Responses of water use efficiency of 9 plant species to light and CO2 and their modeling

Photosynthesis coupled with transpiration determines water use efficiency (WUE) at leaf level, and the responses of WUE controlled by gas exchanges through stomata to environment are the basis of carbon and water cycles in the ecosystem. In this paper, by using Li-6400 Portable Photosynthesis System (LI-COR), WUE at leaf level was analyzed under controlled photosynthetic photons flux density (PPFD) and CO₂ concentration conditions across 9 plant species including maize (Zea mays), sorghum (Sorghum vulgare), millet (Setaria italica), soybean (Glycine max), peanut (Arachis phyogaea), sweet potato (Ipomoea batatas), rice (Oryza sativa), Masson pine (Pinus massoniana) and Schima superba. We had developed a new model to estimate the water use efficiency in response to the combined effects of light and CO₂ concentration. Our measured data validated that this model could simulate the changes of water use efficiency very well under combined effect of light and CO₂ concentration. It could be used to estimate contribution of photosynthesis increase and transpiration decline on water use efficiency with the rising of CO₂ concentration. Great differences in water use efficiency occurred in these different plant species under various CO₂ concentration levels. Based on water use efficiency at regional scale, we concluded that plants should be separated into C₃ plants and C₄ plants, and furthermore, C₃ plants should be separated into herbaceous plants and woody plants. Our separation criteria would do a great favor in modeling the evapotranspiration of terrestrial ecosystem with carbon and water balance.     

不同植物叶片水分利用效率对光和CO2的响应与模拟

植物叶片水分利用效率的高低取决于气孔控制的光合作用和蒸腾作用两个相互耦合的过程,模拟水分利用效率对环境变化的响应特征和机制是理解生态系统碳循环和水循环及其耦合关系的基础.研究通过人工控制光强和CO2浓度,对叶片水分利用效率进行了研究.提出了植物水分利用效率在光强和CO2浓度共同作用下的估算模型.数据分析表明,该模型在包括C3和C4植物、草本和木本植物在内的9种植物上能很好地模拟水分利用效率对光强和CO2浓度共同作用的响应.该模型可以用于估算CO2浓度升高条件下光合速率的提高和蒸腾速率的降低对水分利用效率提高的贡献量.CO2浓度变化条件下,水分利用效率在不同植物之间有巨大差异,研究区域尺度植物的水分利用效率时至少需要将植物区分为C4植物和C3植物,其中C3植物区分为草本和木本植物3种生态功能型才能较为准确地估算植物的整体水分利用效率.应用本研究提出的水分利用效率估算模型和植物水分利用效率生态功能型分类标准,可以为建立以植物的水分利用效率为基本参数的陆地生态系统水循环模型和陆地生态系统生产力模型提供重要依据.     

不同植物叶片水分利用效率对光和CO2的响应与模拟

植物叶片水分利用效率的高低取决于气孔控制的光合作用和蒸腾作用两个相互耦合的过程,模拟水分利用效率对环境变化的响应特征和机制是理解生态系统碳循环和水循环及其耦合关系的基础。研究通过人工控制光强和CO₂浓度,对叶片水分利用效率进行了研究。提出了植物水分利用效率在光强和CO₂浓度共同作用下的估算模型。数据分析表明,该模型在包括C3和C4植物、草本和木本植物在内的9种植物上能很好地模拟水分利用效率对光强和CO₂浓度共同作用的响应。该模型可以用于估算CO₂浓度升高条件下光合速率的提高和蒸腾速率的降低对水分利用效率提高的贡献量。CO₂浓度变化条件下,水分利用效率在不同植物之间有巨大差异,研究区域尺度植物的水分利用效率时至少需要将植物区分为C4植物和C3植物,其中C3植物区分为草本和木本植物3种生态功能型才能较为准确地估算植物的整体水分利用效率。应用本研究提出的水分利用效率估算模型和植物水分利用效率生态功能型分类标准,可以为建立以植物的水分利用效率为基本参数的陆地生态系统水循环模型和陆地生态系统生产力模型提供重要依据。     

Leaf photosynthetic light response: a mechanistic model for scaling photosynthesis to leaves and canopies

1. The response of photosynthesis to radiation is an often-studied but poorly understood process, represented empirically in most photosynthesis models. However, in scaling photosynthesis from leaf to canopy, predictions of canopy photosynthesis are very sensitive to parameters describing the response of leaves to Photosynthetic Photon Flux Density (PPFD).
2. In this study, a mechanistic, yet still simple, approach is presented that models the degree of light saturation in leaves explicitly, assuming a heterogeneous environment of PPFD and chlorophyll.
3. Possible mechanisms determining the ratio of chlorophyll to nitrogen are considered, including a direct dependence on PPFD, a mechanism involving the red/far-red ratio of light in the canopy, and an approach based upon maximizing photosynthesis.
4. Comparison of model predictions with two data sets of light, nitrogen and chlorophyll from canopies of Populus and Corylus suggests that the red/far-red mechanism is the most realistic. The data also show that the trees studied do not always optimize their nitrogen partitioning to maximize photosynthetic yield.
5. We then apply the model to the data sets, to predict the shape of light response curves of leaves within canopies and assess the applicability of simple scaling schemes, in which full acclimation of photosynthesis to PPFD justifies the use of big-leaf models. We conclude that, at least for the data used, basic assumptions of such schemes do not hold.     

两种五倍子蚜虫冬寄主藓类植物的光合特性及其与光照、温度和植物体水分含量变化的关系

使用CI-301PS(CID Inc.USA)对生长于冬季和春季的大羽藓(Thuidium cymbifolium)与垂藓(Chrysocladium retrorsum)的净光合速率(Pn)及其与光照、温度和植物体水分含量的关系进行了研究.结果表明,大羽藓和垂藓的光合能力分别达到141和117μmolCO₂kg^-1·dw·s^-1,光合能力从冬季到春季呈上升趋势.这两种藓类植物的光响应曲线比较相似,光饱和点高达800～900μmol·m^-2s^-1,光补偿点为40～50μmol·m^-2s^-1,光合作用最适温度在春季为25～36℃,而冬季为20～30℃.同时对于低温又具有很强的抗性,在冰点以下的温度条件下(-15～0℃)能够保持一定的净光合.净光合速率的水分响应曲线表明,这两种藓类的最适水分含量为200～300(400)%dw,水分含量降低到150%dw时光合作用开始受到抑制,在40%～50%dw时净光合降低到零或略变为负数.结果表明,大羽藓和垂藓都属于耐干旱、强阳性的藓类,但垂藓在这几个方面略逊于大羽藓.     

Gas exchange and photosynthetic water use efficiency in response to light, CO2 concentration and temperature in Vicia faba

Light and temperature-response curves and their resulting coefficients, obtained within ecophysiological characterization of gas exchanges at the leaf level, may represent useful criteria for breeding and cultivar selection and required tools for simulation models aimed at the prediction of potential plant behaviour in response to environmental conditions.

Leaf-scale gas exchanges, by means of an IRGA open-flow system, were measured in response to light intensity (8 levels from 0 up to 2000 μmol m⁻² s⁻¹), CO₂ concentrations (ambient—350 μmol mol⁻¹ and short-term enriched—700 μmol mol⁻¹) and air temperature (from 7 up to 35 °C) on three Vicia faba L. genotypes, each representing one of the three cultivated groups: major, equina and minor. The net assimilation rate response to light intensity was well described by an exponential rise to max function. The short-term CO₂ enrichment markedly increased the values of light response curve parameters such as maximum photosynthetic rate (+80%), light saturation point (+40%) and quantum yield (+30%), while less homogenous behaviour was reported for dark respiration and light compensation point. For each light intensity level, the major and minor genotypes studied showed assimilation rates at least a 30% higher than equina.

The positive effects of short-term CO₂ enrichment on photosynthetic water use efficiency (WUE) indicate a relevant advantage in doubling CO₂ concentration. In the major and minor genotypes studied, similar assimilation rates, but different WUE were observed.

The optimum leaf temperature for assimilation process, calculated through a polynomial function, was 26–27 °C and no relevant limitations were observed in the range between 21 and 32 °C.

Analysis at the single leaf level provided both rapid information on the variations in gas exchange in response to environmental factors and selection criteria for the screening of genotypes.     

Alterations of pigment composition and their interactions in response to different light conditions in the diatom Chaetoceros gracilis probed by time-resolved fluorescence spectroscopy

Maintenance of energy balance under changeable light conditions is an essential function of photosynthetic organisms to achieve efficient photochemical reactions. Among the photosynthetic organisms, diatoms possess light-harvesting fucoxanthin chlorophyll (Chl) a/c-binding protein (FCP) as peripheral antennas. However, how diatoms regulate excitation-energy distribution between FCP and the two photosystem cores during light adaptation is poorly understood. In this study, we examined spectroscopic properties of a marine diatom Chaetoceros gracilis adapted in the dark and at photosynthetic photon flux density at 30 and 300?μmol?photons?m^?2?s^?1. Absorption spectra at 77?K showed significant changes in the Soret region, and 77-K steady-state fluorescence spectra showed significant differences in the spectral shape and relative fluorescence intensity originating from both PSII and PSI, among the cells grown under different light conditions. These results suggest alterations of pigment composition and their interactions under the different light conditions. These alterations affected the excitation-energy dynamics monitored by picosecond time-resolved fluorescence analyses at 77?K significantly. The contributions of Chls having lower energy levels than the reaction center Chls in the two photosystems to the energy dynamics were clearly identified in the three cells but with presumably different roles. These findings provide insights into the regulatory mechanism of excitation-energy balance in diatoms under various light conditions.     

PROTON GRADIENT REGULATION 5 contributes to ferredoxin-dependent cyclic phosphorylation in ruptured chloroplasts

Antimycin A-sensitive cyclic electron flow (CEF) was discovered as cyclic phosphorylation by Arnon et al. (1954). Because of its sensitivity to antimycin A, PROTON GRADIENT REGULATION 5 (PGR5)/PGR5-like Photosynthetic Phenotype 1 (PGRL1)-dependent CEF has been considered identical to the CEF of Arnon et al. However, this conclusion still needs additional supportive evidence, mainly because of the absence of definitive methods of evaluating CEF activity. In this study, we revisited the classical method of monitoring cyclic phosphorylation in ruptured chloroplasts to characterize two Arabidopsis mutants: pgr5, which is defective in antimycin A-sensitive CEF, and chlororespiratory reduction 2-1 (crr2-1), which is defective in chloroplast NDH-dependent CEF. We observed a significant reduction in CEF-dependent pmf formation and consequently ATP synthesis in the pgr5 mutant, although LEF-dependent pmf formation and ATP synthesis were not impaired at photosynthetic photon flux densities below 130?μmol?m^?2?s^?1. In contrast, the contribution of chloroplast NDH complex to pmf formation and ATP synthesis was not significant. Antimycin A partially inhibited CEF-dependent pmf formation, although there may be further inhibition sites. Unlike in the observation in leaves, the proton conductivity of ATP synthase, monitored as g_H⁺, was not enhanced in ruptured chloroplasts of the pgr5 mutant.     

PS II model based analysis of transient fluorescence yield measured on whole leaves of Arabidopsis thaliana after excitation with light flashes of different energies

Our recently presented PS II model (Belyaeva et al., 2008) was improved in order to permit a consistent simulation of Single Flash Induced Transient Fluorescence Yield (SFITFY) traces that were earlier measured by Steffen et al. (2005) on whole leaves of Arabidopsis (A.) thaliana at four different energies of the actinic flash. As the essential modification, the shape of the actinic flash was explicitly taken into account assuming that an exponentially decaying rate simulates the time dependent excitation of PS II by the 10 ns actinic flash. The maximum amplitude of this excitation exceeds that of the measuring light by 9 orders of magnitude. A very good fit of the SFITFY data was achieved in the time domain from 100 ns to 10 s for all actinic flash energies (the maximum energy of 7.5 × 10¹⁶ photons/(cm² flash) is set to 100%, the relative energies of weaker actinic flashes were of ∼8%, 4%, ∼1%). Our model allows the calculation and visualization of the transient PS II redox state populations ranging from the dark adapted state, via excitation energy and electron transfer steps induced by pulse excitation, followed by final relaxation into the stationary state eventually attained under the measuring light. It turned out that the rate constants of electron transfer steps are invariant to intensity of the actinic laser flash. In marked contrast, an increase of the actinic flash energy by more than two orders of magnitude from 5.4 × 10¹⁴ photons/(cm² flash) to 7.5 × 10¹⁶ photons/(cm² flash), leads to an increase of the extent of fluorescence quenching due to carotenoid triplet (³Car) formation by a factor of 14 and of the recombination reaction between reduced primary pheophytin (Phe⁻) and P680⁺ by a factor of 3 while the heat dissipation in the antenna complex remains virtually constant.The modified PS II model offers new opportunities to compare electron transfer and dissipative parameters for different species (e.g. for the green algae and the higher plant) under varying illumination conditions.