Distribution of 14C-labelled Products of Photosynthesis during the Commercial Life of the Tomato Crop

On seven occasions measured amounts of ¹⁴CO₂ were supplied toindividual leaves on tomato plants grown as a crop under commercialconditions. Twenty-four hours later the distribution of radiocarbonproducts was determined using counting procedures. In the vegetativephase lower leaves exported more carbon up than down, and upperleaves a larger proportion downwards. During the developmentof the crop upper leaves continued to export more carbon downthan up, and lower leaves developed a similar tendency. In theyoung fruiting crop all leaves supplied all the trusses, butas the number of trusses increased groups of leaves came tohave primary (but not absolute) responsibility for supplyingsingle trusses. At the time of fruit development the leavesretained for 20–24 hours more than 80 per cent of thecarbon they fixed. Mature leaves and stems both above and belowthe test leaves were significant sinks for ¹⁴C products.     

Changing Patterns of Distribution of the Products of Photosynthesis in the Tomato Plant with Respect to Time and to the Age of a Leaf

Two series of experiments were conducted with tomato plantsgrown in a glasshouse. In the first series the second leaf ofa young plant was exposed to ¹⁴CO₂ for periods up to a maximumof 4 h and the distribution of the ¹⁴C-products was determinedeither at intervals of 10 and 30 min, 1, 2, 3, 4, 5, 6, 12,and 24 h, or 1, 2, 3, 5, 7, 11, and 15 days after the ¹⁴CO₂was initially supplied. Seventeen per cent of the containedradioactive carbon was exported during the first day and a further23 per cent during the following week. At the death of the leaf44 per cent of the carbon originally fixed was present in thedead structure. There is evidence that the roots re-export alarge proportion of the carbon products originally imported. In the second series the tenth leaf of a plant was suppliedwith ¹⁴CO₂, and 24 h later the relative amounts of radioactivitywere determined for every leaf, internode, and fruit truss andfor the roots and shoot apex. The plants were selected so thatleaf 10 was always immediately below truss 1. The experimentwas performed seven times, each time at a different stage bothin the life-cycle of the leaf and the development of the plant.There were differences in the total amounts of radioactivityfound at the end of each experiment but these differences werenot random. The pattern of distribution of the ¹⁴C exportedby leaf 10 changed as the plant developed. The percentage ofthe ¹⁴C initially fixed which was exported by leaf 10 rose toa peak early in its life and then fell sharply but the distributionof ¹⁴C exported followed a different pattern. For a long periodthere was a heavy accumulation of ¹⁴C in the internode immediatelybelow leaf 10 but later in development this was less evidentand the proportions reaching the roots increased. The internodeabove leaf 10 never showed the same degree of accumulation.Truss I dominated the distribution patterns during the phaseof its active growth. These results are discussed and a hypothesisto account for the changing patterns of distribution is presented.     

Photosynthesis and Diffusion Conductance of the Valencia Orange Fruit under Field Conditions

CO₂ uptake and diffusion conductance of Valencia orange fruits(Citrus sinensis L. Osbeck) were measured in the field duringthe growing season of 1977/78 to ascertain if, as in the leaf,stomata control photosynthesis and transpiration under changingenvironmental conditions. Measurements were made on 15 yearold trees grown in a sandy loam soil and receiving either adry or a wet treatment. Fruit diffusive conductance was measuredwith a modified water vapour diffusion conductance meter andgross photosynthesis was measured with a ¹⁴CO₂ uptake meter.Photosynthetically active radiation (PAR) was measured witha quantum sensor. Fruits exposed to light assimilated CO₂ ata rate which was 25–50% of that assimilated by leaves.The uptake was dependent on fruit size, PAR, chlorophyll content,and on diffusive conductance of the fruit epidermis. Epidermalconductance showed a diurnal trend which was similar in shapeto that of the leaf except in the late afternoon. Cuticularconductance of the fruit was calculated and ranged between 0.22and 0.30 mm s^–1. It was speculated that the CO₂ uptakeby the fruit could support the growth of flavedo cell layerswhen exposed to light. Dry soil caused an increase in the ¹⁴CO₂uptake by fruit possibly caused by the increased potential areaof the stomatal opening per unit of fruit surface area.     

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.     

Effects of Leaf Infections by Septoria nodorum Berk. on the Translocation of 14C-labelled Assimilates in Spring Wheat

The influence of infection with Septoria nodorum of leaves belowthe flag leaf on the translocation of ¹⁴C-labelled assimilatesin wheat was followed. In the vegetative phase export of assimilatesfrom a single infected leaf was reduced, but export from a healthyleaf on a heavily infected plant was increased. During the reproductivephase export from leaves was not affected by disease. Heavyleaf infection had little effect on the patterns of distributionof export especially during reproductive growth when only changesin the proportion of assimilates in leaf sheaths and tillerstumps were found. Distribution of export from a healthy flagleaf on an otherwise heavily infected plant was unaltered. Duringvegetative growth changes in the distribution of assimilateswere more marked, the greatest changes occurring when a singleinfected leaf on a healthy plant was exposed to ¹⁴CO₂.     

Root Development and Source-Sink Relations in Carrot, Daucus carota L: II. EFFECTS OF ROOT PRUNING ON CARBON ASSIMILATION AND THE PARTITIONING OF ASSIMILATES

Physiological responses to root pruning were investigated bycomparing ¹⁴CO₂ fixation rates, the partitioning of ¹⁴C-labelledassimilate, and soluble and insoluble carbohydrate levels inthe leaves of carrot plants following the removal of some ofthe fibrous roots, or fibrous roots and part of the tap root.Root pruning reduced ¹⁴CO₂ fixation by 28–45% but leafspecific activity (¹⁴C assimilation g-¹ leaf fresh weight) wasunchanged. The proportion of total assimilate exported to theroot system increased following root pruning and this was atthe expense of the developing leaves. In younger plants (wherethe tap root received 10% of the assimilate) the supply of ¹⁴Cto the tap root was maintained in spite of root pruning. However,shortening the tap root to 3 cm in older plants (in which 30%of the fixed 14C was normally exported to the developing storageorgan), reduced its sink capacity and resulted in slightly greaterretention of ¹⁴C in the mature leaves. Greater concentrationsof insoluble carbohydrate were found in the mature leaves followingroot pruning but soluble sugar content was unaffected. Onlysmall differences were observed in the distribution of ¹⁴C betweensoluble and insoluble carbohydrate fractions when plants werefed ¹⁴CO₂ several days after the root pruning operations. Thesephysiological responses were mainly associated with the removalof fibrous roots and support the view that the fibrous rootsystem is more important than the developing storage organ inregulating growth in young carrot plants.     

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     

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.     

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 Translocation of 14C-Photoassimilate from Normal and Mutant Leaves to the Pods of Pisum sativum L.

In experiments using radioactive carbon dioxide (¹⁴CO₂) a comparisonwas made of the ¹⁴C-photoassimilate translocation potentialsof two normal leaved (genotype AfAfTlTl) and two mutant formsof Pisum sativum (pea). A ¹⁴CO₂ administration method is describedthat permitted ¹⁴C-translocation studies to be conducted underfield conditions. One of the mutants available produced tendrils in place of leaves(afafTlTl). The other mutant studied was without tendrils buthad a much branched petiole with numerous relatively minuteleaflets (afaftltl). These mutants and the normal-leaved cultivarswith which they were compared were not isogenic lines. Lengthybackcrossing would be required before full assessment couldbe made of the possible agronomic value of such mutations. An interim evaluation of these mutants was based on ¹⁴C-distributionassays that were conducted 48 h after feeding ¹⁴CO₂, to specifiedleaves. The indication was that in translocation terms the leafand pod had a well defined respective source and sink relationshipthat was independent of leaf morphology. In each case the podswhich constituted the major ¹⁴C sinks depended on which leafhad been fed ¹⁴CO₂. With regard to sink specific activity asdefined by the quantity of ¹⁴C incorporated per unit dry weightof pod, the mutants were not significantly different from normal. The implication of these findings was that fundamental changesin pea leaf morphology could be made genetically without a markedeffect on the photoassimilate export potential of the leaf.     

Glycolate Pathway and Serine Synthesis in Relation to CO2 Concentration in Tomato Leaves

Isotopic trapping of the carbon flowing through the glycolatepathway by exogenous glycolate, glycine and L-serine was investigatedduring ¹⁴CO₂ photosynthesis at different CO₂ concentrationsin tomato leaves. L-Serine markedly trapped the carbon flowingfrom ¹⁴CO₂. The amounts of ¹⁴C incorporated into serine decreasedat a high CO₂ concentration, but increased with an increasein the CO₂ concentration in the presence of exogenous serineduring 10-min photosynthesis in ¹⁴CO₂. When ¹⁴CO₂ was fed for5 to 40 sec at 1300 ppm CO₂ to tomato leaves which had beengiven L-serine, an increase in the accumulation of ¹⁴C-serinebegan after 20 sec, and the ¹⁴C-serine molecules formed at 20and 40 sec were labeled uniformly. In the presence of exogenousserine during 10-min photosynthesis in 1300 ppm CO₂, isonicotinicacid hydrazide increased the incorporation of ¹⁴CO₂ into glycinewith a corresponding decrease in the accumulation of ¹⁴C-serine,but it did not inhibit serine accumulation completely; an evidencefor that some serine was formed by a pathway other than theglycolate pathway. The effect of the CO₂ concentration on theglycolate pathway is discussed in terms of serine synthesisin the presence of exogenous serine. (Received June 1, 1981; Accepted September 30, 1981)     

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.     

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     

Effect of leaf age on light and dark 14CO2 fixation in a CAM plant, Bryophyllum calycinum

Leaves of different ages from B. calycinum were exposed to ¹⁴CO₂in light during day and night. The labelling pattern on thechromatogram differed with leaf age. Young leaves had similarpatterns to those of C3 plants during both day and night. Matureleaves showed high incorporation of ¹⁴C into C4 acids, especiallyat night. In contrast, no significant difference with leaf agewas observed in the pattern of dark ¹⁴CO₂ fixation products.Study of the enzyme activity and the content of titratable acidat each leaf age suggested that high incorporation of ¹⁴C inC4 acids during the night was due to the simultaneous absorptionof CO₂ by both enzymes RuDPcarboxylase and PEPcarboxylase. (Received November 24, 1977; )     

The Strategy of Carbon Utilization in Uniculm Barley I. THE CHEMICAL FATE OF PHOTOSYNTHETICALLY ASSIMILATED 14C

Following exposure of the youngest mature leaf of uniculm barleyto ¹⁴CO₂, groups of plants were harvested over a 72 h periodto determine the fate of 14C in the photosynthesizing leaf andin growing leaves and roots. Initially, ¹⁴C was mainly presentin sucrose with a little in starch and charged compounds; transportout of the fed leaf was rapid and, by 7 and 24 h, 56 and 93%respectively of the ¹⁴C had been translocated about equallyto growing leaves and roots. Sucrose entering meristems wasquickly metabolized to protein and structural carbohydrate (40and 60% of the ¹⁴C in these organs at 7 and 24 h respectively),while the remainder was converted to short-term storage productsor intermediary metabolites. By the end of the first day c.35% of the 14C originally assimilated had been lost in respiration. The metabolism of the leaf appeared to be organized on a diurnalbasis, for it exported nearly all its carbon within 24 h ofassimilation. In contrast, some of the assimilate imported intogrowing leaves and, to a lesser extent, roots was not immediatelyused for growth but persisted as temporary metabolites and wassubsequently used for growth in the following days.     

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     

Pre-illumination as a Factor in the Dark Fixation of Carbon Dioxide by Leaf Disks of Kalanchoe blossfeldiana var. Feuer Blute

Dark fixation of CO₂ by leaf disks or whole leaves taken fromplants of variety ‘Feuer Blute’ was measured using¹⁴CO₂. Results indicate that dark fixation by leaf disks isindependent of photoperiodic induction of the plant, but isquantitatively related to the amount of light, over a fairlywide range, to which the leaf is exposed in the single precedinglight period.     

The Influence of Leaf Development on the Expression of C4 Metabolism in Flaveria trinervia, a C4 Dicot

The initial products of ¹⁴CO₂ assimilation were determined understeady state illumination of leaves of Flaveria trinervia, aC₄ dicot of the NADP-mialic enzyme subgroup. Leaf age influencedthe partitioning of ¹⁴CO₂ between the C₄ cycle and the reductivepentose phosphate (RPP) pathway. An estimated 10 to 12%of theCO₂ entered the RPP pathway directly in leaves about 20% fullyexpanded, whereas CO₂ was apparently fixed entirely throughthe C₄ pathway in leaves 75% or more expanded. This partitioningpattern was attributed to the bundle sheath compartment in youngleaves having a relatively high conductance to CO₂ (i.e., beingsomewhat leaky). Of the initially labelled C₄ acids, the proportion that wasmalate, relative to aspartate, increased continuously duringleaf expansion (from 60 : 40 to 87 : 13 at full expansion).Concurrently, there was an increase in the whole leaf activityof NADP malate dehydrogenase and a decrease in the activitiesof aspartate and alanine aminotransferases. Low chlorophylla/b values were observed in young leaves, which may coincidewith an enhanced capacity for non-cyclic electron transportin the bundle sheath chloroplasts of such tissue. Both enhancedaspartate metabolism and direct fixation of CO₂ in the bundlesheath could provide a greater sink for utilization of photochemicallyderived NADPH in the bundle sheath of young leaves. Such metabolicchanges are discussed in relation to a possible decrease inCO₂ conductance of the bundle sheath during leaf development. (Received March 4, 1986; Accepted June 25, 1986)     

The Flux of 14C-Labelled Photosynthate through Soyabean Root Nodules during N2 Fixation

Experiments are described which examine the flux of photosyntheticassimilates from leaves to nodules of soyabean during N₂ fixation.The first part, where the respiratory efflux of ¹⁴CO₂ by noduleswas used as a means of assessing the import of labelled photosynthatefrom leaves, shows that most ¹⁴CO₂ loss from nodulated rootsis due to the metabolic activity of nodules. Much less photosynthatewas imported by nodules if the metabolic activity associatedwith N₂ fixation was inhibited by low O₂ concentration. The second part describes the chemical fate of current photosynthateas it is utilized by nodules. Labelled material was detectedin nodules within c.15 min of supplying ¹⁴CO₂ to the leaf. Thisrose to a maximum at c.70 min before declining by 85% withinthe following 4 h. Most (80%) ¹⁴carbon imported by nodules waseither lost as respiratory ¹⁴CO₂ or re-exported as productsof N₂ fixation. Ten per cent of imported carbon was found asstructural material and 10% as starch. Of the ¹⁴C soluble in ethanol, most was found in the neutralfraction (80% declining to 50% as sucrose) with smaller amountsas amino acids, organic acids (each category rising from 10%to 20%) and phosphate esters (<5%). Comparison of the distribution of ¹⁴C among amino acids, amidesand ureides in the nodules with that of xylem exudates indicatedthat selected compounds were exported from nodules. The ¹⁴Cdata indicate that c.80% of the nitrogen exported from noduleswas in the form of ureides (mainly allantoic acid) and only10–12% as amides. Key words: Nodules, ¹⁴C-photosynthate, Respiration, Carbon flux     

The Regulation of Carbon Transport and the Carbon Balance of Mature Tomato Leaves

Rates of carbon transport from a single mature tomato leaf inthe light period (day transport) and the dark period (nighttransport) were estimated from the rate of carbon fixation inthe light period, the rate of respiration in the dark periodand the changes in carbon contents over these two periods. Plantswere grown initially at 40 W m^–2 light intensity witheither 350 vpm (nonenriched plants) or 1000 vpm CO₂ (enrichedplants). Various light flux densities or CO₂ concentrationswere then applied to the experimental leaves in the light periodduring the experiment When leaves were temporarily exposed to contrasting light fluxdensities both day transport and night transport were linearlyrelated to the rate of carbon fixation. If leaves were shadedbelow the light compensation point for up to five days, or transferredto contrasting CO₂ concentrations for up to ten days, the linearrelationship between carbon fixation and carbon transport nolonger held. During acclimatization, therate of wbon fixationwas simply related to thecurrent light flux density and CO₂concentration, but the rate of carbon transport changed withtime. Day and night transports responded differently to changesin environment: night transport was more related to the contentof reserve, particularly starch, than to the rate of concurrentwbon fixation. It is concluded that the rate of carbon transport of a maturetomato leaf in a single photoperiod is regulated not merelyby the rate of concurrent carbon fixation but by the contentof reserve in the leaf. The latter results from previous cumulativewbon fixation and carbon transport. As a result of changingthe rate of carbon transport, a balance of carbon input andoutput was achieved within 10 days of acclimatization in a maturetomato leaf.