冬小麦光合作用对开放式空气CO2 浓度增高(FACE)的非气孔适应

在同样CO2浓度下测定时,开放式空气CO2浓度增高(FACE,580 μmol CO2 /mol)条件下生长的冬小麦叶片的净光合速率、气孔导度和羧化效率都显著低于普通空气(380 μmol CO2 /mol)中生长的对照叶片.与此相一致,FACE叶片的可溶性蛋白、二磷酸核酮糖羧化酶/加氧酶(Rubisco)和Rubisco活化酶含量也都显著低于对照叶片.这些结果表明,在根系生长不受限制的田间条件下,冬小麦叶片的光合作用对高浓度CO2产生了适应现象,其主要原因可能是碳同化的关键酶Rubisco等含量的降低.     

Elevated CO2 effects on canopy and soil water flux parameters measured using a large chamber in crops grown with free-air CO2 enrichment

An arable crop rotation (winter barley-sugar beet-winter wheat) was exposed to elevated atmospheric CO(2) concentrations ([CO(2) ]) using a FACE facility (Free-Air CO(2) Enrichment) during two rotation periods. The atmospheric [CO(2) ] of the treatment plots was elevated to 550 ppm during daylight hours (T>5°C). Canopy transpiration (E(C) ) and conductance (G(C) ) were measured at selected intervals (>10% of total growing season) using a dynamic CO(2) /H(2) O chamber measuring system. Plant available soil water content (gravimetry and TDR probes) and canopy microclimate conditions were recorded in parallel. Averaged across both growing seasons, elevated [CO(2) ] reduced E(C) by 9%, 18% and 12%, and G(C) by 9%, 17% and 12% in barley, sugar beet and wheat, respectively. Both global radiation (Rg) and vapour pressure deficit (VPD) were the main driving forces of E(C) , whereas G(C) was mostly related to Rg. The responses of E(C) and especially G(C) to [CO(2) ] enrichment were insensitive to weather conditions and leaf area index. However, differences in LAI between plots counteracted the [CO(2) ] impact on E(C) and thus, at least in part, explained the variability of seasonal [CO(2) ] responses between crops and years. As a consequence of lower transpirational canopy water loss, [CO(2) ] enrichment increased plant available soil water content in the course of the season by ca. 15 mm. This was true for all crops and years. Lower transpirational cooling due to a [CO(2) ]-induced reduction of E(C) increased canopy surface and air temperature by up to 2 °C and 0.5 °C, respectively. This is the first study to address effects of FACE on both water fluxes at canopy scale and water status of a European crop rotation.     

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.     

开放式空气二氧化碳浓度增高对小麦氮素吸收利用的影响

2001—2003年,利用我国唯一的农田开放式空气CO₂浓度增高 (free-air carbon dioxide enrichment,FACE) 技术平台,研究了FACE条件下冬小麦宁麦9号不同生育期N含量、吸收、分配和N效率的响应.结果表明：与对照相比,FACE处理使不同生育时期植株含N率显著降低,降幅达4.4%～13.4%;不同生育时期吸N量显著增加(7.4%～25.4%),生育中期的增幅明显大于生育前、后期;不同生育时期茎鞘的N积累能力相对增强,叶片N积累能力相对减弱,而对麦穗N积累能力的影响因生育进程而异;FACE处理使小麦不同生育时期N物质生产效率(5.5%～10.3%)、成熟期N收获指数(16.3%)和N籽粒生产效率(9.3%)均显著或极显著增加;增施N肥,使小麦不同生育时期N含量和吸收量呈增加趋势,使N效率呈下降趋势,而对N在各器官中分配的影响较小.     

Flowering phenology in a species-rich temperate grassland is sensitive to warming but not elevated CO2

* Flowering is a critical stage in plant life cycles, and changes might alter processes at the species, community and ecosystem levels. Therefore, likely flowering-time responses to global change drivers are needed for predictions of global change impacts on natural and managed ecosystems. * Here, the impact of elevated atmospheric CO2 concentration ([CO2]) (550 micromol mol(-1)) and warming (+2 masculineC) is reported on flowering times in a native, species-rich, temperate grassland in Tasmania, Australia in both 2004 and 2005. * Elevated [CO2] did not affect average time of first flowering in either year, only affecting three out of 23 species. Warming reduced time to first flowering by an average of 19.1 d in 2004, acting on most species, but did not significantly alter flowering time in 2005, which might be related to the timing of rainfall. Elevated [CO2] and warming treatments did not interact on flowering time. * These results show elevated [CO2] did not alter average flowering time or duration in this grassland; neither did it alter the response to warming. Therefore, flowering phenology appears insensitive to increasing [CO2] in this ecosystem, although the response to warming varies between years but can be strong.     

开放式空气二氧化碳浓度增高对小麦物质生产与分配的影响

2001—2003年,利用农田开放式空气CO₂浓度增高 (FACE) 技术平台,以冬小麦宁麦9号为供试材料,研究开放式条件下CO₂浓度增高对小麦整个生育期干物质生产与分配的影响.结果表明：与对照相比,FACE处理使小麦播种-越冬始期的干物质生产量略有增加（10.8%),使越冬始期-拔节期、拔节期-孕穗期、孕穗期-抽穗期显著增加,分别增加了31.6%、40.5%、27.2%,使抽穗期-成熟期略有减少(-5.5%),使成熟期生物产量显著增加(13.6%);FACE处理对小麦播种-越冬始期的平均叶面积系数(LAI)和净同化率(NAR)均无显著影响,但使越冬始期-抽穗期LAI显著增加,NAR稍有增加,使抽穗期-抽穗后20 d NAR显著下降;FACE处理使不同生育时期叶片占全株质量的比例下降,而使茎鞘占全株质量的比例增加;FACE小麦抽穗期和成熟期茎鞘可溶性糖和淀粉含量及总量均明显增加.     

Canopy CO2 enrichment permits tracing the fate of recently assimilated carbon in a mature deciduous forest

How rapidly newly assimilated carbon (C) is invested into recalcitrant structures of forests, and how closely C pools and fluxes are tied to photosynthesis, is largely unknown. A crane and a purpose-built free-air CO2 enrichment (FACE) system permitted us to label the canopy of a mature deciduous forest with 13C-depleted CO2 for 4 yr and continuously trace the flow of recent C through the forest without disturbance. Potted C4 grasses in the canopy ('isometers') served as a reference for the C-isotope input signal. After four growing seasons, leaves were completely labelled, while newly formed wood (tree rings) still contained 9% old C. Distinct labels were found in fine roots (38%) and sporocarps of mycorrhizal fungi (62%). Soil particles attached to fine roots contained 9% new C, whereas no measurable signal was detected in bulk soil. Soil-air CO2 consisted of 35% new C, indicating that considerable amounts of assimilates were rapidly returned back to the atmosphere. These data illustrate a relatively slow dilution of old mobile C pools in trees, but a pronounced allocation of very recent assimilates to C pools of short residence times.     

开放式空气CO2浓度增高对水稻产量形成的影响

在大田栽培条件下 ,研究开放式空气CO2 浓度增加 (FACE) 2 0 0 μmol·mol-1的处理对水稻产量及产量构成因素的影响 .结果表明 ,FACE处理对水稻株高和主茎叶片数没有明显影响 ,但使水稻生育进程加快 ,全生育期显著缩短 ,增加施N量可减缓FACE处理对水稻全生育期缩短的程度 ;FACE处理能显著增加分蘖数 ,极显著增加穗数 ,提高结实率 ,但使每穗颖花数显著减少 ;FACE处理能显著提高水稻产量 ,在高N条件下增产幅度更大 ;提高FACE处理的每穗颖花数和单位面积颖花数能极显著提高水稻产量 ,增加施N量是提高FACE处理每穗颖花数和单位面积颖花数的重要措施 .     

Photosynthesis, carboxylation and leaf nitrogen responses of 16 species to elevated pCO2 across four free-air CO2 enrichment experiments in forest, grassland and desert

The magnitude of changes in carboxylation capacity in dominant plant species under long‐term elevated CO₂ exposure (elevated pC_a) directly impacts ecosystem CO₂ assimilation from the atmosphere. We analyzed field CO₂ response curves of 16 C₃ species of different plant growth forms in favorable growth conditions in four free‐air CO₂ enrichment (FACE) experiments in a pine and deciduous forest, a grassland and a desert. Among species and across herb, tree and shrub growth forms there were significant enhancements in CO₂ assimilation (A) by +40±5% in elevated pC_a (49.5–57.1 Pa), although there were also significant reductions in photosynthetic capacity in elevated pC_a in some species. Photosynthesis at a common pC_a (A_a) was significantly reduced in five species growing under elevated pC_a, while leaf carboxylation capacity (V_cmax) was significantly reduced by elevated pC_a in seven species (change of ?19±3% among these species) across different growth forms and FACE sites. Adjustments in V_cmax with elevated pC_a were associated with changes in leaf N among species, and occurred in species with the highest leaf N. Elevated pC_a treatment did not affect the mass‐based relationships between A or V_cmax and N, which differed among herbs, trees and shrubs. Thus, effects of elevated pC_a on leaf C assimilation and carboxylation capacity occurred largely through changes in leaf N, rather than through elevated pC_a effects on the relationships themselves. Maintenance of leaf carboxylation capacity among species in elevated pC_a at these sites depends on maintenance of canopy N stocks, with leaf N depletion associated with photosynthetic capacity adjustments. Since CO₂ responses can only be measured experimentally on a small number of species, understanding elevated CO₂ effects on canopy N_m and N_a will greatly contribute to an ability to model responses of leaf photosynthesis to atmospheric CO₂ in different species and plant growth forms.     

Hourly and seasonal variation in photosynthesis and stomatal conductance of soybean grown at future CO(2) and ozone concentrations for 3 years under fully open-air field conditions

It is anticipated that enrichment of the atmosphere with CO(2) will increase photosynthetic carbon assimilation in C3 plants. Analysis of controlled environment studies conducted to date indicates that plant growth at concentrations of carbon dioxide ([CO(2)]) anticipated for 2050 ( approximately 550 micromol mol(-1)) will stimulate leaf photosynthetic carbon assimilation (A) by 20 to 40%. Simultaneously, concentrations of tropospheric ozone ([O(3)]) are expected to increase by 2050, and growth in controlled environments at elevated [O(3)] significantly reduces A. However, the simultaneous effects of both increases on a major crop under open-air conditions have never been tested. Over three consecutive growing seasons > 4700 individual measurements of A, photosynthetic electron transport (J(PSII)) and stomatal conductance (g(s)) were measured on Glycine max (L.) Merr. (soybean). Experimental treatments used free-air gas concentration enrichment (FACE) technology in a fully replicated, factorial complete block design. The mean A in the control plots was 14.5 micromol m(-2) s(-1). At elevated [CO(2)], mean A was 24% higher and the treatment effect was statistically significant on 80% of days. There was a strong positive correlation between daytime maximum temperatures and mean daily integrated A at elevated [CO(2)], which accounted for much of the variation in CO(2) effect among days. The effect of elevated [CO(2)] on photosynthesis also tended to be greater under water stress conditions. The elevated [O(3)] treatment had no statistically significant effect on mean A, g(s) or J(PSII) on newly expanded leaves. Combined elevation of [CO(2)] and [O(3)] resulted in a slightly smaller increase in average A than when [CO(2)] alone was elevated, and was significantly greater than the control on 67% of days. Thus, the change in atmospheric composition predicted for the middle of this century will, based on the results of a 3 year open-air field experiment, have smaller effects on photosynthesis, g(s) and whole chain electron transport through photosystem II than predicted by the substantial literature on relevant controlled environment studies on soybean and likely most other C3 plants.     

Seasonal changes in the effects of free-air CO2 enrichment (FACE) on growth, morphology and physiology of rice root at three levels of nitrogen fertilization

Over time, the relative effects of elevated [CO₂] on the aboveground photosynthesis, growth and development of rice (Oryza sativa L.) are likely to be changed with increasing duration of CO₂ exposure, but the resultant effects on rice belowground responses remain to be evaluated. To investigate the impacts of elevated [CO₂] on seasonal changes in root growth, morphology and physiology of rice, a free‐air CO₂ enrichment (FACE) experiment was performed at Wuxi, Jiangsu, China, in 2002–2003. A japonica cultivar with large panicle was exposed to two [CO₂] (ambient [CO₂], 370 μmol mol⁻¹; elevated [CO₂], 570 μmol mol⁻¹) at three levels of nitrogen (N): low (LN, 15 g N m⁻²), medium (MN, 25 g N m⁻²) and high N (HN, 35 g N m⁻²). Elevated [CO₂] increased cumulative root volume, root dry weight, adventitious root length and adventitious root number at all developmental stages by 25–71%, which was mainly associated with increased root growth rate during early growth period (EGP) and lower rate of root senescence during late growth period (LGP), while a slight inhibition of root growth rate occurred during middle growth period (MGP). For individual adventitious roots, elevated [CO₂] increased average length, volume, diameter and dry weight early in the season, but the effects gradually disappeared in subsequent stages. Total surface area and active adsorption area per unit root dry weight reached their maxima 10 days earlier in FACE vs. ambient plants, but both of them together with root oxidation ability per unit root dry weight declined with elevated [CO₂] during MGP and LGP, the decline being larger during MGP than LGP. The CO₂‐induced decreases in specific root activities during MGP and LGP were associated with a larger amount of root accumulation during EGP and lower N concentration and higher C/N ratio in roots during MGP and LGP in FACE vs. ambient plants. The results suggest that most of the CO₂‐induced increases in shoot growth of rice are similarly associated with increased root growth.     

Optimal nitrogen allocation controls tree responses to elevated CO2

Despite the abundance of experimental data, understanding of forest responses to elevated CO2 is limited. Here I show that a key to previously unexplained production and leaf area responses lies in the interplay between whole-plant nitrogen (N) allocation and leaf photosynthesis. A simple tree growth model, controlled by net growth maximization through optimization of leaf area index (LAI) and plant N, is used to analyse CO2 responses in both young, expanding and closed, steady-state canopies. The responses are sensitive to only two independent parameters, the photosynthetic capacity per leaf N (a) and the fine-root N:leaf N ratio. The model explains observed CO2 responses of photosynthesis, production and LAI in four forest free air CO2 enrichment (FACE) experiments. Insensitivity of LAI except at low LAI, increase in light-use efficiency, and photosynthetic down-regulation (as a result of reduced leaf N per area) at elevated CO2 are all explained through the combined effects on a and leaf quantum efficiency. The model bridges the gap between the understanding of leaf-level and plant-level responses and provides a transparent framework for interpreting and linking structural (LAI) and functional (net primary production (NPP):gross primary production (GPP) ratio, light-use efficiency, photosynthetic down-regulation) responses to elevated CO2.     

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].