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 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.     

柑橘果实的光合特性、产物运输及分配在糖分积累中的作用

测定了温州蜜柑 (CitrusunshiuMarc .cv .Miyagawawase)果实发育进程中干鲜重、果皮光合速率和叶绿素含量的变化 ,并用14 CO2 示踪技术研究了果皮和叶同化生成的光合产物在果实内的运输分配特性。结果表明 :果皮光合速率与叶绿素含量有关 ,随着叶绿素含量的下降 ,果实光合速率也快速下降。在果实完熟之前 ,即使是当果皮积累的干重超过汁囊时 ,叶同化产物仍主要分配到汁囊中 ;而在完熟阶段 ,果皮光合速率接近零 ,果皮成了叶同化产物的主要库。果皮的同化产物 ,主要保留在果皮中 ,输入到汁囊的比率随果实发育而下降 ,但高峰时也有 12 %输入汁囊。与对照相比 ,果实遮光处理后降低了果皮与汁囊的干重和含糖量。上述结果表明果皮光合产物主要用于果皮自身的发育并能减少对叶光合产物的依赖 ,同时也能部分增加汁囊糖的积累     

Assimilate Transport and Partitioning in Plants: Approaches,Methods, and Facets of Research

This review, dedicated to the 100th anniversary of A.L. Kursanov's date of birth, considers the development of phloem transport studies since his book, Assimilate Transport in the Plant, was published in 1976. This book and several other fundamental publications on phloem structure and functions basically shaped this physiological issue; as a result, several international meetings by scientists working in the area were induced, and the proceedings of these meetings were published at regular intervals. Six conferences have been held to date, and six corresponding collections of papers have been published and are reviewed here along with other experimental communications and reviews. This review considers the following topics: (1) the phloem structure and the ultrastructure of specialized phloem cells, (2) the physiological functions of phloem and their regulation, (3) photosynthesis and phloem loading with assimilates, (4) phloem unloading and the related processes of plant growth and development, (5) the mechanisms of sugar and amino acid transport, (6) the levels of transport, (7) transport compartments; (8) xylem–phloem and symplast–apoplast communication; (9) phloem transport vs. the integral plant physiology, (10) transport of xenobiotics, and (11) the trophic transport networks in symbionts.     

Root Confinement and its Effects on the Water Relations, Growth and Assimilate Partitioning of Tomato (Lycopersicon esculentum Mill)

Although it is well established that the root growth in manyspecies is very sensitive to mechanical impedance or to confinementin small volumes, little is known about the consequent effectson growth of the whole plant and the mechanisms involved. Thiswork investigated the effects of root confinement on the waterrelations, growth and assimilate partitioning of tomato (Lycopersiconesculentum Mill) grown in solution culture. Six-week old plants were transferred to either 4500 ml or 75ml containers filled with nutrient solution, and allowed togrow for 14 d. Transpiration, leaf-air temperature differences,and leaf diffusive resistances were measured frequently. Leaf,stem and shoot dry masses, leaf area and root length, were estimatedwhen the treatments were imposed and at the end of the experiment.After 14 d growth the root and shoot hydraulic resistances wereestimated from measurements of leaf water potential and transpirationrate, using a steady-state technique. Confining root growth to the small containers substantiallyreduced shoot and root growth and increased the proportion oftotal dry matter present in the stems. These effects were dueto drought stress. The hydraulic resistance of the root systemwas greatest in the confined plants. This led to more negativeleaf water potentials, increased leaf diffusive resistance,and reduced the net assimilation rate by a factor of 2.5. Transpirationper unit leaf area was less affected. However, cumulative transpirationwas also reduced by a factor of 2.5. mostly because of the smallerleaf area on the confined plants. Root hydraulic resistivitywas measured at 3.1 x 10¹²s m^–1 in the control treatment,but increased to 3.9 x 10¹² s m^–1 for roots in the smallcontainer. The mechanisms by which root confinement caused drought stressand disrupted the pattern of assimilate partitioning are discussedin detail. Assimilate partitioning, Lycopersicon esculentum, root confinement, plant growth, root growth, root resistance, shoot resistance, tomato, transpiration, water-use efficiency     

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     

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     

Regulation of Assimilate Translocation Between Leaves and Fruits in the Tomato

Simultaneous measurement of export from leaves and import tofruits were made on tomato plants reduced to one fully expandedleaf and one fruit. Experimental leaves were exposed to sixlight flux densities (0.5–100 W m^–2) for 24 h whilerapidly growing fruits were kept in the dark at 22 °C. The rates of export of assimilate from these leaves varied from70 to 120 mg C leaf^–1 day^–1 corresponding with ratesof carbon fixation from 3 to 290 mg C leaf^–1 day^–1.Export from leaves with the lowest carbon fixation rates weremaintained by a loss of up to one-sixth of their initial carbon.In contrast, leaves with the highest carbon fixation rates exportedonly half the newly fixed carbon. The rates of import of assimilate to similar-sized fruits (c.16 cm³) were between 80 and 110 mg C fr^–1 day^–1but differed from the export rates of the source leaves. Thespecific growth rates and the specific respiration rates ofthe fruits were related to their initial carbon content at thebeginning of the experiment. Thus, over 24 h, the rate of importwas predetermined by the developmental stage of the fruit unalteredby the rate of current carbon fixation in the source leaf. Translocationof assimilate was regulated by sink demand under both source-and sink-limiting conditions in this short-term situation. The dynamic relationship between assimilate production in leavesand its utilization in fruits is discussed together with therole of sucrose concentration in these organs in regulatingtransport. Lycopersicon esculentumL, tomato assimilate translocation, source-sink relationships     

Mutant dn Influences Dry Matter Distribution, Assimilate Partitioning and Flowering in Lathyrus odoratus L.

The flowering mutant dn in sweet pea was used as a tool to study¹⁴C-assimilate and dry matter partitioning with respect to nutrientdiversion theories on the control of flower initiation. Wildtype plants (Dn^h) are photoperiodic and exhibit late floweringand profuse basal branching in short days while mutant plants(dn) are day neutral, early flowering and devoid of basal laterals.In short days, dn plants exported a significantly greater proportionof assimilate acropetally than (Dn^h) plants and the upper portionof dn plants had a greater dry weight. These differences werereduced dramatically when basal laterals were excised regularlyfrom the (Dn^h) plants although the difference in flowering remained.However, the effect of dn on resource allocation within theapical region may be more important in regard to flowering thanthe effect on acropetal versus basipetal movement. In shortdays, the dn plants partitioned significantly more resourcesinto their internodes and petioles, and less into their leaflets,than Dn^h plants as shown by dry weight and ¹⁴C-assimilate measurements.These differences were apparent from as early as node 7 up tothe node of flower initiation in dn plants (node 30) and theywere not eliminated by removal of basal laterals from Dn^h plants.Differences between dn and Dn^h plants in partitioning and floweringwere largely eliminated under long days. The fact that in thisspecies a single gene influences both resource allocation andflower initiation lends further support to nutrient diversionhypotheses on the control of flowering. Key words: Assimilate partitioning, branching, flowering, mutant, sweet pea     

C-Photosynthate Partitioning and Translocation in Soybeans during Reproductive Development

Partitioning and translocation of ¹⁴C-photosynthates were examined during flowering and seed maturation in soybean (Glycine max [L.]Merr.) plants to quantify allocation to sugars, amino acids, organic acids, and starch and to study transport of C and N from leaves to reproductive sinks. The trifoliolate leaf at the eighth node was exposed to steady state levels of ¹⁴CO₂ for 2 hours, followed by immediate extraction and identification of radioactive assimilates in the fed leaf blade, tissues of the transport path (e.g. petiole and stem), and fruits if they were present. About one-third of the total ¹⁴C recovered from the leaf blades was in starch until late pod-filling, after which the proportion dropped to 16%. Sugars comprised 70% to 86% of the recovered ¹⁴C from soluble assimilates of the source leaf, with highest proportions occurring during late flowering and early pod-filling. Amino acids accounted for 8% to 17% of the ¹⁴C recovered from the soluble fraction, and were most evident during early flowering and mid to late pod-filling. The ¹⁴C-organic acids comprised from 3% to 14% of the soluble ¹⁴C-assimilates in leaves. Petioles consistently contained a higher percentage of recovered radioactivity in sugars (87-97%) and a lower percentage in amino acids (3-12%) than did leaf blades. ¹⁴C-Amino acids in petioles attained their highest levels during mid and late pod-filling, while ¹⁴C-organic acids comprised 2% or less of the recovered radioactivity after pod initiation. The distribution of ¹⁴C-assimilates in the internode below the source leaf was similar to that found in petioles. A comparison of the above data to calculated C and N requirements for seed development suggests that ¹⁴C-amino acids derived from current photosynthesis and translocated from source leaves supply at least 12% to 48% of the seed N depending on the stage of pod-filling.     

Characteristics of a Diurnal Rhythm of Assimilate Translocation in Grapevine Plants

Characteristics of photoassimilate efflux from the leaves of grapevine (Vitis vinifera L., var. Rkatsiteli) plants under natural conditions and upon darkening, defoliation, and ABA treatment at short exposures (2, 6, and 24 h) as well as diurnal pattern of starch content in the leaves were investigated. Translocation of assimilates from the leaves of grapevine usually occurred at night and started after 20:00 or 21:00. However, such pattern of translocation was not strictly determinated and could be changed upon the action of some factors. Darkening, defoliation, and treatment with ABA initiated the translocation of photoassimilates from the leaves in the daytime. Darkening and ABA treatment turned out to be more efficient than defoliation. Darkening and defoliation significantly changed the character of partitioning of photoassimilates in the stem. We corroborated the opinion that in grapevine, the efflux of assimilates from source leaves is also governed by the feedback mechanism and probably much depends on the level of carbohydrates.     

Partitioning Water Potential and Specific Salt Effects on Seed Germination of Four Grasses

In this study water potential (     

The Effect of Calcium Starvation on Assimilate Partitioning and Mineral Distribution of the Phloem

Populus plants were grown in a medium lacking calcium and exposedto ¹⁴CO₂. In contrast to plants in the complete nutrient medium,the percentage amount of ¹⁴C-assimilates increased in the leavesof calcium-deficient plants and decreased in the stem and theroots. When plants were grown without potassium or magnesiumno differences in the amount of ¹⁴C-label occurred in comparisonwith plants in the complete nutrient medium. Translocation wasrecorded by microautoradiography. It was observed that considerableamounts of labelled photoassimilates were unloaded from thephloem in the middle part of the stem in plants of the completenutrient medium. In contrast, during calcium starvation ¹⁴C-labelwas restricted to the phloem of the stem. In addition, the concentrationsof magnesium and phosphorus showed a remarkable increase instem sieve tubes of calcium-deficient plants. When sieve tubesof source leaves from Populus, barley and maize were comparedwith those of sink leaves, the latter showed higher calciumconcentrations. The results suggest that calcium is a necessaryfactor in the regulation of phloem translocation. Key words: Calcium deficiency, phloem translocation, sieve element loading and unloading, X-ray microanalysis     

Assimilate Uptake and Water Loss in Maturing Barley Grains

When detached ears of barley were supplied with [U-¹⁴C] sucrosefor 4 h, ¹⁴C was detected in the endosperms of grains from earsharvested 50 ‘days’ after anthesis, i.e. 10–15‘days’ after cell wall modifications had taken placein the chalazal region of the caryopsis. In 55-‘day’ears which had been supplied with [U-¹⁴C] sucrose for 20 h no¹⁴C was detected in the upper part of the rachis or in grainswith a moisture content of less than about 25%. In the rachisof wheat and barley, the walls of the xylem parenchyma and phloemcells stain strongly with ruthenium red, and in immature rachidesthis stainability is lost when sections are incubated with pectinase.Near the apex of 55-‘day’ ears the pectinaceouswall material in the vascular bundles is resistant to digestionby pectinase, and pectic substances, which are removed by pectinase,fill the lumina of the xylem elements. These observations arediscussed in relation to the dehydration of the maturing grain. Key words: Barley, ¹⁴C-sucrose, Pectic material, Xylem     

Far-Red Light Reflection from Green Leaves and Effects on Phytochrome-Mediated Assimilate Partitioning under Field Conditions

The influence of plant spacing and row orientation on spectral distribution of light received by growing soybean (Gylcine max [L.] Merr.) plants was measured under field conditions. Light absorption, reflection and transmission of individual leaves showed that most of the blue and red was absorbed while most of the far-red was either reflected or transmitted. Plants growing in the field received different ratios of far-red relative to red, depending on nearness and/or orientation of other vegetation. Plants grown in close-spaced rows, or high population densities, received higher far-red/red ratios than did those grown in wide rows, or sparse populations. Heliotropic movements of the leaves also contributed to the far-red reflection patterns associated with row orientation. Under field conditions, differences in far-red/red ratios associated with nearness of competing vegetation became more pronounced with low solar angle near the end of the day. Plants exposed to far-red for 5 minutes at the end of each day in controlled environments, and those grown in close-spaced rows in the field, developed longer internodes and fewer branches. Red, far-red photoreversibility in the controlled environment study indicated involvement of phytochrome. Dry matter partitioning among plant components in the field was related to far-red/red light ratio received during growth and development.     

Dynamics of Nitrogenous Assimilate Partitioning between Cytoplasmic and Vacuolar Fractions in Carrot Cell Suspension Cultures

Bulk vacuole isolation, gas chromatography-mass spectrometry,, and high-performance liquid chromatography have been used to investigate the accumulation and partitioning of assimilated nitrogen supplied as ¹⁵NH₄Cl between vacuolar and extravacuolar (cytoplasmic) fractions of protoplasts from suspension cultures of carrot (Daucus carota L. cv Chantenay). Glutamine was the most abundant amino acid in the vacuole of protoplasts from late-exponential phase cells, whereas alanine, glutamate, and γ-aminobutyric acid were located primarily in the cytoplasmic fraction. In ¹⁵N-feeding studies, newly synthesized glutamine partitioned strongly to the vacuole, whereas glutamate partitioned strongly to the cytoplasm, γ-aminobutyric acid was totally excluded from the vacuole, and alanine was distributed in both compartments. Comparison of the ¹⁵N-enrichment patterns suggests that initial assimilation to glutamine occurs within a subcompartment of the cytoplasmic fraction. The protoplast-feeding technique may be extended to investigate cytoplasmic compartmentation further.     

Effect of Water Stress on Grain Growth and Assimilate Partitioning in two Cultivars of Wheat Contrasting in their Yield Stability in a Drought-Environment

Grain growth pattern and grain weight in relation to its positionbetween and within spikelets showed assimilate limitation tobe a factor reducing grain yield in wheat plants grown on storedmoisture in the field. Estimation of the relative contributionof pre- and post-anthesis assimilates to grain yield by a ¹⁴Clabelling technique indicated that cultivar C 306, known forits stability in yield and drought-tolerance, was characterizedby substantial mobilization of pre-anthesis assimilate (P),both in irrigated (24 per cent) and unirrigated (33 per cent)plants. As against this, in Kalyansona, a high yielding cultivarwith moderate stress tolerance, the contribution of P to grainyield was only 13 and 22 per cent in irrigated and unirrigatedplants, respectively. The mobilization of amino acids, estimatedfrom nitrogen contents at anthesis and maturity, was decreasedby water stress and was accompanied by a corresponding increasedtransport of carbohydrates. Assimilate partitioning, drought tolerance, grain growth, stability, water stress, wheat     

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     

Regulation of Assimilate Partitioning in Soybean : Initial Effects following Change in Nitrate Supply

Increased concentrations of nitrate in a nutrient solution (2, 5, and 10 millimolar KNO₃) were correlated with increased shoot:root ratios of non-nodulated soybeans (Glycine max [L.] Merr.) grown in sand culture. While altering the pattern of C and N partitioning, the N treatments did not affect whole plant photosynthesis over the study period. To determine the mechanism responsible for the observed changes in assimilate partitioning, detailed C and N budgets were worked out with plants from each N treatment over three consecutive 4-day periods of midvegetative growth. The information for the C and N budgets from the 2 and 10 millimolar NO₃⁻ treatments was combined with data on the composition of xylem and phloem exudates to construct a series of models of C and N transport and partitioning. These models were used to outine a `chain-reaction' of cause-and-effect relationships that may account for the observed changes in assimilate partitioning in these plants. The proposed mechanism identifies two features which may be important in regulating the partitioning of N and other nutrients within the whole plant. (a) The concentration of N in the phloem is highly correlated with the N concentration in the xylem. (b) The amount of N which cycles through the root—from phloem imported from the shoot to xylem exported by the root—is regulated by the root's requirement for N: only that N in excess of the root's N requirements is returned to the shoot in the xylem. Therefore, roots seem to have the highest priority for N in times of N stress.     

The Influence of Applied Nitrogen on Export and Partitioning of Current Assimilate by Field-grown Potato Plants

Field-grown potatoes were subjected to N deficiency (no appliedN) or received high levels of N (240 kg N ha^–1) at planting.The effects of these treatments were monitored at five stagesduring growth in terms of the allocation of photosynthate withinthe leaf, and the export and partitioning of carbon to differentsinks. N deficiency significantly raised the starch concentrationin all organs of the plants, particularly in leaves and stems,and as a consequence the total amount of starch in the canopyof the low N plants remained greater than that of the high Nplants until approx. 100 days after planting (DAP). The totalamounts of carbohydrates, protein and amino acids were calculatedfor each treatment and these values were used to derive a balancesheet for major reserves. Net losses of reserves occurred fromthe canopy in both treatments in the period 97–133 DAP,although these were shown to represent < 3 per cent of thetotal gain in tuber dry weight for the season. Partitioning of ¹⁴C assimilates was examined in whole plantsand also in single leaves. Reduced partitioning to the tubers,seen in high N plants throughout their growth, was shown tobe due to decreased percentage export by the leaf and accumulationof exported ¹⁴C by the stems. Partitioning to the tubers inlow N plants increased prior to senescence when 87 per centof the fixed ¹⁴C was exported within 24 h, 80 per cent of thisto the tubers. The equivalent values for the high N plants were77 and 60 per cent respectively. Increased percentage exportcoincided with decreased allocation to starch in the leaf, anda link between these processes is suggested. N also significantlyaltered the allocation of ¹⁴C within the leaf and may have influencedthe degradation of starch in the dark to a greater degree thanits synthesis in the light. The enzymes sucrose phosphate synthase (SPS), and starch synthasewere measured concurrently with partitioning. High N plantsshowed higher rates of activities of each of the enzymes althoughboth enzymes showed a similar pattern of development over theseason, irrespective of N treatment. The data are discussed in the light of conflicting reports concerningthe influence of N on translocation and partitioning. ¹⁴C assimilates, carbohydrates, nitrogen, potato (Solanum tuberosum L.), protein