Effects of free-air CO2 enrichment on the development of the photosynthetic apparatus in wheat, as indicated by changes in leaf proteins

A spring wheat crop was grown at ambient and elevated (550 micromol 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 to 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 a/b 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 photosystem 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 14d. 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 36d the Rubisco content of these leaves was just 70% of that of the controls and by 52d it was only 20%. At 36d, 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 52d, 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.