植物光合作用模型参数的温度依存性研究进展

综述了植物光合作用与温度响应模型研究的进展,围绕光合作用生化模型的4个主要参数：胞间CO₂浓度、RuBP最大碳同化速率(V_{c max})的活化能、RuBP最大再生速率(J_max)的活化能和J_max/V_{c max},讨论了影响光合作用 温度响应曲线的内在机理.随着生长温度的升高,所有物种的V_{c max}活化能均呈增加趋势,而其他参数的变化因物种不同而存在明显差异,说明V_{c max}的活化能可能是决定光合作用温度依存性的首要参数.最后分析了研究中存在的问题并提出研究展望,认为应整合叶片与群落水平的光合作用模型,从叶面积、太阳辐射、冠层结构、冠层小气候和光合能力等方面研究植物群落对全球变化的响应机理.这对于人们理解和准确估算植物生长、群落碳收支和生态系统初级生产力具有重要意义.     

Seasonal changes in temperature dependence of photosynthetic rate in rice under a free-air CO(2) enrichment

BACKGROUND AND AIMS: Influences of rising global CO(2) concentration and temperature on plant growth and ecosystem function have become major concerns, but how photosynthesis changes with CO(2) and temperature in the field is poorly understood. Therefore, studies were made of the effect of elevated CO(2) on temperature dependence of photosynthetic rates in rice (Oryza sativa) grown in a paddy field, in relation to seasons in two years. METHODS: Photosynthetic rates were determined monthly for rice grown under free-air CO(2) enrichment (FACE) compared to the normal atmosphere (570 vs 370 micromol mol(-1)). Temperature dependence of the maximum rate of RuBP (ribulose-1,5-bisphosphate) carboxylation (V(cmax)) and the maximum rate of electron transport (J(max)) were analysed with the Arrhenius equation. The photosynthesis-temperature response was reconstructed to determine the optimal temperature (T(opt)) that maximizes the photosynthetic rate. KEY RESULTS AND CONCLUSIONS: There was both an increase in the absolute value of the light-saturated photosynthetic rate at growth CO(2) (P(growth)) and an increase in T(opt) for P(growth) caused by elevated CO(2) in FACE conditions. Seasonal decrease in P(growth) was associated with a decrease in nitrogen content per unit leaf area (N(area)) and thus in the maximum rate of electron transport (J(max)) and the maximum rate of RuBP carboxylation (V(cmax)). At ambient CO(2), T(opt) increased with increasing growth temperature due mainly to increasing activation energy of V(cmax). At elevated CO(2), T(opt) did not show a clear seasonal trend. Temperature dependence of photosynthesis was changed by seasonal climate and plant nitrogen status, which differed between ambient and elevated CO(2).     

三种高山杜鹃的光合生理生态研究

对大白花杜鹃（Rhododendron decorum）、云南杜鹃（R.yunnanense）和红棕杜鹃（R.rubiginosum）进行了气体交换、叶片性状等研究,以期了解三种杜鹃的光合生理特性及其对环境的适应。结果表明,三种杜鹃的光饱和光合速率（Pmax）与RuBP饱和最大羧化速率（Vc max）、光饱和最大电子传递速率（Jmax）和气孔导度（gs）呈极显著正相关（P≤0.01）,但仅Vc max存在显著的种间差异,说明三种杜鹃的光合能力主要受Vc max影响。叶氮含量、叶片氮在电子传递和在Rubisco中的分配均显著影响Vc max和Jmax。大白花杜鹃的LSP最低,LCP较高,对强光和弱光利用能力都不强,光适应范围较窄。云南杜鹃LCP最低,LSP和Pmax相对较高,对弱光或较强的光照均能利用,光照适应范围相对最广,光合适应能力最强;红棕杜鹃LSP和LCP均为最高,对强光环境的适应性最强。     

Growth temperature can alter the temperature dependent stimulation of photosynthesis by elevated carbon dioxide in Albutilon theophrasti

Stimulation of photosynthesis in response to elevated carbon dioxide concentration [CO2] in the short-term (min) should be highly temperature dependent at high photon flux. However, it is unclear if long-term (days, weeks) adaptation to a given growth temperature alters the temperature-dependent stimulation of photosynthesis to [CO2]. In velveltleaf (Albutilon theophrasti), the response of photosynthesis, determined as CO2 assimilation, was measured over a range of internal CO2 concentrations at 7 short-term measurement (12, 16, 20, 24, 28, 32, 36 degrees C) temperatures for each of 4 long-term growth (16, 20, 28 and 32 degrees C) temperatures. In vivo estimates of VCmax, the maximum RuBP saturated rate of carboxylation, and Jmax, the light-saturated rate of potential electron transport, were determined from gas exchange measurements for each temperature combination. Overall, previous exposure to a given growth temperature adjusted the optimal temperatures of Jmax and VCmax with subsequently greater enhancement of photosynthesis at elevated [CO2] (i.e., a greater enhancement of photosynthesis at elevated [CO2] was observed at low measurement temperatures for A. theophrasti grown at low growth temperatures compared with higher growth temperatures, and vice versa for plants grown and measured at high temperatures). Previous biochemical based models used to predict the interaction between rising [CO2] and temperature on photosynthesis have generally assumed no growth temperature effect on carboxylation kinetics or no limitation by Jmax. In the current study, these models over predicted the temperature dependence of the photosynthetic response to elevated [CO2] at temperatures above 24 degrees C. If these models are modified to include long-term adjustments of Jmax and VCmax to growth temperature, then greater agreement between observed and predicted values was obtained.     

Variation in acclimation of photosynthesis in Trifolium repens after eight years of exposure to Free Air CO2 Enrichment (FACE)

The initial stimulation of photosynthesis observed on elevation of [CO2] in grasslands has been predicted to be a transient phenomenon constrained by the loss of photosynthetic capacity due to other limitations, notably nutrients and sinks for carbohydrates. Legumes might be expected partially to escape these feedbacks through symbiotic N2 fixation. The Free-Air Carbon dioxide Enrichment (FACE) experiment at Eschikon, Switzerland, has been the longest running investigation of the effects of open-air elevation of [CO2] on vegetation. The prediction of a long-term loss of photosynthetic capacity was tested by analysing photosynthesis in Trifolium repens L. (cv. Milkanova) in the spring and autumn of the eighth, ninth and tenth years of treatment. A high and low N treatment also allowed a test of the significance of exogenous N-supply in maintaining a stimulation of photosynthetic capacity in the long-term. Prior work in this Free Air CO2 Enrichment (FACE) experiment has revealed that elevated [CO2] increased both vegetative and reproductive growth of T. repens independent of N treatment. It is shown here that the photosynthetic response of T. repens was also independent of N fertilization under both current ambient and elevated (600 micro mol mol-1) [CO2]. There was a strong effect of season on photosynthesis, with light-saturated rates (Asat) 37% higher in spring than in autumn. Higher Asat in the spring was supported by higher maximum Rubisco carboxylation rates (Vc,max) and maximum rates of electron transport (Jmax) contributing to RuBP regeneration. Elevated [CO2] increased Asat by 37% when averaged across all measurement periods and both N fertilization levels, and decreased stomatal conductance by 25%. In spring, there was no effect of elevated [CO2] on photosynthetic capacity of leaves, but in autumn both Vc,max and Jmax were reduced by approximately 20% in elevated [CO2]. The results show that acclimation of photosynthetic capacity can occur in a nitrogen-fixing species, in the field where there are no artificial restrictions on sink capacity. However, even with acclimation there was a highly significant increase in photosynthesis at elevated [CO2].     

Seasonal change in the balance between capacities of RuBP carboxylation and RuBP regeneration affects CO2 response of photosynthesis in Polygonum cuspidatum

The balance between the capacities of RuBP (ribulose-1,5-bisphosphate) carboxylation (V(cmax)) and RuBP regeneration (expressed as the maximum electron transport rate, J(max)) determines the CO(2) dependence of the photosynthetic rate. As it has been suggested that this balance changes depending on the growth temperature, the hypothesis that the seasonal change in air temperature affects the balance and modulates the CO(2) response of photosynthesis was tested. V(cmax) and J(max) were determined in summer and autumn for young and old leaves of Polygonum cuspidatum grown at two CO(2) concentrations (370 and 700 micromol mol(-1)). Elevated CO(2) concentration tended to reduce both V(cmax) and J(max) without changing the J(max):V(cmax) ratio. The seasonal environment, on the other hand, altered the ratio such that the J(max):V(cmax) ratio was higher in autumn leaves than summer leaves. This alternation made the photosynthetic rate more dependent on CO(2) concentration in autumn. Therefore, when photosynthetic rates were compared at growth CO(2) concentration, the stimulation in photosynthetic rate was higher in young-autumn than in young-summer leaves. In old-autumn leaves, the stimulation of photosynthesis brought by a change in the J(max):V(cmax) ratio was partly offset by accelerated leaf senescence under elevated CO(2). Across the two seasons and the two CO(2) concentrations, V(cmax) was strongly correlated with Rubisco and J(max) with cytochrome f content. These results suggest that seasonal change in climate affects the relative amounts of photosynthetic proteins, which in turn affect the CO(2) response of photosynthesis.     

柚树叶片CO2驯化的光合参数变化

柚树(Citrus grandis)幼树生长在砂和磋石的生长介质,每周供给0.05mmol P(正常P,P)和0．1mmol P(高磷,2P)的营养液．植株分别生长在空气CO2分压(约39Pa)和倍增CO2分压(81±5Pa)下45d,利用CI-301PS(CID,Inc)光合作用测定系统在较高光强(1150μmol·m^-2·s^-1)下测定叶片光合速率并得出的Pn-Pi关系曲线和在较高CO2分压(PCO2,56Pa)下得出Pn-PAR关系曲线计算有关光合参数。结果表明,大气CO2分压下2P植株最大光合速率较P植株高13．3％,倍增CO2分压下,无论P或2P植株最大光合速率较大气CO2分压下相应植株低,但在倍增CO2分压下2P植株较P植株高,且2P植株有较P植株高的表观量子产率和光能利用效率(P<0．05),但并不改变г^*、Rd和Rubisco羧化速率(Vc)和氧速率的比率(P>0．05)在大气CO2分压下2P植株的Vcmax和Jmax较P植株分别高83％和12．5％,在倍增CO2分压下2P植株的Vcmax和Jmax均较P植株高,柚树在高CO2驯化中改变叶N在Rubisco和捕光组分分配系数,但不改变叶N在光合电子传递链的分配系数,结果表明,增加P供给可以促进高CO2分压下光合碳循环中P的周转,提高倍增CO2分压下植株的光合速率,调节柚树叶片的CO2驯化的光合参数。     

Temperature dependence of leaf-level CO2 fixation: revising biochemical coefficients through analysis of leaf three-dimensional structure

CO2 fixation in a leaf is determined by biochemical and physical processes within the boundaries set by leaf structure. Traditionally determined temperature dependencies of biochemical processes include physical processes related to CO2 exchange that result in inaccurate estimates of parameter values. A realistic three-dimensional model of a birch (Betula pendula) leaf was used to distinguish between the physical and biochemical processes affecting the temperature dependence of CO2 exchange, to determine new chloroplastic temperature dependencies for V c(max) and Jmax based on experiments, and to analyse mesophyll diffusion in detail. The constraint created by dissolution of CO2 at cell surfaces substantially decreased the CO2 flux and its concentration inside chloroplasts, especially at high temperatures. Consequently, newly determined chloroplastic V c(max) and Jmax were more temperature dependent than originally. The role of carbonic anhydrase in mesophyll diffusion appeared to be minor under representative mid-day nonwater-limited conditions. Leaf structure and physical processes significantly affect the apparent temperature dependence of CO2 exchange, especially at optimal high temperatures when the photosynthetic sink is strong. The influence of three-dimensional leaf structure on the light environment inside a leaf is marked and affects the local choice between Jmax and V c(max)-limited assimilation rates.     

Modelling photosynthetic responses to temperature of grapevine (Vitis vinifera cv. Semillon) leaves on vines grown in a hot climate

Field measurements of photosynthesis of Vitis vinifera cv. Semillon leaves in relation to a hot climate, and responses to photon flux densities (PFDs) and internal CO(2) concentrations (c(i) ) at leaf temperatures from 20 to 40 °C were undertaken. Average rates of photosynthesis measured in situ decreased with increasing temperature and were 60% inhibited at 45 °C compared with 25 °C. This reduction in photosynthesis was attributed to 15-30% stomatal closure. Light response curves at different temperatures revealed light-saturated photosynthesis optimal at 30 °C but also PFDs saturating photosynthesis increased from 550 to 1200 μmol (photons) m(-2)s(-1) as temperatures increased. Photosynthesis under saturating CO(2) concentrations was optimal at 36 °C while maximum rates of ribulose 1,5-bisphosphate (RuBP) carboxylation (V(cmax)) and potential maximum electron transport rates (J(max)) were also optimal at 39 and 36 °C, respectively. Furthermore, the high temperature-induced reduction in photosynthesis at ambient CO(2) was largely eliminated. The chloroplast CO(2) concentration at the transition from RuBP regeneration to RuBP carboxylation-limited assimilation increased steeply with an increase in leaf temperature. Semillon assimilation in situ was limited by RuBP regeneration below 30 °C and above limited by RuBP carboxylation, suggesting high temperatures are detrimental to carbon fixation in this species.     

Interactive effects of elevated CO2, warming, and drought on photosynthesis of Deschampsia flexuosa in a temperate heath ecosystem

Global change factors affect plant carbon uptake in concert. In order to investigate the response directions and potential interactive effects, and to understand the underlying mechanisms, multifactor experiments are needed. The focus of this study was on the photosynthetic response to elevated CO(2) [CO2; free air CO(2) enrichment (FACE)], drought (D; water-excluding curtains), and night-time warming (T; infrared-reflective curtains) in a temperate heath. A/C(i) curves were measured, allowing analysis of light-saturated net photosynthesis (P(n)), light- and CO(2)-saturated net photosynthesis (P(max)), stomatal conductance (g(s)), the maximal rate of Rubisco carboxylation (V(cmax)), and the maximal rate of ribulose bisphosphate (RuBP) regeneration (J(max)) along with leaf δ(13)C, and carbon and nitrogen concentration on a monthly basis in the grass Deschampsia flexuosa. Seasonal drought reduced P(n) via g(s), but severe (experimental) drought decreased P(n) via a reduction in photosynthetic capacity (P(max), J(max), and V(cmax)). The effects were completely reversed by rewetting and stimulated P(n) via photosynthetic capacity stimulation. Warming increased early and late season P(n) via higher P(max) and J(max). Elevated CO(2) did not decrease g(s), but stimulated P(n) via increased C(i). The T×CO2 synergistically increased plant carbon uptake via photosynthetic capacity up-regulation in early season and by better access to water after rewetting. The effects of the combination of drought and elevated CO(2) depended on soil water availability, with additive effects when the soil water content was low and D×CO2 synergistic stimulation of P(n) after rewetting. The photosynthetic responses appeared to be highly influenced by growth pattern. The grass has opportunistic water consumption, and a biphasic growth pattern allowing for leaf dieback at low soil water availability followed by rapid re-growth of active leaves when rewetted and possibly a large resource allocation capability mediated by the rhizome. This growth characteristic allowed for the photosynthetic capacity up-regulations that mediated the T×CO2 and D×CO2 synergistic effects on photosynthesis. These are clearly advantageous characteristics when exposed to climate changes. In conclusion, after 1 year of experimentation, the limitations by low soil water availability and stimulation in early and late season by warming clearly structure and interact with the photosynthetic response to elevated CO(2) in this grassland species.     

Temperature acclimation in a biochemical model of photosynthesis: a reanalysis of data from 36 species

The Farquhar et al. model of C(3) photosynthesis is frequently used to study the effect of global changes on the biosphere. Its two main parameters representing photosynthetic capacity, V(cmax) and J(max), have been observed to acclimate to plant growth temperature for single species, but a general formulation has never been derived. Here, we present a reanalysis of data from 36 plant species to quantify the temperature dependence of V(cmax) and J(max) with a focus on plant growth temperature, i.e. the plants' average ambient temperature during the preceding month. The temperature dependence of V(cmax) and J(max) within each data set was described very well by a modified Arrhenius function that accounts for a decrease of V(cmax) and J(max) at high temperatures. Three parameters were optimized: base rate, activation energy and entropy term. An effect of plant growth temperature on base rate and activation energy could not be observed, but it significantly affected the entropy term. This caused the optimum temperature of V(cmax) and J(max) to increase by 0.44 degrees C and 0.33 degrees C per 1 degrees C increase of growth temperature. While the base rate of V(cmax) and J(max) seemed not to be affected, the ratio J(max) : V(cmax) at 25 degrees C significantly decreased with increasing growth temperature. This moderate temperature acclimation is sufficient to double-modelled photosynthesis at 40 degrees C, if plants are grown at 25 degrees C instead of 17 degrees C.     

Gas exchange measurements, what can they tell us about the underlying limitations to photosynthesis? Procedures and sources of error

The principles, equipment and procedures for measuring leaf and canopy gas exchange have been described previously as has chlorophyll fluorescence. Simultaneous measurement of the responses of leaf gas exchange and modulated chlorophyll fluorescence to light and CO2 concentration now provide a means to determine a wide range of key biochemical and biophysical limitations on photo synthesis in vivo. Here the mathematical frameworks and practical procedures for determining these parameters in vivo are consolidated. Leaf CO2 uptake (A) versus intercellular CO2 concentration (Ci) curves may now be routinely obtained from commercial gas exchange systems. The potential pitfalls, and means to avoid these, are examined. Calculation of in vivo maximum rates of ribulose-1,5-bisphosphate (RuBP) carboxylase/oxygenase (Rubisco) carboxylation (Vc,max), electron transport driving regeneration of RuBP (Jmax), and triose-phosphate utilization (VTPU) are explained; these three parameters are now widely assumed to represent the major limitations to light-saturated photosynthesis. Precision in determining these in intact leaves is improved by the simultaneous measurement of electron transport via modulated chlorophyll fluorescence. The A/Ci response also provides a simple practical method for quantifying the limitation that stomata impose on CO2 assimilation. Determining the rate of photorespiratory release of oxygen (Rl) has previously only been possible by isotopic methods, now, by combining gas exchange and fluorescence measurements, Rl may be determined simply and routinely in the field. The physical diffusion of CO2 from the intercellular air space to the site of Rubisco in C3 leaves has long been suspected of being a limitation on photosynthesis, but it has commonly been ignored because of the lack of a practical method for its determination. Again combining gas exchange and fluorescence provides a means to determine mesophyll conductance. This method is described and provides insights into the magnitude and basis of this limitation.     

The regulation of Rubisco activity in response to variation in temperature and atmospheric CO2 partial pressure in sweet potato

The temperature response of net CO(2) assimilation rate (A), the rate of whole-chain electron transport, the activity and activation state of Rubisco, and the pool sizes of ribulose-1,5-bisphosphate (RuBP) and 3-phosphoglyceric acid (PGA) were assessed in sweet potato (Ipomoea batatas) grown under greenhouse conditions. Above the thermal optimum of photosynthesis, the activation state of Rubisco declined with increasing temperature. Doubling CO(2) above 370 mubar further reduced the activation state, while reducing CO(2) by one-half increased it. At cool temperature (<16 degrees C), the activation state of Rubisco declined at CO(2) levels where photosynthesis was unaffected by a 90% reduction in O(2) content. Reduction of the partial pressure of CO(2) at cool temperature also enhanced the activation state of Rubisco. The rate of electron transport showed a pronounced temperature response with the same temperature optimum as A at elevated CO(2). RuBP pool size and the RuBP-to-PGA ratio declined with increasing temperature. Increasing CO(2) also reduced the RuBP pool size. These results are consistent with the hypothesis that the reduction in the activation state of Rubisco at high and low temperature is a regulated response to a limitation in one of the processes contributing to the rate of RuBP regeneration. To further evaluate this possibility, we used measured estimates of Rubisco capacity, electron transport capacity, and the inorganic phosphate regeneration capacity to model the response of A to temperature. At elevated CO(2), the activation state of Rubisco declined at high temperatures where electron transport capacity was predicted to be limiting, and at cooler temperatures where the inorganic phosphate regeneration capacity was limiting. At low CO(2), where Rubisco capacity was predicted to limit photosynthesis, full activation of Rubisco was observed at all measurement temperatures.     

羊草(Leymus chinensis)叶片光合参数对干旱与复水的响应机理与模拟

利用典型草原优势植物羊草（Leymus chinensis）对不同水分胁迫与复水响应的植物光合生理生态模拟实验与野外观测资料,分析研究了羊草叶片光合参数Kcmax（Rubisco的最大羧化速率）、Jmax（最大光合电子传递速率）和TPU（磷酸丙糖利用率）对干旱与复水的响应机理。结果表明,无论是模拟实验还是野外观测均显示羊草叶片的光合参数随着土壤水分的增加呈抛物线曲线变化,但各光合参数最大值对土壤水分的响应不同。温室模拟下的羊草光合参数Vcmax,Jmax和TPU在土壤含水量分别在15．56％,15．89％和16．23％时达到最大,而野外观测羊草的光合参数Vcmax,Jmax和TPU在土壤含水量分别为16．89％,17％和16．79％时达到最大。复水后羊草植株叶片光合参数的变化取决于前期干旱的影响,土壤含水量18％～19％和15％～16％处理的羊草复水后光合参数能够恢复正常,前者甚至超过正常水平,说明适宜的水分胁迫在复水后能够提高羊草叶片的光合能力,促进光合作用;土壤含水量10％～12％和7％～9％处理下的羊草复水后光合参数则不能恢复到正常水平。土壤含水量15％～16％可能是羊草光合能力在水分胁迫后能否恢复的阈值。     

Balancing carboxylation and regeneration of ribulose-1,5- bisphosphate in leaf photosynthesis: temperature acclimation of an evergreen tree, Quercus myrsinaefolia

Changes in the temperature dependence of the photosynthetic rate depending on growth temperature were investigated for a temperate evergreen tree, Quercus myrsinaefolia . Plants were grown at 250 μ mol quanta m^–2 s^–1 under two temperature conditions, 15 and 30 °C. The optimal temperature that maximizes the light-saturated rate of photosynthesis at 350 μ L L^–1 CO₂ was found to be 20–25 and 30–35 °C for leaves grown at 15 and 30 °C, respectively. We focused on two processes, carboxylation and regeneration of ribulose-1,5-bisphosphate (RuBP), which potentially limit photosynthetic rates. Because the former process is known to limit photosynthesis at lower CO₂ concentrations while the latter limits it at higher CO₂ concentrations, we determined the temperature dependence of the photosynthetic rate at 200 and 1000 μ L L^–1 CO₂ under saturated light. It was revealed that the temperature dependence of both processes varied depending on the growth temperature. Using a biochemical model, we estimated the capacity of the two processes at various temperatures under ambient CO₂ concentration. It was suggested that, in leaves grown at low temperature (15 °C), the photosynthetic rate was limited solely by RuBP carboxylation under any temperature. On the other hand, it was suggested that, in leaves grown at high temperature (30 °C), the photosynthetic rate was limited by RuBP regeneration below 22 °C, but limited by RuBP carboxylation above 22 °C. We concluded that: (1) the changes in the temperature dependence of carboxylation and regeneration of RuBP and (2) the changes in the balance of these two processes altered the temperature dependence of the photosynthetic rate.     

增温背景下克氏针茅枯黄期物候对降水响应的光合生理机制

基于红外线辐射增温与控水相结合的原位模拟试验资料,分析了克氏针茅(Stipa krylovii)枯黄期对水热变化响应的光合生理机制。结果表明: 增温背景下降水是枯黄期的主要影响因子,增水(减水)导致枯黄始期和枯黄盛期的发生时间均延迟(提前),枯黄期持续时间均延长,减水处理对枯黄期持续时间的延长作用更显著。增温背景下,降水变化显著影响枯黄期的生理生态特性,且在枯黄始期最为显著,净光合速率、气孔导度、蒸腾速率、核酮糖-1,5-二磷酸(RuBP)羧化的最大速率(V_{c max})、RuBP再生能力的最大速率(J_max)均与降水量呈显著正相关。通径分析表明,克氏针茅枯黄期的光合生理机制因水热变化的不同而异。当前环境条件下,枯黄期物候变化的主要影响因子是J_max,主要限制因子是V_{c max}。未来暖干和暖湿气候下枯黄期物候变化的主要影响因子均是V_{c max};但在暖干气候下主要限制因子为J_max,而在暖湿气候下则无限制因子。这表明克氏针茅枯黄期物候的变化取决于气候环境条件变化对其光合能力的影响。     

海拔对高山栎光合气体交换和叶性状的影响

理解影响植物分布的式样及过程是生态学研究的中心内容之一,但对许多物种而言,限制其分布的原因还不清楚。为了认识高山栎分布与生理生态特性的关系,我们在不同海拔的4个观测点研究了帽斗栎的光合气体交换、叶氮含量、叶绿素含量和比叶重。由于高的水气压亏缺和气温,帽斗栎的光合作用和蒸腾作用在午间表现出明显的降低现象。帽斗栎的饱和光合速率、水分利用效率、最大羧化速率、最大电子传递速率和氮利用效率在海拔中部比低海拔或高海拔处的为高。不同海拔的叶氮含量在5月份有差异,8月份则没有明显不同。叶片厚度随海拔增加,但叶绿素含量及光合最适温度随海拔升高而降低。帽斗栎光合作用的海拔变化与叶片的生化效率和氮含量有关,而与比叶重无关。研究结果说明,温度的海拔变化对高山栎的光合作用和叶性状有明显影响,最适宜帽斗栎光合碳获取及生长的海拔范围是3180～3610m。     

Modelling Optimal Temperature Acclimation of the Photosynthetic Apparatus in C3Plants with Respect to Nitrogen Use

A new hypothesis for temperature acclimation by the photosyntheticapparatus is presented. An optimization model is developed toexamined effects of changes in the organization of photosyntheticcomponents on leaf photosynthesis under various growth temperatureswhere the photosynthetic apparatus is not damaged. In this model,photosynthetic rate is limited either by the capacity of ribulosebisphosphate carboxylase (RuBPCase) to consume ribulose bisphosphate(RuBP), or by the capacity of RuBP regeneration. For temperaturedependence of the RuBPCase activity, data fromSpinacia oleraceaL.,which have a temperature optimum of 30 °C, are used. Fortemperature dependence of the capacity of RuBP regeneration,two contrasting curves that have temperature optima of 30 °C(Eucalyptus paucifloraSieb. ex Spreng) and 40 °C (LarreadivaricataCav.) are applied. The temperature dependence of eachprocess is fixed for respective species, but the rate of eachprocess varies with changes in the amounts of components. Thecost of proteins, in terms of nitrogen, required to carry outeach process is calculated when nitrogen is partitioned differentlyamong photosynthetic components. The optimal nitrogen partitioningthat maximizes daily photosynthesis at a given temperature isobtained. The predicted temperature optimum of the photosyntheticrate inLarrea divaricataexhibits large shifts with changes intarget temperature, while shifts are negligible inEucalyptuspauciflora. It is suggested that the shift in temperature optimumof photosynthetic rate is large when the temperature dependencesof the capacities of RuBPCase and RuBP regeneration differ fromeach other.Copyright 1997 Annals of Botany Company Optimization model; nitrogen use efficiency; photosynthetic acclimation; temperature dependence     

Regulation of Ribulose-1,5-Bisphosphate Carboxylase Activity in Alocasia macrorrhiza in Response to Step Changes in Irradiance

The regulation of ribulose-1,5-bisphosphate (RuBP) carboxylase (Rubisco) activity and pool sizes of RuBP and P-glycerate were examined in the tropical understory species Alocasia macrorrhiza following step changes in photon flux density (PFD). Previous gas exchange analysis of this species following a step increase in PFD from 10 to 500 micromoles quanta per square meter per second suggested that the increase in photosynthetic rate was limited by the rate of increase of Rubisco activity for the first 5 to 10 minutes. We demonstrate here that the increase in photosynthetic rate was correlated with an increase in both the activation state of Rubisco and the total k_cat (fully activated specific activity) of the enzyme. Evidence presented here suggests that a change in the pool size of the naturally occurring tight binding inhibitor of Rubisco activity, 2-carboxyarabinitol 1-phosphate, was responsible for the PFD-dependent change in the total k_cat of the enzyme. RuBP pool size transiently increased after the increase in PFD, indicating that photosynthesis was limited by the capacity for carboxylation. After 5 to 10 minutes, RuBP pool size was again similar to the pool size at low PFD, presumably because of the increased activity of Rubisco. Following a step decrease in PFD from 500 to 10 micromoles quanta per square meter per second, Rubisco activity declined but at a much slower rate than it had increased in response to a step increase in PFD. This slower rate of activity decline than increase was apparently due to the slower rate of 2-carboxyarabinitol 1-phosphate synthesis than degradation and, to a lesser degree, to slower deactivation than activation. RuBP pool size initially declined following the decrease in PFD, indicating that RuBP regeneration was limiting photosynthesis. As Rubisco activity decreased, RuBP slowly increased to its original level at high PFD. The slow rate of activity loss by Rubisco in this species suggests a biochemical basis for the increased efficiency for CO₂ assimilation of successive lightfleck use by species such as A. macrorrhiza.     

Small decreases in SBPase cause a linear decline in the apparent RuBP regeneration rate, but do not affect Rubisco carboxylation capacity

The response of net photosynthetic CO(2) uptake (A) to increasing leaf intercellular CO(2) concentration (c(i)) was determined in antisense Nicotiana tabacum plants, derived from six independent transformation lines, displaying a range of sedoheptulose-1, 7-bisphosphatase (SBPase) activities. The maximum in vivo ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) carboxylation (V(c,max)) and RuBP regeneration (J(max)) rates were calculated from the steady-state measurements of the A to c(i) response curves. In plants with reductions in SBPase activity of between 9% and 60%, maximum RuBP regeneration capacity declined linearly (r(2)=0.79) and no significant change in apparent in vivo Rubisco activity (V(c,max)) was observed in these plants. No correlation between V(c,max) and a decrease in capacity for RuBP regeneration was observed (r(2)=0.14) in the SBPase antisense plants. These data demonstrate that small decreases in SBPase activity limit photosynthetic carbon assimilation by reducing the capacity for RuBP regeneration.