Effects of carbohydrate accumulation on photosynthesis differ between sink and source leaves of Phaseolus vulgaris L

Accumulation of non-structural carbohydrate in leaves represses photosynthesis. However, the extent of repression should be different between sink leaves (sugar consumers) and source leaves (sugar exporters). We investigated the effects of carbohydrate accumulation on photosynthesis in the primary leaves of bean (Phaseolus vulgaris L.) during leaf expansion. To increase the carbohydrate content of the leaves, we supplied 20 mM sucrose solution to the roots for 5 d (sugar treatment). Plants supplied only with water and nutrients were used as controls. The carbohydrate contents, which are the sum of glucose, sucrose and starch, of the sugar-treated leaves were 1.5-3 times of those of the control leaves at all developmental stages. In the young sink leaves, the photosynthetic rate at saturating light and at an ambient CO2 concentration (A360) did not differ between the sugar-treated and control leaves. The A360 of sugar-treated source leaves gradually decreased relative to the control source leaves with leaf expansion. The initial slope of the A-Ci (CO2 concentration in the intercellular space) curve, and the Rubisco (ribulose-1,5-bisphosphate carboxylase/oxygenase) content per leaf area showed trends similar to that of A360. Differences in Amax between the treatments were slightly smaller than those in A360. These results indicate that the effect of carbohydrate accumulation on photosynthesis is significant in the source leaves, but not in the young sink leaves, and that the decrease in Rubisco content was the main cause of the carbohydrate repression of photosynthesis.     

Regulation of photosynthesis through source: sink imbalance in citrus is mediated by carbohydrate content in leaves

In citrus, the occurrence of a sink effect on photosynthesis (A) is controversial. Leaf carbohydrates and photosynthetic rates in field-grown trees of Satsuma mandarin ( Citrus unshiu [Mak.] Marc.) cv. Okitsu, were measured to elucidate whether or not the demand for photoassimilates regulates A. The data indicated that the source-sink imbalances induced by different treatments altered both soluble (sucrose, glucose and fructose) and insoluble carbohydrates in leaves, as well as photosynthetic rates. In general, girdling and defruiting increased starch and reduced photosynthesis, whereas source-limiting conditions imposed through partial defoliations had the opposite effect. These results are compatible with the assumption that a lack of sink activity leads to carbohydrate accumulation and feedback inhibition of A, and vice versa. Further evidence supporting a source-sink effect on A was provided by measurements of the dry matter:leaf area ratio, since defoliations, for example, increased this ratio. The in vivo sucrose supplementation to plants with different source:sink ratios (control, defoliated, girdled and defruited plants) increased carbohydrates and reduced photosynthesis. This suggests that sugars may have, per se, the potential to repress photosynthetic rates in intact plants with active sinks. Based on these results we propose that sugar accumulation in citrus leaves causes a feedback inhibition of A.     

Photosynthesis and Plant Growth at Elevated Levels of CO2

In this review, we discuss the effects of elevated CO₂ levelson photosynthesis in relation to the whole plant growth in terrestrialhigher C₃ plants. Short-term CO₂ enrichment stimulates the rateof photosynthesis. Plant mass is also enhanced by CO₂ enrichment.However, the effects of long-term CO₂ enrichment on photosynthesisare variable. Generally, the prolonged exposure to CO₂ enrichmentreduces the initial stimulation of photosynthesis in many species,and frequently suppresses photosynthesis. These responses areattributed to secondary responses related to either excess carbohydrateaccumulation or decreased N content rather than direct responsesto CO₂. Accumulation of carbohydrates in leaves may lead tothe repression of photosynthetic gene expression and excessstarch seems to hinder CO₂ diffusion. Therefore, the specieswhich have the sink organs for carbohydrate accumulation donot show the suppression of photosynthesis. The suppressionof photosynthesis by CO₂ enrichment is always associated withdecreases in leaf N and Rubisco contents. These decreases arenot due to dilution of N caused by a relative increase in theplant mass but are the result of a decrease in N allocationto leaves at the level of the whole plant, and the decreasein Rubisco content is not selective. Leaf senescence and plantdevelopment are also accelerated by CO₂ enrichment. However,they are independent of each other in some species. Thus, variousresponses to CO₂ observed at the level of a single leaf resultfrom manifold responses at the level of the whole plant grownunder conditions of CO₂ enrichment. (Received July 8, 1999; Accepted August 12, 1999)     

Changes in photosynthetic rates and gene expression of leaves during a source-sink perturbation in sugarcane

BACKGROUND AND AIMS: In crops other than sugarcane there is good evidence that the size and activity of carbon sinks influence source activity via sugar-related regulation of the enzymes of photosynthesis, an effect that is partly mediated through coarse regulation of gene expression. METHODS: In the current study, leaf shading treatments were used to perturb the source-sink balance in 12-month-old Saccharum spp. hybrid 'N19' (N19) by restricting source activity to a single mature leaf. Changes in leaf photosynthetic gas exchange variables and leaf and culm sugar concentrations were subsequently measured over a 14 d period. In addition, the changes in leaf gene response to the source-sink perturbation were measured by reverse northern hybridization analysis of an array of 128 expressed sequence tags (ESTs) related to photosynthetic and carbohydrate metabolism. KEY RESULTS: Sucrose concentrations in immature culm tissue declined significantly over the duration of the shading treatment, while a 57 and 88% increase in the assimilation rate (A) and electron transport rate (ETR), respectively, was observed in the source leaf. Several genes (27) in the leaf displayed a >2-fold change in expression level, including the upregulation of several genes associated with C(4) photosynthesis, mitochondrial metabolism and sugar transport. Changes in gene expression levels of several genes, including Rubisco (EC 4.1.1.39) and hexokinase (HXK; EC 2.7.1.1), correlated with changes in photosynthesis and tissue sugar concentrations that occurred subsequent to the source-sink perturbation. CONCLUSIONS: These results are consistent with the notion that sink demand may limit source activity through a kinase-mediated sugar signalling mechanism that correlates to a decrease in source hexose concentrations, which, in turn, correlate with increased expression of genes involved in photosynthesis and metabolite transport. The signal feedback system reporting sink sufficiency and regulating source activity may be a potentially valuable target for future genetic manipulation to increase sugarcane sucrose yield.     

Regulation of leaf photosynthetic rate correlating with leaf carbohydrate status and activation state of Rubisco under a variety of photosynthetic source/sink balances

There is evidence suggesting that in plants changes in the photosynthetic source/sink balance are an important factor that regulates leaf photosynthetic rate through affects on the leaf carbohydrate status. However, to resolve the regulatory mechanism of leaf photosynthetic rate associated with photosynthetic source/sink balance, information, particularly on mutual relationships of experimental data that are linked with a variety of photosynthetic source/sink balances, seems to be still limited. Thus, a variety of manipulations altering the plant source/sink ratio were carried out with soybean plants, and the mutual relationships of various characteristics such as leaf photosynthetic rate, carbohydrate content and the source/sink ratio were analyzed in manipulated and non-manipulated control plants. The manipulations were removal of one-half or all pods, removal of one-third or two-third leaves, and shading of one-third or one-half leaves with soybean plants grown for 8 weeks under 10 h light (24 degrees C) and 14 h darkness (17 degrees C). It was shown that there were significant negative correlations between source/sink ratio (dry weight ratio of attached leaves to other all organs) and leaf photosynthetic rate; source/sink ratio and activation ratio (percentage of initial activity to total activity) of Rubisco in leaf extract; leaf carbohydrate (sucrose or starch) content and photosynthetic rate; carbohydrate (sucrose or starch) content and activation ratio of Rubisco; amount of protein-bound ribulose-1,5-bisphosphate (RuBP) in leaf extract and leaf photosynthetic rate; and the amount of protein-bound RuBP and activation ratio of Rubisco. In addition, there were significant positive correlations between source/sink ratio and leaf carbohydrate (sucrose or starch) content; source/sink ratio and the amount of protein-bound RuBP; carbohydrate (sucrose or starch) content and amount of protein-bound RuBP and the activation ratio of Rubisco and leaf photosynthetic rate. The plant water content, leaf chlorophyll and Rubisco contents were not affected significantly by the manipulations. There is a previous report in Arabidopsis thaliana that the amount of protein-bound RuBP in leaf extract correlates negatively with the activation ratio of Rubisco in the leaf extract. Therefore, the results obtained from the manipulation experiments indicate that there is a regulatory mechanism for the leaf photosynthetic rate that correlates negatively with leaf carbohydrate (sucrose and starch) status and positively with the activation state of Rubisco under a variety of photosynthetic source/sink balances.     

Effects of nitrogen supply on the acclimation of photosynthesis to elevated CO2

A common observation in plants grown in elevated CO₂ concentration is that the rate of photosynthesis is lower than expected from the dependence of photosynthesis upon CO₂ concentration in single leaves of plants grown at present CO₂ concentration. Furthermore, it has been suggested that this apparent down regulation of photosynthesis may be larger in leaves of plants at low nitrogen supply than at higher nitrogen supply. However, the available data are rather limited and contradictory. In this paper, particular attention is drawn to the way in which whole plant growth response to N supply constitutes a variable sink strength for carbohydrate usage and how this may affect photosynthesis. The need for further studies of the acclimation of photosynthesis at elevated CO₂ in leaves of plants whose N supply has resulted in well-defined growth rate and sink activity is emphasised, and brief consideration is made of how this might be achieved.Abbreviations A rate of CO₂ assimilation - C_i internal CO₂ concentration - PCR photosynthetic carbon reduction - Rubisco Ribulose 1,5-bisphosphate carboxylase/oxygenase - RuBP ribulose 1,5-bisphosphate     

Magnesium deficiency results in accumulation of carbohydrates and amino acids in source and sink leaves of spinach

Accumulation of assimilates in source leaves of magnesium‐deficient plants is a well‐known feature. We had wished to determine whether metabolite concentrations in sink leaves and roots are affected by magnesium nutrition. Eight‐week‐old spinach plants were supplied either with a complete nutrient solution (control plants) or with one lacking Mg (deficient plants) for 12 days. Shoot and root fresh weights and dry weights were lower in deficient than in control plants. Mg concentrations in deficient plants were 11% of controls in source leaves, 12% in sink leaves and 26% in roots, respectively. As compared with controls, increases were found in starch and amino acids in source leaves and in sucrose, hexoses, starch and amino acids in sink leaves, whereas they were only slightly enhanced in roots. In phloem sap of magnesium‐deficient and control plants no differences in sucrose and amino acid concentrations were found. To prove that sink leaves were the importing organs they were shaded, which did not alter the response to magnesium deficiency as compared with that without shading. Since in the shaded sink leaves the photosynthetic production of metabolites could be excluded, those carbohydrates and amino acids that accumulated in the sink leaves of the deficient plants must have been imported from the source leaves. It is concluded that in magnesium‐deficient spinach plants the growth of sink leaves and roots was not limited by carbohydrate or amino acid supply. It is proposed that the accumulation of assimilates in the source leaves of Mg‐deficient plants results from a lack of utilization of assimilates in the sink leaves.     

Effects of season-dependent irradiance levels and nitrogen-deficiency on photosynthesis and photoinhibition in field-grown rice (Oryza sativa)

Photoinhibition and acclimation of photosynthesis in rice plants grown under N-sufficient (NS) and N-deficient (ND) field conditions were investigated during the tropical wet (WS) and dry (DS) seasons in the Philippines. Diurnal patterns of CO₂ assimilation were examined. There was a transient peak in CO₂ assimilation in the leaves of the NS plants in the early morning during the DS and the WS, which was not seen in the ND plants in either season. ND leaves had lower Ribulose bisphosphate carboxylase/oxygenase (Rubisco) contents and lower chlorophyll contents. A lowered quantum yield of photosystem II (φPSII) was observed in the ND plants at an intermediate irradiance though no differences between N treatments were seen at high irradiance. Analysis of carotenoids indicated a small increase in the de-epoxidation state of the xanthophyll cycle (DES) at mid-day in the ND leaves compared to NS. Photoinhibition was greater in ND leaves when incident mid-day irradiance was increased by altering the leaf angle. Although Rubisco contents were lower in ND plants, photosynthesis in situ did not decline proportionally. For NS plants, Chlorophyll content, but not Rubisco content, was season-dependent and results are discussed in terms of the interaction between irradiance use and N content of rice leaves.     

Nitrogen reallocation and photosynthetic acclimation in response to partial shading in soybean plants

The first trifoliate of soybean was shaded when fully expanded, while the plant remained in high light; a situation representative for plants growing in a closed crop. Leaf mass and respiration rate per unit area declined sharply in the first few days upon shading and remained rather constant during the further 12 days of the shading treatment. Leaf nitrogen per unit area decreased gradually until the leaves were shed. Leaf senescence was enhanced by the shading treatment in contrast to control plants growing in low light. Shaded leaves on plants grown at low nutrient availability senesced earlier than shaded leaves on plants grown at high nutrient availability. The light saturated rate of photosynthesis decreased also gradually during the shading treatment, but somewhat faster than leaf N, whereas chlorophyll contents declined somewhat slower than leaf N.
Partitioning of N in the leaf over main photosynthetic functions was estimated from parameters derived from the response of photosynthesis to CO₂. It appeared that the N exported from the leaf was more at the expense of compounds that make up photosynthetic capacity than of those involved in photon absorption, resulting in a change in partitioning of N within the photosynthetic apparatus. Photosynthetic nitrogen use efficiency increased during the shading treatment, which was for the largest part due to the decrease in leaf N content, to some extent to the decrease in respiration rate and only for a small part to change in partitioning of N within the photosynthetic apparatus.     

Influence of assimilate demand on photosynthesis, diffusive resistances, translocation, and carbohydrate levels of soybean leaves

Rates of net photosynthesis and translocation, CO₂ diffusive resistances, levels of carbohydrates, total protein, chlorophyll, and inorganic phosphate, and ribulose 1,5-diphosphate carboxylase activity were measured in soybean (Glycine max L. Merrill) leaves to ascertain the effect of altered assimilate demand. To increase assimilate demand, the pods, stems, and all but one leaf (the “source leaf”) of potted plants were completely shaded for 6 or 8 days and the responses of the illuminated source leaf were monitored. Rate of net photosynthesis in the source leaf of the shaded plants was found to increase curvilinearly to a maximum on the 8th day. The source leaf of the control plants (no sink shading) maintained a constant photosynthetic rate during this period. Vapor-phase resistance to CO₂ diffusion did not vary with treatment, but mesophyll (liquid phase) resistance was significantly lower in the source leaf of the shaded plants.     

The Effect of Elevated Partial Pressures of CO2 on the Relationship between Photosynthetic Capacity and N Content in Rice Leaves

The effects of growth CO2 levels on the photosynthetic rates; the amounts of ribulose-1,5-bisphosphate carboxylase (Rubisco), chlorophyll (Chl), and cytochrome f; sucrose phosphate synthase activity; and total N content were examined in young, fully expanded leaves of rice (Oryza sativa L.). The plants were grown hydroponically under two CO2 partial pressures of 36 and 100 Pa at three N concentrations. The light-saturated photosynthesis at 36 Pa CO2 was lower in the plants grown in 100 Pa CO2 than those grown in 36 Pa CO2. Similarly, the amounts of Rubisco, Chl, and total N were decreased in the leaves of the plants grown in 100 Pa CO2. However, regression analysis showed no differences between the two CO2 treatments in the relationship between photosynthesis and total N or in the relationship between Rubisco and Chl and total N. Although a relative decrease in Rubisco to cytochrome f or sucrose phosphate synthase was found in the plants grown in 100 Pa CO2, this was the result of a decrease in total N content by CO2 enrichment. The activation state of Rubisco was also unaffected by growth CO2 levels. Thus, decreases in the photosynthetic capacity of the plants grown in 100 Pa CO2 could be simply accounted for by a decrease in the absolute amount of leaf N.     

Evidence for the sucrose-binding protein role in carbohydrate metabolism and transport at early developmental stage

Despite the large amount of data regarding sucrose-binding proteins (SBP), their functions remain largely unknown and controversial. In this investigation we performed a detailed temporal and spatial characterization of the phenotypes related to photosynthesis, sucrose exudation and carbohydrate metabolism in SBP antisense plants to gain insights into the physiological role of SBP. Significant reductions in net photosynthesis and in stomatal conductance were observed in the SBP antisense lines but were restricted to the vegetative phase, and persisted during a daily time course at this phase. Photosynthesis was saturated at a substantially lower irradiance in source leaves of the antisense lines, suggesting that light utilization is decreased in these plants. A slight reduction in soluble sugars was observed throughout the development of source leaves, partially overlapping a decrease in sucrose synthase activity (EC 2.4.1.13); whereas a transient increase in starch and adinosine diphosphate (ADP)-glucose pyrophosphorylase activity (EC 2.7.7.27) as well as decreased leaf sucrose exudation were detected in the beginning of the vegetative phase. These changes in source leaves were accompanied by reductions in sucrose and starch in sink leaves, hexoses and sucrose in roots and hexoses in shoot apex, which were observed before the occurrence of a significant reduction in height and in leaf number in the transgenic lines. These alterations in growth parameters did not persist throughout the development, but were associated with a delay in flowering time and leaf senescence in the SBP antisense lines. A likely involvement of SBP in sink strength is discussed.     

Combined Effects of Partial Defoliation and Nutrient Availability on Cloned Betula pendula Saplings: II. CHANGES IN NET PHOTOSYNTHESIS AND RELATED BIOCHEMICAL PROPERTIES

The combined effects of partial defoliation and nutrient availabilityon net photosynthesis and related biochemical variables werestudied in cloned Betula pendula Roth saplings. The saplingswere randomly assigned to different nutrient levels (5, 1·5and 0·5 mol N m^–3) in aerated nutrient cultureand to the following defoliation treatments: (1) control (nodamage), (2) damage of the developing main stem leaves (halfof the leaf lamina removed), and (3)removal of the developingmain stem leaves (entire leaf lamina removed). The leaf immediatelybelow the damaged area in the treated plants, and the correspondingleaf in the control plants, were selected for study. Net photosynthesismeasurements and biochemical determinations were made 2, 8 and14 d after assigning the treatments. At intermediate and lownutrient levels the final net photosynthetic capacity was significantlyhigher in the saplings with the topmost leaves removed thanin the undamaged control saplings, indicating that the expressionof compensatory photosynthesis after partial defoliation isnot inhibited by nutrient deficiency. The photosynthetic enhancementwas closely associated with the increased initial activity ofribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco). However,the increased activity of Rubisco was not exclusively the resultof a higher amount of Rubisco. The expression of compensatoryphotosynthesis after partial defoliation in our study cannotunequivocally be attributed to an increased flow of nitrogento the remaining leaves. Key words: Partial defoliation, nutrient availability, net photosynthesis, nitrogen, Rubisco     

Effect of panicle removal on photosynthetic acclimation under elevated CO<Subscript>2</Subscript> in rice

To examine the role of sink size on photosynthetic acclimation under elevated atmospheric CO₂ concentrations ([CO₂]), we tested the effects of panicle-removal (PR) treatment on photosynthesis in rice (Oryza sativa L.). Rice was grown at two [CO₂] levels (ambient and ambient + 200 μmol mol⁻¹) throughout the growing season, and at full-heading stage, at half the plants, a sink-limitation treatment was imposed by the removal of the panicles. The PR treatment alleviated the reduction of green leaf area, the contents of chlorophyll (Chl) and Rubisco after the full-heading stage, suggesting delay of senescence. Nonetheless, elevated [CO₂] decreased photosynthesis (measured at current [CO₂]) of plants exposed to the PR treatment. No significant [CO₂] × PR interaction on photosynthesis was observed. The decrease of photosynthesis by elevated [CO₂] of plants was associated with decreased leaf Rubisco content and N content. Leaf glucose content was increased by the PR treatment and also by elevated [CO₂]. In conclusion, a sink-limitation in rice improved N status in the leaves, but this did not prevent the photosynthetic down-regulation under elevated [CO₂].     

Ethylene insensitivity results in down-regulation of rubisco expression and photosynthetic capacity in tobacco

Little is known about the effect of hormones on the photosynthetic process. Therefore, we studied Rubisco content and expression along with gas exchange parameters in transgenic tobacco (Nicotiana tabacum) plants that are not able to sense ethylene. We also tested for a possible interaction between ethylene insensitivity, abscisic acid (ABA), and sugar feedback on photosynthesis. We measured Rubisco content in seedlings grown in agar with or without added sugar and fluridone, and Rubisco expression in hydroponically grown vegetative plants grown at low and high CO(2). Furthermore, we analyzed gas exchange and the photosynthetic machinery of transformants and wild-type plants grown under standard conditions. In the presence of exogenous glucose (Glc), agar-grown seedlings of the ethylene-insensitive genotype had lower amounts of Rubisco per unit leaf area than the wild type. No differences in Rubisco content were found between ethylene-insensitive and wild-type seedlings treated with fluridone, suggesting that inhibition of ABA production nullified the effect of Glc application. When larger, vegetative plants were grown at different atmospheric CO(2) concentrations, a negative correlation was found between Glc concentration in the leaves and Rubisco gene expression, with stronger repression by high Glc concentrations in ethylene-insensitive plants. Ethylene insensitivity resulted in plants with comparable fractions of nitrogen invested in light harvesting, but lower amounts in electron transport and Rubisco. Consequently, photosynthetic capacity of the insensitive genotype was clearly lower compared with the wild type. We conclude that the inability to perceive ethylene results in increased sensitivity to Glc, which may be mediated by a higher ABA concentration. This increased sensitivity to endogenous Glc has negative consequences for Rubisco content and photosynthetic capacity of these plants.     

Regulation of photosynthesis by sugars in sugarcane leaves

In sugarcane, increased sink demand has previously been shown to result in increased photosynthetic rates that are correlated with a reduction in leaf hexose concentrations. To establish whether sink limitation of photosynthesis is a result of sugar accumulation in the leaf, excision and cold-girdling techniques were used to modify leaf sugar concentrations in pot-grown sugarcane. In excised leaves that were preincubated in darkness for 3h, sucrose accumulation was reduced but accumulated again upon transfer to the light, while hexose concentrations remained lower than in controls (7.7 micromol mg(-1)FW versus 18.6 micromol mg(-1)FW hexose in controls). These results were associated with a 66% and 59% increase in photosynthetic assimilation (A) and electron transport rate (ETR), respectively, compared to controls maintained in the light. Similar increases in photosynthesis were observed when dark-treated leaves were supplied with 5mM sorbitol, but not when supplied with 5mM sucrose. Further analyses of (14)C-labeled sugars indicated rapid turnover between sucrose and hexose. Cold-girdling (5 degrees C) increased sucrose and hexose levels and resulted in a decline of photosynthetic rates over 5d (48% and 35% decline in assimilation rate and ETR, respectively). These sugar-induced changes in photosynthesis were independent of changes in stomatal conductance. This study demonstrates that the down-regulation of photosynthesis in response to culm sugar accumulation reported previously could be due to the knock-on effect of accumulation of sugar in leaf tissue, and supports the contention that hexose, rather than sucrose, is responsible for the modulation of photosynthetic activity.     

Grain yield and kernel weight of two maize genotypes differing in nitrogen use efficiency at various levels of nitrogen and carbohydrate availability during flowering and grain filling

Grain yield per plant (GYP) and mean kernel weight (KW) of maize (Zea mays L.) are sensitive to changes in the environment during the lag phase of kernel growth (the time after pollination in which the potential kernel size is determined), and during the phase of linear kernel growth. The aim of this study was to assess genotypic differences in the response to environmental stresses associated with N and/or carbohydrate shortage at different phases during plant development. The rate and timing of N and carbohydrate supply were modified by application of fertilizer, shading, and varying the plant density at sowing, at silking or at 14 d after silking. The effects of these treatments on the photosynthetic capacity, grain yield and mean kernel weight were investigated in two hybrids differing in N use efficiency. The total above-ground biomass and grain yield per plant of the efficient hybrid responded little to altered environmental conditions such as suboptimal N supply, enhanced inter-plant competition, and shading for 14 d during flowering, when compared to the less efficient genotype. We conclude that grain yields in the efficient genotype are less sensitive not only to N stress, but also to carbohydrate shortage before grain filling. Shading of N deficient plants from 14 d after silking to maturity did not significantly reduce grain yield in the non-efficient genotype, indicating complete sink limitation of grain yield during grain filling. In the efficient genotype, in contrast, grain yield of N-deficient plants was significantly reduced by shading during grain filling. The rate of photosynthesis declined with decreasing foliar N content. No genotypic differences in photosynthesis were observed at high or low foliar N contents. However, at high plant density and low N supply, the leaf chlorophyll content after flowering in the efficient genotype was higher than that in the non-efficient genotype. Obviously, the higher source capacity of the efficient genotype was not due to higher photosynthetic N use efficiency but due to maintenance of high chlorophyll contents under stressful conditions. In the efficient genotype, the harvest index was not significantly affected by N fertilization, plant density, or shading before the grain filling period. In contrast, in the non-efficient genotype the harvest index was diminished by N deficiency and shading during flowering. We conclude that the high yielding ability of the efficient genotype under stressful conditions was associated with formation of a high sink capacity of the grains under conditions of low carbohydrate and N availability during flowering and with maintenance of high source strength during grain filling under conditions of high plant density and low N availability.     

The effect of Albugo Candida (white blister rust) on the photosynthetic and carbohydrate metabolism of leaves of Arabidopsis thaliana

Albugo candida (pers.) O. Kuntze (white blister rust) is a biotrophic fungus which infects cruciferous plants including Arabidopsis thaliana (L) Heynh. We report the effect of this pathogen on the photosynthetic and carbohydrate metabolism of A. thaliana. As infection progressed A. Candida caused a reduction in the rate of photosynthesis when measured at either ambient or saturating concentrations of CO₂. These data suggested that both chlorophyll and Rubisco were lost from regions of infected leaves, and measurements of chlorophyll, Rubisco content and activity supported these observations. The reduction in the rate of photosynthesis was not caused by closure of stomata as transpiration was unaffected by the disease. Infected leaves accumulated both soluble carbohydrates and starch. The activities of sucrose-phosphate synthase, sucrose synthase and ADP glucose pyrophosphorylase did not change in response to infection. However, the activities of both the wall-bound and soluble acid invertases were higher in infected leaves than in controls; a new soluble invertase isoform with a pl of 5-1 appeared in infected leaves. The possible origin of the increase in wall-bound and soluble invertase activities and its effect on the carbohydrate and photosynthetic metabolism of the leaf are discussed.