Spatial Distribution of Photosynthesis during Drought in Field-Grown and Acclimated and Nonacclimated Growth Chamber-Grown Cotton

Inhomogeneous photosynthetic activity has been reported to occur in drought-stressed leaves. In addition, it has been suggested that these water stress-induced nonuniformities in photosynthesis are caused by “patchy” stomatal closure and that the phenomenon may have created the illusion of a nonstomatal component to the inhibition of photosynthesis. Because these earlier studies were performed with nonacclimated growth chamber-grown plants, we sought to determine whether such “patches” existed in drought-treated, field-grown plants or in chamber-grown plants that had been acclimated to low leaf water potentials (ψ_leaf). Cotton (Gossypium hirsutum L.) was grown in the field and subjected to drought by withholding irrigation and rain from 24 d after planting. The distribution of photosynthesis, which may reflect the stomatal aperture distribution in a heterobaric species such as cotton, was assayed by autoradiography after briefly exposing attached leaves of field-grown plants to ¹⁴CO₂. A homogeneous distribution of radioactive photosynthate was evident even at the lowest ψ_leaf of −1.34 MPa. “Patchiness” could, however, be induced by uprooting the plant and allowing the shoot to air dry for 6 to 8 min. In parallel studies, growth chamber-grown plants were acclimated to drought by withholding irrigation for three 5-d drought cycles interspersed with irrigation. This drought acclimation lowered the ψ_leaf value at which control rates of photosynthesis could be sustained by approximately 0.7 MPa and was accompanied by a similar decline in the ψ_leaf at which patchiness first appeared. Photosynthetic inhomogeneities in chamber-grown plants that were visible during moderate water stress and ambient levels of CO₂ could be largely removed with elevated CO₂ levels (3000 μL L⁻¹), suggesting that they were stomatal in nature. However, advanced dehydration (less than approximately 2.0 MPa) resulted in “patches” that could not be so removed and were probably caused by nonstomatal factors. The demonstration that patches do not exist in drought-treated, field-grown cotton and that the presence of patches in chamber-grown plants can be altered by treatments that cause an acclimation of photosynthesis leads us to conclude that spatial heterogeneities in photosynthesis probably do not occur frequently under natural drought conditions.     

Selection and characterization of tobacco plants with novel o(2)-resistant photosynthesis

Plants were obtained with novel O₂-resistant photosynthetic characteristics. At low CO₂ (250-350 μL CO₂ L⁻¹) and 30°C when O₂ was increased from 1% to 21% to 42%, the ratio of net CO₂ uptake in O₂-resistant whole plants or leaf discs compared to wild type increased progressively, and this was not related to stomatal opening. Dihaploid plantlets regenerated from anther culture were initially screened and selected for O₂-resistant growth in 42% O₂/160 μL CO₂ L⁻¹ and 0.18% of the plantlets showed O₂-resistant photosynthesis. About 30% of the progeny (6 of 19 plants) of the first selfing of a fertile plant derived from a resistant dihaploid plant had O₂-resistant photosynthesis, and after a second selfing this increased to 50% (6 of 12 plants). In 21% O₂ and low CO₂, net photosynthesis of the resistant plants was about 15% greater on a leaf area basis than wild type. Net photosynthesis was compared in leaf discs at 30 and 38°C in 21% O₂, and at the higher temperature O₂-resistant plants showed still greater photosynthesis than wild type. The results suggest that the O₂-resistant photosynthesis described here is associated with a decreased stoichiometry of CO₂ release under conditions of rapid photorespiration. This view was supported by the finding that leaves of O₂-resistant plants averaged 40% greater catalase activity than wild type.     

Effects of Irradiance during Growth on Adaptive Photosynthetic Characteristics of Velvetleaf and Cotton

We grew velvetleaf (Abutilon theophrasti Medic.) and cotton (Gossypium hirsutum L. var. Stoneville 213) at three irradiances and determined the photosynthetic responses of single leaves to a range of six irradiances from 90 to 2000 μeinsteins m⁻²sec⁻¹. In air containing 21% O₂, velvetleaf and cotton grown at 750 μeinsteins m⁻²sec⁻¹ had maximum photosynthetic rates of 18.4 and 21.9 mg of CO₂ dm⁻²hr⁻¹, respectively. Maximum rates for leaves grown at 320 and 90 μeinsteins m⁻²sec⁻¹ were 15.3 and 10.3 mg of CO₂ dm⁻²hr⁻¹ in velvetleaf and 12 and 6.7 mg of CO₂ dm⁻²hr⁻¹ in cotton, respectively. In 1 O₂, maximum photosynthetic rates were 1.5 to 2.3 times the rates in air containing 21% O₂, and plants grown at medium and high irradiance did not differ in rate. In both species, stomatal conductance was not significantly affected by growth irradiance. The differences in maximum photosynthetic rates were associated with differences in mesophyll conductance. Mesophyll conductance increased with growth irradiance and correlated positively with mesophyll thickness or volume per unit leaf area, chlorophyll content per unit area, and photosynthetic unit density per unit area. Thus, quantitative changes in the photosynthetic apparatus help account for photosynthetic adaptation to irradiance in both species. Net assimilation rates calculated for whole plants by mathematical growth analysis were closely correlated with single-leaf photosynthetic rates.     

Photosynthesis and Carbon Partitioning in Transgenic Tobacco Plants Deficient in Leaf Cytosolic Pyruvate Kinase

Whole-plant diurnal C exchange analysis provided a noninvasive estimation of daily net C gain in transgenic tobacco (Nicotiana tabacum L.) plants deficient in leaf cytosolic pyruvate kinase (PK_c−). PK_c− plants cultivated under a low light intensity (100 μmol m⁻² s⁻¹) were previously shown to exhibit markedly reduced root growth, as well as delayed shoot and flower development when compared with plants having wild-type levels of PK_c (PK_c+). PK_c− and PK_c+ source leaves showed a similar net C gain, photosynthesis over a range of light intensities, and a capacity to export newly fixed ¹⁴CO₂ during photosynthesis. However, during growth under low light the nighttime, export of previously fixed ¹⁴CO₂ by fully expanded PK_c− leaves was 40% lower, whereas concurrent respiratory ¹⁴CO₂ evolution was 40% higher than that of PK_c+ leaves. This provides a rationale for the reduced root growth of the PK_c− plants grown at low irradiance. Leaf photosynthetic and export characteristics in PK_c− and PK_c+ plants raised in a greenhouse during winter months resembled those of plants grown in chambers at low irradiance. The data suggest that PK_c in source leaves has a critical role in regulating nighttime respiration particularly when the available pool of photoassimilates for export and leaf respiratory processes are low.     

Thermotolerance of Leaf Discs from Four Isoprene-Emitting Species Is Not Enhanced by Exposure to Exogenous Isoprene

The effects of exogenously supplied isoprene on chlorophyll fluorescence characteristics were examined in leaf discs of four isoprene-emitting plant species, kudzu (Pueraria lobata [Willd.] Ohwi.), velvet bean (Mucuna sp.), quaking aspen (Populus tremuloides Michx.), and pussy willow (Salix discolor Muhl). Isoprene, supplied to the leaves at either 18 μL L⁻¹ in compressed air or 21 μL L⁻¹ in N₂, had no effect on the temperature at which minimal fluorescence exhibited an upward inflection during controlled increases in leaf-disc temperature. During exposure to 1008 μmol photons m⁻² s⁻¹ in an N₂ atmosphere, 21 μL L⁻¹ isoprene had no effect on the thermally induced inflection of steady-state fluorescence. The maximum quantum efficiency of photosystem II photochemistry decreased sharply as leaf-disc temperature was increased; however, this decrease was unaffected by exposure of leaf discs to 21 μL L⁻¹ isoprene. Therefore, there were no discernible effects of isoprene on the occurrence of symptoms of high-temperature damage to thylakoid membranes. Our data do not support the hypothesis that isoprene enhances leaf thermotolerance.     

Effects of phosphorus deficiency on the photosynthesis and respiration of leaves of sugar beet

Phosphorus deficiency was induced in sugar beet plants (Beta vulgaris L. var. F5855441), cultured hydroponically under standardized environmental conditions, by removal of phosphorus from the nutrient supply at the ten leaf stage 28 days after germination. CO₂ and water vapor exchange rates of individual attached leaves were determined at intervals after P cutoff. Leaves grown with an adequate nutrient supply attained net rates of photosynthetic CO₂ fixation of 125 ng CO₂ cm⁻² sec⁻¹ at saturating irradiance, 25 C, and an ambient CO₂ concentration of about 250 μl l⁻¹. After P cutoff, leaf phosphorus concentrations decreased as did net rates of photosynthetic CO₂ uptake, photorespiratory evolution of CO₂ into CO₂-free air, and dark respiration, so that 30 days after cutoff these rates were about one-third of the control rates. The decrease in photosynthetic rates during the first 15 days after cutoff was associated with increased mesophyll resistance (r_m) which increased from 2.4 to 4.9 sec cm⁻¹, while from 15 to 30 days there was an increase in leaf (mainly stomatal) diffusion resistance (r_l′) from 0.3 to 0.9 sec cm⁻¹, as well as further increases in r_m to 8.5 sec cm⁻¹. Leaf diffusion resistance (r_l′) was increased greatly by low P at low but not at high irradiance, r_l′ for plants at low P reaching values as high as 9 sec cm⁻¹.     

Growth and Nitrogen Uptake Kinetics in Cultured Prorocentrum donghaiense

We compared growth kinetics of Prorocentrum donghaiense cultures on different nitrogen (N) compounds including nitrate (NO₃ ⁻), ammonium (NH₄ ⁺), urea, glutamic acid (glu), dialanine (diala) and cyanate. P. donghaiense exhibited standard Monod-type growth kinetics over a range of N concentraions (0.5–500 μmol N L⁻¹ for NO₃ ⁻ and NH₄ ⁺, 0.5–50 μmol N L⁻¹ for urea, 0.5–100 μmol N L⁻¹ for glu and cyanate, and 0.5–200 μmol N L⁻¹ for diala) for all of the N compounds tested. Cultures grown on glu and urea had the highest maximum growth rates (μ_m, 1.51±0.06 d⁻¹ and 1.50±0.05 d⁻¹, respectively). However, cultures grown on cyanate, NO₃ ⁻, and NH₄ ⁺ had lower half saturation constants (K_μ, 0.28–0.51 μmol N L⁻¹). N uptake kinetics were measured in NO₃ ⁻-deplete and -replete batch cultures of P. donghaiense. In NO₃ ⁻-deplete batch cultures, P. donghaiense exhibited Michaelis-Menten type uptake kinetics for NO₃ ⁻, NH₄ ⁺, urea and algal amino acids; uptake was saturated at or below 50 μmol N L⁻¹. In NO₃ ⁻-replete batch cultures, NH₄ ⁺, urea, and algal amino acid uptake kinetics were similar to those measured in NO₃ ⁻-deplete batch cultures. Together, our results demonstrate that P. donghaiense can grow well on a variety of N sources, and exhibits similar uptake kinetics under both nutrient replete and deplete conditions. This may be an important factor facilitating their growth during bloom initiation and development in N-enriched estuaries where many algae compete for bioavailable N and the nutrient environment changes as a result of algal growth.     

Drought Stress and Elevated CO(2) Effects on Soybean Ribulose Bisphosphate Carboxylase Activity and Canopy Photosynthetic Rates

Soybean (Glycine max [L.] cv Bragg) was grown at 330 or 660 microliters CO₂ per liter in outdoor, controlled-environment chambers. When the plants were 50 days old, drought stress was imposed by gradually reducing irrigation each evening so that plants wilted earlier each succeeding day. On the ninth day, as the pots ran out of water CO₂ exchange rate (CER) decreased rapidly to near zero for the remainder of the day. Both CO₂-enrichment and drought stress reduced the total (HCO₃⁻/Mg²⁺-activated) extractable ribulose-1,5-bisphosphate carboxylase (RuBPCase) activity, as expressed on a chlorophyll basis. In addition, drought stress when canopy CER values and leaf water potentials were lowest, reduced the initial (nonactivated) RuBPCase activity by 50% compared to the corresponding unstressed treatments. This suggests that moderate to severe drought stress reduces the in vivo activation state of RuBPCase, as well as lowers the total activity. It is hypothesized that stromal acidification under drought stress causes the lowered initial RuBPCase activities. The K_m(CO₂) values of activated RuBPCase from stressed and unstressed plants were similar; 15.0 and 12.6 micromolar, respectively. RuBP levels were 10 to 30% lower in drought stressed as compared to unstressed treatments. However, RuBP levels increased from near zero at night to around 150 to 200 nanomoles per milligram chlorophyll during the day, even as water potentials and canopy CERs decreased. This suggests that the rapid decline in canopy CER cannot be attributed to drought stress induced limitations in the RuBP regeneration capability. Thus, in soybean leaves, a nonstomatal limitation of leaf photosynthesis under drought stress conditions appears due, in part, to a reduction of the in vivo activity of RuBPCase. Because initial RuBPCase activities were not reduced as much as canopy CER values, this enzymic effect does not explain entirely the response of soybean photosynthesis to drought stress.     

Interactive Effects of Ocean Acidification and Nitrogen-Limitation on the Diatom Phaeodactylum tricornutum

Climate change is expected to bring about alterations in the marine physical and chemical environment that will induce changes in the concentration of dissolved CO₂ and in nutrient availability. These in turn are expected to affect the physiological performance of phytoplankton. In order to learn how phytoplankton respond to the predicted scenario of increased CO₂ and decreased nitrogen in the surface mixed layer, we investigated the diatom Phaeodactylum tricornutum as a model organism. The cells were cultured in both low CO₂ (390 μatm) and high CO₂ (1000 μatm) conditions at limiting (10 μmol L⁻¹) or enriched (110 μmol L⁻¹) nitrate concentrations. Our study shows that nitrogen limitation resulted in significant decreases in cell size, pigmentation, growth rate and effective quantum yield of Phaeodactylum tricornutum, but these parameters were not affected by enhanced dissolved CO₂ and lowered pH. However, increased CO₂ concentration induced higher rETR_max and higher dark respiration rates and decreased the CO₂ or dissolved inorganic carbon (DIC) affinity for electron transfer (shown by higher values for K_{1/2 DIC} or K_{1/2 CO2}). Furthermore, the elemental stoichiometry (carbon to nitrogen ratio) was raised under high CO₂ conditions in both nitrogen limited and nitrogen replete conditions, with the ratio in the high CO₂ and low nitrate grown cells being higher by 45% compared to that in the low CO₂ and nitrate replete grown ones. Our results suggest that while nitrogen limitation had a greater effect than ocean acidification, the combined effects of both factors could act synergistically to affect marine diatoms and related biogeochemical cycles in future oceans.     

Transition from Anoxygenic to Oxygenic Photosynthesis in a Microcoleus chthonoplastes Cyanobacterial Mat

Benthic cyanobacterial mats with the filamentous Microcoleus chthonoplastes as the dominant phototroph grow in oxic hypersaline environments such as Solar Lake, Sinai. The cyanobacteria are in situ exposed to chemical variations between 200 μmol of sulfide liter⁻¹ at night and 1 atm pO₂ during the day. During experimental H₂S to O₂ transitions the microbial community was shown to shift from anoxygenic photosynthesis, with H₂S as the electron donor, to oxygenic photosynthesis. Microcoleus filaments could carry out both types of photosynthesis concurrently. Anoxygenic photosynthesis dominated at high sulfide levels, 500 μmol liter⁻¹, while the oxygenic reaction became dominant when the sulfide level was reduced below 100 to 300 μmol liter⁻¹ (25 to 75 μmol of H₂S liter⁻¹). An increasing inhibition of the oxygenic photosynthesis was observed upon transition to oxic conditions from increasing sulfide concentrations. Oxygen built up within the Microcoleus layer of the mat even under 5 mmol of sulfide liter⁻¹ (500 μmol of H₂S liter⁻¹) in the overlying water. The implications of such a localized O₂ production in a highly reducing environment are discussed in relation to the evolution of oxygenic photosynthesis during the Proterozoic era.     

Effects of sulfur on the photosynthesis of intact leaves and isolated chloroplasts of sugar beets

Effects of sulfur on photosynthesis in sugar beets (Beta vulgaris L. cv. F58-554H1) were studied by inducing sulfur deficiency and determining changes in the photosynthesis of whole attached leaves and of isolated chloroplasts. The rates of photosynthetic CO₂ uptake by intact leaves, photoreduction of ferricyanide, cyclic and noncyclic photophosphorylation of isolated chloroplasts, and the rate of CO₂ assimilation by ribulose diphosphate carboxylase, decreased with decrease in total leaf sulfur from 2500 to about 500 μg g⁻¹ dry weight. Sulfur deficiency reduced photosynthesis through an effect on chlorophyll content, which decreased linearly with leaf sulfur, and by decreasing the rate of photosynthesis per unit chlorophyll. There was only a small effect of sulfur deficiency on stomatal diffusion resistance to CO₂ until leaf sulfur decreased below 1000 μg g⁻¹ when stomatal resistance became a more significant proportion of the total diffusion resistance to CO₂. Light respiration rates were positively correlated with photosynthesis rates and dark respiration was unchanged as leaf sulfur concentrations declined.     

Light acclimation during and after leaf expansion in soybean

Soybean plants (Glycine max var. Ransom) were grown at light intensities of 850 and 250 μeinsteins m⁻² sec⁻¹ of photosynthetically active radiation. A group of plants was shifted from each environment into the other environment 24 hours before the beginning of the experiment. Net photosynthetic rates and stomatal conductances were measured at 2,000 and 100 μeinsteins m⁻² sec⁻¹ photosynthetically active radiation on the 1st, 2nd, and 5th days of the experiment to determine the time course of photosynthetic light adaptation. The following factors were also measured: dark respiration, leaf water potential, leaf thickness, internal surface area per external surface area, chlorophyll content, photosynthetic unit size and number, specific leaf weight, and activities of malate dehydrogenase, and glycolate oxidase. Comparisons were made with plants maintained in either 850 or 250 μeinsteins m⁻² sec⁻¹ environments. Changes in photosynthesis, stomatal conductance, leaf anatomy, leaf water potential, photosynthetic unit size, and glycolate oxidase activity occurred upon altering the light environment, and were complete within 1 day, whereas chlorophyll content, numbers of photosynthetic units, specific leaf weight, and malate dehydrogenase activity showed slower changes. Differences in photosynthetic rates at high light were largely accounted for by internal surface area differences with low environmental light associated with low internal area and low photosynthetic rate. An exception to this was the fact that plants grown at 250 μeinsteins m⁻² sec⁻¹ then switched to 850 μeinsteins m⁻² sec⁻¹ showed lower photosynthesis at high light than any other treatment. This was associated with higher glycolate oxidase and malate dehydrogenase activity. Photosynthesis at low light was higher in plants kept at or switched to the lower light environment. This increased rate was associated with larger photosynthetic unit size, and lower dark respiration and malate dehydrogenase activity. Both anatomical and physiological changes with environmental light occurred even after leaf expansion was complete and both were important in determining photosynthetic response to light.     

Uptake of HCO3? and CO2 in Cells and Chloroplasts from the Microalgae Chlamydomonas reinhardtii and Dunaliella tertiolecta

Mass-spectrometric disequilibrium analysis was applied to investigate CO₂ uptake and HCO₃⁻ transport in cells and chloroplasts of the microalgae Dunaliella tertiolecta and Chlamydomonas reinhardtii, which were grown in air enriched with 5% (v/v) CO₂ (high-Ci cells) or in ambient air (low-Ci cells). High- and low-Ci cells of both species had the capacity to transport CO₂ and HCO₃⁻, with maximum rates being largely unaffected by the growth conditions. In high- and low-Ci cells of D. tertiolecta, HCO₃⁻ was the dominant inorganic C species taken up, whereas HCO₃⁻ and CO₂ were used at similar rates by C. reinhardtii. The apparent affinities of HCO₃⁻ transport and CO₂ uptake increased 3- to 9-fold in both species upon acclimation to air. Photosynthetically active chloroplasts isolated from both species were able to transport CO₂ and HCO₃⁻. For chloroplasts from C. reinhardtii, the concentrations of HCO₃⁻ and CO₂ required for half-maximal activity declined from 446 to 33 μm and 6.8 to 0.6 μm, respectively, after acclimation of the parent cells to air; the corresponding values for chloroplasts from D. tertiolecta decreased from 203 to 58 μm and 5.8 to 0.5 μm, respectively. These results indicate the presence of inducible high-affinity HCO₃⁻ and CO₂ transporters at the chloroplast envelope membrane.     

Overexpression of Iron Superoxide Dismutase in Transformed Poplar Modifies the Regulation of Photosynthesis at Low CO2 Partial Pressures or Following Exposure to the Prooxidant Herbicide Methyl Viologen

Chloroplast-targeted overexpression of an Fe superoxide dismutase (SOD) from Arabidopsis thaliana resulted in substantially increased foliar SOD activities. Ascorbate peroxidase, glutathione reductase, and monodehydroascorbate reductase activities were similar in the leaves from all of the lines, but dehydroascorbate reductase activity was increased in the leaves of the FeSOD transformants relative to untransformed controls. Foliar H₂O₂, ascorbate, and glutathione contents were comparable in all lines of plants. Irradiance-dependent changes in net CO₂ assimilation and chlorophyll a fluorescence quenching parameters were similar in all lines both in air (21% O₂) and at low (1%) O₂. CO₂-response curves for photosynthesis showed similar net CO₂-exchange characteristics in all lines. In contrast, values of photochemical quenching declined in leaves from untransformed controls at intercellular CO₂ (Ci) values below 200 μL L⁻¹ but remained constant with decreasing Ci in leaves of FeSOD transformants. When the O₂ concentration was decreased from 21 to 1%, the effect of FeSOD overexpression on photochemical quenching at limiting Ci was abolished. At high light (1000 μmol m⁻² s⁻¹) a progressive decrease in the ratio of variable (F_v) to maximal (F_m) fluorescence was observed with decreasing temperature. At 6^oC the high-light-induced decrease in the F_v/F_m ratio was partially prevented by low O₂ but values were comparable in all lines. Methyl viologen caused decreased F_v/F_m ratios, but this was less marked in the FeSOD transformants than in the untransformed controls. These observations suggest that the rate of superoxide dismutation limits flux through the Mehler-peroxidase cycle in certain conditions.     

The metabolic origins of non-photorespiratory CO2 release during photosynthesis: a metabolic flux analysis

Respiration in the light (R_L) releases CO₂ in photosynthesizing leaves and is a phenomenon that occurs independently from photorespiration. Since R_L lowers net carbon fixation, understanding R_L could help improve plant carbon-use efficiency and models of crop photosynthesis. Although R_L was identified more than 75 years ago, its biochemical mechanisms remain unclear. To identify reactions contributing to R_L, we mapped metabolic fluxes in photosynthesizing source leaves of the oilseed crop and model plant camelina (Camelina sativa). We performed a flux analysis using isotopic labeling patterns of central metabolites during ¹³CO₂ labeling time course, gas exchange, and carbohydrate production rate experiments. To quantify the contributions of multiple potential CO₂ sources with statistical and biological confidence, we increased the number of metabolites measured and reduced biological and technical heterogeneity by using single mature source leaves and quickly quenching metabolism by directly injecting liquid N₂; we then compared the goodness-of-fit between these data and data from models with alternative metabolic network structures and constraints. Our analysis predicted that R_L releases 5.2 μmol CO₂ g⁻¹ FW h⁻¹ of CO₂, which is relatively consistent with a value of 9.3 μmol CO₂ g⁻¹ FW h⁻¹ measured by CO₂ gas exchange. The results indicated that ≤10% of R_L results from TCA cycle reactions, which are widely considered to dominate R_L. Further analysis of the results indicated that oxidation of glucose-6-phosphate to pentose phosphate via 6-phosphogluconate (the G6P/OPP shunt) can account for >93% of CO₂ released by R_L.     

Differential Control of Xanthophylls and Light-Induced Stress Proteins, as Opposed to Light-Harvesting Chlorophyll a/b Proteins, during Photosynthetic Acclimation of Barley Leaves to Light Irradiance

Barley (Hordeum vulgare L.) plants were grown at different photon flux densities ranging from 100 to 1800 μmol m⁻² s⁻¹ in air and/or in atmospheres with reduced levels of O₂ and CO₂. Low O₂ and CO₂ partial pressures allowed plants to grow under high photosystem II (PSII) excitation pressure, estimated in vivo by chlorophyll fluorescence measurements, at moderate photon flux densities. The xanthophyll-cycle pigments, the early light-inducible proteins, and their mRNA accumulated with increasing PSII excitation pressure irrespective of the way high excitation pressure was obtained (high-light irradiance or decreased CO₂ and O₂ availability). These findings indicate that the reduction state of electron transport chain components could be involved in light sensing for the regulation of nuclear-encoded chloroplast gene expression. In contrast, no correlation was found between the reduction state of PSII and various indicators of the PSII light-harvesting system, such as the chlorophyll a-to-b ratio, the abundance of the major pigment-protein complex of PSII (LHCII), the mRNA level of LHCII, the light-saturation curve of O₂ evolution, and the induced chlorophyll-fluorescence rise. We conclude that the chlorophyll antenna size of PSII is not governed by the redox state of PSII in higher plants and, consequently, regulation of early light-inducible protein synthesis is different from that of LHCII.     

Phosphorus application and elevated CO2 enhance drought tolerance in field pea grown in a phosphorus-deficient vertisol

Background and Aims Benefits to crop productivity arising from increasing CO₂ fertilization may be offset by detrimental effects of global climate change, such as an increasing frequency of drought. Phosphorus (P) nutrition plays an important role in crop responses to water stress, but how elevated CO₂ (eCO₂) and P nutrition interact, especially in legumes, is unclear. This study aimed to elucidate whether P supply improves plant drought tolerance under eCO₂.Methods A soil-column experiment was conducted in a free air CO₂ enrichment (SoilFACE) system. Field pea (Pisum sativum) was grown in a P-deficient vertisol, supplied with 15 mg P kg⁻¹ (deficient) or 60 mg P kg⁻¹ (adequate for crop growth) and exposed to ambient CO₂ (aCO₂; 380–400 ppm) or eCO₂ (550–580 ppm). Drought treatments commenced at flowering. Measurements were taken of soil and leaf water content, photosynthesis, stomatal conductance, total soluble sugars and inorganic P content (Pi).Key Results Water-use efficiency was greatest under eCO₂ when the plants were supplied with adequate P compared with other treatments irrespective of drought treatment. Elevated CO₂ decreased stomatal conductance and transpiration rate, and increased the concentration of soluble sugars and relative water contents in leaves. Adequate P supply increased concentrations of soluble sugars and Pi in drought-stressed plants. Adequate P supply but not eCO₂ increased root length distribution in deeper soil layers.Conclusions Phosphorus application and eCO₂ interactively enhanced periodic drought tolerance in field pea as a result of decreased stomatal conductance, deeper rooting and high Pi availability for carbon assimilation in leaves.     

Carbon gain and photosynthetic response of chrysanthemum to photosynthetic photon flux density cycles

Most models of carbon gain as a function of photosynthetic irradiance assume an instantaneous response to increases and decreases in irradiance. High- and low-light-grown plants differ, however, in the time required to adjust to increases and decreases in irradiance. In this study the response to a series of increases and decreases in irradiance was observed in Chrysanthemum × morifolium Ramat. “Fiesta” and compared with calculated values assuming an instantaneous response. There were significant differences between high- and low-light-grown plants in their photosynthetic response to four sequential photosynthetic photon flux density (PPFD) cycles consisting of 5-minute exposures to 200 and 400 micromoles per square meter per second (μmol m⁻²s⁻¹). The CO₂ assimilation rate of high-light-grown plants at the cycle peak increased throughout the PPFD sequence, but the rate of increase was similar to the increase in CO₂ assimilation rate observed under continuous high-light conditions. Low-light leaves showed more variability in their response to light cycles with no significant increase in CO₂ assimilation rate at the cycle peak during sequential cycles. Carbon gain and deviations from actual values (percentage carbon gain over- or underestimation) based on assumptions of instantaneous response were compared under continuous and cyclic light conditions. The percentage carbon gain overestimation depended on the PPFD step size and growth light level of the leaf. When leaves were exposed to a large PPFD increase, the carbon gain was overestimated by 16 to 26%. The photosynthetic response to 100 μmol m⁻² s⁻¹ PPFD increases and decreases was rapid, and the small overestimation of the predicted carbon gain, observed during photosynthetic induction, was almost entirely negated by the carbon gain underestimation observed after a decrease. If the PPFD cycle was 200 or 400 μmol m⁻² s⁻¹, high- and low-light leaves showed a carbon gain overestimation of 25% that was not negated by the underestimation observed after a light decrease. When leaves were exposed to sequential PPFD cycles (200-400 μmol m⁻² s⁻¹), carbon gain did not differ from leaves exposed to a single PPFD cycle of identical irradiance integral that had the same step size (200-400-200 μmol m⁻² s⁻¹) or mean irradiance (200-300-200 μmol m⁻² s⁻¹).     

Estimating Photosynthesis and Concurrent Export Rates in C3 and C4 Species at Ambient and Elevated CO2

The ability of 21 C₃ and C₄ monocot and dicot species to rapidly export newly fixed C in the light at both ambient and enriched CO₂ levels was compared. Photosynthesis and concurrent export rates were estimated during isotopic equilibrium of the transport sugars using a steady-state ¹⁴CO₂-labeling procedure. At ambient CO₂ photosynthesis and export rates for C₃ species were 5 to 15 and 1 to 10 μmol C m⁻² s⁻¹, respectively, and 20 to 30 and 15 to 22 μmol C m⁻² s⁻¹, respectively, for C₄ species. A linear regression plot of export on photosynthesis rate of all species had a correlation coefficient of 0.87. When concurrent export was expressed as a percentage of photosynthesis, several C₃ dicots that produced transport sugars other than Suc had high efflux rates relative to photosynthesis, comparable to those of C₄ species. At high CO₂ photosynthetic and export rates were only slightly altered in C₄ species, and photosynthesis increased but export rates did not in all C₃ species. The C₃ species that had high efflux rates relative to photosynthesis at ambient CO₂ exported at rates comparable to those of C₄ species on both an absolute basis and as a percentage of photosynthesis. At ambient CO₂ there were strong linear relationships between photosynthesis, sugar synthesis, and concurrent export. However, at high CO₂ the relationships between photosynthesis and export rate and between sugar synthesis and export rate were not as strong because sugars and starch were accumulated.     

Does long-term cultivation of saplings under elevated CO2 concentration influence their photosynthetic response to temperature?

Background and Aims Plants growing under elevated atmospheric CO₂ concentrations often have reduced stomatal conductance and subsequently increased leaf temperature. This study therefore tested the hypothesis that under long-term elevated CO₂ the temperature optima of photosynthetic processes will shift towards higher temperatures and the thermostability of the photosynthetic apparatus will increase.Methods The hypothesis was tested for saplings of broadleaved Fagus sylvatica and coniferous Picea abies exposed for 4–5 years to either ambient (AC; 385 µmol mol⁻¹) or elevated (EC; 700 µmol mol⁻¹) CO₂ concentrations. Temperature response curves of photosynthetic processes were determined by gas-exchange and chlorophyll fluorescence techniques.Key Results Initial assumptions of reduced light-saturated stomatal conductance and increased leaf temperatures for EC plants were confirmed. Temperature response curves revealed stimulation of light-saturated rates of CO₂ assimilation (A_max) and a decline in photorespiration (R_L) as a result of EC within a wide temperature range. However, these effects were negligible or reduced at low and high temperatures. Higher temperature optima (T_opt) of A_max, Rubisco carboxylation rates (V_Cmax) and R_L were found for EC saplings compared with AC saplings. However, the shifts in T_opt of A_max were instantaneous, and disappeared when measured at identical CO₂ concentrations. Higher values of T_opt at elevated CO₂ were attributed particularly to reduced photorespiration and prevailing limitation of photosynthesis by ribulose-1,5-bisphosphate (RuBP) regeneration. Temperature response curves of fluorescence parameters suggested a negligible effect of EC on enhancement of thermostability of photosystem II photochemistry.Conclusions Elevated CO₂ instantaneously increases temperature optima of A_max due to reduced photorespiration and limitation of photosynthesis by RuBP regeneration. However, this increase disappears when plants are exposed to identical CO₂ concentrations. In addition, increased heat-stress tolerance of primary photochemistry in plants grown at elevated CO₂ is unlikely. The hypothesis that long-term cultivation at elevated CO₂ leads to acclimation of photosynthesis to higher temperatures is therefore rejected. Nevertheless, incorporating acclimation mechanisms into models simulating carbon flux between the atmosphere and vegetation is necessary.