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.     

The Interdependence of Tillers in Lolium multiflorum Lam.--a Quantitative Assessment

Experiments were made to determine the extent of reciprocaltransfer of products (derived from the assimilation of ¹⁴CO₂)between various parts of the young vegetative grass plant (Loliummultiflorum Lam.). When individual laminae on different tillerswere supplied with ¹⁴CO₂, 47–64 per cent of the fixedcarbon was exported after 24 h. The principal sinks were theroot system and the shoot or tiller to which the fed leaf wasattached. Other tillers also received significant quantitiesof radiocarbon. When whole tillers were supplied with ¹⁴CO₂the percentage of fixed radiocarbon exported within 24 h rangedfrom 14–31 per cent. Of this, 50–74 per cent wasrecovered from the root system (except in the case of exportfrom the youngest tiller) but exchange of material between tillersalso occurred.A reciprocity diagram is presented and it is concludedthat despite the magnitude of the exchange no tiller showedan over-all net gain or loss. The main shoot and the tillersdiffered in the extent of their carbon exchange and in theirdegree of independence. The oldest daughter tiller of the mainshoot was the most independent and the main shoot most interdependent.     

The Distribution of Assimilates in Lolium multiflorum Lam. Following Differential Defoliation

In seedling plants of Lolium multiflorum Lam. the tillers weredefoliated but the main shoot was left intact. Radiocarbon as¹⁴CO₂ was supplied to this shoot at different times followingtiller defoliation and the pattern of distribution of labelledassimilates was determined quantitatively. It was found thata greater proportion (approximately 10–20 per cent) ofexported assimilate was translocated to the cut tillers butalthough the proportion supplied to the root system was lessthe total radiocarbon incorporated by the roots was unchanged.This was brought about by a large increase in the export ofradiocarbon fixed by the intact shoot—up to 100 per centfollowing one treatment. These alterations in the organizationof the defoliated plant lead to a greater efficiency in thecarbon economy and are discussed in relation to the stress imposedby defoliation.     

Long-term Partitioning, Storage and Re-mobilization of 14C Assimilated by Lolium perenne (cv. Melle)

The youngest fully expanded leaves of single tillers of vegetativeperennial ryegrass plants were exposed to ¹⁴CO₂. Thereafter,quantitative and fractional analysis of the partitioning, storageand re-mobilization after defoliation of the ¹⁴C-labelled assimilatewas sequentially conducted over a 22 d period. In undefoliated plants, most ¹⁴C reached its final destinationwithin 5–6 of feeding. Forty per cent of assimilated ¹⁴Cwas subsequently lost through respiration, while 13.5, 8.5 and34 per cent remained in roots, stem bases and tops respectively.At least some ¹⁴C was distributed to tillers throughout theplant, but secondary tillers subtended by the fed tiller madethe greatest demand on ¹⁴C translocated from the fed tiller. A small, but significant portion of ¹⁴C was invested into longterm storage in undefoliated plants, four per cent of the totalassimilated still being present in a labile chemical form inroots and stem bases 22 d after feeding. In plants that wereseverely defoliated 4 d after feeding, depletion of reserve¹⁴C was observed relative to undefoliated plants. The depletiontook place from stem bases, not roots, and both low and highmolecular weight storage compounds were involved. A portionof the depleted ¹⁴C was incorporated into new growth after defoliation. Lolium perenne, perennial ryegrass, assimilate partitioning, storage, re-mobilization, defoliation     

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.     

Assimilation of 14CO2 by the Inflorescence of Poa annua L. and Lolium perenne L.

The relative assimilatory activity of the inflorescence, itsindividual components, and the leaves of flowering tillers ofPoa annua L. and Lolium perenneL. was determined over the periodfrom inflorescence emergence to seed shedding. The pattern of¹⁴CO₂ fixation was similar for both species and the inflorescencewas by far the most important assimilatory organ of the reproductivetiller, particularly over the latter period of seed developmentas leaf senescence progressed. With the exception of the seedsall parts of the inflorescence showed significant assimilatoryactivity and the lemmas and paleas accounted for 40–50per cent of the total ¹⁴C fixed by the inflorescence in bothspecies. The importance of the grass inflorescence as a photosyntheticstructure is discussed in relation to similar studies on cereals. Poa annua, Lolium perenne, carbon dioxide assimilation, inflorescence     

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     

Some Effects of Yellow Rust (Puccinia striiformis) on 14Carbon Assimilation and Translocation in Wheat

A well-developed infection of Yellow Rust on a leaf of springwheat (Jufy I) caused the assimilation of ¹⁴CO₂ by that leafto decrease to 43.5 per cent of that of an uninfected leaf.Over a period of three hours translocation of ¹⁴C from an infectedleaf was only 0.87 per cent of that from a control leaf. Whencontrol plants were kept in the light for periods up to 16 hoursafter assimilating ¹⁴CO₂ translocation continued at a steadyrate, whereas there was only negligible translocation from infectedleaves after the first few hours. The retention of labelledassimilates in the infected leaf could be partly, but not completely,accounted for by a conversion of assimilates to an alcohol-insolubleform. Rust infection had no effect on the distribution patternof ¹⁴C to other leaves from one which had assimilated ¹⁴CO₂.In contrast to the marked retention of assimilate by an infectedleaf, such a leaf was unable to distort the normal distributionby attracting assimilates from the other leaves.     

Pattern and Control of Distribution of 14C-assimilates in Reproductive Plants of Lolium multiflorum Lam. var. Westerwoldicum

The distribution of ¹⁴C-assimilates was examined in reproductiveplants of Lolium multiflorum Lam. var. Westerwoldicum (cv. Tama)from which all emerged tillers had been removed, leaving themain tiller with two expanding leaves, one of them the flagleaf, and two expanded leaves. Export of ¹⁴C from the lowerexpanded leaf was mainly to the tiller in its axil, the steminternode below its node and the roots, whereas the upper expandedleaf supplied predominantly the expanding leaves, the ear, steminternodes, roots and the tiller bud in the axil of the lowerleaf. Defoliation and root-pruning showed that expanding leaveswere able to compete successfully for assimilates, probablythrough the production of substances capable of mobilizing supply.Local application of 1-naphthaleneacetic acid (NAA), gibberellicacid (GA₃) and 6-benzylaminopurine (BAP) to small tiller budsshowed that GA³ and BAP promoted bud growth and ¹⁴C accumulation,but that addition of NAA reduced these effects.     

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     

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.     

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     

Patterns of Assimilate Distribution and Source--sink Relationships in the Young Reproductive Tomato Plant (Lycopersicon esculentum Mill.)

Patterns of distribution of ¹⁴C were determined in 47-day-oldtomato plants (Lycopersicon esculentum Mill.) 24 h after theapplication of [¹⁴C]sucrose to individual source leaves fromleaves 1–10 (leaf 1 being the first leaf produced abovethe cotyledons). The first inflorescence of these plants wasbetween the ‘buds visible’ and the ‘firstanthesis’ stages of development. The predominant sink organs in these plants were the root system,the stem, the developing first inflorescence and the shoot ‘apex’(all tissues above node 10). The contribution made by individualsource leaves to the assimilate reaching these organs dependedupon the vertical position of the leaf on the main-stem axisand upon its position with respect to the phyllotactic arrangementof the leaves about this axis. The root system received assimilateprincipally from leaf 5 and higher leaves, and the stem apexfrom the four lowest leaves. The developing first inflorescencereceived assimilates mainly from leaves in the two orthostichiesadjacent to the radial position of the inflorescence on thevertical axis of the plant; these included leaves which weremajor contributors of ¹⁴C to the root system (leaves 6 and 8)and to the shoot apex (leaves 1 and 3). This pattern of distributionof assimilate may explain why root-restriction treatments andremoval of young leaves at the shoot apex can reduce the extentof flower bud abortion in the first inflorescence under conditionsof reduced photoassimilate availability. Lycopersicon esculentum Mill, tomato, assimilate distribution, source-sink relationships     

An Examination of the Value of 14C-urea as a Source of 14CO2 for Studies of Assimilate Distribution

Twenty-four hours after leaf 3 of a plant of Lolium multiflorumLam, was supplied with a droplet of ¹⁴C-urea and the plant enclosedin a polyethylene bag with an untreated plant, there were significantamounts of radiocarbon recovered from the untreated plant. Theleaf treated with ¹⁴C-urea was the major source of ¹⁴C leakagebut significant losses were also recorded from other parts ofthe plant. Reducing the humidity within the bag decreased theamount of ¹⁴CO₂ which escaped. Losses of radiocarbon from CO₂-treated plants were very low compared with those from urea-treatedplants but the pattern of assimilate distribution within thetwo types of plants was very similar. The possible causes ofthese effects are considered and the usefulness of ¹⁴C-ureaas a source of ¹⁴CO₂ discussed.     

Aspects of the Comparative Physiology of Ranunculus bulbosus L. and Ranunculus repens L.: II. Carbon Dioxide Assimilation and Distribution of Photosynthates

Detailed analysis of the interrelationships between sourcesof photosynthate production and sites of utilization in thetaxonomically closely related species Ranunculus bulbosus L.and R. repens L. showed that leaves whether present on rosette,stem, or stolon had similar levels of ¹⁴CO₂-fixation but thepattern of distribution of radiocarbon to the rest of the plantdiffered. Fruits of R. bulbosus had a lower fixation rate thanleaves but were characterized by total retention of the fixedradiocarbon. Rosette leaves of R. bulbosus supplied the youngleaves, developing apices in the rosette, roots, and corms,whereas the labelled assimilates from cauline leaves were evenlydistributed between reproductive and vegetative parts. The cormwas the major sink both at the flowering and fruiting stages.When plants were treated with ¹⁴CO₂ in the field even higherlevels of radiocarbon moved into the corm than in comparableexperiments under greenhouse conditions. The rosette leaf ofR. repens exported mainly to actively growing stolons in plantswith many stolons bearing rooted ramets although growth of astolon was also substantially supported by photosynthates producedby its own ramets. A proportion of the radiocarbon fixed byleaves of mature ramets was exported and moved in a predominantlyacropetal direction into the stolon apex, stolon axis, and youngramets of the same stolon. The stock in R. repens had a muchlower demand for assimilates than the corm in R. bulbosus. The results are consistent with the concept that R. bulbosusoperates a conservative policy involving the replacement ofthe parent in situ by a daughter from the corm, coupled withextensive fruit production. In R. repens the emphasis is onlateral spread and exploitation of substantial areas of groundby vegetative spread and replacement of the parent by daughtersmany of which may occupy sites some distance from the parent.     

Respiratory Loss of Recently Assimilated Carbon in Wheat

A series of experiments was undertaken to assess the amountof respiration associated with the growth of wheat at differentstages. Plants (or in some cases just the flag leaf) were labelledwith ¹⁴CO₂ and the amount of ¹⁴CO₂ respired during the subsequent48 or 72 h was measured. The evolution of ¹⁴C, expressed asa percentage of the amount initially assimilated (referred toas the R/A value) was used as a measure of the overall efficiencyof dry matter production. Respiratory ¹⁴CO₂ evolution from labelledplants was most rapid in the first 12 h after labelling, thereafterdeclining rapidly. Evolution was also more rapid following labellingsat the end of the light period (dusk) than at the beginningof it (dawn). The R/A values were greatest (42 and 50 per centrespectively for dawn- and dusk-labelled plants) for young plantsand least (13 and 28 per cent respectively) for plants duringmid grain filling. When flag leaves, as distinct from wholeplants, were labelled, R/A values were lower still (9 and 21per cent respectively), indicating that flag leaf assimilatewas used efficiently in grain production. The calculated minimum R/A for the formation of grain material(10 per cent protein, 90 per cent starch) was 6.2 per cent.That the experimentally determined values were greater thanthis is attributed to the turnover of carbon in enzymes, toother maintenance processes, and possibly to the operation ofthe pentose phosphate pathway of glucose oxidation. R/A valueswere lower in those plants labelled at the beginning than thoseat the end of the photoperiod. This was considered to be a consequenceof refixation of respiratory ¹⁴CO₂ during the light. The higherR/A values found for young plants were considered to be a consequenceof the greater percentage of ¹⁴C translocated to the roots (rootsbeing unable to refix respired CO₂) and of greater turnoverof enzymes associated with more active metabolism. Triticum, wheat, respiration, carbon assimilation, carbon loss, grain-filling     

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     

Effect of Gibberellic Acid on the Distribution of Products of Photosynthesis in Sunflower

Single leaves on growing sunflower plants were allowed to assimilate¹⁴CO₂. Gibberellic acid was applied to the same leaf or to theterminal bud or the roots, and the distribution of assimilated¹⁴C was determined at intervals of 1–96 h. Gibberellicacid had no significant effect on initial distribution of ¹⁴Cduring the period of rapid export from the leaf, but enhancedre-export from the roots after translocation from the leaf hadvirtually ceased. Most of the ¹⁴C exported from the roots accumulatedin the shoot tip. The site of application of the hormone wasof relatively minor importance. Wherever it was applied themajor effect was enhancement of movement from the roots to theshoot tip. Application to the terminal bud was most effectivein this respect. There was no evidence that gibberellic aciddirectly affected the transport system, but the data supportthe hypothesis that it increases the strength of the sink inthe shoot tip. Helianthus annuus L., sunflower, assimilate transport, gibberellic acid, phloem transport     

Post-Anthesis Economy of Carbon in a Cultivar of Cowpea

Budgets for transfer of carbon from individual leaves and othersource organs to fruits and nodulated roots were constructedfor stages of the post-flowering development of symbiotically-dependentcowpea (Vigna unguiculata L. Walp. cv. Vita 3-Rhizobium strainCB756). Exportable surpluses of carbon from sources, assessedfrom net exchanges of CO₂ and changes in carbon content, wereallocated to sink organs in proportion to carbon consumption(growth and respiration) and the ability of each sink organto attract assimilates from the sources, as demonstrated by¹⁴C-feeding. The first 10 d after flowering showed high sinkactivity by roots, stem and petioles, low consumption by fruits,with the upper three trifoliate blossom leaves providing thebulk of the required assimilates. The next 10 d showed a sharpdecline in photosynthesis of the leaf subtending the oldestfruit followed by similar declines in leaves at the other fruitingnodes. All leaflets at fruiting nodes abscised during the final10 d period, while the two lower leaves, not subtending fruits,remained green and supplied most of the carbon required by developingfruits and roots. Throughout fruiting all currently-active sourcessupplied all sinks, with only slight evidence of blossom leavesspecializing in nourishing their subtended fruits. Of the carbontranslocated from leaves during fruiting 32% came from the topmostleaf, 28% from the leaf below this, 16% from the next leaf,and the remaining 24% from the lowest three leaves. Some 80%of the fruit's total intake of carbon came from leaves, therest from mobilization of stored carbon (partly sugars and starch)fromother vegetative parts. Key words: Carbon, Translocation, Cowpea     

Turnover of Carbohydrates in Relation to Growth in Apple Trees. II. Distribution of 14C Assimilates Labelled in Autumn, Spring and Summer

One-year-old plants of the apple rootstocks MM. 106 and M.7were allowed to assimilate ¹⁴CO₂ in autumn, spring and summerand the distribution of the tracer within the plant over a growingseason was followed. In MM.106 distribution of the tracer intwo fractions of extractable carbohydrate and the residues representingstructural material, was determined. The results of the radioactivityassay are discussed in relation to seasonal patterns of assimilatesupply and demand in the different regions of the plant. Malus sylvestris L, apple, ¹⁴C assimilates, distribution of carbohydrates