Effect of Nitrogen Deficiency upon Translocation of C in Sugarcane

Withholding nitrogen decreased the percentages of nitrogen and chlorophyll in the blades; reduced the total fixation of radioactive carbon dioxide at 15, 37, and 178 seconds; and changed the relative composition of fixation products. Translocation of radioactive photosynthate from the fed part down the attached blade and into the stalk was less in the plants deprived of nitrogen than in the control plants supplied with nitrogen. Both the percentage of total activity translocated and the velocity of transport were decreased by nitrogen deficiency. During a translocation period of 90 minutes the minus nitrogen blade retained more ¹⁴C-sucrose than the control in the fed part and the blade below the fed part, but it sent less ¹⁴C-sucrose to the sheath of the fed leaf. Thus translocation decreased with nitrogen deficiency not for lack of sucrose but for some other reason. Although withholding nitrogen decreased translocation of labeled carbon in and from attached blades, there was no effect upon transport in detached blades. The effect of nitrogen deficiency upon translocation may be indirect and secondary to the effect upon growth of the plant as a whole.     

Comparison of upward and downward translocation of C from a single leaf of sunflower

When single leaves attached at a given node were allowed to carry on photosynthesis in ¹⁴CO₂ for 30 min, younger plants showed a higher proportion of upward translocation than did older plants. Downward translocation of ¹⁴C-photosynthate was stimulated by ATP pre-treatment of the translocating leaf, while upward translocation was not affected by ATP. A similar phenomenon was observed in the translocation of ¹⁴C-sucrose infiltrated into a leaf with or without ATP. Downward translocation of photosynthate was inhibited by DNP pre-treatment of a fed leaf. Upward translocation, however, was not affected by DNP. Thirty min after infiltration of ¹⁴C-glucose into a leaf, almost all the ¹⁴C translocated upwards was found to be in the form of glucose, while a great part of the ¹⁴C translocated downwards was in the form of sucrose. In the case of translocation of infiltrated ¹⁴C-sucrose, ¹⁴C found both above and below the fed leaf was mainly in the form of sucrose.     

Effects of temperature on carbon fixation and carbon budget partitioning in the zooxanthellal symbiosis of Aiptasia pallida (Verrill)

Effects of temperature on carbon fixation rates and partitioning between Aiptasia pallida (Verrill) and its symbiotic alga Symbiodinium microadriaticum Freudenthal were examined by ¹⁴C incubation studies. Total fixation varied strongly with temperature, with an optimum of 32 °C. More photosynthate was translocated to the host at 12 °C (82%) than at 27 °C (63%). Partitioning among three fractions (alcohol soluble, ether soluble, and alcohol/ether insoluble) varied with temperature in Aiptasia pallida, but not in the alga. Relative partitioning between host and alga increased with time in favor of A. pallida when maintained at 12 °C, but absolute levels of translocation to the host did not change; however, photosynthate retention by the alga did decline substantially. Total fixation declined by ≈ 80% after 10 days at 12 °C. Turnover rates of fixed carbon also varied with temperature, as determined by pulse-chase studies, and the effect varied for the different fractions.These results suggest that zooxanthellae are less thermally adaptable than their hosts, and may be especially susceptible to low temperatures. Thermal effects on biochemical partitioning may have great importance in relation to growth and reproduction of animal hosts of zooxanthellae and the viability of the symbiotic relationship. These effects, combined with the pronounced effect of temperature on total photosynthate production, probably play a major role in limitation of zooxanthellal symbioses to warm waters.     

HORMONAL CONTROL OF TRANSLOCATION OF PHOTOSYNTHETICALLY ASSIMILATED 14C IN YOUNG SOYBEAN PLANTS

The normal supply of growth substances to a young soybean plant was altered by removing the plant's apical meristem and replacing this meristem with an aqueous solution of either indole-3-acetic acid (IAA), gibberellic acid (GA), or water. The length of each experiment was 1 hr. In the middle of it, ¹⁴CO₂ was administered to one of the primary leaves of the plant, and at the end distribution of ¹⁴C in various parts of the plant was determined. It was found that an addition of growth substances stimulated translocation in three different ways. Both IAA and GA increased the total amounts of sucrose-¹⁴C translocated, increased the rate of its translocation, and affected the distribution pattern of translocated sucrose throughout the plant. Experiments using IAA-¹⁴C have shown that the action of IAA is on the longitudinal translocation in the stem and not on the transfer of photosynthate from the mesophyll to the conducting tissues of the leaf.     

Use of positron-emitting tracer imaging system for measuring the effect of salinity on temporal and spatial distribution of C tracer and coupling between source and sink organs

Salinity stress affects photosynthate partitioning between sources and sinks of plants, but how it affects these systems is less well understood. Because sources and sinks are closely tied, any adverse effect under suboptimal conditions on one of these is often misinterpreted for an effect on the other. Carbon partitioning is indispensable for stress resistance and good plant growth. In the present study, carbon partitioning in tomato plants (Lycopersicon esculentum L. cv. Momotarou) in a saline (NaCl) environment was studied by feeding radioactive ¹¹C and stable ¹³C isotopes. Pulse-chases were conducted to measure the spatial and temporal distribution of ¹³C. ¹³C was measured by a standard conventional technique, but ¹¹C distribution was monitored using a positron-emitting tracer imaging system (PETIS). Salt stress resulted in reduced carbon translocation toward roots. The majority of the photosynthate accumulated in the leaf. We also observed that the reduction in translocation of carbon occurred well before the salt stress symptoms of reduced photosynthesis and reduced plant growth in salt-exposed plants. The effect on sink activity was also shown by a decrease in stem diameter. In addition, PETIS analysis of ¹¹C translocation indicated that carbon translocation to roots was inhibited under salt conditions without a direct effect on leaf Na accumulation or osmotic stress. These results suggest that NaCl has direct effects on plants, inhibiting carbon partitioning within a few hours of salt exposure without inhibition of source 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.     

Diurnal variation in the functioning of cowpea nodules

Nitrogenase (EC 1.7.99.2) activity of nodules of cowpea (Vigna unguiculata [L.] Walp), maintained under conditions of a 12-hour day at 30°C and 800 to 1,000 microeinsteins per square meter per second (photosynthetically active radiation) and a 12-hour night at 20°C, showed a marked diurnal variation with the total electron flux through the enzyme at night being 60% of that in the photoperiod. This diurnal pattern was, however, due to changes in hydrogen evolution. The rate of nitrogen fixation, measured by short-term ¹⁵N₂ assimilation or estimated from the difference in hydrogen evolution in air or Ar:O₂ (80:20; v/v), showed no diurnal variation. Carbon dioxide released from nodules showed a diurnal variation synchronized with that of nitrogenase functioning and, as a consequence, the apparent `respiratory cost' of nitrogen fixation in the photoperiod was almost double that at night (9.74 ± 0.38 versus 5.70 ± 0.90 moles CO₂ evolved per mole N₂ fixed). Separate carbon and nitrogen balances constructed for nodules during the photoperiod and dark period showed that, at night, nodule functioning required up to 40% less carbohydrate to achieve the same level of nitrogen fixation as during the photoperiod (2.4 versus 1.4 moles hexose per mole N₂ fixed).

Stored reserves of nonstructural carbohydrate of the nodule only partly satisfied the requirement for carbon at night, and fixation was dependent on continued import of translocated assimilates at all times. Measurements of the soluble nitrogen pools of the nodule together with ¹⁵N studies indicated that, both during the day and night, nitrogenous products of fixation were effectively translocated to all organs of the host plant despite low rates of transpiration at night. Reduced fluxes of water through the plant at night were apparently counteracted by increased concentration of nitrogen, especially as ureides, in the xylem stream.

     

Morphology and Physiology of Re-growth of the Tropical Tallgrass Hybrid Pennisetum

Changes in d. wt and ¹⁴C distribution were followed in individualorgans during re-growth of hybrid pennisetum after exposureof the plants to [¹⁴C]carbon dioxide before or after defoliation.The changes were recorded at three temperatures (18/13, 24/19and 30/25 °C day/night) in plants in which the apical meristemhad been left intact or removed at defoliation. Initial re-growth was more rapid and at the expense of stemd. wt losses where the apical meristem had been left intact.Rapid re-growth was associated with breakdown of organic reserves(proteins and polysaccharides) and with the capturing of one-quarterof the net ¹⁴C lost from the whole plant. In decapitated plantsre-growth was delayed until new tillers developed and was rapidthereafter. The pattern of photosynthate mobilization was notaffected by temperature, but the speed of mobilization increasedmarkedly with temperature. The pattern was much the same forphotosynthate fixed before or after defoliation, but a highproportion of the ¹⁴C fixed after defoliation was respired andonly 5 per cent entered subsequent new growth. Pennisetum hybrid, tallgrass, re-growth     

Translocation and accumulation of translocate in the sugar beet petiole

Accumulation of translocate during steady-state labeling of photosynthate was measured in the source leaf petioles of sugar beet (Beta vulgaris L. monogerm hybrid). During an 8-hr period, 2.7% of the translocate or 0.38 μg carbon/min was accumulated per cm petiole. Material was stored mainly as sucrose and as compounds insoluble in 80% ethanol. The minimum peak velocity of translocation approached an average of 54 cm/hr as the specific activity of the ¹⁴CO₂ pulse was progressively increased. The ratio of cross sectional area required for translocation to actual sieve tube area in the petiole was 1.2. A regression analysis of translocation rate versus sieve tube cross sectional area yielded a coefficient of 0.76. The specific mass transfer rate in the petiole was 1.4 g/hr cm² phloem or 4.8 g/hr cm² sieve tube. Histoautoradiographic studies indicated that translocation occurs through the area of phloem occupied by sieve tubes and companion cells while storage occurs in these cells plus cambium and phloem parenchyma cells. The ability of the petiole to act as a sink for translocate is consistent with the concept that storage along path tissue serves to buffer sucrose concentration in the translocate during periods of fluctuating assimilation.     

Effect of Moisture Supply upon Translocation and Storage of C in Sugarcane

Low moisture supply, controlled by 3 methods (adding NaCl to a complete nutrient solution, allowing a cut stalk to wilt, or withholding irrigation in the field), decreased the velocity and percentage rate of translocation of ¹⁴C-photosynthate. The surplus sucrose not used in growth moved more slowly in the phloem and was stored in the stalk.

Low moisture supply depressed translocation of ¹⁴C-photosynthate more severely than it curtailed formation of ¹⁴C-photosynthate in the same leaf: therefore, the effect of moisture supply upon translocation was primary.

Low moisture supply retarded profile development in the stem, and a loss in moisture gradient was associated with a steepened slope of the profile. These results indicate a flow mechanism of translocation rather than diffusion.

Results reported now and previously point to the operation of a slow pressure-flow mechanism particularly during the night but also during the day; superimposed upon this general mass transport is the more rapid process of phototranslocation which is independent of sugar gradients and which can cause the accumulation of sucrose at the storage-sink.

During ripening, storage of sucrose in the stalk may be increased by withholding water because less sucrose is hydrolyzed in transit, less is used in growth, and the slowly moving sucrose has more time for transfer from the phloem to the storage parenchyma.

     

Source-sink relations in maize mutants with starch-deficient endosperms

Partitioning and translocation of photosynthates were compared between a nonmutant genotype (Oh 43) of corn (Zea mays L.) and two starch-deficient endosperm mutants, shruken-2 (sh2) and brittle-1 (bt1), with similar genetic backgrounds. Steady-state levels of ¹⁴CO₂ were supplied to source leaf blades for 2-hour periods, followed by separation and identification of ¹⁴C-assimilates in the leaf, kernel, and along the translocation path. An average of 14.1% of the total ¹⁴C assimilated was translocated to normal kernels, versus 0.9% in sh2 kernels and 2.6% in btl kernels. Over 98% of the kernel ¹⁴C was in free sugars, and further analysis of nonmutant kernels showed 46% of this label in glucose and fructose. Source leaves of mutant plants exported significantly less total photosynthate (24.0% and 36.3% in sh2 and bt1 compared to 48.0% in the normal plants) and accumulated greater portions of label in the insoluble (starch) fraction. Mutant plants also showed lower percentages of photosynthate in the leaf blade and sheath below the exposed blade area. The starch-deficient endosperm mutants influence the partitioning and translocation of photosynthates and provide a valuable tool for the study of source-sink relations.     

Changes in carbohydrate metabolism and assimilate export in starch-excess mutants of Arabidopsis

The aim of this work was to investigate the effects on carbohydrate metabolism of a reduction in the capacity to degrade leaf starch in Arabidopsis. The major roles of leaf starch are to provide carbon for sucrose synthesis, respiration and, in developing leaves, for biosynthesis and growth. Wild-type plants were compared with plants of a starch-excess mutant line (sex4) deficient in a chloroplastic isoform of endoamylase. This mutant has a reduced capacity for starch degradation, leading to an imbalance between starch synthesis and degradation and the gradual accretion of starch as the leaves age. During the night the conversion of starch into sucrose in the mutant is impaired; the leaves of the mutant contained less sucrose than those of the wild type and there was less movement of ¹⁴C-label from starch to sucrose in radio-labelling experiments. Furthermore, the rate of assimilate export to the roots during the night was reduced in the mutant compared with the wild type. During the day however, photosynthetic partitioning was altered in the mutant, with less photosynthate partitioned into starch and more into sugars. Although the sucrose content of the leaves of the mutant was similar to the wild type during the day, the rate of export of sucrose to the roots was increased more than two-fold. The changes in carbohydrate metabolism in the mutant leaves during the day compensate partly for its reduced capacity to synthesize sucrose from starch during the night.     

Seasonal Course of Photosynthesis, Respiration, and Distribution of C in Young Pinus resinosa Trees as Related to Wood Formation

Rates of net photosynthesis and dark respiration, and distribution of ¹⁴C were determined for new (current season's) and old (previous season's) needles at 10 times during the seasonal development of young Pinus resinosa Ait. trees. The seasonal changes in these factors associated with the development of the new shoot were related to known seasonal patterns of wood formation.

Net photosynthesis per gram of needle dry weight (photosynthetic efficiency) was maximum in the old needles at the time of first new needle elongation; at the same time translocation of ¹⁴C from old to new needles was greatest. Photosynthetic efficiency of new needles was maximum at the end of the period of rapid new needle elongation, when the new needles also began exporting much greater quantities of ¹⁴C to other plant parts. In particular, the amount translocated from the new needles to the stem was greatly increased. At this time thick-walled xylem cells were first observed in the stem.

These results, together with those of previous studies, indicate that the production of thick-walled xylem tracheids normally associated with latewood is physiologically correlated with maturation of the current season's needles. Because there is a lesser demand for photosynthate in the new shoot and a high rate of photosynthesis in the whole plant at the time of new needle maturity, a sharply increased amount of photosynthate becomes available for wall synthesis by cambial derivatives.

     

Source pool kinetics for C-photosynthate translocation in morning glory and soybean

The kinetic behavior of translocation profiles indicates that their shape is determined largely by the rate at which tracer enters the sieve tubes in the source leaf. Confirmation of this relationship was sought by investigating the kinetics of ¹⁴C in the immediate source pool for translocated sucrose in soybean (Glycine max L., cv. Bragg) and morning glory (Ipomea nil Roth, cv. Scarlet O'Hara) leaves. Quantitative microautoradiography was used to follow the water-soluble ¹⁴C contents of the companion cells in minor veins after pulse-labeling with ¹⁴CO₂. In both morning glory and soybean, the observed kinetics in the companion cells matched reasonably well those expected from the shape of the translocation profiles.

Marked compartmentation of sucrose was evident in soybean leaves in that the specific radioactivity of total leaf sucrose was greatest immediately after labeling and quickly declined, whereas labeling in the companion cells was low at first and did not reach a maximum for about 35 minutes. In morning glory leaves, the kinetics of sucrose specific radioactivity and of companion cell-labeling more closely paralleled one another.

     

Translocation of radioactive kinetin

Kinetin has generally been thought to be immobile in plants. This was confirmed in the case of laminar applications in this study, but not in regard to petiole, vein, or root applications. Radioactivity from kinetin-8-¹⁴C (Kn^*) moved freely in the vascular system of several types of leaves. This movement was usually distal to the point of application and seemed to occur with the transpiration stream. Basipetal as well as acropetal translocation of radioactive kinetin was achieved in tobacco leaves. The translocated material was extracted from veinal tissue, shown to be radioactive, and to be able to retard senescence. Similar but less decisive results were obtained from agar blocks inserted into the vascular system of leaves receiving Kn^* by petiole uptake.

A bioassay employing disks from primary bean leaves was developed for the qualitative determination of substances like kinetin which possess the ability to retard chlorophyll breakdown and plant senescence. The use of radioactive kinetin provided a refinement in this bioassay because treated non-senescent areas could be correlated with exposed areas on radioautographs made from dried leaf disks.

Root treatments showed that cotton seedlings did not take up Kn^* but that similarly treated tobacco seedlings both absorbed and translocated the isotope readily.

     

Gibberellic-Acid-Promoted Transport of Assimilates in Stems of Phaseolus vulgaris. Effects on Assimilate Accumulation by the Sink

Under conditions of apoplastic unloading from the sieve element-companioncell (se-cc) complexes in fully-elongated stems of Phaseolusvulgaris plants, gjbberellic acid (GA₃ stimulated in vitro uptakeof [¹⁴C]sucrose by the stem tissues. The GA₃, response dependedupon the incubate containing calcium ions and being bufferedat pH 6. The GA₃ action could be accounted for by a reductionin the Michaelis-Menten constant of the uptake process. Promotedtransport by GA₃ in the decapitated stems resulted in all thetissues accumulating higher levels of [¹⁴C]photosynthates. Comparisonof this response with that for in vitro uptake of [¹⁴C]sucroseindicated that GA₃ stimulation of the sucrose uptake processcontributed significantly to the accumulation of photosynthatesby the pith alone. The bulk of enhanced photosynthate accumulationby the remaining stem tissues can be accounted for by a GA,-inducedelevation of the apoplast sucrose concentration. In terms ofonset and change in rate, the time-course kinetics of GA₃ stimulationof [¹⁴C]photosynthate transport and of in vitro [¹⁴CJsucroseuptake were found to be similar. It is proposed that GA₃ promotionof photosynthate accumulation by the pith tissues is a minorcontributing factor to GA₃ regulation of phloem translocation Phaseolus vulgaris L., french bean, stem, assimilate transport, gibberellic acid, rink accumulation     

AtRBOH I confers submergence tolerance and is involved in auxin-mediated signaling pathways under hypoxic stress

Satsuma mandarin fruit (Citrus unshiu Mark.) photosynthesizes as comparable to leaf at about 100 days after full bloom (DAFB). In this study, translocation and accumulation of fruit-fixed photosynthate were investigated by using ¹⁴CO₂. When fruit at 108 DAFB was exposed to ¹⁴CO₂ for 48 h under 135 photosynthetic photon flux density (PPFD), ¹⁴C-sucrose, ¹⁴C-glucose and ¹⁴C-fructose were detected not only in flavedo but juice sac; more than 50?% of fruit assimilated ¹⁴C-sugars were present in juice sac. Thus, majority of rind-fixed photosynthate are infiltrated into juice sac and accumulated there within 48 h after assimilation. Although ¹⁴C-sucrose was predominant at flavedo where high SS (sucrose synthase) activity toward synthesis was present, the amount decreased gradually from the outside (flavedo) to the inside (juice sac) of fruit. In vascular bundle, strong SS toward cleavage and soluble acid invertase activities were involved, and ¹⁴C-fructose was predominant in juice sac. Accordingly, rind-fixed photosynthate is once converted to sucrose, the translocated sugar in Citrus, at flavedo by SS toward synthesis, and loaded on vascular bundle through symplastic and/or apoplastic movement in the albedo tissue. In the vascular bundle, sucrose may be degraded by SS toward cleavage and invertase, and resulting hexoses transported symplastically to the juice sac through juice stalk.     

Translocation of [C]sucrose in wheat

Flag leaves of wheat plants (Triticum aestivum L. em. Thell. cv `Duke') were supplied with ¹⁴C(glucosyl)sucrose. Translocated [¹⁴C]sucrose was isolated, then hydrolyzed. Label appeared in both the hexose moieties indicating that some randomization of label had occurred. However, near the radioactive front essentially all of the ¹⁴C was in the glucose moiety, suggesting that randomization occurred after unloading, supporting the conclusion that sucrose was taken up intact by phloem and translocated unaltered.     

Soil temperature affects carbon allocation within arbuscular mycorrhizal networks and carbon transport from plant to fungus

How soil carbon balance will be affected by plant–mycorrhizal interactions under future climate scenarios remains a significant unknown in our ability to forecast ecosystem carbon storage and fluxes. We examined the effects of soil temperature (14, 20, 26 °C) on the structure and extent of a multispecies community of arbuscular mycorrhizal (AM) fungi associated with Plantago lanceolata. To isolate fungi from roots, we used a mesh‐divided pot system with separate hyphal compartments near and away from the plant. A ¹³C pulse label was then used to trace the flow of recently fixed photosynthate from plants into belowground pools and respiration. Temperature significantly altered the structure and allocation of the AM hyphal network, with a switch from more vesicles (storage) in cooled soils to more extensive extraradical hyphal networks (growth) in warmed soils. As soil temperature increased, we also observed an increase in the speed at which plant photosynthate was transferred to and respired by roots and AM fungi coupled with an increase in the amount of carbon respired per unit hyphal length. These differences were largely independent of plant size and rates of photosynthesis. In a warmer world, we would therefore expect more carbon losses to the atmosphere from AM fungal respiration, which are unlikely to be balanced by increased growth of AM fungal hyphae.     

Pathway and sink activity for photosynthate translocation in <Emphasis Type="Italic">Pisolithus</Emphasis> extraradical mycelium of ectomycorrhizal <Emphasis Type="Italic">Pinus thunbergii</Emphasis> seedlings

The purpose of this study was to identify the pathway and sink activity of photosynthate translocation in the extraradical mycelium (ERM) of a Pisolithus isolate. We labelled ectomycorrhizal (ECM) Pinus thunbergii seedlings with ¹⁴CO₂ and followed ¹⁴C distribution within the ERM by autoradiography. ¹⁴C photosynthate translocation in the ERM resulted in ¹⁴C distribution in rhizomorphs throughout the ERM, with ¹⁴C accumulation at the front. When most radial mycelial connections between ECM root tips and the ERM front were cut, the whole allocation of ¹⁴C photosynthates to the ERM was reduced. However, the overall pattern of ¹⁴C distribution in the ERM was maintained even in regions immediately above and below the cut, with no local ¹⁴C depletion or accumulation. We inferred from this result that every portion in the ERM has a significant sink activity and a definite sink capacity for photosynthates and that photosynthates detour the cut and reach throughout the ERM by translocation in every direction. Next, we prepared paired ECM seedlings, ERMs of which had been connected with each other by hyphal fusion, alongside, labelled the left seedling with ¹⁴CO₂, and shaded none, one or both of them. ¹⁴C photosynthates were acropetally and basipetally translocated from the left ERM to ECM root tips of the right seedling through rhizomorphs in the left and right ERMs, respectively. With the left seedling illuminated, ¹⁴C translocation from the left to the right ERM increased by shading the right seedling. This result suggests that reduced photosynthate transfer from the host to its ERM increased sink activity of the ERM.