Transport of sodium into the xylem exudate of tobacco

When tobacco (Nicotiana tabacum L. var. Virginia Gold) plants were pretreated with Na (²²Na) several days before detopping, from 2.3 to 4.9% of Na previously accumulated in roots appeared in the xylem exudate in 7 days after detopping. Na from the external medium, however, was readily transported to the exudate. Moreover, the amount of the pretreatment Na that was transported to the exudate was not influenced by the presence of Na in the external medium. When Na was present in the external medium after detopping, about 4% (with an NaNO₃ post treatment) to 10% (with an NaCl post treatment) of the Na transported to the xylem in the 7 days following detopping originated in the vacuoles. Nitrate salts of K or Na in the external medium after detopping resulted in transport of large quantities of the respective cation to the exudate, but not in increased transport of the pretreatment Na. A much larger percentage of the K that was accumulated after detopping than of the Na similarly accumulated was transferred to the xylem exudate.     

Effect of EDTA on transport forms of manganese in maize xylem exudate

Xylem exudates were collected from 20-day-old maize seedlings grown for 24 h in labelled manganese chloride solution (10 ppm) with or without EDTA (18 mgl^-1). The distribution of radioactivity on paper ohromatograms of exudates did not show the existence of free Mn. Fractionation of exudate constituents by gel filtration on Sephadex G-26 showed that⁵⁴Mn is associated with three UV (254 nm) -absorbing fractions. Two fractions contain amino acids (M. wt.:≧ 10 000 and ∼ 900). The third fraction (M. wt. ∼ 600) however, appears to be a carrier of a different type, which is practically amino acid free, although it contains 10.23% N (by elementary analysis). The properties of this carrier are discussed. Application of EDTA has favoured the; transport of Mn in xylem sap with fractions of relatively lower molecular weights.     

The effect of manganese and EDTA on the balance of nitrogen in maize xylem exudate

The effect of manganese chloride (10 mg Mn l^-1), EDTA (18 mg l^-1) and a mixture of these compounds on the nitrogen balance in maize xylem exudate was investigated. The compounds were applied as experimental solutions to the roots of 20 day old plants 24 h before excision. Application of Mn resulting in a lowered nitrogen level in the xylem exudate increased the relative content of the organic N-compounds in the exudate, particularly that of free amino acids. EDTA appreciably enhanced the content of total nitrogen in xylem exudate, however no significant changes were found in the proportion of inorganic and organic N-compounds in comparison with the water control. The significant features of the free amino acid exudate fraction of all experimental variants were, among others, the relatively high lysine content and the absence of proline arid sulphur containing amino acids. By gel filtration on Sephadex G-25 of xylem exudates UV (254 nm) absorbing fractions, four in the water control, five in Mn and six in EDTA variants were isolated. The UV absorbing fractions with the exception of one in each experimental variant (K_av 0.73 in water control, K_av 0.61 in Mn-variant, K_av 0.75 in EDTA and Mn + EDTA variant) were of peptide character, proline and sulphur-containing amino acids were missing in them. In the exceptional UV absorbing fractions (K_av 0.61–0.75) in spite of their high N-content (10.23%) after hydrolysis practically no amino acid could be detected.     

Translocation of nickel in xylem exudate of plants

Topped plants of tomato (Lycopersicon esculentum), cucumber (Cucumis sativus), corn (Zea mays), carrot (Daucus carota), and peanut (Arachis hypogaea) were treated with 0.5 to 50 micromolar Ni (containing ⁶³Ni) in nutrient solutions. Xylem exudate was collected for 10 hours or, in the case of corn, for 20 hours at 5-hour intervals. Electrophoresis of nutrient solution distributed all Ni cathodically as inorganic Ni²⁺. Low concentrations of Ni in tomato exudate migrated anodically, presumably bound to organic anion (carrier). However, this carrier became saturated at about 2 micromolar Ni in exudate, and excess Ni ran cathodically. Most of the Ni in cucumber, corn, carrot, and peanut exudate ran anodically, and its migration rate was identical for all exudates. Peanut root sap contained 14 to 735 micromolar Ni. The anodic Ni carriers in root sap and exudate appear identical. The carrier in root sap became saturated near 100 micromolar Ni, as shown by cathodic streaking of Ni exceeding that concentration. It appears that all five species translocate low concentrations of Ni in the same anionic form.     

Effect of calcium on transport of cations to the xylem exudate of tobacco

Summary Roots of detopped tobacco plants (Nicotiana tabacum var. Virginia Gold) were exposed to Na, K, and Ca salts or to water, and cation transfer to xylem vessels was measured. In some cases plants had been exposed to Na in addition to regular nutrient solutions before detopping. Calcium in the external medium greatly depressed the transport of Na from the external medium to the xylem vessels and it often stimulated the transfer of K from the external medium to the xylem vessels. The K/Na ratio in the exudate thus was dependent upon the Ca content of the external medium under these conditions. In contrast, externally applied Ca or Ca deficiency had very little effect on the transfer of preaccumulated K and Na from compartments within roots to the xylem vessels. The K/Na ratio in the exudate under these conditions was not related to Ca levels nor to mild Ca deficiency. The ratios decreased with time after detopping regardless of Ca level. Intact plants accumulated more Na than did root systems of detopped plants in a 6-day period.Riverside University of CaliforniaSoil Science and Agricultural Engineering     

Cation and anion balance in the xylem exudate of tobacco roots

Summary The sum of Na, K, Ca, Mg in the exudate of tobacco generally exceeded the sum of mineral anions. Insufficient organic acids were present to account for the differences and bicarbonate appeared to be the other anion involved. Amino acids were present in very low concentrations relative to mineral cations. When nitrate salts only were in the external solutions, the anions were mostly, but not entirely, nitrate. When chloride salts only were in the external solutions, the cations far exceeded the level of mineral anions in the exudate. It is postulated that nitrate is actively transported when nitrate salts are in the external solution regardless of the cation, but when anions other than nitrate are in the external solution, the cations are actively transported with the anions passively following. Nitrate transport was via a symplasm, but that of the other anions seemed to be different. When bicarbonate is the only anion in the external solution and when present at relatively high concentrations (5 × 10⁻³ M or higher), the volume of exudate is decreased. It appears that the organic acids which were synthesized as a result of the bicarbonate absorption were not transferred to the xylem vessels.     

Movement to the xylem exudate of rubidium accumulated in the apex of corn roots

Apical segments of roots of corn (Zea mays, L.) were excised and mounted in experimental salt solutions containing ⁸⁶RbCl and CaCl₂. Xylem exudates were collected hourly beginning at the 21st hour. At the 24th hour, experimental solutions were replaced with solutions of other chlorides (nonlabeled) of Ca²⁺ ± some monovalent cation.     

Translocation of iron citrate and phosphorus in xylem exudate of soybean

Soybean plants, Glycine max (L.) Merrill, in standard solution received 2.5 μm ferric ethylenediamine di(o-hydroxyphenylacetate (FeEDDHA) and 0 to 128 μm phosphorus. Their stem exudates contained: 32 to 52 μm Fe, 120 to 5000 μm P, and 120 to 165 μm citrate. Electrophoresis of exudates with high P caused Fe trailing that precluded identification of any major form of Fe. Exudate with low P gave an anodic band of Fe citrate as the major Fe compound. Phosphate added to exudate in vitro depressed the Fe citrate peak and cause Fe trailing. EDDHA added to exudate in vitro pulled Fe from Fe citrate; citrate then migrated as a slower form and Fe migrated as FeEDDHA. A modified preculture system, involving 2-day renewals of 0.2 μm FeEDDHA with 3.2, 9.6, or 16 μm P and low levels of other ions, controlled pH depression and produced considerable change in citrate and P levels. The exudates contained: 45 to 57 μm Fe, 200 to 925 μm P, and 340 to 1025 μm citrate. The high citrate was from plants grown with low P. The major form of Fe in the exudates was Fe citrate. This is probably the form translocated in the plants.     

Effect of temperature on potassium and calcium transport from within the root to the xylem exudate of tobacco

Summary Detopped root systems of tobacco plants (Nicotiana tabacum L. var. Virginia Gold) were exposed to air at different temperatures (14 to 23°C). The volume of xylem exudate was related to temperature and decreasing the temperature resulted in a lag in the appearance of exudate. For 4 and 8 days of exudation the amounts of cations transported in the exudate were temperature dependent. The Q₁₀ values were higher than usually encountered in biological reactions; the mean value was around 6. This work was supported in part by National Science Foundation Grant GB8170. University of California     

Abscisic Acid promotes both volume flow and ion release to the xylem in sunflower roots

The effect of abscisic acid (ABA) on the exudation process in excised sunflower root was investigated. A promotion of both ion flux and volume exudation rate was observed. Cutting off the external supply of ions to the roots markedly increased the magnitude of the ABA effect. The promotive effect of ABA on exudation rate was extremely rapid (less than 6 minutes) and seemed to be biphasic. Reasons are given for relating the ABA effect to two separate actions, namely: (a) on water permeability; and (b) on the release of ions both from vacuoles to the cytoplasm and from symplasm to the xylem. Kinetin inhibited movement of ions to the xylem and its effect did not appear to be related to that of the ABA.     

Diurnal and temporal changes in the chemical profile of xylem exudate from Vitis rotundifolia

Positive root pressure in Vitis rotundifolia Michx. cv. Noble was employed to quantify diurnal and temporal changes in the chemical profile of xylem exudate. Xylem fluid osmolarity (7.2 to 16.8 m M ), water flux (8.2 to 18.5 ml h⁻¹) and solute flux (0.7 to 2.2 mmol h⁻¹) from a cut spur exhibited a diurnal pattern with maxima during midday and minima at night. Total osmolarity was similar to the sum of all organic and inorganic entitites quantified, indicating that the major solutes have been identified. Total amino acid and organic acid concentration were about equal (2 to 7 m M ), and sugars accounted for a minor fraction of the total profile (<0.2 m M ). Glutamine represented ca 80% of the organic N and 70% of the total N transported in the xylem fluid. A circadian rhythm in water flux and net flux of most organic and inorganic entities was observed with maxima during midday and minima at night. The increase in xylem fluid osmolarity occurring during midday was primarily a consequence of increased organic acid (oxalic, citric, tartaric, malic and succinic acids) and ion (NH₄⁺, No₃⁻, P and Ca) concentration. A diurnal cycle in amino acid concentration was less clear. The concentration of individual organic and inorganic entities varied asynchronously with time. Xylem solute was comprised of 80% organic and 20% inorganic components when collected 5 min to 2 h after the commencement of bleeding, but the ratio of organic to inorganic components fell to about 50% after 7 days.     

Transport of nitrogen in the xylem of soybean plants

Experiments were conducted to characterize the distribution of N compounds in the xylem sap of nodulated and nonnodulated soybean plants through development and to determine the effects of exogenous N on the distribution of N compounds in the xylem. Xylem sap was collected from nodulated and nonnodulated greenhouse-grown soybean plants (Glycine max [L.] Merr. “Ransom”) from the vegetative phase to the pod-filling phase. The sum of the nitrogen in the amino acid, nitrate, ureide (allantoic acid and allantoin), and ammonium fractions of the sap from both types of plants agreed closely with total N as assayed by a Kjeldahl technique. Sap from nodulated plants supplied with N-free nutrient solution contained seasonal averages of 78 and 20% of the total N as ureide-N and amino acid-N, respectively. Sap from nonnodulated plants supplied with a 20 millimolar KNO₃ nutrient solution contained seasonal averages of 6, 36, and 58% of total N as ureide-N, amino acid-N, and nitrate-N, respectively. Allantoic acid was the predominant ureide in the xylem sap and asparagine was the predominant amino acid. When well nodulated plants were supplied with 20 millimolar KNO₃, beginning at 65 days, C₂H₂ reduction (N₂ fixation) decreased relative to nontreated plants and there was a concomitant decrease in the ureide content of the sap. A positive correlation (r = 0.89) was found between the ureide levels in xylem sap and nodule dry weights when either exogenous nitrate-N or urea-N was supplied at 10 and 20 millimolar concentrations to inoculated plants. The results demonstrate that ureides play a dominant role in N transport in nodulated soybeans and that the synthesis of ureides is largely dependent upon nodulation and N₂ fixation.     

Transport of zinc and manganese to developing wheat grains

An understanding of the transport pathway used by Zn and Mn to enter developing grains may allow measures to increase the Zn and Mn content of wheat grain grown on Zn/Mn deficient soils. For this reason, transport of Zn and Mn into developing grains of wheat ( Triticum aestivum L. cv. Aroona) was investigated. Detached ears (18–22 days post-anthesis) were cultured for 48 h in a solution containing 185 kBq of ⁶⁵Zn and 185 kBq of ⁵⁴Mn. Transport of ⁶⁵Zn to the grain was unaffected by removal of glumes but was slightly reduced after the lemma was removed. Heat girdling the peduncle slightly reduced the amount of ⁶⁵Zn transported to the grain, whilst heat girdling the rachilla reduced transport of ⁶⁵Zn to the grain to a greater degree, suggesting phloem transport to the rachilla. The transport inhibitor CCCP (carbonyl cyanide m -chlorophenyl hydrazone) blocked ⁶⁵Zn transport to grain but not to lemma and glumes. Removing glumes and lemma and heat girdling the peduncle did not affect transport of ⁵⁴Mn, but transport was slightly affected by heat girdling the rachilla, indicating xylem transport. CCCP blocked transport of ⁵⁴Mn into the grain but not to lemma and glumes. It was concluded that xylem-to-phloem transfer of Zn occurs in the rachis and to a lesser extent in peduncle and lemma. The results suggest that the lemma may be an important site for phloem loading when the concentration of Zn within the xylem is high. The data also suggest that Mn was predominantly translocated to the spikelets in the xylem, but that transport to the grain was dependent upon membrane transport before entering the grain. Phloem loading of Mn into the grain vascular system may have occurred at the site of xylem discontinuity in the floral axis.     

Ion gradients in xylem exudate and guttation fluid related to tissue ion levels along primary leaves of barley

The concentration of ions in plant cells and tissues is an essential factor in determining physiological function. In the present study, we established that concentration gradients of mobile ions exist in both xylem exudates and tissues within a barley (Hordeum vulgare) primary leaf. For K⁺ and NO₃^?, ion concentrations generally decreased from the leaf base to the tip in both xylem exudates and tissues. Ion gradients were also found for Pi and Cl^? in the xylem. The hydathode strongly absorbed Pi and re‐translocated it to the rest of the plant, whereas Cl^? was extruded. The ion concentration gradients developed early during leaf growth, increased as the tissue aged and remained under both high and low transpiration conditions. Measurement of the expression profiles of Pi, K⁺ and NO₃^? transporters along the longitudinal axis of the leaf revealed that some transporters are more expressed at the hydathode, but for most transporters, there was no significant variation along the leaf. The mechanisms by which longitudinal ion gradients develop in leaves and their physiological functions are discussed.     

Arabidopsis and sunflower plants with increased xylem area show enhanced seed yield

Plant architecture plasticity determines the efficiency at harvesting and plays a major role defining biomass and seed yield. We observed that several previously described transgenic genotypes exhibiting increased seed yield also show wider stems and more vascular bundles than wild‐type plants. Here, the relationship between these characteristics and seed yield was investigated. Hanging weight on the main stem of Arabidopsis plants provoked significant stem widening. Such widening was accompanied by an increase in the number of vascular bundles and about 100% of yield increase. In parallel, lignin deposition diminished. Vascular bundle formation started in the upper internode and continued downstream. AUX/LAX carriers were essential for this response. The increase of vascular bundles was reverted 3 weeks after the treatment leading to an enlarged xylem area. Aux1, lax1, and lax3 mutant plants were also able to enlarge their stems after the treatment, whereas lax2 plants did not. However, none of these mutants exhibited more vascular bundles or seed yield compared with untreated plants. Weight‐induced xylem area enhancement and increased seed yield were also observed in sunflower plants. Altogether these results showed a strong correlation between the number of vascular bundles and enhanced seed yield under a long‐day photoperiod. Furthermore, changes in the levels of auxin carriers affected both these processes in the same manner, suggesting that there may be an underlying causality.     

Transport of manganese intoSaccharomyces cerevisiae

The uptake of Mn2+ by Saccharomyces cerevisiae at the expense of endogenous sources of energy depends on the stage of culture development and is maximum in the middle of the exponential phase. The ability of cells to take up Mn+ is related to the content of intracellular potassium at all stages of growth, to the content of ATP during the exponential phase and it is not related to the content of inorganic polyphosphates. The uptake is inhibited by oligomycin (25 microgram/ml) by 50-85% and under anaerobic conditions by 10-50%, depending on the stage of growth, indicating the role of aerobic phosphorylation in the process. The uptake of Mn+ is apparently associated with a hydrolysis of low-molecular weight polyphosphates and ATP, as well as with the exit of K+ from cells.     

Nitrate reductase activity and nitrogen compounds in xylem exudate of Juglans nigra seedlings: relation to nitrogen source and supply

Nitrogen (N) limits plant productivity and its uptake and assimilation may be regulated by N source, N availability, and nitrate reductase activity (NRA). Knowledge of how these factors interact to affect N uptake and assimilation processes in woody angiosperms is limited. We fertilized 1-year-old, half-sib black walnut (Juglans nigra L.) seedlings with ammonium (NH₄ ⁺) [as (NH₄)₂SO₄], nitrate (NO₃ ⁻) (as NaNO₃), or a mixed N source (NH₄NO₃) at 0, 800, or 1,600 mg N plant⁻¹ season⁻¹. Two months following final fertilization, growth, in vivo NRA, plant N status, and xylem exudate N composition were assessed. Specific leaf NRA was higher in NO₃ ⁻-fed and NH₄NO₃-fed plants compared to observed responses in NH₄ ⁺-fed seedlings. Regardless of N source, N addition increased the proportion of amino acids (AA) in xylem exudate, inferring greater NRA in roots, which suggests higher energy cost to plants. Root total NRA was 37% higher in NO₃ ⁻-fed than in NH₄ ⁺-fed plants. Exogenous NO₃ ⁻ was assimilated in roots or stored, so no difference was observed in NO₃ ⁻ levels transported in xylem. Black walnut seedling growth and physiology were generally favored by the mixed N source over NO₃ ⁻ or NH₄ ⁺ alone, suggesting NH₄NO₃ is required to maximize productivity in black walnut. Our findings indicate that black walnut seedling responses to N source and level contrast markedly with results noted for woody gymnosperms or herbaceous angiosperms.     

Iron-dependent changes of heavy metals,nicotianamine, and citrate in different plant organs and in the xylem exudate of two tomato genotypes. Nicotianamine as possible copper translocator

The influence of Fe nutrition on the distribution of the heavy metals Fe, Mn, Zn, and Cu and of the heavy metal chelators nicotianamine (NA) and citrate in 6 different shoot and 3 different root parts and in xylem exudate of a NA-containing tomato wild type and its NA-less mutant was investigated. Under the same Fe supply the mutant showed higher Fe, Mn, and Zn concentrations in all organs investigated, with exception of the shoot apex. The Cu concentration in the mutant was only in root parts higher than in the wild type but much lower in leaves. Analyses of xylem exudate showed that Fe, Mn, and Zn were readily translocated by both genotypes from the roots to the shoot at all levels of Fe supply, whereas in the absence of NA, Cu was only poorly transported. Citrate as main Fe chelator in the xylem was present in high concentrations in xylem exudate of the wild type under low Fe supply but in the mutant also at 10 M FeEDTA. NA occurred in xylem exudate of the wild type in concentrations high enough to chelate heavy metal ions.Generally, high Fe supply induced a decrease of Mn, Cu, and Zn concentrations in all organs of the wild type whereas high concentrations were observed in most cases under Fe deficiency. A positive correlation between Fe supply and NA concentration existed only in the shoot apex and in the xylem exudate of wild type plants. From the correlation between Cu and NA translocation and from the high stability constant of the NA-Cu-complex (log K=18.6) it is concluded that NA is a chelator for Cu in the xylem, whereas the translocation of Fe, Mn, and Zn is independent of NA.     

Effects of external cations and respiratory inhibitors on electrical potential of the xylem exudate of excised corn roots

Glass capillary microelectrodes were used to study the electrical potential difference (PD) between the xylem exudate of excised corn roots, Zea mays L. Golden Bantam hybrid, and the external solution. A survey of the effects of various ions on the PD was made. With 1 mm single salt solutions, the PD was between 25 and 50 mv, exudate negative. The PD responded to concentration differences in single salt solutions of K⁺, Na⁺, and Ca²⁺ in a manner suggestive of cation selectivity and cation diffusion potentials. With Ca²⁺ present, the PD was insensitive to concentration changes of other cations. Substitution of NO₃⁻ for Cl⁻ in K⁺ solutions increased the PD by 2 to 5 mv, although in general the PD showed little response to anion concentration changes. The PD was partially abolished by cyanide. The remaining fraction of the PD was sensitive to concentration changes in external K⁺, and we postulate that the PD is the result of both a diffusion potential and an electrogenic pump.