Carbohydrate Metabolism and Translocation in Healthy and Cocoa Swollen Shoot Virus-Infected Cocoa Plants

Analyses of cocoa swollen shoot virus-infected and healthy cocoa (Theobroma cacao L.) plant tissues were made to determine the effect of virus infection on the metabolism and transport of carbohydrates in affected plants. Starch, sucrose and reducing sugars were found to accumulate in infected tissues. Translocation of photosynthates (mainly as sucrose) to the stem and root system, as estimated by the overnight loss of carbohydrates from the leaves and by ¹⁴CO₂ tracer experiments, was as efficient in the infected plants as in the healthy. Infected plants showed a higher diurnal turnover of carbohydrates in their leaves and, on unit leaf area basis, higher levels of ¹⁴C-labelled assimilates suggesting that they have a greater photosynthetic capacity than the healthy plants. The rate of respiration, as determined by the proportions of organic acids, amino acids and other intermediary metabolites formed from translocated ¹⁴C-labelled sugars, was generally higher in infected than in healthy plants. It is concluded from available data showing the presence in infected tissues of mineral nutrients, protein N and amino acids at the same concentrations as in healthy plants, and from the relatively high rates of photosynthesis and respiration that a high rate of metabolic activity is maintained in the host-virus system. Some factors possibly contributing to the stunted growth of infected plants are discussed in the light of these findings.     

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.     

Distribution of Soluble 14C-Labelled Assimilates as Related to Distribution of 14C-Benzyladenine in Shoot Tips of Hedera helix

The relationship between benzyladenine and assimilate distributions in shoot tips of English Ivy (Hedera helix L.) was studied. Exogenous applications of benzyladenine resulted in small amounts of the hormone moving to the apical meristem. The large part of the recovered benzyladenine was found in the young leaves of the shoot tip. Benzyladenine increased the amount of ¹⁴C-labelled assimilates in the shoot tip, but the distribution pattern of those assimilates within the tip was like that of the untreated control.     

The Effect of IAA on Sugar Accumulation and Basipetal Transport of 14C-labelled Assimilates in Relation to Root Formation in Phaseolus vulgaris Cuttings

The effect of indol-3yl-acetic acid on root formation, accumulation of 80% ethanol-soluble sugars and basipetal transport of ¹⁴C-labelled assimilates has been investigated in Phaseolus vulgaris (cv. Canadian Wonder) hypocotyl cuttings. The removal of leaves reduced root formation in the hypocotyl, while excision of the apical bud was less detrimental. The expression of the IAA effect in inducing more roots was dependent on the area of leaves, and was found to be better when all leaves were present. Sugars accumulated slowly at the base of cuttings during a four-day period after excision, and IAA greatly enhanced this accumulation. By comparing sugar content at the base of green and starved cuttings it was established that IAA greatly increased it concurrently with root formation. IAA applied in solution to the hypocotyl greatly enhanced the basipetal transport of ¹⁴C-labelled assimilates and their accumulation at the hypocotyl during a 24-h period. The IAA-induced accumulation was found to be connected with a greater mobilization of labelled assimilates from upper parts of the cutting. Experiments involving pretreatment with IAA and transport in cuttings already possessing root primordia, suggest a dual effect of IAA: (I) a direct effect on transport, and (2) an increase in the root-“sink”. It is concluded that both may be operating in inducing basipetal accumulation of labelled assimilates. It is suggested that one of the roles of IAA in promoting rooting of cuttings is to increase sugar availability at the site of root formation.     

Transport and Distribution of 14CO2 Assimilate in Lolium perenne in Response to Varying Nitrogen Supply to Halves of a Divided Root System

The movement of ¹⁴C assimilate from shoots to roots and its subsequent metabolism in the root of Lolium perenne L. was studied using variable N nutrition supplied to halves of a divided root system. Half of the N-deficient root system was supplied with either high NO₃-N or high NH₄-N for 16 hours or 6 days before ¹⁴CO₂ labeling of the shoots. The distribution of ¹⁴C in sugars, ethanol-soluble nitrogen and organic acids in roots appeared to be related to the N content of the tissue. Supply of high NO₃-N for 6 days resulted in significant internal translocation of N into the low N supplied root half. Both root halves also had similar patterns of ¹⁴C distribution among soluble and insoluble metabolites. However, NH₄-N supply for 6 days did not result in a significant increase of N in the low N supplied roots, thus only the high NH₄-N supplied roots displayed stimulated sugar metabolism, similar to that in both root halves in the high NO₃-N supply treatment. Percent transport of ₁₄C assimilates from shoot to root was influenced by form and level of N supplied to root halves. Root halves supplied with either high N source for 6 days accumulated greater amounts of ¹⁴C assimilate than the corresponding low N root half. However NH₄-N supply appeared to make roots stronger sinks since NH₄ supply resulted in significantly greater ¹⁴C accumulation in both the high NH₄ supplied and the low N root halves than did NO₃-N supply in corresponding root halves. The data suggest that factors other than root metabolism, such as N mediated metabolism in the shoot, may also influence the percent transport of assimilates to the root. Internal distribution of the incoming assimilate within the root system could be regulated by the metabolic activity or assimilate demand of the roots.     

The patterns of growth, assimilation of 14CO2 and distribution of 14C-assimilate within vegetative plants of Loliumperenne at low and high density

The patterns of growth, assimilation of ¹⁴CO₂ and distribution of ¹⁴C-labelled assimilate were followed for 12 wk from sowing in individual plants of Lolium perenne grown in miniswards at either low (500 plants m^-2) or high (5000 plants m^-2) density. At the latter density, plants were characterised by a 50% reduction in RGR, by the production of fewer tillers, especially second- and third-order tillers, and by a reduction in mean tiller weight. All the green and senescing leaves of each tiller assimilated ¹⁴CO₂ and the overall assimilatory capacity of a tiller was directly related to its dry weight. At both densities the plant consisted of a main shoot and established tillers with comparable assimilatory activities and a range of developing tillers that assimilated relatively small amounts of ¹⁴CO₂. As each successive primary tiller developed it was supplied with assimilate from the main shoot and the degree of support was inversely proportional to the dry weight of the tiller. At both densities it was concluded that the first primary tiller could be regarded as an independent assimilatory unit when it reached a dry weight of about 25 mg even though some import of main shoot assimilate continued until the tiller was double this weight. The supply of assimilate to the root system was greatly reduced at both densities compared with previous observations on plants grown singly.     

Seasonal differences in the pattern of assimilate movement in branches of Coffea arabica L

At Ruiru, Kenya, ¹⁴CO₂ was fed to single leaves at different distances from the apex on branches of mature fruiting and non-fruiting trees of Coffea arabica on six occasions from the Short Rains (November) 1966 to the Long Rains (April-May) 1967. The location of ¹⁴C-labelled assimilates in the treated branches was determined 26 h later by autoradiography. The direction of movement of labelled assimilates indicated large seasonal differences in the relative sink strengths of the shoot tip, fruit and trunk-root systems. On vegetative trees the sink strength of trunk-roots was much smaller, as compared with the shoots, at the beginning of the Long Rains than at the end of the previous Short Rains or in the intervening dry season. Assimilate use by growing fruits did not alter the pattern of distribution of assimilates to the other sinks at the end of the Short Rains, but it did restrict assimilate movement to both shoot tips and trunk-roots at the_ beginning of the Long Rains. In the dry season, virtually all assimilates were utilized by growing fruits when these were present. Vegetative secondary shoots provided assimilates to growing fruits and trunk-roots at the end of the Short Rains and in the dry season. Some practical implications are noted.     

Assimilate Partitioning in Red and White Clover Either Dependent on N2 Fixation in Root Nodules or Utilizing Nitrate Nitrogen

During vegetative growth in controlled environments, the patternof distribution of ¹⁴C-labelled assimilates to shoot and root,and to the meristems of the shoot, was measured in red and whiteclover plants either wholly dependent on N₂ fixation in rootnodules or receiving abundant nitrate nitrogen but lacking nodules. In experiments where single leaves on the primary shoot wereexposed to ¹⁴CO₂, nodulated plants of both clovers generallyexported more of their labelled assimilates to root (+nodules),than equivalent plants utilizing nitrate nitrogen, and thiswas offset by reduced export to branches (red clover) or stolons(white clover). The intensity of these effects varied with experiment.The export of labelled assimilate to growing leaves at the terminalmeristem of the donor shoot was not influenced by source ofnitrogen. Internode elongation in the donor shoot utilized nolabelled assimilate. Whole plants of white clover exposed to ¹⁴CO₂ on seven occasionsover 32 days exhibited the same effect on export to root (+nodules),which increased slightly in intensity with increasing plantage. Nodulated plants had larger root: shoot ratios than theirequivalents utilizing nitrate nitrogen. Trifolium repens, Trifolium pratense, red clover, white clover, nitrogen fixation, nitrate utilization, assimilate partitioning     

The Influence of Plant Water Deficit on Distribution of 14C-labelled Assimilates in Cacao Seedlings

The relationship between plant water status and distributionof ¹⁴C-labelled assimilates in cacao (Theobroma cacao L.) wasevaluated after ¹⁴CO₂ pulse labelling leaves of seedlings subjectedto varying levels of water deficiency. The proportion of ¹⁴Cexported by source leaves was strongly affected by seedlingwater status. An increasing proportion of labelled assimilatesremained in source leaves at both 24-h and 72-h harvests aswater stress intensity increased. Water stress reduced the distributionof exported label to leaves and to the expanding flush in particularbut increased the proportion of label in stems and roots. Theresults suggest that current photoassimilates may be temporarilystored in source leaves and stems of cacao seedlings duringperiods of plant water deficit. The stress-induced changes inpartitioning of labelled carbon were in concordance with changesin shoot to root biomass ratios, which was likely due to greaterreduction in growth of above-ground organs to that of roots. Theobroma cacao L, assimilate partitioning, cacao, ¹⁴C-photoassimilate, water stress, water potential     

Retention and allocation of <Superscript>14</Superscript>C assimilates by maintenance leaves and harvest index of tea (<Emphasis Type="Italic">Camellia sinensis</Emphasis> L.)

Partitioning of ¹⁴C-labelled photosynthates to various parts of un-pruned tea clones TV1 and TV25 was assessed in vivo by exposing maintenance leaves to ¹⁴CO₂ at monthly intervals throughout the year. The plants from shoot apex to root tip were divided into twelve components to assess the allocation and retention of ¹⁴C-photosynthates by the maintenance foliage. Out of the total photosynthates produced by the maintenance leaves, only 11.08 % was allocated to the commercially useful harvestable two and a bud shoots which is accepted as the harvest index of tea. The photosynthetically active maintenance leaves retained 19.05 % while 24.56 % was distributed to the branches. The bottom and the top parts of the trunk utilized 7.44 and 7.21 %, respectively. The thick roots at the base of the trunk, medium sized roots, pencil size roots, and feeder roots imported 7.28, 7.72, 7.65, and 8.01 % of ¹⁴C assimilates, respectively. Except retention by leaves, all the plant parts of vigorous clone TV25 required higher percentage of assimilates than TV1. The mean quantities of net photosynthates utilized by the stem and the roots were 69.37 and 30.63 %, respectively.     

Assimilatverteilung in Sommergerstensorten mit unterschiedlicher Resistenz gegenüber dem Echten Mehltau (Erysiphe graminis f. sp. hordei)

Distribution of assimilates in cultivars of spring barley with different resistance against powdery mildew (Erysiphe graminis f. sp. hordei) Transport and distribution of radioactive labelled assimilates in spring barley cultivars with different degrees of resistance to powdery mildew were studied after ¹⁴CO₂-treatment of single leaves. Plants of the cultivars ‘Amsel’ (susceptible), ‘Asse’ (adult plant resistant), and ‘Rupee’ (resistant) were analyzed at the vegetative growth stage (5. leaf unfolded) and the generative growth stage (anthesis). At the vegetative growth stage the assimilate export from the mildew inoculated 5. leaf of ‘Amsel’ and ‘Rupee’ is decreased; in ‘Asse’, there is no considerable change of assimilate distribution due to infection. At the generative growth stage the assimilate export from the infected flag leaf of ‘Amsel’ is reduced when the fungus, is sporulating. In the cultivar ‘Asse’ the assimilates are bound at the infection site until the seventh day after inoculation, then the transport of assimilates to the ear is increased. In ‘Rupee’ mildew inoculation causes an enhanced assimilate transport to the ear. The changes in assimilate distribution due to mildew inoculation are discussed with respect to the different types of host-parasite-interactions and the source-sink-activities in the different cultivars.     

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.     

The influence of root temperature on 14C assimilate profiles in wheat roots

Summary The rate of translocation of ¹⁴C assimilates from leaves to seminal roots in wheat seedlings was considerably reduced by lowering root temperature from 20° to 10° or 5° although the total translocation of ¹⁴C to the roots after 24 h was little affected by temperature. The lowered root temperatures (particularly 5°) resulted in a more uniform distribution of assimilate along the roots than did a temperature of 20°, the ratios of radioactivity/cm in the apical cm, elongating zone, and basal parts of the root after 24 h being 14.0:9.6:1 in 20° roots by contrast with 2.8:1:1 in 5° roots. Temperature effects on assimilate distribution may help explain the observations that for roots grown below 15° ion uptake is sustained in older parts and that roots grown at a low temperature are thicker than roots grown at a higher one.     

Transfer of <Superscript>14</Superscript>C-photosynthate to the sporocarp of an ectomycorrhizal fungus <Emphasis Type="Italic">Laccaria amethystina</Emphasis>

Sporocarps of ectomycorrhizal fungi are strong carbon sinks for the source in host trees, but the details of carbon transfer from the host to the sporocarp are unknown. In this study, single seedlings of Japanese red pine (Pinus densiflora) colonised by Laccaria amethystina were grown on floral foam plates fitted in rhizoboxes, resulting in fruiting on the substrate. The seedlings were photosynthetically labelled with ¹⁴CO₂; ¹⁴C-labelled photosynthate transfer from leaves to sporocarps was then chased using a time-course autoradiography technique. ¹⁴C was transferred to healthy, fresh sporocarps in a purple colour ranging from primordial to elongate sporocarps, but hardly to senesced ones that had faded to white or grey, or browned. This suggested that C is transferred only to physiologically active sporocarps. Two seedlings associated with a growing sporocarp were labelled again 7 and 16 days after the first labelling, respectively. ¹⁴C accumulation in the sporocarps rose in a stepwise manner after the second labelling, indicating that sporocarps mainly used recently rather than previously photosynthesised C.     

Independent translocation of 14C-labelled assimilates and of the floral stimulus in Lolium temulentum

Summary It is widely accepted that the floral stimulus produced in leaves is carried to the shoot apex passively in the phloem with the assimilate stream. Three kinds of evidence presented here suggest that the floral stimulus moves independently of the assimilates.Simultaneous determination of the velocities of translocation out of the seventh leaf blade, in comparable plants under the same conditions, yielded estimates of 1–2.4 cm/hr for the floral stimulus, and 77–105 cm/hr for ¹⁴C-labelled assimilates.The effect of the size of the seventh leaf on its ability to export assimilates or to initiate flowering was quite different. Leaves with only 14–26% of their final blade area emerged exported little assimilate, yet were highly active in inducing flowering.The effect of DCMU applications at a range of concentrations on the translocation of assimilates was quite different from their effect on the flowering response.     

The Translocation of 14C-Labelled Assimilates by Dwarf Bean Plants Infected with Pseudomonas phaseolicola (Burk), Dows

The distribution of ¹⁴C-labelled assimilate after infectionof the dwarf bean plant with Pseudomonas phaseolicola was followed.Infection of a single unifoliate leaf did not affect the totalfixation of ¹⁴CO₂ by unifoliates during the assimilation period.Fixation was maximal in unifoliates in the early stages of growthbut declined as trifoliates expanded. Unifoliates on infectedplants retained a greater proportion of assimilated ¹⁴carbonthan leaves on healthy plants.The pattern of distribution ofexported assimilate was not altered in the early stages of infection,the root and apex acting as the major sinks. As the diseasedeveloped, the first trifoliate leaf, unlike similar leaveson healthy plants, continued to import assimilate apparentlyat the expense of the root. Fixation by the first trifoliateand the distribution of assimilate from this leaf were not alteredby infection of a single unifoliate leaf. At no stage duringdevelopment of the disease was there any evidence of translocationof assimilate to either inoculated or non-inoculated unifoliates.     

Transport 14C-markierter Assimilate im Phloem von Pelargonium zonale und Phaseolus vulgaris

Zusammenfassung 6–10 min nach Beginn der ¹⁴CO₂-Assimilation befindet sich ¹⁴C-Aktivität im Stiel des ¹⁴CO₂-exponierten Blattes belichteter Pflanzen von Pelargonium zonale und Phaseolus vulgaris. Die Verteilungsrichtung der Assimilate im Sproß scheint vor allem durch das Alter des ¹⁴C-assimilierenden Blattes bestimmt, jedoch nicht einseitig festgelegt zu sein; geringe, aber faßbare Mengen an ¹⁴C-Saccharose wereden innerhalb von 20 min (bei Phaseolus) und 180 min (bei Pelargonium) auch aus relativ jüngeren in ältere Blätter transportiert.Neben ¹⁴C-Saccharose wurden im Blattstiel-und Stengelgewebe stets markierte Zuckerphosphate, Hexosen und organische Säuren nachgewiesen. Stärke war nur zu einem verschwindend geringen Anteil an der Gesamtmarkierung der analysierten Transportstrecken beteiligt.

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Translocation of ¹⁴C-labelled assimilates in the phloem of Pelargonium zonale and Phaseolus vulgaris

Summary After the laminae of leaves of intact plants had been exposed to ¹⁴CO₂ the translocation of ¹⁴C-labelled assimilates across the petioles starts very quickly: 6 to 10 min later ¹⁴C-activity could be detected in the basal part of the petioles. The way of distribution within the plant seems to be influenced mainly by the age of the ¹⁴CO₂-assimilating leaf, however, but not in the sense of an unidirectional movement; little but distinct amounts of ¹⁴C were carried also from younger yet full expanded leaves down to older leaves, within 20 min (in Phaseolus) or 180 min (in Pelargonium). Besides sucrose, which was shown by paper chromatography to be the main form of assimilates translocated in these species, we identified sugar phosphates, hexoses, and some organic acids in the petiole and stem tissue as being radioactive also. In our experiments, the petiole segments did not contain any remarkable amount of ¹⁴C-labelled starch.

     

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.     

Translocation of Photosynthate in Curly Top Virus-infected Tomatoes

Photosynthate translocation in single leaflets of healthy and curly top virus-infected tomatoes was investigated using ¹⁴C as a marker. The amount of radioactivity found in plant parts not exposed to ¹⁴CO₂ was substantially lower in diseased than in healthy plants. The time lag for the appearance of ¹⁴C in the petiole was considerably longer in the infected plants than in the healthy. The kinetics of disappearance of ¹⁴C from the lamina during the 24-hour period following labeling showed a strong retention of recent assimilates within the diseased leaf, not accompanied by increased immobilization into insoluble forms. Sucrose was the predominant compound participating in photosynthate transport in both healthy and diseased leaves. The amount of ¹⁴CO₂ fixed was approximately 40% lower in curly top virus-infected leaves than in healthy leaves.     

Distribution of labelled assimilates within a young apple tree after supplying14CO2to a leaf or shoot

Labelled carbon dioxide was supplied for 22 hrs to a leaf of the leader or to the lateral shoot in two-year-old apple seedlings. The distribution of radioactive assimilates within the plant following this treatment was investigated by using radioautography. The transport of labelled assimilates from the young leaf of the leader was very meagre and affected only parts of the stem and the leaves situated in the close vicinity of the treated leaf. The¹⁴C-labelled assimilates from the mature leaf of the leader were transported in a considerable amount to the apex and to the other leaves of the leader. They were also found in an appreciable amount in the stem and the roots, as well as in some lateral shoots. After supplying¹⁴CO₂ to the lateral shoot remarkable transport of labelled assimilates was observed. Radioactivity was detected in the tip and in the youngest leaves of the leader, as well as in the roots. Their path in the stem was studied by dissecting the plant and examining the cross section from each internode. This method revealed that the assimilates from the treated leaf or shoot were transported downward only on one side of the stem in a helical pattern. The lateral shoots situated on the radioactive side of the stem were also labelled, whereas those situated on the opposite (non-radioactive) side were not labelled.