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.     

Effects of plant water deficit on the daily carbon balance of leaves of cacao seedlings

The daily carbon balance of individual source leaves of Theobroma cacao L. seedlings was measured at 2- to 3-day intervals during a 19-day period of increasing plant water deficit and during an 8-day period of recovery following rewatering. In each case, responses of stressed seedlings were compared to those of irrigated controls. Leaves of irrigated cacao seedlings assimilated approximately 41 mg carbohydrate dm^-2 during 12-h photoperiods, and exported an average of 34 mg carbohydrate dm^-2 during 24-h measurement cycles. The rate of carbon export from cacao leaves was sharply reduced as leaf water potential (ψ) declined between -0.8 and -2.0 MPa. Further, the rate of export was closely associated with the net assimilation rate (A), with export capacity being severely reduced as A fell to near zero. Net accumulation of dry matter occurred as long as A remained greater than approximately 20 mg carbohydrate dm^-2 over the 12-h photoperiod, but at lower assimilation rates, export exceeded concomitant assimilation. Carbon export continued at the expense of leaf carbon reserves as photoassimilation fell to near zero during periods of severe water stress (ψ < -2.0 MPa). Night respiration rate was independent of plant water status.     

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     

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     

Implications of water stress-induced changes in the levels of endogenous ascorbic acid and hydrogen peroxide in Vigna seedlings

Vigna cutjang Endl. cv. Pusa Barsati seedlings, subjected to increasing degrees of water stress (−0.5, −1.0, −1,5 MPa), produced an approximately proportional increase in glycolate oxidase activity, hydrogen peroxide (H₂O₂) and proline content but a decrease in catalase activity, ascorbic acid and protein content. Leaf water potential (leaf ψ) and relative water content (RWC) were also lowered with increasing stress. Pretreatment with l -cysteine and reduced glutathione (10-3 M) decreased glycolate oxidase activity, H₂O₂ content, ascorbic acid oxidase activity, proline content and also slightly improved the water status of leaves stressed (−1.0 MPa) for 2 days. Pretreatment of non-stressed seedlings with these antioxidants had little or no effect. These studies indicate that treatment with antioxidants makes the plant tolerant against water stress by modulating the endogenous levels of H₂O₂ and ascorbic acid in stressed tissue.     

A genetic basis for the manipulation of sink–source relationships by the galling aphid <Emphasis Type="Italic">Pemphigus batae</Emphasis>

We examined how the galling aphid Pemphigus batae manipulates resource translocation patterns of resistant and susceptible narrowleaf cottonwood Populus angustifolia. Using carbon-14 (¹⁴C)-labeling experiments in common garden trials, five patterns emerged. First, although aphid galls on resistant and susceptible genotypes did not differ in their capacity to intercept assimilates exported from the leaf they occupied, aphids sequestered 5.8-fold more assimilates from surrounding leaves on susceptible tree genotypes compared to resistant genotypes. Second, gall sinks on the same side of a shoot as a labeled leaf were 3.4-fold stronger than gall sinks on the opposite side of a shoot, which agrees with patterns of vascular connections among leaves of the same shoot (orthostichy). Third, plant genetic-based traits accounted for 26% of the variation in sink strength of gall sinks and 41% of the variation in sink strength of a plant’s own bud sinks. Fourth, tree susceptibility to aphid gall formation accounted for 63% of the variation in ¹⁴C import, suggesting strong genetic control of sink–source relationships. Fifth, competition between two galls was observed on a susceptible but not a resistant tree. On the susceptible tree distal aphids intercepted 1.5-fold more ¹⁴C from the occupied leaf than did basal aphids, but basal aphids compensated for the presence of a distal competitor by almost doubling import to the gall from surrounding leaves. These findings and others, aimed at identifying candidate genes for resistance, argue the importance of including plant genetics in future studies of the manipulation of translocation patterns by phytophageous insects.     

Leaf expansion of four sunflower (Helianthus annuus L) cultivars in relation to water deficits. I. Patterns during plant development

Abstract. The influence of a slow stress and recovery cycle on the pattern of leaf expansion in four diverse sunflower cultivars ( Helianthus annuus L. cvs. Hysun 31, Havasupai, Hopi and Seneca) was studied in a glasshouse. Stress had no significant effect on the time of flower bud emergence and anthesis, or on final leaf number, but delayed the appearance of leaves at high insertions in all cultivars except Hysun 31.
Leaf expansion was markedly reduced as the predawn leaf water potential decreased from −0.35 to −0.60 MPa, and the predawn turgor pressure decreased from 0.3 to 0.2 MPa, and expansion ceased at a predawn leaf water potential of about −1.0 MPa, i.e. when the predawn turgor pressure reached zero.
The leaves most reduced in final size when water was withheld were those at the insertions which grew the most rapidly in unstressed plants. The maximum reduction in final leaf size of 25–35% was similar in all cultivars and was due to retardation of the rate of leaf expansion: the duration of leaf expansion was actually increased by stress. However, leaves that were initiated during stress, but emerged after rewatering, had final leaf areas at least equal to those in the unstressed plants: in the cultivar Seneca, the final size of leaves of high insertion was significantly greater in stressed than unstressed plants, whereas in the three other cultivars the final leaf sizes were similar in both treatments. All four cultivars examined adjusted osmotically to the same degree, but leaf water potentials in one, Seneca, increased more rapidly after rewatering than in the other three, and this may have contributed to the greater relative leaf size in the leaves of high insertion in this cultivar.     

Translocation of photoassimilate from leaves of two popyploid genotypes of tall fescue differing in photosynthetic rates

The photosynthetic rate of a decaploid genotype (1-16-2) of tall fescue ( Festuca arundinacea Schreb.) is about twice that of a common hexaploid genotype (V6-802) (Plant Physiol. 72: 16–21, 1983). Translocation of photosynthate out of the leaves is a possible means of regulating carbon assimilation. To evaluate this possibility, we have examined a) translocation velocity, b) time course of translocation from leaves, c) photoassimilate partitioning pattern into whole plants in pulse and chase experiments, and d) interveinal distances between two ploidy genotypes. Most of the ¹⁴C accumulated in sucrose, and the labelled carbon moved down the leaf blades at similar velocities (6 to 10 cm h⁻¹) in both genotypes. Recent ¹⁴C assimilate was rapidly translocated from the fed area of the leaf blade. For example, the decaploid and the common hexaploid had translocated 40 and 26% of the ¹⁴C, respectively, at 6 h, and 79 and 49% of the ¹⁴C, respectively, at 24 h. Partitioning of ¹⁴C among plant organs was considerably different between the genotypes after a 24 h chase. For example, out of the total ¹⁴C recovered from the whole plant, the decaploid had retained 40% in the labelled leaf with 10, 33 and 29% in other leaves, stem bases and roots, respectively; whereas the hexaploid had retained 91% in the labelled leaf with 4, 3 and 2% in other leaves, stem bases and roots, respectively. However, the higher rate of translocation was correlated with greater interveinal distances in the decaploid genotype. These results suggested that the higher translocation percentage in the decaploid than the hexaploid genotype was due to greater sink activity.     

Translocation in Sugar-beet: I. ASSIMILATION OF 14CO2 AND DISTRIBUTION OF MATERIALS FROM LEAVES

¹⁴CO₂ was supplied to leaves, and movement of labelled carbonto other parts of the plant was assessed. Young growing leavesutilized assimilated carbon for their own growth and did notexport carbon to the rest of the plant, while fully expandedleaves exported much of their photosynthate, both to root andto young leaves. Translocation from a particular leaf was tothe two or three younger leaves on the same side of the plant,and to a sector of root below the source leaf. Specific distributionto growing leaves could be modified by partial defoliation.There was no movement of material to leaves which had emergedbefore the source leaf. Part of the carbon entering a leaf by assimilation (and, foryoung leaves, by translocation) was incorporated into insolublematerial, especially in young leaves. Some of the carbon enteringa developing root was permanently stored as sucrose, althoughmuch also entered insoluble material. Loss from the leaf ofcarbon fixed during a short period of photosynthesis was rapidat first but continued at a decreasing rate for several days.Some carbon fixed into the insoluble fraction was translocatedfrom the leaf later, during senescence. Sucrose was the mainmaterial translocated immediately after photosynthesis.     

Physiology of maize I: A comprehensive and integrated view of nitrogen metabolism in a C4 plant

Maize ( Zea mays L., line F2) plants were grown in the field under high or low fertilization input to monitor the metabolic, biochemical and molecular events occurring in young vegetative leaves and in the different leaf stages along the main axis in plants harvested 15 days after silking. This study shows that in maize which possess large sinks represented by the seeds, nitrogen (N) management is different compared with tobacco in which sink strength is much lower and mostly limited to young developing leaves. Although in young leaves nitrate assimilation predominates in both species, ammonium assimilation exhibits some species-specific differences with respect to inorganic and organic N metabolite accumulation during leaf ageing. These differences are likely to be related to the high sink strength of the ear in maize, which continuously imports carbon and N assimilates during grain filling. Consequently, a number of cytosolic glutamine synthetase isoenzymes are expressed during leaf ageing to maintain a constant flux of reduced N necessary for the synthesis of organic N molecules used either for leaf protein synthesis or directly translocated to the grain. This situation contrasts with that found in tobacco for which leaf ammonium assimilation in the plastids is shifted to the cytosol during the transition from sink leaves to source leaves. These species-specific differences for N assimilation and recycling are discussed in relation to the evolution of leaf photosynthetic activity and leaf senescence, which both seem to be largely dependent on the different sink strength in each species.     

Sucrose Hydrolysis in Relation to Phloem Translocation in Beta vulgaris

Asymmetrically labeled sucrose, ¹⁴C(fructosyl)sucrose, was used to determine whether sucrose undergoes extracellular hydrolysis during phloem translocation in the sugar beet, Beta vulgaris. In addition, the metabolism of various sugars accumulated and translocated was determined in various regious of the plant. These processes were studied in detached regions as well as in the intact, translocating plant in the source leaf, along the translocation path, and in a rapidly growing sink leaf and storage beet. The data show that, unlike sucrose accumulation into the sink tissue of sugarcane, sucrose is neither hydrolzyed prior to phloem loading or during transit, nor is it extracellularly hydrolyzed during accumulation into sink leaves or the storage beet.     

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     

留营养枝对棉株同化物生产,运转,分配及产量的影响

运用＾１４Ｃ示踪技术研究了留营养支棉株＾１４Ｃ同化物生产运转分配规律。结果表明：留营养枝与否对全株＾１４Ｃ总同化量基本没有影响,但留营养２枝棉株果枝叶的相对光合强度（以放射性比强度表示）降低;主茎叶、果枝叶的＾１４Ｃ同化量显著降低;营养２叶具有较高的光合作用强度和向外输送＾１４Ｃ同化物的转运速率,在＾１４Ｃ同化物生产运转分配中占有很重要的地位。留营养枝棉株＾１４Ｃ同化物的转运速率,在＾１４Ｃ同化     

Virus diseases of cacao in West Africa; effect of virus infection on growth and water content of cacao seedlings

Apparently healthy cacao seedlings were compared with those infected before planting with 'swollen shoot' viruses. The leaf area and the fresh and dry weights of each organ were measured. Infected plants were lower in dry weight, leaf area, relative growth rate and net assimilation rate; a smaller proportion of the dry matter was in the leaves and lateral roots, a larger proportion in stems and tap roots. Infection caused extensive necrosis of the lateral roots, and reduced the rate of depletion of reserves in the cotyledons and the water content of the plant. Many of these effects were apparent within a month of infection and planting.     

Effects of source/sink manipulation on net photosynthetic rate and photosynthate partitioning during grain filling in winter wheat

Source-sink relationship, which was influenced by both genotype and environmental factors, contributed to the variation in photosynthesis and photosynthate partitioning of wheat. Source reduction by partial defoliation increased leaf net photosynthetic rate (PN), and sink reduction decreased PN of irrigated wheat. However, the change in PN varied among genotypes. Source reduction enhanced photosynthate translocation into grain in irrigated wheat. However, the enhancement was more evident in cv. Lumai 215953 than incv. Lumai 15. Sink reduction had little effect on the translocation of photosynthate into grain in cv. Lumai 15, but decreased the translocation of photosynthate into grain and increased it into stem in cv. Lumai 215953. In rainfed, non-irrigated wheat, the source or sink manipulation influenced PN only slightly. The source reduction decreased the partitioning of photosynthates into the upper parts (including grains) of plant. However, very little effects of sink reduction on the production of photosynthates occurred in rainfed wheat. This showed that grain sink size was not a factor limiting the production of photosynthates, but controlled the partitioning of photosynthates. Sink reduction decreased photosynthate translocation into grains, and increased it into upper parts of rainfed wheat plant. This revised version was published online in July 2006 with corrections to the Cover Date.     

Enhancement of Phloem exudation from cut petioles by chelating agents

The photosynthetic assimilates in leaves of Perilla crispa attached to the plant were labeled by treating the leaves with (14)CO(2). When subsequently detached, these leaves exuded a negligible amount of radioactivity from the cut petiole into water. Ethylenediaminetetraacetate (EDTA), citric acid, and ethyleneglycol-bis (beta-aminoethyl ether) N,N'-tetraacetate greatly increased exudation of labeled assimilates into a solution bathing the petioles. The optimal concentration of EDTA was 20 mm, and maximal exudation took place between 2 and 4 hours after excision. Up to 22% of the radioactivity fixed in the leaf was exuded into an EDTA solution as compared to an export of 38% from attached leaves. The amount of radioactivity in the exudate was much reduced at low temperature. Presence of EDTA was required in the collecting solution for only 1 to 2 hours; upon transfer to water, exudation continued as in continuous presence of EDTA. Ca(2+) completely inhibited the effect of EDTA.Anatomical studies indicated that callose formation on the sieve plates near the cut surface of the petioles was less in leaves on EDTA than on water.More than 95% of the radioactivity exuded by detached leaves was present in the sugars verbascose, stachyose, raffinose, and sucrose, which are translocated in the phloem of Perilla. Labeled glucose, fructose, and galactinol were detected in the leaf blade and petiole, but not in exudates.The addition of EDTA to a solution bathing the petiole of detached leaves of Chenopodium rubrum and Pharbitis nil also increased the exudation of labeled assimilates. In these two species, label appeared only in a compound that cochromatographed with sucrose.It is concluded that the radioactive products in the solution are actually exuded by the phloem. Possibly EDTA chelates Ca(2+) that otherwise participates in the reactions that seal cut phloem.     

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     

两种相思叶片光合作用对干旱的反应

比较了绢毛相思和大叶相思的叶特性,旱季的田间光合速率和供水短缺对光合速率、气孔传导率和蒸腾速率的影响。绢毛相思的比叶重、单位叶面积的叶绿素含量和叶绿素a/b均较大叶相思高,但叶片含水量略低。绢毛相思的中午时叶片水势为-0.6±0.05MPa,而大叶相思则为-1.18±0.07MPa。绢毛相思叶片水势降低时,叶片鲜重的变化较大叶相思大。旱季10月,两种相思的日平均光合速率相近似,但绢毛相思有较高的气孔传导率和蒸腾速率。干旱处理引起大叶相思叶片水势降低较绢毛相思大。当绢毛相思叶片水势从-0.76MPa降至-1.35MPa,日平均光合速率降低49.4％;而大叶相思,叶片水势从-1.22MPa降低至-2.2MPa,日平均光合速率降低55.0%。大叶相思叶片水势降低的幅度比较大,光合速率降低亦大。     

Effects of Temperature and Photoperiod on Assimilate Partitioning in Potato Plants

The effects of temperature and photoperiod on d. wt partitioningand ¹⁴C translocation were studied in three potato varieties.High temperatures and long days enhanced plant growth in termsof plant height and number of leaves, and also affected d. wtpartitioning between the plant organs. However, no temperatureeffect was noted on total plant d. wt, nor on the export of¹⁴C from the source leaf. Translocation of ¹⁴C to the vegetativeorgans (leaves and stems) was greater at higher temperatures,while translocation to the tubers was less under these conditions.We suggest that, under the temperature regimes studied, themain effect of high temperature is on assimilate partitioningand not on total plant productivity. Differences in responseto high temperatures were observed among varieties, with Norchipshowing the least and Up-to-Date showing the most sensitivity. High temperature, partitioning of assimilates, ¹⁴C-translocation, potato, Solanum tuberosum var. Desirèe, Solanum tuberosum var. Norchip, Solanum tuberosum var. Up-to-Date