Increase of photosynthesis and starch in potato under elevated CO2 is dependent on leaf age

Potato plants (Solanum tuberosum cv. Bintje) were grown in open top chambers under ambient (400 microL L(-1)) and elevated CO2 (720 microL L(-1)). After 50 days one half of each group was transferred to the other CO2 concentration and the effects were studied in relation to leaf age (old, middle-aged and young leaves) in each of the four groups. Under long-term exposure to elevated CO2, photosynthesis increased between 10% and 40% compared to ambient CO2. A subsequent shift of the same plants to ambient CO2 caused a 20-40% decline in photosynthetic rate, which was most pronounced in young leaves. After shifting from long-term ambient to elevated CO2, photosynthesis also increased most strongly in young leaves (90%); these experiments show that photosynthesis was downregulated in the upper young fully expanded leaves of potato growing long-term under elevated CO2. Soluble sugar content in all leaf classes under long-term exposure was stable irrespective of the CO2 treatment, however under elevated CO2 young leaves showed a strongly increased starch accumulation (up to 400%). In all leaf classes starch levels dropped in response to the shift from 720 to 400 microL L(-1) approaching ambient CO2 levels. After the shift to 720 microL L(-1), sucrose and starch levels increased, principally in young Leaves. There is clear evidence that leaves of different age vary in their responses to changes in atmospheric CO2 concentration.     

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。     

Growth under elevated atmospheric CO(2) concentration accelerates leaf senescence in sunflower (Helianthus annuus L.) plants

Some morphogenetic and metabolic processes were sensitive to a high atmospheric CO(2) concentration during sunflower primary leaf ontogeny. Young leaves of sunflower plants growing under elevated CO(2) concentration exhibited increased growth, as reflected by the high specific leaf mass referred to as dry weight in young leaves (16days). The content of photosynthetic pigments decreased with leaf development, especially in plants grown under elevated CO(2) concentrations, suggesting that high CO(2) accelerates chlorophyll degradation, and also possibly leaf senescence. Elevated CO(2) concentration increased the oxidative stress in sunflower plants by increasing H(2)O(2) levels and decreasing activity of antioxidant enzymes such as catalase and ascorbate peroxidase. The loss of plant defenses probably increases the concentration of reactive oxygen species in the chloroplast, decreasing the photosynthetic pigment content as a result. Elevated CO(2) concentration was found to boost photosynthetic CO(2) fixation, especially in young leaves. High CO(2) also increased the starch and soluble sugar contents (glucose and fructose) and the C/N ratio during sunflower primary leaf development. At the beginning of senescence, we observed a strong increase in the hexoses to sucrose ratio that was especially marked at high CO(2) concentration. These results indicate that elevated CO(2) concentration could promote leaf senescence in sunflower plants by affecting the soluble sugar levels, the C/N ratio and the oxidative status during leaf ontogeny. It is likely that systemic signals produced in plants grown with elevated CO(2), lead to early senescence and a higher oxidation state of the cells of these plant leaves.     

Elevated CO2, rhizosphere processes,and soil organic matter decomposition

The rhizosphere is one of the key fine-scale components of C cycles. This study was undertaken to improve understanding of the potential effects of atmospheric CO2 increase on rhizosphere processes. Using C isotope techniques, we found that elevated atmospheric CO2 significantly increased wheat plant growth, dry mass accumulation, rhizosphere respiration, and soluble C concentrations in the rhizosphere. When plants were grown under elevated CO2 concentration, soluble C concentration in the rhizosphere increased by approximately 60%. The degree of elevated CO2 enhancement on rhizosphere respiration was much higher than on root biomass. Averaged between the two nitrogen treatments and compared with the ambient CO2 treatment, wheat rhizosphere respiration rate increased 60% and root biomass only increased 26% under the elevated CO2 treatment. These results indicated that elevated atmospheric CO2 in a wheat-soil system significantly increased substrate input to the rhizosphere due to both increased root growth and increased root activities per unit of roots. Nitrogen treatments changed the effect of elevated CO2 on soil organic matter decomposition. Elevated CO2 increased soil organic matter decomposition (22%) in the nitrogen-added treatment but decreased soil organic matter decomposition (18%) without nitrogen addition. Soil nitrogen status was therefore found to be important in determining the directions of the effect of elevated CO2 on soil organic matter decomposition.     

Assimilation,respiration and allocation of carbon inPlantago major as affected by atmospheric CO2 levels

The response ofPlantago major ssp,pleiosperma plants, grown on nutrient solution in a climate chamber, to a doubling of the ambient atmospheric CO₂ concentration was investigated. Total dry matter production was increased by 30% after 3 weeks of exposure, due to a transient stimulation of the relative growth rate (RGR) during the first 10 days. Thereafter RGR returned to the level of control plants. Photosynthesis, expressed per unit leaf area, was stimulated during the first two weeks of the experiment, thereafter it dropped and nearly reached the level of the control plants. Root respiration was not affected by increased atmospheric CO₂ levels, whereas shoot, dark respiration was stimulated throughout the experimental period. Dry matter allocation over leaves stems and roots was not affected by the CO₂ level. SLA was reduced by 10%, which can partly be explained by an increased dry matter content of the leaves. Both in the early and later stages of the experiment, shoot respiration accounted for a larger part of the carbon budget in plants grown at elevated atmospheric CO₂. Shifts in the total carbon budget were mainly due to the effects on shoot respiration. Leaf growth accounted for nearly 50% of the C budget at all stages of the experiment and in both treatments.Abbreviations LAR leaf area ratio - LWR leaf weight ratio - RGR relative growth rate - R/S root to shoot ratio - RWR root weight ratio - SLA specific leaf area - SWR stem weight ratio     

Elevated CO(2) induces biochemical and ultrastructural changes in leaves of the C(4) cereal sorghum

We analyzed the impact of growth at either 350 (ambient) or 700 (elevated) microL L(-1) CO(2) on key elements of the C(4) pathway (photosynthesis, carbon isotope discrimination, and leaf anatomy) using the C(4) cereal sorghum (Sorghum bicolor L. Moench.). Gas-exchange analysis of the CO(2) response of photosynthesis indicated that both carboxylation efficiency and the CO(2) saturated rate of photosynthesis were lower in plants grown at elevated relative to ambient CO(2). This was accompanied by a 49% reduction in the phosphoenolpyruvate carboxylase content of leaves (area basis) in the elevated CO(2)-grown plants, but no change in Rubisco content. Despite the lower phosphoenolpyruvate carboxylase content, there was a 3-fold increase in C isotope discrimination in leaves of plants grown at elevated CO(2) and bundle sheath leakiness was estimated to be 24% and 33%, respectively, for the ambient and elevated CO(2)-grown plants. However, we could detect no difference in quantum yield. The ratio of quantum yield of CO(2) fixation to PSII efficiency was lower in plants grown at elevated CO(2), but only when leaf internal was below 50 microL L(-1). This suggests a reduction in the efficiency of the C(4) cycle when [CO(2)] is low, and also implies increased electron transport to acceptors other than CO(2). Analysis of leaf sections using a transmission electron microscope indicated a 2-fold decrease in the thickness of the bundle sheath cell walls in plants grown at elevated relative to ambient CO(2). These results suggest that significant acclimation to increased CO(2) concentrations occurs in sorghum.     

Nitrogen and carbon partitioning in soybean under variable nitrogen supplies and acclimation to the prolonged action of elevated CO2

This study was conducted to determine reciprocal effects of low to high doses of nitrogenous fertilizer (N₃₀, N₄₀, N₅₀, N₆₀ and N₇₀ — 30, 40, 50, 60 and 70 kg ha⁻¹ respectively) and CO₂ enriched environment on C and N partitioning in soybean (Glycine max (L.) Merril cv JS-335). Plants were grown from seedling emergence to maturity inside open top chambers under ambient, AC (350±50 mol mol⁻¹) and elevated, EC (600±50 mol mol⁻¹) CO₂ and analyzed at seedling, vegetative, flowering, pod setting and maturity stages. Soybean responded to both CO₂ enrichment and N supply. Leaves, stem and root reserves at different growth stages were analyzed for total C and N contents. Consistent increase in the C contents of the leaf, stem and root was observed under EC than in AC. N contents in the different plant parts were found to be decreased under EC-grown plants specially at seedling and vegetative stage despite providing N doses to the soil. Significant increase observed for C to N dry mass ratio under EC in the root, stems and leaves at seedling and vegetative stage was decreased in the middle and later growth stages possibly due to combined impact of N doses to the soil and increased N₂ fixing activities due to EC conditions. Critical analysis of our findings reveals that the composition and partitioning of C and N of soybean under variable rates of N supply and CO₂ enrichment alter according to need under altered metabolic process. These changes eventually may lead to alteration in uptake of not only N but other essential nutrients also under changing atmosphere.     

Responses of legume versus nonlegume tropical tree seedlings to elevated CO2 concentration

We investigated responses of growth, leaf gas exchange, carbon-isotope discrimination, and whole-plant water-use efficiency (W(P)) to elevated CO(2) concentration ([CO(2)]) in seedlings of five leguminous and five nonleguminous tropical tree species. Plants were grown at CO(2) partial pressures of 40 and 70 Pa. As a group, legumes did not differ from nonlegumes in growth response to elevated [CO(2)]. The mean ratio of final plant dry mass at elevated to ambient [CO(2)] (M(E)/M(A)) was 1.32 and 1.24 for legumes and nonlegumes, respectively. However, there was large variation in M(E)/M(A) among legume species (0.92-2.35), whereas nonlegumes varied much less (1.21-1.29). Variation among legume species in M(E)/M(A) was closely correlated with their capacity for nodule formation, as expressed by nodule mass ratio, the dry mass of nodules for a given plant dry mass. W(P) increased markedly in response to elevated [CO(2)] in all species. The ratio of intercellular to ambient CO(2) partial pressures during photosynthesis remained approximately constant at ambient and elevated [CO(2)], as did carbon isotope discrimination, suggesting that W(P) should increase proportionally for a given increase in atmospheric [CO(2)]. These results suggest that tree legumes with a strong capacity for nodule formation could have a competitive advantage in tropical forests as atmospheric [CO(2)] rises and that the water-use efficiency of tropical tree species will increase under elevated [CO(2)].     

Leaf and canopy photosynthetic characteristics of cotton (Gossypium hirsutum) under elevated CO2 concentration and UV-B radiation

Increases in both atmospheric CO2 concentration ([CO2]) and ultraviolet-B (UV-B) radiation on the Earth's surface are features of current climate change patterns. An experiment was conducted in sunlit, controlled environment chambers known as Soil-Plant-Atmosphere-Research (SPAR) units to determine interactive effects of elevated [CO2] and UV-B radiation on leaf and canopy photosynthetic characteristics of cotton. Six treatments were comprised of two CO2 levels of 360 (ambient) and 720 (elevated) microL L(-1) and three levels of 0 (control), 8, and 16 kJ m(-2) d(-1) biologically effective UV-B radiation. Treatments were imposed for 66 days from crop emergence through three weeks after the first flower stage. Plants grown in elevated [CO2] had significantly greater leaf area, higher leaf and canopy net photosynthetic rates (PN), lower dark respiration rate (Rd), and lower light compensation point (LCP) than plants grown in ambient [CO2]. There was no difference in CO2 compensation point (gamma), maximum rate of Rubisco activity (Vcmax), or light-saturated rate of electron transport (Jmax) between ambient and elevated CO2 treatments. When plants were grown in 8 kJ m(-2) d(-1) UV-B radiation, most of the measured photosynthetic parameters did not differ from control plants. High UV-B (16 kJ) radiation, however, caused 47-50% smaller leaf area, 38-44% lower leaf PN, 72-74% lower Vcmax, and 61-66% lower Jmax compared to the control. There were no interactive effects of [CO2] and UV-B radiation on most of the photosynthetic parameters measured. From the results, it is concluded that decreased canopy photosynthesis due to enhanced UV-B radiation in cotton is associated with both smaller leaf area and lower leaf PN, and loss of Rubisco activity and electron transport are two major factors in UV-B inhibition of leaf PN.     

Effects of elevated CO2 on intra-specific competition in Sinapis alba: an examination of the role of growth responses to red:far-red ratio

Response of plants to elevated CO2 differs markedly between individually- and competitively-grown plants, both in terms of mean size and variation about the mean. Using Sinapis alba, we explored whether these contrasting effects are a consequence of the effect of competition on the red:far red (R:FR) light ratio. Plants were grown at both ambient and elevated (700 microl.l(-1)) CO2 in competitive stands, and as individuals at either a low (0.7) or high (1.25) R:FR ratio at a constant photosynthetic photon fluence rate. Elevated CO2 increased stand biomass by enhancing the growth of canopy dominants, but not the subordinates. As a consequence, elevated CO2 increased the coefficient of variation in size within the stands. Elevated CO2 did not enhance the growth of individually-grown plants at the low R:FR ratio, but did at the high R:FR ratio. Both the poor response of subordinate plants to elevated CO2 and the increased size inequalities of individuals within the stand can be explained in terms of the effect of the R:FR ratio on CO2 responsiveness. The effect of the R:FR ratio on CO2 response may be related to its effect on allocation patterns and nutrient uptake.     

不同生长阶段遮荫对番茄光合作用、干物质分配与叶N、P、K的影响

对番茄植株做了两种不同程度的遮荫处理,观测了夏季午间遮荫对光合速率,干物质积累量及其在根,茎,叶之间的分配,和叶N,P,K的含量以及经济产量的影响,发现不同时期遮荫影响不同。（1）遮荫增加三个阶段（开花早期,盛花期和开花后期）的气孔导度和胞间CO2浓度,显著降低开花早期中午的净化合速率,但盛花期中度遮荫（40%遮荫）使净光合速率随着时间的增加逐渐上升,在开花后期表现更加明显,平均净光合速率比对照高20%以上,蒸腾速率也增加较多。（2）开花早期和盛花期重度遮荫（如本实验中的75%遮荫）显著降低根,茎的干重,而开花后期中度遮荫的根,茎干重高于对照,但遮荫对叶干重的影响不明显。（3）开花早期和盛花期遮荫不明显影响叶片中N,P,K的含量,但开花后期中度遮荫使N,P,K含量增加,（4）开花早期两种遮荫对果实产量影响较小,但盛花期重度遮荫使产量降低,全部产量中无效部分所占的比例上升,开花后期中度遮荫的总产量和有效产量增加,单果重也增加,这些结果表明,在某些时期中度遮荫可以克服夏天辐射过强,气温过高对番茄的不良影响,对番茄生长,干物质积累和提高产量等有利,在生产上有意义。     

植物对开放式CO2 浓度增高(FACE)的响应与适应研究进展

开放式CO2浓度增高(FACE)系统是近年研究植物对高CO2浓度响应和适应的新手段,它比以往密闭和半密闭系统对实验植物生长环境的干扰少.利用FACE系统进行研究更有助于正确地预测未来大气CO2浓度增高对植物的影响.该文结合作者的研究工作简要评介了FACE系统与以往密闭和半密闭式CO2浓度增高实验系统的不同之处以及近年来利用FACE系统所作的最新研究进展.     

Changes in Nitrogen Metabolism of Vigna Radiata in Response to Elevated CO2

With the aim to determine the effects of CO₂ on nitrogen metabolism mungbean (Vigna radiata) plants were grown from seedling emergence to maturity inside open top chambers under ambient CO₂ (CA, 350 ± 25 µmol mol^–1) and elevated CO₂ (CE, 600 ± 50 µmol mol^–1) concentrations at the Indian Agricultural Research Institute, New Delhi. Leaflet blades of the same physiological age were sampled at 20, 35 and 50 d after germination. Total nitrogen concentration in dry mass was consistently lower under CE than in CA. Non-protein nitrogen and protein nitrogen were also decreased under CE Total soluble protein content also decreased up to 35 d after germination under CE. However, a 27 % increase in protein content at 50 d after germination due to CE was observed. A significant decrease in total free amino acid under CE at 20 d after germination was observed. CE also brought about a remarkable decrease in the activity of nitrate reductase in leaves at 20 d after germination but increase at 35 d and 50 d after germination. Nitrogenase activity increased at all growth stages due to CE. Although total harvested leaves of CE plants accumulated more nitrogen, the relative amount of nitrogen on a percentage basis was low, probably due to a comparatively greater accumulation of sugars in the leaves of CE plants.     

不同氮源下生长的柚树叶片光合参数对高浓度CO2驯化作用的比较

比较研究了在不同形式氮源下生长柚树叶片光合对高浓度 CO2 驯化过程中有关参数变化。植株生长在人工混成土壤中 ,分别浇灌含有 2 mmol L- 1N的 NO- 3 - N,NH+ 4 - N和 NH4 NO3- N溶液。空气 CO2 增高处理时向生长植株的开顶透明罩中通入 74.4Pa CO2 ,以空气 CO2 生长的植株为对照。利用 CI- 30 1 ( CID,Inc) CO2 气体交换系统测定叶片光合速率和通过光合作用相关响应曲线计算光合参数。结果表明 ,在 CO2分压倍增下 ,NO- 3 - N生长植株光饱和光合速率较大气 CO2 分压下的高。而生长在 NH+ 4 - N和 NH4 NO3- N的植株光合速率与大气 CO2 分压下的相近 ,表现对高 CO2 的驯化。在空气 CO2 倍增下无论供给何种形式氮源并不影响Γ* ,但可增高 Rd( P<0 .0 5 )。 CO2 分压倍增下供给 NO- 3 - N植株的 Vcmax和 Jmax较大气分压相应的植株高 ,而 NH+ 4 - N和 NH4 NO3- N植株则与大气 CO2分压的相应植株相似 ( P>0 .0 5 )。无论供给何种形式氮源 ,生长在空气 CO2 分压倍增下不改变叶片单位面积干重 ,叶绿素含量和叶片中氮在 Rubisco、生物能学组分和捕光色素复合体组分的分配系数 ;但能改变叶片中氮含量。植物对高 CO2 的驯化可能受到不同形式氮利用性的影响 ,在对高 CO2 驯化过程亦反映叶片中氮在不同光合功能组分     

开放式空气CO2增高对水稻物质生产与分配的影响

在大田栽培条件下,研究开放式空气CO2增加(FACE)200μmol·mol^-1的处理对水稻物质生产与分配的影响．结果表明,FACE处理使移栽至抽穗后20d的干物质积累量显著增加,使抽穗后20d至成熟期的干物质生产量显著减少,生物产量显著提高．移栽至抽穗期的干物质积累量增加是由于叶面积系数和净同化率共同提高所致;抽穗期至抽穗后20d的干物质积累量增加主要是由于叶面积系数的增加所致;抽穗后20d至成熟期的干物质生产量减少主要是由于净同化率的下降所造成．提高茎鞘占全株干物重的比例,降低叶片占全株干物重的比例,对穗占全株干物重的比例无显著影响,能显著提高水稻抽穗期茎鞘中可溶性糖、淀粉的含有率和含量,提高FACE处理的生物产量能极显著提高水稻产量(r＝0．7825)．     

Response of nodulated alfalfa to water supply,temperature and elevated CO2: productivity and water relations

Exposing plants to long-term CO₂ enrichment generally leads to increases in plant biomass, total leaf area and alterations on leaf net photosynthetic rates, stomatal conductance and water use efficiency. However, the magnitude of such effects is dependent on the availability of other potentially limiting resources. The aim of our study was to elucidate the effects of elevated CO₂, applied at different temperature and water availability regimes, on nodulated alfalfa plants. Regardless of water supply, elevated CO₂ enhanced plant growth, especially when combined with increased temperature although no differences were detected until 30 days of treatment. Absence of differences in leaf relative growth rate, and gas exchange measurements, suggested that plants grown in a low water regime adjusted their growth to the amount of available water. Elevated CO₂ enhanced water use efficiency because of reduced water consumption and a greater dry mass production. Increased dry matter production of plants grown under elevated CO₂ and temperature was the result of stimulated photosynthetic rates, greater leaf area and water use efficiency. Lack of CO₂ effect on photosynthesis of plants grown at ambient temperature might be consequence of down-regulation phenomena. Plants grown at 700 μmol mol⁻¹ CO₂ maintained control nitrogen levels, discarding enhanced nitrogen availability as the main factor explaining enhanced dry matter.     

Future CO2 concentrations, though not warmer temperatures, enhance wheat photosynthesis temperature responses

The temperature dependence of C3 photosynthesis is known to vary according to the growth environment. Atmospheric CO2 concentration and temperature are predicted to increase with climate change. To test whether long-term growth in elevated CO2 and temperature modifies photosynthesis temperature response, wheat (Triticum aestivum L.) was grown in ambient CO2 (370 micromol mol(-1)) and elevated CO2 (700 micromol mol(-1)) combined with ambient temperatures and 4 degrees C warmer ones, using temperature gradient chambers in the field. Flag leaf photosynthesis was measured at temperatures ranging from 20 to 35 degrees C and varying CO2 concentrations between ear emergence and anthesis. The maximum rate of carboxylation was determined in vitro in the first year of the experiment and from the photosynthesis-intercellular CO2 response in the second year. With measurement CO2 concentrations of 330 micromol mol(-1) or lower, growth temperature had no effect on flag leaf photosynthesis in plants grown in ambient CO2, while it increased photosynthesis in elevated growth CO2. However, warmer growth temperatures did not modify the response of photosynthesis to measurement temperatures from 20 to 35 degrees C. A central finding of this study was that the increase with temperature in photosynthesis and the photosynthesis temperature optimum were significantly higher in plants grown in elevated rather than ambient CO2. In association with this, growth in elevated CO2 increased the temperature response (activation energy) of the maximum rate of carboxylation. The results provide field evidence that growth under CO2 enrichment enhances the response of Rubisco activity to temperature in wheat.