temperature and incident photosynthetically active radiation in this analysis did not reduce the unexplained variation. There was a negative effect of ozone exposure on leaf area duration at most sites. Direct effects of elevated CO₂ concentration on leaf senescence, both positive and negative, were observed in some experiments. There was evidence in several experiments that elevated CO₂ concentration ameliorated the negative effect of ozone on leaf area duration. It was concluded from these results that an analysis of the interactive effects of climate, CO₂ and ozone on canopy development requires reference to the physiological processes involved.     

Photosynthetic responses in spring wheat grown under elevated CO2 concentrations and stress conditions in the European, multiple-site experiment ‘ESPACE-wheat’

Spring wheat cv. Minaret crop stands were grown under ambient and elevated CO₂ concentrations at seven sites in Germany, Ireland, the UK, Belgium and the Netherlands. Six of the sites used open-top chambers and one used a controlled environment mimicking field conditions. The effect of elevated CO₂ for a range of N application regimes, O₃ concentrations, and growth temperatures on flag leaf photosynthesis was studied. Before anthesis, flag leaf photosynthesis was stimulated about 50% by 650 compared with 350 μmol mol⁻¹ CO₂ at all sites, regardless of other treatments. Furthermore, there was no evidence of a decrease in photosynthetic capacity of flag leaves due to growth at elevated CO₂ before anthesis, even for low N treatments. However, photosynthetic capacity, particularly carboxylation capacity, of flag leaves was usually decreased by growth at elevated CO₂ after anthesis, especially in low N treatments. Acclimation of photosynthesis to elevated CO₂ therefore appears to occur only slowly, consistent with a response to changes in sink–source relationships, rather than a direct response. Effect of elevated CO₂ on stomatal conductance was much more variable between sites and treatments, but on average was decreased by ˜10% at 650 compared with 350 μmol mol⁻¹ CO₂. Carboxylation capacity of flag leaves was decreased by growth at elevated O₃ both before and after anthesis, regardless of CO₂ concentration.     

Effects on nutrients and on grain quality in spring wheat crops grown under elevated CO2 concentrations and stress conditions in the European, multiple-site experiment ‘ESPACE-wheat’

Nutrient element concentrations and grain quality were assessed in spring wheat grown under elevated CO₂ concentrations and contrasting levels of tropospheric ozone at different nitrogen supply rates at several European sites. Carbon dioxide enrichment proved to affect nutrient concentrations in a complex manner. In green leaves, all elements (with exception of phosphorus and iron) decreased. In contrast, effects on the element composition of grains were restricted to reductions in nitrogen, calcium, sulphur and iron. Ozone exposure resulted in no significant effects on nutrient element concentrations in different tissues in the overall analysis. The nitrogen demand of green tissues was reduced due to CO₂ enrichment as shown by reductions in the critical leaf nitrogen concentration and also enhanced nitrogen use efficiency. Reductions in the content of ribulose-bisphosphate carboxylase/oxygenase and repression of the photorespiratory pathway and reduced nitrogen allocation to enzymes driving the photosynthetic carbon oxidation cycle were chiefly responsible for this effect. Thus, nitrogen acquisition by the crop did not match carbon acquisition under CO₂ enrichment. Since crop nitrogen uptake from the soil was already completed at anthesis, nitrogen allocated to the grain after anthesis originated from vegetative pools—causing grain nitrogen concentrations to decrease under CO₂ enrichment (on average by 15% when CO₂ concentrations increased from 360 to 680 μmol mol⁻¹). Correspondingly, grain quality was reduced by CO₂ enrichment. The Zeleny value, Hagberg value and dry/wet gluten content decreased significantly with increasing [CO₂]. Despite the beneficial impact of CO₂ enrichment on growth and yield of C₃ cereal crops, declines in flour quality due to reduced nitrogen content are likely in a future, [CO₂]-rich world.     

Effects of elevated carbon dioxide and ozone on potato tuber quality in the European multiple-site experiment ‘CHIP-project’

Potato (Solanum tuberosum L cv. Bintje) was exposed to ambient and elevated carbon dioxide (CO₂), to ambient and elevated ozone (O₃) and to elevated levels of both gases during two growing seasons, 1998 and 1999. Experiments in open-top chambers (OTC) were carried out in Finland, Sweden, Ireland, United Kingdom, Germany and Belgium and a FACE (Free Air Carbon dioxide Enrichment) experiment was carried out in Italy. In OTCs the plants were grown under ambient CO₂ concentrations or with 550 and 680 μl l⁻¹ CO₂ alone or in combination with ambient or elevated O₃ concentrations (target seasonal mean of 60 nl l⁻¹ 8 h per day). In the FACE systems the plants were exposed to ambient or 550 μl l⁻¹ CO₂. In the OTC experiments the reducing sugar content of potato tubers decreased significantly with increased concentration of O₃. The starch content of potato tubers decreased, with negative impact on tuber quality, but the ascorbic acid concentration increased as a function of the AOT40 (The sum of the differences between hourly ozone concentration and 40 nl l⁻¹ for each hour when the concentration exceeds 40 nl l⁻¹ during a relevant growing season). However, simultaneous exposure to elevated CO₂ counteracted the ozone effect. With increase in the CO₂ exposure, glycoalkaloid and nitrate concentrations decreased yielding improved quality, while the citric acid concentration decreased causing a higher risk for discoloration after cooking. The amount of dry matter and starch increased significantly in the FACE experiment.     

Effect of climatic conditions on tuber yield (Solanum tuberosum L.) in the European ‘CHIP’ experiments

The main objective of the CHIP project was to perform ‘standardised’ investigations of potato (Solanum tuberosum L. cv Bintje) responses to increased O₃ and CO₂ concentrations by means of open-top chambers (OTC) and free air carbon dioxide enrichment (FACE) systems. The experimental sites are located across Europe representing a broad range of different climatic conditions. In 1998 and 1999 a total number of 12 OTC experiments and four FACE experiments were conducted. According to the specific needs for subsequent modelling purposes, environmental data were collected during experiments, i.e. air temperature, global radiation, air humidity (vapour pressure deficit (VPD)), soil moisture and trace gas concentrations. In the present paper, the results of these measurements are summarised. It was shown that the experiments covered a considerable range of growing season mean air temperatures (13.8–19.9 °C) and global irradiances (12.0–21.3 MJ m⁻² per day), the most important driving variables for crop growth simulation models. Analysis of the soils used during the experiments demonstrated that in most cases sufficient nutrient elements were available to guarantee an undisturbed growth. Mean concentrations of CO₂ and O₃ in ambient air and in different treatments illustrate the observed variability of trace gas exposures between different sites and experiments. However, the effects of these parameters on growth and yield are subject of separate papers. The general climatic conditions across Europe are also causing important growth and yield effects. Comparison of marketable tuber yields revealed an increase at higher latitudes. This result was associated with lower temperatures and VPD and longer day lengths at the higher latitudes, which in turn were associated with longer growing seasons.     

Chlorophyll content of spring wheat flag leaves grown under elevated CO2 concentrations and other environmental stresses within the ‘ESPACE-wheat’ project

Spring wheat cv. Minaret was grown in open-top chambers at four sites across Europe. The effect of different treatments (CO₂ enrichment, O₃ fumigation, drought stress and temperature) on the chlorophyll content of the flag leaf was investigated using the MINOLTA SPAD-502 meter. Under optimum growth conditions the maximum chlorophyll content, which was reached at anthesis, was consistent among the sites ranging from 460 to 500 mg chlorophyll m⁻². No significant effect of elevated CO₂ or O₃ was observed at anthesis. Leaf senescence, indicated by the chlorophyll breakdown after anthesis, was relatively constant in the control chambers. Under control conditions, thermal time until 50% chlorophyll loss was reached was 600°C day. Elevated CO₂ caused a faster decline in chlorophyll content (thermal time until 50% chlorophyll loss was reduced to 500–580°C day) indicating a faster rate of plant development at two experimental sites. The effect of ozone on chlorophyll content depended on the time and dose of O₃ exposure. During grain filling, high O₃ concentrations induced premature senescence of the flag leaves (up to −130°C day). This deleterious effect was mitigated by elevated CO₂. Drought stress led to faster chlorophyll breakdown irrespective of CO₂ treatment.     

Changing climate and potential impacts on potato yield and quality ‘CHIP’: introduction, aims and methodology

Climate change effects caused by an increasing concentration of CO₂ and ozone represent an issue of major concern both for scientists and policy-makers. In a concerted program funded by the Commission of the European Union, a European network of experiments (in open-top chambers (OTC), and free air carbon dioxide enrichment systems (FACE)) and modelling was carried out to investigate the effects of increasing atmospheric CO₂ and ozone concentrations, under different climatic conditions, on potato (Solanum tuberosum L. cv. Bintje). This contribution describes the experimental network and the standard protocol set-up for the assessments that served to improve and to validate process oriented potato growth simulation models leading to scenarios of future productivity of potato in Europe.     

Simulation of growth and development processes of spring wheat in response to CO2 and ozone for different sites and years in Europe using mechanistic crop simulation models

The response of crop growth and yield to CO₂ and ozone is known to depend on climatic conditions and is difficult to quantify due to the complexity of the processes involved. Two modified mechanistic crop simulation models (AFRCWHEAT2-O3 and LINTULCC), which differ in the levels of mechanistic detail, were used to simulate the effects of CO₂ (ambient, ambient ×2) and ozone (ambient, ambient ×1.5) on growth and developmental processes of spring wheat in response to climatic conditions. Simulations were analysed using data from the ESPACE-wheat project in which spring wheat cv. Minaret was grown in open-top chambers at nine sites throughout Europe and for up to 3 years at each site.

Both models closely predicted phenological development and the average measured biomass at maturity. However, intermediate growth variables such as biomass and leaf area index (LAI) at anthesis, seasonal accumulated photosynthetically active radiation intercepted by the crop (ΣIPAR), the average seasonal light use efficiency (LUE) and the light saturated rate of flag leaf photosynthesis (A_sat) were predicted differently and less accurately by the two models. The effect of CO₂ on the final biomass was underestimated by AFRCWHEAT2-O3 due to its poor simulation of the effect of CO₂ on tillering, and LAI.LINTULCC overestimated the response of biomass production to changes in CO₂ level due to an overprediction of the effect of CO₂ on LUE. The measured effect of ozone exposure on final biomass was predicted closely by the two models. The models also simulated the observed interactive effect of CO₂ and ozone on biomass production. However, the effects of ozone on LAI, ΣIPAR and A_sat were simulated differently by the models and less accurately with LINTULCC for the ozone effects on LAI and ΣIPAR. Predictions of the variation between sites and years of growth and development parameters and of their responses to CO₂ and ozone were poor for both AFRCWHEAT2-O3 and LINTULCC. It was concluded that other factors than those considered in the models such as chamber design and soil properties may have affected the growth and development of cv. Minaret. An analysis of the relationships between growth parameters calculated from the simulations supported this conclusion. In order to apply models for global change impact assessment studies, the difficulties in simulating biomass production in response to CO₂ need to be considered. We suggest that the simulation of leaf area dynamics deserves particular attention in this regard.     

The European Stress Physiology and Climate Experiment—project 1: wheat (ESPACE-wheat): introduction, aims and methodology

The response of crops to CO₂ enrichment represents an issue of major concern both for scientists and for policy-makers. In a concerted programme funded by the Commission of the European Communities, a Europe-wide experimental and modeling study was carried out to investigate the effects of increasing atmospheric CO₂ concentrations, and of environmental stresses such as ozone or water/nutrient shortage, under different climatic conditions on wheat (Triticum aestivum L.). This contribution describes the experimental network and the standard protocol set-up for the assessments which served to improve and to validate process-orientated wheat growth simulation models.