CO2浓度升高和降水增加协同作用对玉米产量及生长发育的影响

关于[CO₂]升高和降水变化等多因子共同作用对植物的影响报道较少, 制约着人们对植物对全球气候变化响应的认识和预测。玉米(Zea mays)作为重要的C₄植物, 受[CO₂]和降水影响显著, 但鲜有[CO₂]升高和降水增加协同作用对其产量及生长发育影响的报道。该研究利用开顶式生长箱模拟[CO₂]升高(390 (环境)、450和550 μmol·mol^-1), 降水增加量设置为增加自然降水量的15% (以试验地锦州1981-2010年6至8月月平均降水量为基准), 从而形成6个处理: C₅₅₀W_+15%、C₅₅₀W₀、C₄₅₀W_+15%、C₄₅₀W₀、C₃₉₀W_+15%和C₃₉₀W₀。试验材料选用玉米品种‘丹玉39’。结果表明: [CO₂]升高和降水增加的协同作用在玉米的籽粒产量和生物产量上均达到了显著水平(p< 0.05), 二因子均起正作用, 使籽粒产量和生物产量均升高。籽粒产量在[CO₂] 390、450和550 μmol·mol^-1水平下的降水增加处理较自然降水处理分别增加15.94%、9.95%和9.45%, 而生物产量分别增加13.06%、8.13%和6.49%。因为籽粒产量的增幅略大于生物产量的增幅, 所以促进了经济系数的升高。穗部性状变化显著, 其中, 穗粒数、穗粒重、穗长和穗粗等性状值均随[CO₂]升高而升高, 且各[CO₂]水平下均表现为降水增加处理>自然降水处理, 而瘪粒数相反。但是, [CO₂]升高和降水增加的协同作用也促进了轴粗的升高, 对玉米产量的增加起着限制作用。二因子协同作用在净光合速率(P_n)和叶面积上达到了极显著水平(p< 0.01), 而在株高和干物质积累量上达到了显著水平(p< 0.05)。二因子协同作用使玉米叶片的P_n升高, 植株高度升高, 穗位高升高, 茎粗增加, 叶面积变大, 从而促进了干物质积累量的升高, 为玉米增产打下了良好的基础。这表明: 在未来[CO₂]升高条件下, 一定程度的降水增加对玉米的产量具有正向促进作用。     

Effect of elevated CO2 concentration on seedling growth rate and photosynthesis inHevea brasiliensis

To study the effect of elevated CO₂ concentration on plant growth and photosynthesis, two clones ofHevea brasiliensis were grown in polybags and exposed to elevated concentration (700±25ppm) for 60 days. There was higher biomass accumulation, leaf area and better growth when compared to ambient air grown plantso From A/Ci curves it is clear that photosynthetic rates increases with increase in CO₂ concentrations. After 60 days of exposure to higher CO₂ concentration, a decrease in the carbon assimilation rate was noticed.     

Cladode development, environmental responses of CO2 uptake, and productivity for Opuntia ficus-indica under elevated CO2

Opuntia ficus-indica, an extremely productive CAM plant cultivatedin many countries, was exposed to 36, 52, and 72–73 PaCO₂ in field plots and open-top chambers. Initiation of newcladodes (stem segments) was monitored until the canopy closed,after which bimonthly harvests maintained the plants for oneyear at a cladode area per unit ground area that is optimalfor biomass production. Doubling the CO₂ partial pressure slightlyincreased the number of first-order daughter cladodes growingon the basal (planted) cladodes after 3 months and nearly doubledthe number and area of second-order cladodes. When the C02 levelwas doubled, cladodes were 5% thicker after a few months and11 to 16% thicker after one year. Although the productivityenhancement by elevated C02 tended to decrease during the year,the annual above-ground dry-mass gain was 37 to 40% higher whenthe C02 level was doubled, reaching 65 tons hectare^–1year^–1 in a field plot. Well-watered cladodes at day/nightair temperatures of 25°C/15°C and a total daily photosyntheticphoton flux (PPF) of 15 mol m^–2 d^–1 in controlledenvironment chambers had 74% more net CO₂ uptake over 24 h at73 Pa than at 37 Pa CO₂. With doubled CO₂, the percentage enhancementof net CO₂ uptake increased as the PPF was lowered, as the temperaturewas raised, and during drought. Using an environmental productivityindex based on such factors, net CO₂ uptake and hence productivityof O. ficus-indica can be predicted for elevated CO₂ levelsand other variations accompanying global climate change. Key words: Crassulacean acid metabolism, environmental productivity index, gas exchange, global climate change, plant growth     

氮素对高大气CO2浓度下小麦叶片光合作用的影响

通过测定小麦拔节期叶片的光合气体交换参数和光强-光合速率（P_n）响应曲线,研究了氮素对长期高大气CO₂浓度（760 μmol·mol^-1）下小麦叶片光合作用的影响.结果表明：在长期高大气CO₂浓度下,增施氮肥能提高小麦叶片P_n、蒸腾速率（T_r）和瞬时水分利用效率（WUE_i）;与正常大气CO₂浓度相比,高大气CO₂浓度下小麦叶片的P_n和WUE_i增加,气孔导度（G_s）和胞间CO₂浓度(C_i）降低.随光合有效辐射的增强,高大气CO₂浓度下小麦叶片的P_n和WUE_i均高于正常大气CO₂浓度处理,G_s则较低,而C_i和T_r无显著变化.高氮水平下小麦叶片G_s与P_n、T_r、WUE_i呈线性正相关,G_s与C_i在正常大气CO₂浓度下呈线性负相关,但高大气CO₂浓度下二者无相关性;低氮水平下小麦叶片的G_s与P_n、WUE_i无相关性,而与C_i和T_r呈线性正相关,表明高大气CO₂浓度下低氮水平的小麦叶片P_n由非气孔因素限制.     

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.     

Decreased photosynthesis and growth with reduced respiration in the model diatom Phaeodactylum tricornutum grown under elevated CO2 over 1800 generations

Studies on the long‐term responses of marine phytoplankton to ongoing ocean acidification (OA) are appearing rapidly in the literature. However, only a few of these have investigated diatoms, which is disproportionate to their contribution to global primary production. Here we show that a population of the model diatom Phaeodactylum tricornutum, after growing under elevated CO₂ (1000 μatm, HCL, pH_T: 7.70) for 1860 generations, showed significant differences in photosynthesis and growth from a population maintained in ambient CO₂ and then transferred to elevated CO₂ for 20 generations (HC). The HCL population had lower mitochondrial respiration, than did the control population maintained in ambient CO₂ (400 μatm, LCL, pH_T: 8.02) for 1860 generations. Although the cells had higher respiratory carbon loss within 20 generations under the elevated CO₂, being consistent to previous findings, they downregulated their respiration to sustain their growth in longer duration under the OA condition. Responses of phytoplankton to OA may depend on the timescale for which they are exposed due to fluctuations in physiological traits over time. This study provides the first evidence that populations of the model species, P. tricornutum, differ phenotypically from each other after having been grown for differing spans of time under OA conditions, suggesting that long‐term changes should be measured to understand responses of primary producers to OA, especially in waters with diatom‐dominated phytoplankton assemblages.     

Effects of elevated CO2 on growth,photosynthesis and respiration of sweet chestnut (Castanea sativa Mill.)

Two year old sweet chestnut seedlings (Castanea sativa Mill) were grown in pots at ambient (350 µmol·mol^–1) and double (700 µmol·mol^–1) atmospheric CO₂ concentration in constantly ventilated greenhouses during entire growing seasons. CO₂ enrichment caused either no significant change or a decrease in shoot response, depending on yearly weather conditions. Similarly, leaf area was either reduced or unchanged under elevated CO₂. However, when grown under controlled conditions in a growth chamber, leaf area was enlarged with elevated CO₂.The CO₂ exchanges of whole plants were measured during the growing season. In elevated CO₂, net photosynthetic rate was maximum in May and then decreased, reaching the level of the control at the end of the season. End of night dark respiration of enriched plants was significantly lower than that of control plants; this difference decreased with time and became negligible in the fall. The original CO₂ level acted instantaneously on the respiration rate: a double concentration in CO₂ decreased the respiration of control plants and a reduced concentration enhanced the respiration of enriched plants. The carbon balance of a chestnut seedling may then be modified in elevated CO₂ by increased carbon inputs and decreased carbon outputs.     

Temperature dependence of photosynthesis in cotton

Cotton plants (Gossypium hirsutum L., var. Deltapine Smooth Leaf) were grown under controlled environmental conditions over a range of day/night temperatures from 20/15 to 40/35 C. Their photosynthetic characteristics were then measured over a comparable temperature range. Net photosynthesis tended stongly to be greatest, and intracellular resistance to CO₂ transport to be lowest, when the measurement temperature corresponded to the daytime growth temperature, suggesting pronounced acclimation of the plants to the growth temperature. The preferred growth temperature was close to the 25/20 C regime, since net photosynthesis of these plants, regardless of measurement temperature, was higher and intracellular resistance lower, than in plants from any other regime.     

Growth dynamics and population development in an alpine grassland under elevated CO2

Leaf expansion, population dynamics and reproduction under elevated CO₂ were studied for two dominant and four subdominant species in a high alpine grassland (2500 above sea level, Swiss Central Alps). Plots of alpine heath were exposed to 335 l l^-1 and 680 l l^-1 CO₂ in open-top chambers over three growing seasons. Treatments also included natural and moderately improved mineral nutrient supply (40 kg N ha^-1 year^-1 in an NPK fertilizer mix). Seasonal dynamics of leaf expansion, which was studied for the dominant graminoid Carex curvula only, were not affected by elevated CO₂ during two warm seasons or during a cool season. Improved nutrient supply increased both the expansion rate and the duration of leaf growth but elevated CO₂ did not cause any further stimulation. Plant and tiller density (studied in all species) increased under elevated CO₂ in the codominant Leontodon helveticus and the subdominant Trifolium alpinum, remained unchanged in two other minor species Poa alpina and Phyteuma globulariifolium, and decreased in Carex curvula. In Potentilla aurea elevated CO₂ compensated for a natural decline in shoot number. By year 3 the number of fertile shoots in Leontodon and individual seed weight in Carex were slightly increased under elevated CO₂, indicating CO₂ effects on sexual reproduction in these two dominant species. The results suggest that the effects of elevated CO₂ on the population dynamics of the species studied were not general, but species-specific and rather moderate effects. However, the reduction of tiller density in Carex curvula, in contrast to the increases observed in Leontodon helveticus and Trifolium alpinum, indicates that elevated CO₂ may negatively affect the abundance of the species most characteristic of this alpine plant community.     

Acclimation of peanut (Arachis hypogaea L.) leaf photosynthesis to elevated growth CO2 and temperature

Peanut (Arachis hypogaea L. cv. Florunner) was grown from seed sowing to plant maturity under two daytime CO₂ concentrations ([CO₂]) of 360 μmol mol⁻¹ (ambient) and 720 μmol mol⁻¹ (elevated) and at two temperatures of 1.5 and 6.0 °C above ambient temperature. The objectives were to characterize peanut leaf photosynthesis responses to long-term elevated growth [CO₂] and temperature, and to assess whether elevated [CO₂] regulated peanut leaf photosynthetic capacity, in terms of activity and protein content of ribulose bisphosphate carboxylase-oxygenase (Rubisco), Rubisco photosynthetic efficiency, and carbohydrate metabolism. At both growth temperatures, leaves of plants grown under elevated [CO₂] had higher midday photosynthetic CO₂ exchange rate (CER), lower transpiration and stomatal conductance and higher water-use efficiency, compared to those of plants grown at ambient [CO₂]. Both activity and protein content of Rubisco, expressed on a leaf area basis, were reduced at elevated growth [CO₂]. Declines in Rubisco under elevated growth [CO₂] were 27–30% for initial activity, 5–12% for total activity, and 9–20% for protein content. Although Rubisco protein content and activity were down-regulated by elevated [CO₂], Rubisco photosynthetic efficiency, the ratio of midday light-saturated CER to Rubisco initial or total activity, of the elevated-[CO₂] plants was 1.3- to 1.9-fold greater than that of the ambient-[CO₂] plants at both growth temperatures. Leaf soluble sugars and starch of plants grown at elevated [CO₂] were 1.3- and 2-fold higher, respectively, than those of plants grown at ambient [CO₂]. Under elevated [CO₂], leaf soluble sugars and starch, however, were not affected by high growth temperature. In contrast, high temperature reduced leaf soluble sugars and starch of the ambient-[CO₂] plants. Activity of sucrose-P synthase, but not adenosine 5′-diphosphoglucose pyrophosphorylase, was up-regulated under elevated growth [CO₂]. Thus, in the absence of other environmental stresses, peanut leaf photosynthesis would perform well under rising atmospheric [CO₂] and temperature as predicted for this century.     

Genotypic variation in source and sink traits affects the response of photosynthesis and growth to elevated atmospheric CO2

This study aimed to understand the response of photosynthesis and growth to e-CO₂ conditions (800 vs. 400 μmol mol⁻¹) of rice genotypes differing in source–sink relationships. A proxy trait called local C source–sink ratio was defined as the ratio of flag leaf area to the number of spikelets on the corresponding panicle, and five genotypes differing in this ratio were grown in a controlled greenhouse. Differential CO₂ resources were applied either during the 2 weeks following heading (EXP1) or during the whole growth cycle (EXP2). Under e-CO₂, low source–sink ratio cultivars (LSS) had greater gains in photosynthesis, and they accumulated less nonstructural carbohydrate in the flag leaf than high source–sink ratio cultivars (HSS). In EXP2, grain yield and biomass gain was also greater in LSS probably caused by their strong sink. Photosynthetic capacity response to e-CO₂ was negatively correlated across genotypes with local C source–sink ratio, a trait highly conserved across environments. HSS were sink-limited under e-CO₂, probably associated with low triose phosphate utilization (TPU) capacity. We suggest that the local C source–sink ratio is a potential target for selecting more CO₂-responsive cultivars, pending validation for a broader genotypic spectrum and for field conditions.     

温度和CO2浓度升高对荒漠藻结皮光合作用的影响

2007年,对腾格里沙漠东南缘沙坡头地区1956年(51龄)和1981年(26龄)人工植被区及自然植被区的藻结皮净光合速率(P_n)变化,及其与结皮含水量(>100%、40%～60%和<20%)、大气CO₂浓度(360和700 mg·L^-1)和温度(13 ℃、24 ℃ 和28 ℃)的关系进行研究.结果表明：51龄、26龄人工植被区和自然植被区的藻结皮P_n分别为3.4、4.4和3.2 μmol·m^-2·s^-1,且51龄人工植被区藻结皮的P_n显著高于26龄人工植被区和自然植被区;藻结皮含水量对其P_n影响显著,且中等含水量(40%～60%)藻结皮的P_n显著高于低含水量(<20%)和高含水量(>100%);CO₂倍增(700 mg·L^-1)后,中等和高含水量藻结皮的P_n增加了1.8～3.3倍,而低含水量时,藻结皮的P_n变化不明显;高含水量和中等含水量处理时,24 ℃和28 ℃条件下藻结皮的P_n较13 ℃时提高27%～66%,而在低含水量时,不同温度的藻结皮P_n值无显著差异.     

Effects of elevated CO2, warming and precipitation change on plant growth,photosynthesis and peroxidation in dominant species from North China grassland

Warming, watering and elevated atmospheric CO₂-concentration effects have been extensively studied separately; however, their combined impact on plants is not well understood. In the current research, we examined plant growth and physiological responses of three dominant species from the Eurasian Steppe with different functional traits to a combination of elevated CO₂, high temperature, and four simulated precipitation patterns. Elevated CO₂ stimulated plant growth by 10.8–41.7 % for a C₃ leguminous shrub, Caragana microphylla, and by 33.2–52.3 % for a C₃ grass, Stipa grandis, across all temperature and watering treatments. Elevated CO₂, however, did not affect plant biomass of a C₄ grass, Cleistogenes squarrosa, under normal or increased precipitation, whereas a 20.0–69.7 % stimulation of growth occurred with elevated CO₂ under drought conditions. Plant growth was enhanced in the C₃ shrub and the C₄ grass by warming under normal precipitation, but declined drastically with severe drought. The effects of elevated CO₂ on leaf traits, biomass allocation and photosynthetic potential were remarkably species-dependent. Suppression of photosynthetic activity, and enhancement of cell peroxidation by a combination of warming and severe drought, were partly alleviated by elevated CO₂. The relationships between plant functional traits and physiological activities and their responses to climate change were discussed. The present results suggested that the response to CO₂ enrichment may strongly depend on the response of specific species under varying patterns of precipitation, with or without warming, highlighting that individual species and multifactor dependencies must be considered in a projection of terrestrial ecosystem response to climatic change.     

Diminution of photosynthesis in rice (Oryza sativa L.) seedlings under elevated CO2 concentration and increased temperature

The photosynthetic responses to elevated CO₂ concentration (EC) at ambient and ambient +4°C temperature were aßsessed in the second leaf of rice (Oryza sativa L.) seedlings. The duration of different leaf developmental phases, as characterised by changes in photosynthetic pigment contents and photochemical potential, was protracted in the seedlings grown under EC. On the other hand, a temporal shift in the phases of development with an early onset of senescence was observed in the seedlings grown under EC at ambient +4°C temperature. The contents of carotenoids, ß-carotene, and xanthophyll cycle pigments revealed that EC downregulated the protective mechanism of photosynthetic apparatus against oxidative damages, whereas this mechanism assumed higher significance under EC at ambient +4°C temperature. We observed an enhancement in electron transport activity, photochemical potential, and net photosynthesis in spite of a loss in photostasis of photosynthesis under EC. On the other hand, the loss in photostasis of photosynthesis was exacerbated under EC at ambient +4°C temperature due to the decline in electron transport activity, photochemical potential, and net photosynthesis.     

Effects of elevated CO2 and increased nitrogen deposition on photosynthesis and growth of understory plants in spruce model ecosystems

We studied the effects of atmospheric CO₂ enrichment (280, 420 and 560 l CO₂ l^–1) and increased N deposition (0,30 and 90 kg ha^–1 year^–1) on the spruce-forest understory species Oxalis acetosella, Homogyne alpina and Rubus hirtus. Clones of these species formed the ground cover in nine 0.7 m² model ecosystems with 5-year-old Picea abies trees (leaf area index of approx 2.2). Communities grew on natural forest soil in a simulated montane climate. Independently of N deposition, the rate of light-saturated net photosynthesis of leaves grown and measured at 420 l CO₂ l^–1 was higher in Oxalis and in Homogyne, but was not significantly different in Rubus compared to leaves grown and measured at the pre-industrial CO₂ concentration of 280 l l^–1. Remarkably, further CO₂ enrichment to 560 l l^–1 caused no additional increase of CO₂ uptake. With increasing CO₂ supply concentrations of non-structural carbohydrates in leaves increased and N concentrations decreased in all species, whereas N deposition had no significant effect on these traits. Above-ground biomass and leaf area production were not significantly affected by elevated CO₂ in the more vigorously growing species O. acetosella and R. hirtus, but the slow growing H. alpina produced almost twice as much biomass and 50% more leaf area per plant under 420 l CO₂ l^–1 compared to 280 l l^–1 (again no further stimulation at 560 l l^–1). In contrast, increased N addition stimulated growth in Oxalis and Rubus but had no effect on Homogyne. In Oxalis (only) biomass per plant was positively correlated with microhabitat quantum flux density at low CO₂, but not at high CO₂ indicating carbon saturation. On the other hand, the less shade-tolerant Homogyne profited from CO₂ enrichment at all understory light levels facilitating its spread into more shady micro-habitats under elevated CO₂. These species-specific responses to CO₂ and N deposition will affect community structure. The non-linear responses to elevated CO₂ of several of the traits studied here suggest that the largest responses to rising atmospheric CO₂ are under way now or have already occurred and possible future responses to further increases in CO₂ concentration are likely to be much smaller in these understory species.     

The impact of elevated CO2 on growth and photosynthesis in Agrostis canina L. ssp. monteluccii adapted to contrasting atmospheric CO2 concentrations

 The aim of this study was to characterise growth and photosynthetic capacity in plants adapted to long-term contrasting atmospheric CO₂ concentrations (C _a). Seeds of Agrostis canina L. ssp. monteluccii were collected from a natural CO₂ transect in central-western Italy and plants grown in controlled environment chambers at both ambient and elevated CO₂ (350 and 700 μmol mol⁻¹) in nutrient-rich soil. Seasonal mean C _a at the source of the plant material ranged from 610 to 451 μmol CO₂ mol⁻¹, derived from C₄ leaf stable carbon isotope discrimination (δ¹³C). Under chamber conditions, CO₂ enrichment stimulated the growth of all populations. However, plants originating from elevated C _a exhibited higher initial relative growth rates (RGRs) irrespective of chamber CO₂ concentrations and a positive relationship was found between RGR and C _a at the seed source. Seed weight was positively correlated with C _a, but differences in seed weight were found to explain no more than 34% of the variation in RGRs at elevated CO₂. Longer-term experiments (over 98 days) on two populations originating from the extremes of the transect (451 and 610 μmol CO₂ mol⁻¹) indicated that differences in growth between populations were maintained when plants were grown at both 350 and 700 μmol CO₂ mol⁻¹. Analysis of leaf material revealed an increase in the cell wall fraction (CWF) in plants grown at elevated CO₂, with plants originating from high C _a exhibiting constitutively lower levels but a variable response in terms of the degree of lignification. In vivo gas exchange measurements revealed no significant differences in light and CO₂ saturated rates of photosynthesis and carboxylation efficiency between populations or with CO₂ treatment. Moreover, SDS-PAGE/ LISA quantification of leaf ribulose bisphosphate carboxylase/oxygenase (Rubisco) showed no difference in Rubisco content between populations or CO₂ treatments. These findings suggest that long-term adaptation to growth at elevated CO₂ may be associated with a potential for increased growth, but this does not appear to be linked with differences in the intrinsic capacity for photosynthesis. Received: 16 August 1996 / Accepted: 19 October 1996