The Export and Distribution of 14C-labelled Assimilates from each Leaf on the Shoot of Lolium temulentum during Reproductive and Vegetative Growth

In both reproductive and vegetative plants of Lolium temulentumL., the export of ¹⁴C-labelled assimilates from each healthyleaf on the main shoot to terminal meristem, stem, tillers,and roots was measured each time a new leaf was expanded, fora period of 5 to 6 weeks. Some labelled assimilates moved fromeach leaf on the main shoot to every meristem in the same shoot,as well as to the tops and roots of adjacent organically attachedtillers. The terminal meristem of the reproductive shoot, which includedthe developing inflorescence, received 70–80 per centof the carbon assimilated by the emerged portion of the growingleaf, 15–25 per cent of the carbon assimilated by thetwo youngest expanded leaves, and 5–10 per cent of thatfrom each of the older leaves. A similar pattern of carbon supplyto the terminal meristem was found in vegetative shoots, exceptthat older leaves on young vegetative shoots supplied even lessof their carbon to the terminal meristem. The general conclusionis that developing leaves at the tip of the shoot receive aboutthe same proportion of carbon from each leaf as does a developinginflorescence. Young expanded leaves provided most labelled assimilates forstem growth; during both reproductive and vegetative growth,expanded leaves increased their export of labelled carbon tostem, and exported less of their ¹⁴C to roots and sometimesto tillers. In these reproductive and vegetative shoots, grown in a constantexternal environment, the major changes in the pattern of distributionof labelled assimilates appeared to be the result of increasedmeristematic activity in stem internodes; the development ofan inflorescence had no obvious direct effect on the carboneconomy of shoots.     

A Quantitative Analysis of the Uptake of Carbon and of the Supply of 14C-labelled Assimilates to Areas of Meristematic Growth in Lolium temulentum

A quantitative analysis of the ¹⁴C-labelled assimilate suppliedby leaves on the main shoot to terminal meristem, stem, tillers,and roots was conducted during parallel periods of reproductiveand vegetative development in Lolium temulentum. The initial rate of entry of carbon into the shoot varied withthe area and photosynthetic efficiency of the assimilating leaf.Subsequently, respiratory losses of carbon during translocationand incorporation of assimilate at the site of utilization alsovaried. The combined effect of these differences resulted inthe supply of recently assimilated carbon being twofold greaterin reproductive shoots than in vegetative shoots, while withinshoots the carbon supply of the youngest fully expanded leafranged from four-or five-fold greater than the oldest leaf inyoung shoots, to two-or three-fold greater in older shoots.In both reproductive and vegetative shoots, the two or threeyoungest leaves thus dominated the supply of carbon for meristematicgrowth. Meristematic tissue in expanding leaves and leaf primordia atthe terminal meristem of the vegetative shoot received 18–27per cent of the total shoot carbon. This meristem utilized aboutthe same proportion of shoot carbon when it developed into aninflorescence, indicating no major change in the level of meristematicactivity. The proportion of shoot carbon utilized in stem growthincreased as both reproductive and vegetative shoots aged; thisincreased meristematic activity in stem internodes was accompaniedby reduced export of carbon to roots, which received less than10 per cent of the shoot carbon when the experiments ended.The main shoot translocated 20–30 per cent of its recentlyassimilated carbon to developing and rooted tillers, which assinks for carbon were thus as important as the terminal meristemand stem. This outward flow of carbon continued relatively uncheckedwhen donor and receptor shoots developed inflorescences.     

Reversal of the Direction of Photosynthate Allocation in Urtica dioica L. Plants by Increasing Cytokinin Import into the Shoot

The natural cytokinin import from the root into the shoot of Urtica dioica plants was enhanced by supplying zeatin riboside (ZR) solutions of various concentrations to a portion less than 10 % of the root system after removal of their tips. After 6 h ZR pretreatment of the plants, ¹⁴CO₂ was supplied for 3 h to a mature (source) leaf or to an expanding leaf and the ¹⁴C-distribution in the whole plant was determined after a subsequent dark period of 14 h. ZR substantially increased ¹⁴C fixation by the expanding leaves and also enhanced export of carbon and transport to the shoot apex. The effect of the hormone treatment was, however, more pronounced when the ¹⁴CO₂ was supplied to a mature leaf. In the control plants these leaves exported carbon only to the roots: When the amount of the natural daily ZR input from the roots to the shoot was enhanced by 20%, the bulk of the ¹⁴C exported from a mature leaf moved to the shoot apex and only a minor portion of ¹⁴C was still detected in the root fraction. A several-fold increase of the natural daily ZR input into the shoot resulted in a flow of ¹⁴C only to the growing parts of the shoot. The results suggest control of the sink strength of the shoot apex by ZR in Urtica diocia.     

Distribution of 14C-sucrose applied to leaves at different nodes of okra (Abelmoschus esculentum) during flowering and early pod growth

The distribution pattern of ¹⁴C-sucrose from ¹⁴C-sucrose applied to vegetative okra plants and leaves 1–9 on separate plants during the green pod development stage were investigated in relation to duration and leaf position. Results indicated bi-directional transport of assimilates to both apical and basal portions of the stem. Within 48 h ¹⁴C moved to all plant parts; stem and leaves appeared to be strong sinks. In plants fed at the vegetative stage, 48 h after feeding, 66% of the fed activity was exported from the fed leaf. At the pod development stage, about 35% of the activity exported from the fed leaf was present in green pods and 65% in vegetative parts. In plants where leaf 1–9 was fed, irrespective of the position of the fed leaf, the subtending fruit was the strongest sink among the reproductive parts. Leaves and stems were the principal sinks.     

Effect of nitrogen deficiency on photosynthesis and the partitioning of 14C-labelled leaf assimilate in unshaded and partially shaded plants of Lolium temulentum

The effect of a deficiency of applied nitrogen on the rate of leaf photosynthesis, and on the subsequent partitioning of ¹⁴C-labelled leaf assimilate between new leaf, stem, tillers and root, was investigated in single plants of Lolium temulentum L., grown normally in controlled environments, or grown with collars shading the base of the plant. The nitrogen deficiency reduced the rate of leaf photosynthesis, increased the retention of assimilate in the leaf, suppressed the export of assimilate to tillers, and generally increased the export of assimilate to roots and to new leaves. Shading the base of the plant generally had little effect on the rate of leaf photosynthesis, reduced the export of assimilate to roots, and increased the export of assimilate to new leaf and to the stem, which elongated when shading was imposed.     

Analysis of assimilate distribution by 3H-tracing during recovery period after rice seedling transplanting

In order to characterize assimilate distribution in rice (Oryza sativa L.) seedlings during recovery period after transplanting, the experiments were conducted by using ³H-isotope tracer technique with three rice hybrid combinations. The label introduced into the leaf sheath was distributed and redistributed to young roots, tillers, and young leaves, thus promoting these organs to grow at the initial stage after transplanting. During period from 10 to 30 days after labeling, ³H-assimilate redistribution occurred mainly from the sheath of the main stem; some radioactivity moved to tillers and roots emerged after transplanting. The activity and percentage of ³H-compounds transporting to rooting nodes and roots emerged after transplanting were higher in root-pruned plants than in control, indicating that growth of root-pruned seedlings more depended on the export of the compounds from the sheath of the main stem than growth of control seedlings.     

Effect of decapitation on absorption,translocation, and phytotoxicity of imazamethabenz in wild oat (Avena fatua L.)

The release of apical dominance by the physical destruction in situ of the apical meristem and associated leaf primordia (decapitation) promoted the growth of tillers in non-herbicide-treated wild oat plants, as indicated by increased tiller lengths and fresh weights. At 96 h after [¹⁴C] herbicide treatment following decapitation, the absorption of [¹⁴C]imazamethabenz and total translocation of radioactivity were respectively increased by 28% and 49%. By 96 h after [¹⁴C]imazamethabenz application, the radioactivity detected in the roots of decapitated plants was 45% higher than that in the roots of nondecapitated plants while the radioactivity in tillers of decapitated plants was 2.6-fold that in tillers of intact plants. Decapitation together with foliar spraying of imazamethabenz at 200 g ha^–1 further reduced tiller fresh weight, greatly decreased the total tiller number, and thereafter significantly increased overall phytotoxicity by 32% as measured by total shoot fresh weight. The results of this study support the hypothesis that main shoot apical dominance limits translocation of applied imazamethabenz to lateral shoots, rendering tillers less susceptible to growth inhibition by the herbicide.     

14CO2 Assimilation and 14C-photosynthate Translocation in Different Leaves of Sunflower

Chlorophyll and nitrogen contents were highest in leaves of middle position, similarly as photosynthetic efficiency represented by ¹⁴C fixation (maxima in leaf 5 from the top). All the leaves lost ¹⁴C after 2 weeks of ¹⁴CO₂ exposure. However, the reduction in radioactivity was less in young upper leaves than in the mature lower leaves. Leaves exported ¹⁴C-photosynthates to stem both above and below the exposed leaf. Very little radioactivity was recovered from the seeds of plants in which only first or second leaves were exposed to ¹⁴CO₂ implying thereby that the carbon contribution of first two leaves to seed filling was negligible. The contribution of leaves to seed filling increased with the leaf position up to the sixth leaf from the top and after the seventh leaf their contribution to seed filling declined gradually.     

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.     

Enhancement of [C]Sucrose Export from Source Leaves of Vicia faba by Gibberellic Acid

The effect of gibberellic acid (GA₃) on sucrose export from source leaves was studied in broad bean (Vicia faba L.) plants trimmed of all but one source and one sink leaf. GA₃ (10 micromolar) applied to the source leaf, enhanced export of [¹⁴C]sucrose (generated by ¹⁴CO₂ fixation) to the root and to the sink leaf. Enhanced export was observed with GA treatments as short as 35 minutes. When GA₃ was applied 24 hours prior to the ¹⁴CO₂ pulse, the enhancement of sucrose transport toward the root was abolished but transport toward the upper sink leaf was unchanged. The enhanced sucrose export was not due to increased photosynthetic rate or to changes in the starch/sucrose ratio within the source leaf; rather, GA₃ increased the proportion of sucrose exported. After a 10-min exposure to [¹⁴C]GA₃, radioactivity was found only in the source leaf. Following a 2 hour exposure to [¹⁴C]GA₃, radioactivity was distributed along the entire stem and was present in both the roots and sink leaf. Extraction and partitioning of GA metabolites by thin layer chromatography indicated that there was a decline in [¹⁴C]GA₃ in the lower stem and root, but not in the upper stem. This pattern of metabolism is consistent with the disappearance of the GA₃ effect in the lower stem with time after treatment. We conclude that in the short term, GA₃ enhances assimilate export from source leaves by increasing phloem loading. In the long term (24 hours), the effect of GA₃ is outside the source leaf. GA₃ accumulates in the apical region resulting in enhanced growth and thus greater sink strength. Conversely, GA₃ is rapidly metabolized in the lower stem thus attenuating any GA effect.     

Transport of Photoassimilates in Young Trees of Fraxinus and Sorbus: Measurement of Translocation in vivo

Abstract: Translocation of photoassimilates was studied on 2-year-old trees of Fraxinus excelsior and Sorbus aucuparia using the short-lived isotope ¹¹C. Leaflets of different leaves on the same plants were radiolabeled showing that both carbon distribution and speeds of transport may vary with leaf position. Within 2 h after pulse feeding with ¹¹CO₂, mainly the lower leaves distributed radiolabel to the roots in Fraxinus, whereas in Sorbus, the upper leaves were also involved. By repeated pulse applications to selected leaves, temporal profiles of ¹¹C transport were followed on individual plants from April to October. Early in the season, within 2 h after pulse labelling, 30–40% of the fixed radiolabel was exported from leaves in Fraxinus and about 20% in Sorbus. Thereafter export started to decline, particularly in Fraxinus, and the distribution of radiolabel between stem and roots could alter depending on the position of the feed leaf. Speeds of translocation obtained along the rachis and stem showed high variability, but they did not necessarily slow down before the end of the season. The speeds monitored at the rachis of Fraxinus leaves (30–75 cm h^?1) were generally lower than those found on Sorbus (50–130 cm h^?1). As reported in the literature, the two tree species translocate different carbohydrates and show remarkable differences in the ultrastructure of their vascular systems. In that context it is interesting that the temporal profiles of ¹¹C radioactivity obtained from F. excelsior and S. aucuparia could clearly be distinguished by their characteristic shapes. The results are discussed in terms of anatomical characteristics of the conducting tissues and possible differences in phloem loading.     

Distribution between parts of the main shoot and the tillers of photosynthate produced before and after anthesis in the top three leaves of main shoots of Hobbit and Maris Huntsman winter wheat

The top three leaves of main shoots in crops of Hobbit and Maris Huntsman winter wheat were exposed to ¹⁴CO₂ at 22 and 16 days before and at 10 days after anthesis in 1978. The distribution of the ¹⁴C recovered in whole plants at anthesis and at maturity was measured. There was negligible loss of ¹⁴C between these two times, but some redistribution. The percentage in the tillers was negligible when the flag leaf (leaf 1) was exposed to ¹⁴CO₂, and otherwise less than 12% except for ¹⁴C absorbed by the third leaf at 16 days before anthesis, when it averaged 26% but was very variable. When ¹⁴C was supplied before anthesis, about 20% reached the grain whichever leaf had been supplied. The ear structures contained about 10% of that absorbed by the third leaf and 35% of that absorbed by the flag or second leaf. When ¹⁴C was supplied after anthesis, the amounts reaching the grain from the different leaves were: flag 82%, second 68%, third 56%. Most of the remainder was in the stem. The exposed leaf never retained more than 6%. The amount of ¹⁴C that moved from the stem to the grain between anthesis and maturity was about 50% greater in the semi-dwarf variety Hobbit than in Maris Huntsman. There was no significant varietal difference in the percentage of post-anthesis ¹⁴C reaching the grain. The ear structures of Hobbit contained about a third more ¹⁴C than those of Maris Huntsman. An additional 90 kg N ha^-1, which increased grain yield by 46%, had negligible effects on the distribution of ¹⁴C.     

Fixation and distribution of 14C in Populus deltoides during dormancy induction

Photosynthetically fixed ¹⁴C was analyzed in various chemical fractions from leaves and stems of cottonwood (Populus deltoides Bartr. ex. Marsh.) during dormancy induction. Dormancy was induced by 8-h photoperiods and 20/14°C temperature regimes. Within 4 weeks under short days, terminal buds were set and leaf expansion and stem elongation had stopped. ¹⁴C₂ was fed to a leaf at Leaf Plastochron Index 7 for 30 min. Either after this 30 min feeding period or after a 48-h translocation period the plants were sampled, freeze-dried, extracted and analyzed for¹⁴C. ¹⁴C-fixation decreased during dormancy induction from 60% to 17% of the 3.7 MBq ¹⁴C applied at 0 week and 8 weeks, respectively. Percentage distribution of ¹⁴C in chemical fractions of source leaves reflected leaf age and translocation inhibition. In rapidly growing plants, considerable ¹⁴C was incorporated into leaf protein while most of the soluble¹⁴C-sugars were either metabolized or translocated out of the leaf. After terminal bud set, the percentage of ¹⁴C in the protein and residue fractions decreased rapidly and that in the sugar fraction increased. Percent distribution in stems closely reflected changing metabolic pathways of carbon flow as influenced by dormancy induction. For example, the ¹⁴C in structural carbohydrates decreased in 5 weeks under short days from 65 to less than 10% of the ¹⁴C recovered in the chemical fractions, thus indicating cambium inhibition. At the same time the percentage of ¹⁴C in starch and sugar increased indicating storage. Short term (after 30 min) incorporation of ¹⁴C into the protein and starch fractions of leaves changed relatively little throughout the 8-week induction period. In contrast the turnover rates of these fractions (¹⁴C present after 48 h) increased considerably after active growth of the whole plant stopped.     

The Distribution and Identity of Assimilates in Tomato with Special Reference to Stem Reserves

Much of the work on the distribution of ¹⁴C-labelled assimilatesin tomato has been done in winter under low light intensities,and consequently the reported distribution patterns of ¹⁴C maynot be representative of plants growing in high light. Further,there are several somewhat conflicting reports on patterns ofdistribution of ¹⁴C-assimilates in young tomato plants. We soughtto clarify the situation by studying the distribution of ¹⁴C-assimilatesin tomato plants of various ages grown in summer when the lightintensity was high. In addition, the role of the stem as a storageorgan for carbon was assessed by (a) identifying the chemicalfractions in the stem internode below a fed leaf and monitoring¹⁴ C activity in these fractions over a period of 49 d, and(b) measuring concentrations of unlabelled carbohydrates inthe stem over the life of the plant. The patterns of distribution of ¹⁴C-assimilates we found fortomato grown under high light intensity confirmed some of thosedescribed for plants grown under low light, but export of ¹⁴Cby fed leaves was generally higher than reported for much ofthe earlier work. Lower leaves of young plants exported over50% of the ¹⁴C they fixed, although export fell sharply as theplants aged. Initially, the roots and apical tuft were strongsinks for assimilates, but they had declined in importance bythe time plants reached the nine-leaf stage. On the other hand,the stem became progressively more important as a sink for ¹⁴C-assimilates.Older, lower leaves exported more of their ¹⁴C-assimilates tothe upper part of the plant than to the roots, whereas youngleaves near the top of the plant exported more of their assimilatesto the roots. The stem internode immediately below a fed leafhad about twice the ¹⁴C activity of the internode above theleaf. Mature leaves above and below a fed leaf rarely importedmuch ¹⁴C, even when in the correct phyllotactic relationshipto the fed leaf. In the first 3 d after feeding leaf 5 of nine-leaf plants, theorganic and amino acid pools and the neutral fraction of theinternode below the fed leaf had most of the ¹⁴C activity, butby 49 d after feeding, the ethanolic-insoluble, starch and lipidfractions had most of the ¹⁴C activity. Glucose, fructose andsucrose were the main sugars in the stem. Although concentrationsof these sugars and starch declined in the stem as the plantsmatured, there was little evidence to indicate their use infruit production. Stems of plants defoliated at the 44-leafstage had lower concentrations of sugars and starch at maturity,and produced less fruit than the controls. It was concludedthat tomato is sink rather than source limited with respectto carbon assimilates, and that the storage of carbon in thestem for a long period is possibly a residual perennial traitin tomato.Copyright 1994, 1999 Academic Press Lycopersicon esculentum, tomato, assimilate distribution, ¹⁴C, internode storage, sink-source relationships, starch, stem reserves, sugars     

Effect of Rate of Photosynthesis on the Pattern of Assimilate Distribution in the Graminaceous Plant

Infrared gas analysis and a quantitative radiocarbon tracertechnique were used to measure photosynthesis and the distributionof ¹⁴C-labelled assimilate in Lolium temulentum and a uniculmbarley exposed continuously or intermittently to contrastinglight intensities. Plants grown for 10–20 d in low light(<50 W m^–2 of visible light) exported a greater proportionof their assimilate to growing leaves at the terminal meristemand a smaller proportion to their roots and tillers than equivalentplants in high light (150 W m^–2). A single day's exposure(8.4 h photoperiod) to a contrasting light regime elicited achange in the pattern of assimilate distribution in the samediurnal period, but 2–3 d exposure was required for asubstantial shift in the pattern of supply of assimilate tomeristems. The data indicated that in terms of assimilate distributioncomplete adaption to a new light regime is attained in about7 d.     

Interorgan photosynthate partitioning in alfalfa

The net partitioning of current photosynthate among vegetative organs of nodulated alfalfa (Medicago sativa L.) was investigated by determining radiolabel distribution from upper and lower source leaves, at different times of day, and during successively longer chase periods. Photosynthate was exported sooner and more completely by a fully expanded lower than by a fully expanded upper source leaf. The radiolabel pulse from the lower source leaf peaked in the main stem within 1.5 hours, in the crown and nodules after 3 hours, in the unexpanded leaves and apex of the main stem after 6 hours, and in the shoots growing from leaf axils on the main stem after 24 hours. The results suggest that the crown, apex, axillary shoots, and nodules both imported and mobilized photosynthate originating at a lower source leaf, while roots and shoots growing from the crown showed net accumulation. The pulse from the upper source leaf was initially rapidly exported by the main stem and imported by the root between 1.5 and 3 hours, but there was no net change of label content of these organs during the ensuing 21 hours. Rapidly growing organs had the highest concentrations, and the largest organs had the highest content of radiolabel. These results provide new information about the accumulation and circulation of photosynthate within the alfalfa plant.     

Utilization of exogenously supplied 14C-saccharose into primary metabolites and alkaloid production in Catharanthus roseus

Partitioning of exogenously supplied U-¹⁴C-saccharose into primary metabolic pool as sugars, amino acids, and organic acids was analyzed and simultaneous utilization for production of alkaloid by leaf, stem, and root in twigs and rooted plants of Catharanthus roseus grown in hydroponic culture medium was determined. Twigs revealed comparable distribution of total ¹⁴C label in leaf and stem. Stems contained significantly higher ¹⁴C label in sugar fraction and in alkaloids [47 kBq kg⁻¹(DM)] than leaf. In rooted plants, label in ¹⁴C in metabolic fractions in root such as ethanol-soluble, ethanol-insoluble, and chloroform-soluble fractions and in components such as sugars, amino acids, and organic acids were significantly higher than in stems and leaves. This was related with significantly higher content of ¹⁴C in alkaloids in stems and leaves. ¹⁴C contents in sugars, amino acids, and organic acids increased from leaf to stem and roots. Roots are the major accumulators of metabolites accompanied by higher biosynthetic utilization for alkaloid accumulation.     

Transfer of exogenous auxin from the phloem to the polar auxin transport pathway in pea (Pisum sativum L.)

When [1-¹⁴C]indol-3yl-acetic acid ([1-¹⁴C]IAA) was applied to the upper surface of a mature foliage leaf of garden pea (Pisum sativum L. cv. Alderman), ¹⁴C effluxed basipetally but not acropetally from 30-mm-long internode segments excised 4 h after the application of [1-¹⁴C]IAA. This basipetal efflux was strongly inhibited by the inclusion of 3.10^–6 mol· dm³ N-1-naphthylphthalamic acid (NPA) in the efflux buffer. In contrast, when [¹⁴C] sucrose was applied to the leaf, the efflux of label from stem segments excised subsequently was neither polar nor sensitive to NPA. The [1-¹⁴C]IAA was initially exported from mature leaves in the phloem — transport was rapid and apolar; label was recovered from aphids feeding on the stem; and label was recovered in exudates collected from severed petioles in 20 mM ethylenediaminetetraacetic acid. No ¹⁴C was detected in aphids feeding on the stems of plants to which [1-¹⁴C]IAA had been applied apically, even though the internode on which they were feeding transported considerable quantities of label. Localised applications of NPA to the stem strongly inhibited the basipetal transport of apically applied [1-¹⁴C]IAA, but did not affect transport of [1-¹⁴C]IAA in the phloem. These results demonstrate for the first time that IAA exported from leaves in the phloem can be transferred into the extravascular polar auxin transport pathway but that reciprocal transfer probably does not occur. In intact plants, transfer of foliar-applied [1-¹⁴C]IAA from the phloem to the polar auxin transport pathway was confined to immature tissues at the shoot apex. In plants in which all tissues above the fed leaf were removed before labelling, a limited transfer of IAA occurred in more mature regions of the stem.Abbreviations IAA indol-3yl-acetic acid - EDTA ethylenediaminetetraacetic acid - NPA N-1-naphthylphthalamic acid We are grateful to the Nuffield Foundation for supporting this research under the NUF-URB95 scheme and for the provision of a bursary to A.J.C. We thank Professor Dennis A. Baker for constructive comments on a draft of this paper and Mrs. Rosemary Bell for her able technical assistance.     

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.     

The Influence of Leaf Position and Defoliation on the Assimilation and Translocation of Carbon in White Clover (Trifolium repens L.) 1. Carbon Distribution Patterns

The assimilation of carbon (C) by, and distribution of ¹⁴C from,leaves at each end of an unbroken sequence of ten mature leaveson the main stolon of clonal plants of white clover (Trifoliumrepens L.) were measured to identify intra-plant factors determiningthe direction of C movement from leaves. Leaves at two intermediatepositions were also measured. Localized movement of ¹⁴C to sinks at the same node as, or atthe one to two nodes immediately behind, the fed leaf accountedfor 40–50% of the total ¹⁴C exported by all measured leaves.A further 50–60% of exported ¹⁴C was therefore availablefor more-distant sinks, and the direction of translocation ofthis C was determined by the relative total strength or demand(number x size x rate of activity or growth) of sinks forwardof, or behind, the leaf in question. Thus 85% of the ¹⁴C exportedfrom the youngest measured leaf moved toward the base of thestolon, while about 60% of the ¹⁴C exported from the oldestleaf moved acropetally. Defoliating plants to leave just one mature leaf on the mainstolon (at any one of the same four positions studied in undefoliatedplants), and no leaves on branches, resulted in: (1) increasednet photosynthetic rate in all residual leaves: (2) increased%export of fixed C from one of the four leaves; (3) increasedexport to the main stolon apex from all except the eldest leaf;(4) increased export to branches from three of the four leaves;and (5) decreased export to stolon tissue and roots from allleaves, within 3 d of defoliation. These responses would seemto ensure the fastest possible replacement of lost leaf areaand, thus, restoration of homeostatic growth. The observed patternsof C assimilation and distribution in both undefoliated anddefoliated white clover plants are consistent with the generalrules of source-sink theory; the distance between sources andcompeting sinks, and relative sink strength, emerge as the mostimportant intra-plant factors governing C movement. These resultsemphasize the need to consider plant morphology, and the modularnature of plant growth, when interpreting patterns of resourceallocation in clonal plants, or plant responses to stressessuch as partial defoliation. Trifolium repens L, white clover, photosynthesis, assimilate translocation, defoliation