Modelling Plant Responses to Elevated CO2: How Important is Leaf Area Index?

BACKGROUND AND AIMS: The problem of increasing CO(2) concentration [CO(2)] and associated climate change has generated much interest in modelling effects of [CO(2)] on plants. While variation in growth and productivity is closely related to the amount of intercepted radiation, largely determined by leaf area index (LAI), effects of elevated [CO(2)] on growth are primarily via stimulation of leaf photosynthesis. Variability in LAI depends on climatic and growing conditions including [CO(2)] concentration and can be high, as is known for agricultural crops which are specifically emphasized in this report. However, modelling photosynthesis has received much attention and photosynthesis is often represented inadequately detailed in plant productivity models. Less emphasis has been placed on the modelling of leaf area dynamics, and relationships between plant growth, elevated [CO(2)] and LAI are not well understood. This Botanical Briefing aims at clarifying the relative importance of LAI for canopy assimilation and growth in biomass under conditions of rising [CO(2)] and discusses related implications for process-based modelling. MODEL: A simulation exercise performed for a wheat crop demonstrates recent experimental findings about canopy assimilation as affected by LAI and elevation of [CO(2)]. While canopy assimilation largely increases with LAI below canopy light saturation, effects on canopy assimilation of [CO(2)] elevation are less pronounced and tend to decline as LAI increases. Results from selected model-testing studies indicate that simulation of LAI is often critical and forms an important source of uncertainty in plant productivity models, particularly under conditions of limited resource supply. CONCLUSIONS: Progress in estimating plant growth and productivity under rising [CO(2)] is unlikely to be achieved without improving the modelling of LAI. This will depend on better understanding of the processes of substrate allocation, leaf area development and senescence, and the role of LAI in controlling plant adaptation to environmental changes.     

Raising yield potential of wheat. II. Increasing photosynthetic capacity and efficiency

Past increases in yield potential of wheat have largely resulted from improvements in harvest index rather than increased biomass. Further large increases in harvest index are unlikely, but an opportunity exists for increasing productive biomass and harvestable grain. Photosynthetic capacity and efficiency are bottlenecks to raising productivity and there is strong evidence that increasing photosynthesis will increase crop yields provided that other constraints do not become limiting. Even small increases in the rate of net photosynthesis can translate into large increases in biomass and hence yield, since carbon assimilation is integrated over the entire growing season and crop canopy. This review discusses the strategies to increase photosynthesis that are being proposed by the wheat yield consortium in order to increase wheat yields. These include: selection for photosynthetic capacity and efficiency, increasing ear photosynthesis, optimizing canopy photosynthesis, introducing chloroplast CO(2) pumps, increasing RuBP regeneration, improving the thermal stability of Rubisco activase, and replacing wheat Rubisco with that from other species with different kinetic properties.     

Carbon and nitrogen assimilation in relation to yield: mechanisms are the key to understanding production systems

Improved understanding of crop production systems in relation to N-supply has come from a knowledge of basic plant biochemistry and physiology. Gene expression leads to protein synthesis and the formation of metabolic systems; the ensuing metabolism determines the capacity for growth, development and yield production. This constitutes the genetic potential. These processes set the requirements for the supply of resources. The interactions between carbon dioxide (CO(2)) and nitrate () assimilation and their dynamics are of key importance for crop production. In particular, an adequate supply of, its assimilation to amino acids (for which photosynthesized carbon compounds are required) and their availability for protein synthesis, are essential for metabolism. An adequate supply of stimulates leaf growth and photosynthesis, the former via cell growth and division, the latter by larger contents of components of the light reactions, and those of CO(2) assimilation and related processes. If the supply of resources exceeds the demand set by the genetic potential then production is maximal, but if it is less then potential is not reached; matching resources to potential is the aim of agriculture. However, the connection between metabolism and yield is poorly quantified. Biochemical characteristics and simulation models must be better used and combined to improve fertilizer-N application, efficiency of N-use, and yields. Increasing N-uptake at inadequate N-supply by increasing rooting volume and density is feasible, increasing affinity is less so. It would increase biomass and N/C ratio. With adequate N, at full genetic potential, more C-assimilation per unit N would increase biomass, but energy would be limiting at full canopy. Increasing C-assimilation per unit N would increase biomass but decrease N/C at both large and small N-supply. Increasing production of all biochemical components would increase biomass and demand for N, and maintain N/C ratio. Changing C- or N-assimilation requires modifications to many processes to effect improvements in the whole system; genetic engineering/molecular biological alterations to single steps in the central metabolism are unlikely to achieve this, because targets are unclear, and also because of the complex interactions between processes and environment. Achievement of the long-term objectives of improving crop N-use and yield with fewer inputs and less pollution, by agronomy, breeding or genetic engineering, requires a better understanding of the whole system, from genes via metabolism to yield.     

Exploring the spatial distribution of light interception and photosynthesis of canopies by means of a functional-structural plant model

Background and Aims

At present most process-based models and the majority of three-dimensional models include simplifications of plant architecture that can compromise the accuracy of light interception simulations and, accordingly, canopy photosynthesis. The aim of this paper is to analyse canopy heterogeneity of an explicitly described tomato canopy in relation to temporal dynamics of horizontal and vertical light distribution and photosynthesis under direct- and diffuse-light conditions.

Methods

Detailed measurements of canopy architecture, light interception and leaf photosynthesis were carried out on a tomato crop. These data were used for the development and calibration of a functional–structural tomato model. The model consisted of an architectural static virtual plant coupled with a nested radiosity model for light calculations and a leaf photosynthesis module. Different scenarios of horizontal and vertical distribution of light interception, incident light and photosynthesis were investigated under diffuse and direct light conditions.

Key Results

Simulated light interception showed a good correspondence to the measured values. Explicitly described leaf angles resulted in higher light interception in the middle of the plant canopy compared with fixed and ellipsoidal leaf-angle distribution models, although the total light interception remained the same. The fraction of light intercepted at a north–south orientation of rows differed from east–west orientation by 10 % on winter and 23 % on summer days. The horizontal distribution of photosynthesis differed significantly between the top, middle and lower canopy layer. Taking into account the vertical variation of leaf photosynthetic parameters in the canopy, led to approx. 8 % increase on simulated canopy photosynthesis.

Conclusions

Leaf angles of heterogeneous canopies should be explicitly described as they have a big impact both on light distribution and photosynthesis. Especially, the vertical variation of photosynthesis in canopy is such that the experimental approach of photosynthesis measurements for model parameterization should be revised.     

Using continuous stable isotope measurements to partition net ecosystem CO2 exchange

Ecosystem-scale estimation of photosynthesis and respiration using micrometeorological techniques remains an important, yet difficult, challenge. In this study, we combined micrometeorological and stable isotope methods to partition net ecosystem CO2 exchange (FN) into photosynthesis (F(A)) and respiration (F(R)) in a corn-soybean rotation ecosystem during the summer 2003 corn phase. Mixing ratios of (12)CO2 and (13)CO2 were measured continuously using tunable diode laser (TDL) absorption spectroscopy. The dynamics of the isotope ratio of ecosystem respiration (R), net ecosystem CO2 exchange (deltaN) and photosynthetic discrimination at the canopy scale (delta canopy) were examined. During the period of full canopy closure, F(N) was partitioned into photosynthesis and respiration using both the isotopic approach and the conventional night-time-derived regression methodology. Results showed that deltaR had significant seasonal variation (-32 to -11% per hundred) corresponding closely with canopy phenology. Daytime deltaN typically varied from -12 to -4% per hundred, while delta canopy remained relatively constant in the vicinity of 3% per hundred. Compared with the regression approach, the isotopic flux partitioning showed more short-term variations and was considerably more symmetric about F(N). In this experiment, the isotopic partitioning resulted in larger uncertainties, most of which were caused by the uncertainties in deltaN. and the daytime estimate of deltaR. By sufficiently reducing these uncertainties, the tunable diode laser (TDL)-micrometeorological technique should yield a better understanding of the processes controlling photosynthesis, respiration and ecosystem-scale discrimination.     

Effects of Elevated CO2 and High Temperature on Single Leaf and Canopy Photosynthesis of Rice

The increase of atmospheric CO2 concentration is indisputable. In such condition, photosynthetic response of leaf is relatively well studied, while the comparison of that between single leaf and whole canopy is less emphasized. The stimulation of elevated CO2 on canopy photosynthesis may be different from that on single leaf level. In this study, leaf and canopy photosynthesis of rice ( Oryza sativa L. ) were studied throughout the growing season. High CO2 and temperature had a synergetic stimulation on single leaf photosynthetic rate until grain filling. Photosynthesis of leaf was stimulated by high CO2, although the stimulation was decreased by higher temperature at grain filling stage. On the other hand, the simulation of elevated CO2 on canopy photosynthesis leveled off with time. Stimulation at canopy level disappeared by grain filling stage in beth temperature treatments. Green leaf area index was not significantly affected by CO2 at maturity, but greater in plants grown at higher temperature. Leaf nitrogen content decreased with the increase of CO2 concentration although it was not statistically significant at maturity. Canopy respiration rate increased at flowering stage indicating higher carbon loss. Shading effect caused by leaf development reached maximum at flowering stage. The CO2 stimulation on photosynthesis was greater in single leaf than in canopy. Since enhanced CO2 significantly increased biomass of rice stems and panicles, increase in canopy respiration caused diminishment of CO2 stimulation in canopy net photosynthesis, keaf nitrogen in the canopy level decreased with CO2 concentration and may eventually hasten CO2 stimulation on canopy photosynthesis. Early senescence of canopy leaves in high CO2 is also a possible cause.     

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.     

基于SVAT模型的冬小麦光合作用和蒸散过程研究

在已建立的土壤－植被－大气传输（SVAT）模型中,冠层光合作用／气孔导度耦合子模型可区分遮荫叶和受光叶光合作用强度的差异;作物生长模型考虑了生长呼吸和维持呼吸,模拟与实测结果对比发现,日总蒸散量实测和模拟的根均方差（RMSD）为0.65mm,平均绝对差（MAPD）为14％;对冠层上部净光合作用率日变化过程而言,实测和模拟结果具有较好的一致性。利用模型模拟了冬小麦全生育争光合作用率和蒸散的演变过程。最后,分析了冬小麦蒸散和水分利用效率对不同最大叶面积指数,大气CO2浓度和叶片N含量的响应。     

Bundle sheath leakiness and light limitation during C4 leaf and canopy CO2 uptake

Perennial species with the C(4) pathway hold promise for biomass-based energy sources. We have explored the extent that CO(2) uptake of such species may be limited by light in a temperate climate. One energetic cost of the C(4) pathway is the leakiness () of bundle sheath tissues, whereby a variable proportion of the CO(2), concentrated in bundle sheath cells, retrodiffuses back to the mesophyll. In this study, we scale from leaf to canopy level of a Miscanthus crop (Miscanthus x giganteus hybrid) under field conditions and model the likely limitations to CO(2) fixation. At the leaf level, measurements of photosynthesis coupled to online carbon isotope discrimination showed that leaves within a 3.3-m canopy (leaf area index = 8.3) show a progressive increase in both carbon isotope discrimination and as light decreases. A similar increase was observed at the ecosystem scale when we used eddy covariance net ecosystem CO(2) fluxes, together with isotopic profiles, to partition photosynthetic and respiratory isotopic flux densities (isofluxes) and derive canopy carbon isotope discrimination as an integrated proxy for at the canopy level. Modeled values of canopy CO(2) fixation using leaf-level measurements of suggest that around 32% of potential photosynthetic carbon gain is lost due to light limitation, whereas using determined independently from isofluxes at the canopy level the reduction in canopy CO(2) uptake is estimated at 14%. Based on these results, we identify as an important limitation to CO(2) uptake of crops with the C(4) pathway.     

大气CO2浓度和温度升高对水稻叶片及群体光合作用的影响

大气ＣＯ２浓度升高对植物光合作用的影响研究多集中在单叶水平,在高ＣＯ２及高温下对植物单叶及群体光合进行比较的研究少有报道,而群体水平的研究则是预测生态系统反应所不可缺少的。采用田间开顶式培养室研究了大气ＣＯ２浓度和温度升高对水稻（ＯｒｙｚａｓａｔｉｖａＬ．）叶片及群体光合作用的影响。发现ＣＯ２浓度和温度对水稻叶片光合作用有协同促进作用,而对群体光合作用的促进则随时间的推移而减弱;单叶光合受到的促进作用大于群体光合;叶面积指数只在营养生长期受到促进,冠层叶片含氮量受ＣＯ２影响降低。群体呼吸（包括茎杆）增加及冠层叶片早衰可能是后期ＣＯ２对群体光合促进作用下降的原因。     

Growth Response of a Chrysanthemum Crop to the Environment. II. A Mathematical Analysis Relating Photosynthesis and Growth

Vegetative crops of chrysanthemum were grown for 5 weeks inthree replicate daylit assimilation chambers. Weekly harvestswere made from each crop for growth analysis, and on seven occasionsduring the 5-week period continuous measurements of the netCO₂ exchange rate of each crop were made over a 24 h period.A semi-empirical model for canopy photosynthesis was fittedto these data. The photosynthesis model was then incorporatedinto a simple, dynamic growth model. Using fitted values ofthe canopy photosynthesis parameters, the daily total radiationintegrals, and the experimentally observed relationship betweenthe leaf area index and crop dry matter per unit ground area,the crop growth model was used to simulate growth over the 5-weekperiod. The predicted and measured crop dry weights were inclose agreement over the whole period.     

The impact of modifying photosystem antenna size on canopy photosynthetic efficiency—Development of a new canopy photosynthesis model scaling from metabolism to canopy level processes

Canopy photosynthesis (A_c) describes photosynthesis of an entire crop field and the daily and seasonal integrals of A_c positively correlate with daily and seasonal biomass production. Much effort in crop breeding has focused on improving canopy architecture and hence light distribution inside the canopy. Here, we develop a new integrated canopy photosynthesis model including canopy architecture, a ray tracing algorithm, and C₃ photosynthetic metabolism to explore the option of manipulating leaf chlorophyll concentration ([Chl]) for greater A_c and nitrogen use efficiency (NUE). Model simulation results show that (a) efficiency of photosystem II increased when [Chl] was decreased by decreasing antenna size and (b) the light received by leaves at the bottom layers increased when [Chl] throughout the canopy was decreased. Furthermore, the modelling revealed a modest ~3% increase in A_c and an ~14% in NUE was accompanied when [Chl] reduced by 60%. However, if the leaf nitrogen conserved by this decrease in leaf [Chl] were to be optimally allocated to other components of photosynthesis, both A_c and NUE can be increased by over 30%. Optimizing [Chl] coupled with strategic reinvestment of conserved nitrogen is shown to have the potential to support substantial increases in A_c, biomass production, and crop yields.     

Variation in the degree of coupling between delta13C of phloem sap and ecosystem respiration in two mature Nothofagus forests

Day-to-day variability in the carbon isotope composition of phloem sap (delta13Chd) and ecosystem respiratory CO2 (delta13CR) were measured to assess the tightness of coupling between canopy photosynthesis (delta13Chd) and ecosystem respiration (delta13CR) in two mature Nothofagus solandri (Hook. f.) forests in New Zealand. Abundant phloem-tapping scale insects allowed repeated, nondestructive access to stem phloem sap 1-2 m above ground. delta13Chd was compared with delta13C predicted by an environmentally driven, process-based canopy photosynthesis model. Keeling plots of within-canopy CO2 were used to estimate delta13CR. By including a lag of 3 d, there was good agreement in the timing and direction of variation in delta13Chd and predictions by the canopy photosynthesis model, suggesting that delta13Chd represents a photosynthesis-weighted, integrative record of canopy photosynthesis and conductance. Significant day-to-day variability in delta13CR was recorded at one of the two forests. At this site, delta13CR reflected variability in delta13Chd only on days with <2 mm rain. We conclude that the degree of coupling between canopy photosynthesis and ecosystem respiration varies between sites, and with environmental conditions at a single site.     

Development of the Monsi-Saeki theory on canopy structure and function

BACKGROUND AND AIMS: Monsi and Saeki (1953) published the first mathematical model of canopy photosynthesis that was based on the light attenuation within a canopy and a light response of leaf photosynthesis. This paper reviews the evolution and development of their theory. SCOPE: Monsi and Saeki showed that under full light conditions, canopy photosynthesis is maximized at a high leaf area index (LAI, total leaf area per unit ground area) with vertically inclined leaves, while under low light conditions, it is at a low LAI with horizontal leaves. They suggested that actual plants develop a stand structure to maximize canopy photosynthesis. Combination of the Monsi-Saeki model with the cost-benefit hypothesis in resource use led to a new canopy photosynthesis model, where leaf nitrogen distribution and associated photosynthetic capacity were taken into account. The gradient of leaf nitrogen in a canopy was shown to be a direct response to the gradient of light. This response enables plants to use light and nitrogen efficiently, two resources whose supply is limited in the natural environment. CONCLUSION: The canopy photosynthesis model stimulated studies to scale-up from chloroplast biochemistry to canopy carbon gain and to analyse the resource-use strategy of species and individuals growing at different light and nitrogen availabilities. Canopy photosynthesis models are useful to analyse the size structure of populations in plant communities and to predict the structure and function of future terrestrial ecosystems.     

水分亏缺对冬小麦净光合速率影响程度研究

水分亏缺对冬小麦净光合速率影响程度研究王慧（西北大学城市与资源学系,西安７１００６９）ＥｆｆｅｃｔｏｆＷａｔｅｒＤｅｆｉｃｉｔｏｎＮｅｔＰｈｏｔｏｓｙｎｔｈｅｓｉｓＲａｔｅｏｆＷｉｎｔｅｒＷｈｅａｔ．ＷａｎｇＨｕｉ（ＤｅｐａｔｍｅｎｔｏｆＵｒｂａｎ...     

用光合-蒸散耦合模型模拟冬小麦CO2通量的日变化

根据SPAC理论建立了一个冬小麦光合和蒸散的耦合模型.冬小麦CO2通量包括冠层光合、呼吸和土壤呼吸.冠层光合采用了Farquhar光合作用生化模型,并通过冠层阻力的参数化将光合作用与蒸腾作用耦合起来.用涡度相关方法观测了CO2通量,对模型进行了验证,结果显示模型可以较好地模拟CO2通量日变化过程.对模型的敏感性分析发现日间CO2通量最敏感的参数是初始量子效率.其次,CO2通量对光响应曲线凸度、CO2补偿点、凋萎点和叶面积指数的变化也有着较强的敏感性;夜间CO2通量敏感的参数是最适温度下Rubisco催化能力和暗呼吸参数.     

The diel imprint of leaf metabolism on the δ13 C signal of soil respiration under control and drought conditions

Recent (13) CO(2) canopy pulse chase labeling studies revealed that photosynthesis influences the carbon isotopic composition of soil respired CO(2) (δ(13) C(SR)) even on a diel timescale. However, the driving mechanisms underlying these short-term responses remain unclear, in particular under drought conditions. The gas exchange of CO(2) isotopes of canopy and soil was monitored in drought/nondrought-stressed beech (Fagus sylvatica) saplings after (13) CO(2) canopy pulse labeling. A combined canopy/soil chamber system with gas-tight separated soil and canopy compartments was coupled to a laser spectrometer measuring mixing ratios and isotopic composition of CO(2) in air at high temporal resolution. The measured δ(13) C(SR) signal was then explained and substantiated by a mechanistic carbon allocation model. Leaf metabolism had a strong imprint on diel cycles in control plants, as a result of an alternating substrate supply switching between sugar and transient starch. By contrast, diel cycles in drought-stressed plants were determined by the relative contributions of autotrophic and heterotrophic respiration throughout the day. Drought reduced the speed of the link between photosynthesis and soil respiration by a factor of c. 2.5, depending on the photosynthetic rate. Drought slows the coupling between photosynthesis and soil respiration and alters the underlying mechanism causing diel variations of δ(13) C(SR).     

用光合-蒸散耦合模型模拟冬小麦CO2通量的日变化

根据 SPAC理论建立了一个冬小麦光合和蒸散的耦合模型。冬小麦 CO2 通量包括冠层光合、呼吸和土壤呼吸。冠层光合采用了 Farquhar光合作用生化模型 ,并通过冠层阻力的参数化将光合作用与蒸腾作用耦合起来。用涡度相关方法观测了 CO2通量 ,对模型进行了验证 ,结果显示模型可以较好地模拟 CO2 通量日变化过程。对模型的敏感性分析发现日间 CO2 通量最敏感的参数是初始量子效率。其次 ,CO2 通量对光响应曲线凸度、CO2 补偿点、凋萎点和叶面积指数的变化也有着较强的敏感性 ;夜间 CO2 通量敏感的参数是最适温度下 Rubisco催化能力和暗呼吸参数     

植被冠层尺度生理生态模型的研究进展

随着人们对植物生命活动各个过程研究的不断深入,以植物生理过程、物理过程为基础的各种生理生态学模型逐渐发展起来,而植被冠层尺度生理生态学过程模型已成为生态系统模型的核心之一。目前植被冠层尺度的大叶模型、多层模型、二叶模型以其成熟的理论基础及对植被冠层的光合作用、蒸腾作用较为成功的模拟,得到了广泛的应用。3个模型都以光合作用-气孔导度-蒸腾作用耦合模型为基础,但又具有各自的特点。本文对3种模型的结构及特点进行了总结,并对其进行了比较,简要介绍了目前植被冠层尺度生理生态学模型的应用及存在的问题和发展状况。     

Canopy Photosynthesis and Crop Growth Rate of Eight Temperate Forage Grasses

The rates of canopy and individual leaf photosynthesis, ratesof growth of shoots and roots, and the extinction coefficientfor light of eight temperate forage grasses were determinedin the field during early autumn. Canopy gross photosynthesiswas calculated as net photosynthesis plus dark respiration adjustedfor temperature using a Q₁₀ = 2. The relationships between canopygross photosynthesis and light intensity were hyperbolic, andthe initial slopes of these curves indicated that light wasbeing utilized efficiently at low light intensities. The initialslope depended on the distribution of light in the canopy andthe quantum efficiency of the individual leaves. The maximumrate of canopy gross photosynthesis reflected the maximum rateof individual leaf photosynthesis. Although the maximum rateof canopy gross photosynthesis was correlated with crop growthrate, there was no significant relationship between daily grossphotosynthesis and crop growth rate. Indeed, daily gross photosynthesisvaried by only 22 per cent, whereas the daily growth of shootsand roots varied by 120 per cent. This poor correlation is influencedby a negative correlation (P < 0.01) between the maximumrate of canopy gross photosynthesis and the initial slope ofthe curve relating canopy gross photosynthesis and light intensity.Difficulties in the interpretation of measurements of dark respirationappeared to confound attempts to relate daily net photosynthesisto crop growth rate, individual leaf photosynthesis, and theextinction coefficient for light.