Regulation Of Carbon Partitioning Research Articles

Overexpression of ZmSUS1 increased drought resistance of maize (Zea mays L.) by regulating sucrose metabolism and soluble sugar content.

ZmSUS1 improved drought tolerance of maize by regulating sucrose metabolism and increasing soluble sugar content, and endowing transgenic maize with higher relative water content and photosynthesis levels. Sucrose synthase (SUS), a key enzyme of sugar metabolism, plays an important role in the regulation of carbon partitioning in plant, and affects important agronomic traits and abiotic responses to adversity. However, the function of ZmSUS1 in plant drought tolerance is still unknown. In this study, the expression patterns of ZmSUS1 in different tissues and under drought stress were analyzed in maize (Zea mays L.). It was found that ZmSUS1 was highly expressed during kernel development but also in leaves and roots of maize, and ZmSUS1 was induced by drought stress. Homozygous transgenic maize lines overexpressing ZmSUS1 increased the content and activity of SUS under drought stress and exhibited higher relative water content, proline and abscisic acid content in leaves. Specifically, the net photosynthetic rate and the soluble sugar contents including sucrose, glucose, fructose and SUS decomposition products including UDP-glucose (UDP-G) and ADP-glucose (ADP-G) in transgenic plants were significantly improved after drought stress. RNA-seq analysis showed that overexpressing of ZmSUS1 mainly affected the expression level of carbon metabolism-related genes. Especially the expression level of sucrose metabolism-related genes including sucrose phosphatase gene (SPP), sucrose phosphate synthase gene (SPS) and invertase gene (INV) were significantly up-regulated in transgenic maize. Overall, these results suggested that ZmSUS1 improved drought tolerance by regulating sucrose metabolism and increasing the soluble sugar content, and endowing transgenic maize with higher relative water content and photosynthesis levels, which can serve as a new gene candidate for cultivating drought-resistant maize varieties.

Trehalose treatment alters carbon partitioning and reduces the accumulation of individual metabolites but does not affect salt tolerance in the green microalga Dunaliella bardawil.

The effects of trehalose (Tre), a non-reducing disaccharide, on metabolic changes, antioxidant status, and salt tolerance in Dunaliella bardawil cells were investigated. Algal suspensions containing 1, 2, and 3M NaCl were treated with 5mM Tre. While the content of pigments, reducing sugars, proteins, glycerol, and ascorbate pool accumulated with increasing salinity, the content of non-reducing sugars, starch, amino acids, proline, hydrogen peroxide, and lipid peroxidation level decreased significantly. Tre-treated cells showed a decrease in pigments content, reducing sugars, starch, proteins, amino acids, proline, glycerol, and the activity of non-specific peroxidase and polyphenol oxidase, but an increase in non-reducing sugars, oxidized ascorbate, and ascorbate peroxidase activity occurred unchanged in the ascorbate pool. However, the density and fresh weight of the cells remained statistically unchanged in all Tre-treated and untreated cultures. These results suggest that D. bardawil cells potentially tolerate different salt levels by accumulating metabolites, whereas Tre treatment changes carbon partitioning and significantly reduces beneficial metabolites without altering salt tolerance. Therefore, the regulation of carbon partitioning rather than the amount of assimilated carbon may play an important role in inducing salinity tolerance of D. bardawil. However, Tre is not able to enhance the salt tolerance of halotolerants and is even economically damaging due to the reduction of unique metabolites such as glycerol and β-carotene.

Manipulation of Source-Sink Relationship in Pertinence to Better Fruit Quality and Yield in Fruit Crops: A Review

Profuse flowering and fruiting creates high demand for a limited source of carbohydrates, which affects fruit set and development. These plenty number of flowers and fruits must be reduced to allow the trees to produce fruit that meets market expectations for size, colour and quality. The concept of source-sink relationship and the regulation of carbon partitioning in plants is useful for better understanding of crop physiology and the influence of yield limiting factors on crop production. This relationship between source leaves and the various sink organs not only affects fruit production in the current season, but also the long term performance due to perennial nature of fruit trees. Modelling early season whole-tree carbohydrate supply and demand has greatly improved the understanding of response of trees to manipulation strategies relating source-sink alliance. Various strategies has been adopted to manipulate source-sink relationship like flower bud inhibition, flower thinning, fruitlet thinning, defoliation, girdling, pruning, application of hormones and nutrients etc. Thus, balance between vegetative and reproductive growth is an important aspect for improvement in yield and quality of fruit crops.

Manipulation of a glycolytic regulator alters growth and carbon partitioning in the marine diatom Thalassiosira pseudonana

The regulation of carbon partitioning in diatoms is key to understanding mechanisms related to their high productivity, and the mechanisms controlling carbon flux towards different metabolic fates are currently not well understood. A unique variant of 6-phosphofructo-2-kinase/fructose 2,6 bisphosphatase (PFK2/F2BP), a regulator of glycolytic flux previously uncharacterized in single-cell photosynthetic eukaryotes, was identified in diatom genomes and overexpressed in Thalassiosira pseudonana. Overexpression resulted in increased activity of the glycolytic rate-determining enzyme phosphofructokinase (PFK) and altered carbon partitioning among intracellular carbohydrate, lipid, and protein levels. Higher PFK activity in transformant lines correlated with reduced growth rate and an extension of the G1 phase of the cell cycle, demonstrating a link between carbon metabolism and the control of cell cycle processes. While the relationship between growth and carbon flux under environmental stress (such as nutrient limitation) is well documented in diatoms, our data demonstrate that manipulation of carbon metabolism, independent of an environmental trigger (such as nutrient limitation), is sufficient to delay cell cycle progression. The inverse relationship between growth and PFK activity (as an indicator of glycolytic flux) differs from those described for other unicellular eukaryotes and supports an alternate organization and regulation of central carbon metabolism in diatoms, including a prominent regulatory role for PFK2/F2BP.

Regulation of Carbon Partitioning in the Seed of the Model Legume Medicago truncatula and Medicago orbicularis: A Comparative Approach.

The proportion of starch, protein and oil in legume seeds is species dependent. The model legume, Medicago truncatula, has predominantly oil and protein stores. To investigate the regulation of seed oil production we compared M. truncatula with M. orbicularis, which has less oil and protein. The types of protein and fatty acids are similar between the two species. Electron microscopy indicated that the size and distribution of the oil bodies in M. orbicularis, is consistent with reduced oil production. M. orbicularis has more extruded endosperm mucilage compared to M. truncatula. The cotyledons have a greater cell wall content, visualized as thicker cell walls. The reduced oil content in M. orbicularis is associated with increased expression of the MtGLABRA2-like (MtGL2) transcription factor, linked to an inverse relationship between mucilage and oil content in Arabidopsis. The expression of the pectin biosynthesis GALACTURONOSYLTRANSFERASE (GAUT) genes, is also increased in M. orbicularis. These increases in extruded mucilage and cell wall storage components in M. orbicularis are accompanied by reduced expression of transcriptional regulators of oil biosynthesis, MtLEAFY COTYLEDON1-LIKE (MtL1L), MtABSCISIC ACID-INSENSITIVE3 (MtABI3), and MtWRINKLED-like (MtWRI), in M. orbicularis. The reduced oil in M. orbicularis, is consistent with increased synthesis of cell wall polysaccharides and decreased expression of master transcription factors regulating oil biosynthesis and embryo maturation. Comparative investigations between these two Medicago species is a useful system to investigate the regulation of oil content and carbon partitioning in legumes.

An update on source-to-sink carbon partitioning in tomato.

Plant growth and carbon metabolism are closely associated since carbohydrate in the form of sucrose generated by photosynthesis, provides the primary source of building blocks and energy for the production and maintenance of biomass. Regulation of carbon partitioning between source and sink tissues is important because it has a vast influence on both plant growth and development. The regulation of carbon partitioning at the whole plant level is directly linked to the cellular pathways of assimilate transport and the metabolism and allocation of sugars, mainly sucrose and hexoses in source leaves, and sink organs such as roots and fruit. By using tomato plant as a model, this review documents and discusses our current understanding of source–sink interactions from molecular to physiological perspectives focusing on those that regulate the growth and development of both vegetative and reproductive organs. It furthermore discusses the impact that environmental conditions play in maintenance of this balance in an attempt to address the link between physiological and ecological aspects of growth.

Carbon partitioning to the terpenoid biosynthetic pathway enables heterologous β-phellandrene production in Escherichia coli cultures.

Escherichia coli was used as a microbial system for the heterologous synthesis of β-phellandrene, a monoterpene of plant origin with several potential commercial applications. Expression of Lavandula angustifolia β-phellandrene synthase (PHLS), alone or in combination with Picea abies geranyl-diphosphate synthase in E. coli, resulted in no β-phellandrene accumulation, in sharp contrast to observations with PHLS-transformed cyanobacteria. Lack of β-phellandrene biosynthesis in E. coli was attributed to the limited endogenous carbon partitioning through the native 2-C-methylerythritol-4-phosphate (MEP) pathway. Heterologous co-expression of the mevalonic acid pathway, enhancing cellular carbon partitioning and flux toward the universal isoprenoid precursors, isopentenyl-diphosphate and dimethylallyl-diphosphate, was required to confer β-phellandrene production. Differences in endogenous carbon flux toward the synthesis of isoprenoids between photosynthetic (Synechocystis) and non-photosynthetic bacteria (E. coli) are discussed in terms of differences in the regulation of carbon partitioning through the MEP biosynthetic pathway in the two systems.

Carbon partitioning in photosynthesis

The work seeks to raise awareness of a fundamental problem that impacts the renewable generation of fuels and chemicals via (photo)synthetic biology. At issue is regulation of the endogenous cellular carbon partitioning between different biosynthetic pathways, over which the living cell exerts stringent control. The regulation of carbon partitioning in photosynthesis is not understood. In plants, microalgae and cyanobacteria, methods need be devised to alter photosynthetic carbon partitioning between the sugar, terpenoid, and fatty acid biosynthetic pathways, to lower the prevalence of sugar biosynthesis and correspondingly upregulate terpenoid and fatty acid hydrocarbons production in the cell. Insight from unusual but naturally occurring carbon-partitioning processes can help in the design of blueprints for improved photosynthetic fuels and chemicals production.

Regulation of biomass growth and carbon partitioning in poplar – molecular characterization of a candidate gene

Improvement of plant feedstock for bioenergy production can be achieved by modifying wood chemical properties and increasing biomass productivity. We previously identified a candidate gene for carbon partitioning and growth on chromosome 13 (cpg13) of poplar. Cpg13 was identified as the regulator of carbon partition and growth within a QTL interval that explains 56% of the variation in cellulose to lignin ratio, as well as 20-25% of the heritable variation in biomass. Putative homologues of cpg13 in Arabidopsis are annotated as proteins of unknown function; therefore, the functional characterization of cpg13 is essential. At present, evidence of the functional role of cpg13 is being obtained by the analysis of poplar transgenic plants transformed with RNAi, overexpression and GFP-fused – cpg13 constructs. Preliminary data indicates that down-regulating cpg13 positively impacts growth, while a negative effect is detected in lines overexpressing cpg13. Transgenic poplar 35S::cpg13:GFP shows localization in cell wall, consistent with in silico predictions. Comparative genomics indicate moderate similarity with methyltransferase. Analysis of the impact of differentially regulating cpg13 on lignin and cellulose is currently under way in the mutants. Furthermore, attempts to purify the cpg13 protein are in progress, to define its role through a series of biochemical function assays.

The Effect of Elevated Concentrations of Fructose 2,6-Bisphosphate on Carbon Metabolism during Deacidification in the Crassulacean Acid Metabolism Plant Kalanchöe daigremontiana.

In C(3) plants, the metabolite fructose 2,6-bisphosphate (Fru 2,6-P(2)) has an important role in the regulation of carbon partitioning during photosynthesis. To investigate the impact of Fru 2,6-P(2) on carbon metabolism during Crassulacean acid metabolism (CAM), we have developed an Agrobacterium tumefaciens-mediated transformation system in order to alter genetically the obligate CAM plant Kalanchöe daigremontiana. To our knowledge, this is the first report to use genetic manipulation of a CAM species to increase our understanding of this important form of plant metabolism. Transgenic plants were generated containing a modified rat liver 6-phosphofructo-2-kinase gene. In the plants analyzed the activity of 6-phosphofructo-2-kinase ranged from 175% to 198% of that observed in wild-type plants, resulting in Fru 2,6-P(2) concentrations that were 228% to 350% of wild-type plants after 2 h of illumination. A range of metabolic measurements were made on these transgenic plants to investigate the possible roles of Fru 2,6-P(2) during Suc, starch, and malic acid metabolism across the deacidification period of CAM. The results suggest that Fru 2,6-P(2) plays a major role in regulating partitioning between Suc and starch synthesis during photosynthesis. However, alterations in Fru 2,6-P(2) levels had little effect on malate mobilization during CAM fluxes.

Source-sink regulation by sugar and stress.

The regulation of carbon partitioning between source and sink tissues in higher plants is not only important for plant growth and development, but insight into the underlying regulatory mechanism is also a prerequisite to modulating assimilate partitioning in transgenic plants. Hexoses, as well as sucrose, have been recognised as important signal molecules in source-sink regulation. Components of the underlying signal transduction pathways have been identified and parallels, as well as distinct differences, to known pathways in yeast and animals have become apparent. There is accumulating evidence for crosstalk, modulation and integration between signalling pathways responding to phytohormones, phosphate, light, sugars, and biotic and abiotic stress-related stimuli. These complex interactions at the signal transduction levels and co-ordinated regulation of gene expression seem to play a central role in source-sink regulation.

Comparative evaluation of the effects of gaseous pollutants, acidic deposition, and mineral deficiencies on the carbohydrate metabolism of trees

With a focus on conifers, recent literature on the interaction of pollution, acidic deposition and mineral (mainly Mg) deficiency on plant carbohydrate metabolism is reviewed. By combining most recent knowledge about the regulation of carbon partitioning and allocation of photoassimilate with data on symptom-related pool sizes of carbohydrates, biochemical indications for shifts in allocation are presented. In summary, the data, although in part conflicting, indicate that the impact of pollutants at moderate levels (up to three times ambient concentration) should not be responsible for the decline symptoms observed.

Nitrate Activation of Cytosolic Protein Kinases Diverts Photosynthetic Carbon from Sucrose to Amino Acid Biosynthesis

The regulation of carbon partitioning between carbohydrates (principally sucrose) and amino acids has been only poorly characterized in higher plants. The hypothesis that the pathway of sucrose and amino acid biosynthesis compete for carbon skeletons and energy is widely accepted. In this review, we suggest a mechanism involving the regulation of cytosolic protein kinases whereby the flow of carbon is regulated at the level of partitioning between the pathways of carbohydrate and nitrogen metabolism via the covalent modulation of component enzymes. The addition of nitrate to wheat seedlings (Triticum aestivum) grown in the absence of exogenous nitrogen has a dramatic, if transient, impact on sucrose formation and on the activities of sucrose phosphate synthase (which is inactivated) and phosphoenolpyruvate carboxylase (which is activated). The activities of these two enzymes are modulated by protein phosphorylation in response to the addition of nitrate, but they respond in an inverse fashion. Sucrose phosphate synthase in inactivated and phosphoenolpyruvate carboxylase is activated. Nitrate functions as a signal metabolite activating the cytosolic protein kinase, thereby modulating the activities of at least two of the key enzymes in assimilate partitioning and redirecting the flow of carbon away from sucrose biosynthesis toward amino acid synthesis.

Regulation of carbon partitioning between sucrose and nitrogen assimilation in cotyledons of germinating Ricinus communis L. seedlings

The interactions between carbon and nitrogen metabolism in cotyledons of germinating Ricinus communis L. seedlings were investigated. The endosperm was removed for 6-d-old seedlings and their cotyledons were supplied with 50 mM glucose and mM potassium phosphate without a nitrogen source, or supplemented with 10 mM glutamine or 5 mM NH4Cl. [U(14)C] Glucose labelling patterns were used to investigate the effect of fluxes. Addition of glutamine or NH4Cl led to a 3.5- to 5-fold increase of labelling in amino acids (most of which were exported) and increased (14)CO2 release. Glutamine also led to a stimulation of glucose uptake, sucrose synthesis and export. Measurements of metabolites showed that glutamine or NH4Cl led to a decrease of a-ketoglutarate, pyruvate, phosphoenolpyruvate, glycerate-2-phosphate and glycerate-3-phosphate, a small increase of triose-phosphate and fructose-1,6-bisphosphate, a small decrease of hexose-phosphate (in the case of glutamine), and an increase of UDP glucose. In both treatments, fructose-2,6-bisphosphate doubled, and inorganic pyrophosphate decreased slightly. Similar results were obtained in detached cotyledons, except that glutamine did not alter the rate of glucose uptake or sucrose synthesis. The increased rate of sucrose synthesis after supplying glutamine to intact seedlings is ascribed to an increased rate of sucrose export from the cotyledons due to enhanced water flow in the phloem, brought about by loading of glutamine. The doubling of the rate of glycolysis after adding glutamine or NH4Cl in intact seedlings or excised cotyledons is ascribed to activation of the terminal enzymes in glycolysis, pyruvate kinase and phosphoenolpyruvate carboxylase. The resulting decrease of phosphoenolpyruvate and glycerate-3-phosphate leads to activation of phosphofructokinase. It also relieves inhibition of fructose-6-phosphate,2-kinase, leading to increased fructose-2,6-bisphosphate and activation of pyrophosphate; fructose-6-phosphate phosphotransferase.

Expression of a maize sucrose phosphate synthase in tomato alters leaf carbohydrate partitioning.

We isolated a complementary DNA sequence for the enzyme sucrose phosphate synthase (SPS) from maize utilizing a limited amino acid sequence. The 3509-bp cDNA encodes a 1068-amino acid polypeptide. The identity of the cDNA was confirmed by the ability of the cloned sequence to direct sucrose phosphate synthesis in Escherichia coli. Because no plant-specific factors were necessary for enzymatic activity, we can conclude that SPS enzyme activity is conferred by a single gene product. Sequence comparisons showed that SPS is distantly related to the enzyme sucrose synthase. When expressed from a ribulose bisphosphate carboxylase small subunit promoter in transgenic tomatoes, total SPS activity was boosted up to sixfold in leaves and appeared to be physiologically uncoupled from the tomato regulation mechanism. The elevated SPS activity caused a reduction of starch and increase of sucrose in the tomato leaves. This result clearly demonstrates that SPS is involved in the regulation of carbon partitioning in the leaves.

Tansley Review No. 27 The control of carbon partitioning in plants.

This review reports on the processes associated with carbon transfer and metabolism in leaves and growing organs and the role of long-distance transport and vascular links in the regulation of carbon partitioning in plants. Partitioning is clearly influenced by both the supply and demand for photosynthate and is moderated by vascular connections and the storage capacity of the leaves and pathway tissues. However there appears to be little more than circumstantial evidence either that short distance transfer of carbon within either the source or the sink, or that long-distance transport in the phloem, are limiting photosynthesis or growth directly. Although individual biochemical and physiological processes relating to photosynthesis and growth may be well understood, the factors primarily responsible for the control of carbon partitioning in plants have not been clearly identified. There is a need for a greater understanding of organ initiation and development (source and sink formation and potential size), the clear identification of whether growth is sink or source limited (including possible sink-controlled photosynthesis) and a detailed assessment of the role of storage in buffering developmental and environmental changes in sink and source activity. Also more information is needed on the role of hormonal and nutritional factors in regulating source and sink activity (organ interactions not directly associated with carbon transfer). CONTENTS Summary 341 I. Introduction 342 II. General source-sink relationships 342 III. Control at the source 345 IV. The utilization of photosynthate: sink characteristics and limitations 353 V. Vascular constraints and temporary storage 360 VI. Concluding comments 366 Acknowledgements 366 References 367.

Energy and redox status, and carbon allocation in one- to three-year-old spruce needles

Metabolite pools were determined in lyophilized homogenates of one- to three-year-old needles from 13-year-old spruce trees. These were exposed in open-chambers for five consecutive years to SO 2 (30 μg m −3) and O 3 (50 to 180 μg m −3), singly and in combination. The samples were taken shortly before bud break (mid-March). The energy (ATP/ADP) and redox status (NAD(P)H/NAD(P)) increased with needle age. This increase was most pronounced in needles exposed to both pollutant gases. To evaluate carbohydrate metabolism, levels of starch, sugars (sucrose, fructose, glucose), sugar phosphates (dihydroxyacetone phosphate, glyceraldehyde 3-phosphate (triose phosphates, TP), fructose 6-phosphate, fructose 2,6-bisphosphate: F2,6BP) and 3-phosphoglyceric acid (PGA) were measured. The concentrations of starch, glucose, fructose and TP on a dry weight basis increased with needle age, while those of sucrose and F2,6BP decreased. Interestingly, the total amount of soluble sugars remained constant in needles from control chambers and large fluctuations mainly occurred after a single application of the pollutants. Needles from fumigated trees generally showed somewhat lower starch and sugar, and fructose 6-phosphate levels. The content of TP increased with needle age while that of sucrose and F2,6BP decreased. This inverse behavior was even more prominent in the presence of both pollutants and is discussed in the context of recent knowledge on the regulation of carbon partitioning. There was a significant increase in the ratio of TP/PGA in samples from the combined exposure treatments. This also indicates an increase in the redox charge and corroborates the data on both the redox status of the pyridine nucleotide system and the significantly increased ATP/ADP ratio. The results are indicative of a switch in metabolism, from anabolic to predominantly catabolic reactions, which is most pronounced in the samples fumigated with both O 3 and SO 2. The data are compared to those obtained from naturally exposed trees in the southern Black Forest (Kälbelescheuer/Haldenhof, near Freiburg, Germany).

Regulation of Carbon Partitioning in Photosynthetic Tissue

Regulation of Carbon Partitioning in Photosynthetic Tissue

REVIEWS

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