Carbohydrate Export Research Articles

Co-overexpression of SWEET sucrose transporters modulates sucrose synthesis and defence responses to enhance immunity against bacterial blight in rice.

Enhancing carbohydrate export from source to sink tissues is considered to be a realistic approach for improving photosynthetic efficiency and crop yield. The rice sucrose transporters OsSUT1, OsSWEET11aand OsSWEET14 contribute to sucrose phloem loading and seed filling. Crucially, Xanthomonas oryzae pv. oryzae (Xoo) infection in rice enhances the expression of OsSWEET11a and OsSWEET14 genes, and causes leaf blight. Here we show that co-overexpression of OsSUT1, OsSWEET11aand OsSWEET14 in rice reduced sucrose synthesis and transport leading to lower growth and yield but reduced susceptibility to Xoo relative to controls. The immunity-related hypersensitive response (HR) was enhanced in the transformed lines as indicated by the increased expression of defence genes, higher salicylic acidcontentand presence of HR lesions on the leaves. The results suggest that the increased expression of OsSWEET11a and OsSWEET14 in rice is perceived as a pathogen (Xoo) attack that triggers HR and results in constitutive activation of plant defences that are related to the signalling pathways of pathogen starvation. These findings provide a mechanistic basis for the trade-off between plant growth and immunity because decreased susceptibility against Xoo compromised plant growth and yield.

Phloem and Xylem Responses Are Both Implicated in Huanglongbing Tolerance of Sugar Belle.

Although huanglongbing (HLB) is a devastating citrus disease, improved tolerant cultivars, such as Sugar Belle (SB) mandarin, have been identified. To understand the responses that HLB-affected SB undergoes, we compared 14CO2 fixation, carbohydrate export, phloem callose accumulation, relative expression of plant defense activators, and anatomical changes between healthy and infected SB trees versus susceptible Pineapple (PA) sweet orange. Eight- to ten-week-old leaves of infected SB showed a 2.5-fold increase in 14CO2 fixation and a 13% decrease in 14C-carbohydrate export, whereas HLB-affected PA presented a decrease of 33 and 50%, respectively. The mean distance of a callose deposit to its closest neighbor was 36% smaller in infected SB versus healthy, whereas in HLB-affected PA, it was 33% higher. Expression of papain-like cysteine proteases (PLCPs) was upregulated in SB but downregulated in PA. Infected SB showed minor alterations in the number of xylem vessels, a 16% larger xylem vessel lumen area, and a 14% increase in the proportional area of the xylem. In contrast, PA showed a 2.4-fold increase in the xylem vessel number and a 2% increase in the proportional xylem area. Three complementary mechanisms of tolerance in SB are hypothesized: (i) increased carbohydrate availability induced by greater CO2 fixation, mild effect in carbohydrate export, and local accumulation of callose in the phloem; (ii) activation of defense response via upregulation of PLCPs, and (iii) increased investment in the xylem structure. Thus, phloem and xylem modifications seem to be involved in SB tolerance.

Exogenous Ca2+ priming can improve peanut photosynthetic carbon fixation and pod yield under early sowing scenarios in the field.

Harnessing cold-resilient and calcium-enriched peanut production technology are crucial for high-yielding peanut cultivation in high-latitude areas. However, there is limited field data about how exogenous calcium (Ca2+) application would improve peanut growth resilience during exposure to chilling stress at early sowing (ES). To help address this problem, a two-year field study was conducted to assess the effects of exogenous foliar Ca2+ application on photosynthetic carbon fixation and pod yield in peanuts under different sowing scenarios. We measured plant growth indexes, leaf photosynthetic gas exchange, photosystems activities, and yield in peanuts. It was indicated that ES chilling stress at the peanut seedling stage led to the reduction of Pn, gs, Tr, Ls, WUE, respectively, and the excessive accumulation of non-structural carbohydrates in leaves, which eventually induced a chilling-dependent feedback inhibition of photosynthesis due mainly to weaken growth/sink demand. While exogenous Ca2+ foliar application improved the export of nonstructural carbohydrates, and photosynthetic capacity, meanwhile activated cyclic electron flow, thereby enhancing growth and biomass accumulation in peanut seedlings undergoing ES chilling stress. Furthermore, ES combined with exogenous Ca2+ application can significantly enhance plant chilling resistance and peanut yield ultimately in the field. In summary, the above results demonstrated that exogenous foliar Ca2+ application restored the ES-linked feedback inhibition of photosynthesis, enhancing the growth/sink demand and the yield of peanuts.

Citrus photosynthesis and morphology acclimate to phloem-affecting huanglongbing disease at the leaf and shoot levels.

Huanglongbing (HLB) is a phloem-affecting disease in citrus that reduces growth and impacts global citrus production. HLB is caused by a phloem-limited bacterium (Candidatus Liberibacter asiaticus). By inhibiting phloem function, HLB stunts sink growth, including the production of new shoots and leaves, and induces hyperaccumulation of foliar starch. HLB induces feedback inhibition of photosynthesis by reducing foliar carbohydrate export. Here, we assessed the relationship of bacterial distribution within the foliage, foliar starch accumulation, and net CO2 assimilation (Anet ). Because HLB impacts canopy morphology, we developed a chamber to measure whole-shoot Anet to test the effects of HLB at both the leaf and shoot level. Whole-shoot level Anet saturated at high irradiance, and green stems had high photosynthetic rates compared to leaves. Starch accumulation was correlated with bacterial population, and starch was negatively correlated with Anet at the leaf but not at the shoot level. Starch increased initially after infection, then decreased progressively with increasing length of infection. HLB infection reduced Anet at the leaf level but increased it at the whole-shoot level, in association with reduced leaf size and greater relative contribution of stems to the photosynthetic surface area. Although HLB-increased photosynthetic efficiency, total carbon fixed per shoot decreased because photosynthetic surface area was reduced. We conclude that the localized effects of infection on photosynthesis are mitigated by whole-shoot morphological acclimation over time. Stems contribute important proportions of whole-shoot Anet , and these contributions are likely increased by the morphological acclimation induced by HLB.

Phloem transport limitation in Huanglongbing-affected sweet orange is dependent on phloem-limited bacteria and callose.

Huanglongbing (HLB), caused by Candidatus `Liberibacter asiaticus' (CLas), is a phloem-limited disease that disrupts citrus production in affected areas. In HLB-affected plants, phloem sieve plate pores accumulate callose, and leaf carbohydrate export is reduced. However, whether HLB causes a reduction in carbohydrate phloem translocation speed and the quantitative relationships among callose, CLas population and phloem translocation are still unknown. In this work, a procedure was developed to concurrently measure sugar transport, callose deposition and relative pathogen population at different locations throughout the stem. Increasing quantities of CLas genetic material were positively correlated with quantity and density of callose deposits and negatively correlated with phloem translocation speed. Callose deposit quantity was position and rootstock dependent and was negatively correlated with phloem translocation speed, suggesting a localized relationship. Remarkably, callose accumulation and phloem translocation disruption in the scion were dependent on rootstock genotype. Regression results suggested that the interaction of Ct values and number of phloem callose depositions, but not their size or density, explained the effects on translocation speed. Sucrose, starch and sink 14C label allocation data support the interpretation of a transport pathway limitation by CLas infection. This work shows that the interaction of local accumulation of callose and CLas affects phloem transport. Furthermore, the extent of this accumulation is attenuated by the rootstock and provides important information about the disease mechanism of phloem-inhabiting bacteria. Together, these results constitute the first example of a demonstrated transport limitation of phloem function by a microbial infection.

The Complementary Roles of Chloroplast Cyclic Electron Transport and Mitochondrial Alternative Oxidase to Ensure Photosynthetic Performance.

Chloroplasts use light energy and a linear electron transport (LET) pathway for the coupled generation of NADPH and ATP. It is widely accepted that the production ratio of ATP to NADPH is usually less than required to fulfill the energetic needs of the chloroplast. Left uncorrected, this would quickly result in an over-reduction of the stromal pyridine nucleotide pool (i.e., high NADPH/NADP+ ratio) and under-energization of the stromal adenine nucleotide pool (i.e., low ATP/ADP ratio). These imbalances could cause metabolic bottlenecks, as well as increased generation of damaging reactive oxygen species. Chloroplast cyclic electron transport (CET) and the chloroplast malate valve could each act to prevent stromal over-reduction, albeit in distinct ways. CET avoids the NADPH production associated with LET, while the malate valve consumes the NADPH associated with LET. CET could operate by one of two different pathways, depending upon the chloroplast ATP demand. The NADH dehydrogenase-like pathway yields a higher ATP return per electron flux than the pathway involving PROTON GRADIENT REGULATION5 (PGR5) and PGR5-LIKE PHOTOSYNTHETIC PHENOTYPE1 (PGRL1). Similarly, the malate valve could couple with one of two different mitochondrial electron transport pathways, depending upon the cytosolic ATP demand. The cytochrome pathway yields a higher ATP return per electron flux than the alternative oxidase (AOX) pathway. In both Arabidopsis thaliana and Chlamydomonas reinhardtii, PGR5/PGRL1 pathway mutants have increased amounts of AOX, suggesting complementary roles for these two lesser-ATP yielding mechanisms of preventing stromal over-reduction. These two pathways may become most relevant under environmental stress conditions that lower the ATP demands for carbon fixation and carbohydrate export.

Maize Brittle Stalk2-Like3, encoding a COBRA protein, functions in cell wall formation and carbohydrate partitioning.

Carbohydrate partitioning from leaves to sink tissues is essential for plant growth and development. The maize (Zea mays) recessive carbohydrate partitioning defective28 (cpd28) and cpd47 mutants exhibit leaf chlorosis and accumulation of starch and soluble sugars. Transport studies with 14C-sucrose (Suc) found drastically decreased export from mature leaves in cpd28 and cpd47 mutants relative to wild-type siblings. Consistent with decreased Suc export, cpd28 mutants exhibited decreased phloem pressure in mature leaves, and altered phloem cell wall ultrastructure in immature and mature leaves. We identified the causative mutations in the Brittle Stalk2-Like3 (Bk2L3) gene, a member of the COBRA family, which is involved in cell wall development across angiosperms. None of the previously characterized COBRA genes are reported to affect carbohydrate export. Consistent with other characterized COBRA members, the BK2L3 protein localized to the plasma membrane, and the mutants condition a dwarf phenotype in dark-grown shoots and primary roots, as well as the loss of anisotropic cell elongation in the root elongation zone. Likewise, both mutants exhibit a significant cellulose deficiency in mature leaves. Therefore, Bk2L3 functions in tissue growth and cell wall development, and this work elucidates a unique connection between cellulose deposition in the phloem and whole-plant carbohydrate partitioning.

Mitochondria Matter: Systemic Aspects of Nonalcoholic Fatty Liver Disease (NAFLD) and Diagnostic Assessment of Liver Function by Stable Isotope Dynamic Breath Tests.

The liver plays a key role in systemic metabolic processes, which include detoxification, synthesis, storage, and export of carbohydrates, lipids, and proteins. The raising trends of obesity and metabolic disorders worldwide is often associated with the nonalcoholic fatty liver disease (NAFLD), which has become the most frequent type of chronic liver disorder with risk of progression to cirrhosis and hepatocellular carcinoma. Liver mitochondria play a key role in degrading the pathways of carbohydrates, proteins, lipids, and xenobiotics, and to provide energy for the body cells. The morphological and functional integrity of mitochondria guarantee the proper functioning of β-oxidation of free fatty acids and of the tricarboxylic acid cycle. Evaluation of the liver in clinical medicine needs to be accurate in NAFLD patients and includes history, physical exam, imaging, and laboratory assays. Evaluation of mitochondrial function in chronic liver disease and NAFLD is now possible by novel diagnostic tools. “Dynamic” liver function tests include the breath test (BT) based on the use of substrates marked with the non-radioactive, naturally occurring stable isotope 13C. Hepatocellular metabolization of the substrate will generate 13CO2, which is excreted in breath and measured by mass spectrometry or infrared spectroscopy. Breath levels of 13CO2 are biomarkers of specific metabolic processes occurring in the hepatocyte cytosol, microsomes, and mitochondria. 13C-BTs explore distinct chronic liver diseases including simple liver steatosis, non-alcoholic steatohepatitis, liver fibrosis, cirrhosis, hepatocellular carcinoma, drug, and alcohol effects. In NAFLD, 13C-BT use substrates such as α-ketoisocaproic acid, methionine, and octanoic acid to assess mitochondrial oxidation capacity which can be impaired at an early stage of disease. 13C-BTs represent an indirect, cost-effective, and easy method to evaluate dynamic liver function. Further applications are expected in clinical medicine. In this review, we discuss the involvement of liver mitochondria in the progression of NAFLD, together with the role of 13C-BT in assessing mitochondrial function and its potential use in the prevention and management of NAFLD.

Proteomic Responses to Alkali Stress in Oats and the Alleviatory Effects of Exogenous Spermine Application.

Alkali stress limits plant growth and yield more strongly than salt stress and can lead to the appearance of yellow leaves; however, the reasons remain unclear. In this study, we found that (1) the down-regulation of coproporphyrinogen III oxidase, protoporphyrinogen oxidase, and Pheophorbide a oxygenase in oats under alkali stress contributes to the appearance of yellow leaves (as assessed by proteome and western blot analyses). (2) Some oat proteins that are involved in the antioxidant system, root growth, and jasmonic acid (JA) and indole-3-acetic acid (IAA) synthesis are up-regulated in response to alkalinity and help increase alkali tolerance. (3) We added exogenous spermine to oat plants to improve their alkali tolerance, which resulted in higher chlorophyll contents and plant dry weights than in plants subjected to alkaline stress alone. This was due to up-regulation of chitinase and proteins related to chloroplast structure, root growth, and the antioxidant system. Spermine addition increased sucrose utilization efficiency, and promoted carbohydrate export from leaves to roots to increase energy storage in roots. Spermine addition also increased the IAA and JA contents required for root growth.

Exogenous Calcium Alleviates Nocturnal Chilling-Induced Feedback Inhibition of Photosynthesis by Improving Sink Demand in Peanut (Arachis hypogaea).

Arachis hypogaea (peanut) is a globally important oilseed crop with high nutritional value. However, upon exposure to overnight chilling stress, it shows poor growth and seedling necrosis in many cultivation areas worldwide. Calcium (Ca2+) enhances chilling resistance in various plant species. We undertook a pot experiment to investigate the effects of exogenous Ca2+ and a calmodulin (CaM) inhibitor on growth and photosynthetic characteristics of peanut exposed to low night temperature (LNT) stress following warm sunny days. The LNT stress reduced growth, leaf extension, biomass accumulation, gas exchange rates, and photosynthetic electron transport rates. Following LNT stress, we observed larger starch grains and a concomitant increase in nonstructural carbohydrates and hydrogen peroxide (H2O2) concentrations. The LNT stress further induced photoinhibition and caused structural damage to the chloroplast grana. Exogenous Ca2+ enhanced plant growth following LNT stress, possibly by allowing continued export of carbohydrates from leaves. Foliar Ca2+ likely alleviated the nocturnal chilling-dependent feedback limitation on photosynthesis in the daytime by increasing sink demand. The foliar Ca2+ pretreatment protected the photosystems from photoinhibition by facilitating cyclic electron flow (CEF) and decreasing the proton gradient (ΔpH) across thylakoid membranes during LNT stress. Foliar application of a CaM inhibitor increased the negative impact of LNT stress on photosynthetic processes, confirming that Ca2+–CaM played an important role in alleviating photosynthetic inhibition due to the overnight chilling-dependent feedback.

Comparative study on the different responses of maize photosynthesis to systemic regulation under light heterogeneity

Photosynthetic performance of a leaf is widely recognized to be systemically regulated by distal parts within the same plant. However, the effects of systemic regulation on different plant materials cannot be generalized. In this work, two cultivars of maize (Zea mays L.), ‘Rongyu 1210’ (RY) and ‘Zhongdan 808’ (ZD), were selected for a comparative study on the different responses of photosynthesis to light-dependent systemic regulation. After the growth of plants in heterogeneous light, the net photosynthetic rate of newly developed leaves increased in RY but decreased in ZD. A distinct capacity of CO2 fixation and assimilation between these two cultivars is also suggested. In ZD, the area of vascular bundles declined obviously, suggesting a restriction on carbohydrate export, which is also indicated by an increase in starch content. Resulting excessive accumulation of carbohydrates is proposed to inhibit the carbon assimilation, and eventually the photosynthesis. A decline in the area of bundle sheath cells also suggests a restriction on carbon assimilation. In contrast, these restrictions were unlikely to present in RY. This study reveals that the response of leaf photosynthetic performance to light heterogeneity is largely dependent on the systemic regulation of carbon assimilation, as well as carbohydrate export in maize.

Relationship among night temperature, carbohydrate translocation and inhibition of grapevine leaf photosynthesis

Important processes such as plant respiration, repair of damaged photosystems and carbohydrate translocation occurs during night. The lack of photosyntate translocation can influence photosynthetic efficiency of the leaves in the subsequent day. Night temperature can play and important role influencing the carbohydrate translocation. The aim of the work was to test the effect of night temperature on the photosynthetic activity, and to assess the correlation between the accumulation of carbohydrates in the leaves due to the reduction of their overnight translocation. In two experiments, taking into account whole canopy and single leaf CO2 assimilation of Vitis vinifera, night temperature was correlated with the variation of CO2 assimilation between the afternoon and the subsequent morning. In a controlled experiment, such variation was correlated with the accumulation of carbohydrates in the leaves in particular for low night temperatures, suggesting a mechanism of end product feedback down regulation. High night temperature caused a poor export of carbohydrates from the leaf, but the increase of respiration caused a consistent decrease of the carbohydrate content in the leaf. In conclusion, night temperature plays a pivotal role in the carbon translocation during the night. Further to its impact on source-sink relationship, this mechanism can have an important impact on the CO2 assimilation of the plant during the day that can be negatively affected either by relatively low and relatively high temperature during the night.

24-Epibrassinolide Pre-Treatment Modifies Cold-Induced Photosynthetic Acclimation Mechanisms and Phytohormone Response of Perennial Ryegrass in Cultivar-Dependent Manner

We investigated the modification of cold-induced mechanisms of photosynthetic apparatus adjustment and phytohormone response by brassinosteroid 24-epibrassinolide (EBR) and its consequences for frost tolerance of perennial ryegrass (Lolium perenne L.). We recorded the responses of two cultivars with contrasting frost tolerances to foliar hormone application, both in non-acclimated plants and plants cold acclimated for 3 and 6 weeks at 4 °C. In non-cold-acclimated plants of both cultivars, EBR induced increases in carbon fixation and lowered sucrose levels. Temporary suppression in quantum efficiency of PSII of photosystem II and activities of ribulose-1,5-bisphosphate carboxylase/oxygenase and sucrose phosphate synthase, a consequence of energy dissipation in non-photochemical quenching, was observed in the leaves of the highly frost-tolerant cultivar after 3 weeks of cold acclimation. After 6 weeks of cold acclimation, EBR accelerated recovery of photosynthesis, reflecting adjustment to cold conditions, and increased frost tolerance. As carbohydrate export from leaves is favored during cold acclimation, EBR application did not increase frost tolerance of the moderately tolerant cultivar, reflecting the downregulation of photosynthesis due to high leaf sucrose concentrations. It is also likely that EBR participated in the enhancement of frost tolerance by regulation of stress-related signaling compounds such as JA and ethylene but not SA, in winter ryegrass undergoing cold acclimation. Taken together, our results demonstrate that EBR-induced changes are temperature dependent. The beneficial effect of EBR is not universal under cold conditions, as genetically determined mechanisms are apparently dominant relative to EBR action.

Down-regulation of tomato PHYTOL KINASE strongly impairs tocopherol biosynthesis and affects prenyllipid metabolism in an organ-specific manner.

Tocopherol, a compound with vitamin E (VTE) activity, is a conserved constituent of the plastidial antioxidant network in photosynthetic organisms. The synthesis of tocopherol involves the condensation of an aromatic head group with an isoprenoid prenyl side chain. The latter, phytyl diphosphate, can be derived from chlorophyll phytol tail recycling, which depends on phytol kinase (VTE5) activity. How plants co-ordinate isoprenoid precursor distribution for supplying biosynthesis of tocopherol and other prenyllipids in different organs is poorly understood. Here, Solanum lycopersicum plants impaired in the expression of two VTE5-like genes identified by phylogenetic analyses, named SlVTE5 and SlFOLK, were characterized. Our data show that while SlFOLK does not affect tocopherol content, the production of this metabolite is >80% dependent on SlVTE5 in tomato, in both leaves and fruits. VTE5 deficiency greatly impacted lipid metabolism, including prenylquinones, carotenoids, and fatty acid phytyl esters. However, the prenyllipid profile greatly differed between source and sink organs, revealing organ-specific metabolic adjustments in tomato. Additionally, VTE5-deficient plants displayed starch accumulation and lower CO2 assimilation in leaves associated with mild yield penalty. Taken together, our results provide valuable insights into the distinct regulation of isoprenoid metabolism in leaves and fruits and also expose the interaction between lipid and carbon metabolism, which results in carbohydrate export blockage in the VTE5-deficient plants, affecting tomato fruit quality.

Carbohydrate regulation of photosynthesis and respiration from branch girdling in four species of wet tropical rain forest trees.

How trees sense source-sink carbon balance remains unclear. One potential mechanism is a feedback from non-structural carbohydrates regulating photosynthesis and removing excess as waste respiration when the balance of photosynthesis against growth and metabolic activity changes. We tested this carbohydrate regulation of photosynthesis and respiration using branch girdling in four tree species in a wet tropical rainforest in Costa Rica. Because girdling severs phloem to stop carbohydrate export while leaving xylem intact to allow photosynthesis, we expected carbohydrates to accumulate in leaves to simulate a carbon imbalance. We varied girdling intensity by removing phloem in increments of one-quarter of the circumference (zero, one--quarter, half, three-quarters, full) and surrounded a target branch with fully girdled ones to create a gradient in leaf carbohydrate content. Light saturated photosynthesis rate was measured in situ, and foliar respiration rate and leaf carbohydrate content were measured after destructive harvest at the end of the treatment. Girdling intensity created no consistent or strong responses in leaf carbohydrates. Glucose and fructose slightly increased in all species by 3.4% per one-quarter girdle, total carbon content and leaf mass per area increased only in one species by 5.4 and 5.5% per one-quarter girdle, and starch did not change. Only full girdling lowered photosynthesis in three of four species by 59-69%, but the decrease in photosynthesis was unrelated to the increase in glucose and fructose content. Girdling did not affect respiration. The results suggest that leaf carbohydrate content remains relatively constant under carbon imbalance, and any changes are unlikely to regulate photosynthesis or respiration. Because girdling also stops the export of hormones and reactive oxygen species, girdling may induce physiological changes unrelated to carbohydrate accumulation and may not be an effective method to study carbohydrate feedback in leaves. In three species, removal of three-quarters of phloem area did not cause leaf carbohydrates to accumulate nor did it change photosynthesis or respiration, suggesting that phloem transport is flexible and transport rate per unit phloem can rapidly increase under an increase in carbohydrate supply relative to phloem area. Leaf carbohydrate content thus may be decoupled from whole plant carbon balance by phloem transport in some species, and carbohydrate regulation of photosynthesis and respiration may not be as common in trees as previous girdling studies suggest. Further studies in carbohydrate regulation should avoid using girdling as girdling can decrease photosynthesis through unintended means without the tested mechanisms of accumulating leaf carbohydrates.

Diel patterns of leaf carbohydrate concentrations differ between seedlings and mature trees of two sympatric oak species

A fundamental aspect of the carbon cycle is the exchange of carbon between plants and the atmosphere. It is, therefore, important to understand factors that affect differences in gas exchange and carbon balance within and among species. Concentrations of nonstructural carbohydrates are often used as a proxy for carbon balance. We determined diel patterns of leaf carbohydrate concentrations in relation to irradiance (sun vs. shade) in seedlings and mature trees of two sympatric oak species (Quercus virginiana Mill. and Quercus hemisphaerica Bartram ex Willd.). For seedlings, we also measured leaf gas exchange. Higher sun exposure significantly increased photosynthesis and carbohydrate concentrations in both species. Carbohydrate concentrations of seedling leaves showed strong diel fluctuations, whereas concentrations in mature tree leaves did not. This contrast might be attributed to faster carbohydrate export from leaves of mature trees. The difference in sink strength between seedlings and adults may be related to the decreasing ratio of leaf mass to plant mass with ontogeny, increasing the demand for carbohydrates per unit leaf mass. Seedlings and mature trees are clearly functionally different and care must be taken when extrapolating results from seedling experiments to mature trees.

Source-Sink Partitioning of Mineral Nutrients and Photo-assimilates in Tomato Plants Grown under Suboptimal Nutrition

A huge number of greenhouse soils in Korea have accumulated mineral elements which induce many nutritional and pathological problems. The present study was performed to the effects of the reduced fertilization on plant growth, and uptake and partitioning of minerals (N, P, K) and soluble carbohydrates using highly minerals-accumulated farmer's greenhouse soil. On the basis of the recommended application for tomato crop, the application rates of N, P and K were 110(50%)-5.2(5%)-41.5(35%)kg <TEX>$ha^{-1}$</TEX>, respectively, using Hoagland's nutrient solution. Tomato growth rates during the whole experiment were not significant between treatments, but it was found that a decrease in daily growth represented after 60 days of treatment (DAT). The reduced application led to a drastic decrease in the concentration of N, P and K in fruits, and, thus, this resulted in lower uptake after 40 DAT. The lower phloem export and utilization of soluble carbohydrates caused an accumulation of extra-carbohydrates in leaves, stems and fruits in the reduced application. The reduced fertilization induced the capture of N, P and K in leaves and of soluble carbohydrates in stems compared to the conventional application. In this study, we suggest that it is possible to delay the first fertigation time in minerals-accumulated soils without an adverse impact on crop growth, but it is necessary to regularly monitor mineral status in soil to ensure a balanced uptake, synthesis and partitioning of minerals and carbohydrates.

The State Budget in Q3 2013

According to data released by the RF Federal Treasury, in Q3 2013 the decline in federal budget revenues was occurring at a slightly slower rate than previously. Thus, over the course of January-September 2013, the federal budget’s revenue and its oil and gas revenues dropped by 1.7 p.p. of GDP and 0.9 p.p. of GDP respectively on the same period of 2012, while over the course of the first half year of 2013 they had dwindled by 2.1 p.p. of GDP and 1.6 p.p. of GDP respectively on the first half year of 2012. In January-August 2013, the consolidated budget revenue of RF subjects amounted to 12.1% of GDP, which represented a 0.3 p.p. of GDP rise on their consolidated budget revenue in January-June 2013. At the same time, Russia’s budget system has very small potential for further growth in revenues, which is especially true of the consolidated budgets of those RF subjects that are not engaged in the export of carbohydrates. Therefore, apart from taking measures designed to optimize federal budget spending, the RF Government must pay special attention to the state of regional budgets and, maybe, to revise its approach to the sphere of inter-budget relations.

Diurnal dynamics of maize leaf photosynthesis and carbohydrate concentrations in response to differential N availability

Diurnal changes of the carbohydrates in maize leaves reflect transitions in photosynthesis, carbohydrate metabolism and translocation rates, and can provide insights into the control of carbohydrate export from source leaves. Relationships of diurnal courses of leaf photosynthetic rates (A), nonstructural carbohydrates (NSC), and ear dry matter accumulation were investigated in field grown maize under different nitrogen (N) regimes. Nitrogen applications increased plant N uptake and grain yield as well as leaf area, chlorophyll content and soluble protein concentration. No differences in A among N treatments were observed during periods of low light intensities early and late in the day. The largest difference of A among N treatments appeared at 12:00PM at silking, and shifted to 15:00PM at 15 and 30 days after silking (DAS). In general, leaf glucose levels tended to be lower in N fertilized than unfertilized plants. At silking, the diurnal pattern of glucose was similar to that of A, reaching a peak between 9:00AM and 12:00PM. Clear diurnal pattern of leaf sucrose and starch concentrations were found at each sampling date, albeit less pronounced as development progressed from silking, to 15 and 30 DAS. Sucrose concentrations were lower in unfertilized than in N fertilized plants for most of the 24-h cycle at all three developmental stages. At silking and 15 DAS, starch concentrations were significantly greater in unfertilized compared to N fertilized plants. Ear leaf NSC dynamics observed here suggest that factors (i.e. source–sink dynamics) other than carbon assimilation limit maize yield under low N conditions.

Integrated operation of the photorespiratory cycle and cytosolic metabolism in the modulation of primary nitrogen assimilation and export of organic N-transport compounds from leaves: A hypothesis

Photorespiration is generally considered to be an essentially dissipative process, although it performs some protective and essential functions. A theoretical appraisal indicates that the loss of freshly assimilated CO2 due to photorespiration in well-watered plants may not be as high as generally believed. Even under moderately adverse conditions, these losses may not exceed 10%. The photorespiratory metabolism of the source leaves of well-watered and well-nourished crop plants ought to be different from that of other leaves because the fluxes of the export of both carbohydrates and organic N-transport compounds in source leaves is quite high. With a heuristic approach that involved the dovetailing of certain metabolic steps with the photorespiratory cycle (PR-cycle), a novel network is proposed to operate in the source-leaves of well-watered and well-nourished plants. This network allows for the diversion of metabolites from their cyclic-routes in sizeable quantities. With the removal of considerable quantities of glycine and serine from the cyclic route, the number of RuBP oxygenation events would be several times those of the formation of hydroxypyruvate. Thus, to an extreme extent, photorespiratory metabolism would become open-ended and involve much less futile recycling of glycine and serine. Conversion of glyoxylate to glycine has been proposed to be a crucial step in the determination of the relative rates of the futile (cyclic) and anabolic (open-ended) routes. Thus, in the source leaves of well-watered and well-nourished plants, the importance of the cyclic route is limited to the salvaging of photorespiratory intermediates for the regeneration of RuBP. The proposed network is resilient enough to coordinate the rates of the assimilation of carbon and nitrogen in accordance with the moisture and N-fertility statuses of the soil.