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.     

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.     

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%).     

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.     

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.     

Spatial and temporal distribution of the root system and root nutrient content of an established Miscanthus crop

Although a high biomass yield is obtained from established Miscanthus crops, previous studies have shown that fertilizer requirements are relatively low. As little information on the role of the Miscanthus roots in nutrient acquisition is available, a study was conducted to gather data on the Miscanthus root system and root nutrient content. Therefore in 1992, the root distribution pattern of an established Miscanthus crop was measured in field trials using the trench profile and the auger methods. Also, in 1994/1995, seasonal changes in root length density (RLD) and root nutrient content were monitored three times during the vegetation period.

The trench profile method showed that roots were present to the maximum depth measured of 250 cm. The top soil (0–30 cm) contained 28% of root biomass, while nearly half of the total roots were present in soil layers deeper than 90 cm. Using the auger method, we found that RLD values in the topsoil decreased with increasing distance from the centre of the plants. Below 30 cm, RLD decreased markedly, and differences in root length in the soil between plants were less pronounced. The total root dry weight down to 180 cm tended to increase from May 1994 (10.6 t ha⁻¹) to November 1994 (13.9 t ha⁻¹) and then decreased again until March 1995 (11.5 t ha⁻¹). Nutrient concentrations in the roots decreased with increasing depth. The concentrations of N (0.7–1.4%) and K (0.6–1.2%) were clearly higher than those of P (0.06–0.17%). The mean values for N, P and K contents of the roots of all three sampling dates in 1994/1995 were 109.2 kg N ha⁻¹, 10.6 kg P ha⁻¹ and 92.5 kg K ha⁻¹.

Although our results showed that RLD values for Miscanthus in the topsoil are lower than for annual crops, the greater rooting depth and the higher RLD of Miscanthus in the subsoil mean that nutrient uptake from the subsoil is potentially greater. This enables Miscanthus crops to overcome periods of low nutrient (and water) availability especially during periods of rapid above-ground biomass growth.     

Effects of elevated CO2 and cutting frequency on the productivity and herbage quality of a semi-natural grassland

Monoliths of a fertile, although N limited, C₃ grassland community were subjected (or not) to an atmospheric CO₂ enrichment (600 μmol mol⁻¹), owing to the Mini-FACE system from August 1998 to June 2001, at two contrasting cutting frequencies (3 and 6 cuts per year). The present study reports the effects of elevated CO₂ on the above-ground productivity and on the herbage quality. Elevated CO₂ did not affect the dry matter (DM) yield of the swards in 1999. In 2000, the second year, there was a positive CO₂ effect (+26%) both on the DM and on the nitrogen yields (+30%). With the frequently cut monoliths, the DM of the legume component of the sward was strongly increased by elevated CO₂. This effect became also significant in July 2000 for the low cutting frequency treatment. These results are in good agreement with the concept of an increased legume development and symbiotic N₂ fixation triggered by an increased ecosystem scale demand of N under elevated CO₂. At a low cutting frequency, the DM of the forbs was strongly increased in elevated compared with ambient CO₂. This increased development of the forbs apparently led to a competitive decline of the grasses. Therefore, the total DM yield response to CO₂ was smaller at a low (+15%) compared with a high (+36%) cutting frequency in 2000. An increase in the water soluble sugar content of the bulk forage under elevated CO₂ and a corresponding decline in cell wall contents (NDF) were observed. In June 1999, the decline in NDF was correlated with an increased in-vitro DM digestibility. The forage quality was also indirectly affected by elevated CO₂ through changes in leaf:stem ratio and in botanical composition. At a low cutting frequency, the increased forb content favoured the herbage quality because of a higher digestibility of the forb shoots and, indirectly, through the reduction in the mass of the grass stems. These results emphasise the role of species dynamics for elevated CO₂ impacts on semi-natural grassland productivity and herbage quality.     

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.     

Yield and water-use efficiency of water- and nitrogen-stressed wheat crops increase with degree of co-limitation

Availability of water and nitrogen are key constraints to primary productivity in arid and semiarid ecosystems. Theoretically, plant growth is maximised when all resources are equally limiting. This paper tested the hypothesis that for a given amount of available water, the gap between actual and attainable yield of dryland crops in semiarid southern Australia is inversely proportional to the degree of nitrogen and water co-limitation.

Field and simulation experiments were combined in an analysis involving three steps. Step 1 assessed the capacity of a crop simulation model to estimate yield and its responses to water and nitrogen inputs in the semiarid Mallee region. Step 2 derived a boundary function relating grain yield and water availability using simulations with long-term weather records. Step 3 explored the link between degree of co-limitation and deviations between actual yield and the boundary function. Degree of co-limitation (C_WN) was calculated as a function of model-derived nitrogen (NSI) and water stress indices (WSI), i.e. C_WN = 1 − |NSI − WSI|. Stress indices range from zero (no stress) to 1 (maximum stress), and C_WN tends to 1 when both resources impose constraints of similar magnitude to crop growth.

The field experiment combining locations, seasons and management practices generated a range of grain yield from 0.6 to 3.8 t ha⁻¹. Water availability, i.e. seasonal rainfall plus change in soil water content from sowing to harvest, ranged from 127 to 370 mm. Nitrogen fertiliser varied from nil to 36 kg N ha⁻¹ and inorganic nitrogen in the soil profile at sowing ranged from 29 to 497 kg ha⁻¹. For these ranges of conditions, the relationship between simulated and measured yield was statistically undistinguishable from the y = x function.

A factorial modelling experiment combining sites, seasons, initial soil water content and dose of nitrogen fertiliser was used to derive a boundary function which provided an objective and independent upper limit for the field data. Actual yield was below the boundary function in most cases. The difference between actual and attainable yield was inversely proportional to C_WN. This study thus supported the hypothesis that yield and water-use efficiency of water- and nitrogen-stressed crops increase with increasing degree of co-limitation.     

Monitoring nitrogen forms in soil/plant systems under different fertilizer managements. A preliminary investigation

A field experiment was carried out on maize (Zea mays, L.) to study the effects of different fertilizer management on nitrogen status in soil and plant response. Three different fertilizers, mineral (MN), mineral plus buffalo manure (MN + BM) and organo-mineral with peat (OMP), were added at the usual (140, 61 and 116 kg ha⁻¹) and the reduced (70, 31 and 58 kg ha⁻¹) rates of N, P and K. respectively. Soil samples were analyzed for N by both the Kjeldahl method and the electro-ultrafiltration technique (EUF). The soil Kjeldahl-N concentrations were scarcely affected by the different fertilizer treatments, while the EUF-N concentrations were closely correlated with the amounts of N added. The EUF also discriminated between the NO₃-N and the sum of the ammonium and the easily extractable organic N forms (EUF-N_org + NH₄). The largest proportions of EUF-Norg + NH₄ were found in the untreated plots and in the plots treated with buffalo manure. The different fertilizer treatments significantly affected grain yield, which ranged from a minimum of 6.3 t ha⁻¹ from the untreated plots, to a maximum of 11.9 t ha⁻¹ from those supplied with 140 kg N, 61 kg P and 116 kg K ha⁻¹ by OMP fertilizer. The highest agronomic efficiency index for N was exhibited in the OMP treatment at the reduced rate. The grain yield was closely correlated with the total extractable EUF-N, but different relationships were found between the rate of N added, the level of EUF-NO,-N in soil and grain yield for the different fertilizer treatments.     

Effect of preceding crop combination and N fertilization on yield of six oil-seed rape cultivars (Brassica napus L.)

Information about the effect of the cropping history on the seed yield of oil-seed rape is extremely scarce. In 1992/93 and 1994/95, the effects of different preceding crop combinations (winter barley and winter wheat as preceding crops, oil-seed rape and wheat as pre-preceding crops) on the yield of six double low oil-seed rape cultivars were examined in a field trial at Hohenschulen Experimental Farm, north-west Germany. In addition, eight nitrogen treatments (different amounts and distribution patterns) were tested for their potential to reduce negative effects of the preceding crops. Following the cropping sequence of oil-seed rape then wheat, oil-seed rape yielded only 3.12 t ha⁻¹; after oil-seed rape then barley, the yield was 3.43 t ha⁻¹ compared with 3.77 t ha⁻¹ following wheat then barley and 3.71 t ha⁻¹ following wheat then wheat. The number of seeds per m² showed a similar pattern, whereas the thousand-seed weight partly compensated for the reduced seed number. It was highest if oil-seed rape was grown 2 years previously. The cultivars differed significantly in their yield potential. Express (3.79 t ha⁻¹) yielded 0.6 t ha⁻¹ more than Falcon (3.18 t ha⁻¹). Increasing amounts of fertilizer-N (80–200 kg N ha⁻¹) increased the seed yield from 3.21 t ha⁻¹ to 3.84 t ha⁻¹. Changes in the distribution pattern within one fertilizer amount had no effect on seed yield. In addition, no interactions between preceding crop combination and the different cultivars or N treatments occurred. It is concluded that crop management cannot totally eliminate the negative effects of an unfavourable cropping history on the seed yield of oil-seed rape.     

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.     

Delaying establishment of controlled atmosphere or CO2 exposure reduces ‘Fuji’ apple CO2 injury without excessive fruit quality loss

Storage of ‘Fuji’ apple fruit in a high CO₂ (3 kPa) and low O₂ (1.5 kPa) controlled atmosphere (CA) reduced firmness and titratable acidity (TA) loss during long term storage. This CA environment also induced development of internal CO₂-injury (brown-heart) and slowed the disappearance of watercore. The symptoms of internal CO₂-injury were first detected 15 days after CA establishment and the severity increased during the first 4 months of CA-storage. Delaying establishment of CA conditions for 2–12 weeks significantly reduced the severity of CO₂-injury. Delaying CO₂ accumulation to 3 kPa for 1–4 months during CA (1.5 kPa O₂+0.05 kPa CO₂) storage also reduced development of CO₂-injury symptoms. Delaying CA or CO₂ accumulation resulted in lower firmness and TA compared to establishment of CA within 72 h of harvest. However, the delay treatments did result in firmness and TA that were significantly higher compared to values for fruit stored in air. The incidence and severity of senescent injuries (flesh browning and core flush) detected during the late period of storage were greater in air- than CA-stored fruit. The results indicate the susceptibility of ‘Fuji’ apples to CO₂-injury is highest during the first weeks of storage after harvest. Delaying establishment of CA or exposure to elevated CO₂ after harvest may be a practical strategy to reduce CO₂-injury while maintaining other important quality attributes at acceptable levels.     

Biomass yield and energy balance of giant reed (Arundo donax L.) cropped in central Italy as related to different management practices

In order to evaluate the possibility of reducing energy input in giant reed (Arundo donax L.) as a perennial biomass crop, a field experiment was carried out from 1996 to 2001 in central Italy. Crop yield response to fertilisation (200–80–200 kg ha⁻¹ N–P–K), harvest time (autumn and winter) and plant density (20,000 and 40,000 plants per ha) was evaluated. The energy balance was assessed considering the energy costs of production inputs and the energy output obtained by the transformation of the final product. The crop yield increased by +50% from the establishment period to the 2nd year of growth when it achieved the highest dry matter yield. The mature crop displayed on average annual production rates of 3 kg dry matter m⁻², with maximum values obtained in fertilised plot and during winter harvest time.

Fertilisation mainly enhanced dry matter yield in the initial period (+0.7 kg dry matter m⁻² as years 1–6 mean value). The biomass water content was affected by harvest time, decreasing by about 10% from autumn to winter. With regard to plant density, higher dry matter yields were achieved with 20,000 plants per ha (+0.3 kg dry matter m⁻² as years 1–6 mean value).

The total energy input decreased from fertilised (18 GJ ha⁻¹) to not fertilised crops (4 GJ ha⁻¹). The higher energetic input was represented by fertilisation which involved 14 GJ ha⁻¹ (fertilisers plus their distribution) of total energy costs. This value represents 78% of total energy inputs for fertilised crops.

Giant reed biomass calorific mean value (i.e., the calorific value obtained from combustion of biomass sample in an adiabatic system) was about 17 MJ kg⁻¹ dry matter and it was not affected by fertilisation, or by plant density or harvest time. Fertilisation enhanced crop biomass yield from 23 to 27 dry tonnes per ha (years 1–6 mean value). This 15% increase was possible with an energy consumption of 70% of the overall energy cost. Maximum energy yield output was 496 GJ ha⁻¹, obtained with 20,000 plants per ha and fertilisation. From the establishment period to 2nd–6th year of growth the energy production efficiency (as ratio between energy output and energy input per ha) and the net energy yield (as difference between energy output and energy input per ha) increased due to the low crop dry biomass yield and the high energy costs for crop planting. The energy production efficiency and net energy yield were also affected by fertilisation and plant density. In the mature crop the energy efficiency was highest without fertilisation both with 20,000 (131 GJ ha⁻¹) and 40,000 plants per ha (119 GJ ha⁻¹).     

Tillage, cover crops, and nitrogen fertilization effects on soil nitrogen and cotton and sorghum yields

Sustainable soil and crop management practices that reduce soil erosion and nitrogen (N) leaching, conserve soil organic matter, and optimize cotton and sorghum yields still remain a challenge. We examined the influence of three tillage practices (no-till, strip till and chisel till), four cover crops {legume [hairy vetch (Vicia villosa Roth)], nonlegume [rye (Secaele cereale L.)], vetch/rye biculture and winter weeds or no cover crop}, and three N fertilization rates (0, 60–65 and 120–130 kg N ha⁻¹) on soil inorganic N content at the 0–30 cm depth and yields and N uptake of cotton (Gossypium hirsutum L.) and sorghum [Sorghum bicolor (L.) Moench]. A field experiment was conducted on Dothan sandy loam (fine-loamy, siliceous, thermic, Plinthic Paleudults) from 1999 to 2002 in Georgia, USA. Nitrogen supplied by cover crops was greater with vetch and vetch/rye biculture than with rye and weeds. Soil inorganic N at the 0–10 and 10–30 cm depths increased with increasing N rate and were greater with vetch than with rye and weeds in April 2000 and 2002. Inorganic N at 0–10 cm was also greater with vetch than with rye in no-till, greater with vetch/rye than with rye and weeds in strip till, and greater with vetch than with rye and weeds in chisel till. In 2000, cotton lint yield and N uptake were greater in no-till with rye or 60 kg N ha⁻¹ than in other treatments, but biomass (stems + leaves) yield and N uptake were greater with vetch and vetch/rye than with rye or weeds, and greater with 60 and 120 than with 0 kg N ha⁻¹. In 2001, sorghum grain yield, biomass yield, and N uptake were greater in strip till and chisel till than in no-till, and greater in vetch and vetch/rye with or without N than in rye and weeds with 0 or 65 kg N ha⁻¹. In 2002, cotton lint yield and N uptake were greater in chisel till, rye and weeds with 0 or 60 kg N ha⁻¹ than in other treatments, but biomass N uptake was greater in vetch/rye with 60 kg N ha⁻¹ than in rye and weeds with 0 or 60 kg N ha⁻¹. Increased N supplied by hairy vetch or 120–130 kg N ha⁻¹ increased soil N availability, sorghum grain yield, cotton and sorghum biomass yields, and N uptake but decreased cotton lint yield and lint N uptake compared with rye, weeds or 0 kg N ha⁻¹. Cotton and sorghum yields and N uptake can be optimized and potentials for soil erosion and N leaching can be reduced by using conservation tillage, such as no-till or strip till, with vetch/rye biculture cover crop and 60–65 kg N ha⁻¹. The results can be applied in regions where cover crops can be grown in the winter to reduce soil erosion and N leaching and where tillage intensity and N fertilization rates can be minimized to reduce the costs of energy requirement for tillage and N fertilization while optimizing crop production.     

Pod photosynthesis and drought adaptation of field grown rape (Brassica napus L.)

In rape (Brassica napus L., cv. Global) seed growth mainly depends on husk CO₂ assimilation. In irrigated plants, the net photosynthetic rate (A_max) was 10–13 μmol CO₂ m⁻² s⁻¹ in non-maturing pods and correlated with nitrogen content. The stomatal conductance of water vapour (g_H2O) was 0.3 mol m⁻² s⁻¹ in non-maturing pods. The photosynthetic nitrogen use efficiency (NUE) was 8.3 μmol CO₂g⁻¹ N s⁻¹, about one-third of that in leaves. The photosynthetic water use efficiency (WUE; A_maxg_H2O⁻¹) was similar in pods and leaves. In severely droughted plants, the photosynthetic rate was reduced to 38%. The seed growth rate, however, was not influenced by intermittent periods of water stress, indicating translocation of assimilates to the seeds. The drought resistant character of the pods was due to low specific area, succulence, low stomatal conductance causing a small decrease of ΔΨ day⁻¹ during soil drying and maintenance of high relative water content during severe drought. A mathematical formulation of the pod water release curve was undertaken. © (1997) Elsevier Science B.V.     

N2 fixation and N supply in organic pea (Pisum sativum L.) cropping systems as affected by weeds and peaweevil (Sitona lineatus L.)

Grain legumes, especially peas, could play a key role in organic cropping systems. They could provide nitrogen (N) to the system via N₂ fixation and produce grain rich in protein while improving soil N for the succeeding crop. Thus, maximising N₂ fixation and optimising grain N production together with N contribution to soil is a challenging issue for organic pea crops. However, pest, disease and weed infestation are less easy to control in organic systems than in conventional systems. Therefore, the effects of weed infestation and pea weevil (Sitona lineatus L.) attacks on N nutrition and N₂ fixation of organic pea crops were examined by on-farm monitoring over two years. The magnitude of the net contribution of the crops to the soil N balance in relation to their productivity was also assessed. In many situations, weed infestation together with pea weevil damage severely limited the nitrogen nutrition and grain yield. Percentage of N derived from fixation (%Ndfa) increased with weed biomass because weeds appeared more competitive than peas for soil N. But %Ndfa decreased with pea weevil leaf damage score. The interaction between these two biotic factors affected N yields and the net contribution of the crops to soil N. This latter ranged from −133 kg N ha⁻¹ to 69 kg N ha⁻¹ depending on %Ndfa and nitrogen harvest index (NHI). Optimising both grain N and net balance would require a reduction in root nodule damage by weevil larvae in order to maximise %Ndfa and a reduction in the NHI through the choice of cultivar and/or suitable crop management.