A model of leaf photosynthesis and respiration for predicting carbon dioxide assimilation in different environments

Summary A model of leaf photosynthesis and repiration was developed which adequately predicted carbon dioxide assimilation responses by a C ₃ species, Atriplex patula, to light, [CO₂], [O₂] and temperature in controlled environments. Methods were developed for estimating input parameters using laboratory, controlled environment and field data.     

Glyphosate effects on carbon assimilation and gas exchange in sugar beet leaves

The mechanism responsible for the inhibition of net carbon exchange (NCE) which was reported previously (DR Geiger et al. 1986 Plant Physiol 82: 468-472) was investigated by applying glyphosate [N-(phosphonomethyl)glycine] to exporting leaves of sugar beet (Beta vulgaris L.). Leaf internal CO₂ concentration (C_i) remained constant despite decreases in stomatal conductance and NCE following glyphosate treatment, indicating that the cause of the inhibition was a slowing of carbon assimilation rather than decreased conductance of CO₂. Throughout a range of CO₂ concentrations, NCE rate at a given C_i declined gradually, with the time-series of response curves remaining parallel. Gas exchange measurements revealed that disruption of chloroplast carbon metabolism was an early and important factor in mediating these glyphosate effects, perhaps by slowing the rate of ribulose bisphosphate regeneration. An increase in the CO₂ compensation point accompanied the decrease in NCE and this increase was hastened by stepwise lowering of the ambient CO₂ concentration. Eventually the CO₂ compensation point approached the CO₂ level of air and the difference between internal and external CO₂ concentrations decreased. In control and in glyphosate-treated plants, both carbon assimilation and photorespiration at atmospheric CO₂ level were inhibited to a similar extent of air level of O₂. Maintaining leaves in low O₂ concentration did not prevent the decline in NCE rate.     

Glyphosate inhibits photosynthesis and allocation of carbon to starch in sugar beet leaves

Application of glyphosate (N-[phosphonomethyl] glycine) to exporting leaves of sugar beet (Beta vulgaris, L.) during the day lowered stomatal conductance and carbon fixation. Allocation of newly fixed carbon to foliar starch accumulation was nearly completely inhibited, being decreased by the same amount as net carbon fixation. In contrast, decreasing net carbon fixation in untreated leaves by lowering CO₂ concentration caused starch accumulation to decrease, but only in the same proportion as net carbon fixation. Shikimate level increased 50-fold in treated leaves but the elevated rate of carbon accumulation in shikimate was only 4% of the decrease in the rate of starch accumulation. Application of steady state labeling with ¹⁴CO₂ to exporting leaves confirmed the above changes in carbon metabolism, but revealed no other major daytime differences in the ¹⁴C-content of amino acids or other compounds between treated and control leaves. Less ¹⁴C accumulated in treated leaves because of decreased fixation, not increased export. The proportion of newly fixed carbon allocated to sucrose increased, maintaining export at the level in control leaves. Returning net carbon exchange to the rate before treatment restored starch accumulation fully and prevented a decrease in export during the subsequent dark period.     

Essentiality of mitochondrial oxidative metabolism for photosynthesis: optimization of carbon assimilation and protection against photoinhibition

The review emphasizes the essentiality of mitochondrial oxidative metabolism for photosynthetic carbon assimilation. Photosynthetic activity in chloroplasts and oxidative metabolism in mitochondria interact with each other and stimulate their activities. During light, the partially modified TCA cycle supplies oxoglutarate to cytosol and chloroplasts. The marked stimulation of O2 uptake after few minutes of photosynthetic activity, termed as light enhanced dark respiration (LEDR), is now a well-known phenomenon. Both the cytochrome and alternative pathways of mitochondrial electron transport are important in such interactions. The function of chloroplast is optimized by the complementary nature of mitochondrial metabolism in multiple ways: facilitation of export of excess reduced equivalents from chloroplasts, shortening of photosynthetic induction, maintenance of photorespiratory activity, and supply of ATP for sucrose biosynthesis as well as other cytosolic needs. Further, the mitochondrial oxidative electron transport and phosphorylation also protects chloroplasts against photoinhibition. Besides mitochondrial respiration, reducing equivalents (and ATP) are used for other metabolic phenomena, such as sulfur or nitrogen metabolism and photorespiration. These reactions often involve peroxisomes and cytosol. The beneficial interaction between chloroplasts and mitochondria therefore extends invariably to also peroxisomes and cytosol. While the interorganelle exchange of metabolites is the known basis of such interaction, further experiments are warranted to identify other biochemical signals between them. The uses of techniques such as on-line mass spectrometric measurement, novel mutants/transgenics, and variability in metabolism by growth conditions hold a high promise to help the plant biologist to understand this     

The influence of water content and leaf anatomy on carbon isotope discrimination and photosynthesis in Sphagnum

The relative effect of diffusional resistance due to water films (r_wf) and leaf anatomy (r_p) on rates of net photosynthesis and on-line measures of carbon isotope discrimination (Δ=Δδ¹³C) was investigated in Sphagnum. Sphagnum species differ in the exposure of photosynthetic cells at the leaf surface. In S. affine, photosynthetic cells are widely exposed at the surface, whereas in S. magellanicum, photo-synthetic cells are enclosed within water-filled hyaline cells. This difference is expected to lead to variation in diffusive resistance within leaves (r_p). Net photosynthesis and on-line Δ were measured at two water contents: greenhouse water content (wet) and blotted dry (dry). Without correcting for respiration, on-line Δ values differed significantly between wet (23.7%_o) and dry (30.9%_o) plants. However, there was no significant difference between species means and no species × water content interaction. Corrections for respiration lowered Δ values by approximately 8.1%_o and reduced the mean difference to 3.1%_o, but did not alter the rank order of treatments. Net photosynthesis also decreased by 16% in wet plants, but there was no significant difference between the two species. In addition, five populations of S. affine and S. magellanicum grown in a common garden were analysed for their organic matter carbon isotope composition (δ¹³C). These values varied more within each species (0.9–1.2%_o) than between the two species (0.6%_o). Therefore, we conclude that variation in surface water films leads to a greater difference in resistance to CO₂ uptake and carbon isotope discrimination than that due to variation in leaf anatomical properties in Sphagnum.     

Studies for the transformation of carbocycles into carbohydrates: approach toward the synthesis of higher sugar derivatives

A highly stereocontrolled synthesis of a beta-D-ribo-hept-6-ulopyranosuronamide derivative, a useful intermediate for the synthesis of other higher sugars, has been developed using naturally occurring (-)-quinic acid as a chiral starting material. The transformation of carbocycle to carbohydrate, a key step in this sequence, occurred in a one-pot reaction: an ozonolysis carried out under mild conditions.     

Intracellular metabolite gradients and flow of carbon during photosynthesis of leaf protoplasts

Protoplasts were isolated from wheat and spinach leaves and after photosynthesis in the presence of ${}^{14}\text{CO}{}_2$ fractionated into a chloroplast and a nonchloroplast fraction. The kinetics of the distribution of labeled metabolities between the fractions indicated carbon flow from the chloroplasts into the cytosol and the vacuole. After 10 min of photosynthesis, more than 90% of the assimilated soluble carbon was outside the chloroplasts. Different metabolite levels indicated intracellular metabolite compartmentation and metabolite gradients. During photosynthesis, gradient coupling facilitated uphill export of triosephosphate from the chloroplasts into the cytosol. The driving force appeared to be an opposite phosphate gradient. Different ratios of phosphoglycerate to triosephosphate inside and outside the chloroplasts indicated the existence of a transenvelope pH gradient. Sucrose appeared to be synthesized outside the chloroplasts and is probably exported into the vacuole. The behavior of malate also suggested transfer into the vacuole. The malate/ aspartate ratio was higher outside the chloroplasts that inside suggesting import of reducing equivalents into the chloroplasts through the dicarboxylate translocator.     

The photosynthesis – leaf nitrogen relationship at ambient and elevated atmospheric carbon dioxide: a meta-analysis

Estimation of leaf photosynthetic rate (A) from leaf nitrogen content (N) is both conceptually and numerically important in models of plant, ecosystem, and biosphere responses to global change. The relationship between A and N has been studied extensively at ambient CO₂ but much less at elevated CO₂. This study was designed to (i) assess whether the A–N relationship was more similar for species within than between community and vegetation types, and (ii) examine how growth at elevated CO₂ affects the A–N relationship. Data were obtained for 39 C3 species grown at ambient CO₂ and 10 C3 species grown at ambient and elevated CO₂. A regression model was applied to each species as well as to species pooled within different community and vegetation types. Cluster analysis of the regression coefficients indicated that species measured at ambient CO₂ did not separate into distinct groups matching community or vegetation type. Instead, most community and vegetation types shared the same general parameter space for regression coefficients. Growth at elevated CO₂ increased photosynthetic nitrogen use efficiency for pines and deciduous trees. When species were pooled by vegetation type, the A–N relationship for deciduous trees expressed on a leaf-mass basis was not altered by elevated CO₂, while the intercept increased for pines. When regression coefficients were averaged to give mean responses for different vegetation types, elevated CO₂ increased the intercept and the slope for deciduous trees but increased only the intercept for pines. There were no statistical differences between the pines and deciduous trees for the effect of CO₂. Generalizations about the effect of elevated CO₂ on the A–N relationship, and differences between pines and deciduous trees will be enhanced as more data become available.     

Oxidation and assimilation of carbohydrates by Micrococcus sodonensis

Perry, Jerome J. (North Carolina State University, Raleigh), and James B. Evans. Oxidation and assimilation of carbohydrates by Micrococcus sodonensis. J. Bacteriol. 91:33-38. 1966.-Micrococcus sodonensis is a biotin-requiring strict aerobe that cannot utilize carbohydrates as sole sources of carbon and energy. However, addition of mannose, glucose, sucrose, or maltose to a medium on which the organism can grow resulted in an increase in total growth. M. sodonensis oxidized these sugars without induction, thus indicating the presence of constitutive enzymes for their transport, activation, and metabolism. Under appropriate nonproliferating cell conditions, glucose was readily incorporated into essential constituents of the cell. When glucose-1-C(14) and glucose-6-C(14) were oxidized by nonproliferating cells, the label was found in both the protein and nucleic acid fractions of the cell. The respiratory quotients of cells oxidizing glucose in saline and in phosphate buffer indicated assimilation of sugar carbon in buffer and virtually no assimilation in saline. The ability of M. sodonensis to completely oxidize glucose and to grow on intermediates of glucose oxidation but not on glucose suggests that glucose may suppress or repress some reaction(s) necessary for growth, and that growth substrates either derepress or circumvent this block.     

Translocation and the control of photosynthesis in sugar beet

Summary Evidence is presented that the size and activity of sinks for the products of photosynthesis exert a large measure of control over carbohydrate levels in the leaves of the sugar beet plant. Further data is presented to show that a correlation exists between the level carbohydrate in the leaf and the rate of photosynthesis. The rate of photosynthesis can thus be linked directly to the sinks of the plant by their ability to control rates of translocation from leaves.     

The OzT8 locus in rice protects leaf carbon assimilation rate and photosynthetic capacity under ozone stress

Tropospheric ozone (O₃) is a phytotoxic air pollutant whose current background concentrations in parts of East Asia have caused estimated rice yield losses of up to 20%; currently, however, little is known about the mechanisms of O₃ tolerance in rice. We previously identified a quantitative trait locus (QTL) in rice called OzT8, which was associated with relative dry weight under ozone stress. The photosynthetic response in SL46, a Nipponbare (NB)–Kasalath chromosome segment substitution line (SL) containing the OzT8 locus, was compared to the parent NB in multiple ozone fumigation experiments (100 ppb, 8 h d^–1, 23 d). By day 23, SL46 showed significantly less reduction of photosynthetic capacity compared to NB; the maximum carboxylation rate of ribulose 1·5‐bisphosphate carboxylase/oxygenase (Rubisco) decreased by 24% in SL46 compared to 49% in NB, and the maximum electron transport rate decreased by 16 and 39%, respectively. The midday carbon assimilation rates also showed a similar trend, but there was no genotypic difference in stomatal conductance. These results indicate that the OzT8 locus confers ozone tolerance via biochemical acclimation, not avoidance, making it a potentially valuable target for breeding of ozone tolerance into future rice lines. The sequence of photosynthetic response of rice to ozone stress and related tolerance factors are also discussed.     

Carbon dioxide assimilation in oxygenic and anoxygenic photosynthesis

This article represents a summary of our contemporary understanding of carbon dioxide assimilation in photosynthesis, including both the oxygen-evolving (oxygenic) type characteristic of cyanobacteria, algae and higher plants, and the non-oxygen-evolving (anoxygenic) type characteristic of other bacteria. Mechanisms functional in the regulation of the reductive pentose phosphate cycle of oxygenic photosynthesis are emphasized, as is the reductive carboxylic acid cycle-the photosynthetic carbon pathway functional in anoxygenic green sulfur bacteria. Thioredoxins, an ubiquitous group of low molecular weight proteins with catalytically active thiols, are also described in some detail, notably their role in regulating the reductive pentose phosphate cycle of oxygenic photosynthesis and their potential use as markers to trace the evolutionary development of photosynthesis.Abbreviations NADP-GAPDH-NADP glyceraldehyde 3-phosphate dehydrogenase - FBPase fructose 1,6-bisphosphatase - FTR ferredoxin-thioredoxin reductase - Rubisco ribulose 1,5-bisphosphate carboxylase/oxygenase - SBPase sedoheptulos 1,7-bisphosphatase - PRK phosphoribulokinase - NADP-MDH-NADP malate dehydrogenase - CF1-ATPase chloroplast coupling factor - G6PDH glucose 6-phosphate dehydrogenase Most of the references cited in this article are reviews. For references to specific material, readers should consult the appropriate review.     

Short-term photosynthesis measurements predict leaf carbon balance in tropical rain-forest canopy plants

Diel (24 h) courses of net CO₂ exchange of leaves were determined in eight species of tropical rainforest plants on Barro Colorado Island, Panama, during 1990 and 1991. The species included three canopy trees, one liana, two epiphytes and one hemiepiphyte. One of the species studied was growing in a rain-forest gap. Daily carbon gain varied considerably across species, leaf age, and season. The analysis of data for all plants from 64 complete day/night cycles revealed a linear relationship between the diurnal carbon gain and the maximum rate of net CO₂ uptake, A_max. Nocturnal net carbon loss was about 10% of diurnal carbon gain and was positively related to A_max. We conclude that short-term measurements of light-saturated photosynthesis, performed at periodic intervals throughout the season, allow the annual leaf carbon balance in these rain-forest plants to be predicted.     

Effect of oxygen concentration on leaf photosynthesis and resistances to carbon dioxide diffusion

Summary Net photosynthesis of tropical legume leaves increased by 44% and that of tropical grass leaves was unaffected when oxygen concentration was reduced from 21 to 0.2%. Stomatal resistance to carbon dioxide diffusion was unaltered in both cases but mesophyll resistance of legume leaves decreased with oxygen concentration. It is proposed that the decrease in mesophyll resistance is accompanied by decreases in excitation and carboxylation resistances.