Water Potential, Translocation and Assimilate Partitioning

Lang, A. and Thorpe, M. R. 1986. Water potential, translocationand assimilate partitioning.—J. exp. Bot. 37: 495–503. The effect of water status upon translocation and assimilatepartitioning is examined both from theory and in an experimentwith young Phaseolus plants. Theory predicts that translocationis unlikely to be directly affected by water status. However,water potential differences within plants should influence translocationflow, with regions at lower potentials attracting disproportionatelylarge shares of assimilate. This prediction is supported in the experiment with Phaseolusin which the pattern of partitioning in the root changed rapidlyin response to bathing portions of it in solutions of differentosmolarity. The relevance of these findings to the growth of plants undernatural conditions is considered and evidence is presented thatwater potential gradients may be an Important factor in thecontrol of partitioning Key words: Phloem translocation, xylem transport, partitioning, water potential, control, osmotic potential     

Silencing of the tomato Sugar Partitioning Affecting protein (SPA) modifies sink strength through a shift in leaf sugar metabolism

Limitations in our understanding about the mechanisms that underlie source‐sink assimilate partitioning are increasingly becoming a major hurdle for crop yield enhancement via metabolic engineering. By means of a comprehensive approach, this work reports the functional characterization of a DnaJ chaperone related‐protein (named as SPA; sugar partition‐affecting) that is involved in assimilate partitioning in tomato plants. SPA protein was found to be targeted to the chloroplast thylakoid membranes. SPA‐RNAi tomato plants produced more and heavier fruits compared with controls, thus resulting in a considerable increment in harvest index. The transgenic plants also displayed increased pigment levels and reduced sucrose, glucose and fructose contents in leaves. Detailed metabolic and enzymatic activities analyses showed that sugar phosphate intermediates were increased while the activity of phosphoglucomutase, sugar kinases and invertases was reduced in the photosynthetic organs of the silenced plants. These changes would be anticipated to promote carbon export from foliar tissues. The combined results suggested that the tomato SPA protein plays an important role in plastid metabolism and mediates the source‐sink relationships by affecting the rate of carbon translocation to fruits.     

Partitioning Control by Water Potential Gradient: Evidence for Compartmentation Breakdown in Grape Berries

Xylem exudate was obtained from berries of Riesling grapes atdifferent stages of development after the onset of ripeningusing a pressure bomb technique. The osmotic potential of theexudate bore a 1:1 relationship to that of juice from the sameberries which were afterwards crushed and centrifuged. Thisresult provides the first direct evidence of compartmentationbreakdown in grape berries after the onset of ripening. Changesin berry deformability which occur at the same time and measurementsof the dynamics of exudation flow lead to the same conclusionregarding compartmentation breakdown. The breakdown in compartmentation occurs at the same time asthe rate of phloem translocation to the fruit suddenly increases.A mechanism was recently proposed to account for this increase.It required the existence of a water potential difference betweensource and sink such as would result from compartmentation breakdownin the sink tissues. The results, therefore, may be taken toindicate that this mechanism is indeed involved in the controlof assimilate partitioning in Vitis. Evidence in other publicationssuggests that the mechanism may be reasonably widespread inplants. Key words: Assimilate partitioning, phloem translocation mechanism, Vitis vinifera L., water potential gradients     

The role of protein kinases in the regulation of plant growth and development

We review the role of protein kinases in plant hormone-mediatedsignalling, nutrient signalling and cell cycle control and in the crosstalkbetween these different contributors to plant growth regulation. The areas ofhormone-mediated signalling covered include ABA-mediated responses to osmoticstress, wounding and pathogen attack, as well as ethylene and cytokininsignalling pathways. These areas involve members of several major protein kinasefamilies, including the SNFl-related protein kinase-2 (SnRK2) subfamily, thecalcium-dependent protein kinase (CDPK) family, the mitogen activated protein(MAP) kinase family, the glycogen synthase kinase (GSK)- 3/shaggy family and thereceptor-like protein kinase (RPK) family. In the section on nutrient signallingwe review the role of SnRK1 protein kinases in the global regulation of carbonmetabolism, including aspects of sugar sensing and assimilate partitioning, andwhat is known about nitrogen and sulphur nutrient signalling. In the cell cyclesection, we summarise progress in the elucidation of cell cycle control systemsin plants and discuss the interaction between cell cycle control anddevelopment. We expand further on the hypothesis of crosstalk between differentsignalling pathways in a separate section in which we discuss evidence forinteraction between plant growth regulators and the cell cycle, betweendifferent nutrient signalling pathways, between nutrient and cell cyclesignalling and between nutrient and ABA signalling.     

Photoassimilate partitioning in nodulated soybean III. The effect of changes in nodule activity shows that carbon supply to the nodule is not linked to nodule nitrogen metabolism

The hypothesis that photoassimilate partitioning to the soybean nodule is controlled by the rate of N export (via an effect on the turgor of the unloading pathway) was tested. Cessation of N2 fixation due to exposure of the nodulated root to Ar:O2 for over 3 h did not affect the partitioning of photoassimilate to the nodule. In contrast, anaerobic conditions (100% N2) resulted in a temporary cessation or marked slowing of carbon import into the nodule and root organs, reflecting an O2 requirement of the unloading process. Carbon accumulation by the nodulated root was less affected by a rhizosphere treatment of 2% O2, although partitioning was decreased over a period of hours. Treatment with 100% O2 also caused an immediate diversion of photoassimilate from the root to the shoot system, although the extent of this diversion was variable. Treatment with stepped increases in O2 did not affect partitioning. It is concluded that the unloading kinetics of the nodule were not disturbed by changes in nodule N metabolism, and therefore that control of assimilate partitioning to the nodule is not influenced by a turgor mechanism involving a balance between the primary export and import solutes. However, photoassimilate import was matched to the respiratory demand of the root system.     

The role of sink demand in carbon partitioning and photosynthetic reinvigoration following shoot decapitation

Photosynthesis, growth, and carbon partitioning of vigorous coppice shoots were compared with the slower growing intact shoots of Populus maximowiczii × nigra L. MN9 to determine the relationship between carbon partitioning and photosynthetic rate. Relative height growth rate of coppice shoots was 2.2 times that of intact shoots with net photosynthetic rate 1.9 times that of intact shoots. Coppice leaves exported a larger proportion of newly-fixed assimilate (11% compared with 6%) after a 4-h chase. The greater export from coppice leaves was correlated with a greater proportion of [¹⁴C]-labelled photosynthate deposited as starch in stems 4 cm below the point of label application. Coppice leaf assimilate levels were reduced to 15% that of leaves on intact plants, but coppice leaves had twice the concentration of labelled sucrose. Carbohydrates constituted 55% of the water-soluble [¹⁴C]-labelled photosynthate in leaves of coppice shoots compared with 40% in intact shoots. The results suggest that carbon allocation and partitioning in coppice shoots were altered towards production and export of new assimilate, and support the hypothesis that photosynthetic rate is responsive to sink demand for assimilates.     

Control of Import and Export of Photosynthate in Leaves

The flow of photosynthetic assimilate in leaves was traced usingradioactive ¹¹CO₂. In younger leaves the direction of flow canchange in response to changes in assimilate potential in therest of the plant whereas inolder leaves it cannot. The irreversibilityof flow in older leaves is apparently a result of their havinga loading process able to maintain a sieve tube assimilate potentialhigher than elsewhere in the plant. Systematic shifts in the responses of leaves to a successionof short term changes in sink demand are best understood interms of radial exchanges between the sieve tubes and storagepools of limited capacity in the surrounding tissue. These takeplace throughout the length of the sieve tubes and may giverise to flows in opposite directions, at the same time, at differentpoints in the same sieve tube. Behaviour indicates that flow between different pools of assimilatein the plant can be thought of as being a functionof differencesin their assimilate potentials, with changes in potential elicitingchanges in flow. Key words: Translocation, carbon partitioning control, leaves     

Nitrate Uptake and Partitioning by Corn Root Systems : Differential Effects of Ammonium among Genotypes and Stages of Root Development

The relative effects of ammonium on nitrate uptake and partitioning during induction were compared among decapitated seedlings of three corn (Zea mays L.) genotypes at two developmental stages. This study tested the hypothesis that root systems efficient at translocating products of ammonium assimilation away from sites of nitrate uptake or reduction would exhibit less inhibition of nitrate uptake by ammonium compared to root systems with inefficient N translocation efficiency. Inhibition of nitrate uptake by ammonium was relatively slight at day 5 ranging from 0% to 20% among the three genotypes, as compared to greater inhibition, from 20% to 37%, at day 8. Five-day-old roots exhibited negligible xylem translocation capacity in comparison with those grown for 8 days. Thus, although the capability to translocate ammonium assimilates out of the root increased between days 5 and 8, inhibitory effects of ammonium also increased. In the absence of ammonium, nitrate uptake per unit root mass increased between days 5 and 8. This increased activity of the uptake system was proportionally more sensitive to ammonium.

Partitioning of entering nitrate into the reduction process was positively correlated with lateral root development of the inbred root systems at 5 and 8 days. This is supportive of a localization of a major portion of nitrate reduction occurring in root apical regions. Nitrate reduction was the partitioning process most severely inhibited by ammonium in all cases, ranging from 39% to 55% inhibition. In contrast, ammonium-inhibition of nitrate accumulation in the root tissue and translocation via xylem vessels varied with genotype and root age.

Two mechanisms of ammonium-inhibition of nitrate are implicated, one which directly affects nitrate reduction and the uptake system associated with it, and another which may involve potassium as an intermediate regulator of nitrate accumulation in the root tissue and nitrate translocation out of the root tissue.

     

Shoot Apex Demand Determines Assimilate and Nutrients Partitioning and Nutrient-uptake Rate in Tobacco Plants

Our previous experiment revealed that apex-removed plants have larger root systems but a lower K＋-uptake rates than intact tobacco plants. Since the apex is not only a center of growth and metabolism, but also an important place of auxin synthesis and export, the aims of this study were to distinguish whether the apex demand or auxin synthesized in the apex regulates assimilate and nutrients partitioning within plant, and to explain the reason for the lower K＋-uptake rate of the apex-removed plant. In comparison with the control plant, covering the shoot apex with a black transparent plastic bag reduced net increases in dry matter and nutrients; however, the distribution of the dry matter and nutrients between shoot and roots and nutrient-uptake rates were not changed. Removal of the shoot apex shifted the dry mass and nutrients distributions to roots, and reduced the rate of nutrient uptake. Application of 1-naphthylacetic acid （NAA） could partly replace the role of the removed apex, stimulated assimilate and nutrient deposition into the treated tissue, and enhanced the reduced plasma membrane ATPase activity of roots to the control level. However, treatment of the apex-removed plants with NAA could not rescue the reduced nutrient uptake rate and the shifted assimilates and nutrients partitioning caused by excision of the apex. Higher nutrient uptake rate of the intact plants could not be explained by root growth parameters, such as total root surface area and number of root tips. The results from the present study indicate that strong apex demand determined assimilates and nutrients partitioning and nutrient-uptake rate in tobacco （Nicotiana tabacum） plants.     

Source-sink regulation by sugar and stress.

The regulation of carbon partitioning between source and sink tissues in higher plants is not only important for plant growth and development, but insight into the underlying regulatory mechanism is also a prerequisite to modulating assimilate partitioning in transgenic plants. Hexoses, as well as sucrose, have been recognised as important signal molecules in source-sink regulation. Components of the underlying signal transduction pathways have been identified and parallels, as well as distinct differences, to known pathways in yeast and animals have become apparent. There is accumulating evidence for crosstalk, modulation and integration between signalling pathways responding to phytohormones, phosphate, light, sugars, and biotic and abiotic stress-related stimuli. These complex interactions at the signal transduction levels and co-ordinated regulation of gene expression seem to play a central role in source-sink regulation.     

Effect of Heat Stress on Assimilate Partitioning in Tomato

Differences in assimilate partitioning in response to heat stresswere observed between the heat-sensitive tomato cultivar, RomaVF and the heat-tolerant cultivar, Saladette. Transport of carbonto the trusses and apex was inhibited in both cultivars, particularlyin Roma VF. Basipetal transport to the roots was inhibited inRoma VF only. Assimilate partitioning within the young trusswas also affected by heat stress in both cultivars, but wasmore pronounced in Roma VF: at higher temperatures more ¹⁴Cwas found in the peduncle and in the older flower buds thanin the younger flower buds. The youngest flower buds were moresensitive than other parts of the truss to heat stress. Theuptake of [¹⁴C]sucrose by detached flower buds from agar mediumwas lower at higher temperatures. Alteration of assimilate partitioningwas also observed following the application of GA₃+Kinetin tothe first truss. The possible relationship between flower-setin tomato and carbohydrate stress is discussed. Lyeopersicon esculentum Mill., carbon translocation, starch     

Re-interpretation of an Experiment on the Role of Assimilated Transport Resistance in Partitioning in Tomato

The importance of transport resistance (distance between sourceand sink) on assimilate partitioning in tomato is questioned.Slack and Calvert ( Journal of Horticultural Science 52 : 309–315,1977) concluded that, in tomato, excising of fruit trusses showeda direct influence of distance from source on assimilate partitioning.A dry matter distribution model for tomato, based on the hypothesisthat distribution is regulated by the sink strengths of theplant organs and that no influence of transport resistance onpartitioning exists, has been described and validated by Heuvelink( Annals of Botany 77 : 71–80, 1996). Using this model,it is shown that the results of Slack and Calvert (1977) canbe explained more simply on the basis of the succession of trusseswith growth shifted with respect to time. Therefore, their resultsdo not prove that transport resistance plays a role in assimilatepartitioning. Allocation; distance; dry matter distribution; model; assimilate pool; partitioning; simulation; transport resistance; tomato     

The proton-sucrose symport

The heterotrophic tissues of the plant are dependent upon carbon and nitrogen import for normal growth and development. In general, oxidized forms of these essential elements are reductively assimilated in the leaf and, subsequently, sucrose and amino acids are transported to the heterotrophic cells in a process known as assimilate partitioning. In many plant species, a critical component of the assimilate partitioning pathway is the proton-sucrose symport. This active transport system couples sucrose translocation across the plasma membrane to the proton motive force generated by the H⁺-pumping ATPase. To date, the proton-sucrose symport is the only known system that can account for sucrose accumulation in the vascular tissue of the plant. This review focuses on recent advances describing the transport properties and bioenergetics of the proton-sucrose symport.     

Effect of shading individual soybean reproductive structures on their abscisic Acid content, metabolism, and partitioning

Pod set in soybean is related to carbon partitioning and may be, at least partially, regulated by abscisic acid (ABA) concentrations. The studies reported here examine the relationship between carbon and ABA partitioning, reproductive abscission and ABA metabolism. The partitioning of radiolabeled ABA and photoassimilates from leaves to flowers and endogenous ABA concentrations were determined in shaded and unshaded reproductive structures. Aluminum foil was gently placed over individual soybean reproductive structures for 48 hours at 0, 4, 12, 17, and 22 days after anthesis (DAA). Shading of flowers at 12, 17, and 22 DAA resulted in significantly reduced concentration of ABA. However, shading had no effect on the catabolism of exogenously supplied [³H] ABA. The shading treatment on the first four of the five dates reduced partitioning of photoassimilates and ABA from the subtending leaf to the flower. Shading of reproductive structures also caused a significant reduction in the amount of assimilate exported from the subtending leaf, at 17 DAA. We conclude that shade-induced premature reproductive abscission in soybean is not stimulated by high levels of ABA within reproductive structures, but that ABA may inhibit abscission of reproductive structures by playing a role in preferential assimilate partitioning.     

Assimilate Partitioning in the Variegated Coleus Leaf

Mature leaves of a variegated cultivar of Coleus blumei Benth. with a green border and central albino region constitute a source-sink system suitable for studies on assimilate partitioning. Leaves treated with ¹⁴CO₂ on a small part of the intact green border export assimilate via the shortest path into the stem. Leaves with all but a small lobe of the green border removed show different partitioning of labeled assimilates when the leaf is exposed to ¹⁴CO₂ (Fisher and Eschrich, 1985): The whole albino region of the leaf is supplied but no tracer is exported into the stem. When the green border is completely removed, ¹⁴CO₂-treatment of the albino region leads to the fixation of CO₂, obviously by PEP carboxylase, as indicated by the occurrence of labeled malate. Results show that the albino region of the variegated leaf constitutes a potential sink when deprived of its green border. In addition, CO₂-fixation by PEP carboxylase in albino tissue seems to indicate a common capacity of leaves which is normally masked by photosynthesis. The difference of assimilate partitioning between leaves with intact and leaves with partly removed green borders demonstrates that the unlabeled assimilates control the movement of labeled assimilates.     

Maternal,Single-Gene Regulation of Assimilate Partitioning in Pea

Assimilate partitioning has been identified as a key process in the control of yield. Although the role of reproductive structures in this process has received intensive study, our understanding of the role of the maternal plant is limited. We suggest that the Sn gene of pea (Pisum sativum L.) is a potentially valuable genetic tool for studying maternal regulation of partitioning. In this study, nearly isogenic lines differing at the Sn locus were compared with respect to seed-filling characteristics and carbon assimilation. Lines with the Sn gene had a slower rate and shorter duration of seed growth than the line recessive for this gene, and these traits could not be ascribed to reduced carbon assimilation. Flowers of the two nearly isogenic lines were manually pollinated to control the genotype of the developing embryo independently of the maternal genotype. The final dry weight of the seed was determined by the genotype of the maternal plant and not by the genotype of the embryo, supporting the hypothesis that the Sn gene acts in the vegetative plant to regulate the partitioning of assimilates between vegetative and reproductive growth. Although the Sn gene has been noted for delaying apical senescence, it also delayed leaf senescence in this study; leaves of the Sn line continued to photosynthesize long past the time that leaves of the recessive line had senesced and after the seeds and pods were dry.     

Assimilate relations in source and sink leaves during acclimation to a CO2-enriched atmosphere

Evidence from previous studies suggested that adjustments in assimilate formation and partitioning in leaves might occur over time when plants are exposed to enriched atmospheric CO₂. We examined assimilate relations of source (primary unifoliolate) and developing sink (second mainstem trifoliolate) leaves of soybean [ Glycine max (L.) Merr. cv. Lee] plants for 12 days after transfer from a control (350 μl l⁻¹) to a high (700 μ l⁻¹) CO₂ environment. Similar responses were evident in the two leaf types. Net CO₂ exchange rate (CER) immediately increased and remained elevated in high CO₂. Initially, the additional assimilate at high CO₂ levels in the light and was utilized in the subsequent dark period. After approximately 7 days, assimilate export in the light began to increase and by 12 days reached rates 3 to 5 times that of the control. In the developing sink leaf, high rates of export in the light occurred as the leaf approached full expansion. The results indicate that a specific acclimation process occurs in source leaves which increases the capacity for assimilate export in the light phase of the diurnal cycle as plants adjust to enriched CO₂ and a more rapid growth rate.     

A model for co-translational translocation: ribosome-regulated nascent polypeptide translocation at the protein-conducting channel

The protein-conducting channel (PCC) must allow both the translocation of soluble polypeptide regions across, and the lateral partitioning of hydrophobic transmembrane helices (TMHs) into, the membrane. We have analyzed existing structures of ribosomes and ribosome-PCC complexes and observe conformational changes suggesting that the ribosome may sense and orient the nascent polypeptide and also facilitate conformational changes in the PCC, subsequently directing the nascent polypeptide into the appropriate PCC-mediated translocation mode. The PCC is predicted to be able to accommodate one central, consolidated channel or two segregated pores with different lipid accessibilities, which may enable the lipid-mediated partitioning of a TMH from one pore, while the other, aqueous, pore allows translocation of a hydrophilic polypeptide segment. Our hypothesis suggests a plausible mechanism for the transitioning of the PCC between different configurations.     

Sink development and dry matter distribution in storage root crops

Events during early development are examined for their contribution to dry matter distribution in the mature plant. It is concluded that timing of initiation of the storage root is not a major cause of differences between cultivars of carrot. These may airse from differences in the control of partitioning of assimilate at storage organ initiation. Root crop species may be affected differently by environmental factors such as density as a result of different thresholds of response of sink development to source stimuli.     

Patterns of 14C-labelled Assimilate Partitioning in Red and White Clover During Vegetative Growth

A quantitative analysis of the ¹⁴C-labelled assimilate suppliedby the expanded leaves on the primary shoot to growing leaves,stem, lateral shoots (branches or stolons) and roots in redand white clover was conducted during vegetative growth. Stem growth of the primary shoot was inhibited in both cloversand utilized no energy resources. The growing leaves at theprimary shoot apex of white clover imported 4 per cent of theshoot's assimilate compared with 10 per cent in red clover.At the basal end of the primary shoot, the tap root of whiteclover imported 16 per cent of the shoot's assimilate comparedwith 22 per cent in red clover. Branches in red clover and stolonsin white clover were by far the largest sinks for primary shootassimilate, importing 39 per cent and 63 per cent of the labelledassimilate, respectively. Analyses of the translocation of assimilate from individualprimary shoot leaves demonstrated that in both clovers olderleaves exported more of their assimilate to branches or stolons,whereas younger leaves exported more of their assimilate toroots, and possibly in white clover, to growing leaves at thetip of the shoot. Of the labelled assimilate exported to branchesor stolons, each primary shoot leaf exported preferentiallyto the branch or stolon in its own axil, but in addition exportedsubstantial quantities of assimilate to all other axillary shoots,particularly those arising from basal axils where the subtendingleaf had died. Trifolium repens, Trifolium pratense, red clover, white clover, assimilate partitioning, perennation