Effect of Ophiobolus graminis infection on the assimilation and distribution of 14C in wheat

The uptake of ¹⁴C and movement of ¹⁴C-labelled assimilates in wheat plants inoculated with Ophiobolus graminis was examined following exposure of the second youngest leaf to ¹⁴CO₂. Autoradiographs of plants with infected seminal roots showed that assimilates were not translocated past the sites of root infection but accumulated in the undamaged portions of infected root systems, in particular the developing crown roots. There was no evidence that assimilates accumulated in the vicinity of O. graminis lesions. The net assimilation of ¹⁴CO₂ by wheat plants over a 5 h feeding period was not significantly affected by O. graminis infection. However, infection reduced the amount of ¹⁴C lost through respiration. Infection delayed the transfer of labelled assimilates from the fed leaf to the remainder of the plant but increased the proportion translocated to the roots. The latter effect was not apparent when infected plants were continuously irrigated during, and for 20 h following, the feeding period.     

Source,sink and hormonal control of translocation in wheat

Summary An analysis of the pattern of movement of ¹⁴C-labelled flag leaf assimilates in wheat (Triticum aestivum l. c.v. Gabo) during grain development, indicated that the greater the requirement for assimilates by the ear the more rapid was the speed of movement of these through the peduncle to the ear and also the lower their concentration. Experiments with [¹⁴C] indoleacetic acid ([¹⁴C]IAA) suggested that auxin production by the grains was not responsible for the control of assimilate translocation through the peduncle. Limiting the supply of available assimilates by shading the lower parts of the plant, did not significantly alter the speed of movement of ¹⁴C-photosynthate through the peduncle, while severing half of the vascular tissue in the peduncle altered the pattern of movement of ¹⁴C to the ear and enhanced the speed of movement of ¹⁴C through the remaining functional conducting tissue. These results are discussed in relation to the mechanism of translocation.     

Effects of Acetylcholine, Cytochalasin B and Amiprophosmethyl on Phloem Transport in Radish （Raphanus sativas）

We investigated the role of the ＂sieve tube-companion cell complex＂ lining the tube periphery, particularly the microfilament and microtubule, in assisting the pushing of phloem sap flow. We made a simple phloem transport system with a living radish plant, in which the conducting channel was exposed for local treatment with chemicals that are effective in modulating protoplasmic movement （acetylcholine, （ACh） a neurotransmitter in animals and insects; cytochalasin B, （CB） a specific inhibitor of many cellular responses that are mediated by microfilament systems and amiprophos-methyl, （APM） a specific inhibitor of many cellular responses that are mediated by microtubule systems）. Their effects on phloem transport were estimated by two experimental devices： （i） a comparison of changes in the amount of assimilates in terms of carbohydrates and ^14C-labeled photosynthetic production that is left in the leaf blade of treated plants; and （ii） distribution patterns of ^14C-labeled leaf assimilates in the phloem transport system. The results indicate that CB and APM markedly inhibited the transfer of photosynthetic product from leaf to root via the leaf vein, while ACh enhanced the transfer of photosynthetic product in low concentrations （5.0×10^-4 mol/L） but inhibited it in higher concentrations （2.0×10^-3 mol/L） from leaf to root via the leaf vein. Autoradiograph imaging clearly reveals that ACh treatment is more effective than the control, and both CB and APM treatments effectively inhibit the passage of radioactive assimilates. All of the results support the postulation that the peripheral protoplasm in the sieve tube serves not only as a passive semi-permeable membrane, but is also directly involved in phloem transport.     

Utilization of Photosynthetically Fixed 14CO2 into Alkaloids in Relation to Primary Metabolites in Developing Leaves of Catharanthus roseus

Partitioning of current photosynthates towards primary metabolites and its simultaneous incorporation in leaf alkaloids was investigated in developing leaves of medicinally important Catharanthus roseus. Of the total ¹⁴CO₂ assimilated, the leaves at positions 1–6 fixed 8, 22, 25, 19, 13, and 8 %, respectively, and stem 3 %. Leaf fresh mass, chlorophyll content, and CO₂ exchange rate increased up to the third leaf. The total alkaloid content was highest in young actively growing leaves, which declined with age. Total ¹⁴C fixed and its content in ethanol soluble fraction increased up to the third leaf and then declined. The ¹⁴C content in primary metabolites such as sugars and organic acids was also highest in the 3^rd leaf. The utilization of ¹⁴C assimilates into alkaloids was maximum in youngest leaf which declined with leaf age. Hence the capacity to synthesize alkaloids was highest in young growing leaves and metabolites from photosynthetic pathway were most efficiently utilized and incorporated into alkaloid biosynthetic pathway by young growing leaves.     

Diurnal export and carbon economy in an expanding source leaf of cucumber at contrasting source and sink temperature

Effects of contrasting temperatures of an expanding leaf (source) and of remaining plant parts (sink) on diurnal export and distribution of carbon were studied in seedlings of Cucumis sativus L., cv. Farbio. The time course of the rate of export was calculated by measuring simultaneously the exchange of ¹⁴CO₂ and the amount of ¹⁴C in the source leaf by means of a Geiger-Müller detector using a steady-state labelling technique. In all treatments average export rate during the night (16 h) was maximally 50% of the average rate during the 8-h day. Temperature affected the diurnal course of export via the source leaf and the sink in different ways. At a source leaf temperature of 25 or 30°C export stopped 12 h after start of the night, whereas at 20°C export continued throughout the night. However, the total amount of carbon exported during a 24 h cycle, expressed as a proportion of the amount of carbon assimilated, was the same at source leaf temperatures of 20 or 30°C. Thus source leaf temperature did not affect the distribution of assimilates between source and sink, in contrast to sink temperature. After 24 h at a sink temperature of 30°C, 20% more ¹⁴C was exported to plant parts below the source leaf than with a sink temperature of 20°C, at the expense of carbon remaining in the source. During the day less starch and more structural dry matter was formed at a source leaf temperature of 30°C than at 20°C. After a complete day/night cycle, however, there was no difference between the treatments. Starch was the primary carbon source during the night, and the decline in the rate of export coincided with the depletion of starch. Thus the decline in the rate of export at a source leaf temperature of 25 or 30°C at 12 h after the start of the night was due to the depletion of starch at that time. Similarly, at 20°C export could continue until the end of the night as the starch degradation supplied assimilates during the whole night.     

Effects of Temperature on the Distribution of 14C-labelled Assimilates in the Flowering Shoot of Carnation

The pattern of distribution of ¹⁴C-labelled assimilates translocatedfrom a leaf on the flower stem of carnation was found to varywith both the ambient air temperature and the localized temperatureof the flower bud. A high bud temperature increased the proportionofassimilates moving into the floral tissues while a low budtemperature increased the proportion accumulating in the stemabove the source leaf. When the air temperature was raised independentlyof the bud temperature, the stem gained assimilates at the expenseof the flower, but if both air temperature and bud temperaturewere raised together, effects of bud temperature predominatedand movement of assimilates into the flower was promoted. Therole of the flower in mediating effects of temperature is discussedwith reference to the distribution of invertase activity inthe shoot.     

Assimilate Partitioning in the Variegated Coleus Leaf

Mature leaves of a variegated cultivar of Coleus blumei Benth. with a green border and central albino region constitute a source-sink system suitable for studies on assimilate partitioning. Leaves treated with ¹⁴CO₂ on a small part of the intact green border export assimilate via the shortest path into the stem. Leaves with all but a small lobe of the green border removed show different partitioning of labeled assimilates when the leaf is exposed to ¹⁴CO₂ (Fisher and Eschrich, 1985): The whole albino region of the leaf is supplied but no tracer is exported into the stem. When the green border is completely removed, ¹⁴CO₂-treatment of the albino region leads to the fixation of CO₂, obviously by PEP carboxylase, as indicated by the occurrence of labeled malate. Results show that the albino region of the variegated leaf constitutes a potential sink when deprived of its green border. In addition, CO₂-fixation by PEP carboxylase in albino tissue seems to indicate a common capacity of leaves which is normally masked by photosynthesis. The difference of assimilate partitioning between leaves with intact and leaves with partly removed green borders demonstrates that the unlabeled assimilates control the movement of labeled assimilates.     

Phloem transport in Picea abies (L.) Karst. in mid-winter

Summary Translocation of ¹⁴C assimilates was studied on four different transport systems of Picea abies branches after induced activation in January. ¹⁴CO₂ assimilation of terminal shoots for 48 h at 25° C resulted in phloem loading and basipetal transport of ¹⁴C photosynthate into the following, older shoot generations. ¹⁴C import was enhanced, when these older shoot generations were kept in the dark. Microautoradiographs of the labelled terminal shoots showed that ¹⁴C assimilates were exported from needles via sieve elements of the leaf traces and loaded into the latest increment of the axial secondary phloem. No ¹⁴C label appeared in the obliterated sieve cells or in the tracheids. In addition, ¹⁴C photosynthate accumulated densely in the chlorophyllous cells of the cortex and in cells of the resin ducts, indicating certain sink activity. In the darkened 2-year-old shoot, imported ¹⁴C photosynthate was concentrated in the functional secondary phloem, while some ¹⁴C label was unloaded into the latest xylem increment. When 6-year-old shoots were exposed to ¹⁴CO₂ for 48 h in the light, ¹⁴C assimilates accumulated in the phloem of the leaf trace and in the latest increment of the axial secondary phloem. However, a substantial amount of radioactivity was unloaded into ray cells and phloem parenchyma cells. Thus, the presence of functioning phloem in needles and twigs of P. abies during winter allows long-distance translocation and radial distribution of assimilates according to existing source-sink relations.     

Assimilate transport in maize (Zea mays L.) seedlings at vertical low temperature gradients in the root zone

Even moderate chilling temperatures may cause important modifications in assimilate movement in maize seedlings from the shoot to the roots, but there is no information on long-distance transport of assimilates in plants subjected to vertical gradients of moderately low temperatures in the root zone. Seedlings of a chilling-tolerant (KW1074) and a chilling-sensitive inbred line (CM109) of maize were grown in a system that allowed the maintenance of temperature gradients between the topsoil (0-10 cm) and the subsoil (10-50 cm). After pregrowth at 24C until the third-leaf stage, plants were subjected to chilling-stress regimes for 6 d (17/17/17C, 17/17/12°C, 12/12/12°C, 12/12/17°C, air/topsoil/subsoil). The time taken for the assimilates to enter the phloem from the second leaf increased at low temperatures for both lines, but to a much greater extent in CM109. Although mainly influenced by air and topsoil temperature, low temperature in the subsoil also affected this trait in CM109. The speed of assimilate transport between the second leaf and the mesocotyl in KW1074 was strongly reduced by cool temperatures in the shoot and topsoil as well as by 12°C in the subsoil in CM109, because the latter line had a larger portion of its root system in the subsoil as compared to KW1074. The portion of assimilates allocated to the root decreased at low temperatures in both lines, but to a greater extent in CM109, and was controlled mostly by the subsoil temperature. After rewarming, values of all measured parameters of assimilate transport returned to near pregrowth levels within a few days.Keywords: Assimilate transport, low temperature stress, root growth, vertical soil temperature gradients, Zea mays L.      

Translocation and incorporation of 14C into the petiole from different regions within developing cottonwood leaves

Summary The ability of a developing cottonwood (Populus deltoides Bartr.) leaf to export ¹⁴C-labeled assimilates begins at the lamina tip and progresses basipetally with increasing LPI. This progression indicates that portions of leaves function quasi-independently in their ability to export ¹⁴C-photosynthate. Although most of the exported radioactivity was recovered in the petiole as water-80% alcohol-soluble compounds, there was also substantial incorporation into the chloroform and insoluble fractions. This observation indicates that assimilates translocated from the lamina are used in structural development of the petiole. Freeze substitution and epoxy embedding were used to prepare microautoradiographs for localization of water-soluble compounds. Radioactivity was found in all cell types within specific subsidiary bundles of the petiole. However, radioactive assimilates appeared to move from the translocation pathway in the phloem toward active sinks in the walls of the expanding metaxylem cells. Translocation in the mature xylem vessels was not observed.     

The influence of externally supplied sucrose on phloem transport in the maize leaf strip

Sucrose (2,5–1000 mmol l^–1), labeled with [¹⁴C]sucrose, was taken up by the xylem when supplied to one end of a 30-cm-long leaf strip of Zea mays L. cv. Prior. The sugar was loaded into the phloem and transported to the opposite end, which was immersed in diluted Hoagland's nutrient solution. When the Hoagland's solution at the opposite end was replaced by unlabeled sucrose solution of the same molarity as the labeled one, the two solutions met near the middle of the leaf strip, as indicated by radioautographs. In the dark, translocation of ¹⁴C-labeled assimilates was always directed away from the site of sucrose application, its distance depending on sugar concentration and translocation time. When sucrose was applied to both ends of the leaf strip, translocation of ¹⁴C-labeled assimilates was directed toward the lower sugar concentration. In the light, transport of ¹⁴-C-labeled assimilates can be directed (1) toward the morphological base of the leaf strip only (light effect), (2) toward the base and away from the site of sucrose application (light and sucrose effect), or (3) away from the site of sucrose application independent of the (basipetal or acropetal) direction (sucrose effect). The strength of a sink, represented by the darkened half of a leaf strip, can be reduced by applying sucrose (at least 25 mmol l^–1) to the darkened end of the leaf strip. However, equimolar sucrose solutions applied to both ends do not affect the strength of the dark sink. Only above 75 mmol l^–1 sucrose was the sink effect of the darnened part of the leaf strip reduced. Presumably, increasing the sucrose concentration replenishes the leaf tissue more rapidly, and photosynthates from the illuminated part of the leaf strip are imported to a lesser extent by the dark sink.Supported by Deutsche Forschungsgemeinschaft     

Modification of Pattern of Photosynthate Movement within and between Shoots of Vitis vinifera L

During the prebloom and bloom stages, no movement of labeled photosynthates occurred from a shoot of Vitis vinifera L. fed with ¹⁴CO₂, to an adjacent shoot on the same spur. Movement of labeled assimilates into the unfed shoot was induced when this shoot was sprayed with 2.89 × 10⁻³m gibberellic acid during the prebloom stage. During the bloom stage darkening or defoliation of the unfed shoot resulted in the import of labeled photosynthates from the adjacent fed shoot. Similarly, movement of ¹⁴C into an untreated shoot was induced by removing the terminal 7.5 centimeters and deblossoming the fed shoot. During the berry set stage, translocation of labeled photosynthates from a newly exporting leaf was upwards to the shoot tip, but the direction of movement was reversed by removal of the shoot tip or by darkening or removal of the leaves below the fed leaf. Translocation of photosynthates was predominantly basipetal from a fed leaf near the base of the shoot during the berry set stage, but upward movement was induced by darkening or defoliation of the upper part of the shoot.     

Influence of gibberellic acid on <Superscript>14</Superscript>CO<Subscript>2</Subscript> metabolism,growth, and production of alkaloids in <Emphasis Type="Italic">Catharanthus roseus</Emphasis>

Changes in growth parameters, carbon assimilation efficiency, and utilization of ¹⁴CO₂ assimilate into alkaloids in plant parts were investigated at whole plant level by treatment of Catharanthus roseus with gibberellic acid (GA). Application of GA (1 000 g m⁻³) resulted in changes in leaf morphology, increase in stem elongation, leaf and internode length, plant height, and decrease in biomass content. Phenotypic changes were accompanied by decrease in contents of chlorophylls and in photosynthetic capacity. GA application resulted in higher % of total alkaloids accumulated in leaf, stem, and root. GA treatment produced negative phenotypic response in total biomass production but positive response in content of total alkaloids in leaf, stem, and roots. ¹⁴C assimilate partitioning revealed that ¹⁴C distribution in leaf, stem, and root of treated plants was higher than in untreated and variations were observed in contents of metabolites as sugars, amino acids, and organic acids. Capacity to utilize current fixed ¹⁴C derived assimilates for alkaloid production was high in leaves but low in roots of treated plants despite higher content of ¹⁴C metabolites such as sugars, amino acids, and organic acids. In spite of higher availability of metabolites, their utilization into alkaloid production is low in GA-treated roots.     

Effect of altering the root-zone temperature on growth, translocation, carbon exchange rate, and leaf starch accumulation in the tomato

Tomato seedlings (Lycopersicon esculentum Mill. cv Vendor) were grown hydroponically with their root systems maintained at a constant temperature for a 2-week period commencing with the appearance of the first true leaf. Based on fresh and dry weight and leaf area, the optimal root-zone temperature for seedling growth was 30°C. The carbon exchange rate of the leaves was also found to increase with rising root-zone temperature up to 30°C. However, a more complex relationship seems to exist between root-zone temperature and the accumulation of ¹⁴C-labeled assimilates in the roots; inasmuch as there is no enhancement in this accumulation at the most growth promoting root-zone temperatures (22-30°C).     

The Effect of Nitrogen Supply to Urtica dioica L. Plants on the Distribution of Assimilate Between Shoot and Roots

In this study the influence of nitrogen nutrition on the patterns of carbon distribution was investigated with Urtica dioica. The nettles were grown in sand culture at 3 levels of NO^?₃, namely 3 (low), 15 (medium) and 22 (high) mM. These levels encompassed a range within which nitrogen did not affect total biomass production. The ratio of root: shoot biomass of the low nitrogen plants was, however, significantly higher than that of the nettles grown at medium and high N supply. Carbon allocation from one leaf of each pair of leaves was examined after a ¹⁴CO₂-pulse and a subsequent ¹⁴C distribution period of one night. Only the youngest two leaf pairs did not export assimilates. Carbon (¹⁴C) export to the shoot apex and to the roots, as measured at the individual nodes responded to the nitrogen status: At medium and high nitrogen supply the 3rd, 4th and 5th leaf pairs exported to the shoot apex, while lower leaves exported to the root. At low nitrogen supply only the 3rd leaf exported towards the shoot apex. The results illustrate the plastic response of carbon distribution patterns to the nitrogen supply, even when net photosynthesis, carbon export from the source leaves and biomass production were not affected by the nitrogen supply to the plant.     

Hormonal Regulation of Assimilate Utilization in Barley Leaves in Relation to the Development of Their Source Function

Growth, photosynthesis, utilization of assimilates, and the development of a source function in leaves were studied in relation to changes in concentrations and ratios of phytohormones. Carbon isotope ¹⁴C was used to trace utilization and outflow of photosynthetic products from the leaf. Concentrations of endogenous phytohormones were determined by solid-phase immunoenzyme assay. It was shown that, in juvenile leaves (one-fifth of their final area), which did not attain a high rate of photosynthesis, up to 80% of assimilates were incorporated into structural polysaccharides (cellulose and hemicellulose) one day after feeding with ¹⁴CO₂. During leaf growth and the development of its source function, the synthesis of structural polysaccharides declined to 10%, but the formation of alcohol- and water-soluble compounds (AWSC) grew to 80%. Monosaccharides and oligosaccharides, which could act as transport forms of carbohydrates, constituted 30% and 40% of AWSC, respectively. The percentage of assimilates utilized for protein synthesis decreased with leaf growth. The revealed changes correlate with the concentration and the ratio of free forms of phytohormones at various stages of leaf development. Development of a source function, a decline in cellulose and hemicellulose syntheses, and an increase in AWSC were related to the decrease in ABA and IAA concentrations and the increase in the ABA/IAA ratio. The ABA level can regulate the pathways of photoassimilate utilization in leaves by partitioning carbon flows either to the synthesis of high-molecular-weight compounds (cellulose, hemicellulose, and proteins), used for cell growth in leaves, or to the synthesis of transport forms of carbohydrates. A high ABA level favors the first pathway while low level switches leaf metabolism to the second one.     

Effects of sodium nitroprusside,the nitric oxide donor,on photosynthesis and ultrastructure of common flax leaf blades

Solutions of sodium nitroprusside, a nitric oxide donor, were introduced at various concentrations into common flax (Linum usitatissimum L.) shoots with the transpirational water flow. Sodium nitroprusside and nitrate were found to exert similar effects on incorporation of ¹⁴C into photosynthetic products, leaf cell ultrastructure, and the export of assimilates from leaves. The results suggest that export of assimilates from leaves might be regulated by the products of incomplete nitrate reduction and that regulation may involve the NO-signaling system.     

Floral stimulus movement in perilla and flower inhibition caused by noninduced leaves

Photoperiodic control of flowering in the short day plant Perilla involves the transmission of a floral stimulus from induced leaves to the shoot apex. We have studied the basipetal movement of this stimulus and of ¹⁴C-labeled assimilates in plants with an induced leaf (donor) grafted into the uppermost internode of a vegetative plant in which the axillary shoots at various nodes along the stem function as receptors.     

Effect of assimilate utilization on photosynthetic rate in wheat

Summary Two weeks after anthesis, when the grain is filling rapidly, the rate of photosynthesis by flag leaves of wheat cv. Gabo was between 20 and 30 mg CO₂ dm^-2 leaf surface hour^-1 under the conditions used. About 45% of flag-leaf assimilates were translocated to the ear, and only about 12% to the roots and young shoots.On removing the ear, net photosynthesis by the flag leaves was reduced by about 50% within 3–15 hours, and there was a marked reduction in the outflow of ¹⁴C-labelled assimilates from the flag leaves.Subsequent darkening of all other leaves on plants without ears led to recovery of flag-leaf photosynthesis, as measured by gas analysis and ¹⁴CO₂ fixation, and to increased translocation of assimilates to the roots and young shoots. Minor changes in the rates of dark respiration accompanied these major, reversible changes in photosynthetic rate.After more than a week in continuous, high-intensity light, the rate of photosynthesis by flag leaves of intact plants had fallen considerably, but could be restored again by a period in darkness, or by inhibiting photosynthesis in the ears by spraying them with DCMU. The inhibition of ear photosynthesis increased translocation of labelled assimilates from the flag leaf to the ears, without affecting leaf sugar levels.The application of TIBA to the culm below the ear inhibited auxin movement throught the culm, but had no influence on flag-leaf photosynthesis.These results suggest that, at least in this system, photosynthesis by the flag leaf is regulated directly by the demand for assimilates from the flag leaf and not indirectly through action in the leaf of auxins produced by the sink organs.     

Distribution of 14CO2 during ontogeny of chickpea (Cicer arietinum) grown at two moisture levels

The distribution of assimilates of ¹⁴CO₂ in ethanol soluble and insoluble fractions was measured at 20-day intervals from 45–135 days after sowing (DAS) in chickpea (Cicer arietinum) grown at two moisture levels. The contribution of pre-flowering assimilates to pods, although very low, was higher under the stress conditions. At the time of harvest, the recovery of ¹⁴C in pods was 0.4 and 0.9% of the total ¹⁴C fed 45 DAS in soluble and 2.5 and 5.1% in insoluble fractions in control and stressed plants, respectively. The %¹⁴C received by nodules continuously decreased with increasing age of plants. Stressed plants diverted more ¹⁴C to nodules, compared to control, during vegetative and flowering stages. During active seed filling, stressed plants diverted more ¹⁴C to reproductive parts and less to nodules, compared to control. Significant amounts of ¹⁴C were retgined by the stem and leaves during the seed-filling period and it appears that there is scope for the remobilisation of pre-flowering, as well as post-flowering assimilates for seed-filling of chickpea.