不同有机肥量对旱地玉米光合特性和产量的影响

在渭北旱塬合阳试验基地进行了4年的旱地有机培肥试验, 研究不同有机肥施用量对旱地玉米光合特性和产量的影响.结果表明： 与单施化肥相比,施有机肥处理的玉米各生育时期的叶片光合速率和气孔导度显著增大,胞间CO2浓度显著减小;随着有机肥施用量的增加,玉米叶片各生育时期的叶片光合速率和气孔导度逐渐增大,胞间CO2浓度逐渐减小.玉米各生育时期的光合作用主要受非气孔因素限制,施用有机肥显著降低了非气孔因素对光合的限制.连续4年施用有机肥,改善了土壤养分状况,使养分不再是玉米光合速率和产量的主要限制因子.     

Growth at elevated CO(2) delays the adverse effects of drought stress on leaf photosynthesis of the C(4) sugarcane

Sugarcane (Saccharum officinarum L. cv. CP72-2086) was grown in sunlit greenhouses at daytime [CO(2)] of 360 (ambient) and 720 (elevated)mumolmol(-1). Drought stress was imposed for 13d when plants were 4 months old, and various photosynthetic parameters and levels of nonstructural carbohydrates were determined for uppermost fully expanded leaves of well-watered (control) and drought stress plants. Control plants at elevated [CO(2)] were 34% and 25% lower in leaf stomatal conductance (g(s)) and transpiration rate (E) and 35% greater in leaf water-use efficiency (WUE) than their counterparts at ambient [CO(2)]. Leaf CO(2) exchange rate (CER) and activities of Rubisco, NADP-malate dehydrogenase, NADP-malic enzyme and pyruvate P(i) dikinase were marginally affected by elevated [CO(2)], but were reduced by drought, whereas activity of PEP carboxylase was reduced by elevated [CO(2)], but not by drought. At severe drought developed at day 12, leaf g(s) and WUE of ambient-[CO(2)] stress plants declined to 5% and 7%, while elevated-[CO(2)] stress plants still maintained g(s) and WUE at 20% and 74% of their controls. In control plants, elevated [CO(2)] did not enhance the midday levels of starch, sucrose, or reducing sugars. For both ambient- and elevated-[CO(2)] stress plants, severe drought did not affect the midday level of sucrose but substantially reduced that of starch. Nighttime starch decomposition in control plants was 55% for ambient [CO(2)] and 59% for elevated [CO(2)], but was negligible for stress plants of both [CO(2)] treatments. For both ambient-[CO(2)] control and stress plants, midday sucrose level at day 12 was similar to the predawn value at day 13. In contrast, sucrose levels of elevated-[CO(2)] control and stress plants at predawn of day 13 were 61-65% of the midday values of day 12. Levels of reducing sugars were much greater for both ambient- and elevated-[CO(2)] stress plants, implying an adaptation to drought stress. Sugarcane grown at elevated [CO(2)] had lower leaf g(s) and E and greater leaf WUE, which helped to delay the adverse effects of drought and, thus, allowed the stress plants to continue photosynthesis for at least an extra day during episodic drought cycles.     

Nocturnal stomatal conductance responses to rising [CO2], temperature and drought

The response of nocturnal stomatal conductance (g(s,n)) to rising atmospheric CO(2) concentration ([CO(2)]) is currently unknown, and may differ from responses of daytime stomatal conductance (g(s,d)). Because night-time water fluxes can have a significant impact on landscape water budgets, an understanding of the effects of [CO(2)] and temperature on g(s,n) is crucial for predicting water fluxes under future climates. Here, we examined the effects of [CO(2)] (280, 400 and 640 μmol mol(-1)), temperature (ambient and ambient + 4°C) and drought on g(s,n,) and g(s,d) in Eucalyptus sideroxylon saplings. g(s,n) was substantially higher than zero, averaging 34% of g(s,d). Before the onset of drought, g(s,n) increased by 85% when [CO(2)] increased from 280 to 640 μmol mol(-1), averaged across both temperature treatments. g(s,n) declined with drought, but an increase in [CO(2)] slowed this decline. Consequently, the soil water potential at which g(s,n) was zero (Ψ(0)) was significantly more negative in elevated [CO(2)] and temperature treatments. g(s,d) showed inconsistent responses to [CO(2)] and temperature. g (s,n) may be higher in future climates, potentially increasing nocturnal water loss and susceptibility to drought, but cannot be predicted easily from g(s,d). Therefore, predictive models using stomatal conductance must account for both g(s,n) and g(s,d) when estimating ecosystem water fluxes.     

Determination of the site of CO₂ sensing in poplar: is the area-based N content and anatomy of new leaves determined by their immediate CO₂ environment or by the CO₂ environment of mature leaves?

Exposure to an elevated CO(2) concentration ([CO(2)]) generally decreases leaf N content per unit area (N(area)) and stomatal density, and increases leaf thickness. Mature leaves can 'sense' elevated [CO(2)] and this regulates stomatal development of expanding leaves (systemic regulation). It is unclear if systemic regulation is involved in determination of leaf thickness and N(area)-traits that are significantly correlated with photosynthetic capacity. A cuvette system was used whereby [CO(2)] around mature leaves was controlled separately from that around expanding leaves. Expanding leaves of poplar (Populus trichocarpa×P. deltoides) seedlings were exposed to elevated [CO(2)] (720 μmol mol(-1)) while the remaining mature leaves inside the cuvette were under ambient [CO(2)] of 360 μmol mol(-1). Reverse treatments were performed. Exposure of newly developing leaves to elevated [CO(2)] increased their thickness, but when mature leaves were exposed to elevated [CO(2)] the increase in thickness of new leaves was less pronounced. The largest response to [CO(2)] was reflected in the palisade tissue thickness (as opposed to the spongy tissue) of new leaves. The N(area) of new leaves was unaffected by the local [CO(2)] where the new leaves developed, but decreased following the exposure of mature leaves to elevated [CO(2)]. The volume fraction of mesophyll cells compared with total leaf and the mesophyll cell density changed in a manner similar to the response of N(area). These results suggest that N(area) is controlled independently of the leaf thickness, and suggest that N(area) is under systemic regulation by [CO(2)] signals from mature leaves that control mesophyll cell division.     

Comparative field water relations of three Mediterranean shrub species co-occurring at a natural CO(2) vent

Annual variations in the water relations and stomatal response of Erica arborea, Myrtus communis and Juniperus communis occurring at a natural CO(2) vent were analysed under Mediterranean field conditions. A distinct gradient of CO(2)concentration ([CO(2)]) exists between two sites near a natural CO(2)-emitting vent, with higher [CO(2)] (700 micromol mol(-1)) in the proximity of the CO(2) spring. Plants at the CO(2) spring site have been growing for generations at elevated [CO(2)]. At both sites, maximum leaf conductance was related to predawn shoot water potential. The effects of water deficits during the summer drought were severe. Leaf conductance and water potential recovered after major rainfalls in September to predrought values. Strong relationships between leaf conductance, predawn water potential, and leaf-specific hydraulic resistance are consistent with the role of stomata in regulating plant water status. Considerable between-species variation in sensitivity of water potentials and stomatal characters to elevated [CO(2)] were observed. Common to all the shrubs were a reduction in leaf conductance and an increase in water potentials in response to elevated [CO(2)]. Elevated [CO(2)] decreased the sensitivity of leaf conductance to vapour pressure deficit. Morphological characters (including stomatal density and degree of sclerophylly) showed site-dependent variations, but degree and sign of such changes varied with the species and/or the season. Measurements of discrimination against (13)C provided evidence for long-term decreases of water use efficiency in CO(2) spring plants. Analysis of C isotope composition suggested that a downward adjustment of photosynthetic capacity may have occurred under elevated [CO(2)]. Elevated [CO(2)] effects on water relations and leaf morphology persisted in the long term, but the three shrubs growing in the same environment showed species-specific responses.     

Long-term growth of soybean at elevated [CO2] does not cause acclimation of stomatal conductance under fully open-air conditions

Accurately predicting plant function and global biogeochemical cycles later in this century will be complicated if stomatal conductance (g(s)) acclimates to growth at elevated [CO(2)], in the sense of a long-term alteration of the response of g(s) to [CO(2)], humidity (h) and/or photosynthetic rate (A). If so, photosynthetic and stomatal models will require parameterization at each growth [CO(2)] of interest. Photosynthetic acclimation to long-term growth at elevated [CO(2)] occurs frequently. Acclimation of g(s) has rarely been examined, even though stomatal density commonly changes with growth [CO(2)]. Soybean was grown under field conditions at ambient [CO(2)] (378 micromol mol(-1)) and elevated [CO(2)] (552 micromol mol(-1)) using free-air [CO(2)] enrichment (FACE). This study tested for stomatal acclimation by parameterizing and validating the widely used Ball et al. model (1987, Progress in Photosynthesis Research, vol IV, 221-224) with measurements of leaf gas exchange. The dependence of g(s) on A, h and [CO(2)] at the leaf surface was unaltered by long-term growth at elevated [CO(2)]. This suggests that the commonly observed decrease in g(s) under elevated [CO(2)] is due entirely to the direct instantaneous effect of [CO(2)] on g(s) and that there is no longer-term acclimation of g(s) independent of photosynthetic acclimation. The model accurately predicted g(s) for soybean growing under ambient and elevated [CO(2)] in the field. Model parameters under ambient and elevated [CO(2)] were indistinguishable, demonstrating that stomatal function under ambient and elevated [CO(2)] could be modelled without the need for parameterization at each growth [CO(2)].     

土壤干旱条件下氮素营养对玉米内源激素含量影响

在田间持水量分别保持于35％、55％和75％±5％的土壤水分条件下,利用盆栽实验研究了土壤干旱和氮素营养对玉米内源激素和气孔导度的影响．结果表明,土壤干旱下氮素营养明显降低了玉米根系木质部汁液ABA浓度,而正常供水下施氮处理间则无显著差异(施氮处理仍较低),同时测定的叶片ABA浓度则呈相反的变化趋势,表现为干旱下施氮处理要高于不施氮处理;施氮处理木质部汁液中ZRs浓度应低于相应的不施氮处理,在调控气孔行为方面并未表现拮抗ABA作用;3种土壤水分条件下,施氮玉米叶片的气孔导度均高于不施氮处理,与木质部汁液ABA浓度呈负相关,说明施氮处理较低的根源ABA浓度是导致其气孔导度较大的主要原因．     

Carbon isotope discrimination and bundle sheath leakiness in three C(4) subtypes grown under variable nitrogen,water and atmospheric CO(2) supply

The changes in composition and productivity of semi-arid C(4) grassland, anticipated with rising atmospheric CO(2), will depend on soil water and nutrient availability. The interactive effects of soil resource limitation and elevated CO(2 )on these grasses, furthermore, may vary among C(4) biochemical subtypes (NADP-ME, NAD-ME, PCK) that differ in bundle sheath leakiness (Phi) responses to drought and nitrogen supply. To address C(4) subtype responses to soil resource gradients, the carbon isotope discrimination (Delta), bundle sheath leakiness (Phi), leaf gas exchange (A, g(s), c(i)/c(a)) and above-ground biomass accumulation were measured on three dominant grasses of semi-desert grassland in south-eastern Arizona. Bouteloua curtipendula (PCK), Aristida glabrata (NADP-ME) and the non-native Eragrostis lehmanniana (NAD-ME) were grown in controlled-environment chambers from seed under a complete, multi-factorial combination of present ambient (370 ppm) and elevated (690 ppm) CO(2) concentration and under high and low water and nitrogen supply. E. lehmanniana (NAD-ME) had the highest photosynthetic rate (A) and lowest Phi compared to the other two grasses when grown under low nitrogen and water availability. However, favourable water and nitrogen supply and elevated atmospheric CO(2) enhanced photosynthetic performance and above-ground biomass production of B. curtipendula (PCK) to a greater extent than in A. glabrata and E. lehmanniana. Contrary to pre vious studies, Phi and Delta in the NADP-ME subtype (A. glabrata) were most affected by changing environmental conditions compared to the other subtypes; deviations from the classic NADP-ME anatomy in Aristida could have accounted for this result. Overall, response of semi-arid grasslands to rising atmospheric CO(2) may depend more on species-specific responses to drought and nitrogen limitation than on general C(4) subtype responses.     

Photosynthesis, productivity, and yield of maize are not affected by open-air elevation of CO2 concentration in the absence of drought

While increasing temperatures and altered soil moisture arising from climate change in the next 50 years are projected to decrease yield of food crops, elevated CO2 concentration ([CO2]) is predicted to enhance yield and offset these detrimental factors. However, C4 photosynthesis is usually saturated at current [CO2] and theoretically should not be stimulated under elevated [CO2]. Nevertheless, some controlled environment studies have reported direct stimulation of C4 photosynthesis and productivity, as well as physiological acclimation, under elevated [CO2]. To test if these effects occur in the open air and within the Corn Belt, maize (Zea mays) was grown in ambient [CO2] (376 micromol mol(-1)) and elevated [CO2] (550 micromol mol(-1)) using Free-Air Concentration Enrichment technology. The 2004 season had ideal growing conditions in which the crop did not experience water stress. In the absence of water stress, growth at elevated [CO2] did not stimulate photosynthesis, biomass, or yield. Nor was there any CO2 effect on the activity of key photosynthetic enzymes, or metabolic markers of carbon and nitrogen status. Stomatal conductance was lower (-34%) and soil moisture was higher (up to 31%), consistent with reduced crop water use. The results provide unique field evidence that photosynthesis and production of maize may be unaffected by rising [CO2] in the absence of drought. This suggests that rising [CO2] may not provide the full dividend to North American maize production anticipated in projections of future global food supply.     

Gas exchange and carbon metabolism in two Prosopis species (Fabaceae) from semiarid habitats: effects of elevated CO2, N supply, and N source

Predicting future plant and ecosystem responses to elevated CO(2) also requires an understanding of the role of other factors, especially soil nitrogen. This is particularly challenging for global aridlands where total N and the relative amounts of nitrate and ammonia vary both spatially and seasonally. We measured gas exchange and primary and secondary C metabolites in seedlings of two dominant aridland shrub species (Prosopis flexuosa [S America] and P. glandulosa [N America]) grown at ambient (350 ppm) or elevated (650 ppm) CO(2) and nitrogen at two levels (low [0.8 mM] and high [8.0 mM]) and at either 1 : 1 or 3 : 1 nitrate to ammonia. Whereas elevated CO(2) increased assimilation rate, water use efficiency, and primary carbon metabolites in both species, these increases were strongly contingent upon nitrogen availability. Elevated CO(2) did not increase secondary metabolites (i.e., phenolics). For these important aridland species, the effects of elevated CO(2) are strongly influenced by nitrogen availability and to a lesser extent by the relative amounts of nitrate and ammonia supplied, which underscores the importance of both the amount and chemical composition of soil nitrogen in mediating the potential responses of seedling growth and establishment of aridland plants under future CO(2)-enriched atmospheres.     

Soil nutrient heterogeneity interacts with elevated CO2 and nutrient availability to determine species and assemblage responses in a model grassland community

Interactive effects of atmospheric CO(2) concentration ([CO(2)]), soil nutrient availability and soil nutrient spatial distribution on the structure and function of plant assemblages remain largely unexplored. Here we conducted a microcosm experiment to evaluate these interactions using a grassland assemblage formed by Lolium perenne, Plantago lanceolata, Trifolium repens, Anthoxanthum odoratum and Holcus lanatus. Assemblages exhibited precise root foraging patterns, had higher total and below-ground biomass, and captured more nitrogen when nutrients were supplied heterogeneously. Root foraging responses were modified by nutrient availability, and the patterns of N capture by interactions between nutrient distribution, availability and [CO(2)]. Greater above-ground biomass was observed under elevated CO(2) only under homogeneous conditions of nutrient supply and at the highest availability level. CO(2) interacted with nutrient distribution and availability to determine foliar percentage N and below : above-ground biomass ratios, respectively. Interactions between nutrient distribution and CO(2) determined the relative contribution to above-ground biomass of four of the species. The responses of dominant and subordinate species to [CO(2)] were dependent on the availability and distribution of nutrients. Our results suggest that soil nutrient distribution has the potential to influence the response of plant species and assemblages to changes in [CO(2)] and nutrient availability.     

干旱胁迫下杨树无性系苗期光合与气孔形态变异研究

为了解不同干旱胁迫处理下,杨树叶片光合特性和气孔形态的变化规律,本研究以4个杨树无性系为材料,对其干旱胁迫下光合指标与气孔形态指标进行测定分析,方差分析结果表明：除气孔器长在处理间差异不显著外,其他性状在无性系间、处理间和无性系与处理交互作用间均达极显著差异水平。随着土壤相对含水量的逐渐降低,无性系净光合速率、蒸腾速率和气孔导度都呈现出不同程度的下降;而胞间二氧化碳浓度呈现为先降后升趋势,表明当土壤相对含水量大于40%时,光合作用的主要限制因子是气孔因素,当土壤相对含水量低于40%时,光合作用的主要限制因子转换为非气孔因素。气孔形态研究结果表明,各指标均随土壤相对含水量的持续降低呈现出持续下降趋势。利用隶属函数法对不同干旱胁迫下4个杨树无性系的抗旱性进行综合评价,结果表明：白城小青黑杨的抗旱性优于其他三个无性系。     

Changes in mass and energy transfer between the canopy and the atmosphere: model development and testing with a free-air CO2 enrichment (FACE) experiment

The rationale for this study is found in the probable higher temperatures and changes in rainfall patterns that are expected in the future as a result of increasing levels of CO2 in the atmosphere. In particular, higher air temperatures may cause an increase in evapotranspiration demand while a reduction in rainfall could increase the severity and duration of drought in arid and semi-arid regions. Representation of the water transfer scheme includes water uptake by roots and the interaction between evapotranspiration and CO2 enrichment. The predicted response of a spring wheat (Triticum aestivum L. cv. Yecora rojo) canopy in terms of energy exchange processes to elevated atmospheric CO2 level was tested against measurements collected at the FACE (Free Air Enrichment Experiment) site in 1994. Simulated and measured canopy conductances were reduced by about 30% under elevated [CO2] under optimum conditions of water supply. Reductions in latent heat fluxes under elevated instead of ambient [CO2] caused reductions in both simulated and measured seasonal water use of 6% under optimum and 2% under suboptimum irrigation. The soil-plant-atmosphere water transfer scheme proposed here offers several advances in the simulation of land surface interactions. First, the stomatal resistance model minimizes assumptions in existing land surface schemes about the effects of interactions among environmental conditions (radiation, temperature, CO2) upon stomatal behavior. These interactions are resolved in the calculation of CO2 in which processes are already well understood.     

A multi-biome gap in understanding of crop and ecosystem responses to elevated CO2

A key finding from elevated [CO(2)] field experiments is that the impact of elevated [CO(2)] on plant and ecosystem function is highly dependent upon other environmental conditions, namely temperature and the availability of nutrients and soil moisture. In addition, there is significant variation in the response to elevated [CO(2)] among plant functional types, species and crop varieties. However, experimental data on plant and ecosystem responses to elevated [CO(2)] are strongly biased to economically and ecologically important systems in the temperate zone. There is a multi-biome gap in experimental data that is most severe in the tropics and subtropics, but also includes high latitudes. Physiological understanding of the environmental conditions and species found at high and low latitudes suggest they may respond differently to elevated [CO(2)] than well-studied temperate systems. Addressing this knowledge gap should be a high priority as it is vital to understanding 21st century food supply and ecosystem feedbacks on climate change.     

The response of photosynthesis and stomatal conductance to rising [CO2]: mechanisms and environmental interactions

This review summarizes current understanding of the mechanisms that underlie the response of photosynthesis and stomatal conductance to elevated carbon dioxide concentration ([CO2]), and examines how downstream processes and environmental constraints modulate these two fundamental responses. The results from free-air CO2 enrichment (FACE) experiments were summarized via meta-analysis to quantify the mean responses of stomatal and photosynthetic parameters to elevated [CO2]. Elevation of [CO2] in FACE experiments reduced stomatal conductance by 22%, yet, this reduction was not associated with a similar change in stomatal density. Elevated [CO2] stimulated light-saturated photosynthesis (Asat) in C3 plants grown in FACE by an average of 31%. However, the magnitude of the increase in Asat varied with functional group and environment. Functional groups with ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco)-limited photosynthesis at elevated [CO2] had greater potential for increases in Asat than those where photosynthesis became ribulose-1,5-bisphosphate (RubP)-limited at elevated [CO2]. Both nitrogen supply and sink capacity modulated the response of photosynthesis to elevated [CO2] through their impact on the acclimation of carboxylation capacity. Increased understanding of the molecular and biochemical mechanisms by which plants respond to elevated [CO2], and the feedback of environmental factors upon them, will improve our ability to predict ecosystem responses to rising [CO2] and increase our potential to adapt crops and managed ecosystems to future atmospheric [CO2].     

Photosynthetic responses of a C3 and three C4 species of the genus Panicum (s.l.) with different metabolic subtypes to drought stress

Young plants of Panicum bisulcatum (C(3)), Zuloagaea bulbosa [NADP-malic enzyme (ME)-C(4)], P. miliaceum (NAD-ME-C(4)) and Urochloa maxima [phosphoenolpyruvate carboxykinase (PCK)-C(4)] were subjected to drought stress (DS) in soil for 6?days. The C(3) species showed severe wilting symptoms at higher soil water potential (-1.1?MPa) and relative leaf water content (77?%) than in the case of the C(4) species (-1.5 to -1.7?MPa; 58-64?%). DS decreased photosynthesis, both under atmospheric and under saturating CO(2). Stomatal limitation of net photosynthesis (P (N)) in the C(3), but not in the C(4) species was indicated by P (N)/C (o) curves. Chlorophyll fluorescence of photosystem II, resulting from different cell types in the four species, indicated NADPH accumulation and non-stomatal limitation of photosynthesis in all four species, even under high CO(2). In the NAD-ME-C(4) and the PCK-C(4) species, DS plants showed increased violaxanthin de-epoxidase rates. Biochemical analyses of carboxylating enzymes and in vitro enzyme activities of the C(4) enzymes identified the most likely non-stomatal limiting steps of photosynthesis. In P. bisulcatum, declining RubisCO content and activity would explain the findings. In Z. bulbosa, all photosynthesis enzymes declined significantly; photosynthesis is probably limited by the turnover rate of the PEPC reaction. In P. miliaceum, all enzyme levels remained fairly constant under DS, but photosynthesis can be limited by feedback inhibition of the Calvin cycle, resulting in asp inhibition of PEPC. In U. maxima, declines of in vivo PEPC activity and feedback inhibition of the Calvin cycle are the main candidates for non-stomatal limitation of photosynthesis under DS.     

Enhancement of rice canopy carbon gain by elevated CO(2) is sensitive to growth stage and leaf nitrogen concentration

Increasing our understanding of the factors regulating seasonal changes in rice canopy carbon gain (C(gain): daily net photosynthesis -- night respiration) under elevated CO(2) concentrations ([CO(2)]) will reduce our uncertainty in predicting future rice yields and assist in the development of adaptation strategies. In this study we measured CO(2) exchange from rice (Oryza sativa) canopies grown at c. 360 and 690 micromol mol(-1)[CO(2)] in growth chambers continuously over three growing seasons. Stimulation of C(gain) by elevated [CO(2)] was 22-79% during vegetative growth, but decreased to between -12 and 5% after the grain-filling stage, resulting in a 7-22% net enhancement for the whole season. The decreased stimulation of C(gain) resulted mainly from decreased canopy net photosynthesis and partially from increased respiration. A decrease in canopy photosynthetic capacity was noted where leaf nitrogen (N) decreased. The effect of elevated [CO(2)] on leaf area was generally small, but most dramatic under ample N conditions; this increased the stimulation of whole-season C(gain). These results suggest that a decrease in C(gain) enhancement following elevated CO(2) levels is difficult to avoid, but that careful management of nitrogen levels can alter the whole-season C(gain) enhancement.