Growth and marketable-yield responses of potato to increased CO2 and ozone

Central to the CHanging climate and potential Impacts on Potato yield and quality project (CHIP) was the consideration of the potential impacts of ozone and CO₂ on growth and yield of future European Potato crops. Potato crops, cv. Bintje, were exposed to ambient or elevated ozone; targeted daily average, 60 nl l⁻¹ for 8 h, and ambient or elevated CO_2; targeted 680 μl l⁻¹ averaged over the full growing season, in open top chambers (OTCs) at six European sites in 1998 and 1999, or to elevated CO₂ (550 μl l⁻¹) in Free Air Carbon dioxide Enrichment facilities (FACE) at two sites in both years. Some OTC experiments included 550 μl l⁻¹. Above and below ground biomass were measured at two destructive harvests; at maximum leaf area (MLA) and at final-harvest. Final-harvest fresh weight yields of marketable-size tubers, >35 mm diameter, from ambient conditions ranged from 1 to 12 kg m⁻². There was no consistent (P>0.1) CO₂×O₃ interaction for growth or yield variables at either harvest. No consistent effects of ozone were detected at the maximum-leaf-area harvest. However, at final harvest, ozone had reduced both above-ground biomass and tuber dry weight (P<0.05), particularly of the largest (>50 mm) size class. These yield losses showed linear relationships both with accumulated ozone exposure; AOT40 expressed as nl l⁻¹ h over 40 nl l⁻¹, and with yields from chambered ambient-ozone treatments (P<0.05) but, because of partial confounding between the treatment AOT40s and the ambient-ozone yields in the data, the two relationships were not completely independent. Yields from ambient-ozone treatments, however, explained a significant (P<0.01) amount of the residual variation in ozone effects unexplained by AOT40. When averaged over all experiments, mean dry weights and tuber numbers from both harvests were increased by elevated CO₂. Only green leaf number at the MLA harvest was reduced. The CO₂ responses varied between sites and years. For marketable-size tubers, this variation was unrelated to variation in ambient-CO₂ treatment yields. Yield increases resulting from the 680 μl l⁻¹ and 550 μl l⁻¹ treatments were similar. Thus elevating [CO₂] from 550 to 680 μl l⁻¹ was less effective than elevating [CO₂] from ambient to 550 μl l⁻¹. On average, CO₂ elevation to 680 μl l⁻¹ increased the dry weight of marketable-size tubers by about 17%, which far exceeded the average ozone-induced yield loss of about 5%. The net effect of raising CO₂ and O₃ concentrations on the European potato crop would be an increase marketable yield.     

CO2 and ozone effects on canopy development of potato crops across Europe

This paper describes the effects of elevated CO₂ (550 and 680 μl l⁻¹) and O₃ (60 nl l⁻¹ O₃ as an 8 h mean), alone or in combination, on canopy development and senescence in potato (Solanum tuberosum L. cv Bintje) across a range of European agro-climatic conditions. The assessments were made within the European CHIP project (CHanging climate and potential Impacts on Potato yield and quality) that was conducted for two growing seasons (1998 and 1999) in free air CO₂ enrichment systems (FACE) and open-top chamber facilities (OTCs) at seven European sites. A comparison of chambered and unchambered experimental plots was included to examine the effects of chamber enclosure. Phenological growth stages, plant height, leaf area index (LAI) and the number of green and yellow leaves were recorded non-destructively throughout the growing season and by a destructive intermediate harvest at maximum leaf area (MLA). In the dynamic growth analysis CO₂ and O₃ effects were studied over three developmental stages: canopy expansion, full canopy and canopy senescence. Chamber enclosures promoted potato crop development (taller plants, more leaves) during the initial growth stages and led to a faster decline of LAI and a higher number of yellow leaves. The growth in ambient plots varied between sites and seasons, as did the scale of the treatment responses. Despite the large background variation, some overall treatment effects could be detected across all sites. Both levels of increased CO₂ reduced final plant height in comparison to ambient concentrations, which indicates a premature ending of the active plant growth. At the stage of full canopy and crop senescence the average number of green leaves was significantly (P<0.05) decreased by 680 μl l⁻¹ CO₂ (OTC experiments) and LAI showed the same tendency (P=0.07). As there was however no indication of a decreased leaf formation during initial growth and at full canopy, this must have been due to an earlier leaf fall. In the FACE experiments LAI had already began to decline at the stage of full canopy at 550 μl l⁻¹ CO₂ but not in ambient CO₂ (DAE×CO₂, P<0.05). These observations strongly indicated that elevated CO₂ induced a premature senescence during full canopy. O₃ did not have an overall detrimental effect on crop development during initial growth nor at full canopy, but did induce a faster reduction of LAI during crop senescence (DAE×O₃, P<0.05). Final plant height was not affected by O₃. There were few CO₂×O₃ interactions detected. There was a suggestion (P=0.06) that O₃ counteracted the CO₂-induced decrease of green leaves at full canopy, but on the other hand during crop senescence the decline of LAI due to elevated O₃ was faster at ambient compared to elevated CO₂ (P<0.05). These responses of canopy development to elevated CO₂ and O₃ help to explain the treatment responses of potato yield within the CHIP project at sites across Europe.     

Photosynthetic and stomatal responses of potatoes grown under elevated CO2 and/or O3—results from the European CHIP-programme

The physiological effects of elevated CO₂ and/or O₃ on Solanum tuberosum cv. Bintje were examined in Open-Top Chambers during 1998 and 1999 at experimental sites across Europe as part of the EU ‘Changing Climate and Potential Impacts on Potato Yield and Quality’ programme (CHIP). At tuber initiation (≈20 days after emergence, DAE) elevated CO₂ (680 μl l⁻¹) induced a 40% increase in the light saturated photosynthetic rate (A_sat) of fully expanded leaves in the upper canopy. This was 16% less than expected from short-term exposures of plants grown under ambient CO₂ (360 μl l⁻¹) to elevated CO₂, indicating that photosynthetic acclimation began at an early stage of crop growth. This effect resulted from a combination of a 12% reduction in stomatal conductance (g_s) and a decline in photosynthetic capacity, as indicated by the significant reductions in the maximum carboxylation rate of Rubisco (Vc_max) and light-saturated rate of electron transport (J_max) under elevated CO₂. The seasonal decline in the promotion of photosynthesis by elevated CO₂ reflected the concurrent decrease in g_s. Vc_max and J_max were both reduced in plants grown under elevated CO₂ until shortly after maximum leaf area (MLA) was attained. Although non-photorespiratory mitochondrial respiration in the light (R_d) increased during the later stages of the season, net photosynthesis was consistently increased by elevated CO₂ during the main part of the season. Photosynthetic rate declined more rapidly in response to elevated O₃ under ambient CO₂, and the detrimental impact of O₃ was most obvious after MLA was attained (DAE 40–50). Several exposure indices were compared, with the objective of determining the critical ozone level required to induce physiological effects. The critical O₃ exposure above which a 5% reduction in light saturated photosynthetic rate may be expected (expressed in terms of cumulative exposure above 0 nl l⁻¹ O₃ between emergence and specific dates during the season (AOT0-cum)) was 11 μl l⁻¹ h; however this value should only be extrapolated beyond the CHIP dataset with caution. The interaction between O₃ and stomatal behaviour was more complex, as it was influenced by both long-term and daily exposure levels. Elevated CO₂ counteracted the adverse effect of O₃ on photosynthesis, perhaps because the observed reduction in stomatal conductance decreased O₃ fluxes into the leaves. The results are discussed in the context of nitrogen deficiency, carbohydrate accumulation and yield.     

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.     

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.     

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.     

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.     

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.     

Chlorophyll concentration of potatoes grown under elevated carbon dioxide and/or ozone concentrations

Potato cv. Bintje was grown in open-top-chambers and free-air-CO₂-enrichment systems at 7 sites across Europe for 2 years (1998–99). The effect of different treatments (CO₂ enrichment and O₃ fumigation) on the chlorophyll content of fully expanded upper and lower canopy leaves was investigated collecting Minolta SPAD-502 meter readings. In both CO₂ treated and O₃ fumigated plants, leaves had lower chlorophyll content than those in ambient air controls; season-long chlorophyll averages were 9.3% lower in the ‘CO₂’ treatments, 9.1% lower in ‘O₃’ treatments and 12.3% lower in ‘CO₂+O₃’ treatments. The analysis of chlorophyll content in three different growth phases (Emergence–Tuber Initiation; Tuber Initiation–Maximum Leaf Area; Maximum Leaf Area–Harvest) showed that in the early growth period, i.e. before tuber initiation there was a slight indication for an higher chlorophyll content at elevated CO₂ (+3.8%) or O₃ (+1.7%). However, from tuber initiation onwards the leaves of plants grown under elevated CO₂ or O₃ showed a progressively lower chlorophyll content (−4.8% for CO₂ treatments and −2.6% for O₃ treatments) indicating a faster senescence of leaves that increased during the late growth period (−12.8% for CO₂ treatments and −12.7% for O₃ treatments) and that was enhanced by CO₂–O₃ interaction (−17.8%).     

Stomatal conductance and ozone exposure in relation to potato tuber yield—results from the European CHIP programme

Measurements of stomatal conductance on field-grown potato (Solanum tuberosum L.) cv. Bintje from the CHIP programme were combined to study the response to environmental factors. 3274 data points were used. Data were obtained from five sites: Jokioinen in Finland, Östad in Sweden, Giessen in Germany, Tervuren in Belgium and Sutton Bonnington in UK. Measurements were made in open-top chamber treatments with ozone and carbon dioxide exposure and in the ambient air. A typical light response curve was obtained with light saturation at approximately 400 μmol m⁻² s⁻¹ photosynthetically active radiation (PAR). The leaf temperature optimum for stomatal conductance was 29 °C. The stomatal conductance declined strongly at leaf-to-air vapour pressure differences >20 hPa. An elevated carbon dioxide concentration (680 μl l⁻¹) reduced the stomatal conductance by up to approximately 20%. Elevated ozone reduced the stomatal conductance towards the end of the growth period, in addition to the negative effect by ordinary senescence on stomatal conductance. A multiplicative model, based on the boundary line technique, was used to estimate the relationship between stomatal conductance and the environmental variables. Test with the data sets from two sites providing sufficient data, Östad and Giessen, showed that the multiplicative model had R²-values of 0.60 and 0.42, respectively, for the relationship between calculated and observed conductance. Test of the model with an independent data set from an open-top chamber experiment with the potato cultivar Kardal showed an R² of 0.59 between calculated and observed conductance. The conductance model was used to estimate the accumulated ozone uptake (CUO₃) by potato leaves from emergence to harvest. The relationship between CUO₃ and relative yield loss, using a threshold for the ozone uptake rate of 7 nmol m⁻² s⁻¹, provided a higher R²-value (0.45) than CUO₃ without any threshold and relationships based on the accumulated exposure over 40 nmol mol⁻¹ (AOT40) or the sum of all hourly average ozone concentrations exceeding 60 nmol mol⁻¹ (SUM06). All four relationships were however statistically significant.     

Growth and yield responses of spring wheat to increasing carbon dioxide, ozone and physiological stresses: a statistical analysis of ‘ESPACE-wheat’ results

One of the major goals of the European Stress Physiology and Climate Experiment (ESPACE-wheat) was to investigate the sensitivity of wheat growth and productivity to the combined effects of changes in CO₂ concentration, ozone and other physiological stresses. Experiments were performed at different sites throughout Europe, over three consecutive growing-seasons using open-top chambers. This paper summarizes the main experimental findings of the effects of CO₂ enrichment and other factors i.e. ozone (O₃), drought stress or nitrogen supply on the biomass and yield of spring wheat (Triticum aestivum cv. Minaret). Final harvest data from different sites and seasons were statistically analysed: (1) to identify main effects and interactions between experimentally controlled factors; and (2) to evaluate quantitative relationships between environmental variables and biological responses. Generally, ‘Minaret’ wheat did not respond significantly to O₃, suggesting that this cultivar is relatively tolerant to the O₃ levels applied. The main effect of CO₂ was a significant enhancement of grain yield and above-ground biomass in almost all experiments. Significant interactions between CO₂ and other factors were not common, although modifications in different N- and water supplies also led to significant effects on grain yield and biomass. In addition, climatic factors (in particular: mean air temperature and global radiation) were identified as important co-variables affecting grain yield or biomass, repectively. On average, the yield increase as a result of a doubling of [CO₂] was 35% compared with that observed at ambient CO₂ concentrations. However, linear regressions of grain yield or above-ground biomass for individual experiments revealed a large variability in the quantitative responses of ‘Minaret’ wheat to CO₂ enrichment (yield increase ranging from 11 to 121%). Hence, CO₂ responsiveness was shown to differ considerably when the same cultivar of wheat was grown at different European locations. Multiple regression analyses perfomed to evaluate the relative importance of the measured environmental parameters on grain yield indicated that although yield was significantly related to five independent variables (24 h mean CO₂ concentration, 12 h mean O₃ concentration, temperature, radiation, and drought stress), a large proportion of the observed variability remained unexplained.     

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.     

Factors influencing visible ozone injury on potato including the interaction with carbon dioxide

Weekly observations and scoring of ozone injury were done on all experimental sites where continuous monitoring of ozone in ambient air was available. Typical acute symptoms were reddish-brown lesions and bronzing, primarily on the upper surface of the leaves. The potato cultivar Bintje, cultivated at all sites, appeared to be sensitive to ozone but less than clover species, which served as model for the short-term critical level (AOT40/5 d). At the mid-European and Nordic latitudes injury appeared after an AOT40/5 d exposure of about 700 and 275 nl l⁻¹ h, respectively. In central Europe the plants were indeed less sensitive than in Scandinavia and the AOT40/5 d values were higher than the proposed critical level of 500 nl l⁻¹ h at a vapour pressure deficit (VPD) exceeding 1.5 kPa and 200 nl l⁻¹ h at a VPD below 1.5 kPa. In addition injury appeared primarily after the stage of maximum leaf area (MLA). In the Nordic countries injury was noticed at very low AOT40/5 d values and occurred before as well as after the stage of MLA. Using artificial neural network models it appeared from the data set that VPD did not play a major role in the ozone sensitivity of potato, but there were indications that daylength was an important parameter in addition to a possible higher uptake at high latitudes. Increasing the CO₂ concentration did not prevent the potato plants from visible ozone damage but it reduced the intensity by 10%. Taking only the OTC experiments of the mid-latitudes into account, there was a correlation between the ozone injury scoring just before harvest and the marketable yield loss. However, in all the cases considered, ozone damage appeared only after the stage of MLA and increased steadily towards harvest. The correlation can be explained by a parallel development of injury and chlorosis (senescence) typical for chronic exposure. There was no relationship between isolated short-term exposures causing acute injury and final yield.     

臭氧与其他环境因子及其复合作用对植物的影响

综述了臭氧（O₃）浓度升高、太阳辐射减弱、UV-B辐射、CO₂浓度升高及其与O₃复合作用对植物形态特征、光合作用、干物质累积及作物产量等生理生化机制的影响。交互作用的试验条件可以更好地模拟自然环境条件。O₃和UV-B辐射对植物几乎没有积极作用。太阳辐射减弱、CO₂浓度升高都会促进植物营养生长。但太阳辐射减弱降低干物质累积和产量,CO₂浓度升高对其有促进作用。CO₂浓度升高在与O₃复合条件下,可部分缓解太阳辐射减弱对植物造成的伤害。而UV-B辐射与O₃复合对植物造成的伤害更大。     

Effect of fruit maturity on efficiency of 1-methylcyclopropene to delay the ripening of bananas

Three bunches of unripe ‘Williams’ banana fruit of different maturity, 173, 156 and 71 days from bunch emergence, were harvested. Fruit from the top, bottom and middle hands from each bunch were fumigated for 24 h with 1-methylcyclopropene (1-MCP) at 0, 5, 50 or 500 nl l⁻¹ at 20^oC. All fruit were then stored at 20^oC in air containing 0.1 μl l⁻¹ ethylene and the time taken for each fruit to ripen (green life) was noted. The green life of fruit treated with 500 nl l⁻¹ 1-MCP varied with fruit maturity. In the two most mature bunches it was 27.9±2.3 days, 4-fold longer than fruit fumigated with 0 nl l⁻¹ 1-MCP (6.7±0.6 days). In the least mature bunch, green life was 39.7±3.0 days, 1.5-fold longer than fruit fumigated with 0 nl l⁻¹ 1-MCP (25.7±2.5 days). Most fruit treated with 500 nl l⁻¹ 1-MCP showed an unacceptable uneven skin colouration when ripe. There was no significant effect on green life of 1-MCP at 50 nl l⁻¹ and 5 nl l⁻¹. Other fruit from these bunches were not exposed to 1-MCP and were held in ethylene-free air until ripe. In the two most mature bunches, these fruit had a significantly shorter green life (11.2±5.6 days in hand 1; 18.9±4.1 days in hands 4 and 6) than fruit that were fumigated with 500 nl l⁻¹ 1-MCP. In the least mature bunch, however, these fruit had a significantly longer green life (56.0±5.9 days) than 1-MCP treated fruit. Since the effectiveness of 1-MCP varied with fruit maturity and in any commercial consignment there is a mixture of fruit maturity, it is concluded that 1-MCP has limited commercial potential for the storage of unripe ‘Williams’ bananas.     

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 CO2 and/or ozone on nutrient concentrations and nutrient uptake of potatoes

Potato crops were grown at seven sites across Europe to test the effects of elevated atmospheric carbon dioxide and/or tropospheric ozone concentrations on growth, yield and various aspects of potato tuber quality within the framework of the EC funded programme Changing Climate and Potential Impacts on Potato Yield and Quality (CHIP). Field exposure systems were used to enrich the atmosphere in CO₂ and/or ozone. At five of the sites, nutrient element conconcentrations (macronutrients: nitrogen, phosphorus, potassium, calcium, magnesium, and micronutrients: mangenese, zinc, iron) in different parts of plants from the various treatments were analysed. Under elevated CO₂, nearly all nutrient elements tended to decrease in concentration. At maximum leaf area, a significant reduction was observed for the concentrations of nitrogen and potassium both in aboveground biomass and in tubers, and for calcium in tubers. Since CO₂ enrichment promoted early tuber growth, these effects could in part be attributed to tuber developmental stage. At maturity, potato grown under CO₂ enrichment exhibited significantly lower concentrations of nitrogen, manganese and iron in aboveground organs, and of nitrogen, potassium and magnesium in tubers which means a reduction of tuber quality. In contrast to CO₂, elevated ozone tended to increase tuber nutrient element concentrations. This was significant for nitrogen and manganese. CO₂ effects on tuber biomass increase were more pronounced than CO₂ effects on nutrient element decrease. Thus, the total amount of nutrient elements taken up by potato crops increased under elevated CO₂. Fertiliser practice in a future, CO₂-rich world will have to be adjusted accordingly.     

Interactive effects of elevated atmospheric CO2 concentrations and plant available soil water content on canopy evapotranspiration and conductance of spring wheat

The present study was conducted to investigate the possible interactive effects of rising atmospheric CO₂ concentration [CO₂] and drought stress on water use of wheat. Spring wheat (Triticum aestivum cv. “Minaret”) was grown either in 1 m diameter lysimeters with 0.4 m soil depth (1998) or in the field (1999) in open-top chambers under two CO₂-concentrations (ambient, ambient + 280 ppm) and two watering regimes (well-watered = WW with a plant available water content PAW > 40 mm and drought stressed = DS, 10 mm < PAW < 30 mm) beginning after first node stage. Canopy evapotranspiration (E_C) was measured continuously from first the node stage until the beginning of flag leaf senescence using four open-system canopy chambers (0.78 m³). Seasonal changes of the absorption of photosynthetically active radiation (APAR) of the canopy and root growth (1999) were also measured.

In both growing seasons leaf area index increased in response to elevated [CO₂] in both water treatments. The related effects of [CO₂] on canopy radiation absorption (APAR) were, however, smaller. E_C was linearily related to APAR in both growing seasons. While elevated [CO₂] reduced the slope of this relation under WW conditions by ca. 20% in both growing seasons, it was not reduced (1998) and even increased (1999) under drought. Canopy conductance (G_C) calculated as E_C divided by vapour pressure deficit of air, showed a non-linear relationship to APAR that was best explained by saturation curves. Under WW conditions, elevated [CO₂] reduced the initial slope of G_C versus APAR as well as G_C at saturating light conditions (ca. −30%), while under DS conditions no effect of elevated [CO₂] could be detected. Under high light conditions (PAR > 400 μmol m⁻² s⁻¹) a critical “threshold value” of PAW (T_PAW, ca. 40 mm) could be identified above which G_C did not respond to PAW. While in 1998 G_C did not respond to elevated [CO₂] at PAW < T_PAW, it was slightly increased at low PAW values in the field experiments of 1999. The reduction of T_PAW by elevated [CO₂] may be explained by enhanced root growth (1999) that would have given the plants better access to soil water resources. The present results suggest that below a critical soil water content elevated [CO₂] will not reduce canopy water loss of wheat or may even enhance it.     

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.     

CO2浓度升高条件下不同氮素水平对小叶章光合特性和生长的影响

为阐明大气CO₂浓度升高和不同氮素水平对湿地植物光合生理特性和生长的影响,本研究以三江平原湿地优势植物小叶章(Calamagrostis angustifolia)为研究对象,通过野外原位控制试验,利用开顶式气室(OTC)模拟环境大气CO₂浓度变化,设置E₀(380 ±20 µmol/mol)、E₁(550 ±20 μmol/mol)和E₂(700 ± 20 μmol/mol)3个CO₂浓度;在每个OTC内设置 N₀(0 g N/m²)、N₁(4 g N/m²)和N₂(8 g N/m²)3个氮素水平。结果表明,N₀条件下,与E₀处理相比,E₁和E₂处理（72 天）后小叶章叶片净光合速率分别降低11%和12%(P<0.05),其叶片可溶性蛋白含量、氮素含量（CO₂熏蒸72 天）、小叶章株高（CO₂熏蒸86 天）均显著低于E₀处理(P<0.05);N₁条件下,与E₀处理相比,E₁和E₂处理（72 天）后小叶章叶片净光合速率降低5%(P>0.05)和10%(P<0.05),其叶片氮素含量(P<0.05)、小叶章株高均低于E₀处理;N₂条件下,E₁和E₂处理（72 天）小叶章净光合速率均呈稍增加的趋势(P>0.05),其叶片可溶性蛋白含量显著增加(P<0.05),氮素含量和小叶章株高无显著变化(P>0.05)。N₀、N₁和N₂条件下,CO₂浓度升高均显著增加了小叶章叶片可溶性糖含量。本研究表明长期CO₂浓度升高可能通过降低小叶章叶片光合酶活性,进而降低了其净光合速率,而施加高浓度的氮肥可以缓解长期高CO₂浓度对湿地植物光合及生长的负面影响。