ALBERT

Notes: Spring wheat [ Triticum aestivum (L). cv. Yecora Rojo] was grown from December 1992 to May 1993 under two atmospheric CO2 concentrations, 550 μmol mol–1 for high-CO2 plots, and 370 μmol mol–1 for control plots, using a Free-Air CO2 Enrichment (FACE) apparatus. In addition to the two levels of atmospheric CO2, there were ample and limiting levels of water supply through a subsurface trip irrigation system in a strip, split-plot design. In order to examine the temporal and spatial root distribution, root cores were extracted at six growth stages during the season at in-row and inter-row positions using a soil core device (86 mm ID, 1.0 m length). Such information would help determine whether and to what extent root morphology is changed by alteration of two important factors, atmospheric CO2 and soil water, in this agricultural ecosystem.Wheat root growth increased under elevated CO2 conditions during all observed developmental stages. A maximum of 37% increase in total root dry mass in the FACE vs. Control plots was observed during the period of stem elongation. Greater root growth rates were calculated due to CO2 enhancement until anthesis. During the early vegetative growth, root dry mass of the inter-row space was significantly higher for FACE than for Control treatments suggesting that elevated CO2 promoted the production of first-order lateral roots per main axis. Then, during the reproductive period of growth, more branching of lateral roots in the FACE treatment occurred due to water stress. Significant higher root dry mass was measured in the inter-row space of the FACE plots where soil water supply was limiting. These sequential responses in root growth and morphology to elevated CO2 and reduced soil water supports the hypothesis that plants grown in a high-CO2 environment may better compensate soil-water-stress conditions.

Notes: A spring wheat crop was grown at ambient and elevated (550 μmol mol−1) CO2 concentrations under free-air CO2 enrichment (FACE) in the field. Four experimental blocks, each comprising 21-m-diameter FACE and control experimental areas, were used. CO2 elevation was maintained day and night from crop emergence to final grain harvest. This experiment provided a unique opportunity to examine the hypothesis that CO2 elevation in the field would lead to acclimatory changes within the photosynthetic apparatus under open field conditions and lo assess whether acclimation was affected by crop developmental stage, leaf ontogeny and leaf age. Change in the photosynthetic apparatus was assessed by measuring changes in the composition of total leaf and thylakoid polypeptides separated by SDS-PAGE. For leaves at completion of emergence of the blade, growth at the elevated CO2 concentration had no apparent effect on the amount of any of the major proteins of the photosynthetic apparatus regardless of the leaf examined. Leaf 5 on the main stem was in full sunlight at emergence, but then became shaded progressively as 3–4 further leaves formed above with continued development of the crop. By 35 d following completion of blade emergence, leaf 5 was in shade. At this point, the chlorophyll alb ratio had declined by 26% both in plants grown at the control CO2 concentration and in those grown at the elevated CO2 concentration, which is indicative of shade acclimation. The ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) content declined by 45% in the control leaves, but by 60% in the leaves grown at the elevated CO2 concentration. The light- harvesting complex of photosystcm II (LHCII) and the chlorophyll content showed no decrease and no difference between treatments, indicating that the decrease in Rubisco was not an effect of earlier senescence in the leaves at the elevated CO2 concentration. Following completion of the emergence of the flag-leaf blade, the elevated-CO2 treatment inhibited the further accumulation of Rubisco which was apparent in control leaves over the subsequent 14 d. From this point onwards, the flag leaves from both treatments showed a loss of Rubisco, which was far more pronounced in the elevated-CO2 treatment, so that by 36 d the Rubisco content of these leaves was just 70% of that of the controls and by 52 d it was only 20%. At 36 d, there was no decline in chlorophyll, LHCII or the chloroplast ATPase coupling factor (CFI) in the elevated CO2 concentration treatment relative to the control. By 52 d, all of these proteins showed a significant decline relative to the control. This indicates that the decreased concentration of Rubisco at this final stage probably reflected earlier senescence in the elevated-CO2 treatment, but that this was preceded by a CO2-concentration-dependent decline in Rubisco.

Notes: The effects of elevated carbon dioxide (CO2) concentration on plant water use are best evaluated on plants grown under field conditions and with measurement techniques that do not disturb the natural function of the plant. Heat balance sap flow gauges were used on individual main stems of wheat (Triticum aestivum L. cv Yecora rojo) grown under normal ambient conditions (control) and in a free-air CO2 enrichment (FACE) system in Arizona with either high (control + high H2O = CW; FACE + high H2O = FW) or low (control + low H2O = CD; FACE + low H2O = FD) irrigation regimens. Over a 30d period (stem elongation to anthesis), combinations of treatments were monitored with,10–40 gauges per treatment. The effects of increased CO2 on tiller water use were inconsistent in both the diurnal patterns of sap flow and the statistical analyses of daily sap flow (Ftot). Initial results suggested that the reductions in Ftot, from CO2 enrichment were small (,0–10%) in relation to the H2O treatment effect (,20–30%). For a 3d period, Ftot of FW was,19–26% less than that of CW (P = 0.10). Examination of the different sources of variation in the study revealed that the location of gauges within the experimental plots influenced the variance of the sap flow measurements. This variation was probably related to positional variation in subsurface drip lines used to irrigate plots. A sampling design was proposed for use of sap flow gauges in FACE systems with subsurface irrigation that takes into account the main treatment effects of CO2 enrichment and the other sources of variation identified in this study. Despite the small and often statistically non-significant differences in Ftot between the CW and FW treatments, cumulative water use of the FW treatment at the end of the first three test periods ranged from 7 to 23% lower than that of the CW treatment. Differences in sap flow between FW and CW compared well with treatment differences in evapotranspiration. The results of the study, based on the first reported sap flow measurements of wheat, suggest that irrigation requirements for wheat production, in the present climatic regimen of the south-western US, may be predicted to decrease slightly because of increasing atmospheric CO2.

Notes: The present study was carried out to test the hypothesis thatelevated atmospheric CO2 (Ca) will alleviate over-excitationof the C4 photosynthetic apparatus and decrease non-photochemicalquenching (NPQ) during periods of limited water availability. Chlorophyll a fluorescencewas monitored in Sorghum bicolor plants grown under a free-aircarbon-dioxide enrichment (FACE) by water-stress (Dry) experiment.Under Dry conditions elevated Ca increased the quantum yield ofphotosystem II (φPSII) throughout the day throughincreases in both photochemical quenching coefficient (qp)and the efficiency with which absorbed quanta are transferred toopen PSII reaction centres (Fv′/Fm′).However, in the well-watered plants (Wets) FACE enhanced φPSIIonly at midday and was entirely attributed to changes in Fv′/Fm′. Underfield conditions, decreases in φPSII under Dry treatmentsand ambient Ca corresponded to increases in NPQ but the de-epoxidation stateof the xanthophyll pool (DPS) showed no effects. Water-stress didnot lead to long-term damage to the photosynthetic apparatus asindicated by φPSII and carbon assimilation measuredafter removal of stress conditions. We conclude that elevated Caenhances photochemical light energy usage in C4 photosynthesisduring drought and/or midday conditions. Additionally,NPQ protects against photo-inhibition and photodamage. However,NPQ and the xanthophyll cycle were affected differently by elevatedCa and water-stress.

Notes: Spring wheat was grown from emergence to grain maturity in two partial pressures of CO2 (pCO2): ambient air of nominally 37 Pa and air enriched with CO2 to 55 Pa using a free-air CO2 enrichment (FACE) apparatus. This experiment was the first of its kind to be conducted within a cereal field without the modifications or disturbance of microclimate and rooting environment that accompanied previous studies. It provided a unique opportunity to examine the hypothesis that continuous exposure of wheat to elevated pCO2 will lead to acclimatory loss of photosynthetic capacity. The diurnal courses of photosynthesis and conductance for upper canopy leaves were followed throughout the development of the crop and compared to model-predicted rates of photosynthesis. The seasonal average of midday photosynthesis rates was 28% greater in plants exposed to elevated pCO2 than in contols and the seasonal average of the daily integrals of photosynthesis was 21% greater in elevated pCO2 than in ambient air. The mean conductance at midday was reduced by 36%. The observed enhancement of photosynthesis in elevated pCO2 agreed closely with that predicted from a mechanistic biochemical model that assumed no acclimation of photosynthetic capacity. Measured values fell below predicted only in the flag leaves in the mid afternoon before the onset of grain-filling and over the whole diurnal course at the end of grain-filling. The loss of enhancement at this final stage was attributed to the earlier senescence of flag leaves in elevated pCO2. In contrast to some controlled-environment and field-enclosure studies, this field-scale study of wheat using free-air CO2 enrichment found little evidence of acclimatory loss of photosynthetic capacity with growth in elevated pCO2 and a significant and substantial increase in leaf photosynthesis throughout the life of the crop.

Notes: The mid-day responses of wheat ear CO2 and water vapour exchange to full-season CO2 enrichment were investigated using a Free-Air CO2 Enrichment (FACE) apparatus. Spring wheat [Triticum aestivum (L). cv. Yecora Rojo] was grown in two experiments under ambient and elevated atmospheric CO2 (Ca) concentrations (approximately 370 μmol mol−1 and 550 μmol mol−1, respectively) combined first with two irrigation (Irr) schemes (Wet: 100% and Dry: 50% replacement of evapotranspiration) and then with two levels of nitrogen (N) fertilization (High: 350, Low: 70 kg ha−1 N). Blowers were used for Ca enrichment. Ambient Ca plots were exposed to blower induced winds as well the Ca× N but not in the Ca× Irr experiment. The net photosynthesis for the ears was increased by 58% and stomatal conductance (gs) was decreased by 26% due to elevated Ca under ample water and N supply when blowers were applied to both Ca treatments. The use of blowers in the Ca-enriched plots only during the Ca× Irr experiment (blower effect) and Low N supply restricted the enhancement of net photosynthesis of the ear due to higher Ca. In the latter case, the increase of net photosynthesis of the ear amounted to 26%. The decrease in gs caused by higher Ca was not affected by the blower effect and N treatment. The mid-day enhancement of net photosynthesis due to elevated Ca was higher for ears than for flag leaves and this effect was most pronounced under ample water and N supply. The contribution of ears to grain filling is therefore likely to increase in higher Ca environments in the future. In the comparison between Wet and Dry, the higher Ca did not alter the response of net photosynthesis of the ear and gs to Irr. However, Ca enrichment increased the drought tolerance of net photosynthesis of the glume and delayed the increase of the awn portion of net photosynthesis of the ear during drought. Therefore, the role of awns for maintaining high net photosynthesis of the ear under drought may decrease when Ca increases.

Notes: Pinus eldarica L. trees, rooted in the natural soil of an agricultural field at Phoenix, Arizona, were grown from the seedling stage in clear-plastic-wall open-top enclosures maintained at four different atmospheric CO2 concentrations for 15 months. Light response functions were determined for one tree from each treatment by means of whole-tree net CO2 exchange measurements at the end of this period, after which rates of carbon assimilation of an ambient-treatment tree were measured across a range of atmospheric CO2 concentrations. The first of these data sets incorporates the consequences of both the CO2-induced enhancement of net photosynthesis per unit needle area and the CO2-induced enhancement of needle area itself (due primarily to the production of more needles), whereas the second data set reflects only the first of these effects. Hence the division of the normalized results of the first data set by the normalized results of the second set yields a representation of the increase in whole-tree net photosynthesis due to enhanced needle production caused by atmospheric CO2 enrichment. In the solitary trees we studied, the relative contribution of this effect increased rapidly with the CO2 concentration of the air to increase whole-tree net photosynthesis by nearly 50% at a CO2 concentration approximately 300 μmol mol−1 above ambient.