植物光合生产力与冠层蒸散模拟研究进展

植物的光合与蒸腾的模拟已经从经验模型发展到过程模型的时代。概括地论述叶片和冠层尺度上,植物生理生态的基本过程,分析近年来几个有代表性的模型在模拟光合作用,蒸腾作用时,对这些听参数化处理的方法,即在叶片水平上,以Ｆａｒｑｕｈａｒ的叶片光合作用的生化模型,Ｂａｌｌ－Ｂｅｒｒｙ的气孔导度模型等为基础。     

陆地生态系统物质交换模型

陆地生态系统物质交换模型是生态系统水、碳、养分循环研究的重要方向和极具发展前景的不可替代的手段.本文对近年来发展的众多陆地生态系统物质交换模型进行了归类:1)根据建模思路不同分为经验模型、过程模型和混合模型;2)根据物质传输机制的不同分为生物物理模型和生物地球化学模型;3)根据对植被结构处理方式的不同分为单层模型、双层模型和多层模型;4)根据模型应用性不同分为诊断性模型和预测性模型;5)根据构建模型的逻辑结构不同分为自下而上模型和由上而下模型;6)根据研究尺度的不同分为生态系统尺度模型、景观尺度模型和区域尺度模型;7)根据植物生理过程的不同分为光合作用模型和蒸腾作用模型.最后对不同类型模型的优缺点进行综合分析,探讨了新的发展方向.     

叶绿素荧光主动与被动联合观测应用前景

叶绿素荧光是研究植物光合生理机制、量化植被光合作用时空格局以及准确理解气候变化背景下陆地生态系统生产力的关键。然而, 目前对于叶绿素荧光主动与被动联合观测的研究还较少。该文对比了叶绿素荧光主动观测与被动观测的优缺点, 展示了叶片尺度和冠层尺度主动与被动联合观测的仪器设备组成, 探讨了主动与被动联合观测在探索叶绿体尺度-叶片尺度-冠层尺度能量在光合、荧光以及热耗散中的分配, 阐明叶绿素荧光与总初级生产力的关联机制, 验证星基日光诱导叶绿素荧光, 解译叶绿素荧光光谱形状4个方面的应用前景。综上, 叶绿素荧光的主动与被动联合观测对于揭示各尺度上荧光与光合作用之间的关联机制, 改善全球尺度植被生产力模型至关重要。     

光合作用—蒸腾作用—气孔导度的耦合模型及C3植物叶片对环境因子的生理 …

以叶片的气体传输过程为基础,将蒸腾作用包括在以往光合作用－气孔导度的耦合模型中,建立了光合作用－蒸腾作用－气孔导度的耦合模型。该模型可以模拟边界层导度对生理过程的影响。模拟了Ｃ３植物叶片对环境因子,如光照、温度、湿度、边界层导度和ＣＯ２浓度等的生理响应（光合作用、蒸腾作用、气孔导度）以及Ｃｉ和水分利用效率的变化。在环境因子变化于较大范围的情况下,模拟结果符合许多实验结论。     

光合作用-蒸腾作用-气孔导度的耦合模型及C_3植物叶片对环境因子的生理响应(英文)

以叶片的气体传输过程为基础,将蒸腾作用包括在以往光合作用气孔导度的耦合模型中,建立了光合作用蒸腾作用气孔导度的耦合模型。该模型可以模拟边界层导度对生理过程的影响。模拟了Ｃ３植物叶片对环境因子,如光照、温度、湿度、边界层导度和ＣＯ２浓度等的生理响应（光合作用、蒸腾作用、气孔导度）以及Ｃｉ和水分利用效率的变化。在环境因子变化于较大范围的情况下,模拟结果符合许多实验结论。     

辐射传输模型多尺度反演植被理化参数研究进展

植被是生态系统最重要的组成成分之一,许多与植被有关的物质能量交换过程都与植被的理化参数密切相关,因此定量估算植被的理化参数含量对监测植被生长状况、森林火灾预警以及研究全球碳氮循环过程等都具有重要意义.在众多定量反演植被理化参数的方法中,基于数学、物理学以及生物学的基本理论建立起来的辐射传输模型受到越来越多的关注.辐射传输模型描述了植被与入射辐射之间的相互作用过程和特征,相对于传统的经验/半经验方法,辐射传输模型物理意义明确,具有稳定性和可移植性强的特点.在分析国内外最新相关研究的基础上,首先从植被叶片、冠层和像元3个不同的尺度阐述反演植被理化参数的辐射传输模型.叶片尺度上主要介绍PROSPECT模型和LIBERTY模型;冠层尺度上主要介绍SAIL冠层辐射传输模型以及PROSPECT与SAIL耦合的PROSAIL叶片-冠层辐射传输模型;像元尺度的植被理化参数反演目前主要采用冠层尺度的辐射传输模型.其次,分析尺度变化下植被理化参数遥感反演所面临的主要问题,如不同尺度下模型参数敏感性的变化、辐射传输模型的选取以及混合像元的影响等.最后,总结展望植被理化参数反演多模型与多种数据源相互结合的研究趋势,以及将来具有高空间分辨率的高光谱遥感卫星升空后所带来的发展前景.     

土壤-植物-大气连续体水热、CO2通量估算模型研究进展

土壤-植物-大气连续体(SPAC)水热、CO2通量的准确估算对理解陆地和大气的物质和能量交换过程有着重要意义.重点阐述了基于过程的土壤-植物-大气连续体水热、CO2通量模型,综述了统计模型、综合模型及基于遥感的模型的发展过程.其中水热通量统计模型包括基于温度和湿度以及基于温度和辐射的方法;CO2通量统计模型包括基于气候因子或蒸散因子以及基于光能利用率的方法.水热通量过程模型包括大叶、双源、多源和多层的水热传输物理模型;CO2通量过程模型包括叶片尺度及由大叶、双叶和多层方法扩展到冠层尺度的生理生态模型以及光合-蒸腾耦合模型.综合模型包括生物物理模型、生物化学模型和生物地理模型.统计模型形式简单,资料易得,对大范围的水热通量模拟具有指导意义;过程模型准确的揭示了水热和CO2通量传输的物理和生理过程,是大尺度综合模型的基础.未来生态系统水热、CO2通量估算模型将集成各种技术手段进行多尺度网络观测和大尺度机理模拟.     

稳定性碳同位素技术在生态学研究中的应用

 植物光合作用是自然界产生碳同位素分馏的最重要过程,也是碳同位素技术在生态学研究中应用的基础。最初,碳同位素主要应用于光合途径的鉴别。随着技术的不断完善和研究的不断深入,目前此项技术在生态学研究的许多领域都得到了广泛的应用。作者从植物叶片、功能群、群落冠层、生态系统以及全球等几个不同的尺度上,对碳同位素技术在生态学研究中的主要应用进行了简要的总结。     

植被光能利用率研究进展

光能利用率是表征植物固定太阳能效率的指标,指植物通过光合作用将所截获/吸收的能量转化为有机干物质的效率,是植物光合作用的重要概念,也是区域尺度以遥感参数模型监测植被生产力的理论基础。传统的研究方法是通过生物量收获法分别确定植物生长和辐射量,求年或生长季比值;涡度相关技术作为目前直接测定植被冠层与大气间的CO2和水热交换量的唯一方法,使从冠层到景观水平的光能利用率估计成为可能。由于植被类型的差异和气候环境的综合影响使光能利用率表现出显著的空间异质性和时间动态性。在全球尺度上,利用耦合大气CO2观测、卫星遥感和大气辐射传输模型的反演模拟,发现净初级生产力的光能利用率存在明显的地理分异。影响光能利用率时空变异性的因子包括植物内在因素(如叶形、叶羧化酶含量)和外在环境因素。针对光能利用率的时空特征及其波动,建立在通量观测及模型分析基础上的跨尺度模拟,将成为今后该领域的研究重点。     

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。     

RATP: a model for simulating the spatial distribution of radiation absorption, transpiration and photosynthesis within canopies: application to an isolated tree crown

The model RATP (radiation absorption, transpiration and photosynthesis) is presented. The model was designed to simulate the spatial distribution of radiation and leaf-gas exchanges within vegetation canopies as a function of canopy structure, canopy microclimate within the canopy and physical and physiological leaf properties. The model uses a three-dimensional (3D) representation of the canopy (i.e. an array of 3D cells, each characterized by a leaf area density). Radiation transfer is computed by a turbid medium analogy, transpiration by the leaf energy budget approach, and photosynthesis by the Farquhar model, each applied for sunlit and shaded leaves at the individual 3D cell-scale. The model typically operates at a 20–30 min time step. The RATP model was applied to an isolated, 20-year-old walnut tree grown in the field. The spatial distribution of wind speed, stomatal response to environmental variables, and light acclimation of leaf photosynthetic properties were taken into account. Model outputs were compared with data acquired in the field. The model was shown to simulate satisfactorily the intracrown distribution of radiation regime, transpiration and photosynthetic rates, at shoot or branch scales.     

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 approximate Bayesian computation to infer photosynthesis model parameters北大核心CSCD

We developed a method, namely Adaptive Population Monte Carlo Approximate Bayesian Computation (APMC), to estimate the parameters of Farquhar photosynthesis model. Treating the canopy as a big leaf, we applied this method to derive the parameters at canopy scale. Validations against observational data showed that parameters estimated based on the APMC optimization are un-biased for predicting the photosynthesis rate. We conclude that APMC has greater advantages in estimating the model parameters than those of the conventional nonlinear regression models.     

Water-use efficiency as a means of modelling net assimilation in boreal forests

Although the processes governing photosynthesis are well understood, scaling from shoot to canopy in coniferous forests is complex. Development of different sap-flow techniques has made it possible to measure transpiration of whole trees and thereby also of whole canopies. There is a strong link between photosynthesis and transpiration, for which reason it would be interesting to test whether measurements of canopy transpiration could also be used to estimate canopy photosynthesis. As a first step towards this, water-use efficiency (WUE) was studied at branch and canopy scales on the basis of branch gas-exchange measurements, with half-hourly and daily temporal resolution. Half-hourly and daily WUE at both branch and canopy scales showed a strong dependency on vapour-pressure deficit ('e). Branch photosynthesis modelled from branch transpiration and 'e mimicked well measured branch photosynthesis. Also, modelled photosynthesis, scaled to canopy and compared to net forest CO₂ exchange measured by the eddy-covariance technique, occasionally showed good agreement. In spite of these seemingly promising results, there was a difference in the response to 'e between branches and between years, which needs to be better understood.     

Using approximate Bayesian computation to infer photosynthesis model parameters

We developed a method, namely Adaptive Population Monte Carlo Approximate Bayesian Computation (APMC), to estimate the parameters of Farquhar photosynthesis model. Treating the canopy as a big leaf, we applied this method to derive the parameters at canopy scale. Validations against observational data showed that parameters estimated based on the APMC optimization are un-biased for predicting the photosynthesis rate. We conclude that APMC has greater advantages in estimating the model parameters than those of the conventional nonlinear regression models.     

Response of wheat canopy CO2 and water gas-exchange to soil water content under ambient and elevated CO2

The nature of the interaction between drought and elevated CO₂ partial pressure (pC_a) is critically important for the effects of global change on crops. Some crop models assume that the relative responses of transpiration and photosynthesis to soil water deficit are unaltered by elevated pC_a, while others predict decreased sensitivity to drought at elevated pC_a. These assumptions were tested by measuring canopy photosynthesis and transpiration in spring wheat (cv. Minaret) stands grown in boxes with 100 L rooting volume. Plants were grown under controlled environments with constant light (300 µmol m^?2 s^?1) at ambient (36 Pa) or elevated (68 Pa) pC_a and were well watered throughout growth or had a controlled decline in soil water starting at ear emergence. Drought decreased final aboveground biomass (?15%) and grain yield (?19%) while elevated pC_a increased biomass (+24%) and grain yield (+29%) and there was no significant interaction. Elevated pC_a increased canopy photosynthesis by 15% on average for both water regimes and increased dark respiration per unit ground area in well‐watered plants, but not drought‐grown ones. Canopy transpiration and photosynthesis were decreased in drought‐grown plants relative to well‐watered plants after about 20–25 days from the start of the drought. Elevated pC_a decreased transpiration only slightly during drought, but canopy photosynthesis continued to be stimulated so that net growth per unit water transpired increased by 21%. The effect of drought on canopy photosynthesis was not the consequence of a loss of photosynthetic capacity initially, as photosynthesis continued to be stimulated proportionately by a fixed increase in irradiance. Drought began to decrease canopy transpiration below a relative plant‐available soil water content of 0.6 and canopy photosynthesis and growth below 0.4. The shape of these responses were unaffected by pC_a, supporting the simple assumption used in some models that they are independent of pC_a.     

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.     

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     

Modelling canopy CO2 fluxes: are ‘big‐leaf’ simplifications justified?

• 1 The ‘big‐leaf’ approach to calculating the carbon balance of plant canopies is evaluated for inclusion in the ETEMA model framework. This approach assumes that canopy carbon fluxes have the same relative responses to the environment as any single leaf, and that the scaling from leaf to canopy is therefore linear.
• 2 A series of model simulations was performed with two models of leaf photosynthesis, three distributions of canopy nitrogen, and two levels of canopy radiation detail. Leaf‐ and canopy‐level responses to light and nitrogen, both as instantaneous rates and daily integrals, are presented.
• 3 Observed leaf nitrogen contents of unshaded leaves are over 40% lower than the big‐leaf approach requires. Scaling from these leaves to the canopy using the big‐leaf approach may underestimate canopy photosynthesis by ~20%. A leaf photosynthesis model that treats within‐leaf light extinction displays characteristics that contradict the big‐leaf theory. Observed distributions of canopy nitrogen are closer to those required to optimize this model than the homogeneous model used in the big‐leaf approach.
• 4 It is theoretically consistent to use the big‐leaf approach with the homogeneous photosynthesis model to estimate canopy carbon fluxes if canopy nitrogen and leaf area are known and if the distribution of nitrogen is assumed optimal. However, real nitrogen profiles are not optimal for this photosynthesis model, and caution is necessary in using the big‐leaf approach to scale satellite estimates of leaf physiology to canopies. Accurate prediction of canopy carbon fluxes requires canopy nitrogen, leaf area, declining nitrogen with canopy depth, the heterogeneous model of leaf photosynthesis and the separation of sunlit and shaded leaves. The exact nitrogen profile is not critical, but realistic distributions can be predicted using a simple model of canopy nitrogen allocation.

     

Scaling CO2-photosynthesis relationships from the leaf to the canopy

Responses of individual leaves to short-term changes in CO₂ partial pressure have been relatively well studied. Whole-plant and plant community responses to elevated CO₂ are less well understood and scaling up from leaves to canopies will be complicated if feedbacks at the small scale differ from feedbacks at the large scale. Mathematical models of leaf, canopy, and ecosystem processes are important tools in the study of effects on plants and ecosystems of global environmental change, and in particular increasing atmospheric CO₂, and might be used to scale from leaves to canopies. Models are also important in assessing effects of the biosphere on the atmosphere. Presently, multilayer and big leaf models of canopy photosynthesis and energy exchange exist. Big leaf models — which are advocated here as being applicable to the evaluation of impacts of global change on the biosphere — simplify much of the underlying leaf-level physics, physiology, and biochemistry, yet can retain the important features of plant-environment interactions with respect to leaf CO₂ exchange processes and are able to make useful, quantitative predictions of canopy and community responses to environmental change. The basis of some big leaf models of photosynthesis, including a new model described herein, is that photosynthetic capacity and activity are scaled vertically within a canopy (by plants themselves) to match approximately the vertical profile of PPFD. The new big leaf model combines physically based models of leaf and canopy level transport processes with a biochemically based model of CO₂ assimilation. Predictions made by the model are consistent with canopy CO₂ exchange measurements, although a need exists for further testing of this and other canopy physiology models with independent measurements of canopy mass and energy exchange at the time scale of 1 h or less.Abbreviations LAI leaf area index - NIR near infrared (700–3000 nm) radiation - PAR photosynthetically active (400–700 nm) radiation - PI photosynthetic irradiance (400–700 nm) - PPFD photosynthetic photon flux area density (400–700 nm) - PS I Photosystem I - PS II Photosystem II - Rubisco ribulose-1,5-bisphosphate carboxylase/oxygenase - RuP₂ ribulose-1,5-bisphosphate