Global photosynthetic capacity is optimized to the environment

Earth system models (ESMs) use photosynthetic capacity, indexed by the maximum Rubisco carboxylation rate (V_cmax), to simulate carbon assimilation and typically rely on empirical estimates, including an assumed dependence on leaf nitrogen determined from soil fertility. In contrast, new theory, based on biochemical coordination and co‐optimization of carboxylation and water costs for photosynthesis, suggests that optimal V_cmax can be predicted from climate alone, irrespective of soil fertility. Here, we develop this theory and find it captures 64% of observed variability in a global, field‐measured V_cmax dataset for C₃ plants. Soil fertility indices explained substantially less variation (32%). These results indicate that environmentally regulated biophysical constraints and light availability are the first‐order drivers of global photosynthetic capacity. Through acclimation and adaptation, plants efficiently utilize resources at the leaf level, thus maximizing potential resource use for growth and reproduction. Our theory offers a robust strategy for dynamically predicting photosynthetic capacity in ESMs.     

Photosynthesis in Encelia farinosa Gray in Response to Decreasing Leaf Water Potential

Photosynthetic responses of intact leaves of the desert shrub Encelia farinosa were measured during a long term drought cycle in order to understand the responses of stomatal and nonstomatal components to water stress. Photosynthetic rate at high irradiance and leaf conductance to water vapor both decreased linearly with declining leaf water potential. The intercellular CO₂ concentration (c_i) remained fairly constant as a function of leaf water potential in plants subjected to a slow drought cycle of 25 days, but decreased in plants exposed to a 12-day drought cycle. With increasing water stress, the slope of the dependence of photosynthesis on c_i (carboxylation efficiency) decreased, the maximum photosynthetic rates at high c_i became saturated at lower values, and water use efficiency increased. Both the carboxylation efficiency and photosynthetic rates were positively correlated with leaf nitrogen content. Associated with lower leaf conductances, the calculated stomatal limitation to photosynthesis increased with water stress. However, because of simultaneous changes in the dependence of photosynthesis on c_i with water stress, increased leaf conductance alone in water-stressed leaves would not result in an increase in photosynthetic rates to prestressed levels. Both active osmotic adjustment and changes in specific leaf mass occurred during the drought cycle. In response to increased water stress, leaf specific mass increased. However, the increases in specific leaf mass were associated with the production of a reflective pubescence and there were no changes in specific mass of the photosynthetic tissues. The significance of these responses for carbon gain and water loss under arid conditions are discussed.     

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.     

Seasonal relationships between leaf nitrogen content (photosynthetic capacity) and leaf canopy light exposure in peach (Prunus persica)

Abstract Field gas exchange measurements on intact peach (Prunus persica (L.) Batsch) leaves indicate that leaf nitrogen content (N_L) and leaf weight per unit leaf area (W_a) are highly correlated with CO₂ assimilation rate (A) and mesophyll conductance (g_m). Therefore, N_L and W_a were used to study seasonal relationships between leaf carboxylation capacity and natural light exposure in tree canopies. From mid-season onwards, N_L and W_a were linearly correlated with light exposure expressed as the amount of time during a clear day that a leaf was exposed to a photosynthetic photon flux density (Q) of ≥ 100 μmol m^?2 s^?1. The data support the hypothesis that whole-tree photosynthesis is optimized by partitioning of photosynthetic capacity among leaves in deciduous tree canopies with respect to natural light exposure.     

Evolutionarily stable leaf area production in plant populations

Using an analytical model, it was shown that for a given amount of nitrogen in the canopy of a stand (N(T)), there exists an evolutionarily stable leaf area index (ES-LAI), and therefore an evolutionarily stable average leaf nitrogen content (n(ES)(av);n(ES)(av) =N(T)/ES-LAI), at which no individual plant in the stand can increase its photosynthesis by changing its leaf area. It was also shown that this ES-LAI is always greater than the optimal LAI that maximizes photosynthesis per unit N(T) of the stand. This illustrates that the canopy structure that maximizes photosynthesis of a population is not the same as the canopy structure that maximizes photosynthesis of individuals within a population. It was further derived that the ES-LAI at given N(T) increases with the ratio between the light-saturated photosynthesis and the N content per unit leaf area (leaf-PPNUE) and that it decreases with the canopy extinction coefficient for light (K(L)), the light availability and the apparent quantum yield (phi). These hypotheses were tested by comparing calculated ES-LAI and n(ES)(av) values to actual LAIs and leaf N contents measured for stands of a large variety of herbaceous plants. There was a close correspondence between the calculated and measured values. As predicted by the model, plants with high leaf-PPNUEs produced more leaf area per unit nitrogen than those with low leaf-PPNUEs while plants with horizontal leaves, forming stands with higher K(L) values, produced less leaf area than those with more vertically inclined leaves. These results suggest that maximization of individual plant photosynthesis per unit of nitrogen plays an important role in determining leaf area production of plants and the resulting canopy structure of stands of vegetation. They further suggest this optimization to be a mechanism by which leaf traits such as leaf-PPNUE and leaf inclination angle are causally related to structural characteristics of the population, i.e. the leaf area index of the stand.     

Photosynthetic capacity of field-grown durum wheat under different N availabilities: A comparative study from leaf to canopy

The effect of N availability on photosynthetic capacity, growth parameters and yield was studied in field-grown durum-wheat plants at both the leaf and canopy levels. Two contrasting nitrogen levels (120 and 0 kg ha^?1) were assayed in a randomised block design with nine replicates each. Total biomass was measured at anthesis and yield and its agronomical components at maturity. Photosynthetic measurements were performed 2 weeks after anthesis in two plots of each N treatment. Flag leaves were measured, using a LI-COR 6400 combined with the chlorophyll fluorescence meter, and the whole canopy by measuring CO₂ and H₂O fluxes in an innovative canopy-chamber system. We showed a clear increase in photosynthetic gas exchange and chlorophyll contents with N fertilisation at both canopy and leaf levels. As a consequence the increase in yield as response to N fertilisation seems the result of a larger green leaf area combined with a higher photosynthetic capacity of the leaves attributable to an increase in the maximum carboxylation velocity of Rubisco. Moreover gas-exchange measurements of the flag leaf during grain filling seem to provide a realistic characterisation, not just of the photosynthetic performance of the crop, but also about the impact of N availability on yield. Thus, measurements performed on the flag leaf matched those at the canopy level, with proportional increases in terms of gas exchange and chlorophyll content, providing a fast, cheap and reliable estimation of canopy photosynthesis and the grain yield attained by the crop.     

Nitrogen allocation and partitioning in invasive and native Eupatorium species

There is a trade-off between nitrogen (N) allocation to photosynthesis and to defence. Invasive species may reduce N allocation to defence because of the absence of natural enemies. Thus, I hypothesised that invasive species may allocate a higher fraction of total leaf N to photosynthesis and have higher light-saturated photosynthetic rate ( P _max) and photosynthetic N-use efficiency (PNUE) than closely related native species. To test these hypotheses, invasive Eupatorium adenophorum and native E.   chinense and E.   heterophyllum were compared in a limestone shrub. Unlike expectation, the invader did not allocate a higher fraction of leaf N to photosynthesis than the natives. However, it was more efficient in photosynthetic N partitioning than the natives. It partitioned a higher fraction of the photosynthetic N to carboxylation and showed higher use efficiency of the photosynthetic N, while the natives partitioned a higher fraction of the photosynthetic N to light-harvesting components. Total leaf N content was not significantly different among the three studied invasive and native species. For the invader, the higher fraction of leaf N allocated to carboxylation resulted in the higher N content in carboxylation and in both carboxylation and bioenergetics, which led to higher P _max, and therefore to higher PNUE, water-use efficiency, respiration efficiency and apparent quantum yield. These physiological advantages of the invader and its higher leaf area ratio may contribute to its invasiveness.     

Canopy nitrogen distribution and the photosynthetic performance of sunflower crops during grain filling — a quantitative analysis

Measurements of the profiles of leaf area and leaf nitrogen were made on five occasions from midflowering to maturity (53, 61, 70, 78 and 83 days after emergence, DAE) in sunflower crops grown at contrasting density (2.4 and 4.8 plants m^-2) and nitrogen supply (0 and 5 g N m^-2 at emergence) in the summer in Buenos Aires, Argentina. As the crops matured, nitrogen was withdrawn unequally from all leaf positions and leaves senesced from the bases of the canopies. A model was used to estimate the daytime net photosynthesis (P_c) of canopies of defined leaf area and nitrogen content under the observed conditions of temperature and irradiance. Comparisons were made between the observed profiles of leaf nitrogen and those that would maximise P_c (the optimal profiles). The observed nitrogen profiles were sub-optimal at mid-flowering, except in the low-density, low-nitrogen treatment. The differences were most marked in the high-nitrogen treatments which held excessive nitrogen in their lower canopies. As the canopies matured and nitrogen was mobilised to the grain, leaf area index and total nitrogen content decreased and optimal profiles changed shape from exponential to linear. During this period observed profiles became more optimal. There was, however, little difference in P_c between observed and optimal profiles. The maximum difference was 3.2% observed in the low-density, high-nitrogen treatment at DAE 53. The comparison of actual and optimal profiles as leaf nitrogen content (mg N) in addition to the more commonly used specific leaf nitrogen (SLN, g N m^-2 leaf) explains this result because relatively large changes inSLN in the small leaves at the top of canopies have little effect on P_c. The study shows that leaf nitrogen content is an appropriate basis for comparison of canopy nitrogen profiles in sunflower.     

Carbon gain in a multispecies canopy: the role of specific leaf area and photosynthetic nitrogen-use efficiency in the tragedy of the commons

Models have been formulated for monospecific stands in which canopy photosynthesis is determined by the vertical distribution of leaf area, nitrogen and light. In such stands, resident plants can maximize canopy photosynthesis by distributing their nitrogen parallel to the light gradient, with high contents per unit leaf area at the top of the vegetation and low contents at the bottom. Using principles from game theory, we expanded these models by introducing a second species into the vegetation, with the same vertical distribution of biomass and nitrogen as the resident plants but with the ability to adjust its specific leaf area (SLA, leaf area:leaf mass). The rule of the game is that invaders replace the resident plants if they have a higher plant carbon gain than those of the resident plants. We showed that such invaders induce major changes in the vegetation. By increasing their SLA, invading plants could increase their light interception as well as their photosynthetic nitrogen-use efficiency (PNUE, the rate of photosynthesis per unit organic nitrogen). By comparison with stands in which canopy photosynthesis is maximized, those invaded by species of high SLA have the following characteristics: (1) the leaf area index is higher; (2) the vertical distribution of nitrogen is skewed less; (3) as a result of the supra-optimal leaf area index and the more uniform distribution of nitrogen, total canopy photosynthesis is lower. Thus, in dense canopies we face a classical tragedy of the commons: plants that have a strategy to maximize canopy carbon gain cannot compete with those that maximize their own carbon gain. However, because of this strategy, individual as well as total canopy carbon gain are eventually lower. We showed that it is an evolutionarily stable strategy to increase SLA up to the point where the PNUE of each leaf is maximized.     

Plant responses to elevated CO<Subscript>2</Subscript> concentration at different scales: leaf,whole plant,canopy, and population

Elevated CO₂ enhances photosynthesis and growth of plants, but the enhancement is strongly influenced by the availability of nitrogen. In this article, we summarise our studies on plant responses to elevated CO₂. The photosynthetic capacity of leaves depends not only on leaf nitrogen content but also on nitrogen partitioning within a leaf. In Polygonum cuspidatum, nitrogen partitioning among the photosynthetic components was not influenced by elevated CO₂ but changed between seasons. Since the alteration in nitrogen partitioning resulted in different CO₂-dependence of photosynthetic rates, enhancement of photosynthesis by elevated CO₂ was greater in autumn than in summer. Leaf mass per unit area (LMA) increases in plants grown at elevated CO₂. This increase was considered to have resulted from the accumulation of carbohydrates not used for plant growth. With a sensitive analysis of a growth model, however, we suggested that the increase in LMA is advantageous for growth at elevated CO₂ by compensating for the reduction in leaf nitrogen concentration per unit mass. Enhancement of reproductive yield by elevated CO₂ is often smaller than that expected from vegetative growth. In Xanthium canadense, elevated CO₂ did not increase seed production, though the vegetative growth increased by 53%. As nitrogen concentration of seeds remained constant at different CO₂ levels, we suggest that the availability of nitrogen limited seed production at elevated CO₂ levels. We found that leaf area development of plant canopy was strongly constrained by the availability of nitrogen rather than by CO₂. In a rice field cultivated at free-air CO₂ enrichment, the leaf area index (LAI) increased with an increase in nitrogen availability but did not change with CO₂ elevation. We determined optimal LAI to maximise canopy photosynthesis and demonstrated that enhancement of canopy photosynthesis by elevated CO₂ was larger at high than at low nitrogen availability. We also studied competitive asymmetry among individuals in an even-aged, monospecific stand at elevated CO₂. Light acquisition (acquired light per unit aboveground mass) and utilisation (photosynthesis per unit acquired light) were calculated for each individual in the stand. Elevated CO₂ enhanced photosynthesis and growth of tall dominants, which reduced the light availability for shorter subordinates and consequently increased size inequality in the stand.     

Construction and maintenance of the optimal photosynthetic systems of the leaf, herbaceous plant and tree: an eco-developmental treatise

BACKGROUND AND AIMS: The paper by Monsi and Saeki in 1953 (Japanese Journal of Botany 14: 22-52) was pioneering not only in mathematical modelling of canopy photosynthesis but also in eco-developmental studies of seasonal changes in leaf canopies. SCOPE: Construction and maintenance mechanisms of efficient photosynthetic systems at three different scaling levels--single leaves, herbaceous plants and trees--are reviewed mainly based on the nitrogen optimization theory. First, the nitrogen optimization theory with respect to the canopy and the single leaf is briefly introduced. Secondly, significance of leaf thickness in CO2 diffusion in the leaf and in leaf photosynthesis is discussed. Thirdly, mechanisms of adjustment of photosynthetic properties of the leaf within the herbaceous plant individual throughout its life are discussed. In particular, roles of sugar sensing, redox control and of cytokinin are highlighted. Finally, the development of a tree is considered. CONCLUSIONS: Various mechanisms contribute to construction and maintenance of efficient photosynthetic systems. Molecular backgrounds of these ecologically important mechanisms should be clarified. The construction mechanisms of the tree cannot be explained solely by the nitrogen optimization theory. It is proposed that the pipe model theory in its differential form could be a potential tool in future studies in this research area.     

Suboptimal nitrogen status sensitizes the photosynthetic apparatus in willow leaves to long term but not short term water stress

The response to drought was compared for willow plants of optimal leaf nitrogen content (100 N) and those of 86% of this content (86 N). Gas exchange measurements revealed that the carboxylation efficiency (CE) of photosynthesis was more sensitive to drought than the photosynthetic capacity in both N regimes. Since the leaf content of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) was found to be much more resistant it is suggested that a decreased specific activity of Rubisco underlies the decreased CE. Although the rate of water consumption was the same for 86 N and 100 N plants the photosynthetic apparatus responded much more rapidly in the 86 N leaves. This increased sensitivity of 86 N leaves was not due to accelerated senescence as judged by comparison with parallel plants subjected to discontinued fertilization; the two categories of treatments resulted in the same loss of leaf nitrogen and Rubisco but drought induced a much more rapid photosynthetic depression. In contrast to the drought situation, 86 N and 100 N plants behaved similarly when compared under short term water stress. First, when single attached leaves were exposed to a sudden drop in air humidity the capacity of CO₂ uptake in both N regimes decreased about 20% over 10 min while the leaf water potential remained high. Second, in freely transpiring leaf discs cut from 86 N and 100 N leaves the same relationship between capacity of O₂ evolution and extent of dehydration was observed. The possible mechanisms underlying the increased susceptibility of 86 N leaves to drought is discussed; the water status of the roots not the leaves is suggested to be the determining factor.Abbreviations CE carboxylation efficiency - 100 N optimal nitrogen regime - 86 N suboptimal nitrogen regime with 86% of the optimal leaf nitrogen content, Rubisco ribulose-1,5-bisphosphate carboxylase/oxygenase     

Relationships between leaf nitrogen and limitations of photosynthesis in canopies of Solidago altissima

Vertical distribution patterns of light, leaf nitrogen, and leaf gas exchange through canopies of the clonal perennial Solidago altissima were studied in response to mowing and fertilizer application in a field experiment. Consistent with the distribution of light, average leaf nitrogen content followed a `smooth' exponential decline along the fertilized stands both in control and mown plots. The nitrogen profile along the unfertilized stands in mown plots, however, was `disrupted' by high-nitrogen leaves at the top of shorter ramets that only reached intermediate strata of the canopies. Hence, in these stands leaf nitrogen was significantly increased in short ramets compared with tall ramets for a given light environment, suggesting suboptimal stand structure but not necessarily suboptimal single-ramet architecture. However, at least under the climatic conditions observed during measurements, such disrupture had no substantial effect on stand productivity: model calculations showed that vertical distribution patterns of leaf nitrogen along ramets only marginally influenced the photosynthetic performance of ramets and stands. This is explained by the observed photosynthesis-nitrogen relationship: the rate of photosynthesis per unit amount of leaf nitrogen did not increase with leaf nitrogen content even under saturating light levels indicating that leaf photosynthesis was not nitrogen limited during the measurement periods. Nevertheless, our study indicates that consideration of how architecture(s) of adjacent individual plants interact could be essential for a better understanding of the trade-offs between individual and canopy characteristics for maximizing carbon gain. Such trade-offs may end up in a suboptimal canopy structure, which could not be predicted and understood by classical canopy optimization models.     

Would transformation of C3 crop plants with foreign Rubisco increase productivity? A computational analysis extrapolating from kinetic properties to canopy photosynthesis

Genetic modification of Rubisco to increase the specificity for CO₂ relative to O₂ (τ) would decrease photorespiration and in principle should increase crop productivity. When the kinetic properties of Rubisco from different photosynthetic organisms are compared, it appears that forms with high τ have low maximum catalytic rates of carboxylation per active site (k^c_c). If it is assumed that an inverse relationship between k^c_c and τ exists, as implied from measurements, and that an increased concentration of Rubisco per unit leaf area is not possible, will increasing τ result in increased leaf and canopy photosynthesis? A steady‐state biochemical model for leaf photosynthesis was coupled to a canopy biophysical microclimate model and used to explore this question. C₃ photosynthetic CO₂ uptake rate (A) is either limited by the maximum rate of Rubisco activity (V_cmax) or by the rate of regeneration of ribulose‐1,5‐bisphosphate, in turn determined by the rate of whole chain electron transport (J). Thus, if J is limiting, an increase in τ will increase net CO₂ uptake because more products of the electron transport chain will be partitioned away from photorespiration into photosynthesis. The effect of an increase in τ on Rubisco‐limited photosynthesis depends on both k^c_c and the concentration of CO₂ ([CO₂]). Assuming a strict inverse relationship between k^c_c and τ, the simulations showed that a decrease, not an increase, in τ increases Rubisco‐limited photosynthesis at the current atmospheric [CO₂], but the increase is observed only in high light. In crop canopies, significant amounts of both light‐limited and light‐saturated photosynthesis contribute to total crop carbon gain. For canopies, the present average τ found in C₃ terrestrial plants is supra‐optimal for the present atmospheric [CO₂] of 370 µmol mol^?1, but would be optimal for a CO₂ concentration of around 200 µmol mol^?1, a value close to the average of the last 400 000 years. Replacing the average Rubisco of terrestrial C₃ plants with one having a lower and optimal τ would increase canopy carbon gain by 3%. Because there are significant deviations from the strict inverse relationship between k^c_c and τ, the canopy model was also used to compare the rates of canopy photosynthesis for several Rubiscos with well‐defined kinetic constants. These simulations suggest that very substantial increases (> 25%) in crop carbon gain could result if specific Rubiscos having either a higher τ or higher k^c_c were successfully expressed in C₃ plants.     

Does Leaf Position within a Canopy Affect Acclimation of Photosynthesis to Elevated CO2? : Analysis of a Wheat Crop under Free-Air CO2 Enrichment

Previous studies of photosynthetic acclimation to elevated CO₂ have focused on the most recently expanded, sunlit leaves in the canopy. We examined acclimation in a vertical profile of leaves through a canopy of wheat (Triticum aestivum L.). The crop was grown at an elevated CO₂ partial pressure of 55 Pa within a replicated field experiment using free-air CO₂ enrichment. Gas exchange was used to estimate in vivo carboxylation capacity and the maximum rate of ribulose-1,5-bisphosphate-limited photosynthesis. Net photosynthetic CO₂ uptake was measured for leaves in situ within the canopy. Leaf contents of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), light-harvesting-complex (LHC) proteins, and total N were determined. Elevated CO₂ did not affect carboxylation capacity in the most recently expanded leaves but led to a decrease in lower, shaded leaves during grain development. Despite this acclimation, in situ photosynthetic CO₂ uptake remained higher under elevated CO₂. Acclimation at elevated CO₂ was accompanied by decreases in both Rubisco and total leaf N contents and an increase in LHC content. Elevated CO₂ led to a larger increase in LHC/Rubisco in lower canopy leaves than in the uppermost leaf. Acclimation of leaf photosynthesis to elevated CO₂ therefore depended on both vertical position within the canopy and the developmental stage.     

Impact of nitrogen allocation on growth and photosynthesis of Miscanthus (Miscanthus × giganteus)

Nitrogen (N) addition typically increases overall plant growth, but the nature of this response depends upon patterns of plant nitrogen allocation that vary throughout the growing season and depend upon canopy position. In this study seasonal variations in leaf traits were investigated across a canopy profile in Miscanthus (Miscanthus × giganteus) under two N treatments (0 and 224 kg ha^?1) to determine whether the growth response of Miscanthus to N fertilization was related to the response of photosynthetic capacity and nitrogen allocation. Miscanthus yielded 24.1 Mg ha^?1 in fertilized plots, a 40% increase compared to control plots. Photosynthetic properties, such as net photosynthesis (A), maximum rate of rubisco carboxylation (V_cmax), stomatal conductance (g_s) and PSII efficiency (F_v'/F_m'), all decreased significantly from the top of the canopy to the bottom, but were not affected by N fertilization. N fertilization increased specific leaf area (SLA) and leaf area index (LAI). Leaf N concentration in different canopy layers was increased by N fertilization and the distribution of N concentration within canopy followed irradiance gradients. These results show that the positive effect of N fertilization on the yield of Miscanthus was unrelated to changes in photosynthetic rates but was achieved mainly by increased canopy leaf area. Vertical measurements through the canopy demonstrated that Miscanthus adapted to the light environment by adjusting leaf morphological and biochemical properties independent of nitrogen treatments. GPP estimated using big leaf and multilayer models varied considerably, suggesting a multilayer model in which V_cmax changes both through time and canopy layer could be adopted into agricultural models to more accurately predict biomass production in biomass crop ecosystems.     

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.

     

塔克拉玛干沙漠南缘豆科与非豆科植物的氮分配

在豆科与非豆科植物光合特性的研究中发现,非豆科植物具有更高的光合速率,与其低的叶氮含量相矛盾。在沙漠中氮素是限制植物生长的关键因子之一,考虑到豆科植物的生物固氮作用和叶氮大部分分配于光合系统,我们假设:(1)非豆科植物具有更低的叶氮含量;(2)分配更少的叶氮于光合系统;(3)具有更高的最大净光合速率(Pmax)和光合氮素利用效率(PNUE)。为了验证这些假设,以塔克拉玛干沙漠南缘的豆科植物骆驼刺(Alhagi sparsifolia)和非豆科植物柽柳(Tamarix ramosissima)、花花柴(Karelinia caspica)为研究对象,比较了它们的叶氮含量、氮分配、Pmax和PNUE等。结果表明:(1)非豆科植物比豆科植物确实有更低的叶氮含量,且差异达到显著水平;(2)非豆科植物分配更少的叶氮于光合系统,但在光合系统内部具有更高效的氮分配机制;(3)非豆科植物具有更高的Pmax和PNUE。在光合系统内部,非豆科植物分配更多的叶氮于羧化系统,而豆科植物分配更多的叶氮于捕光系统。对于非豆科植物而言,其更高的Pmax、PNUE、水分利用效率和表观量子产量,取决于将更多的叶氮投入到羧化和电子传递系统中。这些生理优势决定了塔克拉玛干沙漠南缘非豆科植物高效的资源捕捉和利用能力。     

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.