Photosynthetic Traits in Wheat Grown under Decreased and Increased CO2 Concentration, and after Transfer to Natural CO2 concentration

Wheat plants were grown from sowing to day 18 in 26-dm³ chambers at three different CO₂ concentrations: 150 (-CO₂), 350 (C, control), 800 (+CO₂) mol mol^-1. Afterwards, plants of the three variants were grown at the same natural CO₂ concentration. Plant characteristics were measured just before the transfer (0 days after CO₂ treatment, DAT), and at 5 – 8 DAT on the 1^st leaf, and at 12 – 22 DAT on the 4^th leaf. Decreased or increased CO₂ concentrations caused acclimations which persisted after transplantation to natural CO₂ concentration. At 5 – 8 DAT, stomatal density, stomatal conductance (g_s), CO₂ saturated net photosynthetic rate (P_Nsat0), radiation saturated net photosynthetic rate (P_Nsat1), and carboxylation efficiency () were higher in -CO₂ plants and lower in +CO₂ plants than in C plants. As compared with C plants, the photochemical efficiency () was lower in -CO₂ and higher in -CO₂ plants, however, chlorophyll (Chl) a, Chl b, Chl a–b and carotenoid contents were lower in both -CO₂ and +CO₂ plants. On the 4^th leaf, which emerged on plant after finishing CO₂ treatments, at 12 – 22 DAT, no differences in stomatal density and g, between treatments were observed. In -CO₂ plants, pigment content and P_Nsat0 were higher, was lower, and P_Nsat1 and were not different from C plants. In contrast, in +CO₂ plants, pigment content, P_Nsat1 and were lower, and P_Nsat0 and were unchanged. Leaf area, dry mass, and tiller development increased in +CO₂ plants and decreased in -CO₂ plants. In the interval between 8 and 22 DAT, lower net assimilation rate in +CO₂ than in -CO₂ plants was observed.     

不同氮营养水平下草莓叶片光合作用对高CO2浓度的适应

研究了不同氮素水平（12mmol／L,4mmol／L,0、4mmol／L）下生长的‘丰香’草莓在富C02（700μL/L）和大气CO（390μL/L）下的光合作用。结果表明,高氮（12mmol／L）下,在富CO2环境中生长的‘丰香’草莓叶片未出现光合作用下调,富CO2下草莓叶片的净光合速率、最大羧化速率（Vc.max）、最大电子传递速率（Jmax）、碳同化的电子传递速率（Jc）和光化学猝灭系数（qp）等均显著提高;而在中氮（4mmol／L）、低氮（0．4mmol／L）下,富CO2下生长的草莓叶片的上述参数均出现不同程度的下降。富CO2下,无论氮素水平如何,草莓叶片的光呼吸电子传递速率（Jo）均降低高氮草莓叶片的非光化学猝灭系数（qN或NPQ）降低,光抑制降低,而低氮则相反。上述结果说明,氮素供应不足时草莓叶片在富CO2下光合作用出现下调,因此生产上进行CO2施肥时应适度增加氮素的供应。     

CO2 浓度升高对两种沈阳城市森林树种光合特性的影响

利用开顶式气室, 研究了CO2浓度升高条件下城市森林主要树种油松(Pinus tabulaefomis)和银杏(Ginkgo biloba)主要光合特性的变化。结果表明, 整个生长季, CO2浓度升高(700 mmol.mol-1)条件下2树种叶片的净光合速率、可溶性糖、淀粉和可溶性蛋白含量均接近或高于相应对照(自然CO2浓度)值, 但不同树种增加的幅度不同; 而2树种的叶绿素含量和Chl a/Chl b值对CO2浓度升高反应不一, 表现为CO2浓度升高条件下油松的叶绿素含量较对照值高, Chl a/Chl b值降低, 银杏的叶绿素含量为前期升高, 后期降低, Chl a/Chl b值变化与之正好相反, 说明城市森林组成树种对CO2浓度升高的响应具有复杂性。CO2浓度升高条件下, 两树种均未发生光合适应现象。     

Flavonoid Responses in Wheat Grown at Elevated CO2: Green Versus Senescent Leaves

We compared flavonoids in green, mature, and senescing flag leaves of wheat grown under ambient (AC - 370 mol mol^-1) and elevated (EC - 550 mol mol^-1) concentrations of CO₂ in a FACE (Free Air CO₂ Enrichment) system. The concentrations of flag leaf flavonoids (e.g., isoorientin and tricin) decreased to one third in mature leaves, and the majoritary isoorientin almost disappeared in senescing leaves. Flavonoid concentrations increased in green well-developed flag leaves under EC (46 % isoorientin and 55 % tricin), whereas the differences disappeared in mature and senescing flag leaves. Predictions of changes in litter phenolic concentrations and their effects on decomposition rates under EC based on changes in green leaves need to be revised.     

Diurnal Changes in Photosynthesis,Sugars, and Nitrogen of Wheat and Mungbean Grown Under Elevated CO2 Concentration

The diurnal changes in leaf net photosynthetic rate (P _N) and sugar and nitrogen contents in wheat [Triticum aestivum (L.) cv. HD 2285] and mungbean [Vigna radiata (L.) Wilczek cv. PS 16] were analysed under ambient, AC [350±25 µmol mol^–1] and elevated, EC [600±50 µmol mol^–1] CO₂ concentrations. In both mungbean and wheat P _N of AC- and EC-grown plants compared at the same CO₂ concentration showed that P _N was higher under EC. However, increased P _N in EC-plants declined in the afternoon and approached P _N of AC-plants. Depression in P _N, however, was less in mungbean compared with the large depression in wheat. Greater down regulation of P _N in wheat was associated with the accumulation of large amount of sugars and low nitrogen content in wheat leaves. Mungbean leaves accumulated mostly starch under EC and the difference in N content in AC- and EC-plants was relatively less than in wheat.     

植物光合作用对CO2浓度增高的适应机制

长期在高浓度CO2环境下生长的植物往往会发生光合适应或下调,即在相同CO2浓度下的光合速率明显低于普通空气中生长的对照。虽然关于这种现象已经有许多研究报告和综述文章,但是它的机理还不很清楚。本文结合作者所在研究组的工作,概述了关于植物光合适应机理研究的新进展,提出除了呼吸作用增强和光合产物超常积累的可能作用以外,二磷酸核酮糖（RuBP）羧化限制和RuBP再生限制可能是导致植物光合适应的主要因素。     

Enhanced Water Use Efficiency in Dry Loess Grassland Species Grown at Elevated Air CO2 Concentration

Net CO2 assimilation rate (PN), stomatal conductance (gs), transpiration rate (E), and water use efficiency (WUE) in four perennial C3 species (grasses: Dactylis glomerata, Festuca rupicola, dicots: Filipendula vulgaris, Salvia nemorosa) grown for 231 d in open-top chambers at ambient (CA, 350 µmol mol-1) or elevated (CE, 700 µmol mol-1) CO2 concentrations were compared. When measured at CE, PN was significantly higher in CE plants of all four species than in the CA ones. The increase in PN was less prominent in the two grasses than in the two dicots. The E was significantly higher in the CE-grass F. rupicola and CE-dicot F. vulgaris than in the CA plants. There was no change in E owing to CE in the other grass and dicot. The gs in F. vulgaris and F. rupicola increased, while there was a decrease in D. glomerata and no change in S. nemorosa. WUE increased in all species grown in CE: four- to five-fold in the dicots and less than two-fold in the grasses. The increase in WUE was primarily due to an increase in PN and not to a decrease in E.     

Photosynthetic CO2 Fixation in the Second Leaf of Wheat Seedlings Grown at Different Conditions of Nitrogen Nutrition

The rates of photosynthetic ₂ assimilation were determined in fully expanded second leaves of 21-day-old wheat (Triticum aestivum L.) seedlings grown on media supplied with nitrate or ammonia and on a nitrogen-free medium (NO₃ ^–- or NH₄ ⁺-treatments and N-deficit treatment, respectively). The maximal quantum efficiency of photosynthesis was independent on conditions of nitrogen nutrition. When leaves were exposed to 0.03% ₂ and high-intensity light, the lowest photosynthetic rate was noted for N-deficit treatment and the highest rate was characteristic of NH₄ ⁺ treatment. The elevation of the ₂ concentration in the gas phase to 0.1% stimulated photosynthesis at high-intensity light in all treatments. The rate of ₂ uptake by the leaf of N-deficient seedlings increased with ₂ concentration to a larger extent than in other treatments and approached the ₂ uptake rate characteristic of the NO₃ ^– treatment. In plants grown on a nitrogen-free medium, the leaf accumulated lesser amounts of reduced nitrogen and higher amounts of starch, but the content of chloroplast protein corresponded to that of NO₃ ^– treatment. In the leaf of NH₄ ⁺-treated seedlings, the rate of ₂ assimilation was higher than in the leaf of NO₃ ^– treated plants, regardless of the composition of the gas mixture. The ammonium-type nutrition, as compared to the nitrate-type nutrition, elevated the amount of reduced nitrogen in the leaf and promoted accumulation of chlorophyll and protein, the chloroplast protein in particular.     

CO2倍增对不同氮水平下小麦幼苗根系及叶片NR活性的影响

以小麦品种'小偃22'幼苗为材料,采用开顶式气室和水培实验研究了不同供氮水平(2.5、5.0、10.0和 15.0 mmol·L-1)下小麦幼苗植株生长量、根系形态、有机碳分泌速率和硝酸还原酶(NR)活性对大气CO2浓度升高的响应.结果显示,大气CO2浓度倍增均增加了小麦幼苗各生长阶段根冠生物量以及根系长度、面积、有机碳分泌速率和叶片NR活性.随供氮水平的提高,各生长阶段幼苗根冠生物量、根长和面积以及叶片NR活性呈上升趋势,而有机碳分泌速率呈下降趋势;根冠比变化不同阶段表现不一致,一叶一心期呈下降趋势,二叶一心期和三叶一心期分别以15.0和10.0 mmol·L-1氮水平较高.研究表明,大气CO2浓度升高可促进小麦幼苗根系生长和有机碳分泌速率,提高其氮素同化能力;增加介质供氮有利于高CO2浓度条件下小麦幼苗根冠生长和氮素同化,提高根冠比,减少根系有机碳过度分泌引起的碳损耗.     

Modelling Net Photosynthetic Rate of Winter Wheat in Elevated Air CO2 Concentrations

Winter wheat plants were grown in open top chambers either at 365 µmol mol^–1 (AC) or at 700 µmol mol^–1 (EC) air CO₂ concentrations. The photosynthetic response of flag leaves at the beginning of flowering and on four vertical leaf levels at the beginning of grain filling were measured. Net photosynthetic rates (P _N) were higher at both developmental phases in plants grown at EC coupled with larger leaf area and photosynthetic pigment contents. The widely accepted Farquhar net photosynthesis model was parameterised and tested using several observed data. After parameterisation the test results corresponded satisfactorily with observed values under several environmental conditions.     

Photosynthesis and Plant Growth at Elevated Levels of CO2

In this review, we discuss the effects of elevated CO₂ levelson photosynthesis in relation to the whole plant growth in terrestrialhigher C₃ plants. Short-term CO₂ enrichment stimulates the rateof photosynthesis. Plant mass is also enhanced by CO₂ enrichment.However, the effects of long-term CO₂ enrichment on photosynthesisare variable. Generally, the prolonged exposure to CO₂ enrichmentreduces the initial stimulation of photosynthesis in many species,and frequently suppresses photosynthesis. These responses areattributed to secondary responses related to either excess carbohydrateaccumulation or decreased N content rather than direct responsesto CO₂. Accumulation of carbohydrates in leaves may lead tothe repression of photosynthetic gene expression and excessstarch seems to hinder CO₂ diffusion. Therefore, the specieswhich have the sink organs for carbohydrate accumulation donot show the suppression of photosynthesis. The suppressionof photosynthesis by CO₂ enrichment is always associated withdecreases in leaf N and Rubisco contents. These decreases arenot due to dilution of N caused by a relative increase in theplant mass but are the result of a decrease in N allocationto leaves at the level of the whole plant, and the decreasein Rubisco content is not selective. Leaf senescence and plantdevelopment are also accelerated by CO₂ enrichment. However,they are independent of each other in some species. Thus, variousresponses to CO₂ observed at the level of a single leaf resultfrom manifold responses at the level of the whole plant grownunder conditions of CO₂ enrichment. (Received July 8, 1999; Accepted August 12, 1999)     

Effect of Elevated CO2 Concentration on the Starch Grain Accumulation in Chloroplasts from Soybean Leaves at Different Nodes

Soybean ( Glycine max (L.) Merr. ) plants were grown under ambient and elevated CO2 (plus 350 μL/L) concentration in cylindrical open-top chamber to examine their effects on the ultra- structure of chloroplasts. The upper, lower and mid-node leaves were harvested after 7 days full expansion under different CO2 concentrations and ultrathin section were prepared for transmission electron microscopy. In general, the average content of starch grains and thylakoid membranes in the chloroplasts under the elevated CO2 concentration were always higher than the control. Under higher CO2 concentration, there were smaller and less starch grains in the chloroplasts from upper-node leaves than those from mid-node leaves. The shape of their starch grains changed from elliptical to oval,and their thylakoid membranes and grana remained normal. At lower-node leaves, one or two oval, or three timer starch grains accumulated in the chloroplasts. In the mid-node leaves,however, some chloroplasts under higher CO2 concentration had rather large tim elliptical starch grains which could consequently cause disruption of grana and stroama thylakoids in the chloroplasts, whereas in other chloroplasts, the thylakoid membranes and grana were not deformed as the starch grains were smaller and elliptical. On the other hand, under higher CO2 concentration, the stacking degree of thylakoid membranes and starch grains accumulation in the mid-node leaves were significantly higher than those in the lower-node leaves,and slightly higher than the upper-node leaves. These results, in agreement with the chlorophyll contents and photosynthetic rate which reported by other authors in the past, indicated that the ultrastmcture response of the chloroplasts from different leaf nodes of soybeen under elevated CO2 coneentration were different. The seed yield of soybean at different nodes was decreased gradually from mid-nodes towards both upper- and lower-nodes. The greatest effect of elevated CO2 eoneentrafion on seed yeild was at the mid-node leaves. The variation of seed yields of soybean at different nodes under elevated CO2 concentration was in eoneert with the change in the ultrastmcture of chloroplasts and in turn the change in their photosynthetic rates of leaves at different nodes.     

Photosynthetic Acclimation to Elevated CO2 in Relation to Leaf Saccharide Constituents in Wheat and Sunflower

Wheat (T. durum cvs. HD 4502 and B 449, T. aestivum cvs. Kalyansona and Kundan) and sunflower (Helianthus annuus L. cv. Morden) were grown under atmospheric (360±10 cm³ m^–3, AC) and elevated CO₂ (650±50 cm³ m^–3, EC) concentration in open top chambers for entire period of growth and development till maturity. Leaf net photosynthetic rate (P _N) of EC-grown plants of wheat measured at EC was significantly decreased in comparison with AC-plants of wheat measured at EC. Sunflower, however, showed no significant depression in P _N in EC-plants. There was a decrease in ribulose-1,5-bisphosphate carboxylase (RuBPC) activity, its activation state and amount in EC-plants of wheat, whereas no significant decrease was observed in sunflower. The above different acclimation to EC in wheat and sunflower was related with saccharide constituents accumulated in the leaves. Under EC, sunflower accumulated in the leaves more starch, whereas wheat accumulated more sugars.     

Effect of Elevated CO2 on Photosynthesis and Chlorophyll Fluorescence of Rose Plants Grown at High Temperature and High Photosynthetic Photon Flux Density

Gas exchange and chlorophyll (Chl) fluorescence were measured on young mature leaves of rose plants (Rosa hybrida cvs. First Red and Twingo) grown in two near-to-tight greenhouses, one under control ambient CO₂ concentration, AC (355 µmol mol^–1) and one under CO₂ enrichment, EC (700 µmol mol^–1), during four flushes from late June to early November. Supply of water and mineral elements was non-limiting while temperature was allowed to rise freely during daytime. Leaf diffusive conductance was not significantly reduced at EC but net photosynthetic rate increased by more than 100 %. Although the concentration of total non-structural saccharides was substantially higher in the leaves from the greenhouse with EC, _PS2 (quantum efficiency of radiation use) around noon was not significantly reduced at EC indicating that there was no down-regulation of electron transport. Moreover, CO₂ enrichment did not cause any increase in the risk of photo-damage, as estimated by the 1 – q_P parameter. Non-photochemical quenching was even higher in the greenhouse with EC during the two summer flushes, when temperature and photosynthetic photon flux density (PPFD) were the highest. Hence rose photosynthesis benefits strongly from high concentrations of atmospheric CO₂ at both high and moderate temperatures and PPFD.     

Water Relations and Leaf Anatomy of the Tropical Species,Jatropha gossypifolia and Alternanthera crucis,Grown Under an Elevated CO2 Concentration

In order to address the question of how elevated CO₂ concentration (EC) will affect the water relations and leaf anatomy of tropical species, plants of Jatropha gossypifolia L. and Alternanthera crucis (Moq.) Bondingh were grown in five EC open top chambers (677 mol mol^–1) and five ambient CO₂ concentration (AC) open top chambers (454 mol mol^–1) with seasonal drought. No effect of EC was found on morning xylem water potential, leaf osmotic potential, and pressure potential of plants of J. gossypifolia. In A. crucis EC caused a significant increase in morning xylem water potential of watered plants, a decrease in osmotic potential, and an increase of 24–79 % in pressure potential of moderately droughted plants. This ameliorated the effects of drought. Stomatal characteristics of both leaf surfaces of J. gossypifolia and A. crucis showed time-dependent, but not [CO₂]-dependent changes. In J. gossypifolia the thickness of whole leaf, palisade parenchyma, and spongy parenchyma, and the proportion of whole leaf thickness contributed by these parenchymata decreased significantly in response to EC. In A. crucis EC caused an increase in thickness of whole leaf, bundle sheath, and mesophyll, while the proportion of leaf cross-section comprised by the parenchymata remained unchanged. These effects disappeared with time under treatment, suggesting that acclimation of the leaf anatomy to the chambers and to EC took place in the successive flushes of leaves produced during the experiment.     

CO2浓度升高对两种沈阳城市森林树种光合特性的影响

利用开顶式气室,研究了CO2浓度升高条件下城市森林主要树种油松(Pinus tabulaefomis)和银杏(Ginkgo biloba)主要光合特性的变化.结果表明,整个生长季,CO2浓度升高(700μmol·mol-1)条件下2树种叶片的净光合速率、可溶性糖、淀粉和可溶性蛋白含量均接近或高于相应对照(自然CO2浓度)值,但不同树种增加的幅度不同;而2树种的叶绿素含量和Chl a/Chl b值对CO2浓度升高反应不一,表现为CO2浓度升高条件下油松的叶绿素含量较对照值高,Chl a/Chl b值降低,银杏的叶绿素含量为前期升高,后期降低,Chl a/Chl b值变化与之正好相反,说明城市森林组成树种对CO2浓度升高的响应具有复杂性.CO2浓度升高条件下,两树种均未发生光合适应现象.     

Interactive Effects of Elevated CO2 Concentration and Irrigation on Photosynthetic Parameters and Yield of Maize in Northeast China

Maize is one of the major cultivated crops of China, having a central role in ensuring the food security of the country. There has been a significant increase in studies of maize under interactive effects of elevated CO₂ concentration ([CO₂]) and other factors, yet the interactive effects of elevated [CO₂] and increasing precipitation on maize has remained unclear. In this study, a manipulative experiment in Jinzhou, Liaoning province, Northeast China was performed so as to obtain reliable results concerning the later effects. The Open Top Chambers (OTCs) experiment was designed to control contrasting [CO₂] i.e., 390, 450 and 550 µmol·mol⁻¹, and the experiment with 15% increasing precipitation levels was also set based on the average monthly precipitation of 5–9 month from 1981 to 2010 and controlled by irrigation. Thus, six treatments, i.e. C₅₅₀W_+15%, C₅₅₀W₀, C₄₅₀W_+15%, C₄₅₀W₀, C₃₉₀W_+15% and C₃₉₀W₀ were included in this study. The results showed that the irrigation under elevated [CO₂] levels increased the leaf net photosynthetic rate (P _n) and intercellular CO₂ concentration (C _i) of maize. Similarly, the stomatal conductance (G _s) and transpiration rate (T _r) decreased with elevated [CO₂], but irrigation have a positive effect on increased of them at each [CO₂] level, resulting in the water use efficiency (WUE) higher in natural precipitation treatment than irrigation treatment at elevated [CO₂] levels. Irradiance-response parameters, e.g., maximum net photosynthetic rate (P _nmax) and light saturation points (LSP) were increased under elevated [CO₂] and irrigation, and dark respiration (R _d) was increased as well. The growth characteristics, e.g., plant height, leaf area and aboveground biomass were enhanced, resulting in an improved of yield and ear characteristics except axle diameter. The study concluded by reporting that, future elevated [CO₂] may favor to maize when coupled with increasing amount of precipitation in Northeast China.     

Effects of Elevated Atmospheric CO2 Concentration on Leaf Anatomy and Morphology in Panicum Species Representing Different Photosynthetic Modes

Panicum tricanthum Nees, Panicum antidotale Retz., and Panicum decipiens Nees ex Trin. were selected to represent C3, C4, and C3/C4 intermediate perennial species of Panicum, respectively. Plants grown from seed with 900 ppm [CO2] under natural sunlight and controlled temperatures (30 degrees /22 degrees C) were compared with plants grown with ambient [CO2]. The anatomy of the last fully expanded leaf of the main tiller was studied by light microscopy with computerized graphic image analysis and by transmission electron microscopy. Leaf anatomy did not change qualitatively in response to elevated [CO2], but there were changes in leaf thickness and in the proportions of total transsectional area occupied by mesophyll, bundle sheath cells, vascular elements, and sclerenchyma, according to species. The abaxial stomatal frequency decreased by 22% for P. tricanthum but increased by ca. 30% for the other two species. With 900 ppm CO2, all three species showed a considerable increase in leaf starch content (to >30% of dry matter). Starch granules accumulated in chloroplasts of the mesophyll and bundle sheath cells. Increased leaf glaucousness in response to elevated [CO2] was the result of increased or modified deposition of epicuticular wax on both leaf surfaces, a response to elevated [CO2] that is unusual and one that has not been previously recorded for monocotyledons. The wax patterns were studied by scanning electron microscopy. Panicum decipiens did not respond to elevated [CO2] in a truly intermediate fashion; its responses resembled those of either the C3 or the C4 species. C3/C4 intermediates may thus be interpreted as developmental chimeras and not as species in transition between C3 and C4 modes in an evolutionary sense.