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Notes: A lower than theoretically expected increase in leaf photosynthesis with long-term elevation of carbon dioxide concentration ([CO2]) is often attributed to limitations in the capacity of the plant to utilize the additional photosynthate, possibly resulting from restrictions in rooting volume, nitrogen supply or genetic constraints. Field-grown, nitrogen-fixing soybean with indeterminate flowering might therefore be expected to escape these limitations. Soybean was grown from emergence to grain maturity in ambient air (372 µmol mol−1[CO2]) and in air enriched with CO2 (552 µmol mol−1[CO2]) using Free-Air CO2 Enrichment (FACE) technology. The diurnal courses of leaf CO2 uptake (A) and stomatal conductance (gs) for upper canopy leaves were followed throughout development from the appearance of the first true leaf to the completion of seed filling. Across the growing season the daily integrals of leaf photosynthetic CO2 uptake (A′) increased by 24.6% in elevated [CO2] and the average mid-day gs decreased by 21.9%. The increase in A′ was about half the 44.5% theoretical maximum increase calculated from Rubisco kinetics. There was no evidence that the stimulation of A was affected by time of day, as expected if elevated [CO2] led to a large accumulation of leaf carbohydrates towards the end of the photoperiod. In general, the proportion of assimilated carbon that accumulated in the leaf as non-structural carbohydrate over the photoperiod was small (〈 10%) and independent of [CO2] treatment. By contrast to A′, daily integrals of PSII electron transport measured by modulated chlorophyll fluorescence were not significantly increased by elevated [CO2]. This indicates that A at elevated [CO2] in these field conditions was predominantly ribulose-1,5-bisphosphate (RubP) limited rather than Rubisco limited. There was no evidence of any loss of stimulation toward the end of the growing season; the largest stimulation of A′ occurred during late seed filling. The stimulation of photosynthesis was, however, transiently lost for a brief period just before seed fill. At this point, daytime accumulation of foliar carbohydrates was maximal, and the hexose:sucrose ratio in plants grown at elevated [CO2] was significantly larger than that in plants grown at current [CO2]. The results show that even for a crop lacking the constraints that have been considered to limit the responses of C3 plants to rising [CO2] in the long term, the actual increase in A over the growing season is considerably less than the increase predicted from theory.

Notes: Abstract. The effect of growth temperatures on quantum yield (φ) was examined for leaves at different stages of development within the immature canopies of two crops of field grown maize (Zea mays cv. LG11) sown on 3 May and 20 June 1990. During the period of 23 to 49d after sowing, the crop sown on the 3 May experienced temperatures below 10°C on 19 occasions compared with only two for the crop sown on 20 June. A period of severe chilling at the end of May and the beginning of June was associated with a marked reduction in φ for all leaves in the early-sown crop. This chill-induced depression in φ was greater in recently emerged than more mature leaves in the canopy and was found to be accompanied by modifications in the polypeptide profiles of thylakoids isolated from the leaves. During the chilling period, decreases in some polypeptides, notably in the range of 41–42 and 20kDa apparent molecular size, and increases of polypeptides of c. 15–16kDa were observed compared with leaves developing at warmer temperatures in July. The efficiency of converting intercepted radiation into dry matter (conversion efficiency) was 42% lower in the early- than late-sown crop, but no significant relationship between conversion efficiency and quantum yield was found in either treatment.

Notes: The leaf model of C3 photosynthesis of Farquhar, von Caemmerer & Berry (Planta 149, 78–90, 1980) provides the basis for scaling carbon exchange from leaf to canopy and Earth-System models, and is widely used to project biosphere responses to global change. This scaling requires using the leaf model over a wider temperature range than that for which the model was originally parameterized. The leaf model assumes that photosynthetic CO2 uptake within a leaf is either limited by the rate of ribulose-1,5-bisphosphate (RuBP) regeneration or the activity of RuBP carboxylase-oxygenase (Rubisco). Previously we reported a re-parameterization of the temperature responses of Rubisco activity that proved robust when applied to a range of species. Herein this is extended to re-parameterizing the response of RuBP-limited photosynthesis to temperature. RuBP-limited photosynthesis is assumed to depend on the whole chain electron transport rate, which is described as a three-parameter non-rectangular hyperbolic function of photon flux. Herein these three parameters are determined from simultaneous measurement of chlorophyll fluorescence and CO2 exchange of tobacco leaves, at temperatures from 10 to 40 °C. All varied significantly with temperature and were modified further with variation in growth temperature from 15 to 35 °C. These parameters closely predicted the response of RuBP-limited photosynthesis to temperature measured in both lemon and poplar and showed a significant improvement over predictions based on earlier parameterizations. We provide the necessary equations for use of the model of Farquhar et al. (1980) with our newly derived temperature functions for predicting both Rubisco- and RuBP-limited photosynthesis.

Notes: Photosynthesis is commonly stimulated in grasslands with experimental increases in atmospheric CO2 concentration ([CO2]), a physiological response that could significantly alter the future carbon cycle if it persists in the long term. Yet an acclimation of photosynthetic capacity suggested by theoretical models and short-term experiments could completely remove this effect of CO2. Perennial ryegrass (Lolium perenne L. cv. Bastion) was grown under an elevated [CO2] of 600 µmol mol−1 for 10 years using Free Air CO2Enrichment (FACE), with two contrasting nitrogen levels and abrupt changes in the source : sink ratio following periodic harvests. More than 3000 measurements characterized the response of leaf photosynthesis and stomatal conductance to elevated [CO2] across each growing season for the duration of the experiment. Over the 10 years as a whole, growth at elevated [CO2] resulted in a 43% higher rate of light-saturated leaf photosynthesis and a 36% increase in daily integral of leaf CO2 uptake. Photosynthetic stimulation was maintained despite a 30% decrease in stomatal conductance and significant decreases in both the apparent, maximum carboxylation velocity (Vc,max) and the maximum rate of electron transport (Jmax). Immediately prior to the periodic (every 4–8 weeks) cuts of the L. perenne stands, Vc,max and Jmax, were significantly lower in elevated than in ambient [CO2] in the low-nitrogen treatment. This difference was smaller after the cut, suggesting a dependence upon the balance between the sources and sinks for carbon. In contrast with theoretical expectations and the results of shorter duration experiments, the present results provide no significant change in photosynthetic stimulation across a 10-year period, nor greater acclimation in Vc,max and Jmax in the later years in either nitrogen treatment.

Notes: Lolium perenne, a main component species in managed grassland, is well adapted to defoliation, fertilization, and regrowth cycles; and hence, to changes in the assimilatory carbon source-sink ratio. In the Swiss Free Air CO2 Enrichment experiment the source-sink ratio is (i) increased by elevated partial pressure of CO2 (pCO2), (ii) decreased by enhanced carbon use under high N fertilization, and (iii) gradually increased during regrowth after defoliation. Since sucrose synthesis plays a central role in leaf carbohydrate metabolism in this fructan-accumulating species, we investigated how sucrose-phosphate synthase (SPS) responds to the differing assimilatory carbon fluxes and source-sink ratios in the field. Assimilatory carbon flux, as estimated by leaf gas exchange, strongly depended on pCO2. Surprisingly, the SPS content per leaf area did not increase with pCO2, but increased with N fertilization. During later regrowth, when a dense canopy had formed, the SPS content decreased; in particular, SPS was decreased at high N under elevated pCO2. Further, the higher assimilatory carbon flux through SPS at elevated pCO2 was accompanied by a higher activation state of SPS. The SPS content correlated very strongly with the ratio of free sucrose to free amino acid in leaves, which represents the carbon source-sink ratio. Hence, SPS content in L. perenne appears to be regulated by the current, strongly nitrogen-dependent, source-sink relation.

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: Abstract. The activities of five active-oxygen scavenging enzymes were compared for cold-lability and three were compared for chilling induction in two Zea genotypes of contrasting susceptibility to photoinhibition during chilling. Activities of superoxide dismutase (SOD, EC 1.15.1.1), ascorbate peroxidase (APX, EC 1.11.1.11), monodehydroascorbate reductase (MDHAR, EC 1.6.5.4), dehydroascorbate reductase (DHAR, EC 1.8.5.1), and glutathione reductase (GTR, EC 1.6.4.2) in leaf extracts from plants grown without chilling stress were assayed at 19°C and 5°C. Enzymes from the chilling-susceptible Z. Mays cv. LG11 had lower specific activities at 5°C than did enzymes from the chilling-tolerant Z. diploperennis, except for MDHAR where no significant differences were observed. The activities of SOD and APX from Z. diploperennis were double those of Z. mays at both assay temperatures. Monodehydroa-scrobate reductase and glutathione reductase activities in both species were reduced by 63–78% at a 5°C assay temperature. The dehydroascorbate reductase (DHAR) showed the greatest low-temperature lability losing 96% (Z. diploperennis) and 100% (Z. mays) of its activity at 5°C. To examine possible chilling-induced changes in levels of enzyme activity, plants of both Zea genotypes were transferred to growth chambers at 10°C at moderate light intensities. Glutathione reductase activity was found to increase within 24h in Z. diploperennis, but it decreased slightly in Z. mays. MDHAR activity decreased by 50% in Z. diploperennis but showed only a transient increase in activity in Z. mays.

Notes: Surface ozone concentrations ([O3]) during the growing season in much of the northern temperate zone reach mean peak daily concentrations of 60 p.p.b. Concentrations are predicted to continue to rise over much of the globe during the next 50 years. Although these low levels of ozone may not induce visible symptoms on most vegetation, they can result in substantial losses of production and reproductive output. Establishing the vulnerability of vegetation to rising background ozone is complicated by marked differences in findings between individual studies. Ozone effects are influenced by exposure dynamics, nutrient and moisture conditions, and the species and cultivars that are investigated. Meta-analytic techniques provide an objective means to quantitatively summarize treatment responses. Soybean has been the subject of many studies of ozone effects. It is both the most widely planted dicotyledonous crop and a model for other C3 annual plants. Meta-analytic techniques were used to quantitatively summarize the response of soybean to an average, chronic ozone exposure of 70 p.p.b., from 53 peer-reviewed studies. At maturity, the average shoot biomass was decreased 34% and seed yield was 24% lower. Even in studies where [O3] was 〈 60 p.p.b., there was a significant decrease in biomass and seed production. At low [O3], decreased production corresponded to a decrease in leaf photosynthesis, but in higher [O3] the larger loss in production was associated with decreases in both leaf photosynthesis and leaf area. The impact of ozone increased with developmental stage, with little effect on vegetative growth and the greatest effect evident at completion of seed filling. Other stress treatments, including UV-B and drought, did not alter the ozone response. Elevated carbon dioxide significantly decreased ozone-induced losses, which may be explained by a significant decrease in stomatal conductance.

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.