Cold-Inhibited Phloem Translocation in Sugar Beet: II. CHARACTERIZATION AND LOCALIZATION OF THE SLOW-COOLING RESPONSE

Experiments were undertaken on a simplified sugar beet systemto characterize the phloem translocation response to slow coolingtreatments that were applied to the source leaf petiole. Inthese experiments the temperature was decreased by 4°C every16 min, such that the tissue temperature was lowered from 25°Cto 1°C over a period of 80 min. Our results indicated thatan initial slow cooling treatment, on a given test plant, causedno change in the rate of translocation. However, all subsequentslow cooling regimes that were applied to the same petiole positionelicited a characteristic step-type inhibition. This inhibitionaveraged about 10% of the original translocation rate in allcases with no recovery being observed. The data suggest thatthe initial cooling treatment induced an alteration in the petioletissue which facilitated the inhibition phenomenon during subsequentslow coolings. This alteration was shown to be localized withinthe upstream region of the chilled petiole segment, followingan initial slow cooling, or throughout the chilled petiole segmentafter an initial quick cooling from 25°C to 1°C. Resultsalso show that the alteration is a long-lived phenomenon thathas no detectable influence on the quick-cooling induced transientinhibition of translocation. Key words: Phloem, Translocation, Cooling response, Petiole     

Translocation pathways in the petioles and stem between source and sink leaves of Populus deltoides Bartr. ex Marsh.

Microautoradiography was used to follow the translocation pathways of ¹⁴C-labeled photosynthate from mature source leaves, through the stem, to immature sink leaves three nodes above. Translocation occurred in specific bundles of the midveins and petioles of both the source and sink leaves and in the interjacent internodes. When each of six major veins in the lamina of an exporting leaf was independently spot-fed ¹⁴CO₂, label was exported through specific bundles in the petiole associated with that vein. When the whole lamina of a mature source leaf was fed ¹⁴CO₂, export occurred through all bundles of the lamina, but acropetal export in the stem was confined to bundles serving certain immature sink leaves. Cross-transfer occurred within the stem via phloem bridges. Leaves approaching maturity translocated photosynthate bidirectionally in adjacent subsidiary bundles of the petiole. That is, petiolar bundles serving the lamina apex were exporting unlabeled photosynthate while those serving the lamina base were simultaneously importing labeled photosynthate. The petioles and midveins of maturing leaves were strong sinks for photosynthate, which was diverted from the export front to differentiating structural tissues. The data support the idea of bidirectional transport in adjacent bundles of the petiole and possibly in adjacent sieve tubes within an individual bundle.Abbreviations C central leaf trace - L left leaf trace - LPI leaf plastochron index - R right leaf trace     

Translocation Profiles of [11C] and [13N]-Labelled Metabolites after Assimilation of 11CO2 and [13N]-Labelled Ammonia Gas by Leaves of Helianthus annuus L. and Lupinus albus L.

Tracer amounts of atmospheric [¹³N]-Iabelled ammonia gas, wereabsorbed by leaves of Lupinus albus and Helianthus annuus inboth the light and the dark. Exogenous [¹³N]-ammonia was onlyabsorbed in the dark when the feeding occurred shortly aftera period of illumination and the tissue was not depleted ofits carbohydrate reserves (e.g. starch). Incorporation of the[¹³N]-ammonia appeared to occur via the leaf glutamine synthetase/glutamatesynthase (GS/GOGAT) cycle since 2.0 mol m^–3 MSX, an inhibitorof the GS reduced uptake in both the light and dark. Photosyntheticincorporation of ¹¹CO₂ was not affected by this treatment Therate of movement of [¹³N]-assimilates in the petiole of attachedleaves of Helianthus and Lupinus was similar to that of the¹¹Cl-photo assimilates. Export of both [¹³N] and [¹¹C]-Iabelledassimilates from the leaf and movement in the petiole in boththe light and the dark was inhibited by source leaf anoxia (i.e.nitrogen gas). Translocation was re-established at the samerate when the feed leaf was exposed to gas containing more than2% O₂ which permitted dark respiration to proceed. After aninitial feeding of either ¹¹CO₂ or [¹³N]-ammonia at ambient(21%) O₂ exposure of the source leaf to 2% O2, or 50% O² didnot alter the rates of translocation, indicating that changesin photosynthetic activity in the source leaf due to photorespiratoryactivity need not markedly alter, at least during the shortperiod, the loading and translocation of either [¹¹C ] or [¹³N]-labelledleaf products. Key words: Translocation, CO₂, NH₃, Leaves, Helianthus annuus, Lupinus albus     

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.     

Cold-Inhibited Phloem Translocation in Sugar Beet: IV. ANALYSIS OF THE COOLING-INDUCED REPARTITIONING HYPOTHESIS

Systems identification techniques were used to calculate photo-assimilatetransport characteristics in Beta vulgaris and Pisum sativum,before and after the application of localized perturbationsto the transport path. Changes in photo-assimilate partitioningto various monitored sinks were found to be promoted by slowcool (25 ?C to 1 ?C in 40 min), quick cool (25 ?C to 1 ?C in2 min)/quick warm, and apoplastic osmotic treatments of a localizedregion of the source leaf petiole in Beta. Photo-assimilatepartitioning into both intact and surgically modified (embryo-less)ovules of Pisum was also observed to change following quickcool/quick warm treatments applied to a 2.0 cm region of thepeduncle leading to the monitored pod. No changes in transportproperties were observed through the treated petiole regionof Beta during or following the slow cool treatment (i.e. transittime or system gain). High apoplastic osmolality (addition of1 000 mol m^–3 sorbitol) reduced the transit time of tracermovement through the treated petiole of Beta, while enhancingtracer washout from the bathed tissue region. Our modellingtechniques have shown that the physical or physiological basisfor the sink partitioning changes must be due to alterationsdownstream of the treated pathway zone, suggesting the involvementof physical signals transmitted from the treated region. Theseresults support the previously presented cooling-induced repartitioninghypothesis (Grusak and Lucas, 1986) and demonstrate that pathway-originatedstimuli can alter source-sink photo-assimilate partitioning. Key words: Photo-assimilate partitioning, phloem translocation, Beta vulgaris, Pisum sativum     

Effect of sink region anoxia on translocation rate

Translocation rate, ATP level, and CO₂ production of a developing leaf (sink leaf) were studied in sugar beet (Beta vulgaris) plants prior to and during anaerobic treatment of the sink leaf. Within 3 to 5 minutes after onset of treatment with a N₂ atmosphere, translocation into the sink leaf decreased to near zero and then recovered to a level of about 50% of the control over the next 2 hours. A decline in CO₂ output and ATP levels coincided with the attainment of the new translocation rate. All three quantities returned to near control levels within 60 to 120 minutes after the sink leaf was returned to air. Swelling and ultrastructural changes in mitochondria coincided with the observed ATP level changes during inhibition and recovery periods. The first phase of marked inhibition of translocation did not coincide with low ATP level and appeared to be caused by decreased membrane permeability during the transition to anaerobic metabolism, possibly as a result of a temporary build up of toxic products. The correlation between ATP level and translocation rate suggests that ATP-dependent active transport in the sink leaf augments the driving force for translocation.     

Localized Photosynthate Deposition in Citrus Fruit Segments Relative to Source-Leaf Position

Photosynthate translocation into fruit segments was examinedin ‘Pineapple’ sweet orange Citrus sinensis (L.)Osbeck to determine whether previously reported patterns ofdistribution [Koch (1984) HortScience 19: 260] would changeover time or with alterations in balance between source leavesand sink fruit. In control plants, ¹⁴CO₂ was supplied to a sourceleaf nearest the fruit for 1 h, followed by 5 h translocation.Over 89% of [¹⁴C]assimilates in the fruit were localized in4 segments directly aligned with the source and 73% of thesewere in the center 2 segments. Peel, pulp and seeds showed similarpatterns. Little or no lateral spreading of [¹⁴C]photosynthatesoccurred when an additional 7 days were allowed for translocation,but distribution was slightly broader when the source leaf was8 nodes farther from the fruit. Defoliation and girdling toreduce the source/sink ratio gave variable results if done 18h before experiments, but widened the area receiving [¹⁴C]assimilatesto approximately half the fruit if done 7 days earlier. Thisoccurred only when an entire fruit, was dependent upon a singlesource, leaving the opposite half fruit without an externalsupply of photosynthates. These data show an extreme degreeof preferential translocation and inflexibility which can occurin a transport path. ¹Supported by United States Department of Agriculture CompetitiveResearch Grant 59-2121-1-1-752-0, Regional Project NC-142, andthe Institute of Food and Agricultural Sciences, Universityof Florida. (Received January 12, 1984; Accepted May 14, 1984)     

The influence of plant water deficit on photosynthesis and translocation of 14C-labeled assimilates in cacao seedlings

The effect of plant status on net assimilation and translocation of "C-labeled assimilates in cacao (Theobroma cacao L.) was evaluated. As plant water potential (ψ) decreased from −0.5 to −1.0 MPa, neither net assimilation nor the rate of label translocation out of the l4CO,-fed leaf were affected, but as iji fell between −1.0 and −1.5 MPa, net assimilation decreased sharply and label retention increased greatly. Translocation out of source leaves was strongly correlated with net assimilation (r =−0.93). Translocation velocity, assessed by detection of labeled assimilates in sink leaves, was sensitive to plant water deficit, and it declined linearly (r = 0.97) throughout the range of leaf water potentials observed. The results may be explained by reduction in the velocity of assimilate movement within the sieve elements, reduction in supply of labeled assimilates from source leaves, reduction in sink strength or diversion of assimilates to sites of storage or utilization.     

Changes in apoplastic and intracellular leaf sugars induced by the blocking of export in Vicia faba

Sugar export by broadbean ( Vicia faba L. cv. Aguadulce) was blocked by a cold jacket (1 cm-width, 1°C) applied on the petiole of a mature leaf or by heat-girdling the petiole. A time course study was made on the effects of these treatments on apoplastic and intracellular soluble sugars of the leaf in relation to phloem loading and photosynthesis. Blocking of export by heat-girdling induced an inhibition of phloem loading within 10 min, an accumulation of starch within 30 min and a rise in apoplastic sucrose within 60 min. By contrast, apoplastic hexoses and photosynthesis were not affected by this treatment within 8 h and intracellular sugars were not affected within 2 h. The cold jacket also increased the sucrose content of the apoplast. The increase in apoplastic sucrose induced by the cold barrier is reversed upon rewarming and less marked when the sink/source ratio is increased by defoliating all but the leaves studied. The results are discussed in terms of sink/source relationships. They show that the increase in apoplastic sucrose resulting from inhibition of loading is not part of the events leading from blocking of transport to change in carbon partitioning.     

Real Time Studies of Carbon- 11 Translocation in Moonflower: III. FURTHER EXPERIMENTS ON THE EFFECTS OF A NITROGEN ATMOSPHERE, WATER STRESS, AND CHILLING; AND A QUALITATIVE THEORY OF STEM TRANSLOCATION

Translocation of naturally loaded carbon- 11 is studied in themoonflower Ipomoea alba L. and data are presented to show that:(i) source leaf export is momentarily inhibited by a suddenlyapplied water stress; (ii) pathway translocation appears toproceed by a MÜnch-like mechanism, but in a porous mediumrather than in an array of parallel, only slightly leaky pipes;and (iii) sink unloading seems to proceed normally in a nitrogenatmosphere. A qualitative model which incorporates these findingsis presented.     

Phloem unloading in tobacco sink leaves: insensitivity to anoxia indicates a symplastic pathway

Phloem unloading in transition sink leaves of tobacco (Nicotiana tabacum L.) was analyzed by quantitative autoradiography. Detectable levels of labeled photoassimilates entered sink leaves approx. 1 h after source leaves were provided with ¹⁴CO₂. Samples of tissue were removed from sink leaves when label was first detected and further samples were taken at the end of an experimental phloem-unloading period. The amount of label in veins and in surrounding cells was determined by microdensitometry of autoradiographs using a microspectrophotometer. Photoassimilate unloaded from first-, second-and third-order veins but not from smaller veins. Import termination in individual veins was gradual. Import by the sink leaf was completely inhibited by exposing the sink leaf to anaerobic conditions, by placing the entire plant in the cold, or by steam-girdling the sink-leaf petiole. Phloem unloading was completely inhibited by cold; however, phloem unloading continued when the sink-leaf petiole was steam girdled or when the sink leaf was exposed to a N₂ atmosphere. Compartmental efflux-analysis indicated that only a small percentage of labeled nutrients was present in the free space after unloading from sink-leaf veins in a N₂ atmosphere. The results are consistent with passive symplastic transfer of photoassimilates from phloem to surrounding cells.Symbol VI radio of ¹⁴C in veins and interveinal tissue     

14C-Translocation in Kalanchoe pinnata at two Different Stages of Development

Mayoral, M. L. and Medina, E. 1985. ¹⁴C-translocation in Kalanchoepinnata at two different stages of development.—J. exp.Bot. 36: 1405–1413 Translocation of ¹⁴C-compounds from mature leaves was measuredin plants of Kalanchoe pinnata to determine the interactionbetween plant age and CAM phase when CO² is taken up. Matureleaves of 4 and 12 month old plants were fed with ¹⁴CO² eitherduring CAM phase 1 (midnight) or at the beginning of CAM phase4 (early afternoon). Export of ¹⁴C activity from source leaves,and distribution of ¹⁴C activity in soluble and insoluble compoundswas measured both in source leaves and sink organs. In 4 monthold plants 4 d were needed to export 76% of total ¹⁴C activityincorporated during CAM phase 1, while leaves labelled at thebeginning of CAM phase 4 exported 44% of total ¹⁴C activityafter 4 h, and 80% after 24 h. In both cases the major fractionof total radioactivity translocated was found in the roots inthe form of neutral sugars. Differences in translocation patternsare due to distribution of ¹⁴C in the source leaves, 96 % of¹⁴C taken up during CAM phase 1 is found in the insoluble fractionat the end of the subsequent phase 3, while 93 % of total radioactivitytaken up at the beginning of phase 4 is found in the solublefraction at the end of this phase. In 12 month old plants labelledduring phase 1 very little translocation could be detected atthe end of phase 3, while only 20% of total radioactivity wastranslocated from leaves labelled during phase 4 and measured4 h later. ¹⁴C activity in the older leaves had a similar distributionin soluble and insoluble fractions as the one determined inthe younger plants. Ability to translocate carbon compoundsfrom source leaves during phase 3 was shown by loading matureleaves at dawn with ¹⁴C-sucrose. Here again, mature leaves ofyounger plants showed faster translocation of radioactivitythan those of older plants Key words: Kalanchoe, crassulacean acid metabolism, translocation, sink, source relationships     

Translocation of Assimilates Within and Between Potato Stems

Three aspects of translocation in potato were examined: (i)translocation within stems (ii) translocation between individualstems of a plant (iii) translocation between tubers followinginjection of ¹⁴C sucrose into a single daughter tuber. Assimilatesexported from single leaves of evenly illuminated potato stemsremained confined to the same side of the stem as the sourceleaf in a pattern consistent with the internal arrangement ofvascular bundles in the stem, and tubers borne on stolons verticallybelow the source leaf contained higher concentrations of ¹⁴Cthan those on the opposite side. Consequently ¹⁴C import intothe tubers bore little relationship to tuber growth rates. However,alteration of source/sink relations by pruning stems to a singlesouce leaf resulted in an even distribution of ¹⁴C throughoutthe vascular bundles of the stem and ¹⁴C import into the tubersbore a stronger relationship to tuber growth rates than to thephyllotactic relationship of the tubers with the source leaf. Labelling one stem of a potato plant resulted in little or nomovement of ¹⁴C into tubers on other unlabelled stems. However,removal of the unlabelled stems at ground level induced a significantmovement of ¹⁴C from the labelled stem to the tubers on unlabelledstems, this movement occurring via the mother tuber. Shadingthe unlabelled stems had less effect than stem removal. ¹⁴C sucrose injected into single daughter tubers was translocatedto other tubers on the same stem and also to tubers on a secondstem at the opposite end of the mother tuber. The sucrose wasconverted to starch in these tubers. The results favour the view that each potato stem functionsas an independent unit with potential for assimilate redistributionwithin a stem but with little or no carbon exchange occurringbetween stems, unless under severely altered source/sink patterns. Assimilates, ¹⁴C, autoradiography, potato (Solanum tuberosum L.), tuber growth     

Sink Activity Estimation by Sink Size and Dry Matter Increase During the Ripening Stage of Barley (Hordeum vu/gare) and Rice (Oryza sativa)

During the ripening stage of barley and rice, the sink activitywas defined as the dry matter increase per units sink size,leaf area and time, as follows: NAR = A.SinkW+NAR', where NAR is the net assimilation rate (g d.wt dm^–2d^–1);A is the sink activity [g d.wt g^–1d.wt (ear) dm^–2d^–1]; Sink W is ear wt per plant at heading (g d.wt);and NAR' is net assimilation rate excluding the assimilate ofsink organ (g d.wt dm^–2 d^–1). Plant material with 16 combinations of mutually different sink(ear) and source (leaf) size were produced at heading for eachcrop: parts of each leaf and ear were removed to produce fourgrades in barley, and also a part of each leaf was removed producingfour grades for four rice varieties showing different ear size.NAR and NAR' were determined during 26 and 21 d in barley andrice after heading, respectively. Sink activity (A), representedas the assimilation rate induced by the sink organ, was estimatedfrom the relationship between SinkW and NAR using the previousequation. The sink activity was significantly higher in ricewith a value of 0–0.028 g d.wt g^–1 d.wt (ear) dm^–2d^–1 vs. 0–0.0017 in barley, suggesting that therelative role of leaves for grain filling is considerably higherin rice than in barley. The sink activity obtained in the studymight be introduced into a model to predict the yields of barleyand rice. Hordeum vulgare L, barley, Oryza saliva L, rice, dry matter, NAR, sink, source, sink activity, model     

Long distance translocation of sucrose, serine, leucine, lysine, and carbon dioxide assimilates: I. Soybean

To determine the selectivity of movement of amino acids from source leaves to sink tissues in soybeans (Glycine max [L.] Merr. `Wells'), ¹⁴C-labeled serine, leucine, or lysine was applied to an abraded spot on a fully expanded trifoliolate leaflet, and an immature sink leaf three nodes above was monitored with a GM tube for arrival of radioactivity. Comparisons were made with ¹⁴C-sucrose and ¹⁴CO₂ assimilates. Radioactivity was detected in the sink leaf for all compounds applied to the source leaflet. A heat girdle at the source leaf petiole essentially blocked movement of applied compounds, suggesting phloem transport. Transport velocities were similar (ranged from 0.75 to 1.06 cm/min), but mass transfer rates for sucrose were much higher than those for amino acids. Hence, the quantity of amino acids entering the phloem was much smaller than that of sucrose. Extraction of source, path, and sink tissues at the conclusion of the experiments revealed that 80 to 90% of the radioactivity remained in the source leaflet. Serine was partially metabolized in the transport path, whereas lysine and leucine were not. Although serine is found in greater quantities than leucine and lysine in the source leaf and path of soybeans, applied leucine and lysine were transported at comparable velocities and in only slightly lower quantities than was applied serine. Thus, no selective barrier against entry of these amino acids into the phloem exists.     

Time course of low temperature inhibition of sucrose translocation in sugar beets

Further studies are presented characterizing the time-course response of sucrose translocation in sugar beet (Beta vulgaris L. cv Klein Wanzleben) to low temperature inhibition. Only the temperature of a 2 cm zone of the source-leaf petiole was varied (1° vs 25°, approximately). The half-time of inhibition, defined as the time required for 50% inhibition of the control or pre-cooling rate, varied from 4 to 15 minutes, and the half-time of recovery from 30 to 100 minutes. Maximum inhibition varied from 68 to 92%. Possible uncertainties in evaluating these parameters are discussed. When the duration of the low temperature period was sufficient to permit essentially full recovery, subsequent re-warming of the petiole zone to 25° to 30° effected little or no increase in the translocation rate. It is evident that the interposition between source and sink of a 2 cm petiole zone maintained at a temperature generally inhibitory to physiological processes resulted in little or no impairment to the translocation process, after a suitable thermal adaptation period. Thermally adapted petiole systems de-adapted after periods as short as 1 hour at 25°.     

Effect of light and ontogenetic stage on sink strength in bean leaves

Light (about 3,000 foot-candles) neither increased nor decreased the sink strength of young, rapidly expanding leaves of Phaseolus vulgaris L. cv. Black Valentine, as measured by the comparative rates of import of ¹⁴C-labeled photosynthates by sink leaves in the light versus dark in short term experiments. Although irradiated sink leaves accumulated more ¹⁴C activity, the difference was fully accounted for by photosynthetic reabsorption of respiratory CO₂ derived from substrates translocated to the sink leaves.

Maximum sink strength was attained when the sink leaf reached 7 to 8 cm² in area (9 to 10% of its fully expanded size). Thereafter sink strength declined rapidly and asymptotically to a near zero value at about 45% final area. During this period, however, the rapid decline in translocation was offset by a rapid rise in the photosynthetic rate of the sink leaf, maintaining a near constant relative rate of dry weight increase until the sink leaf had expanded to about 17% of its final area. Although the increasing photosynthetic capacity was associated with a decreasing import capacity, suggesting that the rate of translocation to the sink leaf was controlled by the developing capacity of the sink leaf for photosynthesis, it was not possible to vary the total (true) translocation rate to the sink leaf by varying the photosynthetic rate of the sink leaf in short term light-dark experiments. Despite a high ratio of source to sink in these experiments, no evidence accrued that translocation into young bean leaves was ever sink-limited.

     

Time-dependent Behavior of a Mathematical Model for Munch Translocation: Application to Recovery from Cold Inhibition

The time evolution of a Munch pressure-flow translocation system is calculated using a numerical computer method. Results are obtained for the time course of the system variables following application of a large resistance in the translocation path, intended to simulate a cold block. The resistance factor required to produce translocation inhibition indicates that even moderate inhibition is primarily due to sieve plate pore block-age, rather than to the viscosity increase. The calculated time for recovery from cold inhibition and the shape of the translocation recovery curve agree with experimental results. The time for translocation recovery and the level of velocity recovery depend on the rate of sucrose unloading in the sink; on the sucrose concentration in the sieve tube; on the position, length, and resistance factor of the cold block; and on the hydraulic conductivities.     

RADIOAUTOGRAPHIC STUDY OF TRANSLOCATION IN BEAN LEAVES

The movement of the radioactivity from sucrose, indole-3-acetic acid (IAA) and phosphate has been examined in excised bean leaves. Preferential translocation of the labelled materials toward the base of leaf and petiole was demonstrated, suggesting a natural mobilization gradient down the leaf and petiole. Establishment of other mobilization centers in the leaf by local application of N⁶-benzyladenine diverted the movement of the sucrose label and, to a lesser extent, the phosphate label. There was no apparent mobilization of IAA by benzyladenine. Evidence is provided that there is a continuity of label from the source to the sink regions, and it is suggested that reported instances of noncontinuity of label may be attributable to the refixation of respired C¹⁴O₂ by tissue treated with benzyladenine. The observations appear to substantiate the concept that the unloading of solutes from the phloem can regulate the direction and intensity of translocation.     

Alterations in source-sink patterns by modifications of source strength

Bean plants, trimmed to a simplified “double source, double sink” translocation system (the paired primary leaves serving as the double source and the paired lateral leaflets of the immature first trifoliate leaf as the double sink) were used to study the magnitude and short-term time course of change in the allocation ratio (partition ratio) of assimilates translocated from the labeled primary leaf to its respective “near” and “far leaflet” sinks in response to an increase or decrease in the source strength of the opposite primary leaf (the “control” leaf). If the rates of net photosynthesis in the two primary leaves were similar, assimilates from the labeled source leaf partitioned to the leaflet sinks in the ratio of 5:1 or higher, the dominant sink being the leaflet “nearer” to the labeled source leaf. If the rate of net photosynthesis in the control leaf was increased substantially above that of the labeled source leaf, the rate of translocation from the labeled source to either the near leaflet sink or far leaflet sink remained unaffected, despite, presumably, a higher translocation rate from the control leaf, and hence a higher phloem pressure gradient (or increased cross-sectional area) in the transport pathway from the control leaf to the leaflet sinks. If the control leaf was excised, thus reducing the source leaf area by about a half, the translocation rate from the remaining source leaf rapidly doubled, the partition ratio becoming equal to unity. If the control leaf was darkened, the partition ratio adjusted to an intermediate value. Although export rates from the labeled source leaf were increased either by excising or darkening the control leaf, the rate of net photosynthesis in the labeled leaf remained constant.