The photosynthetic capacity of leaves is related to the nitrogen content primarily bacause the proteins of the Calvin cycle and thylakoids represent the majority of leaf nitrogen. To a first approximation, thylakoid nitrogen is proportional to the chlorophyll content (50 mol thylakoid N mol-1 Chl). Within species there are strong linear relationships between nitrogen and both RuBP carboxylase and chlorophyll. With increasing nitrogen per unit leaf area, the proportion of total leaf nitrogen in the thylakoids remains the same while the proportion in soluble protein increases. In many species, growth under lower irradiance greatly increases the partitioning of nitrogen into chlorophyll and the thylakoids, while the electron transport capacity per unit of chlorophyll declines. If growth irradiance influences the relationship between photosynthetic capacity and nitrogen content, predicting nitrogen distribution between leaves in a canopy becomes more complicated. When both photosynthetic capacity and leaf nitrogen content are expressed on the basis of leaf area, considerable variation in the photosynthetic capacity for a given leaf nitrogen content is found between species. The variation reflects different strategies of nitrogen partitioning, the electron transport capacity per unit of chlorophyll and the specific activity of RuBP carboxylase. Survival in certain environments clearly does not require maximising photosynthetic capacity for a given leaf nitrogen content. Species that flourish in the shade partition relatively more nitrogen into the thylakoids, although this is associated with lower photosynthetic capacity per unit of nitrogen.

Photosynthesis and nitrogen relationships in leaves of C3 plants.

Contents
 	Summary	30	
I.	Allocation in perspective	31	
II.	Topics of this review	32	
III.	Methodology	32	
IV.	Environmental effects	33	
V.	Ontogeny	36	
VI.	Differences between species	40	
VII.	Physiology and molecular regulation	41	
VIII.	Ecological aspects	42	
IX.	Perspectives	45	
 	Acknowledgements	45	
 	References	45	
 	Appendices A1–A4	49	


Summary
We quantified the biomass allocation patterns to leaves, stems and roots in vegetative plants, and how this is influenced by the growth environment, plant size, evolutionary history and competition. Dose–response curves of allocation were constructed by means of a meta-analysis from a wide array of experimental data. They show that the fraction of whole-plant mass represented by leaves (LMF) increases most strongly with nutrients and decreases most strongly with light. Correction for size-induced allocation patterns diminishes the LMF-response to light, but makes the effect of temperature on LMF more apparent. There is a clear phylogenetic effect on allocation, as eudicots invest relatively more than monocots in leaves, as do gymnosperms compared with woody angiosperms. Plants grown at high densities show a clear increase in the stem fraction. However, in most comparisons across species groups or environmental factors, the variation in LMF is smaller than the variation in one of the other components of the growth analysis equation: the leaf area : leaf mass ratio (SLA). In competitive situations, the stem mass fraction increases to a smaller extent than the specific stem length (stem length : stem mass). Thus, we conclude that plants generally are less able to adjust allocation than to alter organ morphology.

https://nph.onlinelibrary.wiley.com/doi/epdf/10.1111/j.1469-8137.2011.03952.x

Biomass allocation to leaves, stems and roots: meta-analyses of interspecific variation and environmental control

In big-leaf models of canopy photosynthesis, the Rubisco activity per unit ground area is taken as the sum of activities per unit leaf area within the canopy, and electron transport capacity is similarly summed. Such models overestimate rates of photosynthesis and require empirical curvature factors in the response to irradiance. We show that, with any distribution of leaf nitrogen within the canopy (including optimal), the required curvature factors are not constant but vary with canopy leaf area index and leaf nitrogen content. We further show that the underlying reason is the difference between the time-averaged and instantaneous distributions of absorbed irradiance, caused by penetration of sunflecks and the range of leaf angles in canopies. These errors are avoided in models that treat the canopy in terms of a number of layers ‐ the multi-layer models. We present an alternative to the multi-layer model: by separately integrating the sunlit and shaded leaf fractions of the canopy, a single layered sun/shade model is obtained, which is as accurate and simpler. The model is a scaled version of a leaf model as distinct from an integrative approach.

/pdf/simple-scaling-of-photosynthesis-from-leaves-to-canopies-1pf2pqerhm.pdf

Simple scaling of photosynthesis from leaves to canopies without the errors of big‐leaf models

AGPase, ADP glucose pyrophosphorylase
GS, glutamine synthetase
GOGAT, glutamate : oxoglutarate amino transferase
NADP-ICDH, NADP-dependent isocitrate dehydrogenase
NR, nitrate reductase
OPPP, oxidative pentose phosphate pathway
3PGA, glycerate-3-phosphate
PEPCase, phosphoenolpyruvate carboxylase
Rubisco, ribulose-1,5-bisphosphate carboxylase/oxygenase
SPS, sucrose phosphate-synthase

This review first summarizes the numerous studies that have described the interaction between the nitrogen supply and the response of photosynthesis, metabolism and growth to elevated [CO2]. The initial stimulation of photosynthesis in elevated [CO2] is often followed by a decline of photosynthesis, that is typically accompanied by a decrease of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco), an accumulation of carbohydrate especially starch, and a decrease of the nitrogen concentration in the plant. These changes are particularly marked when the nitrogen supply is low, whereas when the nitrogen supply is adequate there is no acclimation of photosynthesis, no major decrease in the internal concentration of nitrogen or the levels of nitrogen metabolites, and growth is stimulated markedly. Second, emerging evidence is discussed that signals derived from nitrate and nitrogen metabolites such as glutamine act to regulate the expression of genes involved in nitrate and ammonium uptake and assimilation, organic acid synthesis and starch accumulation, to modulate the sugar-mediated repression of the expression of genes involved in photosynthesis, and to modulate whole plant events including shoot–root allocation, root architecture and flowering. Third, increased rates of growth in elevated [CO2] will require higher rates of inorganic nitrogen uptake and assimilation. Recent evidence is discussed that an increased supply of sugars can increase the rates of nitrate and ammonium uptake and assimilation, the synthesis of organic acid acceptors, and the synthesis of amino acids. Fourth, interpretation of experiments in elevated [CO2] requires that the nitrogen status of the plants is monitored. The suitability of different criteria to assess the plant nitrogen status is critically discussed. Finally the review returns to experiments with elevated [CO2] and discusses the following topics: is, and if so how, are nitrate and ammonium uptake and metabolism stimulated in elevated [CO2], and does the result depend on the nitrogen supply? Is acclimation of photosynthesis the result of sugar-mediated repression of gene expression, end-product feedback of photosynthesis, nitrogen-induced senescence, or ontogenetic drift? Is the accumulation of starch a passive response to increased carbohydrate formation, or is it triggered by changes in the nutrient status? How do changes in sugar production and inorganic nitrogen assimilation interact in different conditions and at different stages of the life history to determine the response of whole plant growth and allocation to elevated [CO2]?

https://www.onlinelibrary.wiley.com/doi/pdf/10.1046/j.1365-3040.1999.00386.x

The interaction between elevated carbon dioxide and nitrogen nutrition: the physiological and molecular background

Plant responses to atmospheric CO2 enrichment with emphasis on roots and the rhizosphere.

The Contribution of Leaves from Different Levels within a Tomato Crop to Canopy Net Photosynthesis: An Experimental Examination of Two Canopy Models

The carbon dioxide exchange rates of single attached tomato leaves were continuously measured over a 24 h period using IRGA techniques. Net photosynthetic rates were measured in a constant light and carbon dioxide environment for 8 h. The leaf was then darkened and the rate of CO2 evolution measured for a further 16 h. Different light levels or carbon dioxide concentrations were used in these assays each day.

Tomato Leaf Photosynthesis and Respiration in Various Light and Carbon Dioxide Environments

Ribulose Bisphosphate Carboxylase Activity and Photosynthesis during Leaf Development in the Tomato

The light-saturated rates of photosynthesis at 300 vpm CO2 and 20°C (Pmax) of leaves at different stages of development have been predicted with a knowledge of the amount and kinetics of tomato leaf RuBPc/o, based on the assumption that under these conditions the enzyme is fully activated and RuBP saturated. When grown in high CO2 (1000 vpm) the amount and activity of RuBPc under unit surface area and Pmax decreased sharply beyond about 25% of full leaf expansion compared with leaves grown in 300 vpm CO2. When almost fully expanded the high CO2 grown leaves contained only about half as much RuBPc as the leaves grown in a normal [CO2] and Pmax was similarly reduced. Overall, the predicted rates of Pmax correlated reasonably well with observed rates, supporting the view that this enzyme may be a factor governing photosynthetic capacity.

Ribulose Bisphosphate Carboxylase and Net CO2 Fixation in Tomato Leaves

Dans cette etude, les effets a long terme d'un enrichissement en CO 2 atmospherique sur la photosynthese sont examines chez des plants de tomate. Des mesures de contenu en ribulose-1,5-bisphosphate carboxylase et de son activite, ainsi que de l'activite photosynthetique sont effectuees au cours du developpement des feuilles

The Greenhouse Effect: Acclimation of Tomato Plants Growing in High CO2, Photosynthesis and Ribulose-1, 5-Bisphosphate Carboxylase Protein

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