Investigation of Bidirectional Movement of Tracers in Sieve Tubes of Salix Viminalis L.

Sieve-tube sap was collected as honeydew from the aphid Tuberolachnussalignus (Gmelin) feeding on a young shoot or a mature stemof willow (Salix viminalis L.). Two radioactive tracers (either³H-glucose and ¹⁴C-sucrose, or ¹⁴CO₂, Na₂ ¹⁴CO₃, and ³²P-orthophosphate)were applied to leaves or to bark abrasions at each end of thestem. In some experiments the tracers were applied after thestart of honeydew production, whilst in others the tracers wereapplied before the aphids were sited. In most cases aphids feeding between the tracer applicationpoints produced doubly-labelled honeydew during a 24–48-hperiod from tracer application. In some instances the firstdrops of honeydew contained one tracer, followed several hourslater by the other tracer, whilst some aphids produced doubly-labelledhoneydew from the first drop. Singly-labelled or inactive honeydewwas found in only a few cases. It was demonstrated that when ³H-glucose and ¹⁴C-sucrose wereapplied at opposite ends of a stem that the tracers moved inopposite directions in the form of sucrose. The data suggest that a simultaneous bidirectional movementof two tracers may occur in the same sieve tube. However, thepossibility that the doubly-labelled honeydew could be producedby lateral movement from one sieve tube carrying one tracerto an adjacent punctured sieve tube carrying the other tracercannot be discounted.     

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     

Changes in the Distribution of 14C-Labelled Assimilates in Sugar-beet with Variation of Temperature

Plants were allowed to assimilate ¹⁴CO₂ for 30 min at 5, 15,25, and 35 °C. The changes in ¹⁴C content of a mature expandedleaf (Leaf 4), young apical leaves, and storage root, were sequentiallyfollowed over a subsequent period of 24 h in continuous light.In a second experiment plants were transferred after ¹⁴CO₂ assimilationto temperatures of 10, 18, 26, and 34 °C, and the partitionof ¹⁴C between the ethanol-soluble and ethanol-insoluble fractionsof the roots and leaves was followed over a period of 72 h. The specific activities of the apical leaves and of the storageroot increased to a maximum 2 h after labelling at 25 °C,4 h at 15 and 35 °C, and 6 h at 5 °C suggesting thatthe optimum temperature for translocation of photosynthate wasabout 25 °C. The ¹⁴C partition to ethanol-soluble and ethanol-insoluble fractionsof the roots and leaves was largely attained in. 9 h. Littlerepartition of ¹⁴C assimilate fractions occurred as a resultof temperature change or growth. Root ethanol-insoluble activity,however, did increase significantly over the 72-h period : possiblecauses of this slow incorporation and their relevance to themechanism of sugar storage are discussed.     

Investigations of Carbon Transport in Plants: I. THE USE OF CARBON-11 TO ESTIMATE VARIOUS PARAMETERS OF THE TRANSLOCATION PROCESS

A technique is described which enables several components ofthe translocation system to be measured routinely, in vivo andsimultaneously. Plants are exposed to a 1-min pulse of ¹¹CO₂and the movement of the ¹¹C pulse through the plant followedusing an array of scintillation counters. It is possible toestimate the time taken for the ¹¹C to enter the sieve tubes.In addition, the speed of translocation can be obtained frommeasurements of the arrival of the ¹¹C pulse at different positionsalong the translocation pathway. The increasing half-width ofthe pulse as it enters and moves through the sieve tubes givessome indication of the extent of the delays in the differentparts of the translocation system. The short half-life of the¹¹C allows the measurements to be repeated on the same plantseveral times per day. A technique is discussed whereby dataon the movement of ¹¹C could be combined with simultaneous measurementsof the rate of uptake of carbon dioxide to provide an estimateof the rate of movement of carbon.     

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     

Electron-Microscopic Evidence of the Vacuolar Nature of Phloem Exudate

We performed electron-microscopic examination of structural diurnal changes in the lumen of sieve tubes and the vacuolar system of corresponding companion cells and changes induced by the experimental blockage of assimilate export from the leaf by its cold-girdling. For these investigations, Cucurbita pepo L. and Helianthus annuus L. plants were used, that is, plant species from groups of symplastic and apoplastic plants, which differ in the type of companion cells and a mode of phloem terminal loading. The examinations showed the complete identity of changes in the electron texture of the sieve-tube lumens and companion-cell vacuoles in both plant species in the course of a day, when the level of assimilates changed, or after export blockage. Similar changes in the structure of the vacuolar labyrinths were stated in the companion cells under normal conditions and after cold-girdling, as related to the rate of sieve-tube loading with the vacuolar exudate. Vacuolar expansion and starch accumulation developing in response to changes in the assimilate level in the evening and after cold blockage of the assimilate export occurred in different types of cells, as dependent on their position in the symplast domains. However, the rate of the process similarly depended on the balance between assimilate synthesis and export. Synchronous changes in the texture of the sieve-tube lumen and companion-cell vacuoles were observed within each complex, but asynchronous changes occurred in different complexes. We suggested this phenomenon for recognizing the particular complexes, when they are grouped in a bundle. We observed no signs of cytoplasm or protein synthetic machinery in the sieve tubes. We concluded that the sieve-tube lumen and vacuoles of companion cells are common in nature. Similar electron texture of the images of the companion-cell vacuolar labyrinth and tube lumens, their connection through the lateral sieve fields, morphological modifications of the companion-cell vacuolar system as dependent on the activity of sieve tube loading—all of these facts imply the continuity of these transport compartments and fluxes in them and the similarity in the composition of the exudates from companion-cell vacuoles and phloem tubes.     

Modifications of Translocation Rate in Studies of Apple Leaf Efficiency

M.7 apple rootstocks were used during the peak period of shootextension for comparisons of dry-matter production per unitleaf area between intact plants and others which had been partiallydefoliated. Dry-matter increment per unit leaf area over a 16-dayinterval was some 70 per cent higher in partially defoliatedplants than in controls. ¹⁴CO₂ was supplied to designated leaves of comparable age andposition. Sample discs were taken from the ‘fed’leaves at intervals up to 9 days from supplying ¹⁴CO₂. Translocationrates were estimated by comparison with leaves on a third setof plants whose petioles were steamed to prevent translocationimmediately on removal of the ¹⁴CO₂ feeding chambers. Translocationrates in partially defoliated plants were enhanced some 30 percent compared with controls. It is suggested that features of the plant outside the studiedleaves may have contributed to the overall efficiency of assimilateproduction and utilization. Malus sylvestris L., apple, dry matter production, leaf efficiency, defoliation, translocation, assimilate distribution, sorbitol, sucrose     

The Effect of Calcium Starvation on Assimilate Partitioning and Mineral Distribution of the Phloem

Populus plants were grown in a medium lacking calcium and exposedto ¹⁴CO₂. In contrast to plants in the complete nutrient medium,the percentage amount of ¹⁴C-assimilates increased in the leavesof calcium-deficient plants and decreased in the stem and theroots. When plants were grown without potassium or magnesiumno differences in the amount of ¹⁴C-label occurred in comparisonwith plants in the complete nutrient medium. Translocation wasrecorded by microautoradiography. It was observed that considerableamounts of labelled photoassimilates were unloaded from thephloem in the middle part of the stem in plants of the completenutrient medium. In contrast, during calcium starvation ¹⁴C-labelwas restricted to the phloem of the stem. In addition, the concentrationsof magnesium and phosphorus showed a remarkable increase instem sieve tubes of calcium-deficient plants. When sieve tubesof source leaves from Populus, barley and maize were comparedwith those of sink leaves, the latter showed higher calciumconcentrations. The results suggest that calcium is a necessaryfactor in the regulation of phloem translocation. Key words: Calcium deficiency, phloem translocation, sieve element loading and unloading, X-ray microanalysis     

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.     

Changes in Partitioning of Current Assimilate During Tuber Bulking in Potato (Solarium tuberosum L.) cv Maris Piper

Potato plants were labelled with ¹⁴CO₂ at six stages duringtuber bulking and changes in current assimilate partitioning(20 h after ¹⁴CO₂ assimilation) were examined in relation toincreasing tuber size on a plant. There was no relationshipbetween the node of origin of a tuber and the amount of ¹⁴Cwhich entered it, and those tubers on a plant which importedthe most ¹⁴C did not all belong to the same stem. However, competitionwithin a node was evident as many tubers borne on second-orderstolons were smaller and contained significantly lower concentrationsof ¹⁴C than those on primary stolons. Two weeks after tuberinitiation there was an almost linear relationship between thefresh weights of the tubers and their ¹⁴C content but the correlationbecame less good at subsequent harvests. Within the tubers theratio of ¹⁴C starch: soluble fell over the course of the experimentand was reflected in an increase in the percentage of ¹⁴C sucroseappearing in the tubers at each harvest. At any one harvest,however, the ¹⁴C starch: soluble ratios were similar but notrelated to the rates of ¹⁴C import of the tubers. The data areconsistent with the view that sucrose is compartmentalized inpotato tubers. Assimilate, ¹⁴Carbon, potato, sugars, tuber     

Partitioning of 14C Labelled Assimilates in Arctium tomentosum

Plants of the biennial Arctium tomentosum were grown from seedto seed-set in an open field under three different treatments:control plants receiving full light intensity, plants with aleaf area reduced by 45 per cent, and shaded plants receivingonly 20 per cent of natural illumination. At various stagesof development the youngest fully expanded leaf of one plantin each treatment was exposed to ¹⁴CO₂ for half an hour. Subsequentdistribution of labelled assimilates in various plant partswas determined after eight hours. In the first year, the mostdominant sink was the tap root irrespective of variation inassimilate supply. During the production of new vegetative growthin the second season, a larger amount of radioactive photosynthatewas recovered from above ground parts, especially during formationof lateral branches. Seed filling consumed 80–90 per centof labelled carbon exported from the exposed leaf. In the secondyear, the most pronounced difference between treatments wasin the degree of apical dominance, being highest in shaded plantsand lowest in the plants with cut leaves. Results from ¹⁴C experimentsagreed fairly well with a ‘partitioning coefficient’derived from a growth analysis of plants grown independentlyunder the same experimental conditions. Reasons for discrepanciesbetween the ¹⁴C results and the partitioning coefficient arediscussed. Arctium tomentosum, burdock, variation in assimilate supply, assimilate distribution, ¹⁴CO₂, labelling, growth analysis     

Translocation of Carbohydrate in Cotton: Movement to the Fruiting Bodies

The translocation of ¹⁴C-labelled assimilate was followed frommain stem leaves on the cotton plant (Gossypium hirsutum) tobolls on the lower sympodia. From the upper leaves the assimilatedescends the stem in well-defined strands in the phloem. Wheresympodia originate close to these strands some of the assimilateis distributed to the bolls on those sympodia. There is evidencethat the involucre, but not the green boll wall, contributesto boll nutrition by assimilation of external CO₂.     

The Redistribution of Assimilate in Field-grown Winter Wheat

The long- and short-term movement of carbon in a field cropof winter wheat was investigated with the radioactive tracercarbon-14. The flag leaves of individual plants assimilateda pulse of ¹⁴CO₂ and the plant parts were assayed subsequently.Groups of plants were pulse-labelled four times during the mainperiod of growth—twice before and twice after anthesis.Plants were harvested and assayed twice weekly after labellingand the time-course of the changes in the amount of ¹⁴C recoveredfrom the leaves, stems and ears was observed for each groupof plants. Concurrently with these long-term studies, otherwheat plants were pulse-labelled, then harvested and assayed24 h later. The partitioning of ¹⁴C between leaves, stem, earand in some cases roots, was measured over the period from thestart of stem elongation to the end of grain filling. Two distinct types of relocation of carbon were observed. Carbonassimilated early in the growth of the plant and used in thegrowth of new leaves was seen to be partly relocated to theear. Carbon assimilated 8 d after anthesis was partly storedin the stem, and 15 d later relocated to the ear. This relocationcorresponded to a decrease in stem dry mass seen in growth analysis.Little other change in the ¹⁴C content of the plants occurred,suggesting that most respiration used current rather than storedassimilate. Key words: Carbon assimilate, redistribution, wheat     

The Response of Groundnut (Arachis hypogaea L.) to Timing of Irrigation: II. 14C-PARTITIONING AND PLANT WATER STATUS

Finite quantities of water were applied at different growthstages of groundnut stands (Arachis hypogaea L.) grown in controlledenvironment glasshouses. Soil moisture deficits were imposedbetween sowing and pod initiation or between pod initiationand final harvest by withholding or applying water. Effectson assimilate production and partitioning and plant water relationswere examined. Leaves were the primary sites of ¹⁴CO₂ fixation, though theircontribution generally declined late in the season, whereasfixation by stems was initially low but increased sharply whenstress was released in the late-irrigated stands. ¹⁴C-fixationby stem apices and pegs also rose sharply following irrigationof the late-stressed stands. Leaves were the primary source of assimilates, though translocationtended to decrease as the season progressed, even in the late-irrigatedstands. Stems were initially the major sinks, but their sinkactivity disappeared almost completely when stress was releasedin the late-irrigated stands. Assimilate import by stem apicesdeclined progressively and pod sink activity was negligiblein the late-stressed stand, but both increased markedly whenearly-season stress was released. Leaf water status showed marked diurnal variation, whereas pegsshowed less variation and maintained much higher turgor levels,largely because of their lower solute potentials. Marked osmoticadjustment occurred in expanding but not in mature leaves, allowingthem to maintain higher turgor levels during periods of severestress. This adjustment was rapidly lost when stress was released.The observed changes in assimilate production and partitioningpreceded detectable changes in bulk turgor levels. Implications for growth, development and yield are discussed. Key words: Groundnut, irrigation, partitioning, water status     

Effects of elevated carbon dioxide on three grass species from montane pasture II. Nutrient uptake, allocation and efficiency of use

Agrostis capillaris L.⁴ Festuca vivipara L. and Poa alpinaL.were grown in outdoor open-top chambers at either ambient (340µmol mol^–1) or elevated (680 µmol^–1)CO² for periods from 79 to 189 d. Under these conditions thereis increased growth of A. caplllarls and P. alpina, but reducedgrowth of F. vivipara. Nutrient use efficiency, nutrient productivity(total plant dry weight gain per unit of nutrient) and nutrientallocation of all three grass species were measured in an attemptto understand their individual growth responses further andto determine whether altered nutrient-use efficiencies and productivitiesenable plants exposed to an elevated atmospheric CO₂ environmentto overcome potential limitations to growth imposed by soilfertility. Total uptake of nutrients was, in general, greater in plantsof A. capillaris and P. alpina (with the exception of N andK in the latter) when grown at 680 µmol mol^–1 CO₂.In F. vivipara, however, uptake was considerably reduced inplants grown at the higher CO₂ concentration. Overall, a doubling of atmospheric CO₂ concentration had littleeffect on the nutrient use efficiency or productivity of A.capillaris. Reductions in tissue nutrient content resulted fromincreased plant growth and not altered nutrient use efficiency.In P. alpina, potassium, magnesium and calcium productivitieswere significantly reduced and photosynthetic nitrogen and phosphorususe efficiencies were doubled at elevated CO₂ with respect toplants grown at ambient CO₂ F. vivipara grown for 189 d showedthe most marked changes in nutrient use efficiency and nutrientproductivity (on an extracted dry weight basis) when grown atelevated CO₂, F. vivipara grown at elevated CO₂ however, showedlarge increases in the ratio of non-structural carbohydrateto nitrogen content of leaves and reproductive tissues, indicatinga substantial imbalance between the production and utilizationof assimilate. Key words: Nutrient, allocation, nutrient use efficiency, grasses, nutrient productivity, elevated CO₂, cliniate change     

Source--sink Relationships and Carbon Metabolism in Tomato Leaves 1. 14C Assimilate Compartmentation

Effects of the interaction between assimilate availability andsink demand on the metabolism of ¹⁴C assimilates in tomato leaveshave been examined in plants where the source—sink relationshipof assimilates was simplified to one leaf and one fruit truss. During experimentation the source leaf was exposed to either80 or 20 W m^–2 (PAR), while the truss was either retainedor removed. Under these four source-sink conditions, a timecourse study was made on ¹⁴C assimilate distribution in thesource leaf over a period of 23 h after pulse feeding with ¹⁴CO₂. While truss removal caused a temporary increase of ¹⁴C sucrosein leaves under both irradiances, the principal assimilatesaccumulated were starch and hexoses. Decreased ¹⁴C export followingtruss removal was observed within a day in well-illuminatedleaves but after 3 days in leaves under low light. The accumulationof ¹⁴C sucrose at the end of the light period was affected bytruss removal in high light leaves only 3 days later. These observations suggest that while the compartmentation ofnewly fixed assimilate was affected rapidly by the change ofsource—sink relationship, carbon export, as measured by¹⁴C loss, was affected only gradually. The possible effect of sucrose accumulation on photosynthesisis discussed.     

A Reanalysis of Particle Motion in Sieve Tubes of Heracleum

From an analysis of the Brownian motion of particles in sieveelements of Heracleum mantegazzianum and Heracleum sphondylium,Barclay and Johnson have suggested that the in situ viscosityof sieve tube sap is four to six times higher than has previouslybeen assumed. In particular, they obtained a value for the sapviscosity of about 10^–2 Pa s, which compares with a valueof 2 x 10^–3 Pa s for a 20 per cent (w/v) sucrose solution.The present paper describes a reanalysis of their data. It isargued that Barclay & Johnson underestimated the Brownianmotion of sieve element particles and so overestimated the sapviscosity. An exact correction was not possible, but it is concludedthat the in situ viscosity of Heracleum sieve tube sap mustbe less than 3 x 10^–3 Pa s, which corresponds to a sucroseconcentration of less than 29 per cent. Hence it may not beunreasonable to suppose that the viscosity of sieve tube sapis determined primarily by the concentration of sucrose, ashas been assumed in theoretical analyses of the Munch hypothesis.It is also concluded that the sieve tubes studied by Barclayand Johnson were not functional, in the sense that they didnot exhibit an axial bulk flow of sap. Heracleum, sieve tubes, Brownian motion, viscosity, Munch hypothesis     

The Effects of Sulphur Dioxide on Phloem Transport in Two Cereals

Gould, R. P., Minchin, P. E. H. and Young, P. C. 1988. The effectsof sulphur dioxide on phloem transport in two cereals.—J.exp. BoL 39: 997–1007. In vivo investigations using ¹¹C-labelled photosynthate revealedthat there is a change in the patterns of tracer profiles whencereal leaves are exposed to SO₂. The change after exposureto SO₂ was interpreted in terms of a decrease in lateral waterflow into the sieve tubes brought about by reduced phloem loadingalong the length of a leaf. Analysis also revealed that thespeed of translocation was reduced, as expected by the Munchmodel of phloem transport. Key words: Sulphur dioxide, phloem transport, cereal leaves     

Assimilation and Translocation in Perennial Grasses

Single leaves, ears, or shoots of timothy (Phleum pratense L.)were exposed in light to ¹⁴CO₂, then left overnight, after whichthe plants were autoradiographed. The following conclusionswere drawn. Actively growing leaves retain all their assimilatesand import from older ones. Fully expanded leaves export butdo not import assimilates. Export begins before leaf expansionis complete, so import and export may for a time be simultaneous.Exports go at first to younger leaves and to roots, accumulatingat meristems. At later stages, exports move downwards ratherthan upwards. Buds and small tillers import from older shoots,but large tillers do not import from other shoots or exportto other large ones. Ears assimilate while still green, andimport assimilates from their associated flag leaves. Exportsfrom other leaves on flowering stems move downwards. These findings agree in general with those from other plants:they are discussed in relation to the vascular system of thegrass plant, and the need for further studies, particularlyquantitative ones, is emphasized.     

Effects of Sulphur Dioxide and Nitrogen Dioxide on Growth and Translocation in Winter Wheat

Gould, R. P. and Mansfield, T. A. 1988. Effects of sulphur dioxideand nitrogen dioxide on growth and translocation in winter wheat.—J. exp. Bot 39: 389–99 Winter wheat (Triticum aestivum L. cv. Avalon) was grown undersimulated autumn conditions for 4 weeks and exposed to a mixtureof SO₂ and NO₂. Biomass was measured after 2, 3 and 4 weeksand the flag leaves of sample plants were labelled with ¹⁴CO₂.Biomass yields revealed an increase in shoot-to-root ratiosunder polluted conditions. The labelling experiments showedthat less assimilate was transported to the roots, whilst morewas allocated to the younger components of the plant. It appearedthat NO₂ and SO₂ also caused labelled photosynthate to be retainedin the labelled leaf. Reducing the photon flux exacerbated theeffects of SO₂ and NO₂ as indicated by changes in biomass andby the distribution of ¹⁴C. Key words: Wheat, SO₂, NO₂, growth, translocation